LIBRARY OF THE UNIVERSITY OF ILLINOIS AT URBANA-CHAMPAIGN 590.5 FI V. 4-8 ; VL5f- CM CD ^~ CO CNI >- BIOLOGY The person charging this material is re- sponsible for its return to the library from which it was withdrawn on or before the Latest Date stamped below. Theft, mufilatlon, and underlining of books ore reasons for discipiinary action and may resuit in dismissal from the University. To renew call Telephone Center, 333-8400 UNIVERSITY OF lUINOIS LIBRARY AT URBANA-CHAMPAIGN APR-efT9M hHr 0 3 1387 L16I— O-1096 A STUDY OF THE DIURNAL SQUIRRELS, SCIURINAE, OF THE INDIAN AND INDOCHINESE SUBREGIONS JOSEPH CURTIS MOORE AND GEORGE H. H. TATE FIELDIANA: ZOOLOGY VOLUME 48 Published by CHICAGO NATURAL HISTORY MUSEUM MAY 28, 1965 BldLOSY LIBRAW 101 B'JRRlLLHAli- FIELDIANA: ZOOLOGY A Continuation of the ZOOLOGICAL SERIES 0/ FIELD MUSEUM OF NATURAL HISTORY VOLUME 48 CHICAGO NATURAL HISTORY MUSEUM CHICAGO, U.S.A. 1965 A STUDY OF THE DIURNAL SQUIRRELS, SCIURINAE, OF THE INDIAN AND INDOCHINESE SUBREGIONS A STUDY OF THE DIURNAL SQUIRRELS, SCIURINAE, OF THE INDIAN AND INDOCHINESE SUBREGIONS JOSEPH CURTIS MOORE Curator, Division of Mammals AND GEORGE H. H. TATE The Late Curator of Mammals, American Museum of Natural History FIELDIANA: ZOOLOGY VOLUME 48 Published by CHICAGO NATURAL HISTORY MUSEUM MAY 28, 1965 Edited by Edward G. Nash Library of Congress Catalog Card Number: 65-22^9 PRINTED IN THE UNITED STATES OF AMERICA BY CHICAGO NATURAL HISTORY MUSEUM PRESS CONTENTS PAGE List of Text Figures 5 Introduction 7 Acknowledgements 8 Scope 10 Material 11 Methods 13 Color 14 Form of Presentation 15 Size and Dimensions 15 Order 16 Taxonomic History 18 Classification 20 Giant Squirrels, Genus Ratufa 29 Species macroura, Ceylonese Giant Squirrel 34 Species indica, Indian Giant Squirrel 41 Species bicolor. Pied Giant Squirrel 49 Indian striped Squirrels, Genus Funambulus 65 Species pennanti, Five-striped Indian Squirrel 72 Species palmarum, Indian Palm Squirrel 77 Species tristriatus, Western Ghats Squirrel 87 Species layardi, Sinhalese Jungle Squirrel 91 Species sublineattis, Dusky Indian Jungle Squirrel 95 Common Tree Squirrels, Genus Callosciurus 99 Species erythraeus, Red-bellied Squirrel 101 Species flavimanus, Belly-banded Squirrel 120 Species /errM^mcMS, Pegu Red Squirrel 158 Species /fn/ai/som, Siamese Squirrel 161 Species caniceps, Tenasserim Squirrel 187 Species phayrei, Shan Highlands Squirrel 201 Species inornatus, Laotian Squirrel 209 Species pygerythrus, Irrawaddy Squirrel 213 Striped Tree Squirrels, Genus Tamiops 231 Species mcclellandi, Burmese Striped Tree Squirrel 235 Species rodolphei, Cambodian Striped Tree Squirrel 243 3 4 CONTENTS PAGE Species smnhoei, Chinese Striped Tree Squirrel 246 Species maritimus, Maritime Striped Tree Squirrel 251 Plain Long-Nosed squirrels, Genus Dremomys 259 Species lokriah, Himalayan Long-nosed Squirrel 263 Species pernyi, Chinese Long-nosed Squirrel 269 Species rufigenis, Red-cheeked Long-nosed Squirrel 278 Species pyrrhomerus, Red-hipped, Long-nosed Squirrel 283 Species evereiti, Montane Bornean Long-nosed Squirrel 289 Indochinese Ground Squirrel, Genus Menetes 293 Species berdmorei, Indochinese Ground Squirrel 294 Chinese Rock Squirrels, Genus Sciurotamias 303 Species davidianus, Chinese Rock Squirrel 304 Species forresti, Chinese Cliff Squirrel 309 Summary 311 Gazetteer 313 References 337 Index 345 LIST OF TEXT FIGURES PAGE 1. Skull of pied giant squirrel, Ratufa bicolor 30 2. Range of Sinhalese giant squirrel, Ratufa macroura 35 3. Range of Indian giant squirrel, Ratufa indica 40 4. Range of pied giant squirrel, Ratiifa bicolor 50 5. Skull of Western Ghats squirrel, Funambulus tristriatus 67 6. Skull of Indian jungle squirrel, Funambulus sublineatus 70 7. Range of five-striped Indian squirrel, Funambulus pennanti 73 8. Range of Indian palm squirrel, Funambulus palmarum 78 9. Range of Western Ghats squirrel, Funambulus tristriatus 88 10. Range of bright-striped squirrel, Funambulus layardi 92 11. Range of Indian jungle squirrel, Funambulus sublineatus 96 12. Skull of Irrawaddy squirrel, Callosciurus pygerythrus 101 13. Ranges of red-bellied squirrel, Callosciurus erythraeus, and the belly- banded squirrel, Callosciurus flavimanus 122 14. Ranges of Pegu red squirrel, Callosciurus ferrugineus, and Thailand tree squirrel, Callosciurus finlaysoni 162 15. Ranges of the Tenasserim squirrel, Callosciurus caniceps; Shan Highlands squirrel, C. phayrei; and Laotian squirrel, C inornaius 203 16. Range of the Irrawaddy squirrel, Callosciurus pygerythrus 214 17. Skull of striped tree squirrel, Tamiops svnnhoei 233 18. Ranges of striped tree squirrels, Tamiops mcclellandi and T. rodolphei . . 236 19. Ranges of striped tree squirrels, Tamiops sunnhoei and T. maritimus (part) 247 20. Range of striped tree squirrel, Tamiops maritimus (part) 252 21. Composite ranges of species of the genus Dremomys 260 22. Skull of Himalayan long-nosed squirrel, Dremomys lokriah 261 23. Skull of Chinese long-nosed squirrel, Dremomys pernyi 261 24. Range of Himalayan long-nosed squirrel, Dremomys lokriah 264 25. Range of Chinese long-nosed squirrel, Dremomys pernyi 270 26. Range of red-cheeked squirrel, Dremomys rufigenis 279 27. Range of red-hipped squirrel, Dremomys pyrrhom^rus 284 28. Orbitonasal length histogram for species of Dremomys 285 29. Length of orbit histogram for species of Dremomys 286 30. Skull of Siamese ground squirrel, Menetes berdmorei 293 31. Range of Siamese ground squirrel, Menetes berdmorei 295 32. Skull of Chinese rock squirrel, Sciurotamias damdianus 304 33. Ranges of Chinese rock squirrel, Sciurotamias damdianus, and Chinese cliff squirrel, S. forresti • 305 5 INTRODUCTION The present study was initiated in connection with the identifi- cation of a large collection of Oriental squirrels acquired by the Archbold Expeditions of the American Museum of Natural History. Contemplating a taxonomic review of the Oriental squirrels, Dr. George Tate undertook to re-examine and attempt to identify with the population to which it belongs, each of the approximately 700 ac- cessible type specimens of squirrels from southeastern Asia and the East Indies. This study involved visiting many of the museums in which the types are deposited, and these visits included study of extensive additional collections of Oriental squirrels. Procedure con- sisted of vocally recording new observations on the types in Ameri- can and European museums, of photographing the skulls of types, and of making detailed measurements of them. Comparative obser- vations on other specimens of Oriental squirrels in European muse- ums in 1951 were also recorded. In some instances specimens were sent on loan for examination (by Tate). Dr. George H. H. Tate died in December of 1953. The other author (Moore) was invited to put the manuscript on the Oriental squirrels into final form for publication. Gradually he found that the manuscript treatment of many genera could not be put into form for publication by editing. He then undertook to change the concept of the paper from a review to a substantial revision based on Tate's transcribed observations of European material and new study of about 8000 Oriental squirrel specimens in museums in the United States. Beginning in September, 1957, grants were soHcited from federal agencies, but not for the preparation of a taxonomic revision alone. It was pointed out that this is a study of a subfamily of ter- restrial mammals unusually variable in color, in a geographic area of exceedingly varied land form and extraordinarily active orogeny and erosion. Federal support of the morphological study and revi- sion was sought and obtained on the grounds that they will reveal evidence on the evolution of the animals in relation to land forms and water barriers and thus may be important to evolutionary theory. Discoveries of special taxonomic importance in this revisionary work have led to several separately published contributions. One is 8 FIELDIANA: ZOOLOGY, VOLUME 48 a classification of the diurnal squirrels of the world (Moore, 1959). Another is an assessment of the taxonomic importance of the num- ber of pairs of functional mammae in the Sciurinae and also the relationship of this character in Sciurinae to climate and to size of broods (Moore, 1961a). The strengthening of the classification of the living Sciurinae provided in these two papers invited considera- tion of the evolutionary implications of the geographic distribution of the morphological characters observed. These implications have been treated in part in the 1959 paper that offered the classification, in part in a paper on the distribution of squirrels in the Indian Sub- region (Moore, 1960), and further in a discussion of spreadings across the Bering and Panamanian land bridges (Moore, 1961b). The use of the plural pronoun "we" should not confuse the reader who has read the above chronology of authorship. The "we" is usually meant only to imply participation of both authors in obtain- ing and collating the information and is not intended to place final responsibility for decisions on more than one pair of shoulders. Vir- tually no decision or opinion stated is Tate's alone unless so indicated in text except for ones that could only have been his because they could have been reached only by a study of the material in European museums. Acknowledgments One or both of the authors are indebted to the following persons in the United States who have welcomed us to their offices and pro- vided us access to the study collections in their charge: Dr. David H. Johnson, United States National Museum, Washington, D.C.; the late Colin C. Sanborn and Mr. Philip Hershkovitz, Chicago Natural History Museum, Chicago, Illinois; Miss Barbara Lawrence, Mu- seum of Comparative Zoology, Cambridge, Massachusetts; Dr. H. Radclyffe Roberts (and Dr. Karl F. Koopman, formerly of) Acad- emy of Natural Sciences, Philadelphia, Pennsylvania; and Dr. W. H. Burt and Dr. Emmett T. Hooper, Museum of Zoology, University of Michigan, Ann Arbor. In Europe the following persons extended similar courtesies and privileges to Tate or helped us with loans and information, for which we both express our appreciation here: Mr. T. C. S. Mor- rison-Scott and Mr. R. W. Hayman, both of the British Museum (Natural History), London; M. Jean Dorst, Museum National d'His- toire Naturelle, Paris; Count Nils Gyldenstolpe, Royal Natural History Museum, Stockholm; Dr. L. Forcart, Naturhistorisches MOORE AND TATE: DIURNAL SQUIRRELS 9 Museum, Basel; Dr. A. M. Husson, Rijksmuseum van Natuurlijke Historie, Leiden; Dr. E. Stresemann, Mr. K. Zimmermann, and Dr. W. Meise, Zoologisches Museum der Humboldt Universitat, Berlin; Dr. R. Mertens, Senckenbergisches Naturforschende Gesell- schaft, Frankfurt-am -Main; Dr. C. Stop-Bowitz, Universitetets Zoo- logiske Museum, Oslo; Dr. A. Holm, Zoologiska Institutionen, Uppsala; and the late Mr. H. J. V. Sody, of Amsterdam. For loans of important material from museums in the Oriental Region we are grateful to the late Dr. Sunder Lai Hora and to Dr. Biswamoy Biswas, both of the Zoological Survey of India; to Mr. Humayun Abdulali, of the Bombay Natural History Society; and to Mr. P. E. P. Deraniyagala, of the National Museums of Ceylon. This investigation was supported from September, 1957, through August, 1961, in part by Public Health Service research grant 5327 and in part by National Science Foundation grants 4447, 6250, and 14265. The surviving author is very appreciative of these. Gratitude is expressed to Dr. Harold E. Anthony of the American Museum of Natural History for the original invitation (to Moore) to take up where Tate had left off, and for support and encourage- ment until his retirement in May, 1958. His successor, Dr. R. G. Van Gelder, continued to make research facilities available for this project through the summer of 1961, provided special support from departmental funds during one four-month period, and through crit- icism of the manuscript has contributed toward accuracy, consist- ency, and clarity of expression in this paper. Acknowledgment for a special kind of helpfulness is made to Mr. Herbert G. Deignan, Curator of Birds, United States National Museum, for a great deal of advice and help in the finding and spell- ing of place names in Thailand. Mr. Melvin A. Traylor, Chicago Natural History Museum, pro- vided me with data on the collection localities of Floyd T. Smith in Szechwan, China. Dr. Charles Vaurie, American Museum of Nat- ural History, welcomed us to study his personal set of 1:1 million topographic maps of the Orient. The skull drawings are from the pen of Mrs. Richard G. Zweifel. The inking and lettering of fourteen of the maps are some of the last work of Mrs. D. F. Levett Bradley, who passed away before com- pleting the series. The maps showing the distribution of Sciurotamias and of Ratufa macroura, R. indica, and of the several species of Callo- sciurus were inked and lettered by Mrs. Zweifel. 10 fieldiana: zoology, volume 48 Scope The number of tribes of diurnal squirrels represented in the Ori- ental Region (5) exceeds that of any other faunal region of the world. The number of genera of diurnal squirrels native to the Oriental Region (17) greatly exceeds that of other faunal regions (see Table 1). Several of the genera listed in Table 1 as Oriental occupy areas mar- ginal to the Oriental Region proper. Prosciurillus, Rubrisciurus, and Hyosciurus are endemic to the island of Celebes, which lies in an area transitional between the Oriental Region and the Australian Region. Sciurotamias occupies an area of China which is almost entirely tran- sitional between the Oriental and Palearctic regions. Nevertheless, none of the 17 genera here listed as Oriental extends more than mar- ginally into another of the faunal regions of the world. These data suggest that the Oriental fauna of diurnal squirrels may be the most important in the world to study. They are also a convenient unit to revise. The Oriental Region is sometimes divided into three subregions: the Indian, the Indochinese, and the Malaysian. In the opinions of some authorities the Philippine Islands constitute a fourth Oriental subregion, but others consider it transitional between the Orien- tal and Australian regions. The diurnal squirrel fauna of the Oriental Region is well divided by the subregional boundaries. One of us has discussed in detail how the boundary between the Indian and Indochinese subregions separates the giant squirrels as species and all other diurnal squirrels as tribes (Moore, 1960). The bound- ary between the Indochinese and Malaysian subregions provides a weaker but still very strong division. This boundary is generally (and here) drawn at the constriction of the Malay Peninsula called the Isthmus of Kra. (See the discussion by Chasen, 1940, p. x, and the map, fig. 13 in the present paper.) Figures 7 and 8 combined may be said to indicate crudely the extent of the Indian Subregion by distribution of the squirrels. Figure 13 does the same for the Indochinese Subregion. The scope of the present paper is revision of the Sciurinae of the Indian and Indochinese subregions. A large part of it constitutes the first revision of the species of Callosciurus of the Indochinese Subregion. The gliding squirrels, Petauristinae, which with the Sciurinae con- stitute the family Sciuridae, are nocturnal and thus more difficult for mammal collectors to obtain. They are rarely collected by orni- thologists. Consequently, in American collections of Oriental squir- rels, specimens of Petauristinae number about one to every 15 of MOORE AND TATE: DIURNAL SQUIRRELS 11 Sciurinae. The Petauristinae, although included in the original plans and draft by Tate, have not yet been studied by Moore, but see McKenna (1963). The geographic area included in the present study is thus pre- cisely that part of the Oriental Region included by EUerman and Morrison-Scott (1951). For this reason the report here on the giant squirrel species, Ratufa hicolor, is confined to the area north of the Isthmus of Kra, although the species range extends southeastward through Malaya, Sumatra, Java, and Bali. There are, however, five other polytypic species with ranges that are widespread in the Indo- chinese Subregion but that extend beyond its border down the Malay Peninsula for varying distances. These five species are treated in full because their ranges do not extend beyond the length of the peninsula, and because they generally have but one subspecies south of the Isthmus of Kra. Four-fifths of the species of the genus Dremomys are widespread, polytypic species of the Indochinese Subregion. There is, however, a poorly known, apparently monotypic species, everetti, confined to Borneo. In the interest of having the revision of the genus Dre- momys in one publication, a treatment of everetti is included here. The forms from south of the Isthmus of Kra that are treated in the present work are labeled "extraterritorial." Material In the lists of material examined for each form, the locality near which a specimen was taken is ordinarily given as written by the collector on the specimen label. This rule has been departed from occasionally when evidence clearly shows that the collector was spell- ing the name of one locality in different ways. Since for some Ori- ental countries one may find the name of a single locality spelled (transliterated to English or French) several different ways on as many maps (e.g., Pak Jong, Pak Chong, Pak Xong), the locality as spelled by the collector may be followed in brackets by the name one of us found on a map and presumed to be the locality meant. Often the collector has recorded on the specimen tag two or more locality names, indicating a village, the town to which it is nearest, and perhaps the provincial capital to which they are both near. If it has not been possible to locate the village on a map, the town or occasionally even the provincial capital has sometimes been plotted as the locality for this specimen on the appropriate species distribu- a o •5 •& S eo g "=» S S ^ fc « js s ^ •" to O >» w ^ S t« •2 S a .. 3 § *5 w .2 S3 t> o ?3 C w hH a, r* eij - •n « •« -2 O 3 a> O) 73 Eh -S igii-s|-s i~ p, s S5 c5 s e 31 2 o ec -e § ^ ^ III »>;s"s •p.- 0-2 § So fi i£sS 5 s e fe 1^ a a I" o x> 6 c 3 13 o •2 c o i- o) 12 MOORE AND TATE: DIURNAL SQUIRRELS 13 tion map. The instances in which this has been done are clearly indicated as follows: the localities not found (as well as any others, such as "India," not suitable for plotting) are placed in quotation marks in the list of material examined, and the nearby places that are plotted instead are italicized (e.g., the village, subdistrict town, district town, and provincial capital "Ban Maeo, Goksatawn," Dan Sai, Loei, Thailand). In our lists of material examined, the institutions in which the study material is housed are cited by initials as follows: AMNH — American Museum of Natural History, New York ANSP — Academy of Natural Sciences, Philadelphia ASB — Asiatic Society of Bengal, Calcutta BM — British Museum (Natural History), London CBK — Private collection of C. Boden Kloss (not seen) CNHM — Chicago Natiu-al History Museum, Chicago IM — Indian Museum, Calcutta MCZ — Museum of Comparative Zoology, Cambridge, Massachusetts MNCN — Museo Nacional de Ciencias Naturales, Madrid MNHN — Museum National d'Histoire Naturelle, Paris MNHUI — Museum of Natural History, University of Illinois, Urbana NM — Naturhistorisches Museum, Basel NMC — National Museums of Ceylon, Colombo NR — Naturhistoriska Riksmuseum, Stockholm RNH — Rijksmuseum van Natuurlijke Histoire, Leiden SMT — Staatliches Museum fiir Tierkunde, Dresden SNG — Senckenbergisches Naturforschende Gesellschaft, Frankfurt-am-Main UMMZ University of Michigan Museum of Zoology, Ann Arbor UZM — Universitetets Zoologiske Museum, Oslo ZI — Zoologiska Insitutionen, Uppsala ZMHU — Zoological Museum of Humboldt University, Berlin Methods In many species the present work has been based on remarkably fine and extensive collections. Some mainland subspecies are known from at least one large series of specimens from a single locality, and smaller numbers from a dozen other widely distributed localities, providing, thus, some indication of individual variation with which to evaluate evidence of geographic variation. Other subspecies are represented by much less material, and future collections may en- able other students to define better the characteristics and ranges of many subspecies. In order to facilitate advancement of knowl- edge of these species, we have attempted to present our knowledge in detail specific enough to enable future students to add their evi- dence to ours, perhaps without the necessity of restudy of the perti- nent material on which we report. 14 FIELDIANA: ZOOLOGY, VOLUME 48 Color. — One of us (Moore) has employed Ridgway's (1912) no- menclature, and where capitalized color names are used in this paper, direct comparison of the pelage (s) with Ridgway's colors is implied. To determine color of a series of study skins, we arranged them on a horizontal plane before a window, preferably at a time between 10 A.M. and 4 P.M. and when the sky was blue, but never when the sun could shine directly on the specimens. The skins were oriented with tails toward the window, and the color comparison was made from a view perpendicular to the surface upon which the specimens lay, and thus from a point directly above the part being compared. Color comparisons were virtually all made at the institution in which the study specimens were stored, but at the United States National Museum, because of cloudy weather and distance to win- dows, some color readings were taken under direct light of an ordi- nary 100-watt, electric light bulb with a tungsten filament. In order to avoid the risk of losing or damaging a copy of Ridg- way by traveling with it, we borrowed and used different copies at the different institutions. Comparisons of material at one museum with material at another by Ridgway color names risks variation between copies of the book, and is thus relatively seldom employed here. Persons seeking to make identifications of considerable im- portance are advised in no case to depend solely upon distinguishing a small difference in colors in Ridgway (1912), but to send skins and skulls to one of the museums listed as having materials of the pre- sumed form for confirmation of the identification. It should perhaps be pointed out that diagnosis of a subspecies of Oriental diurnal squirrel rarely depends upon color of a single part of the pelage which is only one Ridgway color chip different from an adjacent subspecies. The term "agouti" as used here means body pelage with one or more minute contrasting color bands on the individual hairs that, in a broad view of the pelage, produce a speckled effect. Except where otherwise stated, color of agouti pelage is recorded here as the general effect of the visible portions of the pelage, its contrasted ele- ments blended by viewing without magnification with the viewer's eyes deliberately out of focus. The banding of agouti body pelage generally consists of one, two or three light-colored bands (parts of the length of a hair sharply differentiated by color) on any particular hair, but some hairs in agouti pelage have no bands at all. On the individual hairs of the tail, it is the black or blackish bands that are counted unless specifically otherwise stated, and the count of black- ish bands on individual hairs does not include the almost ubiquitously MOORE AND TATE: DIURNAL SQUIRRELS 16 black tip because this tip is often markedly thinner than the remain- der of the hair, and often shorter than the blackish bands. Form of presentation. — Decision to change the study of the Ori- ental squirrels from a review to a revision derived in part from a growing curiosity about the origins of the species and from discovery of indications that better study would reveal firmer evidence on the manner of their evolution. In the Indochinese Subregion, for ex- ample, the exceedingly rich and rather well-collected diurnal squirrel fauna has been evolving on an orogenically and tectonically active mainland that has been sliced into parallel north-south strips of land by large rivers that flow down from the Tibetan Plateau. Thus the intention has not been primarily the laudable desire to bring badly needed order to a chaotic fauna and to complete a con- venient revision by means of which virtually all new specimens might be identified to subspecies. The emphasis has rather been to revise the species for evidence about the evolution of the species from coin- cidence of morphological and geographic relationships. Where the relationships have been obvious enough from the material examined, there has been some tendency simply to plot the subspecies distri- bution on the map, record the material examined, and move on: Where differences have been more difficult to see, as in the subspecies of pygerythrus or the relationships of phayrei, elaborate written analy- ses became necessary and have, of course, been incorporated into the revision. New evidence on relationships between species has been especially sought, and where found is given in detail. To synonymize subspecies without explanation is certainly not a policy which we would wish to advocate. Generally, where there has seemed to be basis enough for a finer splitter to recognize a sub- species that we have lumped with another, we have pointed that out. But the 6}/2 years spent has been too short, the digressions to exploit important discoveries too long, and the pressure from supporters to complete a revisionary manuscript too great, to allow the explana- tion of why each named form that we have synonymized is by our observations undeserving of recognition. Size and dimensions. — Students who have studied species that are more conservative in pelage color and more variable in skull dimensions than diurnal tree squirrels may be surprised at how sel- dom we have needed to utilize measurements. Tree squirrels are, however, well known to be especially conservative in variation in dimensions of the skull. (See Moore, 1959, p. 192 and Table 3.) On hearing that the discovery of a skull character that seemed to 16 FIELDIANA: ZOOLOGY, VOLUME 48 sort tree squirrels into tribes was tempting us to digress, one senior mammalogist assured us almost bitterly, "If you discover one good skull character in tree squirrels, it will be worth every dollar the foundation puts into your project." A number of good skull char- acters at the generic level and above were discovered and reported (Moore, 1959), but at the species and subspecies level in tree squirrels it has been obvious that the return for effort expended would be im- practical for a revision so extensive. In the semi-terrestrial long- nosed squirrels, however, species differences in skulls could be de- tected and are utilized. Scientists who have presented, and to an extent utilized, body and skull dimensions in their revisions of much smaller taxonomic units may find it difficult to endure our failure to emulate their ad- mirable intensive methods in our work. Tate obtained and recorded body and skull measurements and other data for each type specimen. Those of the body he apparently copied from specimen tags, adding plus-or-minus to measurements whose accuracy he doubted. He measured the skulls himself. Some of these data for each genus are presented in Tables 2-4, 6, 8-13, and 15. In these tables total length of skull is the greatest occipitonasal length. Mastoid breadth is the greatest span of the mastoid processes. Nasal length is the greatest length of the longer nasal bone. Diastema is the least distance from the alveolus of the incisor to the alveolus of the fourth upper pre- molar. Length of palate is the distance from the gnathion to the most anterior point on the posterior margin of the palate. Length of bulla is the greatest external length of the auditory bulla. Maxil- lary toothrow is the crown length of the upper row of cheekteeth from the most anterior extension of the crown of the fourth pre- molar to the most posterior point on the crown of the third molar. Orbitonasal length (used in fig. 28) is taken from the most anterior point in the margin of the right orbit to the most anterior point on the right nasal. Length of orbit (used in fig. 29) is taken from the most anterior point on the margin of the orbit (a small notch in the posterior margin of the lacrimal) to the most posterior point on the margin of the orbi to- temporal fossa. Order. — The order of genera progresses from those with no trans- verse septum in the auditory bulla to three such septa, as in the latest classification of the Sciurinae (Moore, 1959). This progression is considered to be from unspecialized to rather highly specialized in this particular character, but it does not imply that Ratufa is unspe- cialized in general adaptation to a niche (which would be untrue) or MOORE AND TATE: DIURNAL SQUIRRELS 17 that Sciurotamias is highly speciaHzed in general adaptation to a niche (though this may be true) . The order of species within a genus, and subspecies within a species, is geographical, from a convenient edge of the range toward the far edges. Under accounts of the genera we give the accepted name, syno- nyms if any, type species, definition, and diagnosis. Where it has seemed especially valuable, we have included data on systematic history and intra-generic relationships, and in several instances have provided keys to identification of the species. In the account of each polytypic species we have given the ac- cepted name, definition, and diagnosis. If it belongs in a polytypic genus we have generally commented on its relationships to other congeneric species with which it is in geographic contact, or which it particularly resembles. Sometimes the species account has seemed the proper place for general comments on the intraspecific variation. In a few instances a systematic history of the species is provided, and in the case of the extraterritorial species Dremomys everetti ob- servation of the rarely seen deciduous upper third premolars stimu- lated some further investigation and reporting of aspects not treated here in the other species. In each account of subspecies and monotypic species, we have given the accepted name, synonyms if any, information on the type specimen (s), and a list of the material examined. If some informa- tion regarding the type is lacking from our account, this may be taken to imply that we did not find that information with the origi- nal materials examined or in the original published account. If our data and observations and opinions on a type (generally Tate's, of course) differ from any given in the original published description of the form, this may be assumed to mean that our information, observation, or opinion (recorded by Tate) differs from that of the original describer. The list of material examined is generally fol- lowed by one or more descriptions. These descriptions may be of three kinds, and the kind is indicated by the heading given to each. "Original Description" designates a quotation from the original pub- lished description, usually of the type specimen, of course. "Type Description" designates observations by one of us of the type speci- men or the cotypes. Unless dated after 1953, these are Tate's ob- servations, expanded from his telegraphic notes. "Pelage Color" designates descriptions from series of specimens considered to repre- sent populations constituting the subspecies under consideration. These are given in Ridgway's color terms, and the observations were 18 FIELDIANA: ZOOLOGY, VOLUME 48 made in the manner described above in the account of color. Dis- cussion of some especially interesting or significant features is some- times contributed under the heading "Variation," "Dimensions," or "Discussion," and field observations from the literature are occasion- ally quoted under the heading "Habits." When only one small paragraph of comment follows the list of material examined, no head- ing is applied to it. Taxonomic History Because the Sciurinae are active by day and often brilHantly colored, they have long claimed the interest of naturalists. In the Oriental Region, Horsfield (1824) was one of the earliest to attempt to summarize knowledge of the squirrels. In his "General enumera- tion of Indian Sciuri" he listed twelve species of non-flying squirrels under the generic name Sciurus, including representatives of the modern genera Callosciurus, Lariscus, Funambulus, and Ratufa. Gray (1867) briefly keyed out and roughly classified the Oriental squirrels by simple pelage and tooth characters but he missed what now seem to be the most obvious of relationships and lumped to- gether in one genus, Macroxus, some species now considered tribally different. John Anderson (1878, pp. 214-277) in order to identify his collections of squirrels from Burma reviewed all of the known Oriental squirrels. His conservatism in admitting Reithrosciurus and Rhinosciurus only to subgeneric rank seems a matter of the custom of his time, but the superficiality of his observations on the diagnostic characteristics of their skulls contrasts sharply with his penetrating observations on pelage color of squirrels in the Indo- chinese Subregion and his shrewd speculations as to the relationships that these indicate. Trouessart (1880) correctly recognized, as Gray (1867) had not, that the forms hicolor, giganteus, indicus, maximus, and macrurus belonged together in a taxon at least subgenerically distinct from typical Sciurus. He recognized Funambulus as of subgeneric value, but placed in it virtually every named Indochinese squirrel with stripes, including berdmorei, mcclellandi, and even quinquestriatus. In an Asiatic and Malaysian subgenus he also proposed to include erythraeus and lokroides with lokriah, pernyi, and rufigenus, among others, and included davidianus in the subgenus Rhinosciurus with laticaudatus. Jentink (1883) recognized as belonging in the genus Sciurus the species erythraeus, lokroides, caniceps, atrodorsalis, quinquestriatus, MOORE AND TATE: DIURNAL SQUIRRELS 19 castaneoventris, and pygerythrus from this region, but he included in the same genus the species lokriah and pernyi (now in Dremomys) and davidianus and berdmorei (now respectively in Sciurotamias and Menetes) and three species now properly belonging to Funamhulus. For the two subregions under discussion he admitted but one genus of diurnal squirrels and no subgenera. By grouping, however, he recognized affinities among species now known to belong to the genera Ratufa, Funamhulus, and Dremomys. Blanford's (1888-91) taxonomic treatment of the squirrels of these two subregions differs little from Jentink's, but Blanford pro- vided keys and more detailed characterizations. Major (1893, p. 189), using primarily characters of the teeth and to a lesser extent those of the skull, produced a classification of the living Sciuridae with six genera in the Sciurinae. One of the largest is Xerus, composed of five subgenera, four of which are entirely African tree and ground squirrels. The fifth subgenus was Eoxerus, and in it were the two commonest species of the Indian Subregion, palmarum and pennanti, now included in Funamhulus, herdmorei, now included in Menetes, and three Malaysian ground squirrel spe- cies of the present genus Lariscus. Under the genus Sciurus, sub- genus Sciurus, and beta group he included tree squirrels of the Indo- chinese Subregion — erythraeu^, atrodorsalis, caniceps, ferrugineus, and lokroides — and several tree squirrels of the Malaysian Subregion as well as one long-nosed species, everetti. Major ends this list with "etc.," which probably implies that long-nosed species rufigenu^, pernyi, and lokriah belong here, too. He holds to genus Sciurus, subgenus Eosciurus, for the several Oriental giant squirrel species as proposed by Trouessart. Thomas (1915) separated Indochinese and Malaysian tree squir- rel species from the genus Sciurus, noting that they are distinguished by the presence of a separate bony blade associated with the shaft of the OS penis. These tree squirrels of the Indochinese Subregion (and Malaysian Subregion) he recognized as Callosciurus and Tomeutes. Robinson and Kloss (1918) contributed a valuable list of the Sciuridae of the Oriental Region. They admitted nine species of Ratufa. In the Indochinese Subregion they admitted 15 species in what is now accepted as Callosciurus. Pocock (1923) followed Thomas' lead with a classification of the Sciuridae based upon further extensive study of the baculum (os penis) and glans penis. He associated the Indian striped squirrels, Funamhulus, and Oriental giant squirrels, Ratufa, with several genera 20 FIELDIANA: ZOOLOGY, VOLUME 48 of tree squirrels of the Ethiopian Region in one subfamily, but placed other Oriental squirrels (excluding Reithrosciurus) in a strictly Ori- ental subfamily, Callosciurinae. Simpson (1945) accepted Pocock's classification but scaled his subfamilies down to tribes. Moore (1959) found new skull characters which generally support Pocock's classi- fication as modified by Simpson, and by which one may allocate several Oriental genera, the bacula of which are still unknown, to the tribe Callosciurini. Ellerman (1940) offered an arrangement of the species and sub- species of Oriental squirrels, and Ellerman and Morrison-Scott (1951) offered an arrangement of species and subspecies of Sciurinae of the Indian and Indochinese subregions. These are checklists and not revisions at the species and subspecies level, and although there are many taxonomic decisions implicit in their arrangement, based on study of specimens, the evidence for these is not given. Zahn (1942) offered a revision of the genera, species, and sub- species of the giant squirrels, striped squirrels, and long-nosed squir- rels of the Oriental Region, based on study of material in European museums. Other authors have published many helpful papers revising small groups of diurnal squirrels of this region. Some of these will be men- tioned in the following species accounts. Some authors have pro- vided important revisions of the Sciurinae included in large faunal papers. Of the latter the papers of G. M. Allen (1940) and Osgood (1934) are particularly helpful. Classification The classification presented here constitutes conclusions derived from the whole present work and might with logic be placed at the end of the work. However, the choice had to be made between logic of presentation and convenience of use. We hope that some students will, indeed, weigh our logic, but we presume, however wrongly, that most students turning these pages, will be primarily interested in using our results. The attained classification is therefore provided in the introduction where it can serve conveniently as a framework for apprehending the work as a whole or the relationships of some particular genus or species. One of us (Moore, 1959) has reported in detail the tribal level skull characters that distinguish the following genera from all others in the Oriental Region: Ratufa (op. cit., p. 169), Funambulus (p. 170), MOORE AND TATE: DIURNAL SQUIRRELS 21 and Sciurotamias (p. 182). The other existing genera of the Sciuri- nae of the Indochinese Subregion may be identified in the key to the genera and subgenera of the subtribe Callosciurina (op. cit., pp. 173-174). Tamiops is treated as a subgenus in that key. Classification of the Sciurinae of the Indian and Indochinese Subregions Tribe Ratufini Moore, 1959. Oriental Region. Genus Ratufa Gray, 1867. Oriental Region. Species macroura Pennant, 1769. Ceylon and tip of Indian Peninsula. Subspecies macroura Pennant, 1769 (including: ceylonicus Erxleben, 1777. ceilonensis Boddaert, 1785. tennenti Blyth, 1849. montanus Kelaart, 1852. macrurus Blanford, 1891.) melanochra Thomas and Wroughton, 1915. dandolena Thomas and Wroughton, 1915 (including sinhala Phillips, 1931.) Species indica Erxleben, 1777. Indian Subregion. Subspecies dealbata Blanford, 1897. indica Erxleben 1777 (including: purpureus Zimmerman, 1777. bombayus Boddaert, 1785. elphinstonei Sykes, 1831. superans Ryley, 1913.) maxima Schreber, 1784 (including: mxilabaricus Scopoli, 1786. bengalensis Blanford, 1897.) centralis Ryley, 1913. Species bicolor Sparrmann, 1778. Indochinese and Malaysian Subregions. Subspecies phaeopepla Miller, 1913 (including: celaenopepla Miller, 1913. marana Thomas and Wroughton, 1924.) leucogenys Kloss, 1916. sinus Kloss, 1916 (not mapped). smithi Robinson and Kloss, 1922. condorensis Kloss, 1920 (not mapped). felli Thomas and Wroughton, 1916. gigantea McClelland, 1839 (including: macruroides Hodgson, 1849. lutrina Thomas and Wroughton, 1916.) hainana J. A. Allen, 1906 (including stigmosa Thomas, 1923.) Tribe Funambulini Simpson, 1945. Ethiopian and Oriental Regions. Subtribe Funambulina Moore, 1959. Indian Subregion. Genus Funambulus Lesson, 1832. Indian Subregion. Subgenus Prasadsciurus, new subgenus Species pennanti Wroughton, 1905. Northern India, W. Pakistan, Nepal. Subspecies pennanti Wroughton, 1905. lutescens Wroughton, 1916. argentescens Wroughton, 1905. 22 FIELDIANA: ZOOLOGY, VOLUME 48 Species palmarum Linnaeus, 1766. Central and southern India and Ceylon. Subspecies palmarum Linnaeus, 1766 (includes: penicillatus Leach, 1814. indicus Lesson, 1835. gossei Wroughton and Davidson, 1919.) comorinus Wroughton, 1905. bellaricus Wroughton, 1916. robertsoni Wroughton, 1916. bengalensis Wroughton, 1916. favonicus Lindsay, 1926. kelaarti Blyth, 1851. brodiei Blyth, 1849. olympius Thomas and Wroughton, 1915. Species trisiriatus Waterhouse, 1837. Western Ghats. Subspecies tristriatus Waterhouse, 1837 (including: dussumieri Milne-Edwards, 1867. annandalei Robinson, 1917.) wroughtoni Ryley, 1913. numxirius Wroughton, 1916 (including thomasi Wroughton and Davidson, 1919.) Species layardi Blyth, 1849. Ceylon and southern India. Subspecies layardi Blyth, 1849. signatus Thomas, 1924, dravidianus Robinson, 1917. Species sublineatus Waterhouse, 1838. Ceylon and southern India. Subspecies sublineatus Waterhouse, 1838 (including: delesserti Gervais, 1841. trilineatus Blyth, 1849.) obscurus Pelzeln and Kohl, 1886 (including kathleenae Thomas and Wroughton, 1915.) Tribe Callosciurini Simpson, 1945. Indochinese and Malaysian Subregions. Subtribe Callosciurina Moore, 1959. Indochinese and Malaysian Subregions. Genus Callosciurus Gray, 1867. Indochinese and Malaysian Subregions. Species erythraeus Pallas, 1778. Indochinese Subregion west of Irrawaddy River. Subspecies erythraeus Pallas 1778 (including wellsi Wroughton, 1921.) erythrogaster Blyth, 1842 (including: punctatissimus Gray, 1867. nagarum Thomas and Wroughton, 1916. kinneari Thomas and Wroughton, 1916. crotalius Thomas and Wroughton, 1916.) bhutanensis Bonhote, 1901 (including crumpi Wroughton, 1916.) intermedius Anderson, 1879 (including aquilo Wroughton, 1921.) sladeni Anderson, 1871 (including: kemmisi Wroughton, 1908. rubex Thomas, 1914. midas Thomas, 1914. vernayi Carter, 1942.) bartoni Thomas, 1914 (including: shortridgei Thomas and Wroughton, 1916. fryanus Thomas and Wroughton, 1916. miliar di Thomas and Wroughton, 1916. MOORE AND TATE: DIURNAL SQUIRRELS 23 caryi Thomas and Wroughton, 1916.) haringtoni Thomas, 1905 (including solutus Thomas, 1914.) Species flavimanus I. Geoff roy, 1831. Indochinese Subregion east of Irra- waddy. Subspecies quinquestriatus Anderson, 1871 (including: heehei J. A. Allen, 1911. imarius Thomas, 1926. Sylvester Thomas, 1926.) gordoni Anderson, 1871. shanicus Ryley, 1914 (including Igriseopedus Blyth, 1847.) hyperythrus Blyth, 1855. (not mapped, no localities) michianus Robinson and Wroughton, 1911 (including haemobaphes G. M. Allen, 1912.) zimmeensis Robinson and Wroughton, 1916 (including primus Allen and Coolidge, 1940.) thai Kloss, 1917. atrodorsalis Gray, 1842. siamensis Gray, 1860 (including tachin Kloss, 1916.) pranis Kloss, 1916. rubeculus Miller, 1903 (including youngi Robinson and Kloss, 1914. Unmapped. Extrater- ritorial.) hendeei Osgood, 1932. castaneoventris Gray, 1842 (including insularis J. A. Allen, 1906.) flavimanus Geoffroy St. Hillaire, 1831 (including: quantulus Thomas, 1927. contumax Thomas, 1927. dactylinus Thomas, 1927. pirata Thomas, 1929. bolovensis Osgood, 1932.) griseimxinus Milne-Edwards, 1867 (including: leucopus Gray, 1867. fumigaius Bonhote, 1907. vassali Bonhote, 1907. phanrangis Robinson and Kloss, 1922.) gloveri Thomas, 1921. bonhoiei Robinson and Wroughton, 1911. styani Thomas, 1894 (including: griseopedus Milne-Edwards, 1874. Herpestes [sic] leucuru^ Hilzheimer, 1905. Herpestes [sic] albifer Hilzheimer, 1906. canigenus Howell, 1927. woodi Harris, 1931.) ningpoensis Bonhote, 1901 (including tsingtanensis Hilzheimer, 1905.) thaiwanensis Bonhote, 1901 (including: centralis Bonhote, 1901. roberti Bonhote, 1901. nigridorsalis Kuroda, 1935.) Species ferrugineus F. Cuvier, 1829. Indochinese Subregion east of Irra- waddy (including keraudreni Lesson, 1830.) 24 FIELDIANA: ZOOLOGY, VOLUME 48 Species finlaysoni Horsfield, 1824. Indochinese Subregion east of Salween River, Subspecies finlaysoni Horsfield, 1824 (including partus Kloss, 1915.) folletti Kloss, 1915. trotteri Kloss, 1916. frandseni Kloss, 1916. albivexilli Kloss, 1916. harmandi Milne-Edwards, 1876 (including pierrei Robinson and Kloss, 1922.) germaini Milne-Edwards, 1867. nox Wroughton, 1908. cinnamomeus Temminck, 1853 (including: splendens Gray, 1861. herberti Robinson and Kloss, 1922.) annellatus Thomas, 1929. williamsoni Robinson and Kloss, 1922. menamicus Thomas, 1929 (1928). sinistralis Wroughton, 1908 (including: dextralis Wroughton, 1908. lylei Wroughton, 1908. grutei Gyldenstolpe, 1917.) boucourti Milne-Edwards, 1867 (including: leucogaster Milne-Edwards, 1867. leucocephalus Bonhote, 1901. floweri Bonhote, 1901. tachardi Robinson, 1916. prachin Kloss, 1920. rajasima Kloss, 1920. cockerelli Thomas, 1928.) boonsongi, new subspecies. Species caniceps Gray, 1842. Indochinese Subregion west of Mekong River. Subspecies concolor Blyth, 1855 (including: lancavensis Miller, 1903. erubescens Cabrera, 1917. telibius Thomas and Robinson, 1921 Extraterritorial.) adangensis Miller, 1903 (including: terutavensis Thomas and Wroughton, 1909. moheius Thomas and Robinson, 1921. mohillius Thomas and Robinson, 1921.) bimaculatus Temminck, 1853 (including: epomophorus Bonhote, 1901. davisoni Bonhote, 1901. sullivanus Miller, 1903. matthaeus Miller, 1903. lucas Miller, 1903. milleri Robinson and Wroughton, 1911. samuiensis Robinson and Kloss, 1914. nakanus Thomas and Robinson, 1921. mapravis Thomas and Robinson, 1921. panjius Thomas and Robinson, 1921. panjioli Thomas and Robinson, 1921. tacopius Thomas and Robinson, 1921. pipidonis Thomas and Robinson, 1921. iabaudius Thomas, 1922. hastilis Thomas, 1923.) MOORE AND TATE: DIURNAL SQUIRRELS 25 casensis Miller, 1903 (unmapped). fallax Robinson and Kloss, 1914 (unmapped). domelicus Miller, 1903 (including bentincanus Miller, 1903.) caniceps Gray, 1842 (including: chrysonotus Blyth, 1847. inexpectatus Kloss, 1916. helgei Gyldenstolpe, 1917. helvus Shamel, 1930.) Species phayrei Blyth, 1855. Indochinese Subregion west of Salween (in- cluding: blanfordi Blyth, 1862, and heinrichi Tate, 1954.) Species inornatus Gray, 1867. Indochinese Subregion east of Mekong (in- cluding imitator Thomas, 1925.) Species pygerythrus Geofifroy St. Hillaire, 1832. Indochinese Subregion west of Irrawaddy. Subspecies pygerythrus Geoffroy St. Hillaire, 1832. janetia Thomas, 1914. owensi Thomas and Wroughton, 1916. Twearsi Bonhote, 1906 (including: bellona Thomas and Wroughton, 1916. virgo Thomas and Wroughton, 1916.) stevensi Thomas, 1908. blythi Tytler, 1854. lokroides Hodgson, 1836 (including: assamensis Gray, 1843. similis Gray, 1867.) Genus Tamiops Allen, 1901. Indochinese Subregion and Malaya. Species mcclellandi Horsfield, 1839. Indochinese Subregion and extraterri- torial subspecies in Malaya. Subspecies mcclellandi Horsfield, 1839 (including: pembertoni Blyth, 1842. manipurensis Bonhote, 1900.) collinus Moore, 1958. inconstans Thomas, 1920. kongensis Bonhote, 1901. barbei Blyth, 1847. leucotis Temminck, 1853 (including novemlineatus Miller, 1903.) Species rodolphei Milne-Edwards, 1867. Thailand, Laos, Cambodia. Subspecies rodolphei Milne-Edwards, 1867 (including: liantis Kloss, 1919. lylei Thomas, 1920. dolphoides Kloss, 1922. holti Ellerman, 1940.) elbeli Moore, 1958. Species svnnhoei Milne-Edwards, 1874. Yunnan, Szechwan, Burma. Subspecies smnhoei Milne-Edwards, 1874 (including: clarkei Thomas, 1920. forresti Thomas, 1920. spencei Thomas, 1921. russeolus Jacobi, 1923. olivaceiis Osgood, 1932.) vestittis Miller, 1915. Species maritimus Bonhote, 1900. Southern China, Vietnam. 26 FIELDIANA: ZOOLOGY, VOLUME 48 Subspecies maritimus Bonhote, 1900 (including: formosanus Bonhote, 1900. sauteri J. A. Allen, 1911.) monticolus Bonhote, 1900. hainanus J. A. Allen, 1906 (including: riudoni J. A. Allen, 1906. laotum Robinson and Kloss, 1922.) moi Robinson and Kloss, 1922. Genus Dremomys Heude, 1898. Indochinese and Malaysian Subregions. Species lokriah Hodgson, 1836. Nepal, Assam, northern Burma. Subspecies lokriah Hodgson, 1836 (including: bhotia Wroughton, 1916. subflaviventris Thomas, 1922.) garonum Thomas, 1922. pagus Moore, 1956. macmillani Thomas and Wroughton, 1916. Species pernyi Milne-Edwards, 1867. Assam to Chekiang above Tropic of Cancer. Subspecies pernyi Milne-Edwards, 1867 (including: griselda Thomas, 1916. lichiensis Thomas, 1922. lentus A. B. Howell, 1927.) howelli Thomas, 1922 (including: mentosus Thomas, 1922. imus Thomas, 1922.) flavior G. M. Allen, 1912. senex G. M. Allen, 1912 (including modestus Thomas, 1916.) calidior Thomas, 1916 (including chintalis Thomas, 1916.) owstoni Thomas, 1908. Species rufigenus Blanford, 1878. Indochinese Subregion and Malaya. Subspecies rufigenis Blanford, 1878 (including: fuscus Bonhote, 1907. adamsoni Thomas, 1914. ornatus Thomas, 1914. laomache Thomas, 1921. belfieldi Bonhote, 1908. opimus Thomas and Wroughton, 1916. Species pyrrhomerus Thomas, 1895. Southern China, northern Vietnam. Subspecies pyrrhomerus Thomas, 1895. riudonensis J. A. Allen, 1906 (including melli Matschie, 1922.) gularis Osgood, 1932. Species everetti Thomas, 1890. Montane Borneo. (Extraterritorial) Genus Menetes Thomas, 1908. Indochinese Subregion. Species berdmorei Blyth, 1849. Indochinese Subregion. Subspecies berdmorei Blyth, 1849 (including amotus Miller, 1914.) peninsularis Kloss, 1919. moerescens Thomas, 1914. decoratus Thomas, 1914. consularis Thomas, 1914. mouhotei Gray, 1861 (including: pyrrhocephalus Milne-Edwards, 1867. koratensis Gyldenstolpe, 1917.) MOORE AND TATE: DIURNAL SQUIRRELS 27 Tribe Tamiasciurini i Simpson, 1945. Nearctic Region and Oriental Region. Genus Sciurotamias Miller, 1901. Montane China, Sikang to Chekiang. Subgenus Sciurotamias Miller, 1901. Montane China, Sikang to Chekiang. Species davidianus Milne-Edwards, 1867. Montane Sikang to Chekiang, China. Subspecies davidianus Milne-Edwards, 1867 (including latro Heude, 1898.) consobrinus Milne-Edwards, 1868-74 (including: collaris Heude, 1898. saltitans Heude, 1898. owstoni J. A. Allen, 1909. thayeri G. M. Allen, 1913.) Subgenus Rupestes Thomas, 1922. Montane Yunnan and Sikang, China. Species forresti Thomas, 1922. Montane Yunnan and Sikang, China. 1 While the present paper was in press. Black (1963) published his dissertation on the North American Tertiary Sciuridae (localities in the United States and Mexico; 12 genera, 5 with living species). From that study Black says he proposes classification of the 37 living genera of Sciurinae of the world. However, besides including names of some North American fossil genera. Black (1963, p. 123) proposed only two changes from the most recent previous classification (Moore, 1959, p. 198-200). One change suggested raising the chipmunk taxon from subtribal to tribal level because ... "I feel that they differ from the marmots and the spermophiles as greatly as they do from the tree squirrels and are fully entitled to tribal status." No evidence, no discussion of relative merits, only opinion based on study of Nearctic fossils representing two of the eight tribes earlier known. The present writer studied all of the living genera of Sciurinae comparatively and quantita- tively, amassed a great deal of evidence, tabulated and discussed this, and con- cluded that in comparison with the marmots and with all other suprageneric taxa of living sciurines, chipmunks merit subtribal rank (Moore, 1959, pp. 162- 166, 180-182). It is recommended that their proposed elevation to tribal rank be rejected. The other change offered by Black (1963) is dissolution of the generally accepted tribe Tamiasciurini. Black (1963, p, 125) reports that 3 of the 26 skulls of Tamiasciurus at the Museum of Comparative Zoology have only two transbullar septa per bulla, but he evidently did not read the method for counting these provided in the caption of Table 1 (Moore, 1959, p. 163). There were 3 speci- mens with only two pairs of septa in 138 specimens at the American Museum of Natural History (Moore, 1959, table 1). If we add these to the 6 that I now find in 361 specimens of Tamiasciurus in Chicago Natural History Museum, the total is 9 per 499, or less than two percent. For the record, totals of specimens in the latter two museums for 2, 2*^, 3 3^^, and 4 pairs of septa are respectively 9, 12, 473, 3, and 1 individuals of Tamiasciurus. Black's (1963, p. 125) proposal to re- duce the rank of the Tamiasciurini seems to be based on too small a sample, faultily observed, for one character (number of transbullar septa), and on an un- justified complaint that the other character (high squamosal) "is extremely hard to evaluate." Consequently, I have retained Tamiasciurini. I placed Sciurotamias in the Tamiasciurini (Moore, 1959, pp. 182-184) only tentatively, trusting it will stimulate publication of pertinent new observations, and I am keeping it there in the present work. GIANT SQUIRRELS Genus RATUFA Gray Ratufa Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 273. Rukaia Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 275. Eosciurus Trouessart, 1880, Le Naturaliste, 2, no. 37, p. 291. Type species. — Ratufa, Sciurus indicus Erxleben;/?wA;am, Sciurus macrourus Pennant; and Eosciurus, Sciurus hicolor Sparrmann. Definition. — The genus Ratufa contains only the species ma- croura, indica, hicolor, and affinis and is endemic in the Oriental Region. Diagnosis. — Consult Figure 1 for some of the following details. (1) The squamoso-alisphenoid suture lies about halfway between the third upper molar and the posterior margin of the base of the zygo- matic process of the squamosal. (2) The chambers of the auditory bullae are not crossed by transbullar septa. (3) The palatines are very short, so that the pterygoid fossa extends as far forward as the third molar and sometimes almost as far as the second molar. (4) The postorbital constriction is less than 0.80 of the interorbital breadth. (5) In fully adult individuals both the sagittal suture and the fronto- parietal suture ankylose completely, so that there remain no visible sutures on the dorsal surface of the skull from the anterior edge of the frontals to the posterior edge of the parietals. (6) In lateral aspect the lateral lip of the infraorbital foramen is vertical to the occlusal plane, or inchned with the top forward, and the top of it reaches the maxillo-premaxillary suture. The above six characters distinguish Ratufa (which constitutes the tribe Ratufini), from other Sciurinae of the Indian and Indo- chinese subregions as follows: Sciurotamias by 1, 2, 3, 4, 5, and 6; sub tribe Callosciurina by 1, 2, 4, and 5; and Funambulus by 1, 2, 3, 4, and 5. Systematic History. — Gray's (1867, p. 273) original characteriza- tion of Ratufa is not meaningful today. It consisted of "Tail elon- gate, longer than the body and head; large-sized." The name Ratufa was evidently meant as a subgeneric designation, and included only 29 Fig. 1. Skull and left mandible of the pied giant squirrel, Ratufa bicolor, AMNH No. 83440, X 1. Note generic characters described in text. For usage of skull anatomy consult Moore (1959, figs. 1-5). 30 MOORE AND TATE: DIURNAL SQUIRRELS 81 the species Sciurus indicus. Farther on in the same paper he placed giant squirrels macroura of Ceylon, bicolor of Java, and ephippium of Borneo in a subgenus Rukaia under a genus Macroxus, but ad- mitted no close relationship between indicus and the other three. When Robinson and Kloss (1918) published a list of the squirrels of the Oriental Region, they employed the nine polytypic species of Ratufa which were then currently accepted : macroura, indica, bicolor, notabilis, ephippium, affinis, gigantea, phaeopepla, and melanopepla. Hayman and Holt (in Ellerman, 1940) followed that arrangement 22 years later. At the same time, however, Chasen (1940) reduced notabilis, ephippium, and melanopepla to subspecific status, and Zahn (1942) reduced phaeopepla, gigantea, and indica to subspecies. Eller- man and Morrison-Scott (1951) accept as species macroura on Cey- lon, indica in the Indian Subregion and bicolor in the Indochinese Subregion. None of these authors, except Zahn, gave reasons for their decision or characters distinguishing the species, and as will be shown, the arrangement we present with reasons for decisions and characters for species, differs from that of Zahn (1942). KEY TO THE SPECIES OF RATUFA 1. Crown sharply separated from nape by a distinct color band between ears. . .2 Crown pelage color connected with that of nape 3 2. Digits black and ear- tufts inconspicuous macroura Digits maroon or buff and ear-tufts conspicuous indica 3. Light mark on thigh; no black mark on chin affinis [extraterritorial] No light mark on thigh; small black mark on chin bicolor Intrageneric relationships. — It is suggested by the key to the spe- cies above and by the species diagnoses which will follow, that ma- croura is more closely related to indica than to bicolor or ajffinis. This is what would be expected, of course, on geographical grounds, and we think that it could also be demonstrated in characters of the skull. It seems desirable to point it out, however, because of a gen- eral similarity of appearance between some forms of macroura on Ceylon and forms of bicolor on Java and Bali (see Moore, 1960, p. 13). We could find no discrete descriptive characters of the skull which distinguish geographically adjacent pairs of these species. Some trends are apparent, but variation in them is considerable. For in- stance, the zygomatic process of the squamosal possesses an elevated ridge which rises to somewhat of a peak, and in lateral view makes (in adults) a distinctive second superior process on the zygomatic arch. This characterizes the USNM macroura material, somewhat variably characterizes the AMNH indica material, occurs in nearby XI 3 T3 C I— I CIS ^3 CO O 5s o S i> bo^ «J "^ ^ ^ ^ ^ ^ ,-H ^ ^ ^ ^ ^ rt ^ ^ -^M o»'-ioou3(Mt-eciosio«oooo;o«o© ~S '-I'-^ooweoNi-i-^-^rHTt'inrrc^ieco eas' eoeccoeoeoeococccoeoeoecweceoeo Ah 1-1 ^2 t-kooot-o-^ooosi-iccot-t-oeo -g 1— ItHt-Ii-HtHt-HtHtHt-HtHt-It-It-It-It-Hi— I J3 CO •f-lS Cit>OOU3in>(Mt>u:>i-lif5ooOi-HTj<00 Sf w «J2; C "3 lOlOosiOOCOt-OOlOt-OoiCiiOO ti? «0rJOO«DN^(MOe0t-O(Mcci(MO 1 § §:2 -S <» S> i S & s s e «> te g-^ "^S '-S -S •S i eo eo 32 MOORE AND TATE: DIURNAL SQUIRRELS 83 Mt. Victoria specimens of R. bicolor gigantea but not in Mt. Popa ones a little farther away. In the rest of bicolor and in afinis this process does not appear to occur with frequency. This skull character might be taken as unexpected support for Zahn's (1942) recognizing macroura, indica, and gigantea as conspe- cific, but not hainana and stigmosa which he included (all grouped because of ear tufts). There is, moreover, in the ANSP material a Philadelphia Zoological Park specimen without data of origin, which is surely a cross between Ratufa indica and R. bicolor, for it displays some of the characters of each. However, since Hill (1939 and 1942) has easily bred both macroura and indica in captivity, it seems likely that this specimen results from a cross made in captivity and cannot be represented as a natural intergrade between indica and bicolor, suggesting that they should be treated as subspecies. In the diag- noses of the species bicolor and indica in the present paper we have offered a total of seven pelage characters which clearly distinguish these two as species. In the present study it has also become clear that the extraterri- torial Ratufa affinis is taxonomically the most distinct of the four species, although one can certainly say that indica is more spectacu- lar. The equatorial giant squirrel affinis is the only one of the four species with three pelage characters distinguishing it from all three other species. Rather little correspondence with the relationships of the species of Ratufa described above is seen in the exceedingly scanty knowledge of bacula of this genus. Dammerman (1931, p. 454) figured and de- scribed the baculum oi R. b. bicolor of Java as "a simple short bone of 9.4-9.9 mm. length, slightly upcurved; the basal end expanded and hollowed; the apex narrowed and flattened horizontally." From the other end of the range of this species, upper Burma, Pocock (1923, fig. 20, E and F) figured a baculum of R. b. gigantea which is very similar to the one from Java, but has the distal, flattened, ventral surface provided with four rows of small bony spicules. Pocock (1923, fig. 20, G and H) illustrated a baculum of R. indica showing it to be strongly curved upward distally, spread at the curvature, tridentate at the tip, and grooved or hollowed on its anteroventral surface. Prasad's (1954, fig. 4, D, E, and F) excellent illustrations of a baculum of R. indica maxima from Coorg or Mysore agree with Pocock's indica (a zoo specimen with no locality data) in the char- acters of base, amount of curvature, and lateral spread at curvature. Prasad's indica, however, does not show a hollow in its anterior sur- 34 FIELDIANA: ZOOLOGY, VOLUME 48 face, ends in a single nob that is slightly bilobed and has rows of bony spicules. Hill (1936, fig. 6, E and F) illustrated the baculum of R. macroura [dandolena] of Ceylon, showing that it has the 90-de- gree bend, widening at the bend, and a tridenticulate tip. In sum it appears that although variable, the bacula of hicolor may be distinguished easily from the equally variable ones of in- dica. However, until more published examples reveal the extent of variation, the existence of a real distinction between the bacula of indica and macroura is much less certain. It must be added that Hill (1940) reports rather striking differ- ences between the glans penes of R. macroura dandolena and R. m. melanochra, and that his illustration of the glans of melanochra is disconcertingly more like that of Prasad's (1954, fig. 4, A, B, and C) R. indica maxima than that of R. macroura dandolena. Much needed are further scientific reports on the glans penes and bacula of these and other subspecies (with the capture locality of each specimen and the identifying characteristics of each captive precisely stated) to demonstrate whether genital characters support or deny the species difference between macroura and indica which we now consider well established by our present study of characters of the pelage. The body and skull dimensions presented in Table 2 are measure- ments made and recorded by one of us (Tate) entirely of type speci- mens. These are offered here not as anything faintly resembling adequate representation of the subspecies or species, but as a rough indication of the dimensions likely to be found in this genus within the Indian and Indochinese subregions. Ratufa macroura (Pennant) Definition. — Ratufa macroura is the Ceylonese giant squirrel, which shares the southern India range of R. indica maxima but occurs with- out a competing giant squirrel species in Ceylon. Diagnosis. — Ratufa macroura has the following pelage characters: (1) The crown is black or blackish. (2) The digits are black or black- ish. (3) The dorsal pelage of feet, ankles, and forearms is yellowish buff. (4) Ear tufts are present but small and short, extending no more than 5 mm. beyond the skin of the ear tips. (5) A light-colored band crosses between the ears, separating crown from nape. The above characters distinguish macroura from the other spe- cies of Ratufa as follows: indica by 2 (ankle part of), 3, and 4; hicolor by 3 and 5; affinis by 1, 2, 4, and 5. MOORE AND TATE: DIURNAL SQUIRRELS 35 Fig. 2. Entire species range of the Sinhalese giant squirrel Ratufa macroura as revealed by material examined. Subspecies: A, macroura; B, melanochra; C, dandolena. Relationship to other species. — Zahn (1942) treated macroura and indica as one species. We know of no evidence of intergradation between macroura and indica, and Abdulali and Daniel (1952) in their study of geographic variation in the species indica evidently observed none. The occurrence of R. macroura in a considerable geographic area within the range of R. indica in southern India with- out evidence of intergradation thus far, requires that the evidence regarding status be considered in detail. There is in the locality records presented here for R. m. dandolena (fig. 2) and R. i. maxima (fig. 3) some fragmentary evidence that in southern India where populations of these forms are geographically sympatric, they may be ecologically segregated. The available ele- vations of Indian localities where R. macroura dandolena were col- 36 FIELDIANA: ZOOLOGY, VOLUME 48 lected are relatively low (1000, 2000, and 3000 feet) ; whereas those for R. indica maxima tend to be higher (3200, 3285, 3300, 3500, 3500, 3500, 4000, 4200, 4500, 5000, 6100, and 6500 feet). Both species were taken in the Palni Hills, two examples of macroura at 3000 feet on the north slope, one of indica at 4000 feet on the north slope, and one of indica on the summit (at Kodaikanal which is 5500 feet). It appears therefore that the two species may be segregated by eleva- tion, a circumstance which is not surprising in pairs of closely related and geographically sympatric species, and which evidently minimizes interspecific competition, enabling the two closely related species to evolve further in the same geographic area. The light coloration of macroura makes it seem more suited to a habitat of dryer, perhaps deciduous forest, and the generally dark color and preponderance of black in R. indica suggest that it is better suited to living in darker, denser, evergreen rain forest high in the mountains or on the windward slopes of the Western Ghats. The geographic distribution of these two species on the southernmost part of the mainland is further suggestive of this sort of ecological segregation. Khajuria (1955) listed both R. i. indica and R. macroura dandolena as members of the "humid element" of the mammal fauna of the Deccan. In view of the evidence advanced here, it appears that R. m. dandolena may now be better regarded as belonging to the mammalian fauna's "semi-humid element" of Khajuria's classi- fication. It may be noted that as revealed by Abdulali and Daniel (1952, color plate) the named forms of indica occupying the Western Ghats form something of a color cline in which the lightest form, which is most like R. m. dandolena, is the most distant from dandolena, and the darkest form, which is least like dandolena, is the one closest to and sympatric with dandolena. Less strikingly but correspondingly, the lightest-colored subspecies of the species macroura is the one which is sympatric with the very dark subspecies of indica, and the blackest subspecies of macroura is geographically the most distant from the area of overlap with the dark subspecies of indica. This status appears to represent the phenomenon known as character displacement described and discussed by Brown and Wilson (1956, p. 49). Intraspecific variation. — Varying sexual dimorphism in size is at- tributed by Phillips (1935) to the several subspecies of macroura. He finds the males larger than the females of the subspecies ma- croura in samples of three males and six females, and in foothills MOORE AND TATE: DIURNAL SQUIRRELS 37 material of dandolena in samples of five males and four females. But in samples of five males and six females of subspecies melano- chra he found the females lai:ger, and in the largest samples (Phillips, 1935, p. 224) of eleven lowland males and nine lowland females, that we here regard as dandolena, he found the sexes very similar in size. We suggest that the supposed sexual dimorphism is an artifact of the small samples, and that larger samples would show the sexes to be alike in size. The averages and maxima that PhilHps (1935, pp. 218-225) gives for body measurements of the several subspecies of macroura do sug- gest the existence of general size differences between the subspecies. The order of size is from melanochra, the largest, down through ma- croura to dandolena (see data in subspecies accounts of the present paper) . Ratufa macroura macroura (Pennant) Sciurus macrourus Pennant, 1769, Indian Zool., 1, pi. 1, and text. Sciurus ceylonicus Erxleben, 1777, Syst. Regn. Anim., p. 416. Sciurus ceilonensis Boddaert, 1785, Elench. Anim., 1, p. 117. Sciurus tennenti Blyth, 1849, Jour. Asiatic Soc. Bengal, 18, p. 600. Sciurus macrourus var. montanus Kelaart, 1852, Prod. Faun. Zeylon., p. 50. Sciurus macrurus Blanford, 1891, Fauna Brit. Ind. . . . Mammal., p. 374. Types. — Sciurus macrourus, not seen, from "Ceylon and Mala- bar," restricted by Phillips (1933) to "the highland jungles of the Central and Uva Provinces," Ceylon; ceylonicus, ceilonensis, mon- tanus, and macrurus are misspellings or alternative names proposed for earlier names given here; S. tennenti not seen, and not found by Robinson and Kloss (1918, p. 185). Material examined. — Pattipola, C. P. [Central Province], Ceylon, 6210 feet (B.M.), two. Discussion. — This is said to be a highland form (Phillips, 1935). It is apparently known only from specimens from the one locality, and is distinguished in these specimens by white tips on the hairs of its black tail, and a grizzled gray line along the sides between the black back and yellowish venter. Dimensions. — Phillips (1935, p. 218) alleges sexual dimorphism in this form, and provides averages of measurements in millimeters of three males and six females: length of head and body, males 346, females 375, length of tail, males 371, females [384], length of hind foot, males 68, females 71; length of ear, males 27, females 25.3. He records weights of two females that were evidently each three 38 FIELDIANA: ZOOLOGY, VOLUME 48 pounds, four ounces. For the average length of tail for the six females above, Phillips (1935, p. 218) actually gives "365 mm. (15.1 in.)