TREESHREWS: AN ACCOUNT OF THE MAMMALIAN FAMILY TUPAIID^. By Marcus Ward Lyon, Jr., Formerly of the Division of Mammals, United States National Museum. INTRODUCTION. This -review of the treeshrews, constituting the mammahan family Tupaiidse, was originally contemplated in 1904 by Mr. Gerrit S. Miller, jr., curator of mammals, United States National Museum, but owing to pressure of other work he was unable to carry it out. In 1910, shortly after I severed my active connections with the Division of Mammals, United States National Museum, Mr. Miller suggested to me the desirability of making a study of the treeshrews. I took up his suggestion and the present paper is the result. At that time he turned over to me some preliminary notes on the group he had made during a visit to European museums when he was primarily engaged in other lines of research. The increase of new material, both in the United States National Museum and in other museums, made it imperative that the entire field be gone over again. The collections in Washington were first studied, and during the summer of 1911 I visited most of the museums which Mr. Miller's previous work showed contained material valuable for this revision. Specifically, the material examined consists of about 800 speci- mens, all of which are listed in the tables of measurements and dis- tributed as follows: British Museum, 355 specimens, 27 types. United States National Museum, 324 specimens, 29 types. Civic Museum of Natural History, Genoa, 37 specimens, no types. Royal Zoological Museum, BerHn, 29 specimens, 1 type. Museum of Natural History, Paris, 20 specimens, 1 type. American Museum of Natural History, New York, 14 specimens, 1 type. Natural History Museum of Geneva, 3 specimens, no types. Natural History Museum of Turin, 1 specimen, no types. In addition to the specimens mentioned above, in most museums, particularly the older ones, there are a number of specimens of very uncertain or generalized localities, which are unsuitable for systematic work, and they are not included in the above figures. Proceedings U. S. National Museum, Vol. 45--No. 1976. 80459°—Proc.N.M.vol.45—13 1 ^ 2 PROCEEDINGS OF THE NATIONAL MUSEUM. vol.45. In addition to examining specimens in these museums, I have also had for study specimens sent to Washington from the following institutions: Museum, Philippine Bureau of Science, 12 specimens, 1 type. Selangor Museum, Selangor, Straits Settlements, 8 specimens, no types. Academy of Natural Sciences, Philadelphia, 6 specimens, no types. I was unfortunate in my time of visiting the Natural History Museum in Leyden. The director. Dr. F. A. Jentink, who has direct charge of the mammals, was on his vacation and I was unable to examine the specimens of Tupaiidse in that museum. The material contained there as listed by Jentink ^ does not appear vitally impor- tant for a systematic review of the group, yet it contains some very interesting and historical specimens, which I regret not having seen. Among them are the unique type of Dendrogale murina and the only skeleton of the genus Ptilocercus that I know of existing in museums and the cotypes of Tana dorsalis. Dendrogale murina is the only species of treeshrew of which I have not seen examples. I take pleasure in here expressing my thanks to the directors of the museums which I personally visited for giving me the privilege of studying the available material in their institutions, or from which material was borrowed. The importance of the explorations of Dr. W. L. Abbott in our knowledge of the treeshrews can not be lost sight of. With the exception of less than a dozen specimens in the United States National Museum the entire series of treeshrews there was collected through his untiring efforts. This means that more than a third of the specimens of treeshrews in all the museums of America and Europe have been personally collected by Doctor Abbott. Among them are 29 types. Indirectly he is also responsible for the tree- shrews collected by Messrs. Kloss and Robinson on the Malay Peninsula, or adjacent islands. The text figures of the skulls and teeth of the various genera were made bj^ Mr. A. J. Engel Terzi. Measurements.