." This figure in millimeters is less than length of head and body in the females, which seems a little improbable, less than length of tail in the males, which seems a little improbable, also, and does not equal 15.1 inches. The figure 15.1 inches does exceed Phillips' averages for length of body of the females and also length of tail in the males. Assuming 15.1 inches to be more correct, we have provided its equiv- alent in millimeters in brackets above. Ratufa macroura melanochra Thomas and Wroughton Ratufa macroura melanochra Thomas and Wroughton, 1915, Jour. Bombay Nat. Hist. Soc, 24, p. 36. Type. — BM No. 15.7.1.4, adult female from Kottawa, elevation 280 feet, Southern Province, Ceylon, collected April 12, 1913, by E. W. Mayor. Material examined.— "Fasdon, Corola," Ceylon (BM), one; "Ana- sigalla," Matugama, W. P. [Western Province] (BM), one; Kalutara, Matugama, Western Province (BM), one. Discussion. — This is apparently a geographic race occupying all the wet lowland area, which lies southwest of the highlands between the highlands and the sea (Phillips, 1935). It is like R. m. macroura in having a yellowish venter and a black back and tail, but differs from it by having no white-tipped tail hairs or gray line along the side. Dimensions. — Phillips (1935, p. 221) provides averages of meas- urements in millimeters of five males and six females: length of head and body, males 369, females 360; length of tail, males 387, females 388; length of hind foot, males 74, females 73.5; length of ear, males 25, females 27; weight, males 3 lb. 6 oz., females 3 lb. 2 oz. Ratufa macroura dandolena Thomas and Wroughton Ratufa macroura dandolena Thomas and Wroughton, 1915, Jour. Bombay Nat. Hist. Soc, 24, p. 36. Ratufa macroura sinhala Phillips, 1931, Ceylon Jour. Sci., Sec. B, 16, p. 215. Types. — Ratufa m. dandolena, BM No. 15.7.1.5, adult female, from Wellawaya, Uva Province, Ceylon, 608 feet, collected July 6, 1913 by E. W. Mayor, sinhala, BM No. 31.1.12.3, adult male col- lected at Nikawewa, near Kantalai, North Central Province, 250 feet, February 21, 1929 by W. W. A. Phillips. MOORE AND TATE: DIURNAL SQUIRRELS 39 Material examined, from Ceylon. — "Mousa Kanda," Gammad- uwa, C.P. [Central Province] (BM), one; Pukpitiya, Gammaduwa, Central Prov., 2000 feet (BM), one; Nikaweratiya, N.W.P. [North- west Province] (USNM), three; Tinnpani [Tirappane], N.C.P. [North Central Province], 275 feet (BM), one; Bintenna, Eastern Province (BM), one, (USNM), one; Mankeni, Eastern Prov. (BM), two; Maha Oya, Eastern Prov. (BM), three; Wellawaya, Uva Prov., 608 feet (BM), one; Kumbukkan, Uva (BM), one; Tellula, Uva Prov., 300 feet (BM), one; Ranna, S.P. [Southern Province] (BM), one; Hambantota District, S.P. (BM), two; "Wawonia" (USNM), seven; "Candelay" (USNM), two. Material examined, from southern India. — Kombu, south Coim- batore (BM), one; Chettiri Range, Salem District, Eastern Ghats, 2000 feet (BM), two; Kurumbapatty, Salem District, Eastern Ghats (BM), two; seven miles north of Gungwadoria, north slope Palni Hills, 3000 feet (BM), one; north slope of Palni Hills, 3000 feet (BM), two; Shrivilliputtur, Madura, southern India, 600-1000 feet (BM), four. Discussion. — This geographic race of Ratufa macroura occupies forests of the dry lowlands of Ceylon and also an area in southern peninsular India. It is distinguished from the black-backed, black- tailed other two subspecies by having dorsal pelage entirely brownish gray except for the black patch of the crown and varyingly a black- ish area across the shoulders "and middle line of rump." The tail hairs have longer light tips than do those of R. m. macroura and these produce a much grayer overlay that obscures the black part of the hairs. The short midventral pelage of tail is consistently whitish. Tails on which the pelage is old, worn, and curling at the tips may be bleached to the color of the venter (yellowish buff). This descrip- tion is based on Thomas and Wroughton (1915a, pp. 36-37) and notes on the United States National Museum material. Phillips (1935, pp. 221-223) regards dandolena as a subspecies confined to forests of the foothills of the central highlands of Central and Uva provinces, and distinct from sinhala which Phillips recog- nizes as occupying the dry lowlands of Ceylon. The characters that he attributes to dandolena on a sample of nine in his arrangement appear to us to describe intergrades between subspecies macroura and Phillips' subspecies sinhala. Since no character distinguishes Phillips' concept of dandolena from both the upland macroura and lowland sinhala, and the intergrade characteristics are not shown to be constant over a considerable geographic area, it seems better to 40 FIELDIANA: ZOOLOGY, VOLUME 48 recognize only the two subspecies, upland macroura and lowland dandolena including the foothills specimens as intergrades. Neither Zahn (1942, p. 13) nor Ellerman and Morrison-Scott (1951, p. 497) have followed Phillips' arrangement of dandolena and sinhala. Dimensions. — Phillips (1935, p. 224) provides averages of meas- urements in millimeters of 11 males and nine females from the low- lands of Ceylon: length of head and body, males 331.2, females 329.8; length of tail, males 330, females 350.6; length of hind foot, males 64.2, females 65.2; length of ear, males 24.1, females 25.1. Weights of one male and one female, respectively, are 2 lb. 5 oz., and 2 lb. 8 oz. + 72*E 4 Tb'E .•• ao-E + WE + ZfU KILOMETERS 500 +08*N Fig. 3. Species range of the Indian giant squirrel, Ratufa indica as revealed by material examined. Subspecies: A, dealbata; B, indica; C, maxima; D, centralis. MOORE AND TATE: DIURNAL SQUIRRELS 41 Ratufa indica (Erxleben) Definition. — Ratufa indica is the principal giant squirrel species which occupies the mainland of the Indian Subregion. Diagnosis. — (1) The ears have maroon tufts which extend about 20 mm. beyond the skin of ear tips, and which are not pointed but in- stead have a great many hairs about the same length. (2) There is a narrow, dark, usually maroon stripe extending ventroposteri- orly from the anterior edge of each ear, and separating the light- colored side of the face from the still lighter-colored bib on the side of the neck. (3) There is a contrastingly-colored line which passes in front of the ear and meets its fellow at the back of the crown be- tween the ears and separates crown from nape. (4) Some or all of the dorsal pelage of the sides, back, and tail is maroon (when the pelage is not badly worn and faded) ; the remainder is black. The above characters distinguish Ratufa indica from the other species as follows: macroura by 1, and 4; bicolor by 1, 2, 3, and 4; and affinis by 1, 2, 3, and 4. Discussion. — We are indebted to Abdulali and Daniel (1952) for their record and analysis of the relationships of pelage color differ- ences to geography in this species, and no deprecation of the value of their work is implied by our taking a stand for recognition of sub- species which are more strongly marked and have greater geographic range and fairly large areas of intergradation. See Figure 3. Moore (1960, p. 13) has drawn attention to how the distribution of this species bears on the Satpura hypothesis. Ratufa indica dealbata (Blanford) Sciurus indicua var. dealbatus Blanford, 1897, Jour. Bombay Nat. Hist. Soc, 11, p. 299, plate A, fig. 1. Type.— B.M. No. 96.11.7.6, adult male from Mahal, Bangs, In- dia, collected April 6, 1896, by R. C. Wroughton. Material examined, all from India. — "Khandeesh " Dangs (B.M.) one; "Mahal" Dangs (B.M.) one (topotype); Mheskatri Surat, Dangs (B.M.), one; Songadh, Nadsari District (B.M.), one. Type description. — Ratufa indica dealbata has the appearance of an albinistic indica. The dorsal coloration of type examined in 1951 is white above, becoming yellowish white on the thighs. The ear tufts and anterior part of face are yellowish white. The tail is almost white except for a darker shade on the basal two to three inches, caused by dark subterminal bands on the hairs which occupy the 42 FIELDIANA: ZOOLOGY, VOLUME 48 greater part of the length of the tail hairs. The under parts are white with a trace of yellowish on the insides of the hind legs. Discussion. — The original recognition of the existence of this form came from Blanford's report on Wroughton's collection of three speci- mens from the Dangs and Wroughton's examination of a captive specimen from the same place. Although Blanford's description and color plate left no doubt as to the distinctiveness of these specimens, subsequent authors (e.g., Ellerman and Morrison-Scott, 1951) have understandably been reluctant to accept this material as character- istic of a population of giant squirrels inhabiting a particular geo- graphic region. Salim Ali however, definitely established the con- tinuing existence of a population of these very pale squirrels in the Dangs fifty years after their original discovery there by Wroughton in 1896. Salim Ali collected six specimens of them in two localities additional to the type locality of Mahal (Abdulali and Daniels, 1952). The area known thus far to be occupied by these cream buff giant squirrels remains small. Nevertheless, the fact that they occupy the northwest extension of the species range, and the striking character of their local color difference, seem to justify recognizing them as a subspecies. Although the status of dealbata as a subspecies is accepted, some uncertainty remains about its color characters. The one principle character is well understood: the dorsal pelage is whitish or cream buff (or as Blanford says "pale rufescent sandy"). Blanford says that this becomes "slightly more rufous on the posterior portion of the body and on the outside of the hind limbs, and . . brown on the outside of the forelimbs and on the basal portion of the tail." (His artist confined this "brown" of the "forelimbs" to the hands, prob- ably correctly.) Whiter than the above were the "forehead, a band down the back, all the tail except the basal portion, and the lower parts. . . ." The tufted ears, Blanford said, were "bright rufus," and although that is how Blanford's colored plate showed them, when one of us (Tate) examined the type specimen in 1951, he re- corded the ear tufts as "yellowish white." Abdulali and Daniel (1952) who had four specimens, say "Ear tufts brown." Tate's notes indicate that the darkness of the basal portion of the tail is due to long, dark, subterminal bands on the tail hairs there. While conforming otherwise to the above description, the little colored figure of dealbata presented by Abdulali and Daniel (1952) shows the color patch between the ears as dark (brown) contrasting MOORE AND TATE: DIURNAL SQUIRRELS 43 with the cream buff dorsal pelage. Abdulali writes (letter of March 6, 1959), however, that "the colour plate is erroneous ... [in showing] a dark patch between the ears; . . . there is no white or pale patch between the ears [either], and the grizzled whitish of the nape and back is separated from the white of the forehead and snout by an almost straight line between the ears." Dimensions. — Blanford's (1897, p. 301) original description in- cludes measurements of a pair of adults of this subspecies taken in millimeters by R. C. Wroughton, who collected the animals: length of head and body, male 369, female 370; length of tail, male 417, female 408; length of hind foot, male 75, female 69. Ratufa indica indica (Erxleben) Sciurus indicus Erxleben, 1777, Syst. Regn. Anim., p. 420. Sciurus purpureus Zimmermann, 1777, Spec. Zool. Geogr. Quadr., p. 518. Sciurus bombayus Boddaert, 1785, Elenchus Anim., 1, p. 117. Sciurus elphinstoni Sykes, 1831, Proc. Zool. Soc. London, 1831, p. 103, Ratufa indica superans Ryley, 1913, Jour. Bombay Nat. Hist. Soc, 22, p. 436. Types and co-types — Sciurus indicus, lost; purpureus, lost; bom- bayus, lost; elphinstoni, BM Nos. 79.11.21.578-579, two adult males marked "Dekkan, India," and 16.3.9.12, an old female from "West- ern Ghats, Dekkan, India," all collected by Mountstuart Elphin- stone; superans, BM No. 13.6.21.3, an old female taken at Wote- kolli. South Coorg, December 28, 1912, by G. C. Shortridge. Material examined, all from western peninsular India. — Devikop, 2000 feet. South Mahratta (BM), two; "Wotekolli," 2000 and 3200 feet, southern Coorg (BM), three; "Makut," 250 feet, southern Coorg (BM), one; Ghatmatha, Satara District (BM), one; Helwak, Satara District (BM), one; Samasgi, 2000 feet, Kanara Border, southwest Dharwar (BM), two; Yellapur, 1800 feet. North Kanara (BM), one; Landa [Londa], Bombay Pres. (AMNH), one; "Kanara," southern India (BM), three; "India" (USNM), one. Pennant (1771, p. 281) described this squirrel well "from a stuffed skin in Doctor Hunter's cabinet" and said it "Inhabits Bombay," but he gave it no name. The scientific name indicus and the syno- nyms purpureus and bombayus cited above were all frankly proposed as scientific names for Pennant's Bombay squirrel, and all three of their Latin descriptions of it show some direct translation from Pen- nant's English description. The Erxleben description of indicus we translate, ". . . tail as long as body, with tip orange. Length of [head and] body 16 inches, tail 17. Tips of ears tufted. Head, back, sides, 44 FIELDIANA: ZOOLOGY, VOLUME 48 upper part of legs, thighs and tail dull purple. Inferior part of legs and thighs, with the belly, yellow . . ." Discussion. — Possibly from just south of the Bangs for something like 100 to 150 miles to Poona, but definitely at Bhimashankar, a point 40 miles directly east of Bombay, Abdulali and Daniel (1952) have demonstrated that the forest is occupied by giant squirrels that have hazel dorsal pelage instead of bay, and that have tails whitish distally for about half of their length. South-southwest of Bhima- shankar about 25 miles, those authors found at Khandala 11 out of 12 specimens to be bay instead of hazel, but at Khandala an aver- age of 0.40 (0.30 to 0.52) of the tail was whitish. Farther south they found bay dorsal color continued and whitish tip to the tail averaged only 0.25. This evidence suggests intergradation between the cream buff dealbata and the bay indica, and Abdulali and Daniel (1952) included the hazel material in the subspecies indica. However, if the hazel color and half white tail should prove to be constant in further material from the Western Ghats from Bhimashankar up to the vicinity of the Dangs, there would be a case for recognizing the giant squirrel of this area as a subspecies, R. i. elphinstoni. Abdulali writes, however (letter of March 6, 1959), that the additional ma- terial received by the Bombay Natural History Society since his 1952 paper was written had yet included "no hazel coloured speci- mens from between Bhimashankar and the Dangs . . ." Dimensions. — Riley (1913 p. 436) provides ranges of some meas- urements taken in millimeters on an unstated number of indica speci- ments from Dharwar and North Kanara: length of head and body, 340-380; length of tail, 370-446; length of hind foot, 73-77; length of ear, 25-33; greatest length of skull, 68-74; basilar length, 53.5-59; length of toothrow, 14.3-15.5; length of diastema, 15.5-18; length of nasals, 24.5-26; and zygomatic breadth, 46-49.5. Southward in the range of R. i. indica, a sample of 14 specimens considered identical with R. i. indica in color, was described as a new subspecies (Ryley, 1913, p. 436), R. i. superans, because an unstated number of the sample was discretely and notably larger than an un- stated number of ordinary subspecies indica from Dharwar and N. Kanara. Abdulali and Daniel (1952) reported another sample of three large specimens from near Hassan, and accepted superans as a good subspecies. Ellerman (1940), Zahn (1942), and Ellerman and Morrison-Scott (1951) have also accepted this as a good subspecies, but we are doubtful. Size alone in some mammals can evidently change with density of population (Scheffer, 1955), and in absence MOORE AND TATE: DIURNAL SQUIRRELS 45 of other differences "superans" is regarded here as atypical samples of the typical subspecies. Ratufa indica maxima (Schreber) Sciurus maximus Schreber, 1784, Saugethiere, 4, p. 784, pi. 217B. Sciurus malabariciis Scopoli, 1786, Del. Flora Fauna Insul., 2, p. 85. Sciurus indicus var. bengalensis Blanford, 1897, Jour. Bombay Nat. Hist. Soc, 11, p. 303, plateB, fig. 2. Types. — S. maximus, MNHN No. 93, adult female, from coast of Malabar, India, collected by Sommerat; bengalensis, BM No. 44.7.4.7, adult from "India." Material examined, all from southwestern peninsular India. — "Cotengady Estate," 3500 feet [Nelliampathi Hills, Palghat Distr.], Cochen (BM), four; Kodaikanal, top of Palni Hills, Madras (BM), one; "Anamaad," 3200 feet, Malabar (BM), one; "Kukkal, Shola," 6100 feet (BM), one; near "Machur," 4500 feet (BM), one; "Otta- coolie Estate," 3300 feet, Malabar [Nelliampathi Hills, Palghat Distr.] (BM), one; Penmudi [Ponmudi], Travancore (BM), one; "Northern slopes Palni Hills," 4000 feet (BM), one; Trivandrum, Travancore (BM), one; "India" (MCZ), one; Kellengode [Kollan- god], 3500 feet, S. India (AMNH), two; Nelliampathi Hills, 3500 feet (AMNH), one; Kil, Kotajiri, 5700 feet, Nilgiri Hills (BM), two; "Attikan," 5000 feet, Mysore (AMNH), one; "Mysore" (AMNH), four; Avalanche, 6500 feet, Nilgiri Hills (AMNH), one; Kalhatti [Falls], 4200 feet, Nilgiri Hills (AMNH), one; Mudumalai, 3285 feet, base of Nilgris (AMNH), two; Ootacamund, Madras, India (AMNH), one. Disctission. — Blanford's (1897) subspecies bengalensis was de- scribed from a single specimen with locality unknown. Ryley (1913, p. 436) reported a sample of 10 from Coorg to be identical in color with the type of bengalensis. This color seen in Blanford's color plate of the type, and the color plate of Abdulali and Daniel (1952) is different from indica only by having a black tail. Since bengalen- sis seems to be present in pure form only in a small area, and since the one character distinguishing it from indica is possessed also by the adjoining subspecies to the south, maxima, it is difficult to re- gard bengalensis as anything but a rather attenuated area of inter- gradation between subspecies indica and maxima. Obviously, it does not fit the characters of either of these, but it is intermediate. From fifty miles south of Coorg in the Nilgiri Hills the American Museum of Natural History has material from Ootacamund and 46 FIELDIANA: ZOOLOGY, VOLUME 48 Mudumalai which possesses the "bengalensis" color pattern, and Abdulali and Daniel (1952) report two other such specimens from the Wynaad Nilgiris. They report two further specimens from the Nilgiri Hills which differ from the hengalensis pattern only in having additionally a patch of black on the upper forelimbs (not the shoul- ders) but point out that this may perhaps be disregarded because it also occurs inconsistently in the subspecies indica. They then re- port that from Kotagiri, only ten miles east of Ootacamund, the material is colored in the centralis pattern (i.e. differing from typ- ical indica in having a black tail and separate black shoulder patches). From Avalanche, about 10 miles in the opposite direction from Oota- camund, and from Kalhatti [Falls], which lies between two localities for the hengalensis pattern, the American Museum has specimens which approach the maxima color pattern more closely, but the feet are bay, the black of the shoulders is discontinuous, and the Kal- hatti one has entirely red hind legs. (Incidentally, these approach Scopoli's type description of malabaricus, but could hardly be more obviously intermediate between all of these other bay-footed color patterns and maxima.) The point is made above that in the Nilgiri Hills the hengalensis color pattern intergrades through examples of the centralis pattern toward and very closely approaching the color pattern of maxima. Possibly they may mix so that all three pat- terns could be observed at one locality, or more than one pattern in one brood. If they do not, the details of intergradation would make an interesting field study. Pelage color. — From the Nelliampathi Hills less than 50 miles south of the Nilgiri Hills the American Museum of Natural History has three specimens, two from Kellangode, all of which possess the color pattern which is typical for Ratufa indica maxima. This pattern is fore feet, fore legs, and hind feet Light Ochraceous-Buff ; the shoul- ders and nape black entirely across from one upper fore leg to the other; tail, rump, and thighs entirely black with an incipient narrow black line extending forward in the middorsum; bib Light Ochra- ceous-Buff (XV) and face forward of the red subaural stripe about Apricot Buff (XIV); crown, subaural stripe, 20 mm. ear tufts, a small anterior portion of the nape, and the shanks of the hind legs Morocco Red; sides Ox-blood Red; ventral pelage (which is long and dense enough to hide the dark basal color of the hairs in two of the three) Warm Buff (XV); interaural patch Light Buff. Diagnosis. — The subspecific characters of maxima are: the con- tinuous black across the shoulders, the black rump, the light feet. MOORE AND TATE: DIURNAL SQUIRRELS 47 and the black tail tip, in that order of importance. The specimen from the Palni Hills which Abdulali and Daniel (1953, p. 731) de- scribe as a variant of maxima does not therefore seem importantly different, Moore wrote Humayun Abdulali to ask if in Bombay Natural History Society material maxima always has a single, con- tinuous shoulder patch and centralis always two separate shoulder patches. He responded (letter of March 6, 1959) that this is so. Habits. — Hutton (1949, p. 691) recorded observations of maxima in the High Wavy Mountains about sixty miles southwest of Ma- dura, some of which contribute toward understanding of factors which may influence their distribution: "A very common animal throughout the [broad-leaved] evergreen forests. It is not found in the deciduous forest. . . . They often use the same nests for some years . . . keeping them repaired and adding to them each year. I have examined several nests. . . . Their average size was 2 feet in diameter and the inside looked as if it would hardly accommodate an adult squirrel, being barely a foot across. They bring forth two lots of young each year, . . . one lot in September and another lot in March, the average per litter being two. Once only have I seen three. The adults . . . have no sense of territory. . . . When the young are past the milk-diet stage, both parents help in the feeding. Quite often I have seen these squirrels foraging together; one on the ground collecting nuts, etc., which the one in the tree knocks down. Very occasionally one comes across a squirrel larder at the base of some tree, which may contain upwards of two measures of nuts and other hard fruit." Dimensions. — Wroughton (1910, p. 889) provides several meas- urements of an unstated number of R. i. maxima, perhaps only one: length of hind foot, 80; greatest length of skull, 77; basilar length, 60; upper molar series, 15; length of diastema, 16.5; length of nasals, 24; zygomatic breadth, 49; interorbital breadth, 30. Ratufa indica centralis Ryley Ratufa indica centralis Ryley, 1913, Jour. Bombay Nat. Hist. Soc, 22, p. 436. Type.— BM No. 12.11.29.85, adult male from Bori, Hoshangabad Central Provinces, India, 1600 feet, collected February 13, 1912 by C. A. Crump. Material examined, all from India. — Amarakantak, source of Nar- budda River (BM), one; Bori, 1800 feet, Hoshangabad (BM), five (topotypes); Balapalli, 1000 feet, Palkonda Hills (BM), one; Kolle- gal, Billigirirangan Hills, Coimbatore (BM), two; Chamarajnagar, 48 FIELDIANA: ZOOLOGY, VOLUME 48 about 2500 feet, South Mysore (BM), one; "Gerung River, Manip- we" (BM), one; Koduru, 550 feet, Balapalli Range (BM), one; Luia, 1000 feet, Chaibassa, Bihar and Orissa (BM), three; Sangajata, 1300 feet, Chaibassa, Bihar and Orissa (BM), one; Dhain, 1400 feet, Hoshangabad (BM), two; no data (AMNH), two; Amraoti, Bastar [Berar] (AMNH), two. Pelage color. — The subspecies centralis as exemplified by the Am- raoti material is: Ox-blood Red on the dorsum, sides, ear tufts, and hind legs; black on the forelegs, shoulders (but separated by 30 mm. of red), and tail (which, however, pales at the tip to Light Buff XV) ; Cinnamon-Rufous (XIV) on the feet; Light Buff on the bib and in- teraural patch, but Buffy Brown (XL) on the side of the face; Liver Brown (XIV) on the crown. Dimensions. — Miss Ryley (1913, pp. 436-437) provides ranges in millimeters of body and skull measurements of an unstated number of her hypodigm of about 20 specimens of centralis: length of head and body, 309-343; length of tail, 382-433; length of hind foot, 72- 79; length of ear, 25-30; greatest length of skull, 69.5-74; basilar length, 50-58; length of tooth row, 14-15; length of diastema, 14-16; length of nasals, 21.5-24; and zygomatic breadth, 42-46. Abdulali and Daniel (1952, p. 473) provide ranges and averages of two measurements in millimeters for 19 specimens of centralis: length of head and body, 300-380, average 340; length of tail, 390- 450 (but an exceptional 270), average 405. Discussion. — This well-marked although variable subspecies oc- cupies low hills of central India, as its name suggests, and follows these eastward almost to Bengal. It also occurs in the Eastern Ghats, however, and southward along this chain to the state of Mysore and reaches the Western Ghats in the Nilgiri Hills where it intergrades in a confusing three-way mixture with maxima of the south and indica of the northwest. Habits. — At Sangajata and Luia, Bihar, collector C. A. Crump took 13 specimens and wrote the following comments (Wroughton, 1915c, p. 107) : "This handsome squirrel is gregarious and one of the most locally distributed animals I have ever come across. At Luia it inhabits a piece of jungle perhaps a mile square, and outside of this it is useless to search for it. Near Sangajata the favorite haunts were patches of jungle near the river, and though I passed through much of the Forest here, nowhere else, outside of the spots men- tioned, did I see the large conspicuous nests made by this species. The call is a loud rattle. . . . This squirrel lives among the most MOORE AND TATE: DIURNAL SQUIRRELS 49 lofty trees, it can take huge leaps, and is equally at home on a smooth trunk or scrambling among the slenderest twigs. I have never seen it lower than about 12 feet from the ground . . ." Ratufa bicolor (Sparrman) Definition. — Ratufa bicolor is the bibbed giant squirrel which ranges over the greater part of the Malaysian Subregion and Indo- chinese Subregion at least as shown in Figure 4. Diagnosis. — Ratufa bicolor has the following characters of the pel- age: (1) A large whitish or buff, bib-like area sharply marked off on the side of the neck is continuous with the color on the side of the face. (2) In most subspecies a broad, moustache-like, black line ex- tends from the black rostrum down and back through the vibrissal patch and the whitish or buffy pelage on the side of the face. (3) There is a small black spot in the whitish pelage of the chin, often divided midsagittally and finely into two parts. The above characters distinguish Ratufa bicolor from the other species of Ratufa as follows: affinis by 2 and 3; indica by 1, 2, and 3; macroura by 1, 2, and 3. Intraspecific variation. — In Table 2 some measurements of nine specimens of the species bicolor provide a bare indication of the dimensions one may find in species bicolor in the Indochinese Sub- region. The whole range of this species of giant squirrel, as revealed by the listings of Chasen (1940) and of Ellerman and Morrison-Scott (1951), extends from Nepal to Hainan, South Viet Nam, and Bali. No definition has previously been proposed for this broad concept of the species bicolor. It was somewhat of a practice in the past to include the forms south of Malacca Strait (bicolor, baliensis, palliata, etc.) in the species bicolor, but the mainland and insular forms north of that strait (peninsulae, phaeopepla, gigantea, etc.) were regarded as belonging in one or more separate species. Robinson and Kloss (1918, pp. 118, 195) thus recognized in what is now regarded the one species, several species separable at the Malacca Strait, the Isthmus of Kra, and along mainland lines less well defined. Although by 1922 Robinson and Kloss had evidently come to consider the above- named forms conspecific, Ellerman still accepted their 1918 classifi- cation for his "Families and Genera of Living Rodents" in 1940. No one has advanced any justification for recognizing a species divi- sion at the Malacca Strait in the first place, nor for abandoning it in the second; and we have empirically sought evidence on this. 50 FIELDIANA: ZOOLOGY, VOLUME 48 Fig. 4. Distribution of the mainland subspecies of Ratufa bicolor which occur north of the Isthmus of Kra: M, phaeopepla; N, leucogenys; 0, smithi: P, felli; Q, gigantea; R, hainana. The pelage characters common to the forms of Ratufa bicolor southwest of the Malacca Strait are: (1) The dorsal body pelage is generally light tipped. (2) the tail hairs are generally light tipped. (3) The black moustachial line is obsolescent. Numbers 1 and 2 provide striking contrast with the pelage characters of all the forms (of species bicolor) north of Malacca Strait except smithi. Pelage characters common to the forms under discussion on both sides of the Malacca Strait are: (1) The ventral body pelage is bicolored. (2) There is a contrastingly light bib on the face and neck. (3) A MOORE AND TATE: DIURNAL SQUIRRELS 51 black triangular mark is found on the chin. (4) There is a light flash mark on the dorsal pelage of the fore legs. (5) A light mark also ornaments the hind foot. In view of the number and character of the differences and similarities, a better case is deemed to exist for regarding these forms as conspecific. North of the Malacca Strait another place where the pied giant squirrels were regarded as divided into geographically juxtaposed species is at the Isthmus of Kra, where melanopepla and phaeopepla seemed to come together (Miller, 1913, p. 25; Robinson and Kloss, 1918, pp. 194-195; Ellerman, 1940, p. 389). The division of forms at the Isthmus of Kra was regarded as but doubtfully of species level by Robinson and Kloss (1918, p. 195), and restudy of the original series involved and consideration of new material leads us to con- clude that it is at best a subspecies division. See the discussion here in the account of subspecies R. b. phaeopepla. The next former alleged species division line farther north, which separated the alleged species phaeopepla and gigantea may be dis- tinguished in refined form on our map, Figure 4, separating subspe- cies gigantea and hainana from subspecies phaeopepla, leucogenys, and smithi. The differences that we find involved here are: (1) Ear tufts occur in gigantea and hainana but not in the other subspecies. (2) Flash marks occur on the fore legs of phaeopepla, leucogenys, and smithi, but not on the other subspecies. (3) The white of the bib and cheeks rises up in front of the eyes in gigantea and hainana but does not in the other subspecies. These taxonomic differences are discussed in further detail below, particularly in the subspecies ac- count of gigantea. One may note that much more taxonomic differ- ence has been found between these subspecies groups here than between those at the Isthmus of Kra. Relationships to other species. — Zahn (1942) has taken the present geographic line of separation as the one that parts the Indian species from the species bicolor. For our part, however, we find that separa- tion to be at the subregional boundary, which may loosely be said to be the barrier imposed by the Ganges River and its valley (but see Moore, 1960). The maroon species indica is separated from the pied species of giant squirrel across the subregional boundary by a more impressive assortment of characters, each of which has wide geo- graphic appHcabihty beyond the subspecies mentioned: (1) A promi- nent whitish or buffy mark separates the crown from the nape in indica centralis but not in bicolor gigantea. (2) A dark line descending from the ear divides the bib into cheek and neck components in cen- 52 FIELDIANA: ZOOLOGY, VOLUME 48 trails but not gigantea. (3) The ears of centralis have large Maroon tufts, but those of gigantea have small black tufts. (4) Light-colored pelage covers the feet, wrists, and ankles of centralis but in gigantea these are black. (5) Large areas of the fresh dorsal pelage are Maroon in centralis, but the back of gigantea is only black. (6) A black or blackish chin spot is present in gigantea but not in centralis. (7) A black or blackish moustachial line extends ventroposteriorly from the rostrum in gigantea but not in centralis. This line is not to be confused or equated with the facial stripe described above as char- acter number two, as Zahn (1942) did equate them. On the basis of the above differences we consider the range of the species Ratufa bicolor to go as far north and west as Nepal. We pre- sume that bicolor does not cross the Ganges River or intergrade any- where with Ratufa indica. We regard all of the giant squirrels of the Indochinese Subregion to belong properly to the species bicolor. The range of the species as indicated by the material we have examined extends from its northwestern locality in Nepal some 1500 miles eastward across Assam, upper Burma, southern Yunnan, Tonkin, and southern Kwangsi to Hainan, The northwest to southeast ex- tent of its range, including the extraterritorial part, is from Nepal to Bali. Indonesia, some 3000 miles. In the Academy of Natural Sciences collection at Philadelphia we found a giant squirrel specimen with characters which seem defi- nitely a mixture of those of species indica and bicolor. Nothing is known of its geographic origin, and it came to the Academy from the Philadelphia Zoological Garden. W. C. 0. Hill (1939, 1942) has re- ported successful breeding in both species macroura and species in- dica in captivity, and in view of the origin of the Academy specimen it seems likely that it represents a species cross between captive speci- mens of indica and bicolor. A less likely origin would be a relict patch of forest, from which no other museum specimens were avail- able to us, naturally inhabited by an interbreeding mixture of indica centralis and bicolor gigantea. Ratufa bicolor phaeopepla (Miller) Ratufa phaeopepla Miller, 1913, Smithsonian Misc. Coll., 61, no. 21, p. 25. Ratufa celaenopepla Miller, 1913, Smithsonian Misc. Coll., 61, no. 21, p. 26. Ratufa phaeopepla marana Thomas and Wroughton, 1924, Jour. Bombay Nat. Hist. See, 24, p. 227. Types. — Ratufa phaeopepla, USNM No. 124235, adult male from Sungei Balik, Tenasserim, Burma, collected February 25, 1904, by MOORE AND TATE: DIURNAL SQUIRRELS 53 W. L. Abbott; celaenopepla, USNM No. 124149, adult male from Domel Island, Mergui Archipelago, Burma, collected January 26, 1904, by W. L. Abbott; marana, BM No. 14.7.19.107, adult female from Mt. Popa, dry zone of upper Burma, collected September 26, 1913, by G. C. Shortridge. Material examined, from Thailand. — Me Ping River 195 miles N. of Bangkok (BM), one; Lai Yoke [Ban Sai Yoke] (BM), one; 20 miles west of Kampengpet, 450 feet (BM), one; Me Yam Valley, N. Phre 185 meters (BM), one; Me Taw Forest, Raheng (BM), one; Bank of Me Ping River, south of Hkampenpet, 120 to 180 feet (BM), two; Me Wong River, 53 miles E. of Um Pang, 800 feet (BM), three, (AMNH), three; 28 miles E. of Um Pang, 1750 feet (AMNH), two; "Wang Pratart Farm," Kam Peng Pet Prov. (CNHM), three; Ban Tong Ting (USNM), one; Raheng (USNM), one; Borphloy, Kan- chanaburi (USNM), one; Klong Klung, Kamphaengphet (USNM), two, (CNHM), five; Doi Phu Kha (USNM), one; Pak Koh (NR), one; Koon Tan (NR), three. Material examined, from Tenasserim, Burma. — Bankachon, V.P. (BM), four; Mergui (BM), one; Thagyet, Little Tenasserim River (BM), one; Thowngyoh [Theng-gan-ngok] (BM), one; Maliwun (AMNH), one; Taok Plateau (AMNH), one; "Bankusun" (BM), two; Victoria Point (BM), one, (USNM), two; Kisseraing Island (BM), two, (USNM), one; King Island (BM), one; Domel Island (USNM), two; Sullivan's Island (BM), one; Red Point (USNM), two; Sungei Balik (USNM), six; Telok Besar (USNM), five. Material examined, from Upper Burma. — Kodugwe, Pegu Yoma (AMNH), one; Camp Pinmezali, Pegu Yoma (AMNH), two; Mt. Popa (AMNH), five, (BM), two; 20 miles N. of Toungoo (BM), one; 30 miles N. to N.W. of Toungoo (BM), two; Kokkogon (AMNH), one; "Shenege Chaung, 1000 feet, Suiges Ruen," Mandalay (BM), one; Taho, Kareni (USNM), one. Dimensions. — From the describer's hypodigm of 16 specimens of phaeopepla, all collected by W. L. Abbott in southernmost Burma, we find the modal weight in 15 recorded on specimen tags by the collector to be four pounds. Only the pregnant female mentioned below weighed more, and the lowest weight, 3^ lb., is of an imma- ture male. The collector's measurements of phaeopepla were: length of head and body, 365-415 (average 386); length of tail, 390-500 (466). Twelve of the 16 were males. The type of celaenopepla was a male weighing five pounds, according to W. L. Abbott's label. See weights reported for R. b. gigantea. 54 FIELDIANA: ZOOLOGY, VOLUME 48 Re-examination of Miller's phaeopepla series of 16 specimens used in the original description, reveals 11 that are mature on the basis of ankylosis and tooth wear. The 11 range from 73.2 to 77.6 and average 75.5 in greatest skull length. The peninsulae available to Miller were only three, admissibly mature by the same criteria, but brought up to 12 by three adults more recently obtained from Kuala Lumpur, one from Bangnara, and five from Ban Don, the series at the U. S. National Museum ranges from 68.3 to 74.0 and averages 71.9. This difference from phaeopepla seems at least feebly to sus- tain Miller's proposition of a taxonomic break at the Isthmus of Kra, but we consider this to be at no better than the subspecies level. Pelage color. — The newer U. S. National Museum material from Bangnara and from Waterfall, Trong, Thailand, includes three speci- mens with molt lines across the back showing that the new pelage of peninsulae is much blacker than the freshest pelage of Abbott's phaeopepla series. This seems also to validate Miller's statement that fresh pelage color distinguishes phaeopepla from peninsulae; al- though the fresh pelage of a molting immature from Rumpin River in Miller's hypodigm of peninsulae seems to us no darker than some in his hypodigm of phaeopepla. Diagnosis. — Miller only distinguished celaenopepla from phaeo- pepla as having the fresh dorsal pelage much blacker. His type of celaenopepla is far darker than any U. S. National Museum phaeo- pepla, but none of the latter demonstrates by having a sharp molt line the ultimate blackness that may be reached by phaeopepla. Ratufa bicolor phaeopepla occupies the southern two-thirds of Burma, a range about 800 miles long. It must be noted that Miller's material was all from the very southern end of the range of phaeo- pepla, and since this range as we see it, borders upon that of the conspecific gigantea, felli, hainana, and leucogenys, it requires also to be carefully distinguished from those subspecies. Ratufa bicolor phaeopepla is distinguished from gigantea and hainana by possess- ing a buffy flash mark spread across the dorsal surface of its fore legs, and by lacking distinct ear tufts; from felli by the dorsal pelage be- ing blackish with no outstanding break in color from crown to tip of tail; from leucogenys by having bib and flash mark on fore leg more richly colored, Salmon Color (XIV) instead of the nearly white Cartridge Buff (XXX) ; from peninsulae by larger adult size. Discussion.— W. L. Abbott noted that the USNM No. 124247 which he collected March 27, 1904, from Telok Besar in southern Tenasserim weighed 4 3/^ lb. and had but a single fetus in the uterus. MOORE AND TATE: DIURNAL SQUIRRELS 55 It seems at least noteworthy that specimens from along the Me- wong River east of Um Pang [Ban Le Kathe], Thailand, have the pelage of their napes faded to form a prominent pale spot as is com- mon in leucogenys. Since the bibs and flash marks on the fore legs of three seem also somewhat light for phaeopepla, these may be con- sidered intergrades with leucogenys. The specimen from Borphloy [King Bo Ploei], Thailand, is likewise intermediate. Habits. — Field observations from the northern and southern lim- its of the subspecies range: "Local and not very plentiful on Mt. Popa, only occurring in the heavy evergreen forest near the top of the mountain." (G. C. Shortridge in Wroughton, 1915a, p. 472.) "Fairly plentiful, but more difficult to obtain in thick evergreen jungle than in the deciduous forests further north, as in other places these squirrels become rust coloured during the hot season. Weight 3-4 lbs." (G. C. Shortridge in Wroughton, 1915b, p. 712.) Ratufa bicolor leucogenys (Kloss) Ratufa melanopepla leucogenys Kloss, 1916, Proc. Zool. Soc. London, 1, p. 43. Type. — BM No. 15.11.4.43, old female from Lem Ngop, southeast Siam, collected January 15, 1915, by C. B. Kloss. Material examined, from southeastern Thailand. — Hinlap, 900 feet (BM), one; Krabin CBM), one; Sakerat (NR), one; Chanta- boon (AMNH), one; Twenty miles west of Kempenpet (AMNH), one; Twenty -eight miles east of Um Pang (AMNH), two; Me Wong River, fifty-three miles east of Um Pang (AMNH), three; Klong Yai (USNM), two; "Huey Yang," Sriracha (USNM), one; Kao Sabab (USNM), four; Nong Dom Ta (USNM), one; Nongkhor (USNM), two; Nong Mong Muang (USNM), one; Hoopbon [Ban Hup Bon] (USNM), one; Bor Phloy, Kanjanaburi (USNM), one; Ban Nong Bua (CNHM), one; Lem Ngop (CNHM), one. Material examined, from Cambodia. — Mt. Pra (BM), one; "Diai, Melton" (BM), one; Samba [Sambor] (BM), one. Original description. — "Characters. — Like R. m. peninsulae . . . but yellow of cheeks, fore limbs, and under surface markedly paler than the respective areas in that form, yellow on thighs more exten- sive and continued along the sides of the feet onto their upper sur- faces, where it occupies a considerable area, while the yellow of the fore limb extends to the bases of the toes above." Pelage color. — We made the following color notes from a sample of six specimens in the United States National Museum, and the 56 FIELDIANA: ZOOLOGY, VOLUME 48 Ridgway color terms used by Kloss are added in parentheses: bib and flash mark on fore leg are Cartridge Buff (Ivory Yellow) ; venter Apricot Buff (Pale Orange-Yellow). Diagnosis. — R. b. leucogenys differs from phaeopepla in having a paler bib and flash mark on the front leg, and in having a larger pale mark on the hind foot; from hainana by possessing a flash mark on the front leg and lacking ear tufts; from smithi by having no whitish tips on the dorsal pelage. Dimensions. — Measurements of three adults provided by Kloss in millimeters in the original description: length of head and body, 370, 360, 345; length of tail, 435, 435, 450; length of hind foot, 79, 80, 79; greatest length of skull, 71, 73, 71; condylobasal length, 60.5, 61, 58.5; palatal length, 27, 27.7, 26.3; diastema, 16, 16, 14.7; length of maxillary toothrow, 14, 14.2, 14; interorbital breadth, 26.5, 29.5, 26.7; zygomatic breadth, 46, 44, 44.5. Sex is respectively female, male, male. Ratufa bicolor sinus (Kloss) Ratufa melanopepla sinus Kloss, 1916, Proc. Zool. Soc. London, 1, p. 44. Type.— BM No. 15.11.4.41, adult female from Koh Kut Island, southeast Siam, collected December 26, 1914, by C. Boden Kloss. Material examined. — Koh Kut Island, southeast Siam (BM), one, (USNM), three. Diagnosis. — This subspecies is most like leucogenys to which it is geographically adjacent, and differs from it by having uniformly clear black dorsal pelage and richer colored ventral pelage, about Orange-Cinnamon we note, but Kloss describes it as varying from Ochraceous Buff to Ochraceous Orange and Ochraceous Tawny in the center of the abdomen. He noted that the nasal bones were rather longer in sinus than in leucogenys, their posterior terminations being more in line with those of the premaxillaries. Dimensions. — The original description provided measurements in millimeters of six adults, three of each sex. Ranges and averages of these are: length of head and body, 355-380 (366); length of tail, 415-450 (431); length of hind foot, 75-79 (77); greatest length of skull, 71-73 (71.9); condylobasal length, 59.5-61.2 (60.3); palatal length, 27-28 (27.3); length of diastema, 15.5-16.3 (15.8); length of maxillary toothrow, 14-14.2 (14.1) ; interorbital breadth, 28-30 (28.5) ; zygomatic breadth, (five specimens) 44^6 (45.1). Ratufa bicolor sinus is not mapped in Figure 4. MOORE AND TATE: DIURNAL SQUIRRELS 57 Ratufa bicolor smithi Robinson and Kloss Ratufa bicolor smithi Robinson and Kloss, 1922, Ann. Mag. Nat. Hist., [ser.] 9, 9, p. 89. Type.— BM No. 26.11.17.3, old female from Langbian Peaks, south Annam, 6000 feet, collected April 22, 1918, by C. Boden Kloss. Material examined. — Tay Ninh, Cochin China (BM), one; Tay- ninh Mt., Cochin China, 1000 meters (MNHN), one; Djiring, south Annam, 3500 feet (BM), three; Ban Me Thuot, Annam (CNHM), two; Langbian Peaks, south Annam (USNM), one. Discussion. — This is a strikingly distinct subspecies with a small, poorly known range in southern Vietnam. The describers had ex- amined nine specimens from three localities: Arbre Broy^ at 5400 feet elevation, Dalat at 4500 feet, and Dran at 3000 feet in the Lang- bian Mountains. It is notable that none of these localities, or of those from which we have examined material, is from below 3000 feet elevation, and that the greatest diameter of its known range barely exceeds 150 miles. Pelage color. — The characters which distinguish it are: buffy tips, to as long as 15 mm., on the hairs of the back, producing about Light Ochraceous Buff in USNM No. 269797; the light tipping of the hairs intensifies to produce a distinct patch on the posterior part of the crown, about Light Buff in the same specimen; the pelage of the sides, limbs, and tail is black when the pelage is fresh. The buffy tipping extends onto the thighs and base of the tail to a small extent and sometimes onto the forelimbs. The black parts do fade to brown to some extent, particularly toward the tip of the tail. The color of the hair tips on the back varies, probably from fading and wear, but the molt line across the back of USNM 269797 shows very little color change from old pelage to new. Ventral pelage Antimony Yel- low to Warm Buff, according to the describers, and bib, flash mark on fore legs, and frequently the hind feet, are the same color. Diagnosis. — Closely related to leucogenys, particularly by the pale nape patch, but also by the flash mark on the fore leg and the hind foot color, smithi is nevertheless sharply distinguished from it by the light-tipped pelage running the length of its back. From hainana, smithi differs by possession of the flash mark on the fore legs and by lacking ear tufts. Dimensions. — Kloss' measurements in millimeters of the type are: length of head and body, 415; length of tail, 500; length of hind foot, 89; length of ear, 31. The describers say that the skull of the largest 58 FIELDIANA: ZOOLOGY, VOLUME 48 of nine specimens measured 79 and 50 mm. in greatest length and breadth of skull respectively. Ratufa bicolor condorensis (Kloss) Ratufa melanopepla condorensis Kloss, 1920, Jour. Nat. Hist. Soc. Siam, 4, p. 71. (Not R. condurensis Miller, 1907.) Type. — C. Boden Kloss No. 2706 (not found by the present re- visers), adult female from Main Island, Pulo Condore, off the coast of South Viet Nam (not Pulo Condur of the Malacca Straits). This insular form shows close relationships to leucogenys and smithi in consistent possession of a light nape patch. It differs from both of these primarily in size, for condorensis is a pigmy giant squirrel, the skulls of seven specimens measuring 60, 61.5, 62, 62.5, 62.5, 63, 64mm. A greatest skull length of smithi (Robinson and Kloss, 1922, p. 93) is 76 mm., and for three leucogenys is 71, 71, 73 mm. (Kloss, 1916, p. 69). This small insular form differs from smithi by lacking light tips to the pelage of the back, but otherwise resembles both smithi and leucogenys closely in pelage color. No material representing this subspecies was examined during the present study. This subspecies is not shown on the map in Figure 4. Ratufa bicolor felli (Thomas and Wroughton) Ratufa felli Thomas and Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, p. 226. Type. — BM No. 15.5.5.55, adult male from Yin, East bank of Lower Chindwin, Burma, collected June 13, 1914 by G. C. Short- ridge. Material examined, all from Burma. — Dudaw-Taung, 650 meters, Pakokku Chin Hills (AMNH), two; Ainggyi, Pakokku Chin Hills (AMNH), one; Mt. Victoria, 500 meters (AMNH), one; Yin, Lower Chindwin (BM), 4; Okma, east bank of Chindwin R., N. Burma (AMNH), two. Pelage color. — This appears to be a well-marked subspecies with a small, and poorly known, geographic range. It differs from gigantea and hainana to the north by having no ear tufts, by having an inva- sion of buffy from the insides of the fore legs virtually across the dor- sal surface, which is otherwise black, by having a more pronounced border of the ventral pelage. It shares with hainana a somewhat richer ventral pelage color than that of gigantea, and also the light mark on the side of the hind foot, and it shares with peninsulae to MOORE AND TATE: DIURNAL SQUIRRELS 59 the south, all of these characters. It differs from all of its conspecific geographic neighbors, however, in that its dorsal pelage from the shoulders to the rump and the pelage of its sides from the fore legs to the hind legs is pale even when fresh. This ^vesfelli a general dorsal color pattern like that of Ratufa bicolor palliata of Sumatra: black on the head, nape, limbs, and tail, and light brown between the fore and hind quarters. In the present subspecies, however, the color of the sides is just like that of the back, not distinctly lighter as is the case in palliata. In one example of felli with apparently fresh glossy pelage the middle dorsal area is very close to but a little browner than Light Ochraceous Buff (XV), whereas in another it is about Cinna- mon-Buff (XXIX). The head and limbs are about Blackish Brown (1) XLV, and the nape and rump are intermediate. The intermedi- ate pale brown of the rump continues out the tail for about a deci- meter in the two with fresh pelage. The ventral pelage is blackish at the bases of the hairs on the hind legs as well as on the body, and is tipped with Cream Color to Warm Buff on an individual. The bib and blaze on the fore leg is Cream Color. It must be noted that the Mt. Victoria specimen is black dorsally like gigantea, but has no ear tufts, and is intermediate in the extent of the white blaze or flash mark across the fore leg, in having a small light mark on the side of the hind foot, and in the intensity of color of the ventral pelage. This specimen is either an intergrade between felli and gigantea or a challenge to the validity of felli as a subspecies. In view of the striking character of the felli material, we assume the former. Ratufa bicolor gigantea (McClelland) Sciurus giganteus McClelland, 1839, Proc. Zool. Soc. London, 1839, p. 150. Sciurus macruroides Hodgson, 1849, Jour. Asiatic Soc. Bengal, 18, p. 775. Ratufa gigantea lutrina Thomas and Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, p. 226. Types. — Sciurus giganteus, BM No. 79.11.21.336, young adult from Upper Assam, India, collected by J. McClelland; macruroides, BM No. 43.1.12.76-77, adult cotypes from Nepal collected in 1830 and 1840 by Hodgson; lutrina, BM No. 15.5.5.52, old female from Tatkon, near Kindat, Upper Chindwin River, Burma, collected July 5, 1914 by G. C. Shortridge. Material examined, from easternmost India. — Duragiri, 1600 feet, Garo Hills (BM), one; "Langting," 1500 feet, Cachar Hills (BM), two; Mokokchung, 4500 feet, Naga Hills (BM), one; "Naga Hills" 60 FIELDIANA: ZOOLOGY, VOLUME 48 Assam (BM), three; "Lakhani," Naga Hills, Assam, 1000 feet (BM), one, (AMNH), one; Tura, Assam (AMNH), two, (CNHM), two; "Chang chang Pani," Assam, 900 feet (AMNH), two; Ukhrul, Mani- pur, 6000 feet (BM), 1; Nongpoh, Khasi Hills, Assam (AMNH), 1, (CNHM), one; "Nepal" (BM), three; Pemionchee, Sikkim, 7000 feet (BM), one; Matanga River, 2500 feet, N. Kamrup (BM), two; Pashok, 3500 feet, Darjeeling (BM), one; Chin Hills 35 miles west of Kindat, 1500 feet (BM), one; Sevoke, Bengal (CNHM), two; Sangsir, 1400-2000 feet, Bengal (CNHM), three, (USNM), two; Tarkhola, Sikkim (CNHM), two; Karong, Manipur (CNHM), one. Material examined, from Tibet. — "Dening," 2250 feet, Mishmi Hills (BM), one; "Piki/' Mishmi Hills 1440 feet (BM), one; "Drexi," Mishmi Hills, 5140 feet (BM), one. Material examined, from Burma. — "Haingyan," 5000 feet. Chin Hills (BM), one; Hkamti, 500 feet, west bank upper Chindwin (BM), two; Kachin Hills 100 miles north of Myitkiyna, 1500 feet (BM), one; Moungkan, between Homalin and Tamanthe, east bank upper Chind- win River, 420 feet (BM), one; Lonkin (AMNH), one; Heinsun, west bank Chindwin R. (AMNH), one; Linhpah, west bank Chind- win R. (AMNH), one; Kindat (BM), one, (AMNH), one; Kabaw Valley, 20 miles west of Kindat, 600 feet (BM), one; Taukchaun, east bank of Chindwin, 500 feet (BM), one; Tatkon (BM), two; Road from Tamanthe to Sinnaing, west bank Chindwin R. (AMNH), one; Moungkan east bank Chindwin R. (AMNH), six; Hualung, east bank Chindwin R. (AMNH), two; 25 miles W. of Myitkyina (USNM), three; "Sedaw, Mandalay Canal and District," 300 feet (BM), one; Kamaing, Myitkyina Dist. (BM), one; N'bunghku (AMNH), one; Haibum (AMNH), four; Pumsin (AMNH), one; Htingwan, 4000 feet (BM), two; Hkani, 3000 feet, Gaw (BM), one; Singkaling Hkamti (AMNH), three. Pelage color. — From Nepal eastward across Sikkim, Assam, north- ern Burma, Yunnan, northern Thailand, and northern Indochina to Hainan the giant squirrel is characterized by tufted ears and en- tirely black dorsal pelage on the fore legs. In Assam and northern Burma we find that considerable American Museum of Natural His- tory material shows the tips of the ventral body pelage to be rather consistently pale, ranging between Cartridge Buff (XXX) and Light Buff (XV) on an individual (judged in places where the dark gray of the hair bases is entirely hidden by the pale tips). At Maymyo, Wan Tien, and the Nam Ting River, however, the ventral pelage shows enrichment at least to Light Ochraceous Buff, and this is true MOORE AND TATE: DIURNAL SQUIRRELS 61 of the material from northern Thailand. This more intense color is here considered to be but a dilution of the still richer color charac- terizing the ventral pelage of the subspecies described from Hainan, which is consistently Apricot Buff to Ochraceous Orange on each Hainan individual. Thomas (1923, pp. 85-86) has shown that in northern Thailand the giant squirrel has a little patch on the side of each (otherwise quite black) hind foot that is the color of the ventral pelage. He found that this character diminishes greatly in northern Burma and Assam, and is entirely absent in the Mishmi Hills, Sikkim, and Nepal. We note, however, that it continues strong from northern Thailand to the westward and characterizes the subspecies which had been already described from Hainan, and so is not diagnostic for the subspecies stigmosa which Thomas (loc. cit.) erected upon it in Thailand. We consider that this character like that of the rich color of the ventral pelage, may emanate from Ratufa bicolor hainana. Diagnosis.— Consequently, Ratufa bicolor gigantea may be iden- tified by presence of ear tufts, black dorsal pelage on the forelegs, pale ventral pelage (about Cartridge Buff to Light Buff), and scarcity of any buff pelage on the side of the hind foot. The Maymyo and Nam Ting River specimens are apparently intergrades with hainana to the southeast. Variation. — Possibly there is a geographic race which should be recognized as Ratufa bicolor lutrina, occupying an area in northern Burma. However, the characters on which lutrina has been based are virtually identical with the faded condition of the pelage of some undoubted gigantea just prior to molt. If the brown color of lutrina were the color of the fresh pelage, and if it predominated in a geo- graphic area, we would freely accord subspecies status to lutrina as have Zahn (1942), Carter (1943), and Ellerman and Morrison-Scott (1951). There could, of course, quite by chance be taken in a given area. A preponderance of specimens whose pelage had come in black when fresh but had faded to brown at the time of collection. There could also be a greater tendency in a geographic area for the pelage to fade quickly to brown, so that it is brown for a longer time before being shed and replaced by black. This quick change to brown could be effected by a special genetic factor or a special ecological factor or both in combination. Carter (1943) considered that this frequency of brown pelage was related to the animal's occurrence in the savan- nah forest, and he allocated the blacker specimens that he studied from the adjacent more heavily forested districts to the subspecies 62 FIELDIANA: ZOOLOGY, VOLUME 48 gigantea. This necessitated arbitrary placing of a series from the deep forests of Lonkin and Singkaling Hkamti in gigantea even though the series is almost as brown as the alleged lutrina. Since there are small spots of new black pelage coming in in two of the brown specimens from Maunkan, the general brown color of the dor- sal pelage of these "lutrina'' is evidently produced by fading, and on the basis of the scanty information on its distribution now at hand, we give no credence to "lutrina" as a valid subspecies. There is, further, some individual variation in pelage color with- in gigantea which should be mentioned. Near the base of the tail for about a quarter or a third of the tail length the short, appressed ventral hairs of the tail are usually buffy instead of black. In one specimen from Tura this buffy line extends for two-thirds the length of the tail. This specimen also has reddish subterminal color bands on the hairs of the dorsum and tail which contrast strongly enough with the basic blackish brown to be quite noticeable. Further, it has a splash of white-tipped hairs in the black dorsal pelage of each fore leg (about 15 mm. in diameter) and a whitish tipping to the hairs on one side of the hind foot. Another specimen from Tura has none of these variations from proper gigantea characteristics. A specimen from Nongpoh in the Khasi Hills is quite as brown as any of the alleged lutrina, and has even the pale tail tip. A specimen from "Changchang Pani," Assam, is richly colored enough ventrally and possessed of a strongly enough marked hind foot to be an intergrade with hainana were that geographically possible. One Hualung speci- men from the east bank of the Chindwin River has almost whitish subterminal bands on many of the hairs of the dorsum, creating a rather agouti effect. Habits. — In Sikkim collector C. A. Crump took 14 specimens at six locaHties and wrote these notes (Wroughton, 1916a, p. 486): "This squirrel is usually found in pairs and is not gregarious. It is more plentiful between the Terai and the low valleys up to 3,000 feet, being particularly partial to the dense tall jungle bordering rivers. It has a loud clacking note common to most squirrels but calls only on rare occasions. It is an extremely active chmber and as a rule is wary, making off and hiding in the thick foliage immediately dan- ger is feared." Dimensions. — Collector H. Stevens noted weights of the three adult squirrels from Sangsir and Sevoke, Bengal Residency, India, on the specimen tags as four, four, and five pounds. These are all males taken in November and December, and are CNHM Nos. MOORE AND TATE: DIURNAL SQUIRRELS 63 35432, 35434, and 35437. This may be the giant squirrel's best claim to being the largest tree squirrel in the world. He found the female specimen from Tarkhola in the Teesta Valley of Sikkim to weigh 43^ pounds. Discussion. — In January, 1931, Lord Cranbrook journeyed north- west from Putao, Burma, across the drainage of the Mali Hka to the Nmai Hka. While in the drainage of the former he heard (letter to us of May 5, 1961, based on his field notes) or saw giant squirrels daily. But once over the 6500 foot divide into the Nmai Hka Val- ley, he saw no more Ratufa; although "the vegetation and climate seem to be exactly the same on both sides of the divide." (Cran- brook, in Kinnear, 1934, p. 348.) Ratufa bicolor hainana (J. A. Allen) Ratufa gigantea hainana J. A. Allen, 1906, Bull, Amer. Mus., 22, p. 472. Ratufa gigantea stigmosa Thomas, 1923, Jour. Bombay Nat. Hist. Soc, 29, p. 86. Types. — Ratufa g. hainana, AMNH No. 26638, adult male from Cheteriang, Hainan, collected in 1903 or 1904 by agents of Alan Owstan; stigmosa, BM No. 98.10.5.40, adult female from 730 me- ters on Doi Sritepe, Chiengmai, Siam, collected April 10, 1898, by T. H. Lyle. Material examined, from Hainan. — Mt. Wuchi (AMNH), four, (BM), two; Nam Fong (AMNH), one; Mts. S. W. of Kachek (USNM), one. Material examined, from Chinese mainland. — Wa-tien, Yunnan (AMNH), one; Wan Tien (MCZ), one; Shui-kow-kwan, Lungchow, Kwangsi (CNHM), one; Nam-ting River at Burma border, 1700 feet (AMNH), one. Material examined from Indochina. — Bolovens Plateau, Laos (AMNH), two, (BM), one; Chapa, Tonkin (MCZ), two, (BM), two; 150 mi. west of Xieng Khouang, Laos (MCZ), one; Mt. Fansi- pan. Tonkin (MCZ), one; between Harsa and Lao Fou Thai, 3000 feet, Laos (CNHM), one; Muong Yo, 2300 feet, Laos (CNHM), two; Ban Phone (CNHM), two; Thateng (CNHM), two; Banteai (CNHM), one; Hoi Xuan, Annam (CNHM), one; Phong Saly, 4400 feet, Laos (CNHM), one; Tam Dao, 3000 feet. Tonkin (BM), five; Nape [Na Pe], 2500 feet, Laos (BM), three; Phu-Qui, 100 feet, An- nam (BM), one, (MNHN), three; "Nghia-hung," Phu-Qui, 100 feet (BM), two; Xien Tuang-Koo [Xieng Khouang], Laos (BM), one; "Ci Norn," 1200 meters (MNHN), one. 64 FIELDIANA: ZOOLOGY, VOLUME 48 Material examined, from Thailand. — Doi Sutep, Chiengmai (AMNH), one; Doi Pui, Chiengmai (AMNH), one; Doi Hua Mot (USNM), two; Doi Nangka (USNM), two; Doi Nangkeo (MCZ), two; Mt. Angka (MCZ), three; Ching Dao (MCZ), one. Material examined, from Burma. — Maymyo (AMNH), one; Gok- teik, 2133 feet, Northern Shan States (BM), one; Katha (BM), one; Madaya Forest, 55 miles north of Mandalay, 2000 feet (BM), two. Original description. — "Whole upper parts, outsides of limbs, and the tail uniform intense black; ventral surface and inside of limbs rusty yellow, the basal half of the pelage over the chest and belly brownish black, showing more or less at the surface over the central part of the abdominal area; a broad black cheek stripe, and two small spots of black on chin. Ears tufted . . ." J. A. Allen gave this de- scription from only the one type specimen, but it is quite good for the additional material subsequently available from Hainan reported here. From our inspection of the above material, however, his sug- gestion that the nasal bones of hainana are longer than those of gigantea is not sustained. For further discussion of the characters of hainana, see the account of gigantea. Habits. — A collector and observer of this squirrel reports: "Plenti- ful in all big forests especially round Gokteik, quite identical in habits with [Indian] giant squirrels." (G. C. Shortridge in K. V. Ryley, 1914, p. 721.) INDIAN STRIPED SQUIRRELS Genus FUNAMBULUS Lesson Funambulus Lesson, 1835, Illustr. de Zoologie, pi. 43, and 2 pp. of text. Palmista Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 279. Tamiodes Pocock, 1923, Proc. Zool. Soc. London, 1923, p. 215. Type species.— Funambulus, Sciuru^ indicus Lesson {=S. palma- rum Linnaeus) ; Palmista, fixed as F. palmarum Linnaeus by Thomas, 1897, Proc. Zool, Soc. London, 1897, p. 933; Tamiodes, Sciurus tris- triatu^ Waterhouse. Definition. — The genus Funambulus comprises the species pen- nanti, palmarum, tristriatus, layardi, and sublineatus, all of which are small to medium-sized tree squirrels and endemic to the Indian Sub- region. (See their distributions in Figures 7-11.) Diagnosis. — (1) The baculum of Funambulus is a single unit, lack- ing the separate bony blade characteristic of some squirrels (Pocock, 1923). (2) There is (as shown in the lateral views in Figures 5 and 6) no dorso-anterior process of the premaxillary bone rising to abut upon the anterolateral angle of the nasal bone (Moore, 1959, p. 170). (3) There are (as shown in the ventral views in Figures 5 and 6) one or two transverse bony septa crossing the auditory bulla (Moore, 1959, p. 170). (4) There are in parous females but two pairs of func- tional mammae (Moore, 1961a, p. 8). (5) The dorsal pelage is longi- tudinally striped with 3 or 5 light stripes contrasted with brown or black pelage between stripes. (6) The coronoid process of the man- dible is low and only incipiently falcate. (Compare Figures 5 and 6 with Figures 1, 12, 17, 22, 23, 30, and 32.) The above six characters distinguish Funambulus from other Sciurinae of the Indian and Indochinese subregions as follows: from Ratufa by 2, 3, 4, 5, and 6; Callosciurv^ by 1, 2, 5, and 6, Tamiops by 1, 2, 4, 5, and 6, Dremomys by 1, 2, 4, 5, and 6; Menetes by 1, 2, 4, and 5; and Sciurotamias by 2, 3, 4, 5, and 6. Systematic history. — For many years the genera Dremomys, La- riscus and Menetes were confused with Funambulus. A number of forms belonging in those genera were described as forms of Funam- 65 5 I ■2^ 2 . O '^' «-> ■& O eo SO) .- 'C u a> 0) D. O. CO CO « o g •S Q, .2^ 2 I ■J JS-^^* c- t-OOOOOOOOOOCT>OOOOOS0500TH0050SC-t-00 Q -g 1-1 i-H i-H .-H ^ to boS Crt '-iot-i(m -i-Hoc^i-irHi-ieo-^iniMWi-iaj -o ^j^ ^^^^ .^^^^^^^^^^^^ .^ jj,^;g to ■oooT-Hoo;o'«i* -H ^ 00 1-1 (M U3 0:1 CO CO CO -H -H -H -H -H w iOTf-^cJt)-^eooot-coco-^oo(M -H ^ - -H -H -H -H ^oeO'^oec«ocoooooooo>o"5t-u3ooio »OTj<o>iOrj0 t-l,-lrHi-(i-(>-li-lTHTH.-l(M 66 MOORE AND TATE: DIURNAL SQUIRRELS 67 Fig. 5. Skull and left mandible of the Indian striped squirrel, genus Funam- bulus, species tristriatus, AMNH No. 54652, X 1. Note that there is no dorso-ante- rior process of the premaxillary bone rising to abut evenly with the antero-lateral angle of the nasal bone as is present in all other genera of Oriental squirrels. hulus. The relationship was clarified when Thomas (1908, 1914, 1916a) separated Lariscus and "Zetis" (=Menetes) from Funambulus of the Indian peninsula and Ceylon. This distinction was further em- phasized by the marked differences in the bacula discovered by Po- cock (1923), whose consequent placement of these genera in separate subfamilies was recognized by Simpson (1945) as of tribal difference. Prasad (1951, 1954, 1957) discovered that the male reproductive tract of Funambulus is of a non-penile-duct type, extremely different from the type considered characteristic for Sciuridae, and (1957, p. 21) proposed that this genus be separately raised to subfamily rank. Moore (1959, p. 170) found a skull character which links Funambulus with African genera Funisciurus, Paraxerus, and Myosciurus, the male genital tracts of which are not known, and left these four genera all in the tribe Funambulini Simpson. Early systematic treatments of the genus Funambulus are few: Blyth (1849, 1852) dealt with Ceylonese races. Wroughton (1905) discussed the identity of F. palmarum at length and distinguished it from the north Indian F. pennanti. Thomas and Wroughton (1915b) published a key to the Funambulus of Ceylon. Wrough- ton (1916d) worked out the races of the species palmarum and tris- triatus of peninsular India. Pocock (1923) showed the characters of the glans penis and baculum in F. palmarum and F. tristriatus and solely on the differences in these separated the species tristriatus at the generic level as Tamiodes. Phillips (1928) discussed the vari- 68 FIELDIANA: ZOOLOGY, VOLUME 48 ous Funambulus from Ceylon. W. C. 0. Hill (1936) examined the bacula of all Ceylonese Funambulus, and was puzzled to find that the bacula of all the Ceylonese forms thought to belong to species Funambulus palmarum actually display the characters which Pocock (1923) ascribed to Tamiodes tristriatus and differ, just as tristriatus was supposed to differ, from Pocock's (1923) characterization of F. palmarum. Prasad (1957, p. 7) supports Hill's findings with an excellent figure showing the baculum of Ceylonese F. palmarum kelaarti to be like that of tristriatus, also. Furthermore, Prasad (1954, fig. 2) had already shown that the baculum of F. palmarum palmartim of India was also quite a good deal more like Pocock's figure of tristriatus than like Pocock's palmarum. It remains only, then, to note that Prasad's (1957, fig. 2) illustration of the baculum of Funambulus pennanti of northern India is like Pocock's (1923, fig. 20) of F. palmarum. Pocock's specimen alleged to be palmarum was therefore evidently pennanti. Actually, as Hill (1936) and Pra- sad (1954, 1957) have now shown, the bacula of Funambulus palma- rum indicate close relationship to F. tristriatus, and it is F. pennanti which is very different. This solution to the Tamiodes puzzle is almost equally well established by the close correspondence of the characters described by Pocock (1923, p. 215) for the glans penes of his supposed Funambulus palmarum and Tamiodes tristriatus respec- tively to the illustrations of F. pennanti (Prasad, 1957, fig. 1) and F. palmarum (Prasad, 1954, fig. 2). Furthermore, since no one has yet pubUshed a description or figure confirming the male genital characters of tristriatus, it remains possible that Pocock's (1923) tristriatus was in fact a specimen of palmarum and that male genital characters for tristriatus remain unknown.^ Pocock's mistake with pennanti must be a fairly easy one to make, for Didier (1953, p. 69, fig. 3) has evidently made it again, illustrat- ing in detail three views of the baculum of "Funambulus palmarum L. (Hindoustan) ." The figures quite evidently represent the baculum which Prasad (1957) has shown to be that of pennanti. Zahn's (1942) revision of this genus is distinguished particularly by his inclusion in it of the striped squirrels of the subgenus Tamiops. Ellerman and Morrison-Scott (1951) did not follow this superficial arrangement, and Moore (1959, 1961a) has produced new evidence 1 Recognition now of the nature of the error involved in Pocock's concept of tristriatus as a distinct genus justifies earlier abandonment of this concept by vari- ous authors because of Hill's (1936) evidence of its incongruity. The difference recognized by Pocock (1923) still exists, differentiating pennanti (not tristriatus) from the other four species, and will be honored here as of subgeneric significance. MOORE AND TATE: DIURNAL SQUIRRELS 69 that supports Pocock's (1923) in classifying Funambulus with cer- tain African squirrels in the tribe Funambulini but Tamiops with Oriental genera in the tribe Callosciurini. Further evidence is also offered in the present paper in the discussion under Tamiops. Parameters of the populations of subspecies of the genus Funam- hulus as represented by museum material would be an important contribution to knowledge, although it is too intensive for so exten- sive a revision as the present one. Some indication of the variation in size within the genus, however, may be seen in Table 3. The sample representing species palmarum in Table 3 may give a fair indication of where the mean for any of these measurements in that species might fall. Distribution. — This primarily tropical genus inhabits India and Ceylon. Three of its five species are represented by seven forms on Ceylon. Nine other races representing all five species, appear to occur on the mainland only below the Tropic of Cancer, leaving five races (representing only two species) which occur north of it. Intrageneric relationships. — The constancy of the distinctive striped pelage pattern causes the species of Funambulus to seem extremely close in relationship to one another. The most distinct of the five species is pennanti, for not only does it possess five white stripes instead of the three common to the other four species, but its rod-like baculum lacks any suggestion of the bifurcated tip now re- ported to occur in the other four species, and the shape of its glans penis apparently also differs from that of the other four (Hill, 1936, figs. 1-5; Prasad, 1954, fig. 2; Prasad, 1957, fig. 1). One might anticipate finding an interesting amount of difference between the two species of jungle squirrels, sublineatus and layardi, which occur in heavy jungle in southern India and Ceylon. Their glans penes and bacula as illustrated by Hill (1936, figs. 4 and 5) appear to distinguish the two from each other as much as from pal- marum. From a very meager amount of material, one skull of F. s. sublineatus, UMMZ No. 81078, three of F. s. obscurus, MCZ No. 27550, NMC No. 60-x-E, NMC No. 60-x-C, and two of Funambulus I. layardi, NMC No. 59-H, and NMC No. 59-N, and a series of three photographic views each of the types of kathleenae (=S. s. obscurus), sublineatus, delesserti (= sublineatus) , and F. layardi signatus, before the senior author, it appears that the interorbital breadth just be- hind the supra-orbital notches (to avoid a broken place), is 0.81, 0.84, and 0.85 of the greatest length of frontals in the three layardi and is 0.60, 0.63, and 0.68 in three of the sublineatus. If this difference is 70 FIELDIANA: ZOOLOGY, VOLUME 48 constant, it is a large one, and suggests an important difference in ecological niche. The broad frontals with more lateral orientation of the eyes generally characterizes squirrels whose habits take them high in the trees. Narrower frontals and more upward orientation of the eyes characterize the species which are adjusted to living close to, or on, the ground. Since one would expect some niche difference between two closely related squirrels of the same genus which are said to inhabit the same jungles and which differ in size, it is espe- cially interesting to find that Phillips (1935, pp. 238 and 241) reports layardi to inhabit the tops of the tallest trees in the tallest and heavi- est forests, but sublineatus to inhabit the "undergrowth in the jun- gle." See details quoted in species account below of sublineatus. Fig. 6. Skull and left mandible of the Indian striped squirrel genus Funam- bulus, species sublineatus, UMMZ No. 81078, X 1. Note the character that distin- guishes Funambulus from all other Oriental squirrels, absence of any dorso-anterior process of the premaxillary bone rising to abut evenly with the antero-lateral angle of the nasal bone. Note also the characters that have led to sublineatus being con- sidered an incipient pygmy squirrel (Moore, 1959, pp. 186-190). Charles McCann (letter of June 21, 1961) comments on apparent ecological segregation of species on the mainland nearby: "... [near] the Palni Mountains pennanti is found on the plains, palmarum on the hills up to about 4,500 feet, and above occurs the little F. subline- atus feeding chiefly on Rubus species in the undergrowth, its normal habitat. The green viper, Trimersurus macrolepis, appears to be its natural enemy up to 7000 feet." (This locality, latitude 10° 20' N., is about 5° farther south than the southernmost locality from which we have examined pennanti material.) A further aspect of the same species character which distinguishes the above-mentioned available material of sublineatus from layardi (and also from palmarum, pennanti, and tristriatu^) is obsolescence in sublineatus of the supra-orbital notch, which otherwise seems to be well developed throughout the genus. MOORE AND TATE: DIURNAL SQUIRRELS 71 The material on Funambulus in American museums and hence available to Moore in the present study, has been extremely small for the assessment of the validity of subspecies and to a much lesser extent than for other genera has it been possible to push the study beyond Tate's review concept to a revision. Moore (1960) has pointed out that the squirrels confronting Funambulus pennanti across the Garo-Rajmahal Gap are tribally different, and that much zoogeographic importance may attach to detailed further field study of the distribution of F. pennanti and Callosciurus pygerythrus in this region, and where the two also appear almost to meet along the foothills of the Himalayas. See also a note in the account of C. pygerythrus blythi in the present paper. The importance and need of more data on the breeding of F. pennanti has also been shown (Moore, 1961, pp. 21, 27, 29). KEY TO THE SPECIES OF FUNAMBULUS 1. Dorsal pelage marked by five longitudinal light stripes, and baculum a simple, slightly curved rod pennanti Dorsal pelage marked by three longitudinal light stripes, and baculum bifur- cated at distal end 2 2. Fur long, tending to obscure the pelage stripes, supraorbital notches absent or obsolescent, bifurcations of baculum ending in large knobs sublineatus Fur short, stripes quite distinct, supraorbital notches usually well developed; bifurcations of baculum without large terminal knobs 3 3. Midstripe, at least, rather brightly colored (orange-yellow, etc.), ventral pelage richly colored (russet, chestnut, yellow-orange, etc.), bifurcation of baculum obsolescent (or only incipient?) layardi Midstripe and ventral pelage only pale buffy, bifurcation of baculum quite distinct 4 4. Occipitonasal length usually exceeding 40 mm.; length of palate (front of in- cisors to back of palate) usually exceeding half of the occipitonasal length. tristriatus Occipitonasal length usually less than 40 mm.; length of palate usually less than half of occipotonasal length palmarum PRASADSCIURUS new subgenus Definition. — This subgenus includes only the species Funambulus pennanti Wroughton which is endemic to the northern part of the Indian Subregion. See its range as plotted from material we exam- ined in Figure 7. Diagnosis. — The baculum is an attenuate rod without apical bi- furcation. The glans penis is extended apically in a slender point. There are five longitudinal light stripes in the pelage of the dorsum and sides. 72 FIELDIANA: ZOOLOGY, VOLUME 48 Funambulus pennanti Wroughton is the type species. The sub- generic name recognizes the work of the Indian scientist, Dr. M. R. N. Prasad, whose description of the genital characters of pennanti pro- vides the critical clue to clarification of the Tamiodes puzzle. Funambulus pennanti Wroughton Definition. — This species includes all of the five-striped forms of Funambulus and is endemic to the northern part of the Indian Sub- region, most of its range lying north of the Tropic of Cancer. See Figure 7. Diagnosis. — Same as for subgenus Prasadscuirus. The following characters seem also to have diagnostic value in the material at the American Museum of Natural History. The longer pair of lateral stripes extends posteriorly over the rump to the base of the tail, and clearly forward upon the shoulders and nape to reach the ear. The middle stripe extends posteriorly onto the tail. The head of pen- nanti bears two nearly parallel stripes, a faint light stripe from ear to eye, and a stronger, almost white one from beneath the base of the ear forward below the eye, and sometimes quite clearly halfway out to the tip of the snout from the eye. This species possesses no bright, warm color about the anus or extending out the ventral side of the tail. Relationships to other species. — In describing this species, Wrough- ton (1905) remarked, "It would almost seem that palmarum is a South and pennanti a North Indian form . . . they occur together on the West Coast. . . ." Our investigation certainly supports this early concept of distribution of the two species. See Figures 7 and 8. Only F. palmarum rohertsoni and F. p. bengalensis occur within the range of F. pennanti, and except for a southward invasion just east of the Western Ghats to Dharwar (latitude 15°27'N.), the spe- cies pennanti seems to range only north of the twentieth parallel of latitude. A collector (C. A. Crump in Wroughton, 1915c, p. 108) reports observations on this species: "Common at Chainpur and well dis- tributed throughout Hazaribagh where, however, it overlaps with F. palmarum, the latter I think predominating. At several places I shot both these squirrels on the same ground but did not find them to- gether in any one tree." Separately one of us has treated the known interrelationships of these two species in more detail (Moore, 1960, p. 6) and speculated on their origin as species (pp. 9-12). MOORE AND TATE: DIURNAL SQUIRRELS 73 Mr. Charles McCann of the Dominion Museum, Wellington, N. Z., contributes (letter, June, 1961) the following comments on pennanti from his years in India with the Bombay Natural History Society : "Habitat. — An inhabitant of the open plains and scrub country. It is more commensurate with man and is frequently associated with villages, whether on the Deccan Plain or the Sind Desert. Does not occur in jungle country except, perhaps, on the outskirts. "Coat. — Harsher, smoother, and lying more flat than in palmarum. B • 80* -fjZ'N TIBET y 'N. INDIA • • •r 7iJ^/24^SBi^ 0 MILES -WOV 1 ' I I I 6i' ^^OFJbE^Al 4- +I6°N Fig. 7. Geographical distribution of the five-striped Indian squirrel, Funam- bulus pennanti, as revealed by specimens examined, except that one record of argentescens is about 275 miles off the map at Mand, Baluchistan. Subspecies: A, pennanti; B, argentescens; C, lutescens. 74 FIELDIANA: ZOOLOGY, VOLUME 48 "Voice. — Whistle-like, shriller, sharper and more prolonged than that of palmarum. "Nest.— Builds nest under tiles and eaves of buildings and in hollow trunks of trees and among palm leaf sheaths. In the desert, owing to the absence of suitable nesting sites, it builds like palmarum, but the nest is globular and more disorderly. It will sometimes com- mandeer the nest of Uroloncha. Coconut fibre is in great demand when available — hence, its nest-building is destructive to mattresses placed out for airing. "Habit. — More terrestrial than palmarum. Perhaps more om- nivorous than palmarum." Funambulus pennanti pennanti Wroughton Funambulus pennanti Wroughton, 1905, Jour. Bombay Nat. Hist. Soc, 16, p. 411. Type.— BM No. 98.4.2.25, adult female from Mandvi Taluka, Bundha, Surat District, India, collected February 27, 1898, by R. C. Wroughton. Material examined, from Bombay. — Bombay (BM), three; Shen- durni, 900 feet (BM), one; Poona, 1900 feet, Deccan (BM), five; Karad, Satara District (BM), one; Nasith [Nasik] (BM), one; "Mehda," Satara District (BM), one; "Satapur " 20 feet, Dhranga- dhra ^BM), four; Dharwar, South Mahratta (BM), one; Bodbad [Bodvad], 1000 feet (BM), one; Parola, 880 feet (BM) two. Material examined, from Berar. — Akola (UMMZ), one. Material examined, from Central Provinces. — Balaghat District (CNHM), three; "PhichpalH," 1300 feet, Chanda (BM), two; "Thain," 1400 feet, Hoshangahad (BM), two; "Ganoor," 1000 feet, Nimar (BM), one; "Sakot," 1200 feet, Hoshangahad (BM), one; "Hewra," 1000 feet, Nimar (BM), two; Sohajpur, 1000 feet, Hoshan- gahad (BM), one; Chanda, 800 feet (BM), two; "Rithi, C. P." (AMNH), one; Khandwa (AMNH), two; Raipur (AMNH), five; Asirgarh, 1500 feet, Nimar (BM), one. Material examined, from Gujerat. — Danta, 1000 feet (BM), one. Material examined, from Kathiawar. — Junagadh [Junagarh], 480 feet (BM), two; Rajkot, 100 feet (BM), two; "Sadla," 10 feet, Bajana State (BM), two; Sehore [Sihor], 1600 feet, Bhaonagar Province (BM), four; Talala [Talaja], 200 feet, Junagarh State (BM), two; Van Kaneer [Vankaner], 500 feet (BM), three. MOORE AND TATE: DIURNAL SQUIRRELS 75 Material examined, from Sind, West Pakistan. — "Pano Agil," Upper Sind (BM), five; Shikarpore [Shikarpur] (BM), one; Sehwan (BM), one. Material examined, from Rajputana. — Mt. Abu, 4300 feet (BM), one; Sambhar (BM), one. Material examined, from Punjab. — "Azadpur," 550 feet, Delhi (BM), one; Lahore (UMMZ), one; Sirsa, Hissar (UMMZ), five; "Bhadwar," Kangra (UMMZ), three; "Khajjean" Kangra (UMMZ), two. Material examined, from United Provinces. — Pihbhit, 800 feet, Rohilkand (BM), two; "Dela," 1500 feet, Ramnagar, Kumaon (BM), one; "Dachawri," 2500 feet, Kumaon (BM), one; "Jerna," 1500 feet, Ramnagar (BM), one; Ramnagar, 1100 feet, Kumaon (BM), seven; SiwaUk Hills (BM), one; Etawah (UMMZ), two. Material examined, from GwaHor. — Bhind (BM), three; Corepura [Chorepura], 1100 feet (BM), two; "Ghatigoan," 900 feet (BM), four; Morar (BM), one. Material examined, from Nepal. — Tribeni [Ghat], Terai (BM), four; Hetora [Hataura] (AMNH), one; "Pili," Sipora or Sipna [Sipra] Valley (BM), one; "Bandasa," West Terai (BM), two. Material examined, from Bihar. — "Jagodih," 600 feet, Hadari- dagh [Hazaribagh] (BM), two; Daltonganj, 600 feet (BM), one. Material examined, from Bengal, East Pakistan. — Chandpara (AMNH), three; Salbani, 200 feet, Midnapore (BM), two; Sevoke (CNHM), one; Kharagpur (UMMZ), one. Material examined, from Cooch Behar. — Haldibari (MCZ), one. Original description. — ". . . body colouring is very much as in palmarum, comorinus, but along the outside edge of the 'saddle mark' on each side there is a supplementary pale stripe . . . bounded on the outer side by the general body colour . . . where it is commencing to pale down to the meeting line with the belly colour. . . . No band of short, rufous hairs along the midrib under the tail as there is in palmarum. ..." Funambulus pennanti lutescens (Wroughton) Funambulus lutescens Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, p. 429. Type.— BM No. 13.9.18.105, adult female from Deesa, 450 feet, Palanpur, Gugerat, India, collected April 29, 1913, by C. A. Crump. 76 FIELDIANA: ZOOLOGY, VOLUME 48 Original description. — ". . . rather smaller than true pennanti, about the size of argentescens, but much paler than either of these two forms. . . . General colour above pale 'cream buff' on the flanks, the saddle near 'mars brown' . . . the five longitudinal stripes white, very slightly tinged with buff, the lateral pair scarcely distinguish- able. . . . Face drab. Hands and feet buffy white. . . . Below white, the hairs white to their bases. Tail also pale buffy white, each hair with two black rings. . . . "This form ranges south as far as the northern part of Kathiawar, but thereafter the specimens grow darker . . . and show a passage to true pennanti." In lutescens the lateral stripe runs undiminished from the ear to the hind limb; its width 6 mm. Funambulus pennanti argentescens Wroughton Funambulus pennanti argentescens Wroughton, 1905, Jour. Bombay Nat. Hist. Soc, 16, p. 412. Type.— BM No. 5.4.2.3, adult male from Rawalpindi, 1670 feet, Punjab, Pakistan, collected December 10, 1900, by Birrell. Material examined.- — Mand, Baluchistan (BM), one; Gajar, 3200 feet, Mashkai, Kalat (BM), two; Bohara [Bahara], Sind (BM), one; Garo [Gharo], Sind (BM), one; Jacobabad, Sind (BM), two; Kash- mor, North Sind (BM), two; Mirpur Sakro, Sind (BM), three; Jhim- pir Lake, Sind (UMMZ), two; "Khinjar Lake," Sind (UMMZ), three; Karachi, Sind (AMNH), one; Kohat, 1700 feet. North West Frontier Province (BM), three; Peshawar, 1700 feet (BM), two; Amballa [Ambala], Punjab (MCZ), six, (BM), one; "Ara," 2100 feet. Salt Range (BM), two; "Choan," 2400 feet. Salt Range (BM), one; Kallak Kahar [Kallar Kahar], 2113 feet. Salt Range (BM), two; Multan, 600 feet (BM), four; Rawal Pindi [Rawalpindi], 1671 feet (BM), 10; "Kotla," Kangra District (AMNH), six; Sodhi, 2000 feet. Salt Range (BM), one; Gholam (BM), one; Turbat, 600 feet, Kech (BM), three; Malasa, United Provinces (AMNH), one; "Lar- kanat," Naundero [Nandero] (BM), two; Kooloo Valley [Kulu Val- ley] (MCZ), two; Chak-Lala [Chakwal] 1100 feet, Rawalpindi, N. W. Punjab (CNHM), two. Original description. — ". . . Pattern . . . identical . . . with . . . pennanti . . . much paler, however, and almost all rufous tint has dis- appeared, . . . body colour is a pale French Gray . . . stripes and belly bright white. . . ." MOORE AND TATE: DIURNAL SQUIRRELS 77 The pelage is very short. The three white dorsal stripes are each 5 mm. wide. Dorsal pelage is dark reddish brown, grizzled, becom- ing pale gray on the shoulders, neck, and head. The limbs are pale gray. Ventral pelage is white the full length of the hairs. This subspecies is distinguished from F. pennanti pennanti by whiter tail, paler feet (Pale Olive Buff), and paler body color. That the races of F. pennanti can apparently be distinguished by the num- ber of black bands on the tail hairs was noted too late for testing on all the material examined, but we find that it separates the above five lots of p. pennanti from the two of p. argentescens of the AMNH material; two bands in p. pennanti, one in p. argentescens. Funambulus palmarum (Linnaeus) Definition. — This species includes those three-striped squirrels of the Indian Subregion, distributions of which are shown by collecting localities for the specimens we have examined, in Figure 8. (See also the synonyms listed in the subspecies accounts.) Diagnosis. — (1) The baculum is distally bifurcated or makes a right angle turn near the distal end, but the apex or apices are not knobbed. (2) The dorsal pelage is marked by three very distinct longitudinal hght stripes. (3) The occipitonasal length is usually less than 40 mm. (4) Measurement from front of incisor to back of palate is usually less than half the occipitonasal length. (5) The pel- age is relatively short and harsh. (6) The midstripe and ventral pelage are no more richly colored than pale buffy. The above characters distinguish palmarum from the other con- generic species as follows: pennanti by 1 and 2; tristriatus by 3 and 4; layardi by 6; and sublineatus by 1, 2, and 5. Relationship to other species. — Mr. Charles McCann of the Do- mion Museum, Wellington, N. Z., contributes the following remarks (letter of June, 1961) on F. palmarum from his years with the Bom- bay Natural History Society in India: "Habitat. — More a hill forest species than is pennanti, inhabiting deciduous rain forest. Ascends to about 4,000-4,500 ft. In such localities it will approach human environments, taking the place of pennanti. On the Panchgani-Mahableshwar plateau, 4100 ft., (out- skirts of the Western Ghats), palmarum occurs on the hills, but is replaced by pennanti on the Deccan Plain (= 1800 ft.). F. palmarum occurs almost throughout the Western Ghats in deciduous rain forest but does not enter the wet evergreen forest. INDIA • D • r • • J r • • C <; ^ \ • \ 20°N + 12' ( /^ -+7 I y"^ +18° N / 86° 72° V , • S • ,•■■■" + ^ 1 • \ +/4°N \ 82» \ • • •/ I2°N + 74° \ • • / / + + +IO°N BS^ 82° 8°N+ ^^ \ 76° 78° \ ^ ^ \CEYLON 0 1 1 MILES 400 1 < 1 82° Fig. 8. Species range of the Indian palm squirrel, Funambulus palmarum, as shown by specimens examined. Subspecies: A, palmarum; B, bellaricus; C, robert- soni; D, bengalensis; E, comorinus; F, favonicus; G, kelaarti; H, brodiei, and I, olympitis. The combined ranges of species palmarum and pennanti approximate the extent of the Indian Subregion. 78 MOORE AND TATE: DIURNAL SQUIRRELS 79 "Coat. — Soft and somewhat silky and in life more erect than that of pennanti. "Voice. — More bird -like and slightly deeper than that of pennanti. "Nest. — Builds nest like a passerine bird in branches, somewhat globular. "Habit. — More arboreal but does come down to the ground. Nec- tar is one of its foods. Bark searching is a frequent habit, possibly for insects. Fruit appears to be its main diet." Funambulus palmarum palmarum (Linnaeus) Sciurus palmarum Linnaeus, 1766, Systema Naturae, 12th ed., 1, p. 86. Sciurtis penicillatus Leach, 1814, Zool. Misc., 1, p. 6, pi. 1. Sciurus indicus Lesson, 1835, Illustr. de Zoologie, no. 15, pi. 43, 2 pp. text. Funambulus gossei Wroughton and Davidson, 1919, Jour. Bombay Nat. Hist. Soc, 26, p. 730. Types. — Sciurus palmarum, apparently lost, not found at Upsala, locality restricted to vicinity of Madras, India (Wroughton, 1905, p. 410); penicillatus, not found; indicus, not found; gossei (BM) No. 19.6.2.30, an adult male from Kotagiri, 4100-4500 feet, Nilgiri Hills, taken June 20, 1918, by PhiHp Gosse. Material examined, from Madras Province. — "Adyar" [in city of Madras] (BM), two; "Kilcauk" [Kilpauk in city of Madras] (BM), two; "Museum Grounds" [city of] Madras (BM), two; "Panampet" (BM), one; "Tadiarpet" (BM), one; Trichinopoli, 400 feet (BM), two; Chittoor (AMNH), one; Salem District (AMNH), two. Material examined, from other provinces. — Coimbatore (BM), one, (CNHM), one; Polkonda Hills, 1000 feet, S. Cuddapah (BM), two; Shevaroy Hills, 4500 feet, E. Ghats (BM), one; "Sweat Dist." (BM), one; "Machen," 4000 feet (BM), one; Kanara at S. Mahratta border, 2000 feet (BM), one; Seringapatam, 2338 feet, S. Mysore (BM), three; Kolar, 2786-4026 feet, E. Mysore (BM), two; Kurum- dapatti, Salem Distr. (BM), three; Kalhatti [Kalpatta], Nilgiri Hills (AMNH), one; "Kellengode" (AMNH), one; "Honnametti Estate," Biligirirangan Hills, Mysore (AMNH), one; Anaikatti, Nilgiri Hills (AMNH), one; "Gantha," 1300 feet, Coimbatore, southern India (BM), one; "Rookery Kil," 4500 feet, Kotagiri, Nilgiris (BM), three. Although gossei was originally described as belonging to the "tri- striatus group," Ellerman (1940, p. 379) held it a separate species by itself, and Zahn (1942, p. 58) placed it in the species palmarum. 80 FIELDIANA: ZOOLOGY, VOLUME 48 Comparison of the dimensions of the type of gossei in our Table 3 with those of species palmarum and species tristriatus supports Zahn's allocation. Ellerman and Morrison-Scott (1951, p. 494) synonymize gossei under F. p. palmarum, and this seems to us the most reason- able treatment. The color of F. p. palmarum, as evidenced by the above series of three from the Salem District and Chittoor near the type locality: A middorsal area Mummy Brown to Prout's Brown pales anteriorly behind the shoulders and posteriorly on the rump. Through this run three longitudinal lines of Light Buff to Light Ochraceous Buff about 3 to 4 mm. wide. The middle one of these extends from the nape (into which it may extend faintly) to the base of the tail. The lateral ones separated by 11 to 12 mm. from the middle stripe, extend from back of the shoulders onto the rump, failing to match the length of the middle stripe by about 10 mm. anteriorly and perhaps 3 to 5 mm. posteriorly. Five to 7 mm. below the lateral lines, the Mummy Brown dorsal area changes abruptly to the agouti sides, which are Smoke Gray to Apricot Buff. The color of the sides ex- tends onto the shoulders, nape, cheeks, ears, and rostrum, and onto the flanks and all limbs and feet. The crown color is intermediate between that of the sides and back. There is an almost white area about 2 by 10 mm. just medial to each ear, oriented anteroposteri- orly, and fading into the body color posteriorly. There is a light- colored "eye-ring" 1 mm. wide above and below the eye, but not enclosing it at either end. The tail is somewhat less than half the animal's total length, about 20 mm. wide. Its dorsal and lateral hairs are whitish at the base, middle, and tip, with two black bands be- tween and these give the tail as a whole an appearance of jumbled annulation (with about 15 black rings in one instance). On the hairs of the lower surface of the tail the basal and middle portions of each hair are Cinnamon-Rufous proximally on the tail grading out to Apricot Buff at its end. This Cinnamon-Rufous color extends onto the ventral pelage of the body in a small area about the anus. The ventral pelage from this to the chin and on the under sides of all the legs is white with faint washings of Maize Yellow. Habits. — G. C. Shortridge remarks of Funambulus palmarum col- lected in the vicinity of Dharwar (in Wroughton, 1912, p. 1186), "They may often be found in prickly pear thickets when their mouths are usually stained crimson with the juice of the fruit. Both this species and tristriatus feed also on the berries of the Lantana." MOORE AND TATE: DIURNAL SQUIRRELS 81 Funambulus palmarum comorinus Wroughton Funambulus palmarum comorinus Wroughton, 1905, Jour. Bombay Nat. Hist. Sec, 16, p. 411. Type.— BM No. 95.10.9.10, adult female from Katyani, Trevan- drum, Travancore, India, collected January 23, 1895, by H. Ferguson. Material examined. — Nagercoil (BM); one, "Benhope," 3000- 4000 feet, Nilgiris (BM), three; Kotagiri, Nilgiris (BM), two; Tre- vandrum, Trevancore [Travancore] (BM), 11. Original description. — "Differs from [palmarum] by its much so- berer grey colouration and larger skull measurements." This race, generally much darker in color than the true palma- rum, is evidently closely related to the dark races of Ceylon. The width of the median line is between 2 and 3 mm. The brown of the dorsal colored area between each lateral line is considerably narrower than in other races of palmarum. The hairs of the under parts are white or dirty white, with short gray bases. The anal area is brown, as in other Indian forms. Funambulus palmarum bellaricus Wroughton Funambulus palmarum bellaricus Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, p. 647. Type. — BM No. 13.4.10.39, adult male from Vizayanajar, Bellary, India, collected July 20, 1912, by G. C. Shortridge. Material examined. — Vizayanajar, 1500 feet, Bellary, India ''CN- HM), two. Original description. — ". . . differs [from palmarum] in the com- plete absence of yellow suffusion on the forearms, shoulders, and thighs. General colour above rather a coarse grizzle of black and white, giving the effect of pale 'smoke grey.' The whole dorsal area forming *a saddle,' which is coloured much darker. . . . The usual three longitudinal dorsal stripes creamy white. The face more or less suffused with ochraceous, according to season. Feet and hands pale. . . . Tail black and white, very indistinctly barred, and below 'orange rufous.' "Shortridge obtained ... 29 specimens in s. Mysore . . . consid- ered intermediates [between bellaricus and palmarum]." This subspecies is sharply separated from subspecies palmarum by the much greater width of the white median line, 6 mm. com- pared to 3. 82 FIELDIANA: ZOOLOGY, VOLUME 48 Funambulus palmarum robertsoni Wroughton Funambulus palmarum robertsoni Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, p. 647. Type.— BM No. 12.11.29.92, adult male from Pachmarhi, Ho- shangabad, India, collected March 20, 1912, by C. A. Crump. Material examined. — Pachmarhi, 3300 feet, Hoshangbad (CNHM), two; Asirgarh, 1500 feet, Nimar (BM), one; Kol Kaz, 1600 feet, Berar (BM), one; Rarighat, 2500 feet, Hoshangabad (BM), one; Siwal, 1000 feet, Nimar (BM), one; Pachmarhi, 3300 feet, Ho- shangabad (BM), one; "Thain," 2000 feet, Hoshangabad (BM), two; "Nawapara," Raipur, Central Provinces (AMNH), one. Original description. — ". . . Sombre coloured . . . markedly smaller than palmarum . . . saddle brown, with a slight yellow tinge . . . get- ting darker at certain seasons. The dorsal stripes buffy white. . . . Below, in most cases, 'vinaceous cinnamon,' in some dull white. . . ." This subspecies differs from others in having no brown on the head; the gray of the sides and nape continues forward onto the crown and face. The width of the median line is nearly five milli- meters. Funambulus palmarum bengalensis Wroughton Funambulus palmarum bengalensis Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, p. 648. Type.— BM No. 15.4.3.77, adult female from Gayhundi, 1000 feet, Hazaribagh, Bihar, India, collected May 10, 1914, by C. A. Crump. Material examined. — "Jagodih," 600 feet, Hazaribagh, Bihar and Orissa (BM), one; Muhammedganj, Behar (AMNH), one. Original description. — ". . . resembling robertsoni in its small size, but distinguishable by its much larger teeth and ... an ochraceous tinge on flank. Below invariably dull white." The width of the median line is about three millimeters. The pelage of the example from Muhammedganj differs from our description of the typical race: in having a Blackish Brown (2) dor- sal area; in having the stripes extend, albeit faintly, upon the shoul- ders to the nape; in having the mid-stripe but half the width of the laterals; in having the Cinnamon -Rufous of the anal area extend out the ventral pelage of the tail about 20 mm. before paling to Light Ochraceous Buff, then Light Buff; and in having a fuller tail (30 mm. wide with hairs to 25 mm. long). MOORE AND TATE: DIURNAL SQUIRRELS 83 Habits. — A field observer collecting this form makes the follow- ing remarks: "Blanford states this squirrel is not found in forests, but in my experience though partial to the neighbourhood of culti- vation, it may like pennanti be found far into the forests." (C. A. Crump in Wroughton, 1915c, p. 108.) Funambulus palmarum favonicus Thomas and Wroughton Funambulus palmarum favonicus Thomas and Wroughton, 1915, Jour. Bom- bay Nat. Hist. Soc, 24, p. 39. Funambulus palmxirum matugamensis Lindsay, 1926, Jour. Bombay Nat. Hist. Soc, 31, p. 239. Types. — Funambulus p. favonicus, BM No. 15.7.1.2, young (orig- inal description says "adult") female from Udugama, Southern Province, Ceylon, collected April 23, 1913, by E. W. Mayor; F. p. matugamensis, BM No. 27.11.17.1, adult female from Anisigalla, Matugama, 30 miles southeast of Columbo, Ceylon. Material examined. — "Anesagalla" [Anisigalla] 100 feet, Matu- gama, Kalutara, Western Province (BM), 18; "St. George," 300 feet, Matugama, Western Province (BM), one; Kottawa, Southern Province (BM), two; Ranna, Southern Province (BM), one; Udu- gama, Southern Province (BM), six (topotypes). Original description. — ". . . darker than true palmarum, owing to the greater amount of black grizzling. The saddle much darker. . . . The central dorsal stripe white, the lateral ones ochraceous buff. Below white. Tail below 'cinnamon rufous' . . . i.e. red rather than yellow. Feet a fine grizzle of black and buff. . . . When these [18] specimens were laid out in a row their warmer colouring, strong col- our contrasts, and grizzled feet differentiate them from the sober, dull-coloured palmarum palmarum." The top of the face and rostrum of the type is a rather bright reddish brown. The under parts are dull whitish with gray hair bases. The under side of the tail is yellowish-brown. Dimensions. — Phillips (1935, p. 231) provides averages of mea- urements in millimeters for 30 males and 30 females of favonicus. These are respectively: length of head and body, 144.7 and 142; length of tail, 124.4 and 137; length of hind foot, 34.5 and 34.5; length of ear, 15.2 and 16. He comments that many had the ends of their tails "damaged," especially the males. We take this to mean that the tails of many were short because the end was missing. He also gives maxima for these measurements. These are, for males and females respectively: 180 and 150, 157 and 169, 37 and 37, 17 and 17. 84 FIELDIANA: ZOOLOGY, VOLUME 48 Funambulus palmarum kelaarti (Blyth) Sciurus kelaarti Blyth, 1851, Jour. Asiatic Soc. Bengal, 20, p. 166. Type. — IM 9479, a parous female from Hambantotte, Ceylon, collected in 1845 by E. L. Layard. Material examined, all from Ceylon. — Aripo [Arippu], northwest Ceylon^ (BM), two; A'pura [Anuradhapura], North Central Prov- ince (BM), one; Kala-oya [a river]. North Western Province (BM), nine; Putlam [Puttalam], North Western Province (BM), one; Man- keni, Eastern Province (BM), two; Inganiyagala, 100 feet, near Kal- munai, Eastern Province (BM), five; Hambanpota [Hambantota], Southern Province (BM), 11; Wellawaya, 608 feet, Uva Province (BM), two; "Micouralia" (BM), one. Original description. — ". . . kelaarti entirely replaces all the other small Sciuri from Tangalle and Hambantotte, and I fancy extends round to Trincomali. ... It [is] . . . like . . . palmarum of India, but the head is much redder, the halves of the back and belly are more blended, and the animal is altogether smaller. . . ." Previously, Blyth (1849, p. 602) had written substantially the above in a footnote, without, however, giving the squirrel in ques- tion a name. Dimensions. — Phillips (1935, p. 229) presents averages of meas- urements in millimeters taken on eight males and six females of kelaarti. These are respectively: length of head and body, 143 and 139.6; length of tail, 136.7 and 106; length of hind foot, 34.5 and 35.5; length of ear, 18 and 17.5. (Note that Phillips' average measure- ments of the tails of favonicus evidently included ones with part of their natural length missing.) He also gave maxima for these dimen- sions, and these are respectively: 149 and 151, 151 and 160, 37 and 37, 20 and 19. Type description.- — Color characteristics of the type of kelaarti, examined in 1960: The pelage of the crown is Russet (XV) and ends abruptly along a line between the ears. The agouti pelage of the nape and shoulders is close to Dresden Brown, or, if one ignore the yellowing effect of the light bands, Saccardo's Umber (XXIV). The postauricular patch is Pinkish Buff. The eye ring and sides of face below and in front of the eye are Light Ochraceous Buff (XV) . The pelage color approaches Ochraceous Tawny between the eye and ear. ' There is another Arippu in Eastern Province (latitude 08° 19' N., longitude 81° 20' E.) which may better be the source of this material. So treated on dis- tribution map. MOORE AND TATE: DIURNAL SQUIRRELS 85 Dark pelage on the back between the Unes is Mummy Brown at its greatest intensity at the middle of its length, gradually grading into the shoulder color anteriorly, but going through the redder Prouts Brown and then Mars Brown before grading into the Dresden Brown of the rump and thighs. The dark saddle area outside of the light lines borders abruptly on the sides without diminution of intensity. The sides are about Isabella Color (XXX). The dorsal pelage on the limbs is like that of the shoulders but paler. The three light lines on the back are Light Ochraceous Buff, the lateral ones reach five millimeters wide, the middle one only two. The light lines disappear in the shoulder and rump areas but are intense for 70 or 80 milli- meters. The ventral pelage is Naples Yellow (XVI) and borders on the color of the sides rather abruptly. The midvane of the under side of the tail is Tawny (XV). The tail hairs have three blackish bands, the proximal two each about 2 mm. long, the distal one about 4 mm. The tail pelage is worn and a little faded but a black pencil is evident at the tip, and the hairs are about 30 mm. long. Funambulus palmarum brodiei (Blyth) Sciurus brodiei Blyth, 1849, Jour. Asiatic Soc. Bengal, 18, p. 602. Funambulus palmarum brodiei, Thomas and Wroughton, 1915, Jour. Bombay Nat. Hist. Soc, 24, p. 40. Type. — IM No. 9840, collected by E. L. Layard in Ceylon. Material examined. — Cheddikakulam [Cheddikulam], Northern Province, Ceylon (BM), nine, (CNHM), two. Original description. — "Very similar to [tristriatus] but distin- guished by its considerably paler colour, and especially by having a very long pencil tuft {S}/2 in.) at the extremity of tail, quite differ- ent from what is ever seen in tristriatus . . . 'confined to Palmyra-tree district from Puttulam to Jaffna. How much further round the coast I do not know." Type description. — The type of Funambulus brodiei has the gen- eral appearance of palmarum. The three dorsal stripes, however, pass through a dark saddle-like area on the back, about 60 mm. long and 40 mm. in overall width. The width of the dark pelage between median and lateral pale lines is 9 mm., its width beyond the lateral lines about 6 mm. The pale median stripe is 4 mm. wide, the lat- eral ones 6 mm. The dorsal color becomes russet on the head and rostrum. The tail is a grizzle of black and buff. The hands and feet are light grayish buff. The under parts to the chin, and up the sides 86 FIELDIANA: ZOOLOGY, VOLUME 48 of the head to the bases of the ears, also the insides of the Hmbs, are dull buffy yellow. The skull of the type is very badly broken and the nasals missing. The teeth show a moderate degree of wear, but the almost unworn condition of the fourth premolar and first molar suggests that the animal is a young adult. The maxilla spreads over a considerable area on the dorsal surface of the skull. Dimensions. — Phillips (1935, p. 227) records averages of measure- ments made in millimeters on but four males and three females of brodiei, and these are respectively: length of head and body, 150 and 148.7; length of tail, 148 and 139; length of hind foot, 35 and 34; and length of ear, 17 and 14.3. The maxima that he provides with the above dimensions are respectively: 162 and 153, 148 and 146, 37 and 34, 17 and 15. Funambulus palmarum olympius Thomas and Wroughton Funambulus palmarum olympius Thomas and Wroughton, 1915, Jour. Bom- bay Nat. Hist. Soc, 24, p. 41. Type.— BM No. 15.7.1.3, adult female from Urugalla, 1600 feet central highlands, Ceylon, collected February 25, 1914, by E. W. Mayor. Material examined. — Kandy (BM), one; "Lindoehn," 4200 feet (BM), one; Perodeniya [Peradeniya], 1600 feet (BM), 11; "Urugala," 1600 feet, Central Province (BM), nine (topotypes), (CNHM), two. Original description. — "A dark highland form. ResembHng bro- diei but much darker. The saddle is commonly almost black and the central dorsal stripe paler than the lateral ones, often white. Lower side of the midrib of tail is much darker chestnut than in brodiei. Feet are darker even than in favonicus. Below dull white." This is quite the darkest of the races of palmarum; the general dorsal color is blackish gray. The face is brownish-gray — about like the sides of the body. The under parts are dull whitish with gray hair bases. The under side of the tail is yellow-brown. Dimensions. — Phillips (1935, p. 234) provides averages and max- ima of measurements taken in millimeters on 13 males and 11 females, and the averages are respectively: length of head and body, 157.5 and 156.5; length of tail, 150.5 and 135.5; length of hind foot, 38.5 and 37.9; length of ear, 17.5 and 17.5. The maxima are: 166 and 164, length of head and body, 168 and 167, length of tail, 40 and 40, length of hind foot, and 20 and 20 length of ear. MOORE AND TATE: DIURNAL SQUIRRELS 87 Funambulus tristriatus (Waterhouse) Definition. — This species is constituted by those large, dark, three- striped forms inhabiting the evergreen rainforest of the Western Ghats of peninsular India as shown in our map Figure 9. Diagnosis. — (1) Funambulus tristriatus has three light longitudi- nal lines in the dorsal pelage. (2) The occipitonasal length of skull in adults with tooth wear exceeds 40 mm. The above characters distinguish species tristriatus from the con- generic species as follows: pennanti by 1 ; palmarum by 2; layardi by 2; and suhlineatus by 2. Systematic history. — The distinctness of Funambulus tristriatus from palmarum was recognized first by Waterhouse, and subse- quently by Wroughton (1905, p. 410), as "the forest form of palma- rum." Later Wroughton (1916d) distinguished the two species by the larger skulls of tristriatus, and in the same article described sev- eral new races. Later Wroughton and Davidson (1919) proposed two additional forms, one of which extended the range of the species northward along the west coast to Bombay. Pocock (1923) then (in error as shown above) distinguished this tristriatus group generically as Tamiodes. Relationships to other species. — Funambulus tristriatus is consid- erably larger than either palmarum or pennanti (see Table 3) and is much darker, except for the dark race of palmarum on Ceylon. The tail is much fuller — its width across the pelage about 25 mm. The longitudinal pale lines are only three. The median line is weak, dull yellow and 3 mm. wide; the two lateral lines are dull white, and about 4 mm. wide. The under parts are dull buffy white, with dis- tinctly gray bases to the hairs. The scrotal area and under side of tail (bases of tail hairs) are russet. This purely tropical species occurs in the forests of the mountains along the western coast of the Indian peninsula from about latitude 19° N., or the city of Bombay, southward well into Travancore. Study to define further the distribution of subspecies tristriatus and wroughtoni in relation to each other and to species palmarum in Coorg, Mysore, Malabar, and the Nilgiris would be rewarding. Funambulus tristriatus tristriatus (Waterhouse) Sciurus tristriatus Waterhouse, 1837, Mag. Nat. Hist., (new ser.), 1, p. 499. Sciurus dussumieri Milne-Edwards, 1867, Rev. Mag. Zool., 19, p. 226. Funambulus tristriatus annandalei Robinson, 1917, Rec. Indian Mus., 13, p. 41. Fig. 9, Ranges of the geographic races of the Western Ghats squirrel, Funam- bulus tristriatus, as shown by material examined. Subspecies: A, tristriatus; B, wroughtoni; C, numaris. 88 MOORE AND TATE: DIURNAL SQUIRRELS 89 Types. — Sciurus tristriatus, BM No. 55.12.24.112, adult male from "India," here restricted to the Western Ghats south of 12° N. latitude; Sciurus du^sumieri, MNHN No. 1838 (292), adult from Malabar; Funambulus t. annandalei (not seen), IM No. 8498, adult female, from Sasthancotta, west side of Western Ghats, Travancore, collected November 8, 1908, by N. Annandale. Material examined. — "Paumpa," North Travancore (BM), three; "Merchiston," Travancore (BM), two; "Paibuna," North Travan- core (BM), one; "Poothota," Vycoona District, North Travancore (BM), three; "Travancore," 4000 feet (BM), six; "Tyebautuchary," North Travancore (BM), two; "Bonaccord," Glen Brith Estate (BM), one; "India" (MCZ), one; "Kellengode," South India (AMNH), two; Kuttyani [Kuttyadi, Malabar], 250 feet (BM), one; "Mella- cotta," Wynaad (BM), one; "Madras" (BM), five; "Poumodi" [Pon- mudi, Travancore], 2000 feet (BM), two. Width of the lateral line is 4 mm., of the median line 2 mm. in the type specimen. The type of dussumieri has the median pale line 45 mm. in length, and the lateral lines 70 mm. The widths are: median line, 2 mm.; laterals, between 3 and 4 mm. The under side of the tail is rather bright rufescent from the vent almost to the tip. Discussion. — Miss Ryley (1913, p. 437) remarked that the type of tristriatus "... very probably . . . came from Travancore as it agrees best with a small series from that district." Nevertheless, when Robinson (1917, p. 41) described annandalei from Travancore, citing this paper of Miss Ryley, he ignored the above-quoted obser- vation of hers and commented, "In default of authenticated skins from Madras I have taken modern skins from Kanara as typical of F. . . . tristriatus, Waterh., though it is by no means impossible that these will prove to represent yet another form." The Kanara series had in fact already been identified as belonging to a new subspecies of tristriatus described by Wroughton (1916, p. 646), who tersely re- stricted the type locality of tristriatus to Travancore. Wroughton and Davidson (1919, p. 728) then reported, "The British Museum has a series, sent by Capt. H. Ferguson, from Trevandrum [in Trav- ancore], which are undoubted tristriatus, with the type of which they agree in all essential particulars." Zahn (1942, p. 68) records the type locality of tristriatus as Travancore. EUerman (1940, p. 379) and Ellerman and Morrison-Scott (1951, p. 495) employ "Madras, India (by designation)." The use of "Madras" is unfortunate, for although the state of Madras formerly included what was still earlier Travancore, it does not now, and in any case Madras includes a 90 FIELDIANA: ZOOLOGY, VOLUME 48 much broader area than that in which anyone presumes subspecies tristriatus to occur. The correct political designation at the time of our writing is southern Kerala, but Western Ghats south of 12° north latitude is a better description, and in using this we are honoring and slightly refining Wroughton's (1916, p. 645) restriction of the type locality to Travancore. Funambulus tristriatus wroughtoni Ryley Funambulus wroughtoni Ryley, 1913, Jour. Bombay Nat. Hist. Soc, 22, p. 437. Types.— F. wroughtoni, BM No. 13.8.22.48, old female from Sri- mangala, 2782 feet. South Coorg, peninsular India, collected Febru- ary 6, 1913, by G. C. Shortridge. Material examined. — "Cotengody Estate," 3500 feet [Nelliam- pathi Hills], Cochin (BM), four; "Shernelly," 1500 feet. Cochin (BM), four; Mudumalai, 3285 feet, Nilgiri Hills (AMNH), one; "Dirajpet," 3000 feet. South Coorg (BM), three; "Makut," 250 feet. South Coorg (BM), two; "Srimangala," 2782 feet. South Coorg (BM), one; "Wotekotti," 2000 feet. South Coorg (BM), one, (CNHM), two; "Benhope," 3000-4000 feet, Nilgiri Hills (CNHM), two. Original description. — ". . . clearly an ally of [tristriatus] . . . gen- eral colour greyish brown . . . three pale yellow longitudinal stripes . . . the middle one being much shorter and narrower than the lateral ones. . . . Saddle rich chestnut, this being one of the most striking characteristics . . . distinguished by its large size, the conspicuous chestnut color of the dorsal fur, and the black and white appearance of its tail. "Examples from N. Kanara and Dharwar decidedly smaller, and have a different general colour and are nearer to the type of F. tri- striatus. "The series of 35 . . . from Coorg is very constant in colouring, only three . . . having any black on the back, whereas the type, four of the five from Travancore, and about half of the specimens from N. Kanara have the saddle black ..." The pale stripes extend from shoulders to rump; the median stripe is 2 mm. wide; lateral ones are 3 to 4 mm. wide. The one ex- ample in the collections of the American Museum of Natural History has three bands of black on the tail hairs. MOORE AND TATE: DIURNAL SQUIRRELS 91 Funatnbulus tristriatus numarius Wroughton Funambulus tristriattis numaritis Wroughton, 1916, Jour. Bombay Nat. Hist. Soc, 24, p. 646. Funambulus thomasi Wroughton and Davidson, 1919, Jour. Bombay Nat. Hist. Soc, 26, p. 729. Types. — Funambulus t. numarius, BM No. 15.7.3.26, adult male from Helwak, Western Ghats, Satara District, Bombay, India, taken December 7, 1914, by S. H. Prater; thomasi, BM No. 19.6.3.50, Khandala, 2000 feet. Western Ghats, Bombay, India, an adult female taken April 11, 1918, by Philip Gosse. Material examined. — "Potoli," 1800 feet. North Kanara (BM), four; Gersappa [Gersoppa], sea level, Kanara (BM), two; Helwak, Satara District (BM), eight, (CNHM), two (topotypes); "Hulekal," 1500 feet, near Sirsi, Kanara (BM), one; Khandala, 1800 feet (BM), two (topotypes); Khed, Ratnagiri District (BM), two; Sirsi, 1500 feet, Kanara (BM), one; Thana (BM), two; "Kardibetta Forest," 2500 feet, Shimoga District (BM), one; Sagar, 2500 feet, Shimoga District (BM), one; "Devikop," 2000 feet. South Mahratta (BM), one. Original description. — "A local race of tristriatus, slightly smaller. . . . General colour above a grizzle of black, and buff giving a general effect of yellowish 'drab' approaching 'isabella colour,' the saddle . . . darker . . . with three longitudinal pale buff lines, broader and better marked than in true tristriatus. Face coloured like the back with a yellow suffusion, cheeks buff. In the summer coat the . . . saddle becomes jet black, and the dorsal stripes tend to become white, while the . . . face becomes tawny and the cheeks more ochraceous. Below dull white except the anal region, which like the under side of the tail, is a bright 'cinnamon rufous'; above the tail hairs are black with white tips, arranged ... to indicate ... a barring . . . "Shortridge obtained 35 specimens in Dharwar and Kanara which are clearly intermediates between [numarius] and tristriatus. . . ." Funambulus layardi (Blyth) Definition. — This species is constituted by the largest and most colorful squirrels of the genus Funambulus on Ceylon, subspecies layardi and signatus, and in the jungles of Travancore by the sub- species dravidianus. See the species distribution in Figure 10. Diagnosis. — The ventral pelage is entirely orange chestnut to cinnamon or may have the thoracic part yellowish orange. 92 FIELDIANA: ZOOLOGY, VOLUME 48 INDIA CEYtON 8°N + 76° Fig. 10. Distribution of the Sinhalese jungle squirrel, Funambulus layardi, plotted from Ceylon localities recorded by Phillips (1935). Subspecies: A, layardi; B, signatus; C, dravidianus. Relationships to other species. — The species layardi seems to be the ecological equivalent on Ceylon of tristriatus in the Western Ghats. This is to say that species layardi is larger than forms of the species palmarum and sublineatus on Ceylon (see measurements given in pertinent subspecies accounts and in Table 3) and occupies prin- cipally the old evergreen rainforest. The fact that layardi seems also to occur in the southern Western Ghats, invites a comparative investigation of the ecology and distribution of these two species on MOORE AND TATE: DIURNAL SQUIRRELS 93 the mainland. Hutton (1949, p. 692) reports the species layardi very common in the High Wavy Mountains of the southwest corner of Madura District, Madras, on the Travancore border, and tristriatus seems to be absent from this locality. Funambulus layardi layardi (Blyth) Sciurus layardi Blyth, 1849, Jour. Asiatic Soc, Bengal, 18, p. 602. Type.