—-All the measurements are in millimeters. With the exception of those of the head and body and of tail of skins, they have all been made by the writer, including those of the hind foot, which includes the claws. In most cases the measurements of the head and body and tail were made by the collector in the flesh. In the tables of measurements where the head and body and tail measure- ments are followed by ±, those measurements were made by the writer from the dried skin or mounted specimenf Head and body and tail measurenlents of specimens preserved in alcohol were also 1 There are, however, in the Selangor Museum, 4 types, none of which I have seen. 2 Cat. Ost6ol. Manun. Mus. Hist. Nat. Pays-Bas., vol. 9, 1887, and Cat. Syst. Mamm. Mus. Hist. Nat. Pays-Bas, vol. 12, 1888. NO. 1976. TREESHREWS: FAMILY TUPAIID^—LYON. 3 made by the writer. Measurements of the skull followed by ± indicate that the measurement is onl}?" approximate owing to damage to the skull. Hindfoot measurements followed by ± are also only approxunatc. In the tables of measurements the specimens in the United States National Museum will be recognized by simple cata- logue numbers, as 104362, without qualifying initials or footnotes; those in the British Museum by the well-known separation of these numbers into sections by means of periods, as 99. 6. 12.3; specimens in other museums will be found designated by appropriate and self- explanatory initial letters or by footnotes, HISTORICAL ACCOUNT. The earliest published account of treeslirews is that of Ellis,^ one of the surgeons of Captain Cook's expedition. On Tuesday or Wednesday, 25th or 26th of January, 1780, *Ellis remarks: ''Our sportsmen * * * having seen only a few monkies, squirrels, and a cock and hen, the latter of which they shot. According to Linnaeus this island is their native place." The island referred to is Pulo Condore, off the coast of Cochin China. The squirrels men- tioned in the account are not squirrels, but Tupaias. One of them was evidently shot. A rough but very accurate sketch of the animal was made by EUis and a Latin diagnosis of it written in his journal. This description of the animal was published by Gray in 1860.^ Through the courtesy of the officials of the British Museum a repro- duction of a photograph of Ellis's drawing is here printed. There can be no doubt from Ellis's picture or description that his squirrels were Tupaias (pi. 1). Tupaias as such were first brought to the attention of the world by M. Diard, a French naturalist, at one time an assistant of Sir Thomas Stamford Raffles, in November, 1820, under the designation of Sorex glis} Six months later. May, 1821, the genus Twpaia was first proposed by Sir Thomas Stamford Raffles,* and the species ferruginea and tana described, the latter in the present paper being made the type of a nev/ genus. Specimens of Tupaias had been seen by Europeans several years earlier, and one even sent to Europe. Geoffroy ^ remarks: The discovery of this remarkable group of Insectivores has been attributed to both M. Diard and Sir Raffles. The fact is that it belongs to neither of these celebrated travelers, but to Leschenault de la Tour, who had sent in 1807 to the Museum of Paris an individual of the species which has since been called Tupaiajavanica. Never- theless it is only since 1820 that the attention of naturalists has been called to Tupaias, and that these animals have really entered the domain of science. 1 Voyage by Capt. Cook and Capt. Gierke in sMps Resolution and Discovery, 1776, 1777, 1778, 1779, and 1780, vol. 2, 1782, p. 340. - 2 Ann. Mag. Nat. Hist., ser. 3, vol. 5, 1860, p. 71. 8 Asiatic Journ. Month. Reg., vol. 10, p. 478, November, 1820. * Trans. Linn. See. London, vol. 13, p. 256, May, 1821. * Belanger, Voyage aux Indes-Orientales, Zoologie, p. 104, 1835. , PROCEEDINGS OP THE NATIONAL MUSEUM. vol. 45. Geoffrey was naturally quite unaware of the existence of Ellis's manuscript notes and drawings. Since Diard's and Raffles's time the group has become better and better known and its geographic range widely extended. The most important discoveries in regard to the group since 1821 have been the announcement of the genus Ptilocercus by Gray/ 1848, of the group now called Dendrogale by Schlegel and Miiller/ the discovery of treeshrews in the Philippine Islands by Whitehead, about 1879,^ now formmg the genus Urogale, and the discovery of treeshrews in India by W. Elliot, about 1849.* DEFINITION AND RELATIONSHIPS. The Tupaiidse are diurnal insectivorous mammals characterized by a general squuTel-like aspect, more or less arboreal habits, orbits completely encircled by bone, alisphenoid canal present, malar bone with a more or less enlarged perforation, separate radius and ulna, and separate tibia and fibula, dental formula / § C }, Pm f M. f upper molars with typical W pattern. The family is composed of two very distinct groups for a long time regarded as genera, the typical members of the family, Tupaia and the aberrant Ptilocercus. The old genus Tupaia has gradually been seen to be a composite genus, and up to the present time has been divided into three sepa- rate genera: Tupaia, Dendrogale, Urogale. In the present paper two more genera are recognized. These genera are now for the first time grouped to form the subfamily Tupaiinse.