—IM No. 9481 (ASB 341 A), skin only, sex unknown, taken in the "uplands" of Ceylon in 1843 by E. L. Layard. Material examined. — "Mousa Kaida" [Mousakande, Gammad- uwa, Ceylon], 3000 feet (BM), one, (NMC), one; "Thimbulketiya," Ceylon (NMC), one. Type description. — The dorsal color of the type is now (1951) grizzled dark grayish brown. Of the three pale dorsal lines the median one is bright orange-yellow and 2-3 mm. wide; the lateral lines are only slightly paler than the body color and 3 mm. wide. The areas between the median and lateral lines are blackish brown. A faint dark line appears outside each lateral line. The median line extends from nape to rump and is 110 mm. long, the laterals from shoulder to rump, 85 mm. long. The head is colored like the back. The tail is grizzled like the body, its hairs with yellow-buff bases, long black subterminal rings, each 11 mm. in length, and buffy white tips. The end of the tail is black. The under parts are russet, the hairs with gray bases. Dimensions. — Phillips (1935, p. 237) offers averages and maxima of measurements taken in millimeters on three males and six females of F. I. layardi. The averages are respectively: length of head and body, 165 and 155.5; length of tail, 144 and 142; length of hind foot, 37.5 and 37.5; length of ear, 16 and 15.5. The maxima are respec- tively: 168 and 160, 145 and 149, 39 and 39, and 16 and 16. Thomas (1924, p. 241) comments, ". . . the type of layardi came from the Ambigamoa Hills which are in the highlands of the Central Province (7°N.,80°30'E.) . . ." Funambulus layardi signatus Thomas Funambulus layardi signatus Thomas, 1924, Ann. Mag. Nat. Hist., (ser. 9), 13, p. 241. Type. — BM No. 15.3.1.73, young male from Rakwana, Ratna- pura District, 115 feet, Ceylon, collected February 17, 1914, by E. W. Mayor. 94 FIELDIANA: ZOOLOGY, VOLUME 48 Material examined. — Southeast of Ratnapura, Ceylon (BM), one. Original description. — "Similar in all respects to true layardi, but the light median dorsal line, instead of being narrow (about 3 mm.) and pale buffy, is broader (nearly 5 mm.) and of a rich ochraceous or flame-colour." Dimensions. — In Phillips (1935, p. 239) one finds averages of measurements taken in millimeters on eight males: length of head and body, 153.7; length of tail, 140.8; length of hind foot, 35.2; length of ear, 17. The maxima for the same series are respectively: 160, 153, 38, and 19. Funambulus layardi dravidianus Robinson Funambulus layardi dravidianus Robinson, 1917, Rec. Indian Mus., 13, p. 42. Type. — IM No, 10143 (=9773), young individual from western side of the Western Ghats, Travancore, India, taken by Nelson Annandale. The reported presence of deciduous premolars in this type is the basis, acceptable to us, for regarding it as immature (e.g., Moore, 1956, pp. 46, 47). Material examined.— "India" (MNHN), one. Original description. — "Differs from [layardi] in having the top of the head and cheeks rich rufous orange, and under surface yellowish orange instead of dull chestnut. Area between light lines on back, deep lustrous black." Type description. — The general color of the upper parts are as in F. I. layardi of Ceylon, but the head is much more reddish and griz- zled with black. The dorsal area between and outside the three light lines is much blacker. The three lines are colored alike, light orange- buff. The width of the median line is 2 mm., of the lateral lines, 3 mm. The hands, feet, and tail are as in layardi. The under parts differ sharply from the typical subspecies. The red of the crown be- comes almost clear chestnut on sides of the head and on either side of the throat. The throat and neck beneath are cinnamon, changing to yellow hairs (with gray bases) on the thorax. Behind this the pelage becomes dull brownish cinnamon (hairs with gray bases). The insides of the fore limbs are yellow-buff, of the hind limbs, brownish cinnamon. The under side of the base of the tail is brighter reddish cinnamon. Although we have been able to examine but two specimens of this form, Hutton (1949, p. 692) in reporting on it from the southwest corner of Madura District, Madras, on the Travancore border, re- MOORE AND TATE: DIURNAL SQUIRRELS 95 garded it, "A very common little animal found throughout the hills above 3,000 feet." Among other comments he adds, "They often forage on the ground for insects." This was apparently in continu- ous evergreen jungle. Funambulus sublineatus (Waterhouse) Definition. — This species is constituted by the most diminutive forms of the genus Funambulus, and these are all endemic to the southern tip of peninsular India and to Ceylon as shown in Figure 11. Diagnosis. — (1) The fur is long and soft, tending to obscure the three rather faint light stripes. (2) The baculum is bifurcated and the tips end in rather spherical nobs. (3) The supra-orbital notches are obsolescent or absent. Relationships to other species. — On Ceylon Phillips (1935, p. 240) states clearly that sublineatus "... is confined to the jungles of the hill and wet zones and is not found at all in the dry zone." And (p. 241) he says, "Its favorite haunt is a dense bamboo brake or patch of . . . Strobilanthes sp. and undergrowth in the jungle. ... It spends almost as much time upon the ground as in the undergrowth, and though it [is observed] to examine the rough bark of large trees, it is but rarely seen in their upper branches." The larger species, layardi, occurs in these same wet jungles of the same central hill zone and southwestern zone of Ceylon as sub- lineatus, according to Phillips (1935, pp. 237-239), in ". . . damp, heavy forests of a medium altitude, from about 1,000 ft. to 4,000 ft.; it is found only in the tallest trees ... it seems to spend much of the day among the foliage of the tops of the tallest trees. ... It is, however, in no way averse to descending to the ground and may fre- quently be seen examining the boles of the forest giants or making its way through the undergrowth." For other differences between these two species in Ceylon, and comment on it, see "Intrageneric Relationships" in the above account of the genus Funambulus. Funambulus sublineatus sublineatus (Waterhouse) Sciurus sublineatus Waterhouse, 1838, Proc. Zool. Soc. London, 1838, p. 19. Sciurus delesserti Gervais, 1841, L'Institut, p. 171. Sciurits trilineatus Blyth, 1849, Jour. Asiatic Soc. Bengal, 18, p. 602. Types. — Sciurus sublineatus, BM No. 55.12.24.321, young male from the Nilgiri Hills, Madras, India; delesserti, BM No. 217a, adult male from Nilgiri Hills, India; trilineatus, not found. 96 FIELDIANA: ZOOLOGY, VOLUME 48 Fig. 11. Geographic distribution of the dusky Indian jungle squirrel, Funam- bulus sublineatus, as indicated from material examined. Subspecies: A, subline- atus; B, obscurus. Material examined. — "Benhope," 3000-4000 feet, Nilgiri Hills (BM), one; "Bombay Shola," 7000 feet, Kodai Kanal [Kodaikanal], Palni Hills (BM), three; Coonoor, Nilgiri Hills (BM), two; "High Range," Trevandrum, Travancore (BM), one; "Madras" (BM), one; Manantoddy, Wynaad (BM), one; "Rookery Kil (Estate)," 3800 feet Kotagiri, Nilgiri Hills (BM), one; "Travancore," 4500-5000 feet (BM), two; "Huvinakadw Estate," 2843 feet, Kirtta, S. Coorg (BM), one; Kuttyani [Kuttyadi], 250 feet (BM), four; Ponmudi, 2000 feet MOORE AND TATE: DIURNAL SQUIRRELS 97 (BM), two; "Shernelly," 1500 feet, Cochin, S. India (BM), one; "Tiger Shola," 5500 feet (BM), one; "Cumbum," 5000 feet, High Wavy Mts., Madura (CNHM), two; Ootacamund, Nilgiri Hills (UMMZ), one. Original description (translation from the Latin). — ^"Above fusco- olivaceous, washed with yellowish; four black dorsal lines, three whit- ish, running from shoulders to rump; under parts yellowish; tail ringed black and yellow. "... four dark and three pale lines on the back: these lines . . . are very narrow . . . are not continued onto the shoulders, [nor] over the haunches. The general colour olive-brown . . . throat, chest and rump are whitish, . . . belly is yellow ..." Pelage color. — The CNHM study skins are an agouti of about Dresden Brown and are uniformly so from snout to ankles excepting for the three light stripes which are only somewhat paler. The four dark lines embrace the light ones and are slightly wider. The tails seem not at all distichous and taper to a point. Habits. — Hutton (1949, p. 692) reporting on this form in the southwest corner of Madura District, Madras, on the Travancore border, remarks, "It does not like thick forests and so is not often seen in our jungles [on a plateau above 4000 feet]. It is quite com- mon in the Varushnaad [Valley], and in light evergreen forest. It prefers country that has a light rainfall and [is] fairly sheltered. In the forests at the south of the Varushnaad [Valley], however, it is quite common up to 5,000 ft." Funambulus sublineatus obscurus (Pelzeln and Kohl) Sciurus palmarum var. obscura Pelzeln and Kohl, 1886, Verhandl. Zool. Bot. Gesell. Wien, 35, p. 525. Funambulus kathleenae Thomas and Wroughton, 1915, Jour. Bombay Nat. Hist. Soc, 24, p. 38. Types. — Sciurus p. obscura, not seen, from the "Mountains of Ceylon, 1000 meters"; kathleenae, BM No. 15.7.1.1, young male from Kottawa, Southern Province, Ceylon, collected April 11, 1913, by E. W. Mayor. Material examined, all from Ceylon.— Pattipola, 6210 feet. Cen- tral Province (BM), two; West Kaputale [Haputale], 6000 feet, Ohiya (NMC), one, (BM), three; Gammaduwa, 3400 feet, Mousa- kande (MCZ), one, (NMC), one. Original descriptions. — Sciurus p. obscurus, "From the mountains (1,000 meters elev.) ; the ones from the lowlands are of lighter color. 98 FIELDIANA: ZOOLOGY, VOLUME 48 The example under consideration is distinguished from the normal by the much darker coloring. The upper side is brown-gray; each hair shows a reddish yellow ring before its dark tip. The back with four dark chestnut-brown longitudinal stripes which are separated from one another by three much narrower pale yellow longitudinal stripes. The under side is lighter than the upper and shows more reddish-yellow." Funambulus kathleenae, "The readiest means of distinguishing kathleenae from sublineatus is the much greater width of the dark dorsal stripes, which are 7-8 mm. in breadth, as contrasted with 4-5." Habits. — This form is confined to Ceylon. According to Phillips (1935, p. 240), "... this little squirrel [occurs in] the jungles of the hills, from the highest peaks down to about 2,000 or 1,500 ft.; but ... in the southwest generally, it descends to the lower foothills . . . through the damp, hilly jungle of the low-country ... to near the sea coast in [the Southern Province]. It is confined to the jungles of the hill and wet-zones and is not found at all in the dry -zone." Dimensions. — For dimensions of this form Phillips (1935, p. 240) presents averages and maxima of measurements taken in millimeters on eight males and eight females. The averages are respectively: length of head and body, 118 and 112.8; length of tail, 109.5 and 103.5; length of hind foot, 28 and 30.3; length of ear, 14.4 and 14.5. The respective maxima are: 126 and 124, 117 and 113, 32 and 32, and 16 and 18. Maximum weight of either sex is 2}4 ounces. COMMON TREE SQUIRRELS Genus CALLOSCIURUS Gray, 1867 Callosciurus Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 277. Baginia Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 279. Erythrosciurus Gray, 1867, Ann. Mag. Nat. Hist., (ser. 3), 20, p. 285. Heterosciurus Trouessart, Le Naturaliste, 1 , p. 292. Tomeutes Thomas, 1915, Ann. Mag. Nat. Hist., (ser. 8), 15, p. 385. Type species. — Callosciurus, Sciurus rafflesi Vogors and Horsfield, the Sumatran form of C. prevosti Demarest from Malacca; Baginia, Sciurus notatus Boddaert from Java; Erythrosciurus, Sciurus ferru- gineus Cuvier from Burma; Heterosciurus, Sciurus erythraeus Pallas from Assam, India; Tomeutes, Sciurus lokroides Hodgson from Nepal a subspecies of C. pygerythrus Geoffroy from Burma. Definition. — Callosciurus is a genus of tree squirrels endemic to the Malaysian and Indochinese subregions, and is composed of the species erythraeus, ferrugineus, flavimanus, finlaysoni, caniceps, phay- rei, inornatus, and pygerythrus of the Indochinese Subregion (see their ranges in Figures 13-16) and extraterritorial species prevosti, notatus, nigrovitatus, albescens, and melanogaster of the Malaysian Subregion. Diagnosis. — Callosciurus possesses the following characteristics: (1) The least interorbital breadth exceeds the greatest length of nasal. (2) There are no pronounced longitudinal stripes on the back (although there may be on the sides and venter). (3) There is a single, unforked, bony septum across the chamber of the auditory bulla. (4) The third upper premolar is present. (5) The coronoid process of the mandible is high and falcate. (6) The upper edge of the infra-orbital foramen is well separated from the maxillo-premaxil- lary suture. (7) The baculum consists of two separate parts, a shaft and a blade. (8) Orbit length exceeds 13 mm. (9) The supra-orbital notches are obsolescent. Callosciurus is distinguished from the other genera of the Indian and Indochinese subregions by the above characters as follows: from Ratufa by 3, 4, 6, and 7; Funamhulus by 2, 5, and 7; Tamiops by 99 100 FIELDIANA: ZOOLOGY, VOLUME 48 2 and 8; Dremomys 1; Menetes by 1 and 2; and Sciurotamias by 1, 3, and 9. Intrageneric relationships. — Species of the genus Callosciurus are the common tree squirrels of the Indochinese and Malaysian sub- regions. No members of this genus cross the Garo-Rajmahal Gap into the Indian Subregion. Throughout a great deal of the Indo- Chinese Subregion there are two species of Callosciurus present in any one area. We recognize eight species of Callosciurus in the In- dochinese Subregion, and we consider them more closely related to each other than to any of the other species of Callosciurus, which occur only south of the Isthmus of Kra. The evidence so far ad- vanced (Moore, 1961a, p. 14) on which this monophyletally of the eight mainland species is inferred, is that the number of pairs of func- tional mammae is consistently only two; whereas the characteristic number of pairs in the several Malaysian species is three. This may be too weak a distinction to be regarded as subgeneric, but it does seem to distinguish what is a phylogenetic group higher than an ar- tenkreis and composed of two artenkreis. In further study this arrangement will probably be found to be supported in certain gen- eral characteristics of the pelage. For convenience until further study is accomplished, one of these two natural units may be called the mainland unit and the other the Sundaland unit of genus Callo- sciurus. There is some overlap in the distribution of these two units. The Malay Peninsula south of the Isthmus of Kra is occupied by two species of the mainland unit as well as three species of the Sunda- land unit. The mainland unit, we find, consists of one artenkreis of four allo- patric species spread over the whole Indochinese Subregion (eryth- raeus, ferrugineus, flavimanus, and finlaysoni) and a second artenkreis of four allopatric species {pygerythrus, phayrei, caniceps, and inor- natus) which is not so widespread but is on the whole sympatric with the first artenkreis. Although the species listed here bear names employed by Ellerman and Morrison-Scott (1951) and earlier auth- ors, the composition of our species usually differs markedly from earlier concepts. The reasons for the differences will be found in the following accounts. Descriptive skull characters. — (1) Least interorbital breadth ap- proximates orbitonasal length. (2) The occlusal plane is parallel to a plane connecting the tips of the upper incisors and the ventral extremities of the auditory bullae. (3) There is a subsquamosal foramen without vestige, or incipience, of a postglenoid foramen. MOORE AND TATE: DIURNAL SQUIRRELS 101 Fig. 12. Skull and left mandible of the Irrawaddy squirrel, Callosciurus pygerythrus, AMNH No. 163507, X 1. Note the indication of a single septum dividing the auditory bulla. (4) The frontals are level along the line of least interorbital breadth, but rise to slight inflations where they meet the premaxillaries. (5) The temporal ridges do not form sagittal crests (excepting in a small percent of adult melanogaster) . (6) The temporal foramen is usually fairly well developed. (7) The jugal is broad with a fairly prominent postorbital process. (8) The lip of the infra-orbital foramen in lateral aspect is ordinarily concave. (9) The upper incisors are proodont. Callosciurus erythraeus (Pallas) Definition. — Callosciurus erythraeus is a species occupying the area of the Indochinese subregion west of the Irrawaddy River and its tributary the Nmai. It consists of the subspecies erythraeus, erythrogaster, bhutanensis, intermedius, sladeni, hartoni, and haring- toni and the named forms here included in these subspecies. See the species map in Figure 13. Diagnosis. — In this extraordinarily variable species no pelage (or other) characters are known which distinguish all of its subspecies from all the subspecies of its nearest relatives. See the accounts of the species flavimanus and ferrugineus for a discussion of the evi- dence of relationships between erythraeus as here recognized and its closest relatives. Table 4 provides some measurements of body and skull for 17 types of named forms of this species. Relationships to other species. — ^We have been at great pains with Callosciurus in the Indochinese Subregion to ascertain where there is evidence of intergradation between forms and where in absence of intergradation there is also positive evidence for recognizing alio- as J <» yi Ui & s » s CG S § r'^ % S a CO ooOi-Hioioiraoo-^oot~eDi-Ht-Tticx)CD s bo 3e; ^ -ooo •oot-iO'-ioO'-io'-i • CQa> i-H •l-H.-l.-l -i-Hi-lTHi-li-Hi-Hi-lT-li-lT-t -u-tJ oiiot-t-toos'-Hi-iec'O-^eooot-oot^tra cu be w S gOJ (MrHC^ieOIMOtMCMi-IIMIMCOfMlMCCIMi-l -t^oj i-lT-lt-05CD^O(Mt-(MeO-^00 •CCOO-'^ O/O M .^ ^^^^^^^^^^^^^ .^^^ .^ O) tJjXi'S °0 -OCDO •'-l.-IC»'ni(M^t-OWC2'om CO •■^-^eo -coTjfc^iTfccuti-rjiuti-^io a> 8 ~-^ 5 03-tf„t- -C-Ol -OiOOOOOOCOt- -^ot- ^ -tJbf s L_iqj »o . .10110 -iciiousiousioio -utiuo • i. e -^^ -H •§ flO ec«ocoOTjooioo>ooTj-^THCOOOSOOOS©OOOOai0000050iOO>Osa>05050SO>050>00 +i.2 P3 -oot-t-t- •o^c^'-ioNoecc^iui -tomt-ooooooo bfl-S C^ O^ -OiOOO -0)0)00)000)000 -0)0)0)0)00)0) a>PQ 1— l»-l- i-Hi— ll— ItHi— I.— I- 1— I 4J?^ Cf3Tj-l,-|^i-Hr-t rH»-lTHi-(i-li-l»-l>-lrHi-l.-HT-li-(i-H.-li-HrH JOO(Mt>'-(MOi-IO)U3t-t- -t-»-l00«O-^Tj ^"oS* CO -^rHC^ieo -ooc^i-^c^coeoeocoeceo -0(Moj pq 3 CO w _. ^ ii 01 ^-u CO --^lit-OO) -(Ml0C0(MO'<*0)00NO •t-C> o "O -t-> fl o .5 o -co cs o "^ -H oo;oc^i-ieo«o-^(M.-(iC(MW«oi«'^«D":i(N«OTHO'-io -H-H -H-H O O CO lO w o o OJ t> 00 in O Tj" 00 (M .-H i-H rH CJ 00;DOO(M»-ITjooiMO(Mi-ieo(Meo-^-J(NNC "S - B 'C -« ^ » a>0 C» a C» CO s.2"S a -H g ^3 w fl GO (D Oi Eh H 124 MOORE AND TATE: DIURNAL SQUIRRELS 125 agouti than the general dorsal pelage. 3. The agouti guard hairs of the dorsal body pelage have from one to three light bands, gen- erally two, but on some individuals three is common. 4. The tail in ventral view appears scarcely more distichous than in dorsal. 5. There is no pronounced eye ring. The distinctions in color of the pelage between intermedius and quinquestriatus are: 1. In five of the six intermedius the venter is en- tirely red (not bisected by a band of agouti), but in quinquestriatus the most extreme condition of bisection in the whole subregion oc- curs. (The bisecting band is black instead of agouti, the divided ventral pelage is white instead of red, and the outer margins of the white are bordered with black.) 2. C e. intermedium has no black tip to the tail, but quinquestriatus generally has a large and conspic- uous black tip to the tail (which becomes subterminal in some speci- mens wherein the black terminal hairs are generously tipped with brown). 3. The hairs of the tail of intermedium have five blackish bands (additional to a short, black tip), but those of quinquestriatus have only four. 4. The ears of intermedium are no redder than the pelage of the middorsum, but in quinquestriatus the ears are redder than the middorsum (except in the very red specimens from low ele- vations across the Irrawaddy from Myitkyina). 5. The rostrum and the face below and slightly encircling the eye of intermedium is a noticeably cooler gray than the nape, but in quinquestriatus these parts are huffier and warmer in color than the nape (again excepting the very red lowland specimens). The above differences between these two forms are much greater than those between quinquestriatum and gordoni and even somewhat greater than those between quinquestriatus and shanicus, the next form south of gordoni. Since the differences between quinquestriatus and gordoni and between gordoni and shanicus are in both cases very great ones for subspecies in the subgenus Callosciurum of this sub- region, it appears that the differences between quinquestriatum and intermedium are too great and the similarities too general for one to presume their relationship to be subspecific. Since on the basis of degree of difference, forms intermedium and quinquestriatus evidently represent two species, one should note addi- tionally that a geographic intensification of differences between the two species progresses toward the probable point of most frequent contact. In the eastern species, for which flavimanum is the oldest acceptable name, the pelage character which may be called bisected venter is one of the most widespread in the Oriental Region (south 126 FIELDIANA: ZOOLOGY, VOLUME 48 for 1700 miles through six subspecies, east for 1500 miles through five subspecies). In gordoni (not far from the range of intermedius) it intensifies to its strongest agouti expression. In quinquestriatus, closer to the range of intermedius, it becomes stronger still by turn- ing to black. Approaching from the south through subspecies ruhec- ulus, pranis, siamensis, atrodorsalis, thai, zimmeensis and gordoni, or from the east through thaiwanensis, castaneoventris, hendeei, and michianus one finds the rich red venter (divided or undivided) . This is similar to the venter of C. erythraeus intermedius (and all but one form of erythraeus), but upon closest approach to the probable point of latest, or most frequent species contact, in C. flavimanus quinque- striatus the red venter is lost and replaced with black and white stripes. This provides extreme contast with the bright red venter of intermedius. Approaching from the west, in C e. hhutanensis one finds the black tail tip, reddish face, reddish ear tips, and divided venter that occur in eastern subspecies of the species flavimanus. Moving closer to the probable point of most frequent, or recent, con- tact between the species, one finds all of these characteristics gone or greatly reduced in intermedius. The above phenomenon appears to be very similar to what has been called "character displacement" (Brown and Wilson, 1956), differing from their definition of character displacement in that there is now no known zone of overlap of ranges of the two species. It is apparent on examination of the range map (fig. 16) of Cal- losciurus pygerythrus that the former river course of the Irrawaddy- Sittang must have been the eastern barrier to spread by that species, and for the most part, still is today. The Nmai Hka branch of the upper Irrawaddy apparently still is the barrier to eastward spread of pygerythrus in the region where the species pygerythrus is sympatric with C. e. intermedius. Although the Nmai is not shown so large on maps as the Salween, Mekong, or Brahmaputra, it seems likely on the basis of squirrel distribution that it was larger in the past and sub- sequently lost some of its headwaters by stream piracy. Such a former condition would better explain why the Irrawaddy shows so much evidence of having provided the greatest barrier to east and west spread of tree squirrels in the Indochinese Subregion. Callosciurus flavimanus quinquestriatus (Anderson) Sciurus quinquestriatus Anderson, 1871, Proc. Zool. Soc. London, 1871, p. 142. Sciurus beebei J. A. Allen, 1911, Bull. Amer. Mus. Nat. Hist., 30, p. 338. MOORE AND TATE: DIURNAL SQUIRRELS 127 Callosciurus quinquestriatus imarius Thomas, 1926, Ann. Mag. Nat. Hist. (ser. 9), 17, p. 640. Callosciurus quinquestriatus Sylvester Thomas, 1926, Ann. Mag. Nat. Hist. (ser. 9), 17, p. 641. Types. — Sciurus quinquestriatus, IM No. 10163, specimen from Ponsee, east of Bhamo, Burma, in the Kakhyen Hills, taken March 4, 1868, by John Anderson; beebei, AMNH No. 32624, adult male, originally described in error as from Sarawak, but fixed by Goodwin (1953, p. 282) as Sansi Gorge, Burma-Yunnan boundary, Decem- ber 7, 1910, collected by C. William Beebe; imarius, BM No. 20.8.- 7.6, old male, from west flank of Imaw Bum, Kachin, Burma, lati- tude 26°10'N., longitude 98°30'E., October 21, 1919, by F. K. Ward; Sylvester, BM No. 25.10.5.31, adult male (skull lost), from the forests of the Shweli-Salween divide, 9000 feet, west Yunnan, China, collected September, 1925, by G. Forrest. Material examined, from northeastern Burma. — 15 miles north of Myitkyina on east side of Irrawaddy River (USNM), eight; six miles north of Myitkyina (USNM), one; Porg-ka-tong (AMNH), one; Pum-ka-tong (AMNH), one; between Nomoyoung and Seniku (AMNH), one; between Tamu and Chipwi (AMNH), three; be- tween Chipwi and Laukhauna (AMNH), two; between Tamu and Tanga (AMNH), two; Laukhaung (AMNH), four; Pyepat (AMNH), three; between Pyepat and Langyang (AMNH), two; above Tsonma, 8300 feet (AMNH), one; Gangfang, 5200 feet (AMNH), one; Hkam- kawn, 4000 feet (AMNH), one; Hpimaw, 7600 feet (AMNH), two; Htawgaw (AMNH), one; Sima, Myitkyina, 4500 feet (BM), three; Sui Kin, Bhamo (BM), three; Teinzo, northeast of Bhamo (BM), two. Pelage color. — Description of Indian Museum Paratype, a stuffed study skin examined in January of 1960: Dorsal pelage agouti of about Cinnamon Brown (XV) on crown, nape and back, but paling on tail and sides to Dresden Brown. Ears in part redder, about Ochraceous Tawny. Feet like back but digits blacker. Tail hairs with blackish tips plus 4 blackish bands, but tip of tail light, about Buckthorn Brown. Agouti dorsal pelage has two distinct light bands and black on the tips and between light bands. Ventral pelage has two longitudinal light stripes. Light Ochraceous Buff from posterior end of throat to anus, separated and bordered by blackish bands (thus making two "white" and 3 "black" stripes) of the same length. The light bands are about 10 mm. wide on the thorax but 5 mm. or less for most of their length. The blackish middle stripe is similarly 128 FIELDIANA: ZOOLOGY, VOLUME 48 wide but the two lateral ones seem narrower. The middle stripe is definitely agouti on the thorax. The "black" stripes seem faded and are only about Prouts Brown. The hind legs are agouti all the way around, definitely, but blackish pelage apparently extends out the ventral surfaces of the fore limbs. The tail has a subterminal black tip. In the original description Anderson recorded the light stripes as "pure white" and their laterally bordering stripes as black and this black as "expanding on the inside of the thighs." Probably during the intervening 90 years the white has become less white and the black less black, for the American Museum Natural History material agrees emphatically with Anderson's description in these three par- ticulars. Discussion. — At the very beginning of knowledge of this squirrel, Anderson (1871, p. 142) considered it most closely related to [Callo- sciurus flavimanus] gordoni and included the two kinds under one common name, "the belly-banded squirrels." Nevertheless, the uniqueness of the five vividly contrasting black and white longitu- dinal bands of quinquestriatus has evidently discouraged all subse- quent students down to the present from considering that these two squirrels may be conspecific. There is, however, evidence that their ventral markings possess a common pattern which suggests that the amount of genetic difference between quinquestriatus and gordoni is not very great. The following observations are the basis upon which quinquestri- atus and gordoni are here considered conspecific (see fig. 13). The gordoni specimen CNHM No. 32519 from 27 miles north of Bhamo has a blackish midventral stripe and incipient black showing in the edge of the agouti of the animal's side where the agouti meets the red of the venter. In gordoni specimen AMNH No. 43217 from Hui Yao, Yunnan, the edges of the 7 mm. wide midventral agouti stripe, as well as the ventral edges of the agouti sides of the animal, are blackened. AMNH No. 43217 represents no great departure from the characteristics of gordoni, but its midventral stripe is precisely like that of many of the American Museum specimens of quinquestri- atus. Even the ventral edges of the sides of AMNH No. 43217 would need to become but little blacker to equal the blackness of some examples of American Museum material of quinquestriatus. Especially significant among the large series of quinquestriatus is AMNH No. 114907 from between Tomu and Chipwi, Burma. This specimen although preponderantly quinquestriatus in character ap- MOORE AND TATE: DIURNAL SQUIRRELS 129 preaches the characteristics of gordoni in several ways: (1) The mid- ventral stripe is quite like that described for AMNH No. 43217, and the ventral edges of the sides are only a little blacker. (2) The sub- terminal black tip to the tail is short and rather subdued. (3) The two ventral stripes which are ordinarily white in quinquestriatiLs, are Apricot Buff (XIV). The ventral pelage of gordoni, thus, may be said to be three- striped, or to consist of a 6 or 7 mm. agouti longitudinal mid-stripe which separates the red of the remainder of the venter into two broader, rather stripe-like, sections. In quinquestriatus the midven- tral stripe is blackened on its edges or sometimes is entirely black. The outer edges of the remaining ventral pelage is black, making two lateral black stripes of a width approximating that of the middle stripe, but including also the entire ventral pelage of the limbs. The remaining two areas of ventral pelage, generally about ten to twelve millimeters wide, are white. Thus, there are characteristically two white stripes intersticed between three black ones. The dorsal pelage of these two forms differs in but small details. Dorsal pelage of both body and tail is an agouti varying about Buffy Brown (XL) in quinquestriatus but is generally less red in gordoni. The subterminal black tip of the tail is generally longer in quinque- striatus. No evidence of intergradation between C /. quinquestriatus and subspecies [C. erythraeus] intermedius or [C. flavimanus] michianus has reached our attention as such. Probably the Sal ween River is barrier enough to prevent quinquestriatus from intergrading with michianus (see fig. 13). It would seem surprising, however, if the Nmai Hka or its northeastern tributary, the Taron River, would long prevent C. e. intermedius from coming into contact with C. f. quin- questriatus. See discussion above in the account of the species flavi- manus. Callosciurus flavimanus gordoni (Anderson) Sciurus gordoni Anderson, 1871, Proc. Zool. Soc. London, 1871, p. 140. Type, here designated. — IM No. 9268 a parous female taken at Bhamo, upper Burma, on February 24, 1868, by John Anderson. This specimen is here selected from the two cotypes as the type of gordoni because it is adult (whereas the other is not) and is rather typical of the subspecies in ventral color (whereas the other is not) . Material examined, all from a small area of northeast Burma and adjacent Yunnan. — Homoshu Pass, 8000 feet (AMNH), one, 130 FIELDIANA: ZOOLOGY, VOLUME 48 (CNHM), one, (MCZ), one; Taipingpu, 7000 feet, Shweli River (AMNH), two, (CNHM), one, (MCZ), one; "Hui Yao," 5500 feet, Tengyueh (AMNH), one; Watien, Tengyueh (AMNH), two; "Yun- nan" (AMNH), one; Kyu Loi, S. Shan States (ANSP), one; Shweli- Salween Divide, 7000-10,000 feet (CNHM), two, (BM), twelve; Bhamo, Burma (USNM), one topotype; Momauk, nine miles east of Bhamo (USNM), one; Cuchi, Yunnan (USNM), one; "Linchi- apu," Yunnan (USNM), one; Shweli Valley, 9000 feet (BM), two, 8000 feet (BM), eight, 7000 feet (BM), one; hills northwest of Tengyueh, Yunnan (BM), three; "Watan," 5000 feet, Bhamo (BM), five; Machangkai, 6500 feet, Tengyueh (BM), one. Type description. — A description of the type from examination of it in 1959, follows: Ventral pelage Cinnamon Rufous (XIV), but divided all the way down the middle by an agouti stripe 3 mm. to 5 mm. wide. The color of the venter extends over half the length of the throat. The remainder and muzzle are about Isabella Color, and so are circular areas around the four teats (about 10 mm. diam- eter). Dorsal pelage agouti, paler than Citrine Drab (XL). The sides nicely Dark Olive Buff (XL). Dorsal guard hairs have two buffy bands each 1^/^ mm. long separated by a black band 23^ mm. long. Tips are black. Head, legs, and feet like back. Ears like muzzle. Tail color like dorsum both dorsally and ventrally. Not perceptibly distichous below. Tail hairs have black tips and four black bands 3 mm. to 4 mm. long (and four 2 mm. to 3 mm. light bands and a shorter light base) . Tail has short blackish pencil end- ing in buff tip. Notably, we think, in both cotypes the mid ventral agouti stripe is most intensely agouti in the stripe's midlength rather than near the throat and in the present specimen, does not reach the throat. Discussion. — This rather well-marked subspecies appears from the material examined to occupy a very small range in northeastern Burma between the Irrawaddy River at Bhamo and the Salween River. Anderson (1879, p. 240) gives an impression of a somewhat greater range when he says, "I have collected about twenty-five specimens of this squirrel from various parts of Burma, north of the capital [Mandalay], . . ." However, it seems doubtful that gordoni occurs south of the Shweli River in Burma excepting as an intergrade with C. f. shanicus, for shanicus is known from several localities north- east of Mandalay along the road and railroad to Lashio. To the north gordoni appears to be replaced rather abruptly by quinquestriatus, and evidence of intergradation between the two has MOORE AND TATE: DIURNAL SQUIRRELS 131 been described above under the discussion of subspecies quinquestri- atus. Subspecies gordoni differs from michianus and zimmeensis to the east in lacking the incipient black mark on the posterior two- thirds of the middorsum and in possessing a median agouti band which completely bisects the red pelage and which is itself the color of the agouti dorsum. To the south it seems most logical that gor- doni intergrades with shanicus from which it differs by possessing: (1) divided red ventral pelage, (2) a face which is just as dark agouti as its dorsum, and (3) a middorsum which lacks the incipient black on its posterior two-thirds. From subspecies [C. erythraeus] sladeni to the west across the Irrawaddy gordoni differs (so greatly that it is no wonder they have been considered separate species) by possess- ing: (1) the red venter divided, (2) the feet and rostrum agouti in- stead of red like the venter, (3) the tail colored like the back instead of extensively red-tipped. Details of the possibility of intergrada- tion with phayrei are considered under the species phayrei. Callosciurus flavimanus shanicus (Ryley) Sciurus atrodorsalis shanicus Ryley, 1914, Jour. Bombay Nat. Hist. Soc, 22, p. 662. (?) Sciurus griseopectus Blyth, 1847, J, Asiat. Soc. Bengal, 16, p. 873. Types. — Sciurus a. shanicus, BM No. 13.11.18.1, an adult male from Gokteik, 2133 feet. North Shan States, Burma, taken April 23, 1913, by G. C. Shortridge; griseopectus, IM No. 9369, a parous fe- male captive from unknown locality, presented in 1847 by Rajah R. Mullick, Material examined, all from Burma. — Maymyo, 800 meters (AMNH), four, (BM), six; Gokteik, 2133 feet (BM), two, (CNHM), two; Pyaunggaung, 2794 feet (BM), four, (CNHM), one; Kawlaw, 4400 feet, west of Taunggi (BM), one; northeast of Hoiparo, N, Shan States (BM), one; Mongha, N. Shan States (BM), one. Discussion. — Specimens of shanicus were taken at precisely the same elevations as pharyrei at three of the same localities (Maymyo, Gokteik, and Pyaunggaung). At Maymyo in 1937 Gerd Heinrich collected six phayrei between November 27 and December 25 and a series of four shanicus between December 3 and 20, He took one specimen of each form on one of these days (the 20th), Heinrich's four phayrei with substantial toothwear have head-and-body length ranging from 222 to 238 mm. Although three of the shanicus have substantial toothwear, all have head-and-body length between 200 and 211 mm. These data are quantitatively weak but no indication 132 FIELDIANA: ZOOLOGY, VOLUME 48 is seen in them that shanicus is a seasonal or other color phase of phayrei. This and the many differences reported in the account of species phayrei suggest strongly that they belong to separate species. The form shanicus closely approximates gordoni in size and will doubtless be shown to intergrade with gordoni. The color charac- teristics of shanicus show evidence of relationship also to atrodorsalis and zimmeensis, but the Salween River probably prevents intergra- dation. See also comparisons made under C. p. janetta. Pelage color. — The two topotypes at Chicago have dorsal pelage of body, legs, feet, head, and tail Grayish Olive and incipient black shows on the posterior two-thirds of the middorsum on both. The abrupt, brief, black tip of tail has whitish tips. The ventral pelage is Old Gold (XVI) in color, and all of it is agouti. The Chicago Nat- ural History Museum specimen from Pyaunggaung agrees with this, and so do the four specimens in the American Museum of Natural History from Maymyo. //a6*7s.— Shortridge's field notes on this subspecies follow: "The most plentiful squirrel in Hsipaw State and around Maymyo, recall- ing Funamhulus palmarum of India in its habit of collecting around gungalows. The black mark on the back was seldom conspicuous except in immature specimens." (G. C. Shortridge in K. V. Ryley, 1914, p. 721.) Type description. — Examination of the type of Sciurus griseopec- tus Blyth, 1847, reveals (in 1959) the following characteristics: Ven- tral pelage Ochraceous-Tawny to Tawny except for agouti throat and a midventral extension from the throat posteriorly halfway to the anus. Throat is Deep Olive Buff (XL). Muzzle and chin are Chamois (XXX), and so is a circular area 10 mm. diameter around each teat. Proximally the tail is lighter beneath. Dark Olive Buff (XL), than above. Deep Olive. Dorsal pelage agouti and entirely about Buffy Brown (XL). The ears are like the muzzle. Chamois. The dorsal guard hairs have two buffy bands each two mm. long and separated by a 2 or 3 mm. blackish band, and have black tips. The tail hairs are brownish buffy tipped and have four black bands. The tail's dull color may be in part due to the hair being old, but probably when fresh it is only a little less dull. The feet look faintly paler than the middorsum, but not paler than the sides. There are barely evident eye rings tending toward Chamois. The tail is not perceptibly distichous below. Discussion. — In the original description of griseopectus, Blyth (1847) mentioned two points important to identification of the re- MOORE AND TATE: DIURNAL SQUIRRELS 138 gion whence the type must have come: the divided red venter, and "the tail being slightly black-tipped, but with pale ends to the hairs." These conditions occur in ningpoensis, michianus, and zimmeensis, as will be shown below, and in gordoni as reported above. Strong tendency to feet blacker than dorsum mitigates against all of these but ningpoensis. Anderson (1879, p. 239) made something of a case for the type of griseopectus having come from China, and placed it in the synonymy of castaneoventris Gray 1842. As restricted here, castaneoventris has feet too black and tail tip too different. The proper color relationship of foot to dorsum is common in the large AMNH series of ningpoensis, but in ningpoensis the muzzle and chin are rather consistently cool gray, and the tail hairs have five or even six blackish bands instead of four. In ningpoensis also three light bands on a guard hair in the pelage of the dorsum is common, where- as in the type of griseopectus the highest number noted was two. In conclusion, the probability of the type specimen of griseopectus be- ing from China east of the Sal ween River seems slight to us, even though the sample of ningpoensis is so variable, and even though the area of southern China from which no material representing this spe- cies has been available, is extraordinarily large. When material becomes available from an area of intergradation between gordoni and shanicus, it seems likely that individuals quite like the type of griseopectus may be found among them. The ven- tral pelage color would be quite intermediate, and shanicus could contribute the proper muzzle, chin, ear, and foot color and the correct number of blackish bands on tail hairs and light bands on guard hairs of the dorsal pelage. The favorable gordoni influence would be toward browner dorsal pelage and less distichous tail shape. At present writing this identification of griseopectus seems more probable than geographic location in eastern China. Until someone can identify material from known localities with the type of griseopectus, however, it seems best to let the name shanicus stand and to associate grise- opectus with it only uncertainly. Callosciurus flavimanus hyperythrus (Blyth) Sciurus hyperythrus Blyth, 1855, Jour. Asiatic Soc. Bengal, 24, p. 474. Type. — IM No. 9478, an individual from Tenasserim, presented in 1852 by Major Berdmore. Ellerman and Morrison-Scott (1951, p. 479) suggest the Sittang Valley as the geographic source of the type. 134 FIELDIANA: ZOOLOGY, VOLUME 48 Type description. — The following characteristics of the type were observed by one of us in 1959: Ventral pelage, including that on limbs, is entirely Bay or Claret Brown except for the chin which is infiltrated by agouti and is about Hazel. The dorsal pelage is agouti and is about Light Brownish Olive (XXX). This color is ubiquitous except on the ears and the head forward of the ears, where it merges into Hazel. The short pelage on the rather battered ears is clearly deeper red, about Kaiser Brown or even Hays Russet. There is no eye ring. Dorsal guard hairs generally have two light bands each somewhat exceeding one mm. in length and separated 3 or 4 mm. from each other. The tail hairs have five blackish and five light bands and their tips are blackish on the proximal half of the tail but reddish on the distal half. The tail shows a faint tendency to be distichous beneath. Discussion. — Material representing the species flavimanus from lower Burma is so scarce that although we are doubtful regarding whether the type specimen came from the vicinity of the town of Tenasserim or just from somewhere in the province of Tenasserim (which seems from usage to have extended north at least as far as the lower Sittang Valley), it is not possible for us to locate the most probable geographic origin of the type by comparing it with material from various parts of Tenasserim. In any case the type locality is certainly not Moulmein as questioningly suggested by Robinson and Kloss (1918, p. 199) unless they would wish to imply across the Sal- ween River, for Moulmein is designated type locality for C. f. atro- dorsalis and has proven to be quite acceptable as such. Martaban across the Salween would be a perfectly acceptable type locality, but not necessarily any better than the Sittang Valley 70 miles farther north. However, until specimens become available for study from these places it must be left a little uncertain where the type locality of this named form is, and if it is in fact a subspecies, where its whole range may be. Burma between the Sittang and the Salween seems the most reasonable possibility, but it remains to be proven. Callosciurus flavimanus michianus (Robinson and Wroughton) Sciurus castaneiventris michianus Robinson and Wroughton, 1911, Jour. Fed. Malay States Mus., 4, p. 234. Sciurus castaneoventris haemobaphes G. M. Allen, 1912, Proc. Biol. Soc. Wash- ington, 25, p. 177. Types. — Sciurus castaneiventris michianus, BM No. 8.11.14.13, old female, from Mee Chee, Yunnan, collected January 5, 1903, by MOORE AND TATE: DIURNAL SQUIRRELS 135 F. W. Styan; haemobaphes, MCZ No. 13693, adult male, from Chih- ping, southeast Yunnan, collected February 26, 1911 by Kobayashi. Material examined, from Yunnan, China. — "Ssu Shan Chang," 9000 feet, Likiang (AMNH), three; Yunnanfu, 6400 feet (AMNH), two; 15 miles south of Kunming [= Yunnanfu] (USNM), one, (MNHN), one; Yunnanyi, 6500 feet (AMNH), one, (CNHM), one; Hangai, 5900 feet, 40 miles east of Talifu (AMNH), one; Chao Chu, 6700 feet, south end of Lake Talifu (AMNH), one; Shan Kuang, 6000 feet. Lake Talifu (AMNH), two, (CNHM), three, (MCZ), one; Mekong-Salween divide at 28° 20' N. latitude, 7000 feet (BM), one; Chepei, Fumin (AMNH), one; Kao Chiao, Yunnanfu (AMNH), six, (MCZ), one; Lungkai, Makaihsien (AMNH), six; Lungkai, Wutinghsien (AMNH), one; "Kun Yang" (AMNH), one; "Nda Mu Cho" (USNM), one; Makaihsien (MCZ), one; Likiang, 9000 feet (MCZ), three, (USNM), six, (CNHM), four, (BM), one; Chihping [Shihping] (MCZ), four; "Hofuping Mountains," Mekong Valley (USNM), three; Chiihsiung, 160 miles east of Yunnanfu (CNHM), one; Nui Kai, 7200 feet, 30 miles north of Talifu (CNHM), one; Lo Tu Kow [Lukou] (BM), one; six miles north of Luchang (BM), two. This subspecies seems to be bounded on the northeast by the Yangtze, but allocation of the Yung-pei-ting specimen to gloveri is made on this assumption in absence of adequate color notes on the specimen. The southwestern boundary of the range of michianus is probably the Mekong River. To the south in southern Yunnan michianus must surely intergrade broadly with Callosciurus flavi- manus hendeei. Diagnosis. — Subspecies michianus generally differs from gloveri to the north by possessing: (1) an agouti throat and a wedge from it extending posteriorly as an incipient midventral agouti stripe; (2) an incipient blackening of the pelage on the posterior two-thirds of the mid-dorsum; (3) a short, incipient, sub terminal, black tip to the tail; and (4) digits which are often notably blacker agouti than that of the animal's own gray dorsal pelage. Subspecies michianus differs from castaneoventris in characters (2) and (4) above, and in lacking a reddish infusion in the agouti dorsal pelage. From hendeei, michi- anus differs by characters (1) and (2) above, and the feet of hendeei are substantially blacker, and the black tail tip of hendeei is substan- tially blacker and longer, than that of michianus. Across the Me- kong River from C. flavimanus michianus is a subspecies, C. flavi- mauns gordoni, from which michianus differs only by being paler and larger. 136 FIELDIANA: ZOOLOGY, VOLUME 48 Callosciurus flavitnanus zimmeensis (Robinson and Wroughton) Callosciurus atrodorsalis zimmeensis Robinson and Wroughton, 1916, Jour. Fed. Malay States Mus., 7, p. 91. Callosciurus ferrugineus primus Allen and Coolidge, 1940, Bull. Mus. Comp. Zool., 87, p. 157. Types. — zimmeensis, BM No. 9.10.11.20, an adult female taken at Chiengmai, 2100 feet, northern Siam on April 12, 1908, by T. H. Lyle; primus, MCZ No. 35352, old female from Mae Wan River, 1500 feet, near Mount Souket, north Thailand. Material examined, from Yunnan, China. — Mucheng, 7000 feet (AMNH), four, (CNHM), two, (MCZ), one; Hsiaomengtung, 6000 feet (AMNH), one; Tashutangh, 6000 feet (AMNH), one; Chenkang, 3000 feet (AMNH), two; Chaunglung, Salween ferry, 2000 feet (AMNH), four, (CNHM), three, (MCZ), one; and Nam Ting River at Burma border (AMNH), four, (CNHM), two, (MCZ), two; Mengting, 1700 feet (AMNH), one. Material examined, from Burma. — Malipa (AMNH), one; Ban Sob Pa, Salween River (USNM), two. Material examined, from Thailand. — Doi Soutep (AMNH), three, (BM), seven, (USNM), one; Mt. Chieng Dao (AMNH), one, (USNM), three, (ANSP), three; Pa Kwe, Chiengmai (AMNH), two; Mt. Doi Pui, Chiengmai (AMNH), two; Mong Hawt, Chieng- mai (AMNH), two; Ban Kang, base of Doi Angka (USNM), two; Chomtong (USNM), one; Mekhan (USNM), one; Waterfall Me- kang (USNM), one; Mae Hong Song (BM), two, (USNM), one; Doi Hua Mot (USNM), one; Doi Angka (MCZ), 13, (CNHM), one; Chieng Mai (ANSP), two, (BM), four; Mee Tan [Ban Mae Tan Nua] (BM), one; Ban Meh Na (NR), two; Doi Par Sakeng (NR), three; Muang Pai, 600 meters (BM), one; "Muang Sang," 425 me- ters, north of Chiengmai (BM), one; Me Ping River at IS** 00' N., 235 meters CBM), one; "Me Ping River" (BM), four. This subspecies occupies the waveringly wedge-shaped area be- tween the Salween River and the Mekong River from about 18° north latitude to about 25°, and possibly to 26° or beyond. The type locality is rather close to the area of intergradation with atro- dorsalis on the south. Diagnosis. — Subspecies zimmeensis differs from michianus on the east by being smaller and darker, from gordoni on the west by pos- session of the incipient black posterior middorsum and less nicety of the midsagittal division of the red venter, from shanicus by possess- ing a red, divided venter and five dark bands on the tail hairs, from MOORE AND TATE: DIURNAL SQUIRRELS 137 hendeei by having at least incipient bisection of the ventral red pelage and by having the tail colored like the dorsum or with only a thin buffy tipping (in some southern material), and from atrodorsalis and thai by having the posterior two-thirds of the middorsum only in- cipiently black (i.e., without an area of pure black). Discussion. — Note in Figure 13 that zimmeensis material is widely divided into two lots. The difference between the northern (Tem- perate Zone) material of zimmeensis and its neighbor across the Me- kong is rather slight when compared with characters differentiating other subspecies of this revision of Callosciurus flavimanus. If this Temperate Zone sample of zimmeensis had shown greater differences from material from Thailand (300 miles into the tropics) we should have included the northern lot in michianus rather than naming it as a new subspecies. For discussion of the named form primiis see the account of C finlaysoni menamicits. It is interesting that about the Tropic of Cancer where the pop- ulations of species flavimanus have been sampled on both sides of the Salween and Mekong rivers which have run closely parallel to one another for about 400 miles, the samples from either side of the Salween are much more highly differentiated from each other than those from either side of the Mekong. See Table 7. Table 7. Subspecies of C. flavimanus separated by Salween River and Mekong Rivers 300 kilometers north and 300 south of the Tropic of Cancer. E. of Salween W. of Salween, E. of Mekong W. of Mekong quinquestriatiis zimmeensis michianus gordoni zimmeensis michianus thai zimmeensis hendeei Callosciurus flavimanus thai (Kloss) Sciurus atrodorsalis thai Kloss, 1917, Jour. Nat. Hist. Soc. Siam, 2, p. 285. Type. — Field No. 2474/CBK (not seen), from Raheng, central Siam, collected July 1916 by C. Boden Kloss. Material examined, from Thailand. — Mewong River, 1000 feet, 40 miles east of Um Pang (AMNH), two; Mewong River, 800 feet, 53 miles east of Um Pang (AMNH), two, (BM), four; 20 miles west of Kempenpet (AMNH), one, (BM), one; "Siam" (AMNH), one; 138 FIELDIANA: ZOOLOGY, VOLUME 48 Muang Kan Bouri [Kanchanaburi] (USNM), three; Ban Pong (USNM), one; Hue Yah Pla, 2500 feet (USNM), two; Me Taqua, 800 feet (USNM), five, (CNHM), one; Wang Kien (USNM), one; Sai Yoke (BM), one; "Dooweeklo," 1500 feet, Mae Klong (BM), one; Mae Taw Forest, 900 feet, Raheng (BM), five; Lai Yoke [Ban Sai Yoke] (BM), one; 10 miles west of Raheng, 600 feet (BM), one; Raheng, 115 meters (BM), one; 28 miles east of Um Pang, 2000 feet (BM), two; Doi Mei Lai [Doi Mae Lai] (USNM), one; Doi Mae Kong Kha (USNM), two; Banphotphisai, Nakon Sawan Prov. (USNM), four; Slokbai [Ban Salok Bat], Kano, Kamphaeing Phet (USNM), one; "Tumphol," Kha Nu, Kamphaeing Phet (USNM), six; Ban Hua Thanon, Klong Klung, Kamphaeing Phet (USNM), six. Material examined, from Burma. — Lampha, Tenasserim (AMNH), one, (BM), three. The zimmeensis from northern Thailand show some intergrada- tion with the subspecies thai to the west and southwest, in occasional occurrence of some pure black pelage on the posterior two-thirds of the middorsum, and in the general occurrence of long light tips on the tail hairs which give to the posterior half of the tail an overlay of straw color (Pale Ochraceous Buff to Light Ochraceous Buff, XV). Diagnosis. — The subspecies thai is distinguished from zimmeensis, however, by having the following characteristics; (1) regular occur- rence of the solid black band of pelage 20-30 mm. wide along the posterior two- thirds of the mid-dorsum; (2) the rostrum, and occa- sionally the crown, is colored rusty (ranging from only Buckthorn Brown to Hazel) ; whereas in zimmeensis the rostrum and crown are always a cool gray. Callosciurus jiavimanus thai is distinguished from atrodorsalis by: (3) having its feet a blacker agouti than the legs; (4) having its hind- quarters not notably redder than its forequarters; (5) having long straw-colored tips to the tail hairs (Light Buff to Light Ochraceous Buff) instead of reddish (Ochraceous Buff to Cinnamon Rufous). Habits. — Callosciurus jiavimanus thai appears to occupy a rain shadow zone in Thailand and is probably found where the forest is primarily deciduous; whereas the following subspecies, atrodorsalis, apparently occurs where the forest is primarily broad-leafed ever- green rain forest. Callosciurus flavitnanus atrodorsalis (Gray) Sdurus atrodorsalis Gray, 1842, Ann. Mag. Nat. Hist. (ser. 1), 10, p. 263. MOORE AND TATE: DIURNAL SQUIRRELS 139 Cotypes.—BM Nos. 41.1818 and 41.1819, both adult males re- ceived from the East India Company, "from Bhotan," corrected by Ryley (1914, p. 663) to Moulmein. Material examined, all from Burma. — Taok Plateau, 3050 feet (AMNH), eight, (BM) four; Zami R., 100 mi. S. of Moulmein (BM), one; Kokareet [Kawkareik], Tenasserim (USNM), one, (CNHM), one, (BM), three; Mt. Nwalabo, Tavoy District (BM), two; Doonsa (BM) one; Thaungyin Valley (BM), one; "Lathorgu," Myawadi (BM), one; Ban Sob Pa, Salween R. (USNM), two; Myawadi (BM), two; Sookli [Sukli], Myawaddy Rd. (BM), one; Maitho, Thaungyin River, Tenasserim (BM), one; Haungtharaw River (BM), one; "Shan Mepa," Amherst (BM), three. Pelage color. — One striking character that shows the relationship between the subspecies atrodorsalis and thai is, of course, the broad (20-30 mm.) band of pure black pelage on the posterior two-thirds of the mid-dorsum. Many subspecies of flavimanus have an incip- ient manifestation of the black band but only atrodorsalis and thai have it pure black. The agouti dorsal pelage varies considerably in color individually among specimens from the rainforest of the Tenasserim region, de- pending upon the amount of infusion with red. The rostrum in the atrodorsalis at the American Museum of Natural History is redder (Hazel to Kaiser Brown, XIV) than the dorsal pelage on the same individual. The anterior dorsal pelage in the Tenasserim AMNH series ranges from about Hazel in the reddest to about light Brown- ish Olive (XXX) in the grayest. The posterior half of the dorsal pelage (exclusive of the black stripe) is redder on each individual than the anterior half, the range in the same series being about Russet (XV) to Saccardo's Umber (XXIX). The tail is just about as reddish as the hindquarters on each individual. The feet, unusual in this species, are no blacker agouti than the legs. Discussion. — This subspecies occupies the very heavy rainfall area of Tenasserim and is much redder than the adjacent subspecies thai which occupies the contiguous rainshadow area in Thailand. Anderson (1879, p. 233) placed a great deal of stress on the oc- currence of two color phases in atrodorsalis, the more common one with a broad black band on the back, the other without. He found that the difference between the two was not sexual dimorphism and that neither color phase exclusively represents the stage of maturity. Whether it is seasonal, his material was not adequate to reveal, and 140 FIELDIANA: ZOOLOGY, VOLUME 48 we have examined too few of the ones lacking black patches to say. For the geographic range that Anderson attributed to atrodorsalis (Moulmein to Malacca), the difference has now been shown to be geographic, indeed, but he was also concerned with the local varia- tion in the Moulmein area and to the east of Moulmein. Here he declared the evidence was non-geographic but he did not state the evidence in adequate detail for one confidently to agree. Anderson (1879, p. 235) also states that "atrodorsalis is very common in Marta- ban. ..." We have seen no specimens from there, and since neither caniceps nor phayrei are known to have crossed the Salween River, we have presumed (in absence of any substantial evidence) that atrodorsalis does not cross it either. If this is the case, it may well be that the squirrel species Anderson mentions occurring in Marta- ban is in fact hyperythrus. He considered the type of hyperythrus to represent the phase of atrodorsalis without the black band on the back. We observed nothing about the type specimen of hyperythrus which mitigates against this concept (see description above), but it seems quite possible that squirrels taken just across the river from Moulmein at Martaban might in those days have been quite freely labeled "Moulmein," and that such material may in part constitute the phase of atrodorsalis without the black back which Anderson (1879, p. 233) describes from the Moulmein region. This problem will, of course, not be solved by disposition here of the name hyperythrus. What is needed is a series of specimens from Martaban and any points north of there (and west of the Salween) for comparison with the type of hyperythrus and with new material of the scarcer color phase of atrodorsalis (if any) from the Moulmein area and eastward from there toward Kawkereik and Myawadi. Callosciurus flavimanus siamensis (Gray) Sdurus siamensis Gray, 1860, Ann. Mag. Nat. Hist. (ser. 3), 5, p. 500. Sciurus atrodorsalis tachin Kloss, 1916, Jour. Nat. Hist. Soc. Siam, 2, p. 178. Types. — Sciurus siamensis, BM No. 59.7.8.1, young adult with dp* present, from Siam, collected by M. Mouhot; tachin, USNM No. 221566, a young adult female from Pak Bu, Tachin, Siam, taken on October 23, 1916, by C. Boden Kloss. Material examined, all from Thailand. — Bangkok (AMNH), three, (CNHM), six, (MCZ), 12, (UMMZ), 10, (USNM), 22; Pak Jong (AMNH), one; "Doi Souket," 6500 feet (MCZ), one; Paknampoh (MCZ), one; Tachin (USNM), six; Rajaburi (USNM), three; Pota- ram, near Ratburi (USNM), one; Ban Pong, near Rajaburi (USNM), one. MOORE AND TATE: DIURNAL SQUIRRELS 141 Material examined, of intergrades with C. f. thai, all from Thai- land.— Paknampo, Nakorn Sawan (USNM), four; Moung Wat Sy, Nakorn Sawan (USNM), one; Kowkat, Paknampo, Nakorn Sawan (USNM), four; Kowkob, Paknampo, Nakorn Sawan (USNM), one. Material examined, of intergrades with C. f. pranis. — Tra Kha- nun, Kanachanaburi, Thailand (USNM), three; Hinlaem, Tra Khanun, Kanachanaburi, Thailand (USNM), one. The type locality of tachin seems to be the southwest edge of the geographic range of this subspecies, and material from there shows intergradation with pranis. Consequently, material from Tachin, Ratburi (=Rajaburi), Potaram, and Ban Pong are to be expected to show evidences of such intergradation. Diagnosis. — This subspecies is best known from the large series from Bangkok in several museums. C. /. siamensis is distinguished from C. f. thai on its west by having: (1) a maximum of four blackish bands on the tail hairs instead of five; (2) the tail increasingly red toward the tip instead of ivory or yellowish white; (3) the feet no blacker agouti than the legs; (4) the venter duller red, generally about Tawny or Hazel (although we recorded extreme ones as bril- liant as Burnt Sienna), whereas ventral pelage in six out of seven thai is Mahogany Red and the other Burnt Sienna; (5) smaller size; (6) no indication of a broad black longitudinal stripe on the poste- rior two-thirds of its mid-dorsum. From C. f. zimmeensis to the north, siamensis is also distinguished by above characters numbers 1, 2, 5, and 6. From C. f. pranis to the south, siamensis is distinguished by above characters numbers 2, 3, and 6. Callosciurus flavimanus pranis (Kloss) Sciurus erythraeus pranis, Kloss 1916, Jour. Nat. Hist. Soc. Siam, 2, p. 178. Type.— USNM No. 221568, an old male from Koh Lak [Prachuap Kiri Khan], southwest Siam [Thailand], taken November 9, 1916, by C. Boden Kloss. Material examined, all from Thailand. — Koh Lak [Prachuap Kiri Khan] southwestern Siam (USNM), six, (NR), four, (BM), two; Hue Sai (NR), one; Sungei Balik (BM), one; Bok Pyin (USNM), two; Khao Luang, 3400 feet (ANSP), two; "Khao Nok Wua," 2000 feet (ANSP), one. Pelage color. — The dorsal pelage is agouti and as a whole appears to be about Olive Brown and indistinguishable from that of tachin 142 FIELDIANA: ZOOLOGY, VOLUME 48 except that pranis regularly has an incipient expression of the black middorsal stripe that demonstrates its relationship to atrodorsalis. The reddish ventral pelage is regularly divided longitudinally by a narrow band of agouti pelage. The color of the reddish pelage ranges from Tawny Olive (XXIX) to Tawny. It also shows relationship to atrodorsalis and tachin in having the pelage of the rostrum hazel, and the ears tinged with the same color. Discussion. — This as yet poorly known subspecies is intermediate in some characteristics between C. /. atrodorsalis and rubeculus, but the material is difficult to shrug off as series of intergrades, because it seems consistently to possess a lighter, less richly colored venter than either atrodorsalis or rubeculus. Callosciurus flavimanus rubeculus (Miller) [extraterritorial] Sciurus rubeculus Miller, 1903, Smithsonian Misc. Coll., 45, p. 22. Sciurus erythraeus youngi Robinson and Kloss, 1914, Ann. Mag. Nat. Hist, (ser. 8), 13, p. 224. Types.— S. rubeculus, USNM No. 86777, old male from Khao Soi Dao, 1000 feet, Trong, collected February 21, 1899, by W. L. Abbott; youngi, BM No. 21.11.8.11, old female, from Teku Plateau, 5000- 6000 feet, Gunong Tahan, north Pahang, collected July 19, 1911, by H. C. Robinson and C. B. Kloss. Discussion. — Here is another subspecies which takes its name from a type specimen obtained in an area of intergradation with an adjacent subspecies. The typical material from Trong shows some trend toward intergradation with the only adjacent subspecies, pranis. Although C. f. rubeculus is partially isolated on a long peninsula far removed from most of the range of its species, it displays a most intensive expression of those conservative pelage characters which are so widespread in the species as to be regarded "typical" of the species. (No known pelage or other character has been found which delimits the species flavimanus.) These widespread characters are: faintly orange-rimmed ears, agouti feet blacker than the legs and sides, dark red ventral pelage bisected longitudinally by a narrow band of agouti gray pelage, tail hairs with four or five blackish bands. Diagnosis. — This subspecies is here distinguished from the only other conspecific one with which it is in contact, C. f. pranis, by: (1) absence of any suggestion of a broad blackish band on the poste- rior two-thirds of the mid-dorsum; (2) darker, richer ventral pelage; (3) blacker feet; (4) gray rostrum. Subspecies rubeculus is not mapped here. MOORE AND TATE: DIURNAL SQUIRRELS 143 Callosciurus flavimanus hendeei (Osgood) Callosciurus erythraens hendeei Osgood, 1932, Field Mus. Publ. Zool., 18, p. 270. Type.— CNUM No. 32290, an adult male taken at Chapa, Ton- kin, February 14, 1929, by Russell W. Hendee. Material examined, from Tonkin. — Chapa, 5000 feet (CNHM), six, (MCZ), two, (USNM), one, (BM), 12; Bao Ha, 500 feet (CNHM), one, (BM), six; Muong Mo, 750 feet (CNHM), four, (USNM), one; Muong Boum (CNHM), one; Lieng San [Leng Sang], 1500 meters (CNHM), one; Pakha [Pa Kha] (CNHM), two; Isle de la Table (BM), one; Lao Kay (MCZ), three; Thai Nien, 300 feet (BM), four; Lai Chau (USNM), one, (MNHN), one; Langson, 500 feet (BM), one; Tam-dao (MNHN), one, (BM), four; Bac Kan (BM), four; "Chora," 1000 feet (BM), one; Ha-Giang (MNHN), one; Ngan-son, 3000 feet (MNHN), one; Muong Moun (AMNH), two. Material examined, from Laos. — Lao Fou Tchay, 3400 feet (CNHM). one; Phong Saly, 4400 feet (CNHM), four, (USNM), one; Muong Yo, 2300 feet (CNHM), two; Lo Tiao (MCZ), one; 2 km. east of Nam Khueng (MCZ), one; Hou^ Sai (MCZ), one. Material examined, from Annam. — Phu Qui, 100 feet (CNHM), two, (MNHN), one, (BM), seven; Than Hoa (CNHM), one; Hoi Xuan (CNHM), one; "Lung-lunh" (CNHM), two, (BM), one; Lung Van, 1000 meters, pr. de Thanhoa (USNM), one; "Nam Da, near Tatka" (BM), one; Nghia Hung, 100 feet, Phu Qui (BM), seven. Material examined, from China. — "San-ho Hsien," Kweichow (CNHM), one; "Shui-kow-kwan," Lungkow, Kwangsi (CNHM), one. For discussion of characters and relationships see castaneoventris and flavimanus. Callosciurus flavimanus castaneoventris (Gray) Sciurus castaneoventris Gray, 1842, Ann. Mag. Nat. Hist. (ser. 1), 10, p. 263. Sciurus erythraeus insularis J. A. Allen, 1906, Bull. Amer. Mus. Nat. Hist., 22, p. 473. Types. — Sciurus castaneoventris, BM No. 72A, young female and 72B adult male, both from "China," evidently the vicinity of Canton and collected between 1812 and 1831, for they were presented by John R. Reeves (who worked there as a tea taster during that time; see Moore and Tate, 1959) ; S. e. insularis, AMNH No. 26609, parous female from "Lei-Mui-Mon," Hainan, taken January 5, 1903 by agents of Alan Owston. 144 FIELDIANA: ZOOLOGY, VOLUME 48 Material examined, all from Hainan, China. — "Lei Mui Mon" (AMNH), 14; "Utocki" (AMNH), one; "Luidon" (AMNH), one; "Hainan" (AMNH), three; Nodoa (AMNH), 80, (USNM), two, (CNHM), 14, (BM), one, (MCZ), six; Nam Fong (AMNH), seven, (CNHM), two; Kachek (USNM), one; "Ngau Tchi Lea Mts." (MCZ), one, (BM), one. Discussion. — No specimens of the species flavimanus, other than the cotypes collected by Reeves are known from the vicinity of Can- ton, but Robinson and Kloss (1918, p. 199) compared the cotypes with material from Hainan and found them to be indistinguishable, thus rendering J. A. Allen's subspecies insularis from Hainan a syn- onym of castaneoventris. Without comment these authors thereupon substituted Hainan as type locality for the "China" which was the only type locality provided by Gray in his original description of castaneoventris. While that substitution was a considerable improve- ment in its time, as indicated above, it now seems better to accept the vicinity of Canton as type locality, but only provisionally until further material becomes known from there. There were two specimens involved in the original description of castaneoventris. The first mentioned was a rather darker one (BM No. 72B) the original description of which amounts to, "Very like S. hippurus, but only half the size, and the ears are gray." This im- plies red ventral pelage like that of hippurus, for Gray proceeds to contrast the variant second specimen with it, "Var. rather paler; chin grayish, beneath yellowish red." The amount of variation im- plied does not require that the type series came from widely different localities nor that it came from a single locality which is in an area of intergradation between two subspecies (castaneoventris and ning- poensis) . Nevertheless, the location of Canton, from which the type material is presumed to come, in reference to known localities for ningpoensis and subsequent castaneoventris material does suggest this latter possibility, and the Hungshui River which debouches about Canton and Macao, could quite possibly form a partial, natural bar- rier between the two subspecies. If this should prove to be the case, the type series does better represent the southern, not the northern subspecies, and no need is anticipated to alter the well-established usage of castaneoventris for Hainan material and ningpoensis for that from Fukien. In his diagnosis of the subspecies hendeei Osgood (1932, p. 270) stressed no single character which could be depended upon alone to distinguish it from the one on Hainan, but stated all the differences MOORE AND TATE: DIURNAL SQUIRRELS 145 in subjectively comparative terms. It seems worthwhile to provide quantitative differences here as we find them to exist. Pelage color. — Among the large series of castaneoventris from Hai- nan at the American Museum of Natural History there are only five individuals that have a full-length midventral band of agouti pelage. There are 56 that have agouti throats and an agouti wedge extending onto the breast, and 41 that have neither midventral band nor wedge. This contrasts with the series of thirty skins of hendeei at Chicago Natural History Museum in which there were no full-length, ventral, median bands of agouti and only one with an agouti wedge on the breast (No. 32297). The ventral pelage color of the large AMNH series from Hainan is about Mahogany Red in about 30 specimens. There are one or two skins of adults in good pelage as pale as Mars Orange, and the remainder are about Burnt Sienna (except for one individual from Nodoa (AMNH No. 58133) which has such an exceptionally agouti venter that virtually only the insides of the legs have really red pel- age. The possibility of such aberrant individuals elsewhere needs to be borne in mind in this species) . This is a very high degree of con- sistency in ventral pelage color compared with ningpoensis. In the (smaller) Chicago collection of hendeei material, however, the ven- tral pelage color was found to vary from about Morocco Red to Hays Russet, a range of variation which is even more conservative than the castaneoventris. C. f. castaneoventris and C. f. hendeei both have five black bands on the tail hairs. The most constant and obvious color difference between the hen- deei and castaneoventris as represented in the large series at Chicago and New York is the color of the tip of the tail. In the Hainan ma- terial there appears to be no variation away from a white-upon-black- tipped tail. Some description is necessary here. The tail hairs of these C. f. castaneoventris from Hainan, excepting near the distal end of the tail, have five blackish bands and a minute blackish tip as is widespread in the species. But the light subterminal band is un- usually light in this species (about Cartridge Buff) and is progres- sively longer toward the extremity of the tail, often exceeding a centimeter at the extremity, but rarely two centimeters. The most distal black band becomes progressively longer, also, toward the end of the tail, by spread toward the base of the hair. Thus, at the end of the tail the subterminal whitish band of one centimeter is distal to a sub-subterminal black band of from two to five centi- 146 FIELDIANA: ZOOLOGY, VOLUME 48 meters. (Osgood, incidentally, ignored the minute blackish tips of the hairs and described the whitish bands as terminal.) The tips of the tails of the hendeei series at Chicago Natural History Museum are characteristically and uncomplicatedly black. There are four (three from Chapa and one from Lung-lunh), how- ever, which do have long, light bands superimposed on the extrem- ities of the black hairs, rendering them indistinguishable by this criterion from Hainan material. However, 87 percent is a fairly good distinction of subspecific material (Amadon, 1949) on the basis of a single color character. Dimensions. — Size is something on which we can also provide data: Greatest skull length in 36 adults of castaneoventris ranges from 50.2 to 53.5 mm., except for two well outside the curve at 55.0 and 55.9, and 29 (80 percent) are less than 53 mm. long; whereas all 17 skulls of adult hendeei are 53 mm. or more in length. Similarly, the hind foot length of the adults of hendeei ranges from 54 to 59 mm., but in a sample of 61 adult castaneoventris only four range above 53 mm. Callosciurus flavimanus flavimanus (Geoffroy St. Hilaire) Sciurus flavimanus Geoffroy St. Hilaire, 1831, in Belanger, Voyage Indies Orientales, 1, p. 148. Callosciurus flavimanus quantulus Thomas, 1927, Proc. Zool. Soc. London, 1927, p. 51. Callosciurus flavimanus contumMX Thomas, 1927, Proc. Zool. Soc. London, 1927, p. 52. Callosciurus flavimanus dactylimis Thomas, 1927, Proc. Zool. Soc. London, 1927, p. 52. Callosciurus flavimanus piraia Thomas, 1929, Proc. Zool. Soc. London, 1928, p. 836. Callosciurus flavimanus bolovensis Osgood, 1932, Field Mus. Nat. Hist., Zool. Ser., 18, p. 276. Types. — S. flavimanus, MNHN No. 177, from Tourane, Annam; quantulus, BM No. 26.10.4.96, adult male from Xieng Quang Koo, Laos, collected December 19, 1925, by J. Delacour and W. Lowe; contumax, BM No. 26.10.4.104, adult female from Kontoum, An- nam, collected February 27, 1926, by J. Delacour and W. Lowe; dactylinus, BM No. 26.10.4.101, old female from Dak-to, Annam, collected March 16, 1926, by J. Delacour and W. Lowe; pirata, BM No. 28.7.1.79, adult female from Nape, 2500 feet, Laos, collected February 11, 1928 by J. Delacour and W. Lowe; bolovensis, CNHM No. 37874, old female from Paksong, Indochina, collected Janu- ary 24, 1932 by J. Delacour. MOORE AND TATE: DIURNAL SQUIRRELS 147 Material examined, from Annam. — Col de Nuages, 400 meters (MNHN), eight, (BM), 12; Thua Luu, 150 feet (BM), two; Dak To (BM), two, (MNHN), two; Kao Hao, Quangtri Province (BM), one; Kontoum [Kontum] (BM), two, (MNHN), two; Hu^ (BM), one; Cua-Ting, Quangtri (BM), one; Quangtri (BM), one, (MNHN), one, (CNHM), six, (USNM), one; Dong Hoi (MNHN), one; Quang- tri River (MNHN), one. Material examined, from Laos. — Thateng (CNHM), 15, (BM) one, (USNM), one; Nap6, 2500 to 3000 feet (BM), five; Xien Kuang Koo [Xien Khouang] (BM), three, (MCZ), two; Pasa (AMNH), one; Don Qua (AMNH), one; Plateau Bolovens (AMNH), four; Phu Kobo (MCZ), two; Bantion [Ban Tayun] (CNHM) one; "Pakhout" (CNHM), one; Pak Song (CNHM), one; Nong Kai [presumed to mean in Laos across the river from Nong Kai, Thailand] (USNM), one. Discussion. — The two AMNH specimens from Pasa and Don Qua, villages on the Nam Khan (river) between Luang Prabang and M. You, exhibit a condition of intermediacy between C. f. hendeei and C. f. flavimanus. There is no geographic barrier between these localities and those of the hendeei. There is no difference from hen- deei in the appearance or size of the Pasa and Don Qua skulls, and body measurements of these and hendeei taken by the same collector (T. Donald Carter) offer no hint of a size difference. The rostrum is colored like the dorsum as in hendeei rather than orange-yellow as in flavimanus. The feet are agouti like the back, a stage midway between the blackened agouti of hendeei and the orange-yellow of flavimanus. However, the Pasa specimen possesses pronounced black- ish agouti on its toes showing more influence of the hendeei character; whereas the Don Qua one lacks the blackish agouti entirely but has a faint infusion of reddish-orange around the ankles (absent in the Pasa specimen) showing more the influence of flavimanus. In both of these Nam Khan (river) specimens the tail is colored like the back but brighter buff approaching the condition of flavimanus, not black- tipped like hendeei. The Museum of Comparative Zoology material from Xien Khou- ang and Phu Kobo seems to be quite good flavimanus with no indi- cation of intergradation ; but five of the six Chicago Natural History Museum specimens from Quangtri have abrupt solid black tail tips indicating some penetration of hendeei characteristics even to that locality. Both of the Nam Khan specimens have four prominent blackish bands on the tail hairs. This is a reduction of one from the 148 FIELDIANA: ZOOLOGY, VOLUME 48 number characteristic of hendeei and one or two more than is com- mon in flavimanus farther south. The magnitude of difference between diagnostic characteristics of flavimanus and hendeei does greatly exceed that between hendeei and any of the Chinese subspecies. The evidence of intergradation described above compells us, however, at this state of our knowledge, to regard subspecies flavimanus and hendeei as conspecific. It is also quite evident to us that flavimanus material is much more variable individually than is that of hendeei, but these conditions in no way justify admission as good subspecies any of the many other names that have been applied in this case. Pelage color. — The characteristics accepted for flavimanus are orange-yellow rostrum and fore feet, and reddish-orange hind feet. The tail may be colored like the dorsum or huffier or approaching the color of the venter. The venter varies among individuals from Mars Orange to Burnt Sienna. This color extends from wrist to ankle and covers the venter except for the agouti scrotum and chin. An occasional individual shows feeble anterior indications of a mid- ventral agouti stripe. Faint indications of a broad, black midstripe on the posterior two-thirds of the dorsum is characteristic. Callosciurus flavimanus griseimanus (Milne-Edwards) Sciurus griseimanus Milne-Edwards, 1867, Rev. Zool. (ser. 2), 19, p. 195. Macroxus leucopus Gray, 1867, Ann. Mag. Nat. Hist. (ser. 3), 20, p. 282. Sciurus leucopus fumigatus Bonhote, 1907, Abstr. Proc. Zool. Soc. London, no. 38, p. 2. Sciurus vassali Bonhote, 1907, Proc. Zool. Soc. London, 1907, p. 9 (footnote, a new name for fumigatus Bonhote, 1907, not of Gray, 1867). Callosciurus ferrugineus phanrangis, Robinson and Kloss, 1922, Ann. Mag. Nat. Hist. (ser. 9), 9, p. 91. Types. — Sciurus griseimanus, MNHN No. 1864-679(167), adult male from Saigon, collected by R. Germain; leucopus, BM No. 60.8.28.13, adult male from Cambodia; vassali, BM No. 6.11.6.25, male, from Bali, near Nhatrang, Annam; phanrangis, BM No. 26.- 11.17.5, adult female from Tour Cham near Phanrang, south Annam, collected May 23, 1918, by C. B. Kloss. Material examined, from Annam. — Djiring, 3000 feet (BM), 11, (MNHN), four; Da Ban (BM), one; Nhatrang (BM), 12; Phanrang (BM), one; Lang Bian Peaks (MCZ), one; Pie de Langbian (MCZ), one; forest of Krongba [Krong Ba] (USNM), one. MOORE AND TATE: DIURNAL SQUIRRELS 149 Material examined, from Cochin China. — Tayninh (BM), four, (MNHN), six; An Binh (BM), three; Trang Bom (BM), two; An Blu (MNHN), one; "Arboretum de" Trang Bom, Province de Bien Hoa (USNM), three; Bien Hoa (USNM), five; (MNHN), one; "Montaojur de Nrii," Chua Chan (USNM), one; "Prang," (USNM), one; Baria (CNHM), two; Ninh Hoa (CNHM), two; Phan Rang (CNHM), two; "Gougah" (CNHM), two; Ban Me Thuot (CNHM), two; "Eaktur" (CNHM), one. Diagnosis. — The distinguishing characteristics of griseimanus are: (1) very Hght dorsal pelage, some Mouse Gray, some Drab; (2) seem- ingly white feet, about Cream Buff in some. Cartridge Buff in others; (3) tails not tipped with a contrasting color, but unusually disposed to show transverse annulations, numbering about ten; (4) number of blackish bands on tail hairs at least four, often five. There is, in griseimanus, as in flavimanus, a great deal of varia- tion that is individual or microgeographic or both. Pelage color. — The ventral pelage color in eleven specimens at the CNHM varies from Sanford's Brown in two, through Ochraceous Buff in two. Cinnamon Buff in five, to something lighter still in two immatures from Phan Rang. Bisection of the ventral pelage by an agouti stripe is incipient in four of these eleven. Discussion. — A specimen (USNM 256749) indistinguishable from hendeei was, according to the label, taken by Poilane on Nua Chua Chan, a small mountain close to Bien-hoa, Cochin China, deep in the range of subspecies griseimanus, from which hendeei differs greatly. A companion specimen of hendeei was taken by Poilane (USNM 256748) at Lung Van, Than Hoa Province, Annam, well within the proper range of hendeei. It seems rather more likely that the first specimen somehow became erroneously labeled than that a squirrel which is indistinguishable from hendeei could in fact have been taken on Nua Chua Chan (see map of species flavimanus, figure 13). The color difference between subspecies griseimanus and flavi- manus is just as striking as that between subspecies flavimanus and hendeei, but it is color intensity which differs more than pattern, and in view of the color patterns common to these first two forms and absence of any geographical barrier between them, it would seem unrealistic to us even in the absence of noticed intergradation, to regard griseimanus and flavimanus as more distinct than subspecies. 150 FIELDIANA: ZOOLOGY, VOLUME 48 Callosciurus flavimanus gloveri (Thomas) Callosciurus castaneoventris gloveri Thomas, 1921, Jour. Bombay Nat. Hist. Soc, 27, p. 502. Type.— Callosciurus castaneoventris gloveri, BM No. 13.9.13.3, young female, from Nagchuka, 10,000 feet, west Szechwan, col- lected August 14, 1908, by W. R. Zappey. Material examined, from Szechwan, China. — West of Fulin, 5000 feet (USNM), one; Ningyuanfu, 6200 feet (USNM), one; "Tanken" (USNM), one; 40 miles E. of Hokow (ANSP), one; Hokow (=Nag- chucka) (ANSP), one topotype, Nagchucka, 10,000 feet (BM), one topotype; "Toloko," south of Muli (MCZ), one, (ANSP), one; Na- chukar [Nagchuka] (MCZ), three topotypes, (USNM), one; Rama La Pass, 13,000 feet (MCZ), three; Mili [same as Muli] (CNHM), three, (USNM), three; Baurong (CNHM), four, (ANSP), one. Material examined, from Yunnan, China. — Yungning (CNHM), two; Yung-pei-ting, "N. 27° 10', E. 100° 50'" [Yungpeh] (BM), one. Material examined, from Tibet. — "U-long-si Gorge," 14,000 feet (USNM), two. Pelage color. — Color notes from material examined in the Museum of Comparative Zoology and Chicago Natural History Museum that includes three topotypes: The ventral pelage varies from Hay's Rus- set (XIV) through Vinaceous-Rufous to Ferruginous from individ- ual to individual, but is consistent on any one specimen from throat to tail and wrists to ankles, and is not in any instance divided by a mid-saggital agouti band, even incipiently. Dorsal pelage an agouti of about Mouse Gray (LI) at the more northern localities of Nag- chuka and Rama La Pass, Grayish Olive (XLVI) at Baurong, and Dark Olive Buff (XL) at the more southern localities of Mili and Yungning. Inner side of pinna is Apricot Buff (XIV) in the more northern material and Cinnamon Rufous to Vinaceous Rufous in the southern, but at Baurong there is just a hint of warm color in the gray. The pelage of the feet is colored like that of the dorsum. The tail is colored like the dorsum for its full length, and the tail hairs (at Yungning, Mili, and Baurong, at least) have six dark bands, and tips of Ochraceous Buff (at least in the north) . Discussion. — This rather variable subspecies shows relationship to intermedius to the west and bonhotei to the east in having undivided ventral pelage and uniformity in color of tail. It differs from both of them in having reddish ears and unblackened feet. Its dorsal pelage is also paler than that of intermedius and lacks the reddish infusion. f MOORE AND TATE: DIURNAL SQUIRRELS 151 Callosciurus flavimanus bonhotei (Robinson and Wroughton) Sciurus castaneiventris bonhotei Robinson and Wroughton, 1911, Jour. Fed- erated Malay States Mus., 4, p. 234. Type.— BM No. 8.8.11.25, an adult female taken at Chin Chien San, Szechwan, China, by F. W. Styan. Material examined, all from Szechwan, China. — Lian-feng-Kiang, Omei Shan (CNHM), one; Yaichowfu [Yachowfu], Omei Shan (CNHM), two, (USNM), one, (ANSP), three; "Ho Ni Pa" (CNHM), nine; Fi Shan Kwan (CNHM), eight; Lu Chang Pu (CNHM), two; Yung Cha Shan (CNHM), one; Chung Chiao [Chiang] Miao (CNHM), one; Hsiao Yang Chi (CNHM), one; Fu Fu Su (CNHM), three; Ninguanfu (USNM), one, (BM), one; "Tai Tsin Tang" (USNM), one; "Wan Nien Si" (USNM), one; "Lu Din" (USNM), one; Kia- ting (USNM), two; Mt. Omei (USNM), four; Suifu (USNM), two; "Shin Kai Si," Mt. Omei (USNM), five; Wan hsien (MCZ), one; "Chen Yen Say" (BM), one; "Yusu-Ching-Hsien" (BM), one; "Szechwan" (BM), two. There is a specimen referable to gloveri as well as one referable to bonhotei, both taken at Ningyuanfu and both showing some indica- tions of intergradation. As the map shows, there is one other point at which individuals bearing the characteristics of these two sub- species were taken very close together. Pelage color. — The agouti dorsal pelage of the bonhotei material at Chicago Natural History Museum is generally about Dresden Brown (XV) with the reddest variant being Sudan Brown (III). The tail hairs have six dark bands and sometimes seven. The ven- tral pelage is Burnt Sienna and is not divided in any of this series. Very little warm color can be found on the ears. Discussion. — Although somewhat variable, the material reported above appears to represent a distinct enough geographic subspecies which is richer in ventral pelage and darker in dorsal pelage than the adjacent gloveri, and also differs by having its feet slightly blackened and the reddish orange color almost absent from its ears. Resem- blance to hendeei of eight specimens from Fi Shan Kwan and two from Lu Chang Pu in characters of dorsal pelage is noted. However, these are the most northwestern and most southeastern of the col- lecting localities found. In the Lu Chang Pu two the tail has a long, black subterminal mark with the tip itself buffy or whitish, which is also like the northern hendeei known to us. In the other 19 ex- amples of bonhotei in the Chicago Natural History Museum series, 152 FIELDIANA: ZOOLOGY, VOLUME 48 however, difference from hendeei in dorsal pelage characters is fairly substantial. It is interesting to note that the northern limit suggested by the collecting localities mapped here may be close to correct. The Sage West China expedition of the American Museum of Natural History collected west and north out of Wenchwan for a total of about two months without taking squirrels of this species. They obtained series of other squirrels (Dremomys, Sciurotamias, and Tamiops) and good stands of coniferous forest and bamboo forest in the vicinity of their camps are evident in photographs shown to us by T. Donald Carter of that expedition. Most of their collecting was above 6000 feet. Callosciurus flavimanus styani (Thomas) Sciurus styani Thomas, 1894, Ann. Mag. Nat. Hist. (ser. 6), 13, p. 363. Macroxus griseopectus Milne-Edwards, 1874, Recherches pour servir a I'His- toire Naturelle des Mammiferes ... p. 305 (not griseopectus Blyth, 1847). Herpestes leucurus Hilzheimer, 1905, Zool. Anz., 29, p. 299. Herpestes albifer Hilzheimer, 1906, Abh. Ber. Mus. Nat. Heimatk. Magde- burg, 1, p. 177 (substitute for H. leucurus, preoccupied). Callosciurus caniceps canigenus Howell, 1927, Jour. Washington Acad. Sci., 17, p. 81. Callosciurus erythraeus woodi Harris, 1931, Occas. Papers Mus. Zool., Univ. Michigan, no. 228, p. 1. Types, all from China. — S. styani, BM No. 86.10.28.5, young female "from between Shanghai and Hangchow [about 100 miles], probably Kahing ..." [Kashing]; griseopectus, collected by A. David in Chekiang; leucurus, "2 Felle."; albifer, "2 Felle. In Hankau gefauft. 1905"; C. canigenus, USNM No. 241509 an adult male taken at Hayenhsien, Hangchow Bay, Chekiang, December 10, 1925, by Arthur de C. Sowerby; woodi, UMMZ No. 55827, adult male, taken at Lungtan in the Purple Mountains 25 miles east of Nanking, Kiangsu, south side of Yangtse River, December 14, 1923, by Norman A. Wood. Material examined, all from China. — Tehan (UMMZ), one; Wan Mts., 100 mi. S.W. of Nanking, Anhwei (UMMZ), five, (MCZ), two; Purple Mts., Lungtan, 25 mi. E. of Nanking, Kiangsu (UMMZ), three; Tunglu, Chekiang (AMNH), one, (MCZ), nine; Mokanshan, Chekiang (ANSP), three; Hai Yen Hsien, Hangchow Bay, Chekiang (USNM), two; Kangpu, near Hangchow, Chekiang (USNM), one; Nimrod Sound, coast of Chekiang (ZMHU), one; "Anhwei" (NR), 17; "Chien San," Anhwei (BM), three; Lushan Hills, Kiukiang, MOORE AND TATE: DIURNAL SQUIRRELS 153 Kiangsi (BM), one; "Hung Fa Chao," Shanghai (UMMZ), one; "Kin King," Chekiang (USNM), one. We are in agreement with G. M. Allen (1940, p. 633) on the above synonymy, which he has fairly and adequately discussed (pp. 634, 635). Pelage color. — The series from the Wan Mountains are dorsally an agouti of Light Brownish Olive (XXX) in most, but one is Buck- thorn Brown (XV) . There are five blackish bands on the individual tail hairs. There is an abrupt, short tip of black hairs present on the tail, but this is obscured to some extent by whitish or buffy tips even on the otherwise entirely black hairs. No orange is seen on the ears nor black on the feet. The ventral pelage is Light Ochraceous Buff to Pale Ochraceous Buff. Chin or chin and throat are agouti like the dorsum. There are two from the Purple Mountains with ven- tral pelage that is Pale Pinkish Buff to almost White, and their dorsal pelage is agouti of about Dark Olive Buff (XL) which is very slightly lighter than the lightest of the series from the Wan Mountains. The other Purple Mountains specimen, however, has blackish feet, dorsal pelage agouti of about Dresden Brown, and ventral pelage of Mo- rocco Red. All three from the Purple Mountains have chin or chin and throat agouti and ears concolorous with dorsum. Discussion. — Thomas (1894) distinguished styani from "castaneo- ventris" to the south by the ventral pelage being a "peculiar reddish cream-colour ('pinkish-buff' of Ridgway) instead of the rich rufous ('orange-rufous') of the older known form." A good deal more mate- rial has since become available, and although some variation toward ningpoensis characteristics is seen in styani, the difference is still as Thomas said "extremely striking." When the material is compared in series, a general difference is also evident in the color of the dorsal pelage, but there is some overlap among individuals. As Thomas said also, in other pelage characters the two subspecies seem to be quite alike. Harris (1931) proposed the name woodi for material from east of Nanking in the Purple Mountains which has lighter pelage in both dorsum and venter than previously found in styani. However, some years later he obtained another specimen (UMMZ No. 87081) from the same vicinity with dorsal pelage about Dresden Brown (XV) and ventral pelage that is about Morocco Red (I). This must be regarded as a most exceptional specimen for styani, and it certainly invalidates woodi as a subspecies. 154 FIELDIANA: ZOOLOGY, VOLUME 48 Since the subspecies styani is known from quite close to the south of the Yangtze Kiang in several places but as yet from no localities north of that great river, it seems possible that the lower course of the Yangtze in eastern China has been a barrier to northward spread of C. flavimaniLs. Callosciurus flavimanus ningpoensis (Bonhote) Sciurus castaneoventris ningpoensis Bonhote, 1901, Ann. Mag. Nat. Hist, (ser. 7), 7, p. 163. Sciurus tsingtanensis Hilzheimer, 1905, Zool. Anz., 29, p. 298 ("corrected" to tsingtauensis, 1906, Abhandl. Berichte Mus. Natur. Heimatk. Magde- burg, 1, p. 172). Types. — Sciurus c. ningpoensis, BM No. 86.10.28.3, old individ- ual from hills 30 miles from Ningpo, collected March, 1884, by F. W. Styan; tsingtanensis, ZMHU (Kreyenberg) No. 238, a male collected August 16, 1902, by Kreyenberg (at Nimrod Sound, coast of Che- kiang, according to G. M. Allen, 1940, p. 632) just south of Shanghai (Tsingtao is 400 miles farther north). Material examined, all from China. — Ningpo, Chekiang (AMNH), eight, (BM), seven, (MCZ), two, (CNHM), two; Fuching Hsien [Futsingl, Fukien (AMNH), 53, (MCZ), six; (CNHM), four; Yen- ping, Fukien (AMNH), six, (CNHM), two; Chungan Hsien, Fukien (AMNH), four; Kushan [Kaoshanshih] near Foochow, Fukien (USNM), one; "Peiliang," near Foochow, Fukien (USNM), one; Foochow [later Minhowl (CNHM), one, (BM), six; "Fukien" (NR), one, (CNHM), six; "Ah Chiung," Fukien (BM), three; "Mayhos," China (BM), one; Pootoo I. [Putu Shan], Chusan Archipelago, Che- kiang (BM), one. Pelage color. — The series of eight topotypes of ningpoensis in the American Museum of Natural History is consistently lighter colored in dorsal pelage than the majority of the 63 specimens from several localities in Fukien Province. While the range in ventral pelage color is very similar between the Ningpo and the Fukien material, the frequency of the palest color, about Apricot Buff, in the Ningpo is 50 per cent and in the Fukien three per cent. The richest ventral pelage color in the Ningpo eight is about Ferruginous, occurring in three specimens. Forty-one per cent of Fukien AMNH specimens have a richer pelage than Ferruginous, the richest color attained be- ing about Sanford's Brown. In 16 per cent of the AMNH Fukien series there is a longitudinal midventral band of gray agouti pelage entirely bisecting the red pel- MOORE AND TATE: DIURNAL SQUIRRELS 155 age. In 56 per cent there is an agouti wedge on the throat and breast dividing only the anterior portion of the red pelage. In 30 per cent there is no division of the red pelage. Of the eight Ningpo specimens five have an agouti wedge and three no division at all. The ventral pelage of the Tunglu series of nine at the Museum of Comparative Zoology is generally about Sanford's Brown with one about Orange Cinnamon (XXIX) and another Ochraceous Buff (XV), although the AMNH Tunglu specimen is quite as pale as styani, Light Ochra- ceous Buff at its richest. It should be said that material included in this subspecies seems to have six black bands to the fully grown-out tail hairs, besides the hardly noticeable blackish tip. And black-tipped guard hairs with three (as well as two, one, and no) light bands, occur commonly in the dorsal pelage. Discussion. — A pale subterminal band on the tail hairs is so long and noticeable that it gives a strong impression that the tail hairs are pale-buff tipped. The terminal hairs of the tail are each black for most of their length proximal to the mentioned subterminal light band. This creates a sub-sub terminal black spot in the tail. This spot is particularly noticeable from the under side and quite like that found also characterizing gordoni and michianus. It rarely shows any disposition to spread onto the sides of the tail as is seen in hendeei and castaneoventris. This black tail spot thus suggests closer rela- tionship of ningpoensis to michianus (and through it to zimmeensis and gordoni) than to the subspecies honhotei and gloveri north of the Yangtze Kiang. The above evidence of relationship finds striking support in the common possession by ningpoensis, michianus, zim- meensis, and gordoni, of the agouti wedge on the breast or a full- length mid ventral agouti band. North of the ranges of these four subspecies, gloveri and honhotei each have undivided red ventral pel- age and uniformly colored tail. To the south of the same four sub- species, the agouti wedge on the breast (as well as the full-length midventral agouti band) becomes scarce in castaneoventris and ab- sent in hendeei. Correspondingly, the tails of these two subspecies are distinguished from the black spot characterizing ningpoensis, michianus, zimmeensis, and gordoni. In hendeei and castaneoventris black is not confined to a terminal spot but extends anteriorly along each outer edge of the tail, progressively diminishing anteriorly. Callosciurus flavimanus thaiwanensis (Bonhote) Sciurus thaiwanensis Bonhote, 1901, Ann. Mag. Nat. Hist., (ser. 7), 7, p. 165. 156 FIELDIANA: ZOOLOGY, VOLUME 48 Sciurus thaiwanensis centralis Bonhote, 1901, Ann. Mag. Nat. Hist., (ser. 7), 7, p. 166. Sciurus thaiwanensis roberti Bonhote, 1901, Ann. Mag. Nat. Hist., (ser. 7), 7, p. 166. Callosciuru^ erythraeus nigridorsalis Kuroda, 1935, Jour. Mammal., 16, p. 281. Types. — S. thaiwanensis, BM No. 94.11.22.5, adult male from Baksa, Formosa, collected October 20, 1893, by P. A. Hoist; cen- tralis, BM No. 94.11.22.4, adult female from Lak Ku Li, central Formosa, collected June 29, 1894, by P. A. Hoist; roberti, BM No. 62.12.24.13, adult male from northwest Formosa, collected by Robert Swinhoe (skin only); nigridorsalis (not seen), Marquis Yamashina Collection No. 22, adult female from Riran, Taito, southeast For- mosa, collected August 1, 1932, by H. Orii. Material examined, all from Taiwan (Formosa) . — Teraso [Teraso Soan] (AMNH), two; Chip Chip (AMNH), three; Mt. Arisan, 8000 feet (MCZ), one, (BM), four; Kagi District (CNHM), three; "For- mosa" (UMMZ), three; Tainan (UMMZ), one; Racu Racu [Raku- raku] Mountains, 7000 feet (BM), two; Tapposha [Tappan-sha], Central Formosa (BM), five; "Ho Ho Mountain," 5000 feet (BM), one; "Lak-ku-H," Central Formosa (BM), three; Baksa (BM), one; "Lau Long" (BM), one; South Cape of Formosa (BM), two; Nan- Tou Hsien, Wu-Sheh (AMNH), one; Ping Tung (AMNH), one; Taipei Hsien, Wu Lai, 16 miles south of Taipei (AMNH), five; "I-Lan Hsien," Chiao-chi (AMNH), two; Taipei (AMNH), one; "Taipei Hsien," Ping Ling, Taipei (AMNH), five; I-Lan Hsien, I-Lan, Taipei (AMNH), two. Discussion.— Bonhote (1901) described three Formosan forms (of flavimanus) from only three localities and apparently very few, if any, more than three specimens. Furthermore, the localities from which Marquis Nagamichi Kuroda (1935, pp. 280-282) reported specimens in support of Bonhote's concepts are exceedingly few: one for roberti, two for centralis, one for thaiwanensis, and two for the subspecies he himself proposed, nigridorsalis. Kuroda did have, and enumerate, at least modest series from these localities, and does to some extent describe the variation noted in his series. The material that we have been able to examine, however, does not support all of Kuroda's findings. The only Baksa we find is about 95 miles north of the southern tip of Taiwan. We presume this Baksa to be the type locality of thaiwanensis. This type locality is farther north than the entire known range of nigridorsalis, although Kuroda says that thaiwanen- MOORE AND TATE: DIURNAL SQUIRRELS 157 sis "is found only in the southernmost parts of the island" implying south of the range of his nigridorsalis. If Kuroda's implication is correct, thaiwanensis would appear to be restricted to the peninsula of the southern cape, failing by 40 or 50 miles to reach its own type locality. Kuroda (1935) did not establish whether thaiwanensis, in his restricted concept, may in fact extend from the south peninsula westward along the south coast and then northward toward Baksa. The specimen from Ping Tung, having a red venter, strongly sug- gests that the gray-bellied population does not so extend westward and northward. If the locality we find for Baksa is the correct one, the type is therefore but a variant specimen from well within the geographic range of the centralis population. This concept of the type is supported by the other Baksa specimen in the British Mu- seum which one of us (Tate) found to be like the centralis population. The above four specimens from Teraso and South Cape support Kuroda's finding that a population exists on the South Cape in which the venter is consistently (May, November, January) all gray or has but small patches of reddish at the bases of the limbs. The greatest known distance between two localities from which only the grey-bellied south cape population is known, is between the "South Cape" and Koshun, ten miles. The distance between the only two known localities for nigridorsalis, Taito and Daibusan, is thirty miles. Only one specific locality has been attributed by a taxonomist to the dorsally reddish subspecies roberti, that is, Taiheizan. This reddish infusion is quite evident in one male specimen from Taipei Hsien, Ping Ling, but absent from two other males taken there the same day. The reddish infusion is quite evident in two specimens from Taipei Hsien, Wu Lai, but not in four others from the same place. The alleged differences between all these subspecies may be geographically constant enough over a large enough range to merit recognition as respectable geographic subspecies, but it seems to us that the constancy in some cases and the range in all remains to be demonstrated. Diagnosis. — Bonhote (1901, p. 165) pointed out certain features that characterize all C. flavimanus on Formosa, and which may be altered to read: (1) Hairs of the dorsal pelage have 2 or 3 light bands. (2) The ears are somewhat orange on the concave surface. (3) The tail is agouti (with 3 to 5 blackish bands on each hair) for the prox- imal half-length but blackens progressively out the distal half, where it also has on each hair a long, yellowish tip. (4) In addition, the variation in ventral pelage is like that described above for castaneo- 158 FIELDIANA: ZOOLOGY, VOLUME 48 ventris except that the percentage of each kind of variant differs. In a sample of 23 skins four have red throats and no, or virtually no, agouti in the midline, eleven have agouti throat and breast wedge, six have complete midventral bisection of the red by an agouti line, and two have no, or virtually no, red. (5) There is a strong tendency for the feet to be either black or blackish agouti. Numbers 1, 2, and 5 above are characters widespread in the spe- cies flavimanus and emphasize this relationship. Numbers 3 and 4 particularly emphasize relationship to subspecies castaneoventris, not to the geographically nearer ningpoensis. Callosciurus ferrugineus (F. Cuvier) Sciurics ferrugineus F. Cuvier, 1829 Table General et Methodique (appended to Geoffrey St. Hilaire and F. Cuvier, Histoire Naturelle de les Mammi- feres, 3, p. 238). Sciurus keraudreni Lesson, 1830, Centurie Zoologique, pi. 1 and text. Types. — S. ferrugineus, not found at MNHN in 1951, collected by Duvaucel "from India"; keraudreni, not found at MNHN in 1951, taken in Pegu, Burma. Material examined, all from Burma. — Mt. Popa, 4960 feet (AMNH), seven, (BM), four; "Kodugwe," Pegu Yoma (AMNH), three; "Yetho River," Pegu Yoma (AMNH), two; "Owegoo Dis- trict," Pegu Yoma (BM), one; Chaunglon, Tkyaulla District (BM), one; Daingmhu, 200 feet, 40 miles north of Pegu (BM), two; Gok- teik, 2133 feet, northern Shan states (BM), one; Kaing River, S. Py- inmana (BM), two; Lawksawk State (BM), one; "East India" (MCZ), two; 700 feet, northern Zamaya Reservation, 70 miles north of Pegu (BM), one; 300 feet, southern Zamaya Reservation, 60 miles north of Pegu (BM), one; Rangoon (MNHN), one, (BM), five; "Tankton," Rangoon (BM), one; Definition. — Callosciurus ferrugineus is here regarded as mono- typic (but including a named form keraudreni which may prove to occur west of the lower Irrawaddy and to be a good subspecies) and occurs only in Burma and principally between the Sittang Valley and the lower Irrawaddy. See the mapped distribution in Figure 14. Diagnosis. — The dorsal pelage of Callosciurus ferrugineus is all a glossy, rich, dark red except that it darkens to blackish on the feet, and that in the tip of the tail there is a small tuft of white hairs. The ventral pelage is entirely a lighter red than the back. These char- acters distinguish ferrugineus from all other species of Callosciurus. MOORE AND TATE: DIURNAL SQUIRRELS 159 Relationships to other species. — It is obvious that the wholly red squirrels, ferrugineus, which live in the forests of the Pegu Yoma of Burma above Rangoon (see fig. 14) very closely resemble the wholly red squirrels of Thailand, Cambodia, and Laos. Nor can it be re- garded as surprising that earlier authors (Ellerman, 1940; Ellerman and Morrison-Scott, 1951) have placed them in a single species. Their present ranges as known to us from collected specimens, are some 200 miles apart (see fig. 14) and separated by one great river barrier, the Sal ween. (Geologically earlier these separately evolving red squirrel populations must have been separated also by the com- bined Irrawaddy and Sittang rivers, as will be discussed in detail below) . The orange-footed form C. erythraeus sladeni of upper Burma closely resembles the orange-footed C. /. flavimanus of Laos and Viet- nam and is separated from it by more miles and one more barrier river, the Mekong, than C. ferrugineus is from the all red [C. finlay- soni] menamicus of northern Thailand. But since C. e. sladeni and C /. flavimanus are shown above to be separately evolved, one might anticipate something of the same sort of separate origin of the all red ferrugineus from that of the all red menamicus, williamsoni, and cinnamomeus. Chhibber (1934) reports that the Chindwin and Irrawaddy were formerly quite separate rivers and that a small lateral branch of the Chindwin must have by head erosion performed an act of stream piracy which diverted the whole upper Irrawaddy north of Manda- lay westward into the Chindwin. The former lower course of the Irrawaddy River was clearly down the Sittang Valley almost straight south from Mandalay to the sea. See, for example, de Terra's (1944, p. 71) sketch map of the physiography of upper Burma. Chhibber (1934, p. 20) implies that this piracy occurred after certain geological events which "characterised the close of the Tertiary ..." Prior to the piracy performed by the Chindwin, then, the Chind- win and Irrawaddy held between them a narrow strip of land con- sistently about 100 miles wide and at least 650 miles long, from the Tibetan Plateau to the sea. In the upper 50 miles of this strip the squirrel species Callosciurus erythraeus is a gray agouti dorsally and red only on the venter (intermedius) . Southward from there the spe- cies descends from the mountains to forests of the wide valleys, and the pelage of the rostrum, feet, and tail are red (sladeni). The south- ernmost known specimens of sladeni, from Yin (about 22° 43' N.), are very red. 160 FIELDIANA: ZOOLOGY, VOLUME 48 It seems a most reasonable hypothesis that in the 400-mile length of this inter-riparian strip remaining south of Yin, the population of these squirrels became redder until entirely red. When the stream piracy was accomplished by the Chindwin, then, the wholly red pop- ulation would have become cut off completely from contact with any population with which it could interbreed. It seems probable that enough further differentiation has taken place in ferrugineus during this isolation that interbreeding would no longer take place if natural access to the adjacent population of sladeni were restored. No en- tirely red specimens are known from north of the new river barrier. While the stream piracy barred ferrugineus from spreading to the north, it opened an area to the east across the empty former bed of the Irrawaddy. It is interesting to note that ferrugineus has spread across this barrier very little. Of the 13 collecting localities from which we have examined specimens, only two are across this former barrier: Gokteik and Lawksawk. No specimens which might be in- tergrades between ferrugineus and either shanicus or phayrei are known to us, although specimens of all three forms are now known from the single collecting locality of Gokteik. No evidence at all is known that C. flavimanus shanicus has crossed the former barrier to the west. Possibly its ecology is too specialized. Better details from the field on both geographical and ecological distribution of ferru- gineus, phayrei, shanicus, and janetta on both sides of the broken barrier would be interesting. Pelage color. — The dorsal pelage is generally a glossy Morocco Red, mid-dorsally deepening to about Claret Brown. The tail is Maroon, and within its tip a small cluster of white hairs regularly occurs. Ventral pelage is also red but a little lighter, about Mahog- any Red in most specimens but as light as Burnt Sienna in some. The dorsal pelage color extends onto the feet but darkens near the digits which are almost black. It is the blackish feet and white tuft of hair in the tip of the tail that distinguish ferrugineus from the all red squirrels of the species finlaysoni. Discussion. — Lesson's plate of keraudreni indicates a much greater length of the white tail tip than is the case in ferrugineus. There is in the British Museum a specimen. No. 79.11.21.644, which is labeled "keraudreni." It is from Arakan in west Burma, and this may pro- vide a notable westward extension of the species. Almost two inches of the tip of its tail are white. It thus agrees substantially with Les- son's plate and differs somewhat from true ferrugineus in which the white tip is very much smaller. Confirmation of the occurrence of MOORE AND TATE: DIURNAL SQUIRRELS 161 dark-footed, all red, squirrels occupying a geographic area east of the present lower course of the Irrawaddy in forest of the Arakan district, especially if the long, white tail tip is characteristic of it there, would justify recognition of keraudreni as a subspecies of fer- rugineus. Nothing that the writer has seen in southern material of C. eryth- raeus erythrogaster suggests intergradation farther south with a red subspecies. However, no barrier is known to prevent such inter- gradation now or formerly, and the area in the Arakan Yoma from which no specimens have been studied and whence evidence of inter- gradation might yet be obtained, is quite large (fig. 13). If there is a geographic population of red squirrels south of the range of erythro- gaster, its true relationships will remain uncertain until someone can show evidence that it does or does not intergrade with erythrogaster. Lacking such evidence, we are assuming that if a red population does live there, it became transferred west of the main barrier of the Irra- waddy in the delta by a shift of the main flow to a more eastern chan- nel, and subsequent silting up of the old channel until it could be crossed through vegetation by squirrels. We recognize that species level differentiation of C ferruginens from C. erythraeus is tenuous and hypothetical. Field observations by an experienced field collector are available: "On Mt. Popa this species only occurred in the thick jungle on the higher slopes of the mountain. Seen from a distance in a tree it ap- pears black, the white tail tip showing up conspicuously." (G. C. Shortridge in Wroughton, 1915a, p. 473.) Callosciurus finlaysoni (Horsfield) Definition. — The species Callosciurus finlaysoni is constituted by subspecies finlaysoni, folletti, trotteri, frandsoni, alhivexilli, harmandi, germaini, nox, cinnamomeus, annellatus, williamsoni, menamicus, sin- istralis, bocourti, and boonsongi (and see their synonymies). This species occupies the forests of Thailand, Cambodia, and the part of Laos along the Mekong as shown in Figure 14. Diagnosis. — There are populations of this squirrel with entirely white pelage, other populations with entirely black pelage, and still others with entirely red pelage, and there is no single character now known to be diagnostic for the species. Table 8 shows the dimen- sions of some type specimens of forms here included in the species finlaysoni, and the color characters are discussed in the subspecies accounts. 1 ^ Q3 / Si { 1 + 22-H + 20" N P' o> THAILAND ir^fl _r.©<^ 2^^4y^iV^^£4^ J5THMU5 OFKRA lOO'E Fig. 14. Geographic distribution of two species, the Pegu red squirrel, Cal- losciurus ferrugineus, A, and the Thailand tree squirrel, Callosciurus finlaysoni, B to P, as plotted from material examined. Subspecies of finlaysoni: B, finlaysoni; Cfolletti; D, trotter; E, frandseni; F, albivexilli; G, hardmandi; (off the S. E. corner of the map about 80 miles SSE on a small island is) germaini; I, nox; J, cinna- momeus; K, annellatus; L, williamsoni; M, menamicus: N, sinistralis; 0, bocourti; and P, boonsongi. 162 163 FIELDIANA: ZOOLOGY, VOLUME 48 Relationships to other species. — There is in Thailand, Cambodia, and Laos a complex of squirrels so extraordinarily differentiated that their variation has to be seen to be believed. Judging by collections, various local populations of it exhibit pelage that is entirely white, other populations entirely red, others entirely black, black above with white below, and other more complicated patterns. About 28 to 30 subspecies names have been applied to color variations of this presumed species. After having studied the outstanding collection of squirrels of this complex in the United States National Museum listed in the follow- ing pages, and having plotted as carefully as we could the many col- lecting localities of it, we feel that this has substantially advanced understanding of these squirrels, but that a great deal of mystery remains to be dispelled. We feel that we have improved knowledge of this complex not by discovering what its true relationships are, but by showing better how some of the problems need to be attacked locally in greater detail. Thus, even though we accept finlaysoni here as a species, it is pro- visionally and with some serious reservations. Its red subspecies williamsoni may quite well intergrade with C flavimanus flavimanus, but evidence that it does is scanty (see figs. 13 and 14). There are indications that C. finlaysoni sinistralis crosses (or intergrades?) with C. flavimanus thai, but just north of the range of thai, sinistralis seems to occur sympatrically with C. flavimanus zimmeensis or to in- terdigitate with it in a substantial area where no evidence of inter- breeding has come to our attention. Callosciurus finlaysoni is one of the species reported earlier to possess but two pairs of functional mammae (Moore, 1961a, p. 14) before the present revision of this species was accomplished, and it may be worth mention that no exceptions to this were observed in the larger series at the United States National Museum during this revision. The following two records of brood size hint that in finlay- soni brood size may also be small. Bonhote (1901b, pp. 53, 54) notes one specimen from above Uttaradit on the Mae Nan that would now be known as subspecies menamicus, which was taken "pregnant with two young" on April 3, 1900; and another specimen which would now be known as subspecies sinistralis which was "pregnant with two young" when taken on February 3, 1900, at Kamphaeng Phet on the Mae Ping. This species finlaysoni is for the most part a tree squirrel of the lowland forests. It is rather small, and it shares nearly all of its «a9 I s a 9< CQ CO i2'5fe '-ioc rt g •jra> 00050(M(MC(M-*rHCgO(MCDeO«ON(M«0 -t-t- • bo ^ fjrt e<3(MOJecoo":i«D"rj«c>"^Ti IM O "3 • • <£> • CO (M (N -rj" (M 00 t- 05 - - t- t- ■* O ,ii >^"oS O500505-«J<(M - -(M -i-IIM-*C0Oi-li-li-HOu:' • • • -t^i-H '^ i_3 ^ i CQ 00 I ffi ns O w 41 -H .t3 0 «>t~--rJ<-^t-Tjiit>05CD00(Micooo-*ooiraT)<«Dt-Ti'005omoooo50 tS ;Ci00t-t-Oi-lOO»C-«Di-l»Oe0t^00OIM00->^05t-(Mi-l E-( i-fi-ti-li-IC0050 (M050500C0. Eh a> B o S 5 PQ < CQ ^^& eooocji-HtN^oeci-Hoooeoco-^iniifit-oooiOi-ii-ias • ■C9(h OJOSOSOSOOOOOSOOSOOOlOOOOSOiOiOOOOS • rtrl l-Hl-ll-Hl-ll-ll-l ,_|^rt §^ -u^ ■^c»o©'-(OJoa>ojrHoooi-iO'^ot-ot- bOTa C5 O00©O0000O05a>O0JOO©OO05OOOOOO (UPQ'-H 1— li-H 1-H rH i-HlHl-tl-Hi-l i-(t-Ii-I 1— ItH j3 0) 41 -tJ^J COOt-U3T}i5505i-IOOiOOD3lO(NaJt-«OlO bO^ I S e<5Tjoooooo»oeo«OiH,-i^rH;ocot-t-eo«DcoeciM(Mc^eoeo— li— I'-lrH ,H»-lrHi— Ii-Ht-Hi-Ht-Ii— li— (I— li— (t-HtHi— (r-lTHi-H MM QjZ i-HtHT-l»-lTHT-lr-(iHi-lT-lTHi-lT-lrHi-liHT-lTHTH.-lr-ti-li-H -1^ a -2x3 c i^OCS '-l'-l'-l(M00O(Mi-H(M(M(M^(M(M03(M»-lir0(M'^(MO00 ^ ^+^0) (^ac<^c<^(^^l-^(^a<^J<^^<^^(^^<^^(^^(^^(^^l^^(^^l^^(^^(^^<^^(^^l^l<^^ 5 m >> _H+j oe<5t>eoio-«tt--^0(M-rH-«iiosOieoNC? n ffi 186 MOORE AND TATE: DIURNAL SQUIRRELS 187 Callosciurus caniceps (Gray) Definition. — Callosciurus caniceps is a species that includes sub- species concolor and adangensis that are extraterritorial, being entirely south of the Isthmus of Kra, and himaculatus, domelicus, and caniceps in the Indochinese Subregion. See also the synonymy of each of these. The range of the species caniceps is shown in Figure 15. Diagnosis. — Within the Indochinese Subregion this species is dis- tinguished from all others by the combined characters of cool gray ventral pelage and a short, abruptly marked off, intensely black tip to the tail. Relationships to other species. — The range of this tree squirrel is from Malaya north to the Salween River in Burma and to 19° 25' N. latitude (Doi Chieng Dao) in northern Thailand by our records. It is also distributed eastward across Thailand to the Mekong, but curiously we find no record of it in eastern Thailand south of about 14° north latitude (Ban Kabin Buri) nor any record from Cambodia. Throughout its range species caniceps is sympatric either with species flavimanus or with finlaysoni. It is usually more of a lowland species than is flavimanus in the regions in which these two are sym- patric, but where the two tend to occur together in generally low country caniceps is the larger animal. The species finlaysoni occurs with caniceps in the lowlands throughout central Thailand, and here also caniceps seems always to be the larger. See the accounts of species phayrei and inornaius for relationships of species caniceps to these. Table 9 provides some body and skull dimensions of 23 of the type specimens of named forms that belong in this species. Callosciurus caniceps concolor Blyth [Extraterritorial] Sciurus concolor Blyth, 1855, Jour. Asiatic Soc. Bengal, 24, p. 474. Sciurus lancavensis Miller, 1903, Smithsonian Misc. Coll., 45, p. 16. Callosciurus erubescens Cabrera, 1917, Bol. Real. Soc. Espan. Hist. Nat., 17, p. 518. Callosciurus concolor telibius Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 121. Types. — Sciurus concolor, "From the vicinity of Malacca," In- dian Museum (not seen); erubescens, MNCN No. 1.669 adult ob- tained in Selangor in May or June of 1900 by J. Waterstradt (not seen) ; lancavensis, USNM No. 104390, young male taken on Lankawi Is. December 1, 1899, by W. L. Abbott; telibius, BM No. 20.12.4.67, old male taken on Pulau Telibun, Trang, S.W. Siam, Jan. 2, 1917. 188 FIELDIANA: ZOOLOGY, VOLUME 48 Material examined, from Malaya. — Straits Settlements [Malacca] (MCZ), one, (UMMZ), three, (BM), one; Kuala Lumpur (USNM), one, (BM), one; Gin ting Bidai, 2300 feet, Selangor (BM), one; "Pahang River" (BM), one; Semangko Pass, Selangor-Pahang boundary (BM), one; "Panjum" Kuala Lipis, Pahang (BM), two; Telom River, 400 feet (BM), four; Gunong Ijau, 4700 feet, Perak (BM), one; Maxwell's Hill, 3600 feet, Perak (BM), one; Temengeh [also Temongoh and Tumangoh], 400 feet, upper Perak (BM), one; Ulu Selama [also Ulu Selamar], Perak (BM), five; Sungei Lebeh, Kelantan (NR), one; Kroh, 1100 feet, upper Perak (BM), nine; Gunong Gajah, Kedah (BM), one. Material examined, from lower peninsular Thailand.— Bangnara [Bang Nara = Narathiwat] (USNM), 16, (MCZ), one, (AMNH), one, (CNHM), one, (UMMZ), one; Lankawi Island (BM), 12, (USNM), four; Bukit Patani (USNM), two; Patani (BM), two; Biserat, Jalor (BM), five; Ban Sai Kau [Nawngchik District] (BM), two; Singora [Songkhla] (USNM), two; "Pak Bayoon" (USNM), two; Nakon Sritamarat [Nakhon Si Thammarat] (USNM), eight, (ANSP), one, (CNHM), one; Pulo Telibun (BM), three. Original description. — "Lower-parts dull ash-colour: the rest griz- zled throughout with black and dull ruddy-ferruginous; the latter somewhat brighter on the middle of the back, croup, and upon the tail, which last is conspicuously ringed with black and dull ferrugi- nous, and has a black tip mingled with hoary- white." Diagnosis. — The above description fits quite well the material in the United States National Museum. The diagnostic characters are (1) the reddish suffusion of color occurring on the middorsum but not on the sides of the neck and body, (2) the actual absence of a sharply marked off, quite black tip of the tail, and (3) the cool sil- very gray color of the venter. Discussion. — It may be seen that material from the northwest edge of the range of this subspecies from Pulo Lankawi (named form lancavensis) and Pulo Telibun (named form telibius) and the north- east edge at Nakon Si Thammarat is intermediate between concolor and bimaculatus because it has the sharply marked-off, black tip of the tail, but it is listed here for convenience because it is otherwise like concolor. See Figure 15. Habits. — Near Kuala Lumpur Harrison and Traub (1950, p. 339) tabulate data showing that Callosciurus caniceps was frequently taken in the forest, was observed in numbers in the town of Kuala MOORE AND TATE: DIURNAL SQUIRRELS 189 Lumpur, and was observed in abandoned clearings on the edges of country settlements, but that it was not found in the mountain forest about 5000 and 6000 feet of elevation on Gunong Brinchang near Cameron Highlands. On the other hand Callosciurus flavimanus was not taken in the lowland forest about Kuala Lumpur, nor in the town, nor in abandoned clearings but was observed in the above mountain forest. They also report of caniceps, "This squirrel is very common in secondary forest and in association with man. It builds nests in trees and feeds on fruit. Thirty-seven specimens were ex- amined from the three forest sites," Later Harrison (1954, p. 161) reported further on its food, "Although this squirrel is abundant in waste land only a few shot specimens have been examined [for food]. The stomachs contained fruit and other vegetable matter, and one out of three contained a proportion of insects." Robinson (in Bonhote, 1903, p. 21) comments from his own and Annandale's field notes: "This species is emphatically the village squirrel of the Patani States, and it is very exceptionable to find it otherwise than in the immediate proximity of dwellings. It is ex- ceedingly abundant in the cocoanut groves and orchards, and com- mits great ravages among the fruit, being particularly destructive to the jack fruit or nangka (Artocarpus integrifolia). It is commonly seen on the trees in the early morning, up till about 9 a.m., and after about 4 p.m., and in the heat of the day remains hidden in the crowns of the palms, where it also forms nests similar to the drey of the Brit- ish species. In South Perak, if it occurs, it must be very rare, and we never saw a specimen, but in the neighbourhood of Kuala Lumpur it, or a closely allied species, is fairly abundant. An entirely black variety was seen at Biserat on several occasions." Audy and Harrison (1953, p. 10) comment "These . . . squirrels are abundant in the understory layer of forest and secondary forest, and come down to ground level freely. They range into scrub and cul- tivated country, and are well known in the gardens of town houses." Callosciurus caniceps adangensis (Miller) [Extraterritorial] Sciurus adangensis Miller, 1903, Smithsonian Misc. Coll., 45, p. 17. Sciums concolor terutavensis Thomas and Wroughton, 1909, Ann. Mag. Nat. Hist. (ser. 8), 4, p. 535. Callosciurus moheius Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 122. Callosciurus moheius mohillius Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 122. 190 FIELDIANA: ZOOLOGY, VOLUME 48 Types. — Sciurus adangensis, USNM No. 104389, a young male taken on Adang Island, Butang Islands, southwestern Thailand, on December 14, 1899, by W. L. Abbott; terutavensis, BM No. 9.11.1.54, an old male taken from Telok Udang, Teratau Island, Thailand, March 7, 1909, by H. C. Robinson; moheius, BM No. 20.12.4.72, a female taken on Pulau Mohea (northern half), 7° 14' N. latitude about 30 miles east of Siamese coast, February 2, 1919; mohillius, BM No. 20.12.4.75, a male taken from the southern half of Pulau Mohea February 3, 1919. Material examined, all from islands off southwestern peninsular Thailand.— Pulo Terutau (BM), nine, (USNM), eight; Terutau Is- land [Pulo Terutaul (BM), nine; Pulo Adang, Butang Islands (USNM), two; northern part, Mohea Island (BM), three; southern part Mohea Island (BM), three. Discussion. — This subspecies consists as presently known, of in- sular populations off the most southwestern coast of Thailand in the area of intergradation between concolor and bimaculatus. Their pel- age characters include the sharply demarked tail tip of bimaculatus and the unornamented side pelage and venter of concolor. Although they vary somewhat among themselves in pelage color from one is- land to another, the diagnostic bright buffy midline of the ventral aspect of the tail links them together and distinguishes them from both concolor and bimaculatus. Callosciurus caniceps bimaculatus (Temminck) Sciurus bimaculatus Temminck, 1853, Equisses Zoologiques sur la cote de Guine, p. 251. Sciurus epomophorus Bonhote, 1901, Ann. Mag. Nat. Hist. (ser. 7), 7, p. 272. Sciurus davisoni Bonhote, 1901, Ann. Mag. Nat. Hist. (ser. 7), 7, p. 273. Sciurus sullivanus Miller, 1903, Smithsonian Misc. Coll., 45, p. 17. Sciurus matthaeus Miller, 1903, Smithsonian Misc. Coll., 45, p. 19. Sciurus lucas Miller, 1903, Smithsonian Misc. Coll., 45, p. 20. Sciurus epomophorus milleri Robinson and Wroughton, 1911, Jour. Federated Malay States Mus., 4, p. 233. Sciurus concolor samuiensis Robinson and Kloss, 1914, Ann. Mag. Nat. Hist. (ser. 8), 13, p. 226. Callosciurus epomophorus nakanus Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 120. Callosciurus epomophorus mapravis Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 120. Callosciurus epomophorus panjius Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 119. MOORE AND TATE: DIURNAL SQUIRRELS 191 Callosciurtis epomophorus panjioli Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 120. Callosciurus epomophorus tacopius Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 121. Callosciurus epomophorus pipidonis Thomas and Robinson, 1921, Ann. Mag. Nat. Hist. (ser. 9), 7, p. 121. Callosciurtis epomophorus tabaudius Thomas, 1922, Jour. Bombay Nat. Hist. Soc, 28, p. 1067. Callosciurus epomophorus hastilis Thomas, 1923, Jour. Bombay Nat. Hist. Soc, 29, p. 377. Types. — Sciurus bimaculatus, RNH No. 13363, an adult male from the peninsula of Malacca; epomophorus, BM No. 85.8.1.192, old female, from Salanga Island, Thailand, collected March 4, 1879, by J. Darling; davisoni, BM No. 85.8.1.187, young female, from Ban- kasun, Tenasserim, Burma, collected June 20, 1877, by W. Davison; sullivanus, USNM No. 104377, old female, from Sullivan Island, Burma, collected February 1, 1900, by W. L. Abbott; matthaeus, USNM No. 111920, adult female, from St. Matthew Island, Burma, collected December 11, 1900, by W. L. Abbott; lucas, USNM No. 104385, adult female, from St. Luke Island, Burma, collected Janu- ary 20, 1900, by W. L. Abbott; milleri, BM No. 0.10.4.5, young male, from Trong, Thailand, collected February 20, 1896, by W. L. Ab- bott; samuiensis, BM No. 21.11.8.10, adult male, from Samui Island, Thailand, collected May 12, 1913, by H. C. Robinson and E. Sei- mund; nakanus, BM No. 20.12.4.43, old male, from Naka Island, Thailand, collected February 4, 1918, by H. C. Robinson and C. B. Kloss; mapravis, BM No. 20.12.4.51, adult female, from Maprau Island, Thailand, collected February 10, 1918, by H. C. Robinson and C. B. Kloss; panjius, BM No. 20.12.4.36, old male, from Pan- jang Island, Thailand, collected January 20, 1918, by H. C. Robin- son and C. B. Kloss; panjioli, BM No. 20.12.4.41, adult male, from Panjang Anak Island, Thailand, collected January 29, 1918, by H. C. Robinson and C. B. Kloss; tacopius, BM No. 20.12.4.56, old male, from Takopah Island, Thailand, collected February 16, 1919, by H. C. Robinson and C. B. Kloss; pipidonis, BM No. 20.12.4.53, adult female from Pipidon Island, Thailand, collected February 3, 1919, by H. C. Robinson and C. B. Kloss; tabaudius, BM No. 28.8.21.2, an adult female taken on Tavoy Island, Mergui Archi- pelago, Burma, October 21, 1921, by C. Primrose; hastilis, BM No. 23.4.10.6, adult male, from Hastings Island, Burma, collected Oc- tober 21, 1921, by C. Primrose. 192 FIELDIANA: ZOOLOGY, VOLUME 48 Material examined, from mainland of peninsular Thailand. — Tha Lo [Ban Tha Lo], Bandon (USNM), three; Bandon [Amphoe Ban Don=Ban Makham Tia] (USNM), two; Trang (USNM), eight, (ANSP), one, (CNHM), one; Ban Ta [Tha] Yai (USNM), one; Kao [Khao] Luang (USNM), one, (CNHM), one; "Waterfall," Trang (USNM), two; Kao [Khao] Chong (BM), three, (USNM), one; Kao [Khao] Soi Dao (USNM), two; Poonga [Phangnga] (MCZ), one, (BM), two; Tang Pra [Laem Ao Kham], Takuatung (BM), four; Taptien [Tapting], Trang (BM), five; Chong, Trong (BM), three; Gap Patalong, Trang (BM), one; "Lem Ma," Trang (BM), three; Nam Chuh, Pak Chan (BM), one; "Renong River" [Ranong] (BM), two; Klong [Khlong] Wang Hip, Tung Sawng [Thung Song] (BM), one; Mamoh, Pakchan (BM), two; Torsan [Ban Tha San], Chun- tawn [M. Chum Pon] (BM), one; Nong Kok, Grabi [Ban Nong Kok, Krabi] (BM), one. Material examined, from Siamese islands west of peninsula. — Tongka [Koh Phuket] (BM), eight; Telok Palas [Ban Ao To Nong, Koh Phuket] (BM), two; DeLisle, or Piam, Island (BM), four; "Pasir Raja Is." (BM), six; Pipidon Island (BM), two; Maprau Is- land (BM), four; Panjang Island (BM), two; Panjang Anak [Island] (BM), three; Naka Island (BM), three; Takopah Island (BM), three; Pulo Sarik, Tongka [Ko Sire, near Ko Phuket] (BM), five. Material examined, from Siamese islands east of peninsula. — Koh Samui [Ko Samui] (USNM), four. Material examined, from Tenasserim mainland, Burma. — Bok Pyin (USNM), one; Maliwun (USNM), two; Victoria Point (USNM), three, (BM), one; Sungei Balik (USNM), one; Tanjong Badak (USNM), two; Champang (USNM), one; Boyce's Point (USNM), one; Bankasoon (BM), 14 topotypes; Taroar [Tagoot], Malay Pen- insula (BM), one; "Kussoom," N.W. peninsula, south of Tenasserim (BM), one; Tenasserim Town (BM), one. Material examined, from Mergui Archipelago, Burma. — St. Mat- thew's Island (USNM), four; St. Luke's Island (USNM), one; Has- tings Island (BM), three; Sullivan Island (BM), two, (USNM), five; Clara Island (USNM), one; James Island (USNM), three; Tavoy Island (BM), two; King Island (BM), six; "Nathay Mine" (BM), six; Kisseraing Island, 100 feet (BM), three; "Sir John Malcolm Island," 100 feet (BM), two. Original description. — Temminck's species bimaculatus clearly be- longs here, for his description of it nicely provides the pelage charac- ters diagnostic for this subspecies: "The upper parts of the head. MOORE AND TATE: DIURNAL SQUIRRELS 193 neck, all the back and the tail as far as the tip bear a . . . pelage regu- larly ringed with ashy and black . . . the sides of the neck, the upper part of the legs and flanks are of the red of rust; the lower parts are of a whitish gray." Another very early specimen erroneously referred by Horsfield (1824, no. 7, p. 8) to Sciurus affinis Raffles, 1822, is sometimes {e.g., Ellerman, 1940, p. 355) treated as if it were the type of another affinis, even though Sciurus affinis Raffles is at the same time recog- nized as valid for a species of Ratufa. The description by Horsfield, and his specimen do clearly belong to what is now another genus, Callosciurus, and in the present species and subspecies, but Sciurus affinis Horsfield, 1824, is a primary homonym of Sciurus affinis Raffles, 1822, and is permanently rejected (Mayr, Linsley and Usin- ger, 1953, p. 312), Diagnosis. — The present subspecies is characterized by being gray where concolor is reddish (i.e., on the middorsum) and reddish on the sides where concolor is gray. There is considerable variation in in- tensity of both the gray and the red, and some of it is geographic, but these differences adequately distinguish the present subspecies from the localities presently indicated. Callosciurus caniceps bi- maculatus also has a sharply black-tipped tail, and as mentioned above, this also occurs on material which is geographically intergrade between concolor and bimaculatus. The type specimen of bima^u- latu^ does also have a white tip distal to the black tip, but in the opinion of the one of us (Tate) who examined it, the white tip is an individual aberration. The reddish infusion of the color on the sides which characterizes bimaculatus extends over the side of the neck posterior to the ear and along the side of the body, posteriorly extending ventrally onto the groin. It may or may not be continuous from the neck across the shoulder to the side of the body and from the side onto the thigh. In areas where the red side color is most intense, it tends to be so continuous; in areas where the red color pales, the side of the neck and groin retain the most color almost as a large spot in each place. Koh Phuket is a focal geographic area for intensification of the red. Pelage color. — The venter is generally gray with an overlay of whitish hair tips, and a darker gray midline of agouti pelage, but in areas where the sides are intensely red, there may be a general suf- fusion of reddish invading the ventral pelage. Habits. — One highly qualified observer collecting this squirrel in most southern, peninsular Burma comments: "Very plentiful every- 194 FIELDIANA: ZOOLOGY, VOLUME 48 where, both around habitations and in the thickest forest. A thick- set rather clumsy looking squirrel. Wherever there are plantations this squirrel destroys large numbers of cocoanuts, by drilling a cir- cular hole in the side and extracting the contents. Weight. 10-14 ozs." (G. C. Shortridge in Wroughton, 1915b, p. 713.) Discussion. — In the collections of the named forms of the main- land north of subspecies concolor that are available in the United States National Museum, we laid out 16 specimens of "davisoni" from the Isthmus of Kra near the tenth parallel of latitude north (see Figure 15), and 14 specimens of "milleri" from Trong, represented in Figure 15 by the southernmost all-black dots. We could find no consistent difference between these either in color or size, both lots proving quite variable and the number of individuals showing inter- mediacy in any particular character often exceeding the number that show a discrete difference. Although St. Matthew's Island and Sullivan Island of the Mergui Archipelago are both outside the nine meter depth line (Fourth Sur- vey of India, U. S. Army Map Service, 1944, sheet N. C. 47) the named forms from them, matthaeus and sullivanus, are indistinguish- able from each other by comparison of the original hypodigms (5 and 6 specimens respectively) and are indistinguishable from the series of thirty specimens of bimaculatus from the adjacent mainland. St. Luke's Island and Hastings Island are satellite islands of St. Mat- thew's Island, each less than a mile from it and connected to it by shallows less than nine meters deep. The original hypodigm of two specimens of the named form from St. Luke's I., lucas, are distin- guishably darker than all of the mainland sample of bimaculatiLS, but not the St. Matthew's I. sample. In lumping the Hastings I. form, hastilis, with bimaculatus {= davisoni) , we follow Ellerman and Morrison-Scott (1951, p. 486). Callosciurus caniceps casensis (Miller) [Extraterritorial] Sciurus casensis Miller, 1903, Smithsonian Misc. Coll., 45, p. 19. Type. — USNM No. 104370, a young female taken on Chance Island, Thailand, December 28, 1899, by W. L. Abbott. Material examined. — Chance Island, off west coast of peninsular Thailand (USNM), four. The original hypodigm of five specimens, all we have seen, vary little among themselves. They have the ventral side of the tail strik- ingly paler than those of any specimens of geographically nearby MOORE AND TATE: DIURNAL SQUIRRELS 195 forms of caniceps and grayer than the somewhat yellowish ones of altinsularis, which are from High Island 110 miles to the north. Callosciurus caniceps altinsularis (Miller) Sciurus altinsularis Miller, 1903, Smithsonian Misc. Coll., 45, p. 21. Type.— VSNM No. 111975, an old female taken on High Island, Burma, December 31, 1900, by W. L. Abbott. Material examined. — High Island, Mergui Archipelago, Burma (USNM), four. The four specimens and the type laid out together, vary hardly at all among themselves, and do not differ ventrally from several of the himaculatus in the United States National Museum, but the dor- sal pelage as a whole is slightly but distinctly paler than any of the himaculatiLs and paler than any of the nearby insular forms except casensis (which, however, is much more brilliantly colored). Callosciurus caniceps fallax (Robinson and Kloss) Sciurus concolor fallax Robinson and Kloss, 1914, Ann. Mag. Nat. Hist. (ser. 8), 13„ p. 225. Type. — BM No. 21.11.8.9, an adult male taken on Pennan (or Pangnan) Island, Thailand, 9° 45' N. latitude off east coast, on May 30, 1913. Material examined. — Koh Pangan, Thailand (USNM), four. The material examined is consistent within the series and distin- guishable from bimaculatus by having paler tails (quite like those of casensis). It differs from casensis by quite dull pelage, the ruddiness of the sides being barely noticeable, and differs from altinsularis by much darker pelage on both dorsum and venter, and very much greater size. It seems a bit odd that the principal distinguishing character of fallax should be one shared only with two insular forms on the far side of the Malay peninsula, casensis 140 miles west and very slightly south, and altinsularis about 140 miles northwest. Callosciurus caniceps domelicus (Miller) Sciurus domelicus Miller, 1903, Smithsonian Misc. Coll., 45, p. 18. Sciurus bentincanu^ Miller, 1903, Smithsonian Misc. Coll., 45, p. 19. Types. — Sciurus domelicus, USNM No. 104381, an adult female taken on Domel Island, Mergui Archipelago, February 24, 1900, by 196 FIELDIANA: ZOOLOGY, VOLUME 48 W. L. Abbott; bentincanus, USNM No. 104383, an adult female taken on Bentinck Island, Mergui Archipelago, March 11, 1900, by W. L. Abbott. Material examined. — Domel Island (USNM), four; Bentinck's Island (USNM), three; Kisseriang Island (USNM), one. The sample of five specimens from Domel Island representing this form is a darker gray ventrally than any of the material of hi- maculatus examined (by Moore). Since the Bentincks Island mate- rial is also rather dark dorsally and intensely red laterally like the material from Domel Island, and is separated geographically by Domel Island from the mainland range of himaculatus, it is here in- cluded in the subspecies domelicus. The subspecies domelicus also occurs on Kisseriang Island which lies between Domel Island and the mainland. Callosciurus caniceps caniceps (Gray) Sciurus caniceps Gray, 1842, Ann. Mag. Nat. Hist. (ser. 1), 10, p. 263. Sciurus chrysonotus Blyth, 1847, Jour. Asiatic Soc. Bengal, 16, p. 873. Sciurus epomophorus inexpectatus Kloss, 1916, Jour. Nat. Hist. Soc. Siam, 2, p. 178. Sciurus helgei Gyldenstolpe, 1917, Kungl. Svensk. Vet. Handl., 57, no. 2, p. 34. Sciurus caniceps hehus Shamel, 1930, Jour. Mammal., 11, no. 1, p. 72. Sciurus epomophorus fluminalis Robinson and Wroughton, 1911, Jour. Fed- erated Malay States Mus., 4, nos. 3 and 4, p. 233. Types. — Sciurus caniceps, BM No. 213a (41.1817), an adult male from Tenasserim; chrysonotus (not seen), from Tenasserim Valley; inexpectatus, USNM No. 221557, a young female (deciduous upper fourth premolar still present) from Koh Lak, Pran, latitude 11° 45' N., southwest Siam, collected November 15, 1916 by C. B. Kloss; helgei, NR No. 71, a young male from Koh Lak, Siam, collected Novem- ber 29, 1914, by N. Gyldenstolpe; fluminalis, BM No. 7.11.13.17, an adult male taken at the Meping Rapids, Siam, August 8, 1907, by T. H. Lyle. Material examined, all from Koh Tau, Thailand (USNM), 12. Material examined, from the mainland of Thailand. — Khao Luang (ANSP), five, (CNHM), one; Hat Sanuk, near Koh Lak, Rajburi (BM), six; Pran [Prachuap Khiri Khan] (USNM), five; Koh Lak [Prachuap Khiri Khan] (CNHM), one, (USNM), five, (NR), one, (BM), eight; Sam Roi Gop [Sathani Sam Roi Yot] (USNM), four; Kwe Koi [Mae nam Khwae Noi] (USNM), one; Muang Kan Buri MOORE AND TATE: DIURNAL SQUIRRELS 197 [Kanchanaburi] (CNHM), one, (USNM), two; Ban Pong, Rajburi (USNM), one; "Nong Mong, Muang Krabin" [Ban Kabin Buri] (USNM), two; "Lam Ton Lang, Krabin" [Ban Kabin Buri] (USNM), two; Krabin [Ban Kabin Buri] (BM), three; Kao Lem [Khao Laem] (USNM), two; Pak Jong [Ban Pak Chong] (AMNH), four, (CNHM), one, (USNM), two; "Lam Klong Lang, Pak Chong" [Ban Pak Chong] (USNM), one; Muek Lek [Ban Muak Lek] (CNHM), one; Lat Bua Kao [Ban Lat Bua Khao] (USNM), one; Siken, near Korat [Ban Si Khiu] (USNM), one; Chainat [Muang Chainat] (BM), one; Pak- nampo [Ban Pak Nam Pho] (MCZ), one, (BM), two; Nakon Sawan [Nakhon Sawan] (BM), two; Bung Borapet [Boraphet] (USNM), one; Um Pan [Ban Um Phang=Ban Le Kathe] (AMNH), two, (BM), two; Longlung [near Ban Nong Pla Lai] (BM), one; Wung Pratart Farm, Kampengpet Prov. (CNHM), three; Klong Klung [Ban Khlong Khlung] (CNHM), five; Kampengpet [Changwat Kam- phaeng Phet] (AMNH), one, (CNHM), two, (BM), one; Sawan Kaloki [SawankhaIok=Ban Wang Mai Khon] (BM), three; Pak Koh (NR), two; Ban Hue Hom (NR), one; Mee Tan [Ban Mae Tan Nua] (BM), two; Mt. Chieng Dao (USNM), two, (AMNH), three; "Watpa" (ANSP), one; "Ubol Chanumon" (ANSP), one; "Nong Dom" (UMMZ), three; "Me Ping River" (AMNH), one; (BM), four; "Ban Kon" (NR), one; "Kao Phlyng" (NR), one; Phan Mt., Sakon Nakhon (USNM), two; Lomlo Mt., Ban Maeo (USNM), 17; "Phak Khinak Mt.," Dan Sai (USNM), one; "Nam Lang Mt.," Ban Khok (USNM), one; Kowjeen, Pak Tho, Rajburi (USNM), two; "Pukongchai," Korat (USNM), one; Hin Laem, Tra Khanun, Kanchanaburi (USNM), 16; Ban Klua Klang, Prachuap Kiri Khan (USNM), three; "Ban Sop Luak," Chiang Saen Kao, Chiang Rai (USNM), one; "Mushi Tar Shang," Korat (USNM), one; "Ban Hua Thanon," Klong Klung (USNM), six; Kowkat, Paknampho (USNM), 16; Kowkob, Paknampho (USNM), two; "Moung Wat Sy Tie," Nakorn Sawan (USNM), one; "Sretahn, Wung Sapueng," Loei (USNM), one. Material examined, from Burma. — Tavoy (BM), two; "Shan Mepa," Amherst (BM), one; "Tikotaw" Amherst (BM), one; Lakya (AMNH), two; Kawlichuang (AMNH), one; Lampha (AMNH), one, (BM), three; Moulmein (BM), three; Kawkareik (AMNH), one, (BM), one; Myawadi (BM), one; "Lothorgu," Myawadi (BM), one; "Thoungyin River, 1500 feet" (BM), three; "Thoungyin above Myawadi" (BM), three. 198 FIELDIANA: ZOOLOGY, VOLUME 48 Original description. — Gray's type description of caniceps is very bare: "Pale gray, grisled; back yellowish, beneath paler gray; tail long, gray, black varied, ringed, hair with three broad black bands." Pelage color. — Dorsal pelage color of American Museum Tenas- serim material is about Orange Rufus (II) and that of Doi Chieng Dao about Ochraceous Orange (XIV). Ventral pelage color is an agouti of about Deep Gull Gray (LI 1 1) in this Tenasserim and mid- dle Siam material, and about Light Gull Gray in the northernmost. Every American Museum specimen has a darker agouti, longitudinal midventral stripe about four to six millimeters wide. The throat, chin and under sides of legs are like the general color of the venter. The tail hairs have five black bands besides the black tips. The tips of the tails are all abruptly black. Dorsally the tail and all legs are agouti of about Mouse Gray (LI). The dorsal pelage of the feet, snout and ear tips is notably lighter and corresponds closely to the ventral pelage color. The bright orange color covers the back and sides, fading into agouti gray as it approaches the venter, on the crown, and on the proximal one- tenth of the tail. Habits.- — Gyldenstolpe (1914, p. 11) remarks interestingly, "In Siam this species was only common in the bamboo-forests in the North and seemed to live in rather high altitudes. In the bamboo- forests on the Korat plateau in Eastern Siam it was never observed." Discussion. — Although the named form inexpectatus is distin- guishable when one compares series of study specimens in a museum, it seems to occupy but a small geographic area between the ranges of subspecies bimaculatus and subspecies caniceps, the rainshadow area of peninsular Thailand north of about 11° North latitude. Since its observed characters in the material examined are but tran- sitional between those of bimaculatus and caniceps, it is here regarded as intermediate between those two subspecies and not fully belong- ing to either, but for convenience the records of it are lumped with caniceps. The named form helvus from Koh Tao is rather distinct, but the real interest which attaches to the Koh Tao material is that it shows relationship as close to caniceps as to the geographically nearer bimaculatus, and like the sample of "inexpectatus" is actually intermediate. Inclusion of it with subspecies caniceps emphasizes the interest which attaches to its close relationship to caniceps across a much greater water gap. It is interesting, too, that the sample from Koh Phangan fallax, which lies between the Koh Tao popula- tion and the bimaculatus of the nearest mainland is good bimaculatus, but shares with the Koh Tao sample a character which differs from mainland bimaculatus (pale underside of tail). \ MOORE AND TATE: DIURNAL SQUIRRELS 199 It certainly seems worth mentioning here that in the American Museum of Natural History material the roughly topotypical cani- ceps specimens from the heavy rainfall area of upper Tenasserim are not darker colored than ones from the much drier, rainshadow area of Siam almost directly east (localities Kawkereik, Lampha, Kaw- lichaung, and Lakya in Burma, versus Um Pang, Me Ping, and Kam- pengpet, Siam). The northernmost material that we have seen for caniceps, from Doi Chieng Dao, is notably lighter in both dorsal and ventral pelage, and is also slightly larger than this more southern American Museum material. If these differences could be shown to be consistent over a considerable geographic area, the name flumi- nalis Robinson and Wroughton would be available for the subspecies. The orange-colored dorsal pelage of Callosciurus c. caniceps has been reported to change to agouti gray color seasonally. The follow- ing records from examination of museum specimens in London (by Tate) and Chicago, Boston, and New York (by Moore) provide some evidence of such seasonal change: January, 12 orange, no gray; Feb- ruary, four orange, no gray; March, 12 orange, one changing, one gray; April, one orange, two changing, no gray; May, no data; June, one orange, three changing, nine gray; July, no orange, one changing, two gray; August, no data; September, no orange, one gray; October, one orange, two changing, no gray; November, three orange, one gray; December, two orange, one changing, no gray. It thus appears that in the November through April or May dry season the pelage is more often orange, but during the June through October season of rains, the dorsal pelage is more often gray. Some of the specimens that we have recorded above as "chang- ing" may possibly be only intermediate in some other sense. How- ever, one (CNHM No. 47332) in April was definitely molting the orange and replacing it with gray, and one December specimen (AMNH No. 54692) seems to be changing from gray to orange with- out showing a molt line, by the simultaneous appearance of orange in large patches throughout the length of the back. In this latter specimen the guard hairs of the remaining patches of agouti gray pelage seem under magnification quite as fresh as those of the in- cipiently orange blotches. Furthermore, the guard hairs of the incipi- ently orange blotches are agouti and only warmly colored on the two light bands (more deeply colored on the proximal one), and more of the orange color seems attributable to the thinner under hairs. How- ever, in a fully orange specimen of January 19th (AMNH No. 54707) O CO CO Eh 0) E o CO I b. •a o in cc oo t~ io t~ 05 O > > o +j o "oS ^!: ■" -fJ 73 ^ a-^ ID . CO a> g-s CI] Cl} e CD o-^ ^ S J> 200 MOORE AND TATE: DIURNAL SQUIRRELS 201 from nearby, the guard hairs are not agouti but are entirely orange excepting the thin, short, black tip and a short paler base, and there seems to be very little pelage wear (breakage of these guard hairs). In a fully orange March 3rd specimen, the pelage is as in the preced- ing specimen but shows wear (has the tips of a good many guard hairs broken off). These few superficial observations suggest that at the end of the rainy season, orange color may replace the black and white bands on the agouti guard hairs rather than orange guard hairs replacing black and white banded guard hairs by molt. Thus it may be that this tropical tree squirrel molts but once annually while accomplishing change in color twice a year, although it seems well established that the well-studied north temperate tree squirrels Tami- asciurus hudsonicus, Sciurus vulgaris, and Sciurus carolinensis are known to molt twice a year (each spring and fall). Callosciurus phayrei (Blyth) Sciurus phayrei Blyth, 1855, Jour. Asiatic Soc. Bengal, 24, p. 472. Sciurus blanfordi Blyth, 1862, Jour. Asiatic Soc. Bengal, 31, p. 333. Callosciurus griseimanus heinrichi Tate, 1954, American Mus. Novitates, no. 1676, p. 1. Types. — Sciurus phayrei, BM No. 62.7.16.7, an adult male taken at Martaban, Burma, by Major Phayre; blanfordi BM No. 63.5.- 9.9, an immature taken at Mt. Ava about 20 mi. S.W. of Mandalay; heinrichi AMNH No. 163466, an adult female taken at Maymyo, Mandalay, Burma, at 800 meters elevation on December 6, 1937, by Gerd Heinrich. Material examined, all from Burma, south to north. — Martaban (BM), one cotype; Thatone [Thaton] (BM), three; Pahpoon [Papun] northern Tenasserim (BM), two; "Bantam, Kiu Pang Valley," Sal- win Distr. (USNM), one; five miles east of Toungoo, 100 feet (CNHM), two; 40 miles north of Toungoo, 500 feet (BM), four; "Ta-ho, Kareni" (USNM), one; Mt. Ava (BM), one cotype; seven miles southeast of Mandalay (USNM), one; Maymyo, 800 meters (AMNH), six; Gokteik, 2100 feet (BM), three; Kokkoaing, 40 miles N.N.E. of Mandalay, 800 feet (BM), one; Ngapyiniun, 250 feet, opposite Kyaukmyaung, on Irrawaddy R. (MNHN), one, (BM), six; Pyaunggaung, 2800 feet (BM), three; Hsipaw, 1350 feet (BM), two; Se'en, 1400 feet (BM), five, (CNHM), two. Table 10 provides some body and skull dimensions of some of the type specimens of named forms that belong in species Callosciurus phayrei. 202 FIELDIANA: ZOOLOGY, VOLUME 48 Definition. — The species phayrei is accepted here as monotypic and is confined to the region between the valley of the Sittang and upper Irrawaddy rivers and Salween River as shown in Figure 15. Diagnosis. — The species phayrei has the following distinctive color characters of the pelage. (1) The rostrum, ears, crown, and side of neck are agouti gray like the dorsum. (2) The contrasting color of the manus stops abruptly at the wrist. (3) The tail is tipped briefly atid rather abruptly with black. (4) The ventral pelage varies from a rich orange to a very pale orange but is never red or gray. (5) All four feet are yellowish buff to pale orange. (6) Long, yellow bases of the ventral tail hairs provide a rich yellow midstripe 12 to 15 mm. wide along the under side of the tail. (7) There is usually at least a faint blackish band 20 mm. wide along (and within) the lateral mar- gins of the ventral pelage. (8) There are five blackish bands on the fully grown out tail hairs. The above characteristics distinguish species phayrei from the subspecies of other species that are geographically closest to phayrei, as follows: C. erythraeus sladeni by 1, 2, 3, 4, 5, 6, and 7; C. flavi- manus shanicus by 1, 4, 5, 6, and 7; C. flavimanus zimmeensis by 1, 2, 4, 5, and 7; C . ferrugineous by 1, 2, 3, 4, 5, 6, 7, and 8; C. c. caniceps by 4. 5, 6, and 7; C. pygerythrus janetta by 1, 2, 5, 7, and 8. Relationships to other species. — This species appears to inhabit the rain forest vegetation area from Martaban at the mouth of the Sal- ween River north to about Toungoo, and to occupy the strip of trop- ical deciduous forest along the escarpment of the Shan Highlands from about Toungoo north to at least fifty miles north of Mandalay (according to the map of vegetation zones of Burma by de Terra (1944, p. 80). On ecological grounds it seems very likely that the range of phayrei would extend up the long corridor of rain forest of the Salween River valley. Whether it also occurs throughout the Shan Highlands seems more doubtful for the Maymyo, Gokteik, Pyaungaung, and Se'en localities seem to lie along the southeastern margin of an eastward jutting of the tropical deciduous forest into the "mixed forest with grassland" of the Shan Highlands (de Terra, 1944, p. 80), and may not represent habitat typical of the Shan Highlands. Relationships between C. phayrei and C. erythraeus sladeni. — While our record of the distribution of phayrei from material ex- amined leaves large gaps where its occurrence remains to be ascer- tained, its range seems to be confined between two rivers, on the south and east by the Salween and on the west by the Irra- sourf(a{ii/^6E^ Fig. 15. Geographic distribution of three species, as determined from mate- rial examined: the Tenasserim squirrel, Callosciurus caniceps, circular symbols; the Shan Highlands squirrel, Callosciurus phayrei, black squares; and the Laotian squirrel, Callosciurus inornatus, triangles. Subspecies of caniceps: half black [extra- territorial], concolor; vertically divided [extraterritorial], adangensis; all black, bimaculatus; dotted circles, domelicus; and open circles, caniceps. 203 204 FIELDIANA: ZOOLOGY, VOLUME 48 waddy. Across the upper Irrawaddy phayrei seems to be ecologically replaced by sladeni as the tree squirrel of medium size in the forests in the broad fertile valleys and relatively low hills of this interri- , parian area of Burma. See map figs. 13 and 15. i Besides occupying what is apparently the same ecological niche on opposite sides of the Irrawaddy River phayrei and sladeni share a number of characteristics of the pelage: (1) The dorsal pelage of all four feet is colored like the venter. (2) The dorsal pelage of the body and proximal part of the tail are an agouti gray with three light bands on many of the guard hairs of the middorsum and with five black bands on the hairs of the tail at its midlength. (3) The tail is not agouti at its tip. (4) The ventral pelage is orange-red or orange but never divided down the midline by an agouti stripe. (5) The ears are the same color as the dorsum. The pelage characters which distinguish phayrei and sladeni are : (1) The specimens of sladeni almost invariably have the rostrum, and sometimes the crown back as far as the ears, colored like the feet; whereas the rostrum and crown of phayrei are as agouti gray as its back. (2) The color of the manus in sladeni extends up onto the forearm, meeting the agouti gray abruptly for half the forearm's length; whereas in phayrei the manus color terminates abruptly at the wrist. (3) In sladeni the tail has a rather long, reddish tip, but in phayrei the tail tip is short, fairly abrupt, and black. These three characters, in the absence of any known area of intergradation, sug- gest something more than subspecific difference between the two. Even though the evidence is excellent that the subspecies sladeni and haringtoni are conspecific, and although the pelage difference between sladeni and haringtoni appears to exceed that between sladeni and phayrei, the latter two are not known to intergrade: their differ- ences are great enough that they probably would not interbreed if brought together naturally, for example, by a shift of the course of the Irrawaddy and elimination of the old channel. (Intergradation between sladeni and haringtoni is apparently accomplished only through another subspecies and interbreeding might not occur at all between individuals of these two forms themselves if they could be brought together under natural conditions.) The Sittang Valley separates the range of C. phayrei from that of C. ferrugineus to a great extent, although the specimen of phayrei from Ava shows that phayrei has crossed the valley a short distance eastward and a specimen each from Gokteik and Lawksawk indicate that ferrugineus has crossed the valley westward. It is assumed MOORE AND TATE: DIURNAL SQUIRRELS 205 that this condition of Hmited or incipient sympatry has come about entirely since the capture of the headwaters of the "Irra-Sittang" by the "Chindwin-waddy" {i.e., the removal of the river barrier from the Sittang Valley). Their difference in pelage color is virtually total, ferrugineus being an all red species with some blackening on the feet and tail. This fact and those of sympatry with no evidence of intergradation demonstrates that phayrei and ferrugineus are sep- arate species. Similarities between C. caniceps caniceps and species phayrei. — From the material examined, it appears that the subspecies caniceps extends northwestward in typical form to the vicinity of the Salween River and stops abruptly there; and immediately on the other side of the Salween River the species phayrei apparently replaces caniceps ecologically. The lack of intergradation between these two forms and the niceness of their separation by the existing geographic bar- rier of the river are such that no previous student has regarded them as conspecific. Nevertheless, the number of evidences of close rela- tionship is great: (1) The back, sides and dorsum of the tail of phayrei are colored like those of gray phase caniceps. (2) The tail tip in phayrei is black like that of caniceps. (3) The head, ears, and rostrum of phayrei are gray as in gray phase caniceps. (4) All four feet are strikingly light in both forms, lighter than their respective ventral pelage. (5) The tail hairs have five black bands additional to black tips in both forms, and the guard hairs of the dorsum have two white or very pale buffy bands in phayrei as in the gray phase of caniceps. (6) The frontal projects forward as a thin process between the nasal and the premaxillary on each side, often to a distance equaling least width of nasal in many subspecies of Callosciurus; but it does not do this in caniceps or in the material representing phayrei in the American Museum of Natural History. (7) Subspecies caniceps, unlike most mainland forms of large Callosciurus, is strongly pro- odont, and the seven Maymyo specimens of phayrei are at least somewhat proodont. Distinctions between species phayrei and C. caniceps caniceps. — (1) The ventral pelage is Light Ochraceous Buff to Ochraceous BufT (in the Maymyo series) even approaching Ochraceous Orange on the underside of the hind legs, compared to the whitish over pale gray of caniceps, and lacks any vestige of the midventral agouti stripe of caniceps. (2) All four feet are dorsally Light Ochraceous Buff in phayrei (Maymyo series) but about Pale Gull Gray (LIII) in cani- ceps (Mt. Chieng Dao series). (3) The ventral aspect of the tail of 206 FIELDIANA: ZOOLOGY, VOLUME 48 phayrei reveals long, yellow bases of ventral tail hairs so that the tail has a Warm Buff midstripe 12 to 15 mm. wide beneath but is cool gray dorsally. In caniceps some faint yellow color shows in the dorsal aspect but ventrally virtually none. (4) In phayrei there is a 20 mm. wide blackish band on either side in the ventral pelage about 60 mm. long. This is very faint in the Maymyo series but also in three Gokteik, one Pyaunggaung, and three Se'en specimens in the British Museum. The others in the British Museum have the ven- tral stripes strongly marked and agree completely in appearance with the type. Both variants were thus taken at the Se'en and Pyaung- gaung localities and all in May. It is not apparently a seasonal or sexual difference, nor is it clearly a geographic variation. (5) The ears appear to be notably shorter in the Maymyo series of phayrei than in caniceps. (6) Subspecies phayrei lacks the seasonal orange dorsal pelage of caniceps. C. caniceps caniceps and C. phayrei apparently maintain these striking pelage color differences to the opposite banks of the Salween River which separates them, and in absence of any indication of in- tergradation, and on the degree of difference in pelage characters, they are here confidently regarded separate species. C. pygerythrus janetta is geographically adjacent to C. phayrei on the west and resembles it more closely than do any of the other sub- species of C. pygerythrus, excepting possibly the typical subspecies pygerythrus (which likewise is only west across the Sittang River from C. phayrei and resembles the rainforest population of phayrei almost as closely as janetta does the deciduous forest population). The close relationship these observations suggest, demands direct comparison of janetta with phayrei. Just as phayrei shares certain color characters with C. c. caniceps as its closest relative to the south or east, phayrei shares some also with C. p. janetta revealing it to be the closest relative to the west. These latter are: (1) All of the dorsal pelage is a cool agouti gray excepting hands and feet and tip of tail. (2) The dorsal pelage of all four feet is not agouti but is colored like that of the venter and con- trastingly lighter than that of the dorsum. (3) There is an abrupt short, black tip to the tail. (4) There is a narrow buffy mid ventral stripe along the tail. (5) There is some darkening of the ventral pelage in a broad band across the abdomen, tending to fade near the midline. The first three of these five characters are shared also by C. c. caniceps. 1 r MOORE AND TATE: DIURNAL SQUIRRELS 207 The important color differences between phayrei and janetta are: (1) There are five blackish bands to fully grown out tail hairs in phayrei, but four in janetta. (2) There is a flash mark on each hip of janetta, none in phayrei. (3) The hind feet of phayrei are nearly Warm Buff whereas those of janetta are but Cream Color. (4) The ventral pelage of phayrei is contrastingly richer in color than the mid- ventral longitudinal tail stripe, but in janetta they are about the same, or the venter is the more pale. (5) The side of the neck is pale, almost cream color, in janetta, but hardly lighter agouti than the nape in phayrei. There is an important general size difference between phayrei and janetta which is fairly well indicated by hind foot measurements of both these forms taken in the field by the same collector, Gerd Hein- rich. These are: janetta, 40, 40, 40, 43 mm.; phayrei, 43, 45, 46, 47, 48, 48 mm. C. phayrei has been taken at Ava west of the Irra- Sittang Valley and janetta east of it at Mandalay. Since there is at least this small area of sympatric distribution, then, and no evidence of intergradation in the available material, C. pygerythrus janetta and C. phayrei are here accepted as separate species. The form phayrei is separated from C. erythraeus atrodorsalis and C. e. zimmeensis by the apparently quite effective barrier of the Sal- ween River. Since phayrei also differs in color characters of the pel- age from both zimmeensis and atrodorsalis a great deal more than it does from C. e. sladeni or C. c. caniceps, it seems unquestionable that phayrei is of a distinct species, and no details for comparison with zimmeensis and atrodorsalis seem necessary here. C. phayrei has here been accepted as different at the species level from all tree squirrel forms to the south, west, east, and northeast of its known range that, like phayrei, ordinarily have only two pairs of functional mammae. It is separated from these other species to the south and west by the Salween River, and from a species to the northeast by the Irrawaddy River, and was formerly separated from two species to the east by a river barrier (the Irra-Sittang) . There remains only to ascertain the relationship of phayrei to the form im- mediately to the north and northwest. This is C. flavimanus shanicus which occurs in the area between the Salween and Irrawaddy rivers and has no barrier separating it from phayrei. The trenchant differences between phayrei and shanicus in pelage color are: (1) All four feet of shanicus are blackish agouti like its dor- sum, but those of phayrei are yellow or yellow-orange like its venter. (2) The ears of shanicus have the reddish rims, but those of phayrei 208 FIELDIANA: ZOOLOGY, VOLUME 48 are plain agouti like its back. (3) The ventral aspect of the tail of shanicus is as plain as the dorsal, but that of phayrei is ornamented by a longitudinal yellow midstripe. (4) The back of shanicus has a faintly expressed, broad, blackish, longitudinal band on the posterior two-thirds of the middorsum, but the back of phayrei is plain agouti like its sides, nape and crown. (5) The ventral pelage of shanicus is (somewhat obscurely) bisected by a longitudinal narrow stripe of agouti pelage, but that of phayrei has no such division. (6) the rostrum of shanicus shows more or less suffusion of rusty red, but that of phayrei is cool agouti like its crown. (7) The ventral pelage is generally a dull gray, but that of phayrei is a bright yellow-orange (sometimes with broad black margins extending between the fore and hind limbs). The first six of the above color characteristics of shanicus are gen- erally shared with C. f. gordoni, zimmeensis, atrodorsalis, thai, and other forms of C. fiavimanus to the east and south indicating a con- specific relationship with them while separating shanicus from phay- rei at the species level. Species distinction between these two is further indicated by both of them having been taken at precisely the same altitudes at three of the same localities, Maymayo, Gokteik, and Pyaunggaung, and there is a 100-mile north-south sympatry in their ranges indicated by mappable collecting localities. (See account of C /. shanicus for further details.) Thus, the form phayrei is clearly distinct at the species level from all other tree squirrels of its subgenus which surround it on every side, and if one accepts blanfordi and heinrichi as synonyms of it, phayrei is a monotypic species. Shortridge's field note on phayrei (in K. V. Ryley, 1914, p. 721) may shed light on the distributional relationship between phayrei and shanicus: "A particularly active species, its leaps from tree to tree almost rivaling those of Ratufa. Around Hsipaw town and Se'en, even more plentiful than . . . shanicus. Not observed at Maymyo." In contrast he found shanicus the most abundant squir- rel in Hsipaw State habitually near bungalows around Maymyo. What evidence we have suggests the possibility that C. f. shani- cus and C. phayrei may be in active competition with one another where their ranges meet or overlap and even that phayrei may be replacing shanicus in accordance with the "competitive exclusion principle" (Hardin, 1960). C ferrugineus also seems to be involved in this, but more locally and perhaps to a very minor extent making it three-way competition. MOORE AND TATE: DIURNAL SQUIRRELS 209 Gallosciurus inornatus (Gray) Macroxus inornatus Gray, 1867, Ann. Mag. Nat. Hist. (ser. 3), 20, p. 282. Callosciurus imitator Thomas, 1925, Proc. Zool. Soc. London, 1925, p. 502. Types. — M. inornatus BM No. 62.8.18.4, an old female from mountains in Laos, taken by C. Mouhote; imitator, BM No. 25.1.1.- 68, adult male from Thai-Nien, latitude 22° N. on the Song Koi, Tonkin, 300 feet, collected February 26, 1924, by H. Stevens. Material examined, from Annam. — Hoi Xuan (CNHM), two; "Muong Sen," prov. Vinh (BM), one, (MNHN), one; Phu Qui (BM), three, (MNHN), eight; Lao Bao (MNHN), one; "Ipuing" (MNHN), one. Material examined, from Tonkin. — Phong Tho, 1000 feet (AMNH), one, (CNHM), one; Bac Tan Tray, 700 feet (AMNH), one; Lai Chau, 500 feet (AMNH), two, (ANSP), one, (CNHM), one, (USNM), three; M. Mouen (AMNH), two, (ANSP), eight; Na Hai (AMNH), two; Ba Nam Nung (CNHM), one; Muong Mo (CNHM), 12; Muong Moun 1200 feet (CNHM), nine, (MCZ), one, (UMMZ), one; Muong Boum (CNHM), 11, (MCZ), two; Nong Lum (CNHM), one; Lieng San, 1500 meters (CNHM), six; Chapa (CNHM), four, (MCZ), two, (BM), 10; Pakha (CNHM), one, (BM), two. Material examined, from Laos. — Muong Yo, 2300 feet (CNHM), 14, (USNM), two; Nap^, 2000 feet (BM), four; Pasa (AMNH), one; Xieng Khouang (BM), one; Don Qua (AMNH), one; Muong Koa [Khoua] (USNM), one; Nam Khueng (MCZ), two; Phu Kobo (MCZ), two; Lo Tiao (MCZ), two; Col de Taloun (MCZ), one; Phong Saly 4400 feet (MCZ), one, (UMMZ), one, (CNHM), three; "Tha Ngon," Vientiane (CNHM), one; Vientiane (USNM), one. Systematic history. — Gray's original description of inornatus fits the squirrel known since 1925 as imitator, quite well in every way excepting the last item on the tail: "Fur olive-grey . . . throat, inner side of limbs, and [venter] pale bluish grey, washed with whitish ; feet like back; tail longer than the body and head, colored like the back, with elongated white- tipped black hairs at the tip; hairs of the tail yellow, with three black bands ..." One of us (Tate) examined the type in 1951, noted it to be very much like the type of C. pygerythrus stevensi, but did not compare it with "imitator," and photographed and measured the skull. There is no doubt from the skull that this is Callosciurus, and there is no other Callosciurus species known from Laos which the type of inornatus reasonably could represent. Badly worn tails sometimes have the ends of the hairs broken off enough to 210 FIELDIANA: ZOOLOGY, VOLUME 48 reduce the count of black color bands, and tails with the hairs not yet grown out to full length do not show the full number of black bands. Thomas (1925) in describing imitator from Tonkin did not com- pare it with inornatus Gray and seemed unaware of the existence of inornatus, for he remarked the peculiarity of its possible relationship to the C. caniceps complex and noted its "superficial resemblance" to C. pygerythrus stevensi. Robinson and Kloss (1918) had overlooked inornatus in their nominal list of the squirrels of the Oriental Region. Pelage color. — Ventral pelage generally Light Violet Gray from chin to wrists and ankles. In the 68 specimens at Chicago Natural History Museum there is occasionally a collar of agouti intruded into this ventral pelage and a wedge of it extending posteriorly on the mid- line or a midstripe of it bisecting the bluish ventral pelage. The chin pelage is almost invariably bluish gray. Excepting for minor indi- vidual variations of lighter or darker, there seems to be no other vari- ation in the ventral pelage of this great series excepting the only summer specimen CNHM No. 32381 from Tha Ngon, Vientiane, Laos. This exception has a rather general suffusion of agouti through- out the ventral pelage. (Since the Vientiane specimen is from an area on the edge of the known species range and rather distant from sources of other material examined, one wonders whether the varia- tion might be geographic. A perhaps similar condition in an AMNH specimen from Na Hai, near Dien Bien Phu, suggests that the vari- ation may be individual.) The dorsal pelage is agouti, with two or often three light bands on the individual hairs, and Deep Olive (XL) in color. It varies aston- ishingly little in the large series at Chicago Natural History Museum. (There is some reddish suffusion the length of the dorsum, however, in 10/12 of the Muong Mo series and all 11 from Muong Boum, but it is poorly developed in the Lieng Sen six and Muong Mouen eight and virtually absent from the Phong Saly and Muong Yo series.) The ears are the same as the dorsal pelage; the feet are only a little more contrasty agouti. The tails are colored like the dorsal pelage and regularly but sparingly black- tipped (50 or 60 mm. black on light tipped hairs) throughout. The tail hairs regularly have five blackish bands on the individual hairs when the hairs are fully grown out, but one AMNH specimen has six. Table 10 provides measure- ments of two specimens which give some indication of the dimensions of this animal. This is a smaller squirrel than Callosciurus erythraeus hendeei with which it is almost completely sympatric (and which also shows re- MOORE AND TATE: DIURNAL SQUIRRELS 211 markably little geographic variation in color characteristics) . Com- pare Figures 13 and 15. Diagnosis. — In describing imitator {=inornatus) Thomas noted its striking similarity to Callosciurus caniceps concolor far to the south in Malaya and to Callosciurus pygerythrus stevensi far to the northwest in Assam. C. inornatus has, however, a considerable number of somewhat varying pelage characteristics which together easily dis- tinguish it from C. p. stevensi: (1) The ordinary number of blackish bands on the tail hairs is five rather than four. (2) The tail is col- ored dorsally the same as the animal's back instead of paler and grayer. (3) The tail is lighter colored ventrally than dorsally but is yellowish brown instead of cool gray. (4) There is no flash mark of lighter color on the dorsal pelage of the hip as is common in stevensi. (5) The feet and forelegs are colored like the back instead of being notably more gray. (6) The subterminal light bands on the tail hairs are too short (ca. 2 mm.) and too yellow to give any special impres- sion as to the color of the tail hair tips, whereas in stevensi their white ness and length (ca. 3 mm.) give an impression of whitetipped hairs. Although C. caniceps concolor is most like C. inornatus in pelage characteristics, inornatus is geographically remote from concolor. On the other hand, C. inornatus is separated from the range of C. c. caniceps only by the Mekong River. Because of this proximity to caniceps it is important to consider the amount of difference be- tween inornatus and C. c. caniceps: (1) C. inornatus does not have a seasonal change of the dorsal pelage to orange but remains agouti gray (as do the subspecies of C. caniceps to the south of C. c. cani- ceps). (2) C. inornatus has agouti feet concolorous with the pelage of its dorsum instead of contrastingly pale gray. (3) C. inornatus has a vaguely delineated black pencil to its tail, obscured by whitish tips on the elongate hairs that compose it and varying amounts of light banding basal to the black on these hairs, unlike the crisply all black and sharply defined black tail tip of C. c. caniceps. (4) C. in- ornatus is substantially smaller than caniceps, greatest skull length of adults being about 51 mm. compared to 58 mm. in caniceps. (5) The rostrum and ear tips are colored like the back in inornatus (the tip of the rostrum tending a little toward black) instead of be- ing quite whitish as in C. c. caniceps. (6) C. inornatus seems gener- ally to lack the rather prominent longitudinal stripe of agouti which bisects the bluish gray venter of caniceps. Relationships to other species. — C. inornatus is almost wholly sym- patric with C. erythraevs hendeei, and was taken without any evidence 212 FIELDIANA: ZOOLOGY, VOLUME 48 of interbreeding in many of the same localities, and (when the ele- vation was given on the specimen tag) at the same elevation. Since inornatus is, furthermore, a smaller squirrel than hendeei, we do not question (nor has anyone) that they are specifically distinct. C. inornatus is separated from all of the subspecies of C. pygerythrus by two great barrier rivers, the Salween and the Mekong, and earlier the Irrawaddy must have constituted a third. It is shown above that inornatus generally has a good many differences in pelage color from the subspecies of C. pygerythrus that it most closely resembles. We recognize that the barrier strength of the great rivers may weaken to the north, that some other tree squirrel species are sep- arated into populations of no more than subspecific rank by any one of these river barriers in the north, and that C. pygerythrus stevensi is the northernmost subspecies of its species. These things do seem to support Ellerman and Morrison-Scott (1951, p. 488) in treating the species pygerythrus as the nearest relative of inornatus. On the amount of pelage color difference between inornatus and C. p. ste- vensi, the great gap in known distribution between them (a gap from which specimens of other tree squirrel species are nevertheless repre- sented in the collections available to us), and the interposition of the several major rivers as filter barriers, however, we cannot accept a subspecific relationship between inornatus and stevensi and conclude that these two forms are specifically distinct. Whether the species Callosciurus pygerythrus is indeed the closest relative of Callosciurus inornatus is to be questioned next. The dif- ferences between C. inornatus and C. c. caniceps described above are numerous and trenchant, but those between C. inornatus and C. cani- ceps concolor are exceedingly few. This close similarity between in- ornatus and concolor becomes significant in view of the following considerations which are emerging from the present study: (1) The Malay Peninsula below the Isthmus of Kra is a large mountainous area of little-differentiated tropical rainforest, virtually without sea- sons, and this region appears to provide many niches for tree squir- rels in which they remain conservative in pelage color. (2) Tonkin and upper Laos constitute an area which seems to have fewer niches for tree squirrels, but where they are similarly conservative in pelage color. (3) Thailand, excluding the peninsular portion, in strong con- trast with Malaya, contains large tropical lowland forest areas which differ sharply in annual rainfall, and have prominent alternating dry and rainy seasons, and in it tree squirrels develop the most fantasti- cally colored pelage. MOORE AND TATE: DIURNAL SQUIRRELS 213 It seems reasonable to conceive that when CaUosciurus caniceps jfirst over-ran Thailand and spread throughout the area that is the present range of C. caniceps (Malaya to northern Thailand), it would then have had color characters much like concolor or inornatus throughout this range. Only two changes are needed then to achieve present conditions. (1) A stream piracy of the Mekong River by one of its own tributaries could have removed a substantial area from west to east of the Mekong River. A hypothetical example: the course of the Mekong might once have been through the valley which the Nam Tha now drains but was subsequently diverted to its present channel by head erosion of a recurved western tributary. This would have the effect of isolating a large population of this new species C. caniceps and freeing it to spread elsewhere east of the river. A population so separated from the species caniceps would reason- ably be expected to have spread to the limits now known for inor- natus and to have conserved rather well the pelage characters with which it arrived. (2) The population of species caniceps remaining west of the Mekong in the main part of Thailand would have evolved other pelage characteristics which would be the trenchant and extra- ordinary ones now observable in C. c. caniceps. That C inornatus thus may be directly derived from C. caniceps instead of C. pygerythrus does not appear to have been seriously con- sidered before, but may now be regarded as the more probable rela- tionship as a matter of minimum hypothesis. If inornatus is derived from species caniceps, there is an interesting niche reversal where sympatric with species flavimanus, for C. c. caniceps is the larger squirrel than C. /. tachin, and atrodorsalis and thai, but C. f. hendeei is the larger squirrel than imitator. CaUosciurus pygerythrus (Geoffroy St. Hillaire) Definition. — The species pygerythrus includes subspecies pyger- ythrus, janetta, owensi, mearsi, stevensi, hlythi, and lokroides, and their synonyms. These occur principally east of the Sittang and upper Irrawaddy valleys to the edges of the Indochinese Subregion as shown in Figure 16. Diagnosis. — The fully grown-out tail hairs of this species have but four blackish bands. There is a seasonal flash mark in the pel- age of the thigh which contrasts with the ordinary agouti dorsal pelage by being lighter (about cream color in most subspecies, but ochraceous buff in some) . These characters distinguish pygerythrus from sympatric species and adjacently allopatric ones. 214 FIELDIANA: ZOOLOGY, VOLUME 48 CHIMA Fig. 16. Geographic distribution of the Irrawaddy squirrel, Callosciurus pygerythrus, as determined from material examined. Subspecies: A, pygerythrus; B, janetta; C, owensi: D, mearsi; E, stevensi; F, hlyihi; and G, lokroides. Dotted lines separate subspecies as known from specimens from the localities plotted but are not offered even as potential boundary lines between the subspecies. Relationships to other species. — This is a species of tree squirrel that is of smaller size than a species of tree squirrel generally attains in a hospitable geographic area which it occupies alone. Throughout virtually all of its range pygerythrus is sympatric with a larger spe- cies of tree squirrel, either Callosciurus erythraeus or C. ferrugineous. Compare their dimensions as indicated by Tables 11 and 4. This geographic range is constituted by those parts of Burma, India, and Pakistan which lie west of the Irrawaddy and Sittang rivers and east of the Brahmaputra River. An additional strip west of the Brahma- putra extends along the southern face of the Himalayas beyond the MOORE AND TATE: DIURNAL SQUIRRELS 215 middle of Nepal. There is also a very small area that this species has invaded eastward across the Sittang Valley in the vicinity of Mandalay. It is shown in the account of Callosciurus phayrei that phayrei is the closest relative of Callosciurus pygerythrus and that through phay- rei, pygerythrus is next most closely related to Callosciurus caniceps. It is interesting that John Anderson (1879, p. 231) ventured much this same opinion about a definite interrelationship of pygerythrus, phayrei, and caniceps, from examination of a substantial series, but he was speculating that they are all one species. Intraspecific variation. — In subspecies lokroides the term "winter pelage" may be perfectly acceptable to distinguish the duller pelage of winter from the very much brighter pelage of summer. However, Raven collected subspecies mearsi in dull "winter" pelage without flash marks on the hip in March at several localities directly across the Chindwin River from where he was collecting subspecies owensi at the same time in bright "summer" pelage with flash marks. Short- ridge (in Wroughton, 1916, p. 293) remarked no pronounced differ- ence in the character of the forest on the two sides of the Chindwin River at these places, between Hkampti and Tamanthe. The pelage of two subspecies being out of phase across the river barrier raises some question as to whether the terms "winter" and "summer" pel- age are in fact suitable terms to be applied to these subspecies of lower latitude and elevation. Perhaps "eclipse" pelage would be better ter- minology for the dull pelage, where a distinctive alternation occurs. We have examined and compared summer and winter specimens of the subspecies janetta, which occupies the dry scrub forest of the dry zone of central Burma, but the differences between these are so slight that one can hardly claim there are seasonally bright and dull pelages in janetta at all. The flash mark on the hip is present in all specimens of janetta. Part of what we recognize here as mearsi west of the lower Chindwin and lower Irrawaddy also occurs in the ex- treme dry zone of central Burma. Material of mearsi collected in winter in this zone is, of course, distinct from janetta, but surprisingly resembles it in having a flash mark on the hip; whereas the large winter season series of mearsi from the deciduous forest zone seems to be in dull seasonal pelage and has no flash mark on the hip. Thus, we have a species which as a whole seems to alternate dull pelage (and no flash mark) with bright pelage (and flash mark) . But within this species is one subspecies, janetta, and part of another, mearsi, that unlike the rest of the species occur in the scrub forest of a nearly 2 o (-1 MS C 3 05C0Ot--^eCOC0Tj-*oocoa50ix>t-oo a 03 1,2 O OS 00 00 t- -^ i-i CO P3 (M 1-1 r-( ITS i-H CO T— 1 CO CO "^ "^ "^ "^ 1— 1 1— 1 T-l 1— 1 1— 1 1— 1 • Tj< 00 (N 4^ UtI O ■ 00 I-H • a o CO (D a. >> Eh a> s o pa 4J bO O C • lO 00 t- O (M w ^3 CO CO o Oi ^ O t:1< lO Tf r73 ^ 00 U5 «3 Tt tH Ti. . -H 4i -H -H c OOi(MC^]«DOOOO • t- «0 Oi «£> CD *- 1—1 I-H 1-H L, ^ -H 41 -H -H -H f. 000«OU5-«*00000 a>o>i-ioooooocoi-iooo iHi-l(Mi-li-li-l(MC> i rtj3 .^ « . S-kJ? '-iiOMO«o«Doot-«D-^t--^Oi-it-eoeot-oocOOi50c^rHCO -(M -coos '-h'Y 535 • • ■«£>t-'X>ia5«DCOt-C-.'l>t>'l>t^ -C-I -t-^tD's-'S I— I -C "-H , OS I 5 eSg t-OOvO(Mt-OoO'-IOOOiU3000t-0000 ^ Q+j t-t- •t>t-co«o«o?Doooooot~t~t~t-oot-t-t~ -C J3- ^ O g rt ooaiasosoioot^ -OiOiMNooooooooiaj 2+^ -g '" > tJjTJ'C •o-^ifl'-Ht- -OS CD -cot-eo -t- •t-'-H an tiSoa, - -lo-rjiini-^Ti* --"tyj ■ lei la iti -lo ■ tA ^ '"^, W — ,0) o Ph (-1 T-H o 55 ■ eo (M -* ■ i-H • c^ • ■ t-^ ->^* -to • 10* 10 -^ C Ht • -cocoeo -co -co • • -coco -co -coc^ -wO) »-] e ^ e i3 .S o -H -H (MOooooaiOooooomT-iookn)Tj -H -H -H -H -H 4^ 4^ ,— (I— IC I I I I KILOMETERS Fig. 18. Ranges of the Burmese striped tree squirrel, Tamiops mcclellandi, A to F; and the Cambodian striped tree squirrel, Tamiops rodolphei, G and H, plotted from collecting localities of material examined. Subspecies of mcclellandi: A, mcclellandi; B, collinus; C, kongensis; D, barbei; E, leucotis; and F, inconstans. Subspecies of rodolphei: G, rodolphei; and H, elbeli. 236 MOORE AND TATE: DIURNAL SQUIRRELS 237 and through Yunnan to northern Vietnam (where it is represented by the subspecies inconstans) . There it seems to be sympatric with, or interdigitated with, species maritimus. Intraspecific variation. — The species mcclellandi occurs in forests from Sikkim, Bhutan, Assam, and northernmost Burma southward in forested hill country probably throughout Burma, Thailand ex- cepting the Mekong plain, and the Malay States. Its outer pair of light stripes in the dorsal pelage is strikingly light and prominent in comparison to the subdued inner pair, and it is characteristic that the outer light stripes continue strongly marked across the shoulder area and connect with the light facial stripe. While five subspecies of mcclellandi receive recognition here, the authors wish to point out that there is a greater distinction between the form mcclellandi and the other four than there is between any two of the other four. Because of this, one of the present authors (Tate) elected to recognize mcclellandi as a monotypic species and the other four forms as constituting the species barbei. This may prove to be correct if study of further material discloses no intergra- dation between mcclellandi and collinus. One might suppose it to be equally logical in our present state of knowledge, however, to con- sider the distinction between mcclellandi and the other four forms as of subspecific value and that the "other four forms" constitute a vari- able but single subspecies. From the material that we have studied, it seems to us that leucotis, barbei, and kongensis are distinct geo- graphic subspecies occupying equally distinct physiographic areas. The other form, collinus, is somewhat less distinct, and also the considerable geographic area it occupies does not appear to be physio- graphically distinct. However, this form is more than a broad zone of intergradation between mcclellandi and kongensis, for it possesses a color character shared by neither, the blackness of the inner paired dark lines. It is here considered, therefore, that the distinction be- tween subspecies mcclellandi and the other four is at an advanced subspecific (and perhaps incipient species) state of differentiation; whereas the distinctions between any adjacent two of the subspecies collinus, kongensis, barbei, and leu/^otis are at a subspecies, but less advanced, state of differentiation. Tamiops mcclellandi mcclellandi (Horsfield) Sciurus mcclellandi Horsfield, 1839, Proc. Zool. Soc. London, 1839, p. 152. Sciurus pembertoni Blyth, 1842, J. Asiatic Soc. Bengal, 11, p. 887. Sciurus macclellandi manipurensis Bonhote, 1900, Ann. Mag. Nat. Hist. (ser. 7), 5, p. 51. 238 FIELDIANA: ZOOLOGY, VOLUME 48 Types. — Sciurus mcclellandi, two cotypes, BM Nos. 79.11.21.372- 373, adult male and a young adult, the former marked "lectopara- type," both from "Assam"; pembertoni, BM No. 79.11.21.374, adult female, from Bhotan; manipurensis, BM No. 85.8.1.273, old male from Aimole, Manipur. Material examined, from Assam, India. — Tura, Garo Hills (AMNH), one; Tura Mt., Garo Hills (CNHM), three; Pynursla, Khasi Hills (CNHM), two; Mawphlang, Khasi Hills (CNHM), three; Cherrapunji (CNHM), one; Sangau, Lushai Hills (CNHM), three; Karong, Manipur (CNHM), six; Takubama, Naga Hills (CNHM), four; Kohima, Naga Hills (CNHM), two; Sadiya, Mish- mi Hills (AMNH), three, (CNHM), one; Dreyi, Mishmi Hills (AMNH), one; Tezu, Mishmi Hills (AMNH), five. Material examined, from Sikkim, India. — Darjeeling (CNHM), one; Rungbong Valley, 4000 feet (MCZ), one; Chungtang (CNHM), three, (USNM), one; Lingtam (USNM), three, (CNHM), 28; "Sik- kim" (MNHN), one. Material examined, from Burma. — 6 to 25 miles N. of Myitkyina (USNM), five; 170 miles N. of Myitkyina (CNHM), one; Seniku- Shingaw Road (AMNH), one; Laukhaung-Pyepat Road (AMNH), one; Nanyaseik (AMNH), three; Lonkin (AMNH), one; Mansum (AMNH), one; Dalu (AMNH), four; Tagahka (AMNH), five; Luck- changa (AMNH), one; Dagung Hka (AMNH), one; Haibum (AMNH), three; Singkaling Hkamti (AMNH), one; Kawai, W. Bank (AMNH), one; Kaunghein (AMNH), four; Nonswa (AMNH), one; Mawlaik (AMNH), one; Mt. Victoria, Pakokku Chin Hills (AMNH), nine. Type description. — Typically mcclellandi, studied in 1951, has the median black line about 4 mm. wide and extending from nape to rump. The lateral pair of dark lines extends from the shoulders to the rump but are a little wider, 5 mm. They are partly obscured at their inner edges by the overlying yellowish buffy hair tips of the intervening area. Laterally they are edged by the pair of strongly colored yellow-buff lines which are the primary elements in the dor- sal pattern. Then two yellow lines reach from the rostrum over the whiskerpatch, beneath the eyes and ears, become broadened at the sides of the neck and connect with the outer pair of lines on the back, which finally fade out at the rump. The maximum width of this line is at the side of the neck. External to this pair of yellow lines is the merest suggestion of the outermost pair of dark lines about 1 inch in MOORE AND TATE: DIURNAL SQUIRRELS 239 length and less than 2 mm. in width. Underparts buffy-tipped (with- out any trace of red) with the bases of the hairs gray. Hands and feet grayish buff. Tail with bases of the hairs brownish buff, subterminal bands black, tips buffy white. Extreme tip of tail blackish. This is the basic form against which all others must be compared. The length of the head and body is ca. 110 mm., of tail ca. 100 mm., hind foot ca. 28 mm. It is apparently a much smaller Tamiops than any member of the species swinhoei. Discussion. — While there are slight differences in pelage color which may be locally geographic in expression within this typical subspecies, these are negligible by comparison with the differences between T. m. mcclellandi and its nearest conspecific neighbor T. m. collinus. A series collected high on Mt. Victoria in Burma is not only lighter in ventral pelage color than typical mcclellandi but some- what larger in size. Nevertheless, these differences would make of it only some sort of a micro-race by comparison with those differen- tiating collinus and mcclellandi. Habits. — In a letter of June 10, 1961, Lord Cranbrook contributes the following field notes on this species from his collections on the Adung Valley expedition: "No. 60. The commonest squirrel in the district (Myitkyina — Sumpra Bum). Usually seen high up in tall trees, moving in short rushes and then staying motionless, sometimes head downward, often some minutes at a time. I have never seen one in low bushes, always high up. The tail is usually held straight out [behind] and very seldom 'flicked' or held over the back. Fright- ened, they freeze flattened out with tail extended." These notes were not included in Cranbrook's account in Kinnear (1934). Another collector wrote: "This squirrel is found in most of the forests above 5,000 feet. It is common but owing to its power of concealment is generally not very easy to find. It is very nimble and appears to glide rather than walk along the boughs and slender twigs. I observed it in pairs and sometimes in small parties often sharing the tree with D. lokriah. It seldom comes to the ground. The call is a quickly repeated 'chick' much harsher in tone than that of F. pennanti." (C. A. Crump in Wroughton, 1916a, p. 488.) Tamiops mcclellandi collinus Moore Tamiops mcclellandi collinus Moore, 1958, American Mus. Novitates, no. 1879, p. 1. Type. — AMNH no. 163528, an old male from 800 meters elevation, Maymyo, Burma, collected November 28, 1937, by Gerd Heinrich. 240 FIELDIANA: ZOOLOGY, VOLUME 48 Material examined. — Maymyo, 800 meters, Burma (AMNH), 14; Mengting, 1700 feet, Yunnan (AMNH), four; Mu-cheng, 5000 feet, Yunnan (AMNH), two; Nam Ting, Yunnan (MCZ), one; Chiang Saen Kao, Chiang Rai, Thailand (USNM), one; Sawan Mt., Ban Seid, Loey, Thailand (USNM), one; Lomloe Mt., Maeo, Goksatawn, Loey, Thailand (USNM), eight; Vientiane, Laos (USNM), five; Chieng Dao, Thailand (CNHM), one; Doi Chieng Dao, 4000 feet, N. Siam (ANSP), three; Chieng Sen [Chiang Saen], N. Siam (ANSP), one; Don Qua, Laos (ANSP), one. This subspecies is more strongly striped than mcclellandi, less red than barhei, darker dorsally than kongensis, and much more strongly striped than inconstans. It is not in geographic contact with leu- cotis, but is ventrally more orange and less yellow than leucotis. Tamiops mcclellandi inconstans (Thomas) Tamiops inconstans Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, p. 306. Type. — BM No. 12.7.25.31, an old male from southern Yunnan (possibly near Mengtsze), collected January 31, 1910, by H. Orii. Material examined.— "Yunnsin" (BM), one; "Tonkin" (MCZ), three, (CNHM), one; Bao Ha, Tonkin (CNHM), two; Pakha, An- nam [Tonkin] (CNHM), one. Original description. — "This very distinct little squirrel is char- acterized by its unusually inconspicuous striping above and by the strong yellowish buff y of its lower surface — in fact, it is above one of the dullest and below one of the brightest of the genus." We find the ventral pelage to be Light Salmon Orange where the colored hair tips entirely cover the gray hair bases. Tamiops mcclellandi kongensis (Bonhote) Sciurus mcclellandi kongensis Bonhote, 1901, Proc. Zool. Soc. London, 1901 p. 55. Type. — BM No. 0.10.7.18, adult female from Raheng, Siam, col- lected February 7, 1900, by T. H. Lyle. Material examined, all from Thailand.- — Chieng Dao [Ban Chiang Dao] (USNM), one, (CNHM), one; Mae Hong Sorn [Ban Muai To] (USNM), two; Doi Hua Mot (USNM), one; Doi Phra Chao [Khao Pha Cho] (USNM), one; Mae Wan River near Doi Saket (MCZ), one; Nan [Muang Nan] (USNM), four; Doi Sutep [Doi Suthep] (AMNH), one; Doi Angka [Doi Ang Ka=Doi Inthanon] (MCZ), six, (USNM), two; Koon Tan [Doi Khun Tan or Sathani Khun Tan] MOORE AND TATE: DIURNAL SQUIRRELS 241 (USNM), four, (ANSP), two, (NR), three, (CNHM), one; Chum Poo [Sathani Tha Chomphu] (NR), two; Mesarieng [Ban Mae Sari- ang] (USNM), one; Ta Chang Tai [Tha Chang Tai] (USNM), two; Raheng [Ban Rahaeng] (USNM), five; Chiengmai [Muang Chiang Mai] (ANSP), two, (MCZ), three; "Wung Pratart Farm" Kam Peng Pet Prov. [Kampheng Phet] (CNHM), 14; 40 miles E. of Um Pang [Ban Um Phang] (AMNH), one; Pitsanulok (AMNH), one; Pak Tha (NR), one; Pak Koh (NR), one; "Den Chai" (NR), one; Pran [Pran Buri] (USNM), two; Huey Yang, Sriracha [Sathani Huai Yang] (USNM), one; "Muang Qua Yie," Nakorn Sawan (USNM), one; "Wat Sytie," Paknampho, Nakorn Sawan (USNM), one; "Ban End," Chiengmai (USNM), one; Pujeg, Pak Tho, Rajburi (USNM), two; Kowjeen, Pak Tho, Rajburi (USNM), one; Tungnockarien, Jombing, Rajburi (USNM), three; Borploy, Kanjanaburi (USNM), two; "Tarn bol, Tak tog, Bantak," Tak [Ban Rahaeng] (USNM), two; Tra Khanun, Hin Laem, Kanachanaburi (USNM), nine; Ban Klua Klan [Ban Khlua Klang], Prachuap Kiri Khan (USNM), one; Klong Klung [Ban Khlong Khlung], Kampheng Phet (USNM), one; "Ban Thoong Cheark, Slog Bartara, Kha Nu," Kamphengphet (USNM), two; "Srisawat" (ANSP), two. In this race the dimensions of the various dorsal lines remain al- most exactly as in harhei. The chief difference occurs in the overall pallid coloration. There is a general infusion of light gray, instead of the reddish orange wash of barbei and the yellow wash in leucotis. The pale hues of the back in the present race are very pale buffy yellow, and the underparts are about Ochraceous Buff. This sub- species seems to have differentiated in the rain shadow zone of Thai- land, and has a range apparently identical with that of the pale race of Menetes herdmorei consular is. Tamiops mcclellandi barbei (Blyth) Sciurus barbei Blyth, 1847, J. Asiatic Soc. Bengal, 16, pt. 2, p. 875. Type. — Not seen. From Zami River, Ye Province, 100 miles south of Moulmein, Tenasserim, Burma. Material examined, all from Burma. — Kawkereik, Tenasserim (AMNH), two; Lampha, Tenasserim (AMNH), one; Tavoy (CNHM), one; Tenasserim Town (CNHM), one; Taok Village, Tenasserim (AMNH), two; Zami River, Ye Province, 100 miles south of Moulmein (BM), one. 242 FIELDIANA: ZOOLOGY, VOLUME 48 Discussion. — While material from Kawkereik at 175 feet eleva- tion has ventral pelage about Ochraceous-Salmon at the throat and Cinnamon-Rufous or Ferruginous on the remainder of the venter, material from nearby Taok at 1100 feet elevation is much paler ven- trally and slightly grayer dorsally so as to be really intermediate between barhei and kongensis. Like the Taok material, also, is a specimen from neighboring Lampha. Ventral pelage of these is near- est to Capucine Orange. The Kawkereik and Lampha material was collected December 12 and 28 respectively, the Taok January 16. The entire dorsal pelage of the Kawkereik material is infused with reddish which shows as Light Salmon Orange in the outer light dor- sal stripes. Habits. — Field observations are offered from southernmost Burma: "Completely arboreal. I have never observed this species actually on the ground. Common around villages on the Tenasserim River but not observed further south. More plentiful in native fruit gar- dens than in jungle. Weight, 13^-2 ozs." (G. C. Shortridge in Wroughton, 1915b, p. 713.) Tamiops mcclellandi leucotis (Temminck) [Extraterritorial] Tamias [sic] leucotis Temminck, 1853, Esquisses Zool. Cote Guine, p. 252. Sciurus novemlineatus Miller, 1903, Proc. Biol. Soc. Washington, 16, p. 147. Types. — Tamias leucotis "Mallaca peninsula" and not seen by the present authors; novemlineatus, USNM No. 84403, adult male from Trong, 1500 feet, collected February 19, 1897, by W. L. Abbott. Material examined, from Malay States. — "Selangor" (USNM), two; Ginking B., Selangor (USNM), one; Brinchong Hill, 5500 feet, Cameron Highlands, Pahang (USNM), one; Gunong Mengkuang, 4800 feet, Lebah, Selangor (USNM), one; Telom, 3800 feet, Perak near the Pahang border (USNM), one. Material examined, from peninsular Thailand. — Trong [Trang], 500 feet (USNM), one; Khaw Nom Phu, 2000 feet, Trong [Trang] (USNM), one; Ban Kiriwong (USNM), five; Ban Chit [Ban Nong Chik] (USNM), one. Material examined, from Tenasserim, Burma. — Telok Besar (USNM), three. Discussion. — This subspecies lacks the general reddish infusion in the dorsal pelage displayed by T. b. barbei, and the ventral pelage of leucotis is much the yellower, being Ochraceous Buff in the yellow- MOORE AND TATE: DIURNAL SQUIRRELS 243 est instances and a little more orange than that in several others, matching no Ridgway color precisely. Surprisingly the series of three from Trong show strong evidence of intergradation with barbei both dorsally and ventrally, the ventral pelage being Apricot Buff on the throat and Orange Cinnamon on the venter. Yet from north of this, 200 miles closer to barbei range, at Telok Besar in the south- ern tip of Burma, the specimens show no dorsal infusion of red, and the ventral pelage is also correct for leiLCotis. Presumably less than 100 miles north of this leucotis intergrades with both barbei and kongensis. Habits. —Concerning the distribution and habits of this subspecies Robinson {in Bonhote, 1903, p. 21) wrote, "This beautiful little spe- cies was not met with on the Eastern side of the Peninsula. In Perak and Selangor it is certainly a mountain form, and I do not think it occurs much below 3000'. It was very common at Telom, and was also very abundant on the mountains round the Semangko Pass. It is very largely an insectivorous species, and seems to keep chiefly to the trunk and main branches of the trees, running along them with its tail pressed close against the bark." Chasen (1940, p. x) in commenting on the distinctive mountain fauna encountered above 3000 feet elevation in the Malaysian Subregion, notes that, "A Himalayan element (e.g., Rattus bowersi, Tamiops, Dremomys rufi- genis, Sciurus erythraeus) is the dominant feature of the high-levels in Perak, Pahang and Selangor, but it fades out in the south, and is absent in Negri Sembilan and on the Johore hills; it is also weak on the isolated summit of Kedah Peak in the north." Tamiops rodolphei (Milne-Edwards) Definition. — The species rodolphei is constituted by subspecies rodolphei and elbeli and the named forms here included in them. Its distribution, shown in Figure 18, is in Cambodia, southern Laos, southern Vietnam, and eastern Thailand. Diagnosis. — Species rodolphei is distinguished from the other spe- cies of this genus by: (1) The outer and inner pairs of light lines are of equal width. (2) The inner pair of light lines approximates the brightness of the outer pair. (3) The blackish middorsal line is almost invariably divided for part of its length by a very thin, pale brown line. Relationships to other species. — Tamiops rodolphei appears to re- place Tamiops mcclellandi with no known overlap or intergradation in eastern Thailand. In southern Vietnam it overlaps or interdigi- 244 FIELDIANA: ZOOLOGY, VOLUME 48 tates with the known southern extremity of the range of the species maritimus. Compare Figures 18 and 20. Tamiops rodolphei rodolphei (Milne-Edwards) Sciurus (Tamias) rodolphei Milne-Edwards, 1867, Revue Mag. Zool., 19, p. 227. Tamiops macclellandi liantis KIoss, 1919, J. Nat. Hist. Soc. Siam, 3, p. 370. Tamiops lylei Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, p. 307 (hom- onym). Tamiops macclellandi dolphoides Kloss, 1922, J. Nat. Hist. Soc. Siam, 4, p. 101. Callosciurus holti Ellerman, 1940, Families and Genera of Living Rodents, 1, p. 355 (new name for lylei). Types.— Sciurus (Tamias) rodolphei MNHN No. 1864-683 (318), adult from Saigon, Indo-China; dolphoides, apparently lost, Kloss field no. 2723/CBK, adult female from Kompong Som Bon, near Sr4 Umbel, southwest Cambodia; liantis, USNM No. 221542, young adult female from Sata Hip, near Cape Liant, southeastern coast of Siam; lylei= holti, EM No. 6.10.7.9, young adult male from sea coast fifty miles south of Bangkok, Siam. Material examined, from Thailand. — Sriracha [Ban Si Racha] (USNM), two, (CNHM), one; Chantabun [Chanthaburi] (ANSP), two, (USNM), one; Kao Sabab [Khao Sa Bap] (USNM), two; Lem Ngop [Ban Laem Ngop] (USNM), one; Kao Seming, Krat [Khao Saming, Trat] (ANSP), two, (USNM), one; Nongkhor [Ban Nong Kho] Chon Buri Prov. (USNM), two. Material examined, from Indochina. — Ban Me Thuot [Buon Ma Thuot], Annam (CNHM), four; Bien Hoa, Cochin China (USNM), two; Gougah [Lien Gongah] Annam (CNHM), 11, (MCZ), one; Thateng (USNM), one, (CNHM), 11; Pak-S6, Laos (CNHM), two; Paksong, Laos (CNHM), one; "Phu Kongutoul" (CNHM), one; Quangtri Phuoc Mon, Annam (CNHM), three; Ninh Hoa, Annam (CNHM), one; "Cochin China" (CNHM), one; Plateau Bolovens, 2500-3000 feet, Laos (ANSP), three, (AMNH), three; Tayninh, 100 feet, Cochin-China (MNHN), one; Angkor, 200 feet, Cambodia (MNHN), two; N. V. Kampot, 100 feet, Cambodia (MNHN), one. The head and nape are almost uniformly dull brown in this sub- species. The tail is exceptionally slender due to shortness of its hairs. The underparts tend toward yellow or orange, somewhat as in mcclellandi kongensis or m. collinus, never buffy or whitish as in m. mcclellandi and in subspecies of maritimus. MOORE AND TATE: DIURNAL SQUIRRELS 245 Tamiops rodolphei elbeli Moore Tamiops rodolphei elbeli Moore, 1958, Amer. Mus. Novitates, no. 1879, p. 4. Type.—VSNM No. 294876, adult female, skin and skull, collected by Robert E. Elbel on January 18, 1952, at village of Ban Lad, Pookeio District, Chaiyaphum Province, Thailand (ca. latitude 16° 40' N., longitude 101° 45' E.). Material examined. — Ban Lad, Pookeio, Chaiyaphum, Thailand (USNM), two; Ban Non Toulek, Pookeio, Chaiyaphum, Thai- land (USNM), three; Khon Kaen, Thailand (USNM), seven; "Watpa" (ANSP), one. Diagnosis. — Tamiops rodolphei elbeli is distinguished from all sub- species of Tamiops mcclellandi, swinhoei, or maritimus by the mid- dorsal black stripe, for this stripe is divided at least part of its length by a fine line of yellowish brown down its middle. It differs from Tamiops rodolphei rodolphei, which occurs to the east and south of it, in the brilliance of its Isabella Color crown and nape compared with the Buffy Brown (XL) to Snuff Brown (XXIX) crown and nape of T. r. rodolphei (including three virtual topo types from Bien Hoa, Cochin China). Dimensions. — The skull measurements (in millimeters) of the type are: occipitonasal length, 32.6; condylobasal length, 30.0; right maxillary toothrow, 5.4; palatal length, 15.8; mastoid breadth, 14.9; greatest breadth, 20.1; nasal length, 10.2; and orbitonasal length, 13.6. Field measurements in millimeters of the total length, tail length, and hind foot of the type of T. r. elbeli are, respectively, 250, 130, and 30. The same characters recorded for a male and two females of T. r. elbeli from Ban Non Toulek are, respectively: 220, 250, 220; 110, 129, 102; and 30, 32, 30. Variation. — Of 17 skulls of Tamiops m. collinus and kongensis examined in the United States National Museum from localities geo- graphically near to elbeli, five have the posterior surface of the ex- tremities of the upper incisors beveled so that it shows just a slight concavity in the side view. Each of the other 12 is cut so that it shows a notch. Of 13 skulls of T. r. elbeli examined, four have just a slight concavity, and nine have a straight bevel. The lateral mar- gins of the basioccipital are curled ventrad and rise slightly to a low point or process in T. r. elbeli, but in T. m. collinus and kongensis and also T. r. rodolphei these edges remain low ridges and do not form a point or process. 246 FIELDIANA: ZOOLOGY, VOLUME 48 Discussion. — This subspecies comes from the eastern part of Thai- land, the Mekong plateau 2000 feet in elevation, which is subtended on the north and east by the Mekong River and south by the Phanom Dang Raek, a range of mountains. Excepting for Gyldenstolpe (1914) who collected mammals on its southwestern edge about Korat, evi- dently no mammalogist had collected on this plateau before R. E. Elbel got into the northwestern quarter of it. The material repre- senting Tamiops r. elbeli all comes from Chaiyaphum Province and Khonkaen Province. Tamiops swinhoei (Milne-Edwards) Definition. — This species includes subspecies swinhoei and the several named forms here included in it, and subspecies vestitus. Figure 19 shows the distribution of species swinhoei as known to us from specimens. Diagnosis. — This is apparently an extremely variable species geo- graphically, and although swinhoei seems to be generally larger than the other species of Tamiops and to have longer, denser fur, diag- nostic distinction by these characters does not seem possible in the material we have examined. These two tendencies may be related to the fact that species swinhoei lives farther north or at higher ele- vations than the other species of Tamiops. The outer pair of light lines is less brilliant than those of mcclellandi and often broader, and they more generally stop at the shoulder instead of being connected with the cheek stripe. Relationships to other species. — The range of this species appar- ently does not overlap that of mcclellandi to any great extent. The only localities from which both species have been taken are where the edges of their ranges meet. At these places there has been a notable difference in the elevations at which the two species were taken. In northeastern Burma near Imaw Bum Anthony (1941, p. 93) took the species mcclellandi between 1000 and 2000 feet ele- vation in definitely tropical habitat and at 4500 feet, but he collected the species swinhoei at 9000 and 10,000 feet. Northward about 130 miles from Imaw Bum, Cranbrook collected mcclellandi at 1500 feet elevation but at nearby Adung Valley took swinhoei at 8000 and 9000 feet. At Mucheng, Yunnan, China, about 175 miles south of Imaw Bum, Edmund Heller collected the species mcclellandi at 5000 feet elevation and swinhoei at 7000 feet. These three localities are the westernmost known for the species swinhoei and the northeastern- most known for mcclellandi. See Figure 19. ] MOORE AND TATE: DIURNAL SQUIRRELS 247 9i't •t- "m't r ■"■■ / KH'C in ...••■■' (B) / ^ 0 MiLES •■■ J?" ■:;^\',..^ /' ; HOPEH k 0 s» "■■. + KILOMETERS C;' V^' KANSU \ j j ( ..r •••■••* riCA) ! HONAN ';5^^^K \ '+. ; '"" ■\ v.-—. v;;— -^ ;anhw£i j V SIKANG C< • ••.. ..•• H J' ■; •jpj HUPEH 5 fJl . f A) J-'\ t— YUNNAN ■ ■ "'^4. i..,.'" ■■■' ~' "■ '■ r CHEKIANGV .■■■ • "7 ■ ■' • • • tf/ :. (D)« > ; FU KlENj^ /^ BURMA j^ •(a) s TONKIN ^ KWANG5/ 'f^l^ FOR>|S/ Fig. 19. Distribution of the Chinese striped tree squirrel, Tamiops sivinhoei (entire range) and the maritime striped tree squirrel, Tamiops maritimus (north- eastern range) as obtained from plotting collecting localities of materials examined. Subspecies of swinhoei: A, swinhoei; and B, vestitus. Subspecies of maritimus: C, maritimus; and D, mx)nticolus. On the apparent southern edge of the range of T. swinhoei Dela- cour and Lowe evidently obtained a series of 17 from 8000 to 10,000 feet elevation on Mt. Fan Si Pan near Chapa in northern Tonkin, but at lower elevations in the same vicinity took T. maritimus haina- nus. In reporting this, Osgood (1932, p. 292) recognized these ex- amples of swinhoei as a race of monticolus. Eastward in the range of swinhoei there is an extraordinary gap in examined material of this species, as shown on the map. The eastern distribution (T. s. ves- titus) lies entirely in Wang's (1956) oak-dominated deciduous broad- leaved forest type. The area of this forest type extends westwardly about to the western distribution (T. s. swinhoei), but it appears that not a single one of the twenty-odd localities that we have for T. swin- hoei here is in that general forest type area. Possibly this constitutes 248 FIELDIANA: ZOOLOGY, VOLUME 48 something of a barrier, and the absence of records of Tamiops from middle China really indicates absence of the genus, for Sciurotamias has been collected there. Compare Figures 19 and 33. Of course, Sciurotamias is very much a ground squirrel and would possibly have increased in the same places where Tamiops may have been elimi- nated by deforestation. Fu (1936) found "Tamiops macclellandi vestitus" in the Sung-shan, which he describes as a famous mountain in western Honan with luxuriant forests, and he says that this squir- rel lives in holes in trees. Other than this we find no literature references to Tamiops swinhoei in very central China. Tamiops swinhoei swinhoei (Milne-Edwards) Sciurus maccleUandi var. swinhoei Milne-Edwards, 1874, Recherches . . . des mammiferes, 1, p. 308. Tamiops clarkei Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, p. 304. Tamiops maritimus forresti Thomas, 1920, Ann. Mag. Nat. Hist. (ser. 9), 5, p. 305. Tamiops spencei Thomas, 1921, J. Bombay Nat. Hist. Soc, 27, p. 503. Tamiops macclellandi russeolns Jacobi, 1923, Abh. Berich. Mus. Dresden, 16, no. 1, p. 11. Tamiops monticolus olivacetLS Osgood, 1932, Field Mus. Nat. Hist. Publ., Zool. Ser., 18, p. 292. Types. — Sciurus maccleUandi var. swinhoei, type not found in the MNHN in Paris, but there are four possible cotypes taken at Moupin by Pere David in 1870, MNHN 1870-608 (309); 1870-39 (311); 1870-38; and 1870^0 (308); russeolus, three cotypes (skins without skulls) from Tsalila, a pass near Atuntse, between the upper Yangtse and upper Mekong Rivers, SMTB 5887-9 collected September, 1916- 1917 [1914] by Hugo Weigold; clarkei, BM No. 20.1.16.6, adult male from forests in Yangtse Valley, latitude 27° 20' north, 8000 feet col- lected September, 1918, by George Forrest; forresti, BM No. 20.1.- 16.4, adult male from Lichiang Range, Yunnan, 27° 20' latitude north, 11,000 feet collected July, 1918 by George Forrest; spencei, BM No. 20.8.8.6, female from 28° 22' north latitude, 97° 40' east longitude, at 10,000 feet; olivaceous, BM No. 32.4.19.6, adult male from Lo Gui Ho, Tonkin, 5000 feet, collected December 30, 1916 by P. M. Leonard. Material examined, from Tibet. — Tsarong, 7000 feet, Londjre Valley (USNM), one. Material examined, from Sikang, China. — Mupin [Muping] (MNHN), one, (USNM), two; Luting Shan [Lutingkiao] (CNHM), MOORE AND TATE: DIURNAL SQUIRRELS 249 one; Wuchi [Wushi] (CNHM), one; Kulu (CNHM), one; "Ta Chiao," 12,000 feet near Tatsienlu (MCZ), one; 35 miles east of Hokow (ANSP), one; Madischung Valley, 10 miles from Hokow (ANSP), one. Material examined, from Kansu, China. — Chulungapu, 8000 feet, Upper Tebbuland (CNHM), one, (MCZ), six. Material examined, from Szechwan, China. — "Lian-feng-Kiang," Omei Shan (CNHM), one; 25 miles west of Wench wan, 5-9000 feet (AMNH), nine, (USNM), three; Wa Shan (USNM), one; Weichow [Weikiu] (USNM), one; "Tsungshi (Ngoloshi)" (ANSP), one, (MCZ), one. Material examined, from Burma. — Adung Valley, 8-9000 feet, Kachin Prov. (CNHM), three; Imaw Bum (AMNH), two; Road to Chimeli Pass, 10,000 feet (AMNH), two. Material examined, from Yunnan, China. — "Mt. Djinaloko," Yangtze Valley slopes, 10,000 feet (USNM), one; "Hofuping Mts.," Mekong Valley (USNM), eight; "Sila Mts.," Salween-Mekong Di- vide, 8000 feet (USNM), one; N. slopes of Likiang Snow Range (USNM), one; Likiang, 10,000 feet (AMNH), three; 40 miles N. of Likiang (MCZ), one; "Tao Mung Chung" southwest of Lu-tien (MCZ), one; Mu Cheng, 7000 feet (AMNH), three, (MCZ), one; Ngu-luko (CNHM), one; "Yunnan" (CNHM), two. Material examined, from Indochina. — Lo-Gui-Ho, Tonkin (CNHM), three; Chapa, Tonkin (CNHM), five. Pelage color. — Compared with mcclellandi, T. s. swinhoei is often distinguished by its much larger size and long, soft pelage. The general dorsal color is yellowish brown. The black middorsal line is much wider than in mcclellandi, 9 to 10 mm. The width of the pair of black lateral lines is ca. 7 mm., and their length ca. 75 mm. The outer light line is in swinhoei dull yellow brown (scarcely brighter than the yellow brown lines on either side of the median line) . This line is interrupted at the shoulder, not connecting with the cheek line which extends from beneath the ear forward to the whisker patch. The outermost dorsal line is black. Its dimensions are about 28 by 5 mm. The underparts are buffy white with the bases of the hairs slate gray. Discussion. — A circle only 100 miles in radius will include the type localities of russeolus, clarkei, forresti, and spencei, and one of 150 miles radius will include the type localities of these four and that of swinhoei as well. These localities are on high elevations in the 250 FIELDIANA: ZOOLOGY, VOLUME 48 mountains segregated by the deep, north-south river gorges which dissect the general area. Two of the type locaHties, furthermore, those of russeolus and forresti, are not even so separated from each other. We cannot concur in G. M. Allen's (1940, p. 669) acceptance of clarkei as a good subspecies. His distribution map does not in- clude in the range of T. s. swinhoei the Yunnan locality of Mucheng, nor in the range of clarkei the Yunnan locality of Taomungchung southwest of Lutien, although he identified material from these places respectively as T. s. swinhoei and T. s. clarkei. The inclu- sion of Mucheng brings the range of swinhoei 300 miles farther south than he had it shown, and the inclusion of Taomungchung southwest of Lutien would extend his shown range of clarkei nearly 200 miles east. The specimen from Taomungchung seems to us as different from Likiang material as either is from a series of swinhoei from near Wenchwan, and that if it were represented by a series, it would have as good a claim on subspecific difference from typical swinhoei as does the material from Likiang. Another subspecific name for one specimen from one locality could be justifiably added to the list of synonyms by the next author, however, unless some further collect- ing is done in the range of this species to demonstrate the geographic relationships of this and some of the other considerable variation which is already represented by names in the presently available material. Habits. — In a letter of June 10, 1961 Lord Cranbrook contributed the following observations on this species from his field notes of 30 years earlier in the Adung Valley of Burma: "No. 206. Shot in a tree amongst scrub jungle just below snow line (March). When first seen quite low down in bushes and low tree rhododendrons, very unlike the habits of the striped squirrels [T. mcclellandi] seen further south, which keep entirely to high trees. Tail held straight out be- hind, not flicked. No. 229. Shot running along a fence around a field." For other details see Cranbrook's account of the expedition in Kinnear (1934). Tamiops swinhoei vestitus (Miller) Tamiops vestitus Miller, 1915, Proc. Biol. Soc. Washington, 28, p. 115. Type.—USNM No. 199561, adult male from Hsin-lung-shan, 65 miles northeast of Peking, collected February 15, 1915, by Arthur de Carle Sowerby. Material examined, all from Chihli [Anhwei] Province, China. — Tungling (AMNH), six, (CNHM), six, (MCZ), one; Eastern Tombs MOORE AND TATE: DIURNAL SQUIRRELS 251 (AMNH), six, (MCZ), two; Hsing-lung-shan, 65 miles N.E. of Pe- king (USNM), 12; 80 miles E. of Peking (USNM), two. Pelage color. — This race seems to be a pallid offshoot of the moun- tain dwelling subspecies swinhoei. As is also found in Tamiops mari- timus of southern China, there is a striking difference between summer and winter pelage. This is exhibited, for example, by seven specimens taken in April while still in the winter pelage in which only the midstripe is black, and five specimens taken in late July and early August in the summer pelage which differs by the mid- stripe and inner dark pair of stripes all being black. The outer and inner pairs of light lines are whitish, the outer pair whiter than the inner. The underparts are buffy white, their hair bases gray. Discussion. — Jentink (1883) records two specimens of this sub- species in the Leiden Museum from Tingchow, which is apparently 120 miles southwest of Peiping. It is also this form which Fu (1936) reported in the Sung Shan of western Honan. Tamiops maritimus (Bonhote) Definition. — This evident species includes subspecies maritimus, monticolus, hainanus, and moi, and see the synonyms of these. The known distributions of maritimus and monticolus are shown in Fig- ure 19, those of hainanus and moi in Figure 20. Diagnosis.— The pelage is comparatively short and thin, the gen- eral coloration tends to be more olive, and the inner pair of light stripes tends to be nearer the color of the nape, than in the other species of Tamiops. Relationships to other species. — This is a relatively low altitude species occupying the southeastern coastal region of China and all of Vietnam and Laos, and according to G. M. Allen (1940) appar- ently penetrating to middle China. At the southern end of its range it ascends the Langbian Peaks to considerable elevation, but it is evidently also found at the foot. At Pak Hin Bun, Laos, Robinson found it along the Mekong River and on islands in the River. Al- though we have examined specimens of it from two other Laos local- ities along the Mekong, Vientiane and the westernmost locality, Lo Tiao, it is not known from west of that river. Its geographic rela- tionships to Tamiops swinhoei east of the Mekong in China remain too much a mystery. It is at least of interest that Thomas (1920, p. 305) once thought that maritimus penetrated Yunnan to the Liki- ang Range. When it is discovered what the distributions of these striped squirrel species are in central and southern Yunnan, it should 252 FIELDIANA: ZOOLOGY, VOLUME 48 become possible to interpret with greater confidence the information already available from northern Yunnan and nearby Burma, Sikang, and Szechwan. -■"7 +«*N + + + \ WE (orE I06'E I09E \., YUNN-AN ,--v^.,^KWAKG5I v./'-. ( > LA03 -^, Y 1^ r THAILAND N., ro ^'^N, SOU.'BtCHlU^SEA^ [ 4-15'M + )+ f \ + 1 ■%. 1 4 \ I ^ C^ \ A-iz'H f \ 0 ^'v CAMBODJA j i + + 1/+ 0 + MILES "^ SW r 1 ' i' 1 r 500 KILOMETERS Fig. 20. Southern part of distribution of the maritime striped tree squirrel species, Tamiops maritimns, as determined by plotting collecting localities of mate- rial examined. Subspecies: A, maritimus; B, monticolus; C, hainanus; and D, moi. Tamiops maritimus maritimus (Bonhote) Seiurus macclellandi maritimus Bonhote, 1900, Ann, Mag. Nat. Hist. (ser. 7), 5, p. 51. Seiurus mjxcclellandi formosanus Bonhote, 1900, Ann. Mag. Nat. Hist. (ser. 7), 5, p. 52. Tamiops sauteri J. A. Allen, 1911, Bull. Amer. Mus. Nat. Hist., 30, p. 339. MOORE AND TATE: DIURNAL SQUIRRELS 253 Types. — Sciurus m. maritimus, BM No. 94.9.1.11, adult from Foochow, Fukien, China, collected April 1893 by C. B. Rickett; formosanus, BM No. 62.12.24.18, adult female from northern For- mosa, collected April 1862 by Robert Swinhoe; sauteri, AMNH No. 31621, adult male from Chip-Chip, northern Formosa, collected November 1908 by H. Sauter. Material examined, from F^ikien Province, China. — Yuki (AMNH), three, (CNHM), two, (MCZ), two; Fuching Hsien (AMNH), 10, (MCZ), one; Kucheng (CNHM), one; 70 miles S.W. of Yen-Ping- Fu [Yenping], 500 feet (USNM), six; "N.W. Fukien" (MCZ), two; "Fukien" (CNHM), two. Matrial examined, from Formosa. — Kagi Dist., Central Formosa (CNHM), one; Chip-Chip (AMNH), five; Shineigun (MCZ), one; "Formosa" (CNHM), one. G. M. Allen (1940) reports a specimen in the British Museum from Chiong Lok, 100 miles west of Swatow, Kwangtung, and we have included this on our map. Original description. — "This is the form which most nearly ap- proaches the typical Sc. MacClellandi, from which it differs in being far greyer and more concolorous. The median dorsal stripe is by no means well marked and very short, not being continued to the root of the tail. The two subdorsal [inner light] stripes are of the same colour as the back, while the outermost light stripes are of a dull white, very narrow and short, not being continued to the root of the tail, and only starting at the shoulder." Tamiops m. maritimus, as we know it here, appears to occupy the vegetation area which Chi-Wu Wang (1956) has mapped as Rain Forest. Tamiops maritimus monticolus (Bonhote) Sciurus macclellandi monticolus Bonhote, 1900, Ann. Mag. Nat. Hist. (ser. 7), 5, p. 52. Type.— BM No. 97.3.2.6, adult from Ching Feng Ling, 1500 to 2000 feet, 100 miles northwest of Foochow, Fukien, China, collected December 1896 by C. B. Rickett. Material examined, from Fukien Province, China. — Chungan Hsien (AMNH), 17, (MCZ), four; mountains near Yenping (AMNH), 11, (MCZ), one, (USNM), four, (CNHM), six. Material examined, from Kwangsi. — Yao-shan (CNHM), two. Because of loss of data from our own examinations of material in European museums on this species, we are departing in this instance 254 FIELDIANA: ZOOLOGY, VOLUME 48 from the practice of citing and mapping localities only of material examined, and are including here and mapping the additional records reported by G. M. Allen (1940, pt. 2, p. 684): Material examined, from Fukien. — Pucheng (BM), two; Kuaton [Kaotien] (BM), eight; Chungfengling [Chingfengling] (BM), eight; Kienyang (BM), one; Tingchow (BM), two. Material examined, from Anhwei. — Chinteh [Tsingteh] (BM), 10. Material examined, from Hupeh.^ — Changyang (BM), one. Material examined, from Chekiang. — Ningpo (BM), one. Original description. — "Brighter than [maritimus] to which it is most nearly allied, and from which it differs in having the median dorsal stripe more distinct and always continued to the root of the tail. The outermost light stripes are very broad and distinct and continued to the root of the tail." Discussion.— We find contrary to G. M. Allen (1940, pt. 2, p. 684) that Bonhote was right in differentiating between the coastal samples of this species and those inland in the mountains. However, he did err in trying to distinguish them primarily upon a character of the middorsal stripe. The better character which he merely mentions is that the outer pair of dorsal light stripes continues posteriorly to the base of the tail and are broader and more distinct throughout than in maritimus. Bearing in mind, of course, the pelage differences related to season, we note that in the summer monticolus material from the mountains about Chunganhsien, the dark stripes are gen- erally blacker than in the winter material from there; although there is also one taken on December 9 from the mountains near Yenping, on which the dark stripes are quite as black as any with summer dates. Comparing only winter series, six January and three March maritimus specimens from Fuchinghsien with five December, one January, two February, and one March monticolus ones from near Yenping, we find the two series easily distinguished from each other. Eight of the nine Yenping ones in winter pelage are individually dis- tinguished when mingled in a series of maritimus. The ten maritimus from Fuchinghsien and three from Yuki in winter pelage remain equally distinguishable when placed in the series of eight Yenping monticolus. The Yenping material in winter pelage is also consist- ently redder in the dark stripes and its inside light stripes are lighter and huffier than the nape whereas those of the Fuchinghsien material are the olive gray of the nape. These latter characters are not dis- played by the four Chunganhsien winter pelage examples of mon- ticolus, however. This may be a formerly more distinct northern MOORE AND TATE: DIURNAL SQUIRRELS 255 subspecies of maritimus in the process of being obliterated, or our material may be from a zone of intergradation between the two good subspecies. Tamiops maritimus monticolus appears to occur in an area which coincides rather well with the extent in eastern China of two of Chi- Wu Wang's (1956) general vegetation types: 1. Evergreen broad- leaved forest of evergreen oaks, schima and laurels, with Pinus mas- soniana in secondary stands. 2. Mixed mesophytic forest. Dimensions. — In six maritimus from 500 feet elevation 70 miles southwest of Yenping, Fukien, taken in November and December of 1921, and four monticolus taken in April of 1922 at 2000 feet eleva- tion near Yenping, all by Arthur de Carle Sowerby, hind foot and head-and-body size show no differentiation, but five of the maritimus have tails recorded between 120 and 125 mm., whereas the longest three of the monticolus range from 110 to 117 mm. Tamiops maritimus hainanus (J. A. Allen) Tamiops macclellandi hainanus J. A. Allen, 1906, Bull. Amer. Mus. Nat. Hist., 22, p. 476. Tamiops macclellandi riudoni J. A. Allen, 1906, Bull. Amer. Mus. Nat. Hist., 22, p. 477. Tamiops macclellandi laotum Robinson and Kloss, 1922, Ann. Mag. Nat. Hist. (ser. 9), 9, p. 92. Types. — Tamiops m. hainanus, AMNH No. 26664, an adult fe- male from Lei Mui Mon, mountains of Hainan Island, China, col- lected December 31, 1902, by agents of Alan Owston; riudoni, AMNH No. 26672, an adult male from Riudon, lowlands of Hainan Island, China, collected March 5, 1903, by agents of Alan Owston; laotum, BM No. 26.11.17.6, adult male from Pak Hin Bun, Mekong River, Laos, collected March 2, 1920, by Herbert C. Robinson. Material examined, from Hainan, China. — Nodoa (AMNH), 57, (CNHM), 14, (MCZ), six, (USNM), one; Nam Phong (AMNH), 10, (CNHM), four, (USNM), one; Riudon (AMNH), two; Lei Mui Mon (AMNH), nine, (USNM), two; Kachek (USNM), one. Material examined, from Tonkin, Vietnam. — Pakha [Pa Kha] (CNHM), one; Muong Boum (CNHM), six; Lieng San [Leng Sang] (CNHM), five; Nong Lum (CNHM), three; Chapa [Cha Pa] (CNHM), seven, (USNM), two, (MCZ), one; Phong Tho (CNHM), one; Muong Mo (CNHM), seven, (USNM), one, (MCZ), one; Ba Nam Chi (CNHM), one; Lai Chau (CNHM), two, (USNM), two, (AMNH), one; Muong Moun (CNHM), five, (AMNH), two; Pa 256 FIELDIANA: ZOOLOGY, VOLUME 48 Ham (CNHM), two; Bac Tan Tray (AMNH), one, (ANSP), one; Moung Mouen (ANSP), two. Material examined, from Annam, Vietnam. — Hoi Xuan (CNHM), four. Material examined, from Laos. — Lao Fou Tchay (CNHM), one; Phong Saly (CNHM), five; Muong Yo (CNHM), five; Bonn Tai [Bun Tai] (CNHM), one; Lo Tiao (MCZ), two; Xieng Khouang (AMNH), two, (MCZ), three; Tha Ng'on, Vientiane (CNHM), one; Saravane (CNHM), one; Phu Kobo (MCZ), three; Col de Taloun (MCZ), two; "Indochina" (CNHM), one. The extensive series in winter pelage now available from Hainan and Tonkin show no color difference from the winter pelage material of maritimus, but as G. M. Allen (1925, p. 7) pointed out and Osgood (1932, p. 290) amplified, there is a size difference. Hind feet in mari- timus measure about 32 to 35 mm. in the dried skins, and those of hainanus measure about 28 to 30 mm. in the dried condition. The skulls of maritimus are also larger. No other examples of the reddish riudoni have appeared to justify the supposition that this represents a lowland race, and the type apparently is a strongly erythristic in- dividual. Robinson and Kloss (1922) attributed no diagnostic characters to their laotum, and although Osgood (1932, p. 291) retained it as "well characterized by very pale color which reaches its extreme develop- ment in southern Laos," he had only one specimen from southern Laos. Tamiops maritimus moi (Robinson and Kloss) Tamiops macclellandi moi Robinson and Kloss, 1922, Ann. Mag. Nat. Hist, (ser. 9), 9, p. 92. Type.— BM No. 26.11.17.7, adult male from Langbian Peaks, 5500 to 6500 feet, southern Annam, collected April 25, 1918, by C. Boden Kloss. Material examined, all from southern Vietnam. — Pie de Langbian (MCZ), four; Langbian Peak (MCZ), two, (CNHM), 10; Dalat (CNHM), one; Gougah (MCZ), one, (CNHM), two. Original description. — "Like T. m. laotum, but darker above; up- per parts more suffused with ferruginous, so that in addition to being more rich-coloured generally, the yellow stripes are ochraceous in- stead of buff, while the rump and the outer pair of dorsal stripes are a brighter brown." MOORE AND TATE: DIURNAL SQUIRRELS 257 Discussion. — This is only a faintly distinguishable subspecies. The winter pelage is like that of hainanus excepting for the two dark paired dorsal stripes being faintly redder. We find that the hind foot of moi is larger than that of hainanus, however, in the seven MCZ specimens, which range between 31 and 32.2 mm. in the dried skins. The skull does not appear to be appreciably larger than that of hainanus. PLAIN LONG-NOSED SQUIRRELS Genus DREMOMYS Heude Dremomys Heude, 1898, Mem. Hist. Nat. Emp. Chinois, 4, part 2, p. 54. Zetis Thomas, 1908, J. Bombay Nat. Hist. Soc, 18, p. 244. Type species. — Dremomys, Sciurus pernyi Milne-Edwards; Zetis, Sciurus rufigenis Blanford. Definition. — The genus Dremomys consists of four species of the Indochinese Subregion, lokriah, pernyi, rufigenis, and pyrrhomerus, and one species of the Malaysian Subregion, everetti. See Figure 21. Diagnosis. — The genus Dremomys possesses the following char- acteristics: (1) There is one bony septum crossing the chamber of the auditory bulla. (2) The teeth are orthodont or slightly proodont. (3) The length of the nasal bone exceeds the least interorbital breadth. (4) The upper cheek teeth are not specialized with an extremely deep valley that remains as a dirt-filled lake after the cusps have been planed away by use. (5) The coronoid process of the mandible is well developed and strongly falcate. (6) The baculum is composed of two separate parts, shaft and blade. (7) The dorsal pelage has no longitudinal light stripes. The genus Dremomys may be distinguished from other genera of Sciurinae of the Indian and Indochinese subregions by the above characteristics as follows: from Ratufa by 1, 2, 3, and 6; Funambulus by 5, 6, and 7; Callosciurus by 3; Tamiops by 3 and 7; Menetes by 4, 5, and 7; and Sciurotamias by 1 and 2. Systematic history. — Dremomys was poorly characterized by Heude. Thomas (1908) discussed the genus under the name Zetis in greater detail, and gave a key to the species (pp. 248-249). Later Thomas (1916a) published another synoptic key using the generic name Dremomys. Pocock (1923) and Ellerman (1940) gave the genus full standing, and Archbold and Tate (1935, p. 1, footnote 1) designated Dremomys pernyi as the genotype. Besides the species pernyi, Heude included three other species in the genus which he described as new: saltitans, collaris and latro. The animals to which these three names were applied, however, have since been placed in the genus Sciuro- 259 260 FIELDIANA: ZOOLOGY, VOLUME 48 ^OUrHEA^r A3IA ^^Y OF 3EN(^Al. SPECIES OF DREMOMYS ■ lokriah ^ pernyi Srufigenis BpyrrhomcruS fKkvcretti 0 KI10MET£RS SOO Fig. 21. Composite, slightly generalized distribution of the five species of the long-nosed squirrel genus Dremomys. tamias (G. M. Allen, 1940, p. 646). It was recognized in the early nineteen hundreds that Dremomys was the proper genus for the spe- cies lokriah and rufigenis, and various authors have at one time or another (correctly) placed the species everetti and pyrrhomerus in this genus. These four species and the original one pernyi constitute the genus Dremomys. Range. — On the continent it is a wide-ranging genus, extending from Nepal in the Himalayas 2000 miles east across the breadth of China to the coastal province of Chekiang, and from only 250 miles above the equator in the Malay Peninsula north through Siam, Burma, Indochina and China nearly 2000 miles to northern Szech- wan. It occurs also on Borneo, Hainan, and Formosa. Dremomys is a long-nosed squirrel, characteristically with plain, drab dorsal pelage. It is generally reported to be terrestrial or semi- terrestrial in habit, and to occur only in the mountains or hills where Fig. 22. Skull and left mandible of the Himalayan long-nosed squirrel, Dremomys lokriah, AMNH No. 114936, Xl. Fig. 23. Skull and left mandible Xl of the Chinese long-nosed squirrel, Dremomys pernyi, AMNH No. 43999, but lower jaw of AMNH No. 114938. 261 O ^ I pi, i.s 0) O S S.S t> I 0) CO D. ^ a s s -^ O oooooot>oot-oooooooooas050505ajo5asoo "S)J3 UtiC^JIMt— ©Otra(MOUtiOO-^l J^ -t=-§ t>ooo5t-a5«ot- •oooot-'rH«DTj<»-(eoifii-«s<-«i<(M oOw le "ti(Meooo(Mec«DOO(M--l'-l'-l,-l ■ueB ■■^THOt-CS«Dt-0>t000«DO"^l'3Wc0rH'^»-IC0 be 2 (U^ •,-(T-li-l.-lT-lrHrHrt^.-lrHi-l(Mr-(t-l,-IC^^,H^ jjj — ,^ 10-^«00 •t>O(M00t-C(m;d«doo • -t-ooiflt^eoo -"^ • ■qG -oooooseoot-eo -TjitoeOTfeot- -t- .S o -co 0) O "^ -H «Ot-000000'-IOO"3rJ<«DOiO'-IOOOT-lTjt^OiOOOt>050000C<3(Mi;DOOOiOi-(rH«D(M«0 i-Hi-HtHi— IC> -o O (D CO o -^ w _ fl .5 a> Q ° 1—1 QJ < s 4J> ^ 00 t- (M 00 ix> 00 00 00 00 ^ •f.-y 0.-lrHtraCO(Mt-40r-lU500 bfee 00?D«0i:D«0t-ut>t-«000O j|j2 -^t-ioosoicot-moooio X CO -gj"^ 00,-((N00«0OOC0<0(M«D ^1^ T-l,-(,-l,_(i-(r-l,-(»Hi-(i-(T-l (ij'^;g «Dt~oo(MC^ -oo ■■^,-1 i^5S ooii-Hoo -oico -c^jcc pq — 43 cs-tr'. ^c^ t- t-oo o • -u bo Ofi i-HOi -00 ■ ix> ^^ •«£> T3 -tJ Wti. -H + -^ (M 00 tr- io lo uTi ira -H -H -H ^ 0050iO'-tOOiOiOOO«D Eh y-l ,-< r-t ,-, ^ y-> ,-|_i^,-| -co -H o to o 00 <7i O OJ tH 00 t- (M »-H OJ T-( I— I -H -H -H O O O O lO ■<* O ;D O OS i-H CO CO tH (M 1-1 CO IM 5 s i 1 ? to -ST 1 s ?^ ^ S ■g PS. -s 8 -« -^ 3 cii bfi 3 o Eh 298 MOORE AND TATE: DIURNAL SQUIRRELS 299 around Maliwun it was occasionally seen running across tracks and among long grass, and bamboo scrub, especially in the early eve- nings, but I have also seen it in the thickest forest. It is very like Tupaia in its movements, hiding at the smallest noise and not read- ily making a second appearance. Weight — 73^ ozs." (G. C. Short- ridge in Wroughton, 1915b, p. 713) Menetes berdmorei peninsularis Kloss Menetes berdmorei peninsularis Kloss, 1919, Jour, Nat. Hist. Soc. Siam, 3, p. 375. Type.— BM No. 26.10.19.5, adult male, from Ban Koh Klap [Ban Na], Nakon Sritamarat [Nakhon Si Thammarat], near Bandon, pen- insular Thailand, collected July 3, 1913, by H. C. Robinson and Cecil B. Kloss. Material examined. — Kampenpet [Ban Kamphaeng Phet, latitude r 11' N.], Thailand (BM), one. Although the type was described as having the median and lat- eral black lines strongly marked, it has been previously pointed out (Robinson and Kloss, 1915, p. 121) that the Menetes from elsewhere (Bandon) are extremely variable in this respect. Menetes berdmorei moerescens Thomas Menetes berdmorei moerescens Thomas, 1914, Jour. Bombay Nat, Hist. Soc, 23, p. 24. Type. — BM No. 6.11,6.32, young female from Bali, near Nha- trang, latitude 12° N., south Annam, Material examined. — Kontoum [also, Kon-toum and Kontum], Annam (MNHN), two, (BM), 4; Dak-To [also Dak- to], Annam (MNHN), one, (BM), two; Ban Me Thuot [Buon Ma Thuot], An- nam (CNHM), four; "Eaktur," E. of Ban Me Thuot, Annam (CNHM), two; Ninh Hoa, Annam (CNHM), one. Original description. — "Markings about as in decoratus, but less conspicuously contrasted, owing to general body colour being darker and duller, more olive brown, underside and tips of tail hairs yellow- ish. Size rather larger than in other forms. Muzzle of skull unusu- ally slender." Menetes berdmorei decoratus Thomas * Menetes berdmorei decoratus Thomas, 1914, Jour. Bombay Nat, Hist. Soc, 23, p. 23. 300 FIELDIANA: ZOOLOGY, VOLUME 48 Type. — BM No. 14.4.3.4, old female, from Mt. Popa, south cen- tral Burma, below 4000 feet, collected April 20, 1913, by G. C. Shortridge. Material examined, all from Burma. — Mt. Popa, 4961 feet (BM), eight; Mt. Popa (AMNH), five, (CNHM), two (topotypes); Pegu Yoma Range, north of Rangoon (AMNH), eight; North Zamayi Res., 70 miles north of Pegu, 700 feet (BM), one; South Zamayi Res., 60 miles north of Pegu, 500 feet (BM), one; 30 miles N.W. of Toun- goo, Burma (CNHM), two. Original description. — "Median dorsal and upper lateral dark streaks prominent, all the markings very strongly defined, the main dark lateral band broad and glossy black; an additional blackish streak edging the belly; general body colour clear grizzled olive, undersurface and tips of tail hairs pure white." Discussion. — A glance at the map shows that the Pegu Yoma range is a southward extension from the prominent Mt. Popa. Speci- mens from these mountains tend to have the black striping heavily accentuated. It appears likely that the present race decoratus is bounded on the west by the Irrawaddy River and on the east by the Salween River. Eastward beyond the Salween the relatively less striped race berdmorei occurs. Intensification of the black striping recurs elsewhere only in pen- insularis and moerescens, both of which are from mountainous regions of heavy rainfall. Habits. — This form has been observed in the field and commented upon as "Occurring on Mt. Popa among rocks and stones, that are surrounded by thick scrub, and often close to cultivation, up to 4,000 feet. Very shy, running into holes and crevices at the slightest sound or movement. It is essentially a ground squirrel, seldom, if ever, ascending trees, though by no means confining itself to open coun- try." (G. C. Shortridge, in Wroughton, 1915a, p. 474) Menetes berdmorei consularis Thomas Menetes berdmorei consularis Thomas, 1914, Jour. Bombay Nat. Hist. Soc, 23, p. 24. Type. — BM No. 2.6.6.6, young female, from Nan [Muang Nan], 200 meters, Siam, collected October 6, 1901, by T. H. Lyle. Material examined, from Thailand. — "Kowpla," Paknampho [Ban Pak Nam Pho], Nakonsawan (USNM), one; Airawan Mtn. [Khao Erawanl, Lop Buri (USNM), one; Mt. Angka [Doi Inthanon], 4300 feet [1310 meters] (MCZ), six, (CNHM), one; "central Siam" MOORE AND TATE: DIURNAL SQUIRRELS 301 (MCZ), one; Doi Nangkeo [Doi Phi Phan Nam] (MCZ), two; Chien- grai [Muang Chiang Rai] 410 meters (BM), three; "Tahkamen" [west of Bangkok] (BM), one; halfway between Pichet [Ban Nai Muang] and Paknampo [Ban Pak Nam Pho], 33 meters (BM), one; 100-125 miles north of Bangkok, 20-25 meters (BM), two; "Nam Phi" Nan, 225 meters (BM), one; Nakon Sawan [Nakhon Sawan], 29 meters (BM), one; Me Wong (river), 53 miles east of Um Pang [Ban Le Kathe] (BM), three, (AMNH), three; Koon Tan [Doi Kuhn Tan, or Sathani Kuhn Tan] (NR), one; Chum Poo [Sathani Tha Chomphy] (NR), one; "Vieng Nun," northern Siam (NR), one; Muang Prom [Muang Phrom Buri] (BM), two; Me Ping River (AMNH), one; below Paknampo [Ban Pak Nam Pho], 26 meters (BM), one; Melamoung [Me Lamung] (AMNH), one; north of Raheng [Ban Rahaeng] (NM), one; Raheng [Ban Rahaeng], 120 meters (BM), one; Sokotai [Ban Thank], 64 meters (BM), one; 20 miles west of Kempanpet [Changwat Kamphaeng Phet] (AMNH), one. Material examined, from Burma. — Taok, Thuaungyin Valley [Thaungyin Valley], 1100 feet (BM), one; Myawadi [Mya Wadi], Tenasserim (BM), two; Kawkereik [Kawkareik], Tenasserim (AMNH), one. Original description. — "No median dorsal stripes or upper dark lateral ones, the only dark streak being that between the light lat- eral streaks, and even this is absent in January and February. Under surface yellowish white." Menetes berdmorei mouhotei (Gray) Sduriis mouhotei Gray, 1861, Proc. Zool. Soc. London, 1861, p. 137. Sciurus pyrrhocephalus Milne-Edwards, 1867, Rev. Mag. Zool. (ser. 2), 19, p. 225. Menetes berdmorei koratensis Gyldenstolpe, 1917, Handl. K. Svenska Vetensk. Akad., 57, no. 2, p. 39. Types. — Sciurus mouhotei, BM No. 61.4.12.13, from Cambodia; pyrrhocephalus, MNHN No. 1864-682, adult female, collected in Saigon, Cochin China, by Rudolphe Germain; koratensis, NR No. 4, adult male, from Sakerat [Ban Chakkrarat], east Siam, collected January 9, 1912, by Nils Gyldenstolpe. Material examined, from Thailand. — "Khow Wing," Siracha [Ban Si Racha], Chonburi (USNM), one; Satahip (Ban Sattahip], sea level (BM), one; Chantabun [Chanthaburi] (USNM), four, (AMNH), four; 302 FIELDIANA: ZOOLOGY, VOLUME 48 Klong Yai (BM), two, (USNM), three; Lem Ngop [Ban Laem Ngop] (BM), three, (USNM), one; Ok Yam [Ok Pyam] (BM), one, (USNM) three; Koh [Ko] Kut I. (USNM), six, (BM), six; Koh [Ko] Chang I. (BM), one, (USNM), two; Sriracha [Ban Si Racha] (USNM), three, (CNHM), one; Hup Bum [Ban Hup Bum], 500 feet (152 meters) (BM), two; Ban Sadet [Ban Phan Sadet] near Sriracha (USNM), one; Nonokhor [Ban Nong Kho], near Sriracha (USNM), one; "Nong Mong," Muong Karbin [Ban Kabin Buri] (USNM), one, (BM), one; Kao Sabab [Khao Sa Bap] (USNM), one; Pak Jong [Ban Pak Chong] (USNM), one, (AMNH), three, (MCZ), one; Aranya [Ban Aranya- prathet] (USNM), one; Lat Bua Kao [Ban Lat Bua Khao] (USNM), six; Sakarat [Ban Chakkrarat] (NR), one; 50 miles south of Bangkok, seacoast (BM), two. Material examined, from Indochina.^ — Plateau Bolovens, Laos (AMNH), one; An-Bierh [An Binh] (BM), one; Honquan [Hon quan] (BM), two, (CNHM), one; Phu-Quoc I. (BM), one; Tay ninh, 100 feet (30 meters) (BM), two; Lagna River (AMNH), one; Dinquan [Din Quan] (USNM), one; "Cambodia" (BM), two; Sien- Reap [Siem reap] (MNHN), one; Angkor, 150 feet (46 meters) (BM), one; Cam Chay [Kam Chay] Mts. (BM), two. Original description. — "Grisled grey-brown, with pale rings; lips, chin, throat, and under side of body and inside of limbs white; upper part of the sides with a longitudinal black streak, edged above and below with a narrow white line; tail blackish, whitish washed, hairs elongate, brown, with two broad black rings and a white tip. . . ." Habits. — Little seems to be known about the natural history of Menetes. Gyldenstolpe (1914, p. 15) makes an interesting comment, which in his experience appears to apply to this form and consularis. "This species was very common in the dry forests both in Northern and Eastern Siam. It was always observed near the villages, or in the compounds in the towns, but never far into the jungles." CHINESE ROCK SQUIRRELS Genus SCIUROTAMIAS Miller Sciurotamias Miller, 1901, Proc. Biol. Soc. Washington, 14, p. 23. Rupestes Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, p. 398. Type species. — Sciurotamias, Sciurus davidianus Milne-Edwards; Rupestes, Rupestes forresti Thomas. Definition. — The genus Sciurotamias is constituted by the two monotypic subgenera Sciurotamias and Rupestes, both endemic to northern China. Diagnosis. — (1) There are two dorsal processes on the zygomatic process of the squamosal. (2) The ectopterygoid ridge of the ali- sphenoid is obsolescent. (3) The upper incisors are strongly opistho- dont, (4) Three bony septa cross the chamber of the auditory bulla. The above characters all distinguish Sciurotamias from each of the other genera of the Sciurinae of the Indian and Indochinese sub- regions. Subgenus SCIUROTAMIAS Miller Sciurotamias Miller, 1901, Proc. Biol. Soc. Wachington, 14, p. 23 Type species. — Sciurus davidianus Milne-Edwards. Definition. — The subgenus Sciurotamias includes only the type species davidianus, a polytypic, montane, ground squirrel of north- ern China. Diagnosis. — The subgenus Sciurotamias apparently may differ from subgenus Rupestes in several notable characters: (1) The squa- mosal is high. (2) The zygoma ascends to a point well above halfway on the height of the rostrum. (3) The superior process of the jugal is very low and anterior to the midlength of the orbit. (4) There is no pronounced temporal ridge. (5) The postorbital process of the frontal is more than 2 mm. long. (6) There is a peg-shaped third upper premolar. (7) There are generally faint whitish postauricular streaks on the pelage but no whitish longitudinal stripes on the sides of the body. 303 304 FIELDIANA: ZOOLOGY, VOLUME 48 Fig. 32. Skull and left mandible of the montane rock squirrel of China, Sci- urotamias davidianus, AMNH No. 45370, X 1. Note indications of the three trans- bullar septa which distinguish this genus from the Oriental tribe Callosciurini and the Holarctic tribe Sciurini as well. Sciurotamias davidianus Milne-Edwards Definition. — The species davidianus is composed of subspecies davidianus and consobrinus, wiiich include several named forms here as synonyms. The geographic distribution known to us from speci- mens examined is shown in Figure 33. Diagnosis. — Since the subgenus is monotypic, the diagnosis of the species is identical to that for the subgenus. Relationships to other species. — Sciurotamias is said to inhabit cliffs and rocky ground of the mountains avoiding trees, and nesting in deep crevices. It would be interesting to know the extent to which it differs from Dremomys p. pyrrhomerus in habitat. The two seem not to have been collected at the same localities; although their ranges appear to be closely approximated. G. M. Allen (1940, p. 646) has stated that Dremomys is arboreal in contrast to Sciurotamias, but he evidently inferred this from Dremomys being taken in deep forest. References to its habits in Malaya and Borneo indicate that MOORE AND TATE: DIURNAL SQUIRRELS 305 ^IKANi^ ■\ ,.^JEHOL /-• (' / •^ -.../•• A :'■'"/ A^/'hoPEH ("^'f^iKLt •4 ( ^*')SHMJSlY'--^\) KAN^U / B ••vJ->^-X, t/ ^ZECHWAN r^-" HUFEH f •-.'•'' •'.'■^^ 1 .1 YUNNAN •^•'•> KWEICHOW ; .,^..^, >— '"^' p KILOMCTtRS i^ 0 ■^— , MILK ,\. I -i Fig. 33. Geographic distribution of the Chinese rock squirrel, Sciurotamias davidianus (round symbols), and Chinese cliff squirrel, Sciurotamias (Rupestes) forresti (square symbols), as revealed by plotting the collecting localities of mate- rial examined. Subspecies of species davidianus: A, davidianus; B, consobrinus. it inhabits forested areas of mountains, but is primarily terrestrial there. In 1934, at a single locality in the mountains 25 miles west of Wenchwan, Szechwan, T. Donald Carter collected Dremomys pernyi and Sciurotamias d. consobrinus. Since this is the only place which we have located where both of these species are known to have been collected, his comments on their habitats seem especially significant. He recalls clearly (September 1956) in conversation over the collected material laid before him, that the Dremomys pernyi in- habited the steep, forested slope of the narrow valley of Chengou Creek, and he recalled them particularly in the second growth near 306 i^'%i FIELDIANA: ZOOLOGY, VOLUME 48 the foot of the slope where there had been some removal of timber. They did ascend trees, he said, but seemed to be on the ground a good deal, and he collected them on the ground, particularly among moss-covered logs of down timber. The Sciurotamias lived in small areas of rather level, boulder-strewn grassland where the narrow valley bottom widened out occasionally. There were no trees on this, and Sciurotamias lived here like a ground squirrel. The tree- less character of these level areas along the creek bottom Mr. Carter attributed to perhaps seasonal flooding by the torrential mountain stream. The evidence presented in the accompanying maps shows that the ranges of both Dremomys pernyi and Dremomys pyrrhomerus lie to the south of the distribution of Sciurotamias davidianus (excepting for the isolated example of the latter from Hwangtsaopa, Kweichow) . The above example of apparently sharp segregation into two very different biotic communities in an area of overlap of species range, may not, however, be a fair indication of the variety of habitat uti- lized by either species in the parts of its range which are not shared with closely related species. It may be worth noting in this connection that the distribution of Sciurotamias davidianus shown here coincides with the long north- east to southwest portion (but not the southeast) of the "Deciduous broad-leaved forest predominated by deciduous oaks" mapped by Wang (1956, p. 513) and that that of S.forresti lies in an area in which Wang shows two alternating vegetation types: 1. Tundra and alpine vegetation, and 2. Montane coniferous forest predominated by spruce and fir. Dremomys p. pyrrhomerus occurs south of this in the area that Wang maps as "Evergreen broad-leaved forest of evergreen oaks, shima, and laurels, with [pines] in secondary stands." D. p. pernyi is sympatric with S. forresti but apparently allopatric with S. davidianus. D. p. flavior is allopatric with D. p. pyrrhomerus but occupies a more western part of the evergreen broad-leaved forest area as mapped by Wang. Although Sciurotamias is as long-snouted proportionately as Dre- momys lokriah, for example, a skull of Sciurotamias is readily distin- guished from those of Dremomys species by being broader, arched, and flattened, so that it is shaped quite like skulls of the chipmunk, Eutamias. The posterior upper premolar is notably smaller than any of the upper molars in Sciurotamias whereas in Dremomys, its size does approximate that of one or more of the molars. The fora- men ovale is circular in Sciurotamias, narrowly oval in Dremomys. MOORE AND TATE: DIURNAL SQUIRRELS 307 Sciurotamias has three bony septa across the chamber of the auditory bulla; whereas members of the tribe Callosciurini have one or none. The upper incisors are strongly opisthodont in Sciurotamias; where- as those of members of the Callosciurini are orthodont or, more fre- quently, proodont (defined in Moore, 1959, p. 162). Sciurotamias differs also in having cheek pouches. See Table 15 for an indication of the body and skull dimensions of species davidianvs. Sciurotamias davidianus davidianus (Milne-Edwards) Sciurus davidianus Milne-Edwards, 1867, Rev. et Mag. Zool. (ser. 2), 19, p. 196. Dremomys latro Heude, 1898, Mem. Hist. Nat. Chinois, 4, pt. 2, pp. 54-55. Types. — Sciurus davidianus, MNHN No. 1863-655, collected by Pere Armand David in the "mountains of Pekin," Hopei, China; latro, not seen, but thought to be from Shantung, China (Allen, 1940, p. 662). Material examined, from Hopeh, China (except as noted). — "Chi- hli" (NR), 15; near Ching Wang Tas [Chinwangtao] (USNM), 11; Hsin-lung-shan, 65 miles N.E. of Peking [Jehol, Manchurial (USNM), two; Wu-ting-shan, 75 miles N.E. of Pekin [Jehol, Manchuria] (USNM), two; Western Hills, 15 miles W. of Peking (USNM), one, (RNH), one; Tung-Hng (USNM), one, (AMNH), three, (CNHM), four; Eastern Tombs (AMNH), three, (CNHM), one. Material examined, from Shansi, China. — Mts. 50 miles N.W. of Tai-yuan-fu [Yangku] (USNM), four; Mts. 10 miles S. of Wu-tsai [Wutai] (USNM), one; Mts. 70 miles N.N.W. of Tai-yuan-fu [Yang- ku] (USNM), three; "He-shuin" (MCZ), one; "Sjol" (NR), five. This subspecies is distinguished from the southern race by its lighter, grayer dorsal color, about Deep Grayish Olive, and by its grayish instead of buffy-brownish under parts. The race is appare- ently confined to territory north and east of the Hwang-Ho River. The postauricular patches appear to be less prominent in this race, and the posterior streaking from them, mentioned above, is evident less frequently. We follow Allen (1940, p. 662) in synonymizing latro with da- vidianus. Sciurotamias davidianus consobrinus (Milne-Edwards) Sciurus consobrinus Milne-Edwards, 1868-74, Recherches pour servir a I'Hist. Nat. des Mammiferes, p. 305. 308 FIELDIANA: ZOOLOGY, VOLUME 48 Dremomys collaris Heude, 1898, Mem. concern. I'Hist. Nat. de I'Emp. Chinois, 4, part 2, p. 55, pi. 12, figs. 2-2c. Dremomys saltitans Heude, 1898, Mem. concern. I'Hist. Nat. de I'Emp. Chinois, 4, part 2, p. 55, pi. 12, figs. 4-4c, Sciurotamias owstoni J. A. Allen, 1909, Bull. Amer. Mus. Nat. Hist., 26, p. 428. Sciurotamias davidianus thayeri G. M. Allen, 1913, Mem. Mus. Comp. Zool., 40, no. 4, p. 231. Types. — Sciurus consobrinus, not found at MNHN, a specimen from Muping, Szechwan, China, collected by Pere Armand David; collaris, not seen, possibly at the Sikawei Museum in Shanghai, China; saltitans, not seen, possibly at the Sikawei Museum in Shang- hai, China, thought to have been taken in Hupeh Province, China (Allen, 1940, p. 665) ; owstoni, AMNH No. 27545, an adult female from Si-Tai-pa-shieng Mts., Shensi, China, taken in October 1905 by A. Owston; thayeri, MCZ No. 8008, an adult male from Washan, 6000 feet, Szechwan, China, taken May 17, 1908, by Walter R. Zappey. Material examined, from Shensi, China.^ — Tai-Pa-Shiang Mts., 6000 feet (AMNH), eight; Tai-pei-shan Dist., 3000 feet, 80 miles W.S.W. of Sian-fu [Siking] (USNM), three; base of Tai Pei Shan, Tsingling Mts. (CNHM), six; 15 miles east of Yen-an-fu Fushih (USNM), one; Tsingling Mts. (AMNH), three, (MCZ), one, (CNHM), six; 45 miles south of Fensiangfu, 3000 to 3600 feet (AMNH), two, (MCZ), one, (CNHM), two; nine and 15 miles south of Ching-chien-hsien, 2600 feet (USNM), two; Liu-tsuen, 1500 feet, 15 miles south of Sian-fu (USNM), four. Material examined, from Kansu, China. — Ha Tebbuland, 8500 feet, Wantsangku (MCZ), three. Material examined, from Hupeh, China. — Ma Fu Ling, 5000 feet (MCZ), two; Hsien Shan Hsien, 4000 feet (MCZ), two; "Tan Sweo Nah," 3000 feet (MCZ), one; Fongshan [Fanghsien] (MCZ), one; "Moo Swei Ping" (MCZ), one. Material examined, from Honan, China. — "Honan" (NR), three. Material examined, from Szechwan, China. — "Lu Din Chiao," 5000 feet (USNM), one; Longpan [Sungpan] (USNM), one; near Weichow [Weikiu], 9000 feet (USNM), one; "Tu-pa-keo," Moupine [Muping] (CNHM), one; Wenchuan [Wench wan] (USNM), 19, (AMNH), 24; Mt. Omei (USNM), two; E. of Romitchangu (MCZ), one; Ko-chia-ho-pa, 7500 feet (CNHM), one; Trashi-cho-ten, 8250 feet (CNHM), one; WaShan (MCZ), one; "Nai Su Chen" (CNHM), one; "Goan Shih Dwe" (CNHM), four; "Sungpan" (CNHM), two. Material examined, from Kweichow, China. — Whang Tsao Pa [Hwangtsaopa] (USNM), one. MOORE AND TATE: DIURNAL SQUIRRELS 309 Discussion. — We have been unable to distinguish any really good characters to separate consobrinus, owstoni and thayeri from one an- other. Taken together they are darker and more brownish, Cinna- mon Brown to Mummy Brown, than the northern race davidianus, and their under parts have a wash of Light Ochraceous Buff to Ochraceous Buff which is lacking in the gray-bellied davidianus. In the series of consobrinus from Wenchwan, the white postauricular patches are prominent. It is rather remarkable that a specimen should come from very southern Kweichow over 300 miles south of other Sciurotamias d. consobrinus locations found. In this hiatus, moreover, are a number of localities where Dremomys pernyi and D. pyrrhomerus were taken. Habits. — Fu (1936, p. 258) makes an interesting comment on the habits of this squirrel in western Honan where he collected it in the Sung-shan: "This animal nests in the ground, and is ascribed as a nuisance to peasants on account of its damage on corns. The den and its contents are often destroyed by ploughing." Subgenus RUPESTES Thomas Rupestes Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, p. 398. Type species. — Rupestes forresti Thomas. Definition.— The subgenus Rupestes includes only the one species forresti, which is a relict known only from the mountains of western China as shown in Figure 33. Diagnosis. — (This diagnosis is based upon one skull of a subadult with the occiput and bullae missing, USNM 255138, and four skins.) On the evidence examined Rupestes differs from subgenus Sciuro- tamias in several notable characters: (1) The squamosal reaches up the side of the cranium less than halfway from the base of the zygo- matic process to the base of the postorbital process. (2) The anterior edge of the zygoma ascends to a point only halfway in the height of the rostrum. (3) The superior process of the jugal is high and pos- terior to the midlength of the orbit. (4) There are pronounced tem- poral ridges. (5) The postorbital process of the frontal is very short (less than 1 mm. in this specimen). (6) There are no third upper premolars. (7) There is a rather faint light stripe along the flank between shoulder and hip. Sciurotamias (Rupestes) forresti (Thomas) Rupestes forresti Thomas, 1922, Ann. Mag. Nat. Hist. (ser. 9), 10, p. 399. 310 FIELDIANA: ZOOLOGY, VOLUME 48 Type. — BM No. 22.9.1.54, old female, from Mekong-Yangtze divide, latitude 27° 20' N., 7000 to 9000 feet, northern Yunnan, China, collected June 5, 1921 by George Forrest. Material examined. — E. of Yalung [River], 15,500 feet [4725 me- ters], Szechwan [Sikang] (USNM), one; Chienchwan [Tienchiian], Szechwan [Sikang] (MCZ), one; LiChiang [Likiang], Yunnan (AMNH), one, (BM), one; Mekong-Yangtze divide latitude 27° 20' N., 7000 to 9000 feet, Yunnan (BM), two. Definition. — This relict, rarely taken species is apparently mono- typic, and its range as known from four specimens examined is shown in Figure 33. Diagnosis. — Since the subgenus is monotypic, the diagnosis of the subgenus serves for the species. This species is readily told from davidianus by the presence of a pair of faint whitish lines high on the sides which extend from the shoulders to the hips. These lines are edged on each side by the dark grizzled dorsal coloring. Some 6 mm. beyond them the dark dorsal color changes to the light orange brown of the sides and that in turn to the buffy of the underparts. See Table 15 for some dimensions of the type. SUMMARY Twenty-six species of the seven genera of Sciurinae occurring in the Indian Subregion and Indochinese Subregion of the Oriental Re- gion are, for these geographic areas, here fully revised. Since four of the seven genera (Funambulus, Tamiops, Menetes, and Sciuro- tamias) are endemic, complete generic revisions of these are pre- sented. In order to offer a complete revision of the genus Dremomys, also, the one extraterritorial (monotypic, Malaysian) species, everetti, is included here. Eight species of Callosciurus are recognized in the Indochinese Subregion, and are shown to constitute two superspecies which are broadly sympatric throughout much of the subregion. These two superspecies are observed to form a phylogenetic unit separate from the Malaysian species of Callosciurus, and a complete revision is offered for this eight species unit. In the genus Ratufa complete revision is presented for the two species in the Indian Sub- region. The species Ratufa bicolor which occurs throughout much of the Indochinese Subregion is revised only for this area, although it also has an extensive distribution in the Malaysian Subregion. The principal changes from earlier existing concepts are within the Indochinese subregional unit of the genus Callosciurus, a unit previously unrevised. Considerable advancement in clarification and improvement of previously existing knowledge, however, is pro- vided for every species by mapping the distribution of collecting localities. Future supplementation and correction of the present knowledge is made easy by presenting latitudes and longitudes for the collecting localities presumed found. In those subspecies for which adequate material was examined, particularly in American museums, descriptions are provided from large samples in effort to make them applicable to the populations (not merely the type speci- mens) involved. In the genus Callosciurus in particular, perhaps only because more collecting sites and materials are known, striking evidence is present in the taxonomic account of inability to cross the main courses of the great rivers of the subregion. The evolutionary implications of this and other facts of geographic distribution of diurnal squirrels in this subregion are only sparingly discussed. 311 GAZETTEER It is intended in any discussions in the present paper to employ the geographic names current at the time of final preparation of the manuscript. In the lists citing localities where specimens were col- lected, however, the names and spellings that were recorded on the original specimen tag by the collector have ordinarily been retained. Should any question arise, this permits reinterpretation of the local- ity without resort to the original source. The currency of geographic names of the smaller principalities actually used here necessarily varies a great deal, depending upon the source. For some localities no knowledge is available to us except that they are villages near which specimens were taken, and they are mapped in an account of a collecting trip. Many, however, were taken by collectors whose routes and collecting localities were not published and whose local- ties to be plotted must usually be found on some map. Since villages in the jungle are often moved, localities of 50 years ago may be in many cases more easily found on maps made about that time. Many sources were utilized, but those most helpful and employed most were: Stanford (1917) "Complete Atlas of China" with maps at scale of 1:3 million; a set of maps of Indochina and most of Thai- land at 1:1/2 million published at Luang Prabang in 1931 by le Service Geographique de I'lndochine; "The Times Survey Atlas and Gazetteer of the World," J. G. Bartholomew (1922); its successor, "The Times Atlas of the World, Mid-Century Edition," John Bar- tholomew (1958) ; and maps published by the National Geographic Society, Washington, D. C. Because villages are moved and the names of towns, cities, and even natural geographic features, have been changed by new political regimes, latitude and longitude (from the Greenwich zero meridian, NOT the Paris one of the map of Indochina cited above) are em- ployed. Even this effort at accuracy embraces errors of five miles or so from differences of latitude or longitude between different maps. This may be a matter of some concern in the immediate vicinity of a large river which is potentially an important, long-time barrier to the species. 313 314 FIELDIANA: ZOOLOGY, VOLUME 48 In alphabetizing, place names of more than one word are treated as if but a single word (e.g., Ban Don, Bangnara, Ban Huai Som would come properly in this order) , Occasionally place names are used that may be found on a good map in several places (e.g., Khao Soi Dao is the name of at least three mountains in Thailand) . With the knowledge from the speci- men tag of who collected the specimen and when, and with available knowledge of the collector's itinerary, we have generally been able to decide which must be the correct place. In the gazetteer, only if specimens here studied were collected at two localities of the same name is an effort made to warn the user of the existence of other like place names. Adung Valley, Burma: 28° 10' N., 97° 40' E. Agia, Assam, India: 26° 06' N., 90° 34' E, AiNGGYl, Pakokku Chin Hills, Burma: 21° 05' N., 94° 22' E. AlRAWAN, Mt.: see Khao Erawan Akhe Triangle, Upper Burma: 26° 54' N., 98° 12' E. Akola, Berar, India: 20° 45' N., 77° 00' E. Amarakantak, India: 22° 40' N., 81° 45' E. Amherst, Tenasserim, Burma: 16° 00' N„ 97° 38' E. Amlekhganj, Nepal: 27° 18' N., 85° 00' E. Amraoti, Bastar [Berar], India: 20° 55' N., 77° 50' E. An Binh, Cochin China, Viet Nam: 10° 30' N., 106° 43' E. Angkor, Cambodia: 13° 25' N., 103° 50' E. Angtong [Ang Thong]: see Ban Bang Kaeo Aranya: see Ban Aranyaprathet Arbre Broye, Viet Nam: 11° 57' N., 108° 28' E. Avalanche, Nilgiri Hills, India: 11° 10' N., 76° 35' E. Ayuthia: see Phra Nakhon Si Ayutthaya Bag Kan, Tonkin, Viet Nam: 22° 10' N., 105° 45' E. Bag Tan Tray, Tonkin, Viet Nam: 22° 06' N., 103° 15' E. Baksa, Formosa, China: 23° 10' N., 120° 29' E. Balapalli, Madras, India: 13° 45' N., 79° 35' E. Bali: see Nhatrang Bamanigaon, Assam, India: 26° 10' N., 91° 25' E. Ba Nam Chi, Tonkin, Viet Nam: 22° 07' N., 102° 56' E. Ba Nam Nhung, Tonkin, Viet Nam: 22° 07' N., 103° 00' E. Ban Aranyaprathet, Prachin Buri, Thailand: 13° 40' N., 102° 30' E. Ban Bang Kaeo, Ang Thong, Thailand: 14°35'N.,100°25'E. Ban Chakkrarat, Nakhon Ratchasima, Thailand: 15° 00' N., 102° 25' E. Ban Chang Thuk, Thailand: 14° 45' N., 101° 30' E. Ban Chiang Dao, Thailand: 19° 20' N., 99° 00' E. Ban Choeng Doi, Chiang Mai, Thailand: 18° 50' N., 99° 10' E. Ban Den, Nong Khai, Thailand: circa 18° 00' N., 102° 55' E. Ban Doi Saket : see Ban Choeng Doi Bandon: see Ban Makham Tia MOORE AND TATE: DIURNAL SQUIRRELS 315 Ban Dong: see Ban Pong Bang Hue Hom, Thailand: 17° 53' N., 100° 06' E. Bang Hue Pong: see Sathani Pang Hua Phong Bangkok (160 miles north of), Thailand: 16° 00' N., 100° 45' E. Bangkok: see Krung Thep Bangnara: see Narathiwat Ban Hai Huai Som: see Ban Huai Som Ban Hin Laem, Kanchanaburi, Thailand: 14° 40' N., 98° 40' E. Ban Hin Ngom, Nong Khai, Thailand: 17° 55' N., 102° 50' E. Ban Hoi Mak: see Pak Hin Bun Ban Houei Sai, Laos: 20° 18' N., 100° 25' E. Ban Huai Som, Nan, Thailand: 18° 20' N., 100° 30' E. Ban Huang Som, Trat, Thailand: 11° 50' N., 102° 50' E. Banhuathanon: see Ban Hua Thanon Ban Hua Thanon, Kamphaeng Phet, Thailand: circa 16° 14' N., 99° 35' E. Ban Hu Kwang, Nakon Sawan, Thailand: 15° 55' N., 100° 00' E. Ban Hup Bon, Thailand: 13° 05' N., 101° 05' E. Ban Kabin Buri, Prachin Buri, Thailand: 14° 00' N., 101° 45' E. Bankachon, Tenasserim, Burma: 10° 9' N., 98° 36' E. Ban Kang: see Ban Mae Klang Ban Khana, Nan Province, Thailand: 19° 19' N., 100° 56' E. Ban Khlong Bang Lai, Patiyu, Thailand: 10° 45' N., 99° 10' E. Ban Khlong Khlung, Kamphaeng Phet, Thailand: 16° 10' N., 99° 45' E. Ban Khlong Mon, Samut Sakhon, Thailand: 13° 35' N., 100° 35' E. Ban Khlua Klang, Prachuap Khiri Khan, Thailand: circa 11° 35' N., 99° 39' E. Ban Khok, Naphung, Dansai, Loei, Thailand: 16° 25' N., 101° 25' E. Ban Khok Klap: see Ban Na Ban Khung Samphao, Thailand: 15° 20' N., 100° 00' E. Ban Kiriwong: see Ban Wat Khiriwong Ban Klua Klang: see Ban Khlua Klang Ban Krat, Thailand: 14° 26' N., 103° 04' E. Banlad: see Ban Lat Ban Laem Ngop, Trat, Thailand: 12° 08' N., 102° 35' E. Ban Lat, Phu Kheio, Chaiyaphum, Thailand: 16° 30' N., 101° 55' E. Ban Lat Bua Khao, Nakhon Ratchasima, Thailand: 14° 50' N., 101° 35' E. Ban Le Kathe, Tak, Thailand: 15° 50' N., 98° 50' E. Ban Luang, Chiang Mai, Thailand: 18° 25' N., 98° 40' E. Ban Mae Klang, Thailand: 18° 30' N., 98° 40' E. Ban Mae Mo, Thailand: 18° 15' N., 99° 45' E. Ban Mae Sariang, Mae Hong Son, Thailand: 18° 10' N., 97° 55' E. Ban Mae Tan Nua, Lampeng, Thailand: 18° 20' N., 99° 25' E. Ban Mae Ton: see Pang Mae Ton Ban Maha Chai, Samut Sakhon, Thailand: 13° 30' N., 100° 14' E. Ban Makham Tia, Thailand: 9° 10' N., 99° 20' E. Ban Manao Wan, Thailand: 14° 55' N., 101° 05' E. Ban Manaswan: see Ban Manao Wan Ban Meh Na, Thailand: 19° 12' N., 98° 52' E. Ban Mb Thuot: see Buon Ma Thuot Ban Mi Chai, Nongkhai, Thailand: 17° 50' N., 102° 50' E. Ban Muak Lek, Sara Buri, Thailand: 14° 40' N., 101° 10' E. 316 FIELDIANA: ZOOLOGY, VOLUME 48 Ban Muang Khai, Tha Li, Muang Loei, Thailand: 17° 35' N., 101° 20' E. Ban Mung Ky: see Ban Muang Khai Ban Nai Muang, Phichit, Thailand: 16° 25' N., 100° 20' E. Ban Na, Thailand: 8° 53' N., 99° 17' E. Ban Na Ko, Nan, Thailand: circa 19° 10' N., 100° 54' E. Ban Nam Chut Yai, Ranong, Thailand: 10° 25' N., 98° 45' E. "Ban Na Muang, Na Haeo," Dan Sai, Loei, Thailand: 17° 15' N., 101° 02' E. Ban Nam Yen, Koksathon, Thailand: 17° 10' N., 101° 05' E. "Ban Na Nong," Chuempae [Ban Chum Phae], Khon Kaen, Thailand: 16° 34' N., 102° 03' E. Ban Neng Kho: see Ban Nong Kho Ban Nong Bua, Pasak River, Thailand: 14° 50' N., 101° 05' E. Ban Nong Chik: see Ban Tuyong Ban Nong Don Ta, Thailand: circa 18° 12' N., 104° 05' E. Ban Nong Kho, Chon Buri, Thailand: 13° 10' N., 101° 05' E. Ban Nong Pla Lai, Thailand: 16° 05' N., 98° 45' E. Ban Non Tao Lek, near Chaiyaphum, Thailand: 16° 27' N., 101° 50' E. Ban Non Toulek: see Ban Non Tao Lek Ban Pak Chan, Thailand: 10° 32' N., 98° 51' E. Ban Pak Chong, Nakhon Ratchasima, Thailand: 14° 40' N., 101° 25' E. Ban Pakli: see Ban Sopli Ban Pak Nam Pho, Nakhon Sawan, Thailand: 15°'40' N., 100° 10' E. Ban Pak Tho, Rajburi, Thailand: 13° 21' N., 99° 50' E. Ban Phan Sadet, Thailand: 13° 05' N., 101° 05' E. Ban Phone, Laos: 15° 25' N., 106° 35' E. Banphotphisai: see Ban Hu Kwang Ban Phu Khieo, Chaiyaphum, Thailand: 16° 34' N., 101° 52' E. Ban Pong, Rat Buri, Thailand: 13° 50' N., 99° 55' E. Ban Pua, Nan, Thailand: 19° 11' N., 100° 53' E. Ban Rahaeng, Tak, Thailand: 16° 50' N., 99° 05' E. Ban Sadet: see Ban Phan Sadet Ban Sahng Kaw: see Ban Sawan Ko Ban Sai Kan: see Nawngchik Ban Sai Yok, Kanchanaburi, Thailand: 14° 29' N., 98° 50' E. Ban Sa Kaeo, Prachin Buri, Thailand: 13° 50' N., 102° 05' E. Ban Salok Bat, Kamphaeng Phet, Thailand: 16° 00' N., 99° 45' E. Ban Salui: see Ban Khlong Bang Lai Ban San Tha, Nan, Thailand: circa 18° 21' N., 100° 40' E. Ban Sattahip, Chon Buri, Thailand: 12° 40' N., 100° 55' E. Ban Sawan Ko, Sakon Nakhon, Thailand: 17° 27' N., 104° 05' E. Ban Sichon, Thailand: 9° 00' N., 99° 55' E. Ban Si Khiu, Nakhon Ratchasima, Thailand: 14° 55' N., 101° 45' E. Ban Si Racha, Chon Buri, Thailand: 13° 10' N., 101° 05' E. Ban Sob Pa [Soppa], Burma: 18° 48' N., 97° 27' E. Ban Sompan, Lower Burma: circa 10° 30' N., 98° 30' E. Ban Sopli, Thailand: 18° 00' N., 100° 30' E. Ban Talok Bat: see Ban Salok Bat Ban Tarn Dam: see Ban Than Dam Ban Ta Yai: see Ban Tha Yai Ban Tayun, Plateau Bolovens, Laos: 15° 23' N., 106° 23' E. MOORE AND TATE: DIURNAL SQUIRRELS 317 Ban Tha Chang, Thailand: 14° 35' N., 101° 25' E. Ban Tha Khanum, Kanchanaburi, Thailand: 14° 45' N., 98° 40' E. Ban Than Dam: Near Ban Si Racha Ban Thani, Sukhothai, Thailand: 17° 00' N., 99° 51' E. Ban Tha San, Ranong, Thailand: 10° 30' N., 98° 55' E. Ban Thateng, Laos: 15° 25' N., 106° 23' E. Ban Tha Yai, Thailand: 8° 23' N., 99° 52' E. Bantion: see Ban Tayun Ban Ton Phung, Laos: 20° 18' N., 100° 05' E. Ban Tuyong, Thailand: 06° 50' N., 100° 00' E. Ban Um Phang: see Ban Le Kathe Ban Wang Mai Khon, Sukhothai, Thailand: 17° 19' N., 99° 50' E. Ban Wang Takhian: see Kanchanaburi Ban Wat Khiriwong, Thailand: circa 8° 23' N., 99° 52' E. Ban Wiang Tai, Mae Hong Son, Thailand: 19° 20' N., 98° 25' E. Ban Wichian, Thailand: 15° 40' N„ 101° 05' E. Bag Ha, Tonkin, Viet Nam: 22° 10' N., 104° 23' E. Barapani: see Borpani Baria, Cochin China, Viet Nam: 10° 30' N., 107° 10' E. Baurang, Yunnan [Sikang], China: 28° 52' N., 101° 23' E. "Bhadwar," Kangra, Punjab: 32° 05' N., 76° 18' E. Bhor Phloy Nong Yungchang: see King Bo Phloi Bhuket: see Koh Phuket BiEN HOA, Cochin China: 10° 52' N., 106° 50' E. Bintenna: see Pintenne BOK Pyin, Tenasserim, Burma: 11° 15' N., 98° 46' E. Boraphet, Thailand: 15° 43' N., 100° 14' E. BORI, Hoshangabad, India: 20° 55' N., 79° 05' E. Borpani, Khasi Hills, Assam, India: 25° 39' N., 91° 53' E. Borploy: see King Bo Phloi BOUN Tai: see Bun Tai Boyce's Point, Tenasserim, Burma: 10° 45' N., 98° 29' E. BUKET: see Koh Phuket Bung Borapet: see Boraphet Bun Tai, Laos: 21° 18' N., 102° 00' E. BUON Ma Thuot, Viet Nam: 12° 40' N., 108° 05' E. Burnihat, Assam, India: 25° 37' N., 91° 50' E. Cachar [Silchar], Assam, India: 24° 42' N., 92° 50' E. Cachar Hills, Assam, India: 25° 00' N., 93° 00' E. Chaduquet Point: see Laem Set Kuat Chaiyaphum, Thailand: 16° 27' N., 101° 50' E. Chamaphum: see Ban Phu Kheio Champang, Tenasserim, Burma: 10° 12' N., 98° 27' E. Chamrajnagar, Mysore, India: 11° 55' N., 76° 55' E. Chance Island: see Ko Chan Chandragiri Pass, Nepal: 27° 39' N., 85° 08' E. Changwat Kamphaeng Phet, Kamphaeng Phet, Thailand: 16° 30' N., 99" 30' E. Changwat Mae Hong Son, Thailand: 19° 15' N., 97° 55' E. Chang Yang Hsien, Hupeh, China: 30° 25' N., 111° 13' E. Chantabun: see Chanthaburi 318 FIELDIANA: ZOOLOGY, VOLUME 48 Chanthaburi, Thailand: 12° 35' N., 102° 05' E. Chan Tuk: see Ban Chan Thuk Chao Chu, Yunnan, China: 25° 35' N., 100° 18' E. Chapa: see Cha Pa Cha Pa, Tonkin, Viet Nam: 22° 20' N., 103° 50' E. Chaung, 20 miles n.n.w. of Kindat, Burma: 23° 45' N., 94° 17' E. Chaung-lung, Salween ferry, Yunnan, China: 24° 12' N., 98° 58' E. Chenga Hka, Burma: 26° 07' N., 95° 52' E. Chengou Creek, 25 mi. W. of Wenchwan, Szechwan, China: 31° 20' N., 103° 00' E. Chenkang, Yunnan, China: 24° 06' N., 99° 22' E. Che Pei, Fu Min District: see Fuminhsien Cherrapunji, Assam, India: 25° 13' N., 91° 47' E. Chettiri Range, Salem Dist., Madras, India: 11° 50' N., 78° 25' E. Chiang, Szechwan, China: 28° 47' N., 102° 53' E. Chiang Rai: see Muang Chiang Rai Chiang Saen Kao, Chiang Rai, Thailand: 20° 15' N., 100° 05' E. Chiang Saen Luang: see Chiang Saen Kao Cheinat: see Muang Chainat Chienchwan: see Tienchiian Chieng Dao: see Ban Chiang Dao Chiengmai: see Muang Chiang Mai Chih ping: see Shihpingchow "Chi li ping," Fanghsien, Hupeh, China: 32° 03' N., 110° 54' E. Chimeli Pass, Upper Burma: 26° 08' N., 98° 40' E. Chinbyit, Burma: 22° 03' N., 94° 40' E. Ching-chien-shien: see Tsingkien Ching Feng Ling, Fukien, China: 27° 11' N., 118° 13' E. Chinteh: see Tsingteh Chinwangtao, Hopeh, China: 39° 56' N., 119° 37' E. Chin Wang Tas: see Chinwangtao Chip-Chip, Formosa: circa 23° 55' N., 120° 45' E. Chipwi, northeastern Burma: 25° 52' N., 98° 07' E. Chomtong: see Ban Luang Chora, Tonkin, Viet Nam: 22° 18' N., 105° 52' E. Chulungapu, Upper Tebbuland, Kansu, China: circa 34° N., 101° 30' E. Chumphon, Thailand: 10° 30' N., 99° 10' E. Chum Poo: see Sathani Tha Chomphu Chunganhsien, Fukien, China: 28° 02' N., 117° 53' E. Chungan Hsien: see Chunganhsien Chung Chiao Miao: see Chiang Chung King, Szechwan, China: 29° 34' N., 106° 35' E. Chungtang, Sikkim, India: 27° 37' N., 88° 38' E. Chuntown: see Chumphon Clara Island, Mergui Archipelago, Burma: 10° 50' N., 97° 55' E. CoiMBATORE, Madras, India: 10° 55' N., 76° 58' E. Col des Nuages, Annam, Viet Nam: 16° 11' N., 108° 10' E. Col de Taloun, Laos: 19° 50' N., 102° 15' E. COORG, India: 12° 00' N., 76° 00' E. "Cotengady Estate," [Nelliampathi Hills, Palghat District], Cochin, India: 10° 40' N., 76° 35' E. MOORE AND TATE: DIURNAL SQUIRRELS 319 CuCHAi: see Kuchai Da Ban, Phanrang Province, Annam: 12° 38' N., 109° 06' E, Dacca, East Pakistan: 23° 43' N., 90° 26' E. Dagung Hka, Burma: 26° 05' N., 95° 55' E. Dak-to, Annam, Viet Nam: 14° 40' N., 107° 45' E. Daingmhu, 40 mi. N. of Pegu, Burma: 17° 42' N., 96° 14' E. Dalat, Annam, Viet Nam: 11° 58' N., 108° 25' E. Dalu, N. Burma: 26° 20' N., 96° 12' E. Dao Soi Dao: see Khao Soi Dao Darjeeling, Bengal, India: 27° 00' N., 88° 18' E. Darugiri, Garo Hills, Assam, India: 25° 37' N., 90° 46' E. DeLisle, Thailand: 09° 40' N., 98° 20' E. Den Chai, near Bang Hue Hom, Thailand: circa 17° 53' N., 100° 06' E. Dening, Mishmi Hills, Assam, India: 28° 01' N., 96° 10' E. Devikop, S. Mahratta, India: 15° 08' N., 75° 00' E. DiKCHU, Sikkim, India: 27° 25' N., 88° 35' E. Djiring, Annam, Viet Nam: 11° 35' N., 108° 06' E. Doi Angka: see Doi Inthanon Doi Chiang Dao: see Doi Luang Chiang Dao Doi Chiengdao: see Doi Luang Chiang Dao Doi Hua Mot: see Chiengmai Doi Inthanon, Chiang Mai, Thailand: 18° 32' N., 98° 32' E. Doi Kang Ma: see Doi Khang Ma Doi Khang Ma, Chiang Mai, Thailand: circa 18° 08' N., 98° 27' E. Doi Khun Tan, Lamphun, Thailand: 18° 30' N., 99° 20' E. Doi Lak Saen, Thailand: circa 18° 08' N., 98° 07' E. Doi Lak Sen: see Doi Lak Saen Doi Langka: see Khao Pha Cho Doi Luang Chiang Dao, Chiang Mai, Thailand: 19° 25' N., 98° 55' E. Doi Mae Kong Ka, Mae Hong Son, Thailand: circa 18° 06' N., 97° 49' E. Doi Me Kong Kha: see Doi Mae Kong Ka Doi Mae Lai, Mae Hong Son, Thailand: circa 18° 09' N., 98° 00' E. Doi Mei Lai: see Doi Mae Lai Doi Nangka: see Khao Pha Cho Doi Nang Keo: see Doi Phi Phan Nam Doi Par Sakeng, northwestern Thailand: 19° 35' N., 99° 03' E. Doi Phi Phan Nam, Chiang Mai, Thailand: 19° 05' N., 99° 25' E. Doi Phra Chao: see Khao Pha Cho Doi Phu Het, Nan, Thailand: circa 18° 20' N., 100° 35' E. Doi Phu Ka, Nan, Thailand: 19° 05' N., 101° 05' E. Doi Phu Kha, Thailand: 19° 05' N., 101° 05' E. Doi Pu Het: see Doi Phu Het Doi Pu Kha: see Doi Phu Kha Doi San Huai Wai, Nan, Thailand: circa 18° 20' N., 100° 35' E. Doi Sritepe: see Doi Suthep Doi Sutep: see Doi Suthep Doi Suthep, Chiang Mai, Thailand: 18° 50' N., 98° 55' E. DoMEL Island, Mergui, Burma: 11° 37' N„ 98° 15' E. Donghai: see Dong Hoi Dong Hoi, Annam, Viet Nam: 17° 32' N., 106° 35' E. 320 FIELDIANA: ZOOLOGY, VOLUME 48 Don Qua, Laos: 19° 45' N., 102° 35' E. DOONSA, near junction Gyn [Gyaing] and Haung Thrau [Haung Tharaw] Rivers, near Moulmein, Burma; 16° 37' N., 98° 00' E. Drachuapchiri Khan Bankhluaklang: see Ban Khlua Klang Dreyi, Mishmi Hills, Tibet: 28° 03' N., 96° 15' E. DuRAGlRl: see Darugiri "Eaktur," E. of Ban Me Thuot: see Buon Ma Thuot Eastern Tombs, Hopeh, China: 40° 00' N., 117° 00' E. Etawah United Provinces, India: 26° 50' N., 79° 00' E. Fanghsien, Hupeh, China: 32° 03' N., 110° 54' E. Fansipan: see Mt. Fan-si-pan Fengsiangfu, Shensi, China: 34° 24' N., 107° 29' E. Feng Yang, Yunnan, China: 24° 45' N., 102° 40' E. FiMNON, S. Annam, Viet Nam: 11° 40' N., 108° 22' E. Fi Shan Kwan, Szechwan, China: 30° 03' N., 103° 05' E. FOOCHOW, Fukien, China: 26° 05' N., 119° 18' E. Fuching: see Futsing FUCHOW, Fukien, China: 26° 05' N., 119° 18' E. Fu Fu Su, Szechwan, China: 29° 35' N., 103° 35' E. Fu Lin, Szechwan, China: 29° 20' N., 102° 45' E. FUMINHSIEN, Yunnan, China: 25° 12' N., 102° 31' E. Futsing, Fukien, China: 25° 40' N., 119° 25' E. Gammaduwa, Ceylon: 07° 35' N., 80° 41' E. Gangfang, Burma: 26° 06' N., 98° 38' E. Gangtok, Sikkim, India: 27° 20' N., 88° 30' E. Ghatmatha, Satara District, Bombay, India: 17° 25' N., 73° 40' E. GOALPARA, Assam, India: 26° 10' N., 90° 37' E. Godavari, Nepal: 27° 35' N., 85° 20' E. GoDAVERi: see Godavari GoKTEiK, Northern Shan States, Burma: 22° 20' N., 96° 52' E. GOLAGHAT, Assam, India: 26° 30' N., 94° 01' E. Gopaldhara, Rungbong Valley, India: 26° 55' N., 88° 12' E. Gorkha, Nepal: see Satthar Gougah: see Lien Gongah GuNGWADORiA, north slope of Palni Hills, India: circa 10° 20' N., 77° 30' E. Gyaing, and Haung Thrau [Haung Tharaw] Rivers, near Moulmein, Burma: 16° 37' N., 98° 00' E. Ha-Gang: see Ha Giang Ha Giang, Tonkin, Viet Nam: 22° 50' N., 104° 58' E. Hahti, Burma: 26° 04' N., 95° 32' E. Hai Bum, Burma: 26° 02' N., 95° 47' E. Hai Yen Hsien, Hangchow Bay, Chekiang: 30° 17' N., 120° 10' E. Hambantota, Ceylon: 06° 07' N., 81° 07' E. Hastings Island, Mergui Archipelago, Burma: 10° 05' N., 98° 20' E. Hathiban, Nepal: 27° 37' N., 85° 15' E. Hathiben: see Hathiban Hataura, Nepal: 27° 25' N., 85° 05' E. Hat Sanuk, Rajburi, Thailand: 11° 52' N., 99° 42' E. Haungtharaw River (where crossed by Moulmein-Kawkareik road), Burma: 16° 30' N., 98° 05' E. MOORE AND TATE: DIURNAL SQUIRRELS 321 Heinsum, Burma: 25° 52' N., 95° 32' E. Helwak, Satara Dist., Bombay, India: 17° 20' N., 73° 40' E. Hetora: see Hataura HiNLAEM: see Ban H in Laem HiN Lap: see Sathani Hin Lap Hkamkawn, Burma: 26° 02' N., 98° 23' E. Hkamti, Burma: 26° 00' N., 95° 42' E. Hkamti Plain, Burma (taken to be the same locality as Putao and Ft. Hertz): 27° 22' N., 97° 27' E. Hkawni, Gaw, Upper Burma: 25° 58' N., 98° 00' E. Hmawbi, 40 km. north of Rangoon, Burma: 17° 05' N., 96° 05' E. Hnathial, Lushai Hills, Assam, India: circa 23° 45' N., 92° 55' E. Hoi Xuan, Annam, Viet Nam: 20° 20' N., 105° 05' E. HOKOW, Szechwan, China: 30° 01' N., 101° 08' E. HOMALIN, Burma: 24° 53' N., 94° 55' E. HOMU-SHU Pass, Yunnan, China: 24° 55' N., 98° 45' E. HOOPBON: see Ban Hup Bon HOSHANGABAD, India: 22° 45' N., 77° 45' E. HouE SAi: see Ban Houei Sai Hpimaw, Salween-Irrawaddy Divide, Burma: 25° 58' N., 98° 38' E. HsiAO-MENG-TUNG: see Siaomengtung Hsiao Yang Chi, Szechwan, China: 28° 47' N., 102° 44' E. Hsien Shan Hsien, Hupeh, China: 31° 30' N., 110° 55' E. Hsing Liao Pa, Kweichow, China: 28° 14' N., 106° 33' E. HsiN Kai: see Sinkai HsiN LUNG SHAN, Jehol, Manchuria: 40° 38' N., 117° 17' E. HsiPAW, Shan States, Burma: 22° 38' N., 97° 20' E. Htawgaw, Burma: 25° 55' N., 98° 25' E. Htingwan, Burma: 26° 31' N., 97° 52' E. Huai Nua Pla, Tak, Thailand: 16° 54' N., 98° 48' E. Hue Sai, near Koh Lak, Thailand: circa 11° 48' N., 99° 47' E. Hue Yah Pla: see Huai Nua Pla HuEY Yang: see Ban Si Racha HULAUNG, Burma: 25° 10' N., 95° 05' E. HUNGAI, 40 miles E. of Talifu, Yunnan, China: circa 25° 30' N., 100° 20' E. Hung Fa Chao: see Shanghai Hup Bon: see Ban Hup Bon Hup Bum: see Ban Hup Bon Hwang Tsao Pa: see Hwangtsaopa HWANGTSAOPA, Kweichow, China: 24° 57' N., 104° 36' E. Ichang, Hupeh: 30° 45' N., 111° 22' E. ICHANG Hsien, Hupeh, China: 30° 42' N., 111° 17' E. I-Lan Hsien, I-Lan, Taipei, Taiwan, China: 24° 25' N., 121° 42' E. Imaw Bum: see Mt. Imaw Bum INTHA, Burma: 24° 30' N., 94° 42' E. IPIN, Szechwan- Yunnan border, China: 28° 43' N., 104° 32' E. Isle [He] de la Table, Tonkin, Viet Nam: 20° 57' N., 107° 30' E. James Island, Mergui Archipelago, Burma: 10° 25' N., 98° 17' E. Jhimpir Lake, Sind, India: 25° 00' N., 68° 00' E. Kabaw Valley, 20 mi. W. of Kindat, Burma: 23° 40' N., 94° 12' E. 322 FIELDIANA: ZOOLOGY, VOLUME 48 Kachek, Hainan, China: 19° 14' N., 110° 27' E. Kadrangyang, Fort Hertz Rd., 75 mi. N. of Myitkyina, Burma: circa 26° 07' N., 97°30'E. Kagi Dist., Formosa: 23° 29' N., 120° 17' E. Kaing R., S. Pyinwana, S. Burma: 19° 44' N., 96° 09' E. Kalewa, Burma: 23° 12' N., 94° 13' E. Kalhatti [Falls], Nilgiri Hills, India: 11° 27' N., 76° 43' E. Kalutara, Matugama, Ceylon: 06° 34' N., 79° 58' E. Kampeng Pet: see Kamphaeng Phet Kamphaeng Phet, Thailand: 16° 30' N., 99° 30' E. Kampot, Cambodia: 10° 40' N., 104° 18' E. Kan Bouri : see Kanchanaburi Kanchanaburi, Thailand: 14° 00' N., 99° 30' E. Kangpu: see Kanpu Kangra, Punjab: 32° 05' N., 76° 18' E. Kanjanaburi: see Kanchanaburi Kanpu, near Hangchow, Chekiang, China: 30° 20' N., 120° 50' E. Kao Chiao, Yunnan, China: 25° 01' N., 102° 20' E. Kao Hao, Quangtri Province, Annam, Viet Nam: 16° 45' N., 107° 11' E. Kao Lem: see Khao Laem Kao Luang, Huey Yang: see Khao Luang, Sathani Huai Yang (not the same locality as Khao Luang, Nakhon Si Thammarat) Kao Sabab: see Khao Sa Bap Kao Seming: see Khao Saming Kaoshanshih, near Foochow, Fukien, China: 25° 25' N., 119° 28' E. Kao Soi Dao: see Khao Soi Dao Kaotien, Fukien, China: 27° 18' N., 117° 25' E. Kao Tung Sawng: see Khao Thung Song Karjine: see Ban Pak Tho Karong, Manipur, Assam, India: 25° 15' N., 94° 05' E. Katha, Burma: 24° 10' N., 96° 20' E. Kauktaung, Burma: 25° 43' N., 95° 25' E. Kaunghein, Burma: 25° 42' N., 95° 24' E. Kawai, Burma: 26° 02' N., 95° 34' E. Kawkareik, Burma: 16° 28' N., 98° 16' E. Kawlaw, Southern Shan States, Burma: 20° 38' N., 96° 34' E. Kawlichaung, Burma: circa 16° 15' N., 98° 25' E. Kaw Nom Plu: see Khao Nam Pliu Kellengode: see Kollangod Kempanpet: see Changwat Kamphaeng Phet Khad Tham Phra, Laos: 20° 15' N., 100° 05' E. Khajjean: see Kangra Kha Nu, Kampeng Phet, Thailand: 16° 07' N., 99° 46' E. Khao Erawan, Lop Buri, Thailand: circa 14° 50' N., 100° 35' E. Khao Khat, Nakhon Sawan, Thailand: 15° 43' N., 100° 07' E. Khao Kop, Nakhon Sawan, Thailand: 15° 43' N., 100° 08' E. Khao Laem, Thailand: 14° 25' N., 101° 30' E. Khao Luang, Sathani Huai Yang, Thailand: 11° 36' N., 99° 41' E. (not the same locality as Khao Luang, Nakhon Si Thammarat) Khao Na Khae: see Khao Thung Song MOORE AND TATE: DIURNAL SQUIRRELS 323 Khao Nam Pliu, Thailand: 7° 35' N., 99° 50' E. Khao Pha Cho, Chiang Mai, Thailand: 19° 00' N., 99° 25' E. Khao Phra Phut, Lop Buri, Thailand: 14° 50' N., 100° 40' E. Khao Sa Bap, Chanthaburi, Thailand: 12° 35' N., 102° 15' E. Khao Saming, Trat, Thailand: 12° 21' N., 102° 27' E. Khao Soi Dao, S.E. Thailand: 15° 30' N., 100° 45' E. (not the same locality as Khao Soi Dao, Trong, Thailand) Khao Thung Song, Peninsular Thailand: 8° 20' N., 99° 48' E. Kharagpur, Bengal, India: 22° 20' N., 87° 20' E. Khet Dong Heing, Laos: 17° 53' N., 101° 34' E. Khlong Wang Hip, Thailand: circa 8° 12' N., 99° 43' E. Khon Kaen, Thailand: 16° 25' N., 102° 40' E. Khun Tan Mts. : see Doi Khun Tan KiATiNG: see Loshan Kienyang, Fukien, China: 27° 21' N., 118° 05' E. KiL: see Kotajiri Kin, Burma: 22° 41' N., 94° 41' E. KiNDAT, Burma: 23° 40' N., 94° 22' E. King Bo Phloi, Kanchanaburi, Thailand: 14° 48' N., 99° 10' E. King Chiang Saen: see Chiang Saen Kao King Island, Mergui Archipelago, Burma: 12° 30' N., 98° 22' E. KissERiANG Island, Burma: 11° 41' N., 98° 28' E. Klong Ban Lai: see Ban Khlong Bang Lai Klong Klung: see Ban Khlong Khlung Klong Menao: see Ban Huang Som Klong Morn: see Ban Khlong Mon Klong Wang Hip: see Khlong Wang Hip Klong Yai, Thailand: 11° 44' N., 102° 52' E. Ko Chan, Mergui Archipelago, Thailand: 9° 25' N., 97° 50' E. Ko Chang, Trat, Thailand: 12° 00' N., 102° 30' E. KODAIKANAL, India: 10° 12' N., 77° 30' E. KODURA, Balapalli Range, Madras, India: 13° 57' N., 79° 32' E. KoH Chang Island: see Ko Chang Ko-chia-ho-pa, Szechwan, China: circa 30° 20' N., 102° 20' E. KOHIMA, Assam, India: 25° 40' N., 94° 12' E. KoH KuT Island: see Ko Kut KOH Lak: see Prachuap Khiri Khan KoH Lan: see Ko Lan KoH Pangan: see Ko Phangan KoH Phai: see Ko Phai KOH Phuket, Southwestern Thailand: 7° 53' N., 98° 24' E. KoH Si Chang: see Ko Si Chang KoH Tao: see Ko Tao KOKKOAING, Burma: 22° 28' N., 96° 18' E. Kokkogon: see Kokkogwa KOKKOGWA, Burma: 19° 59' N., 95° 30' E. Ko Kut, Trat, Thailand 11° 40' N., 102° 35' E. Ko Lak: see Prachuap Khiri Khan Ko LAN, Chon Buri, Thailand: 12° 55' N., 100° 45' E. KOLLANGOD, India: 10° 38' N., 76° 37' E. 324 FIELDIANA: ZOOLOGY, VOLUME 48 KOLLEGAL, Coimbatore, India: 12° 10' N., 77° 10' E. Ko Maphrao, Thailand: 7° 55' N., 98° 25' E. Kontoum: see Kontum KONTUM, Annam, Viet Nam: 14° 25' N., 107° 55' E. KOON Tan: see Doi Kuhn Tan Ko Phai, Chonburi, Thailand: 12° 55' N., 100° 40' E. Ko Phangan, Thailand: 09° 45' N., 100° 00' E. Ko Si Chang, Chon Buri, Thailand: 13° 10' N., 100° 50' E. KOTAJIRI, Nilgiri Hills, India: 11° 25' N., 76° 55' E. Ko Tao, Surat Thani, Thailand: 10° 05' N., 99° 50' E. KOTTAWA, Ceylon: 06° 03' N., 80° 20' E. Kowjeen: see Ban Pak Tho KOWKAT: see Khao Khat Kowkob: see Khao Kop Krabin: see Ban Kabin Buri Krat: see Ban Krat Kratt: see Ban Krat Krong Ba, S. Annam, Viet Nam: 14° 00' N., 108° 18' E. Krung Thep, Phra Nakhon, Thailand: 13° 45' N., 100° 30' E. KUCHAI, Yunnan, China: 27° 20' N., 103° 14' E. Kucheng, Fukien, China: 26° 40' N., 118° 43' E. Ku-LU, Szechwan, China: 28° 05' N., 101° 12' E. KUMBUKKAN, Uva Prov., Ceylon: 06° 47' N., 81° 17' E. Kunming : see Yunnanfu KURUMBAPATTI, India: 11° 45' N., 78° 15' E. KusHAN: see Kaoshanshih Kutung, Yunnan, China: 25° 27' N., 99° 30' E. Kwatum: see Kaotien KwE Koi: see Mae Nam Khwae Noi Kyaukmyaung, Burma: 22° 30' N., 95° 57' E. Kyundaw, Burma: 21° 10' N., 94° 40' E. Lachen, Sikkim, India: 27° 45' N., 88° 32' E. Lachung, Sikkim, India: 27° 43' N., 88° 44' E. Lacon Sritamarat: see Nakhon Si Thammarat Laem Set Kuat, Thailand: 10° 05' N., 98° 40' E. Laem Sing, Chantaburi, Thailand: 12° 30' N., 102° 05' E. Lahkaw Hka, Burma: 26° 25' N., 96° 12' E. Lahore, Punjab: 31° 35' N., 74° 15' E. Lai Chau, Tonkin, Viet Nam: 21° 52' N., 103° 10' E. Laitlynkot, Khasi Hills, Assam, India: 25° 27' N., 91° 51' E. Lai Yoke: see Ban Sai Yok Lakchang Ga, Burma: 26° 16' N., 96° 07' E. Lakhon: see Muang Lampang Lakya, Tenasserim, Burma: circa 16° 15' N., 98° 25' E. Lamaya, Szechwan, China: 29° 52' N., 99° 54' E. Lampha, Burma: 16° 18' N., 98° 19' E. Lam-ra: see Sathani Lamphura Landa [Londa], India: 14° 30' N., 74° 25' E. Langbian Peaks, Annam, Viet Nam: 12° to 12° 15' N., 108° 20' E. Lang Son, Tonkin, Viet Nam: 21° 50' N., 106° 45' E. MOORE AND TATE: DIURNAL SQUIRRELS 325 Langyang, Burma: 25° 54' N., 98° 18' E. Lao Bao, Laos: 16° 37' N., 106° 34' E. Lao Fou Tchay, Laos: 21° 43' N., 102° 25' E. Lao Kay, Tonkin, Viet Nam: 22° 30' N., 104° 00' E. Lat Bua Kao: see Ban Lat Bua Khao Laukhaung, Burma: 25° 53' N., 98° 13' E. Lawksawk, Burma: 21° 13' N., 96° 50' E. Lay Song Hong: see Thale Song Hong LE BOKOR, Cambodia: 10° 40' N., 104° 00' E. Lei-Mui-Mon, Hainan, China: 19° 00' N., 109° 30' E. Lem Ngop: see Ban Laem Ngop Lem Sing Mountain: see Laem Sing Leng Sang, Tonkin, Viet Nam: 22° 25' N., 103° 17' E. Lian-feng-Kiang, Omei Shan, Szechwan, China: 29° 25' N., 103° 25' E. LiCHlANG : see Likiang Lien Gongah, Annam, Viet Nam: 11° 30' N., 108° 20' E. Lieng San: see Leng Sang Likiang, Yunnan, China: 27° 04' N., 100° 12' E. LiMPA, Burma: 25° 45' N., 95° 30' E. Lingtam: see Singtam Litien, Yunnan, China: 27° 14' N., 99° 27' E. Liu-TSUEN, Shensi, China: 34° 02' N., 108° 55' E. Lo-Gui-Ho, Tonkin, Viet Nam: 22° 19' N., 103° 42' E. LOMLOE Mt., Ban Maeo, Goksatawn, Dahnsai, Loey, Thailand: 17° 00' N., 101° 00' E. Long Lung: see Ban Nong Pla Lai LONKIN, Burma: 25° 40' N., 96° 22' E. Lonpan: see Sungpan LosHAN, Szechwan, China: 29° 35' N., 103° 40' E. Lo Tiao, Laos: 20° 20' N., 100° 05' E. Lo Tu Kow: see Lukou Lsien: see Wutingchow Luang Prabang, Laos: 19° 53' N., 102° 10' E. Lu Chang Pu, Szechwan, China: 29° 23' N., 108° 10' E. LuiA, Chaibassa, India: 22° 35' N., 85° 25' E, Lukou, Yunnan, China: 25° 55' N., 98° 50' E. LUNGCHOW, Kwangsi, China: 22° 22' N., 106° 54' E. Lunghkang (Ga), Burma: 26° 36' N., 97° 48' E. LUNGKAi, Yunnan, China: 25° 57' N., 101° 54' E. LUNGTAN, China: 32° 05' N., 119° 20' E. Lung Van: see Van Lung LUTINGKIAO, China: 29° 54' N., 102° 28' E. Luting Shan: see Lutingkiao Ma-Chang-Kai, Tengyueh, Yunnan, China: 24° 39' N., 98° 00' E. Madaya Forest, Burma: 22° 30' N., 96° 06' E, Madischung Valley, Sikang, China: 26° 12' N., 101° 59' E. Mae Hong Song: see Changwat Mae Hong Son Mae Klong, Thailand: 13° 26' N., 100° 01' E. Mae Nam Khwae Noi, Kanchanaburi, Thailand: 14° 00' N., 99° 30' E. Mae Nam Yom, Prae, Thailand: 15° 52' N., 100° 16' E. 326 FIELDIANA: ZOOLOGY, VOLUME 48 Mae Pan, Phrae, Thailand: 18° 00' N., 100° 00' E. Mae Taw Forest, Thailand: 17° 20' N., 99° 35' E. Mae Tha Khwae, Tak, Thailand: 16° 54' N., 98° 52' E. Mae Wan River near Doi Saket, Thailand: 18° 50' E., 99° 35' E. Ma Fu Ling, Hupeh, China: 31° 52' N., 110° 18' E. Maha Oya, Ceylon: 07° 30' N., 81° 19' E. Ma Kai Hsien: see Makai Makai, Yunnan, China: 25° 42' N., 101° 55' E. Malipa, Burma: 23° 40' N., 98° 45' E. Maliwun, Burma: 10° 12' N., 98° 40' E. Mamoh: see Pakchan Mandalay, Burma: 21° 59' N., 96° 08' E. Mangpu, Bengal, India: 26° 58' N., 88° 27' E. M. Angton: see Muang Phrom Buri Mankeni, Ceylon: 08° 00' N., 81° 30' E. Manoram: see Ban Khung Samphao Mansum, Burma: 25° 47' N., 96° 15' E. Manthe, Burma: 25° 18' N., 95° 13' E. Maprau Island: see Ko Maphrao Margherita, Assam, India: 27° 17' N., 95° 47' E. Martaban, Burma: 16° 30' N., 97° 28' E. Matanga River, N. Kamrup, Assam: circa 26° 30' N., 91° 30' E. Matinga: see Matanga River Matsatap, Burma: 27° 30' N., 97° 50' E. ■ Matugama, Ceylon: 06° 30' N., 80° 08' E. Maungkan, Burma: 25° 05' N., 95° 00' E. Mawlaik, Burma: 23° 38' N., 94° 22' E. Mawlyngkueng, Assam, India: 25° 34' N., 92° 03' E. Mawphlang, Assam, India: 25° 27' N., 91° 46' E. Mawryngkueng: see Mawlyngkueng Maymyo, Burma: 22° 05' N., 96° 30' E. M. Chum Pon: see Ban Tha San Mee Nan, 30 miles N.E. of Uttaradit, Thailand: 17° 50' N., 100° 15' E. Mee Pan: see Mae Pan Mee Tan: see Ban Mae Tan Nua Mehongson: see Changwat Mae Hong Son Meiktila, Burma: 20° 50' N., 95° 52' E. Mekong-Salwin [Salween] Divide, Sikang, China: 28° 20' N., 99° 20' E. Mekong Valley, Yunnan, China: 28° 00' N., 99° 23' E. Mekong- Yangtze Divide, Yunnan, China: 27° 30' N., 99° 25' E. Melamoung: see Me Lamung Me Lamung, Thailand: 15° 32' N., 98° 48' E. Me Moh: see Ban Mae Mo Meng-ting, Yunnan, China: 23° 31' N., 99° 06' E. Me Ping River [Mae Ping], 180 miles N. of Bangkok, Thailand: circa 16° 40' N., 99° 55' E. Mesarieng: see Ban Mae Sariang Me Taqua: see Mae Tha Khwae Me Taw Forest: see Mae Taw Forest Me Wong River, 53 mi. E. of Urn Pang, Thailand: 15° 50' N., 99° 38' E. MOORE AND TATE: DIURNAL SQUIRRELS 327 Me Yam River: see Mae Nam Yom MiLi, Szechwan, China: 28° 11' N., 100° 50' E. MiNGUN, Sagaing, Burma: 22° 00' N., 95° 57' E. MiSHMi Hills, Assam, India: circa 28° 00' N., 96° 10' E. M. MouEN: see Moung Mouen MOKANSHAN, Chekiang, China: 30° 26' N., 119° 47' E. MOKLOK, Burma: 25° 37' N., 95° 25' E. MOKOKCHUNG, Assam, India: 26° 24' N., 94° 32' E. MONG Chit: see Ban Tuyong Mono Ha, N. Shan States, Burma: 22° 20' N., 98° 15' E. Mongkhor: see Ban Nong Kho MONGTZE, Yunnan, China: 25° 56' N., 112° 14' E. Moung Boum, Tonkin, Viet Nam: 22° 32' N., 102° 50' E. Moung Mo, Tonkin, Viet Nam: 22° 12' N., 102° 56' E. Moung Mouen, Tonkin, Viet Nam: 21° 30' N., 103° 00' E. Moung Moun, Tonkin, Viet Nam: 21° 36' N., 103° 20' E. Moung Yo, Laos: 21° 28' N., 101° 52' E. Mt. Angka: see Doi Inthanon Mt. Arizan, Formosa: 23° 32' N., 120° 46' E. Mt. Ava, Burma: 21° 52' N., 96° 01' E. Mt. Fan-si-pan, Tonkin, Viet Nam: 22° 19' N., 103° 49' E. Mt. Imaw Bum, Burma: 26° 08' N., 98° 30' E. Mt. Mooleyit: see Mulai-yit Hill Mt. Nwalabo, Burma: 14° 07' N., 98° 28' E. Mt. Omei, Szechwan, China: 29° 25' N., 103° 25' E. Mt. Popa, Burma: 20° 58' N., 95° 20' E. Mt. Tura, Assam, India: 25° 27' N., 90° 30' E. Mt. Victoria, Burma: 21° 17' N., 93° 53 'E. Mt. Wuchi: see Wuchi Shan Moupine: see Muping MUAKLEK [Muak Lek], Sraburi [Sara Buri], Thailand: 14° 35' N., 101° 17' E. Muang Chainat, Thailand: 15° 10' N., 100° 10' E. Muang Chiang Mai, Thailand: 18° 45' N., 99° 00' E. Muang Chiang Rai, Thailand: 19° 55' N., 99° 50' E. Muang Lampang, Thailand: 18° 18' N., 99° 31' E. Muang Nan, Thailand: 18° 47' N., 100° 47' E. Muang Nong Khai, Thailand: 17° 52' N., 102° 44' E. Muang Pai: see Ban Wiang Tai Muang Phichit: see Ban Nai Muang Muang Phitsanulok, Thailand: 16° 50' N., 100° 15' E. Muang Phrom Buri, Anghtong or Sing Buri, Thailand: 14° 50' N., 100° 26' E. Muang Prom: see Muang Phrom Buri Muang Qua Yie: see Nakhon Sawan Mu Cheng, Yunnan, China: 23° 45' N., 99° 10' E. Mudumalai, India: 11° 37' N., 76° 31' E. MuEK Lek: see Ban Muak Lek Mulai-yit Hill, Burma-Thailand border: 16° 05' N., 98° 43' E. MUNSIN, Burma: 25° 33' N., 95° 20' E. MuONG Boum, Tonkin, Viet Nam: 22° 32' N., 102° 50' E. MUONG Karbin: see Ban Kabin Buri 328 FIELDIANA: ZOOLOGY, VOLUME 48 MuONG Mo, Tonkin, Viet Nam: 22° 12' N., 102° 56' E. MuONG MoUN, Tonkin, Viet Nam: 21° 36' N., 103° 20' E. MuoNG Sen, Vinh, Viet Nam: 19° 32' N., 104° 07' E. MuONG Yo, Laos: 21° 28' N., 101° 52' E. Mupin: see Muping MUPING, Szechwan, China: 30° 29' N., 102° 49' E. Myawadi, Burma: 16° 41' N., 98° 31' E. Myitkyina, Burma: 25° 24' N., 97° 26' E. Nagarjun, Katmandu, Nepal: 27° 45' N., 85° 22' E. Nagchuka: see Hokow Naggenjung: see Nagarjun Na Hai, Tonkin, Viet Nam: 21° 12' N., 102° 55' E. Nakhon Fathom, Lum Phaya, Thailand: 13° 50' N., 100° 05' E. Nakhon Sawan, Thailand: 15° 41' N., 100° 07' E. Nakhon Si Thammarat, Thailand: 8° 26' N., 99° 58' E. Nakonrajasima [Nakhon Ratchasima], Thailand: 14° 57' N., 102° 05' E. Nakon Sawan: see Nakhon Sawan Nam Chuh: see Ban Nam Chut Yai Nam Fong, Hainan, China: 19° 24' N., 109° 33' E. Nam Khueng, Laos: 20° 25' N., 100° 18' E. Nam Nen, Tonkin, Viet Nam: circa 21° 40' N., 103° 15' E. Nam Tamai Valley, Burma: 27° 42' N., 97° 54' E. Nam Ting, Yunnan, China: 23° 31' N., 99° 06' E. Nam-ting River at Burma Border, China: 23° 23' N., 98° 43' E. Nan: see Muang Nan Nan-Tou Hsien, Wu Sheh, Taiwan, China: 23° 54' N., 120° 42' E. Nanyaseik, Burma: 25° 37' N., 96° 36' E. NAPfi, Laos: 18° 20' N., 105° 05' E. Narathiwat, Thailand: 6° 25' N., 101° 50' E. Na Tebbu land, Wantsang Ku, Kansu, China: circa 34° 00' N., 101° 30' E. Nauswa, Burma: 25° 40' N., 95° 20' E. Nawakot, Nepal: 27° 56' N., 85° 10' E. Nawocot: see Nawakot N'BUNGHKU, Burma: 25° 54' N., 96° 12' E. Nelliampathi Hills, India: 10° 30' N., 76° 35' E. Ngai Tio, Tonkin, Viet Nam: 22° 35' N., 103° 45' E. Ngan-son, Tonkin, Viet Nam: 22° 25' N., 106° 00' E. Ngapyiniun, Burma: 22° 32' N., 96° 00' E. Ngau-Tchi-Lea (Mts.), Hainan, China: 19° 00' N., 109° 30' E. Nghia Hung, Phu Qui, Annam: 19° 20' N., 105° 20' E. Ngoloshi: see Tsungshi Ngu-luko, Yunnan, China: 27° 03' N., 100° 12' E. Nhatrang, Annam, Viet Nam: 12° 15' N., 109° 15' E. NiKAWERATlYA, Ceylon: 07° 44' N., 80° 08' E. Nikawewa, near Kantalai, Ceylon: 08° 16' N., 81° 01' E. Nimrod Sound, Chekiang, China: 29° 40' N., 121° 50' E. NiNGPO, Chekiang, China: 29° 53' N., 121° 33' E. NiNGYUANFU, Szechwan, China: 27° 55' N., 102° 18' E. NiNGYUEN FU: See Ningyuanfu Ninh Hoa, Annam, Viet Nam: 12° 30' N., 109° 10' E. MOORE AND TATE: DIURNAL SQUIRRELS 329 NODOA, Hainan, China: 19° 31' N., 109° 34' E. NOKREK, India: 25° 28' N., 90° 19' E. NoNG Bua: see Ban Nong Bua NoNG DOM Ta: see Ban Nong Don Ta NONGKAi: see Muang Nong Khai NONGKHOR: see Ban Nong Kho Nong Lum, Tonkin, Viet Nam: 22° 22' N., 102° 53' E. NONGPOH, Assam, India: 25° 55' N„ 91° 51' E. NONGSTOIN, Assam, India: 25° 31' N., 91° 16' E. Nong Yang, Thailand: 13° 10' N., 101° 30' E. NUA Chua Chan, Bien Hoa Prov., Viet Nam: 10° 56' N., 107° 23' E. Nui Kai, Yunnan, China: 26° 05' N., 100° 00' E. Nyetmaw River, Burma: 26° 05' N., 98° 18' E. Okma, Burma: 22° 35' N., 94° 56' E. Ok Pyam: see Pyam Ok Yam: see Pyam OOTACAMUND, Nilgiri Hills, India: 11° 24' N., 76° 44' E. Pa Ham, Tonkin, Viet Nam: circa 21° 35' N., 102° 55' E. Pahpoon: see Papun Pah Toop: see Pha Tup Pakchan: see Ban Pak Chan Pak Chong: see Ban Pak Chong Pakha: see Pa Kha Pa Kha, Tonkin, Viet Nam: 22° 33' N., 104° 18' E. Pak Hin Bun, Laos: 17° 35' N., 104° 40' E. Pak Jong: see Ban Pak Chong Pak Koh, Thailand: 18° 02' N., 99° 53' E. Paknampho: see Ban Pak Nam Pho PAK-Sfi, Laos: 15° 09' N., 105° 45' E. Paksong, Laos: 15° 15' N., 106° 07' E. Pak Tha, Thailand: circa 18° 00' N., 100° 00' E. Pak Tha Pujeg: see Ban Pak Tho Palasbari, Assam, India: 26° 10' N., 91° 35' E. Palghat, Malabar, India: 10° 40' N., 76° 35' E. Palni Hills, Madura Dist., Madras, India: circa 10° 20' N., 77° 30' E. Pang Mae Ton, Chiang Mai, Thailand: 18° 55' N., 99° 15' E. Pang Me Ton: see Pang Mae Ton Pang Nam Un, Nan, Thailand: circa 18° 30' N., 100° 33' E. Pang Sok: see Sathani Pang Sok Pantha, Burma: 23° 50' N., 94° 31' E. Paotaram: see Sathani Photharam Papun, Burma: 18° 05' N., 97° 22' E. Pasa, Laos: 19° 40' N., 102° 25' E. Pasbok, Darjeeling, Bengal, India: 27° 10' N., 88° 25' E. Pattipola, Ceylon: 06° 51' N., 80° 50' E. Pemionchee, Sikkim, India: 27° 20' N., 88° 17' E. Pennan (or Pangan) Island, Siam: see Ko Phangan Phanrang, Annam, Viet Nam: 11° 34' N., 108° 59' E. Pha Tup, Northern Thailand: South of Muang Nan Phawzaw, Burma: 25° 28' N., 95° 20' E. 330 FIELDIANA: ZOOLOGY, VOLUME 48 Phitsanuloke, Siam: 16° 50' N., 100° 15' E. Phong Saly, Laos: 21° 42' N., 102° 09' E. Phong Tho, Tonkin, Viet Nam: 22° 20' N., 103° 20' E. Phra Nakhon Si Ayutthaya, Ayutthaya, Thailand: 14° 20' N., 100° 35' E. Phu Do, Kan Luang, Nakon Phanom, Thailand: circa 16° 55' N., 104° 30' E. Phuket Island: see Ko Phuket Phu Kho, Kan Luang, Nakon Phanon, Thailand: circa 16° 55' N., 104° 30' E. Phu Kobo, Laos: 19° 22' N., 103° 23' E. Phu Phan, Sakon Nakon, Thailand: 16° 52' N., 104° 23' E. Phu Qui, Annam, Viet Nam: 19° 25' N., 105° 22' E. Piam: see DeLisle Picket: see Ban Nai Muang PiBN Ngai, Szechwan, China: 29° 12' N., 108° 10' E. Ping Tung, Taiwan, China: 22° 40' N., 120° 30' E. PiNTENNE, Ceylon: 07° 21' N., 81° 00' E. Pitsanulok: see Muang Phitsanulok Plateau Bolovens, Laos: 15° 10' N., 106° 20' E. Poi, Assam, India: 25° 20' N., 94° 37' E. PONMUDI, Travancore, India: 08° 44' N., 77° 07' E. POOKEIO: see Ban Phu Kheio Pookeiochaiyabhoom: see Ban Phu Kheio PoOTOO: see Puto Shan Pot ARAM: see Sathani Photharam PoWAi : see Poi Pra (Mt.), Cambodia: 12° 38' N., 103° 02' E. Prachuap Khiri Khan, Thailand: 11° 50' N., 99° 50' E. Pran: see Prachuap Khiri Khan and Pran Buri Pran Buri, Prachuap Khiri Khan, Thailand: 12° 22' N., 99° 56' E, Prapoot Mt.: see Khao Phra Phut Prapootabat: see Khao Phra Phut Prome, Burma: 18° 50' N., 95° 14' E. Pua: see Ban Pua Pucheng, Fukien, China: 28° 05' N., 118° 28' E. Pujeg: see Pujek PuJEK, Pak Tho, Rajburi, Thailand: 13° 23' N., 99° 42' E. Pukpitiya : see Puwakpitiya PULO Beman: see Clara Island PULO Ketchen: see St. Lukes Island PULO Lambi: see Sullivan's Island PULO Salakring: see Hastings Island PUM-KATONG, Burma: 25° 23' N., 97° 44' E. PUMSIN, Burma: 25° 57' N., 96° 11' E. PuTO Shan, Chusan Archipelago, Chekiang, China: 30° 00' N., 122° 25' E. Puwakpitiya, Gammaduwa, Ceylon: 07° 36' N., 80° 43' E. Pyam, Kampot, Thailand: 11° 37' N., 102° 56' E. Pyaungbyin, Burma: 24° 15' N., 94° 47' E. Pyaunggaung, Burma: 22° 35' N., 97° 05' E. Pyepat, Burma: 25° 54' N., 98° 15' E. Pynursla, Assam, India: 25° 18' N., 91° 53' E. Quangtri Phuoc Mon, Annam, Viet Nam: 16° 35' N., 107° 10' E. MOORE AND TATE: DIURNAL SQUIRRELS 331 QUANG Tri River, Annam, Viet Nam: circa 16° 46' N., 107° 11' E. Racu Racu Mountains: see Rakuraku Raheng: see Ban Rahaeng Rahnampo: see Ban Pak Nam Pho Rajaburi: see Rat Buri Rajburi: see Ban Pong Rakuraku, Taiwan, China: 23° 27' N., 121° 30' E. Rama la Pass, Szechwan, China: 29° 40' N., 100° 25' E. Rangoon, Burma: 16° 47' N., 96° 10' E. Ranna, Ceylon: 06° 05' N., 80° 55' E. Rat Buri, Thailand: 13° 32' N., 99° 49' E. Ratnamti, Burma: 27° 35' N., 97° 55' E. Red Point, Tenasserim, Burma: 10° 40' N., 98° 30' E. Ringin, Sikkim, India: 27° 30' N., 88° 30' E. RiRi Bazaar, Nepal, India: 27° 55' N., 83° 22' E. Rumitchangu, Szechwan, China: 30° 50' N., 102° 05' E. RUNGBONG Valley, Sikkim, India: 26° 50' N., 88° 13' E. Sadiya, Assam, India: 27° 50' N., 95° 45' E. St. Lukes Island, Burma: 10° 05' N., 98° 15' E. Sakeo: see Ban Sa Kaeo Sakerat : see Ban Chakkrarat Salanga: see Koh Phuket Salangka: see Koh Phuket Salween-Mekong Divide at 28° 20' N.: 28° 20' N., 98° 35' E. Samasgi, Dharwar-Kanara border, India: 14° 40' N., 75° 00' E. Sambor, Cambodia: 12° 45' N., 105° 59' E. Sam roi Gop: see Sathani Sam Roi Yot Sangau, Assam, India: 21° 44' N., 93° 03' E. Sangsir: see Singsir Sata Hip: see Ban Sattahip Sathani Hin Lap, Thailand: 14° 40' N., 101° 10' E. Sathani Lamphura, Thailand: 07° 40' N., 99° 35' E. Sathani Pang Hua Phone, Thailand: 18° 25' N., 99° 15' E. Sathani Pang Sok, Sara Buri, Thailand: 14° 40' N., 101° 20' E. Sathani Photharam, Rat Buri, Thailand: 13° 40' N., 99° 50' E. Sathani Sam Roi Yot, Thailand: 12° 15' N., 99° 55' E. Sathani Tha Chomphu, Lamphun, Thailand: 18° 30' N., 99° 15' E. Sathani Thung Song, Nakhon Si Thammarat, Thailand: 08° 10' N., 99° 40' E. Sattha Hill: see Satthar Satthar, Nepal: 28° 01' N., 84° 35' E. Sawankaloki: see Ban Wang Mai Khon Sawankhalok: see Ban Wang Mai Khon Seechol: see Ban Sichon Sedonchen, Sikkim, India: 27° 30' N., 88° 30' E. Se'en, N. Shan States, Burma: 22° 39' N., 97° 22' E. Seniku, Burma: 25° 34' N., 97° 47' E. Sevoke, Bengal Presidency, Pakistan: 26° 51' N., 88° 32' E. Shandaw, Burma: 17° 55' N., 95° 40' E. Shanghai, China: 31° 13' N., 121° 25' E. Shang Kuan, Yunnan, China: 25° 55' N., 100° 05' E. 332 FIELDIANA: ZOOLOGY, VOLUME 48 Shihpingchow, Yunnan, China: 23° 40' N., 102° 25' E. Shinei, Formosa, China: 23° 18' N., 120° 18' E. Shineigun: see Shinei Shingbwiyang, Hukawng Valley, Burma: 26° 41' N., 96° 12' E. Shoo-O-Lo: see Siolo Shrivilliputtur [Srivilliputtur], Madras, India: 07° 30' N., 77° 38' E. Shuan Lung Chang, Kweichow, China: 28° 05' N., 106° 48' E, Shui-Kow-Kwan: see Lungchow SHWEBO, Burma: 22° 35' N., 95° 42' E. Shweli Valley, Yunnan, China: circa 25° 00' N., 98° 40' E. Shweli-Salween Divide, Yunnan, China: circa 25° 05' N., 98° 45' E. SiAOMENGTUNG, Yunnan, China: 24° 10' N., 99° 15' E. SiEM Reap, Cambodia: 13° 20' N., 103° 51' E. Siken: see Ban Si Khiu Sikeu: see Ban Si Khiu SiMA, Myitkyina, Burma: 25° 05' N., 97° 35' E. Singkaling Hkamti, Burma: 26° 00' N., 95° 39' E. SiNGSiR, Bengal Presidency, India: 27° 04' N., 88° 35' E, Singtam, Sikkim, India: 27° 14' N., 88° 33' E. SiNKAi, Yunnan, China: 24° 12' N., 102° 04' E. Siolo, Szechwan, China: 30° 02' N., 100° 48' E. Siracha: see Ban Si Racha SiRSA, Hissar, Punjab, Pakistan: 29° 30' N., 75° 00' E. SiSAGARHi, Nepal: 27° 25' N., 85° 05' E. Slokbai: see Ban Salok Bat SoKOTAi: see Ban Thani SOOKIA POKHARi, Darjeeling, India: 27° 02' N., 88° 14' E. SOOKLI: see Sukli South Cape of Formosa, China: 21° 54' N., 120° 52' E. Sriracha : see Ban Si Racha SUCHOW, Szechwan- Yunnan border, China: 28° 43' N., 104° 32' E. SuiFU: see Suchow SuiYANG, Kweichow, China: 27° 50' N., 107° 18' E. SUKHOTHAi: see Ban Thani SUKIAPOKHRI : see Sookia Pokhari SUKLI, Myawadi Rd., Burma: 16° 42' N., 98° 22' E. Sullivan's Island, Mergui Archipelago, Burma: 10° 40' N., 97° 58' E. SUMKA Uma, Burma: 25° 57' N., 97° 49' E. SUMPRABUM, Burma: 26° 38' N., 97° 36' E. Sungei Balik, Tenasserim, Burma: 10° 31' N., 98° 33' E. Sungpan, Szechwan, China: 32° 41' N., 103° 21' E. Sunjang: see Suiyang Sung Shan, Honan, China: 34° 35' N., 113° 20' E. Ta Chang Tai: see Tha Chang Tai Ta Chiao, Szechwan, China: circa 30° 03' N., 103° 13' E. Tachin: see Ban Maha Chai Taga Hka, Burma: 26° 22' N., 96° 07' E. Tagoot, Burma: 10° 25' N., 99° 18' E. Ta-ho: see Ta Ho-Hta Ta Ho-Hta, Independent Kareni, Burma: 19° 06' N., 97° 30' E. MOORE AND TATE: DIURNAL SQUIRRELS 333 Tainan, Taiwan, China: 23° 00' N., 120° 12' E. Tai-Pa-Shiang Mts. : see Tapa shan Taipei, Taiwan, China: 25° 05' N., 121° 32' E. Taipei Hsien, Wu Lai, Taiwan, China: 24° 50' N., 121° 32' E. Tai-pei-shan District, Shensi, China: 33° 56' N., 107° 41' E. Tai-ping-pu, Yunnan, China: 24° 57' N., 98° 42' E. Tai-yuan-fu, Shansi, China: 37° 53' N., 112° 29' E. Tak: see Ban Rahaeng Takubama, Assam, India: 25° 50' N., 94° 28' E. Tamabin, Burma: circa 17° 45' N., 96° 30' E. Tamanthi, Burma: 25° 20' N., 95° 12' E. Tam Dao, Tonkin, Viet Nam: 21° 30' N., 105° 38' E. Tamu, Burma: 25° 45' N., 98° 02' E. Tanga, Burma: 25° 50' N., 98° 05' E, Tangshu, Gieu Long Shien, Tibet: 28° 58' N., 98° 08' E. Tang Gu: see Tangshu? Tanjong Badak, Tenasserim, Burma: 10° 06' N., 98° 29' E, Taok, Tenasserim, Burma: 16° 20' N., 98° 28' E. Tag Mung Chung, Lu-tien, Yunnan, China: 27° 12' N., 103° 31' E. Tapa Hka, Burma: 26° 07' N., 96° 15' E. Tapa shan, Shensi, China: 32° 40' N., 107° 30' E. Tappan-sha, Formosa, China: 23° 28' N., 120° 44' E. Tapposha: see Tappan-sha Tarkhola, Sikkim, India: 27° 07' N., 88° 33' E. Taroar: see Tagoot Taron Valley, Burma: circa 28° 00' N., 98° 10' E. Ta-shui-tang: see Tashutang Tashutang, Yunnan, China: 24° 17' N., 99° 15' E. Tasu Bum, Burma: 26° 04' N., 96° 14' E. Tatkon, Burma: 23° 45' N., 94° 23' E. Tatsienlu, Szechwan, China: 30° 03' N., 102° 13' E. Taukchandmai, Burma: 23° 30' N., 94° 35' E. Tavoy, Burma: 14° 07' N., 98° 18' E. Tavoy Island, Mergui Archipelago, Burma: 11° 00' N., 98° 20' E. Tawmaw, Burma: 25° 44' N., 96° 18' E. Tay Ninh, Cochin China, Viet Nam: 11° 12' N., 106° 02' E Tehan, China: circa 30° 20' N., 117° 20' E. Tellula, Ceylon: 06° 36' N., 81° 08' E. Telok Besar, Tenasserim, Burma: 10° 22' N., 98° 35' E. Tempao, Burma: 24° 59' N., 94° 56' E. Tenasserim Town, Burma: 12° 06' N., 99° 03' E. Tengyueh, Yunnan, China: 25° 00' N., 98° 30' E. Teraso : see Teraso Soan Teraso Soan, Formosa, China: 22° 01' N., 120° 51' E. Tha Chang: see Ban Tha Chang Tha Chang Tai, Tak, Thailand: 16° 51' N., 99° 03' E. Thagyet, Burma: 12° 06' N., 99° 06' E. Thai Nguyen, Tonkin, Viet Nam: 21° 30' N., 105° 55' E. Thai Nien: see Thai Nguyen Thale Song Hong, Thailand: 7° 50' N., 99° 27' E. 334 FIELDIANA: ZOOLOGY, VOLUME 48 Thamaung: see Thandaung Thandaung, Burma: 19° 05' N., 96° 38' E. Than Hoa, Annam, Viet Nam: 19° 49' N., 105° 48' E. Tha Rong: see Ban Wichian Thateng: see Ban Thateng Thaton, Burma: 16° 50' N., 97° 18' E. Thatone: see Thaton Thaungyin Valley, Tenasserim, Burma: circa 16° 42' N., 98° 30' E. Theng-gan-ngok, Tenasserim, Burma: 13° 20' N., 98° 55' E. Thobal, Manipur, India: 24° 18' N., 94° 02' E. Thotel: see Thobal Thowngyoh: see Theng-gan-ngok Thua Luu, Annam, Viet Nam: circa 16° 19' N., 108° 00' E. Thuangyin Valley: see Thaungyin Valley TiENCHUAN, Szechwan, China: 30° 05' N., 103° 00' E, TiNGCHOW, Hopeh, China: 38° 30' N., 115° 02' E. Tingchowfu, Fukien, China: 25° 44' N., 116° 27' E. TiNNPANi: see Tirappane TiRAPPANE, Ceylon: 08° 12' N., 80° 31' E. Ti-yu Gomba, Yunnan [Sikang], China: circa 28° 40' N., 101° 10' E. Tongka: see Koh Phuket TOONG, Sikkim, India: 27° 35' N., 88° 40' E. Torsan: see Ban Tha San TOUNGOO, Burma: 18° 55' N., 96° 25' E. Tra Khanun: see Ban Tha Khanum Trang Bom, Cochin China, Viet Nam: 10° 50' N., 107° 00' E. Trashi-cho-ten, Szechwan, China: circa 30° 20' N., 102° 20' E, Trivandrum, Travancore, India: 08° 29' N„ 76° 55' E. TsAO Po, Szechwan, China: 21° 05' N., 103° 31' E. TSE-KOW: see Tseku TsEKU, Yunnan, China: 28° 00' N., 98° 52' E. TsEO JiA Keo, Szechwan, China: circa 28° 18' N., 104° 12' E. TsiNGKiEN, Shensi, China: 37° 12' N., 110° 12' E. TsiNG-LiNG Mts., Shensi, China: circa 33° 00' N., 108° 00' E. TsiNGTEH, Anhwei, China: 30° 23' N., 118° 30' E. TSONMA, Burma: 26° 10' N., 98° 35' E. TsuK KON Shih: see Tungshih TsUNGSHi: see Tungshih TSUNYI HsiEN, Kweichow, China: 27° 35' N., 107° 02' E. TUNG-LiNG: see Eastern Tombs TUNGLU, Chekiang, China: 29° 47' N., 119° 37' E. Tung Nog Karien, Jombing, Rajburi, Thailand: 13° 33' N., 99° 35' E. Tung Sawn: see Sathani Thung Song Tungshih, Hupeh, China: 30° 22' N., 111° 42' E. TUNGTZE, Kweichow, China: 27° 58' N., 106° 51' E. Tung Wong Tien, Kweichow, China: 28° 03' N., 105° 50' E. Tu-PA-KEO, Moupine, Szechwan, China: circa 30° 30' N., 102° 45' E. Tura, Garo Hills, Assam, India: 25°31'N., 90°16'E. Ukhrul, Manipur, India: 25° 10' N., 94° 18' E. Ulongkong, Szechwan, China: circa 29° 55' N., 102° 10' E. MOORE AND TATE: DIURNAL SQUIRRELS 335 Um Pang: see Ban Le Kathe Umran, Assam, India: 25° 65' N., 91° 50' E. Uttaradit, Thailand: 17° 38' N., 100° 06' E. Van Lung, Thanhoa, Annam, Viet Nam: 20° 30' N., 104° 40' E. ViCHiANBURi: see Ban Wichian Victoria Point, Tenasserim, Burma: 10° 00' N., 98° 30' E. Vientiane, Laos: 17° 45' N., 102° 40' E. Wan Mts., Anwei, China: circa 30° 45' N., 117° 55' E. Wang Kien: see Kanchanaburi Wan hsien, Szechwan, China: 30° 49' N., 108° 23' E. Wan-tien, Yunnan, China: 24° 31' N., 99° 21' E. Wa Shan, Szechwan, China: circa 29° 25' N., 104° 20' E. Waterfall, Mekang: see Ban Mae Klang Wa TIEN, China: 25° 20' N., 98° 36' E. Wei Chow: see Weikiu Weikiu, Szechwan, China: 31° 25' N., 103° 28' E. Weisi Pass, Yunnan, China: 27° 14' N., 99° 27' E. Wellawaya, Uva Prov., Ceylon: 06° 43' N., 81° 06' E. Wenchwan, Szechwan, China: 31° 20' N., 103° 31' E. Wen-shui, Kweichow, China: 28° 25' N., 106° 18' E. Western Hills, 15 mi. W. of Peking, Hopeh, China: 39° 57' N., 116° 07' E. WiE Shi Pass: see Weisi Pass Wu-chi: see Wushi WuCHi Shan, Hainan, China: 18° 55' N., 109° 20' E. WUNG Pratart Farm, Kam Peng Pet, Thailand: 16° 10' N., 99° 45' E. WUNTHO, Burma: 23° 56' N., 95° 45' E. WuSHi, Yunnan, China: 29° 05' N., 101° 28' E. WUTAI, Shansi, China: 38° 55' N., 113° 38' E. WuTiNGCHOW, Yunnan, China: 25° 30' N., 102° 26' E. WuTiNG Hsien: see Wutingchow Wu-TING-SHAN, Jehol, Manchuria: 40° 42' N., 117° 21' E. Wu-TSAi: see Wutai XiENG Khouang, Laos: 19° 25' N., 103° 25' E. XiEN QUANG Koo: see Xieng Khouang Yachow: see Yachowfu Yachowfu, Szechwan, China: 29° 59' N., 103° 10' E. Yalung River, S.W. Szechwan, China: circa 27° 10' N., 101° 50' E. Yangwupa, Yunnan, China: 23° 56' N., 102° 10' E. Yao-shan, Kwangsi, China: 24° 00' N., 110° 00' E. Yellapur, Bombay, India: 14° 55' N., 74° 40' E. Yen-an-fu, Shensi, China: 36° 44' N., 109° 25' E. Yenping, Fukien, China: 26° 38' N., 118° 10' E. Yin, Burma: 22° 43' N., 94° 42' E. Yiu Shan: see Yu Shan YUKi, Fukien, China: 25° 28' N., 119° 18' E. Yulong-Vlang: see Ulongkong Yung Cha Shan, Szechwan, China: 29° 23' N., 108° 10' E. Yunghtang, Burma: 27° 38' N., 98° 03' E. Yung-Ning, Yunnan, China: 27° 45' N., 100° 41' E. Yungpeh, Szechwan, China: 26° 40' N., 100° 45' E. 336 FIELDIANA: ZOOLOGY, VOLUME 48 Yung-pei-ting: see Yungpeh YUNNANFU, Yunnan, China: 25° 00' N., 102° 40' E. YUNNANYi, Yunnan, China: 25° 25' N., 100° 37' E. Yu SHAN, Kwangtung, China: 24°42'N.,114°10'E. Zamayi Res., Pegu Yoma, Burma: 18° 21' N., 95° 59' E. Zami River, 100 miles south of Moulmein, Burma: 15° 25' N., 98° 25' E. Zaulep Ga, Burma: circa 25° 57' N., 96° 11' E. Zaungtu, Burma: circa 17° 50' N., 96° 30' E. REFERENCES Abdulali, Humayun, and J. C. Daniel 1952. Races of the Indian giant squirrel (Ratufa indica). Jour. Bombay Nat. Hist. Soc, 50, pp. 469-474, colored map. 1953. A colour variation, and albinism in the giant squirrel Ratufa indica. Jour. Bombay Nat. Hist. Soc, 51: no. 3, p. 731. Allen, Glover Morrill 1925. Squirrels collected by the American Museum Asiatic expeditions. Amer. Mus. Novitates, no. 163, pp. 1-16. 1940. The mammals of China and Mongolia. Natural History of central Asia, 11, pt. 2 (Rodents, Ungulates and Sirenians), pp. 618-1350, figs. 24-75. New York, American Museum of Natural History. Allen, Glover Morrill, and H. J. Coolidge 1940. Mammal and bird collections of the Asiatic primate expedition. Bull. Mus. Comp. Zool., 87, pp. 121-166. 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The squirrels of the Funamhulus palmarum-tristriatus group in the peninsula. Jour. Bom- bay Nat. Hist. Soc, 24, pp. 644-649. 1919. Summary of the results from the Indian Mammal Survey of the Bombay Natural History Society. Part III. Jour. Bombay Nat. Hist. Soc, 26, no. 2, pp. 338-379. 1921, On the erythraeus group of squirrels. Jour. Bombay Nat. Hist. Soc, 27, pp. ns-m. Wroughton, R. C, and W. M. Davidson 1919. Scientific results from the mammal survey. No. 20. C. Two new forms of the "Funambulus tristriatus" group. Jour. Bombay Nat. Hist. Soc, 26. pp. 728-730. Zahn, Walter 1942. Die Riesen-, Streifen-, und Spitsnasenhornchen der Orientalischen Re- gion. Zeitschrift fur Saugetierkunde, 16, no. 1, pp. 1-182, 6 maps. INDEX Abdulali, Humayun 9, 44, 47 adamsoni = Dremomys rufigenis rufigenis 280 adangensis Callosciums caniceps 189 affinis, Ratufa 33 agouti pelage 14 albifer = Callosciurtis flavimanus styani 152 albivexilli Callosciurus finlaysoni 167 Allen. G. M. (1940) 20 altinsularis Callosciurus caniceps 195 amotus = Menetes berdmorei berdmorei 297 Anderson, John (1878) 18 annandalei =Funambulus tristriatus tristriatxis 87 annellatus Callosciurus finlaysoni 172 Anthony, Harold E. 9 aquilo — Callosciurus erythraetis intermedius 110 Archbold Expeditions 7 argentescens Funambulus pennanti 76 assamensis = Callosciurus pygerythrus lokroides 227 atrodorsalis Callosciurus flavimanus 138 Baginia = Callosciurus 99 harbei Tamiops mcclellandi 241 bartoni Callosciurus erythraeus 117 heebei = Callosciurus flavimanus quinqu£striatus 126 belfi£ldi Dremomys rufigenis 282 bellaricus Funambulus palmarum 81 bellona == Callosciurus pygerythrus m^arsi 222 bengalensis Funambulus palmarum 82 bengalensis = Ratufa indica maxima 45 bentincanus = Callosciurus caniceps domelicus 195 berdmorei, Menetes 294 berdmorei Menetes berdmorei 297 bhotia = Dremomys lokriah lokriah 264 bhutanensis Callosciurus erythraeus 108 bicolor, Ratufa 49 bimaculatus Callosciurus caniceps 190 Biswas, Biswamoy 9 Black, Craig C. 27 Blanford (1888-91) 19 blanfordi = Callosciurus phayrei 201 blythi Callosciurus pygerythrus 226 bocourti Callosciurus finlaysoni 179 bolovensis = Callosciurus flavimanus flavimanus 146 bombayus = Ratufa indica indica 43 bonhotei Callosciurus flavimanus 151 boonsongi Callosciurus finlaysoni 184 brodiei Funambulus palmarum 85 Burt, W. H. 8 calidior Dremomys pernyi 276 Callosciurus 99 caniceps 187 erythraeus 101 ferrugineus 158 flavimanus 120, 189 finlaysoni 161 inornatus 209 phayrei 201 pygerythrus 213 caniceps, Callosciurus 187 camceps Callosciurus caniceps 196 canigenus = Callosciurus flavimanus styani 152 careyi = Callosciurus erythraeus bartoni 117 Carter, T. Donald 305 casenszs Callosciurus caniceps 194 345 346 INDEX castaneoventris Callosdurus flavimanus 143 ceilonensis = Ratufa macroura macroura 37 celaenopepla =Ratufa bicolor phaeopepla 52 centralis Ratufa indica 47 ceylonicus = Ratufa macroura macroura 37 chintalis =Dremomys pernyi calidior 276 chrysonotv^ = Callosdurus caniceps caniceps 196 cinnammneus Callosdurus finlaysoni 170 clarkd = Tamiops swinhod swinhod 248 classification 20 cockerelli = Callosdurus finlaysoni bocourti 179 collaris —Sdurotamias davidianus consobrinus 308 collinus Tamiops m^clellandi 239 color names 14 comorinus Funximhulus palmar um 81 concolor Callosdurus caniceps 187 condorenMs Ratufa bicolor 58 consobrinus Sdurotamias davidianus 307 consularis Menetes berdmord 300 contumax = Callosdurus flavimanus flavimanus 146 Cranbrook, Lord 63, 113, 120, 121, 239, 250, 274 crotalius =Callosdurus erythraeus erythrogaster 105 crumpi = Callosdurus erythraeus bhutanensis 108 dactylinus = Callosdurus flavimanus flavimanus 146 dandolena Ratufa macroura 38 davidianus, Sdurotamias 304 davidianus Sdurotamias davidianus 307 Davis, D. Dwight 291 davisoni = Callosdurus caniceps bimacuJatus 190 dealbata Ratufa indica 41 decoratus Menetes berdmord 299 Deignan, Herbert G. 9 delesserti =Funambulus sublineatus sublineatus 95 Deraniyagala, P. E. P. 9 dextralis = Callosdurus finlaysoni dnistralis 177 dimensions 15 diurnal squirrels 10, 12 dolphoides = Tamiops rodolphd rodolphd 244 domelicus Callosdurus caniceps 195 Dorst, M. Jean 8 dravidiamis Funambulus layardi 94 Dremamys 11, 259 everetti 17, 289 lokriah 263 pernyi 269 pyrrhomerus 283 rufigenus 278 dussumieri = Funambulus tristriatus tristriatus 87 elbeli Tamiops rodolphd 245 EUerman (1940) 20 elphinstoni —Ratufa indica indica 43 Eosdurus = Ratufa 29 epomophorus = Callosdurus caniceps bimaculatus 190 erubescens = Callosdurus caniceps concolor 187 erythraeus, Callosdurus 101 erythraeus Callosdurus erythraeus 104 erythrogaster Callosdurus erythraeus 105 Erythrosdurus = Callosdurus 99 everetti 11 everetti, Dremomys 17, 289 evolution 7, 15 evolutionary implications 8 extraterritorial 11 fallax Callosdurus caniceps 195 favonicus Funambulus palmarum 83 felli Ratufa bicolor 58 ferrugineus, Callosdurus 158 finlaysoni, Callosdurus 161 Callosdurus finlaysoni 165 flavimanus, Callosdurus 120 Callosdurus flavimanus 146 Dremomys pernyi 275 INDEX 347 floweri = Callosciurus finlaysoni bocourti 179 fluminalis = Callosciurus caniceps caniceps 196 folletti Callosciurus finlaysoni 166 Forcart, L. 8 formosanus = Tamiops maritimus maritimus 252 forresti, Sciurotamias (Rupestes) 309 forresti = Tamiops sunnhoei sunnhoei 248 frandseni Callosciurus finlaysoni 167 fryanus — Callosciurus erythraeus bartoni 117 fumigatus = Callosciurus flavimanus griseimanus 148 Funambulus 65 layardi 91 palmarum 77 pennanti 72 sublineatus 95 tristriatus 87 = Dremomys rufigenis rufigenis 280 garonum Dremomys lokriah 266 Callosciurus finlaysoni 169 gigantea Ratufa bicolor 59 Callosciurus flavimanus 150 gordowi Callosciurus flavimanus 129 gossei = Funambulus palmarum palmarum 79 Granger, Walter 287 Gray (1867) 18 gnsetmaniis Callosciurus flavimanus 148 = Dremomys pernyi pernyi 271 griseopectus Blyth, 1847 = Callosciurus flavimanus shanicus 131 griseopectus Thomas, 1894 = Callosciurus flavimanus styani 152 = Callosciurus finlaysoni sinistralis 177 Dremomys pyrrhom^rus 288 = Callosciurus flavimanus michianus 134 Ratufa bicolor 63 Tamiops maritimus 255 /lanngtom Callosciurus erythraeus 118 ftarmandt Callosciurus finlaysoni 167 = Callosciurus caniceps bimaculatus 191 Hayman, R. W. 8 heinrichi = Callosciurus phayrei 201 = Callosciurus caniceps caniceps 196 AeipMs = Callosciurus caniceps caniceps 196 Callosciurus flavimanus 143 = Callosciurus finlaysoni cinnamomeus 170 Hershkovitz, Philip 8 //eierosciMrws = Callosciurus 99 Holm, A. 9 = Tamiops rodolphei rodolphei 244 Hooper, Emmett T. 8 Hora, Sunder Lai 9 Horsfield (1824) 18 Dremomys pernyi 273 Husson, A. M. 9 Hyosciurus 10 fti/peri/iferus Callosciurus flavimanus 133 iTwanus = Callosciurus flavimantis quinquestriatus 127 imitator = Callosciurus inornatus 209 imiis = Dremomys pernyi howelli 273 inconstons Tamiops m/;clellandi 240 Indian Subregion 10, 12 indica, Ratufa 41 indico Ratufa indica 43 indiciis = Funambulus palmarum palmarum 79 Indochinese Subregion 10, 12 in«xpecto