HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology S -- N A-^ti University of Kansas Publications Museum of Natural History Volume 4, pp. 1-466, plates 1-41, 31 figures in text December 27, 1951 ^' >'' AMERICAN WEASELS BY E. RAYMOND HALL University of Kansas Lawrence 1951 \ UNIVERSITY OF KANSAS PUBLICAUONS The University of Kansas Publications, Museum of Natural His- tory, are offered in exchange for the publications of learned societies and institutions, universities and libraries. For exchanges and in- formation, address the Exchange Desk, University of Kansas Li- brary, Lawrence, Kansas, U. S. A. Museum of Natural History. — E. Raymond Hall, Chairman, Editorial Com- mittee. This series contains contributions from the Museum of Natural History. Cited as Univ. Kans. Publ., Mus. Nat. Hist. Vol. 1. 1. The pocket gophers (genus Thoniomys) of Utah. By Stephen D. Durrant. Pp. 1-82, 1 figure in text. August 15, 1946. 2. The systematic status of Eumeces pluvialis Cope, and noteworthy records of other amphibians and reptiles from Kansas and Okla- homa. By Hobart M. Smith. Pp. 85-89. August 15, 1946. 3. The tadpoles of Bufo cognatus Say. By Hobart M. Smith. Pp. 93-96, 1 figure in text. August 15, 1946. 4. Hybridization between two species of garter snakes. By Hobart M. Smith. Pp. 97-100. August 15, 1946. 5. Selected records of reptiles and amphibians from Kansas. By John Breukelman and Hobart M. Smith. Pp. 101-112. August 15, 1946. 6. Kyphosis and other variations in soft-shelled turtles. By Hobart M. Smith. Pp. 117-124, 3 figures in text. July 7, 1947. 7. Natural history of the prairie vole (Mammalian genus Microtus). By E. W. Jameson, Jr. Pp. 125-151, 4 figures in text. October 6, 1947. 8. The postnatal development of two broods of great homed owls (Bubo virginianus ) . By Donald F. Hoffmeister and Henry W. Setzer. Pp. 157-173, 5 figures in text. October 6, 1947. 9. Additions to the list of the birds of Louisiana. By George H. Lowery, Jr. Pp. 177-192. November 7, 1947. 10. A check-list of the birds of Idaho. By M. Dale Ar\ey. Pp. 193- 216. November 29, 1947. 11. Subspeciation in pocket gophers of Kansas. By Bernardo Villa-R. and E. Raymond Hall. Pp. 217-236, 2 figures in text. November 29, 1947. 12. A new bat (genus Myotis) from Mexico. By Walter W. Dalquest and E. Raymond Hall. Pp. 237-244, 6 figures in text. December 10, 1947. 13. Tadarida femorosacca (Merriam) in Tamaulipas, Mexico. By Walter W. Dalquest and E. Raymond Hall. Pp. 245-248, 1 figure in text. December 10, 1947. 14. A new pocket gopher (Thomomys) and a new spiny pocket mouse (Liomys) from Michoacdn, Mexico. By E. Raymond Hall and Bernardo Villa-R. Pp. 249-256, 6 figures in text. July 26, 1948. 15. A new hylid frog from eastern Mexico. By Edward H. Taylor. Pp. 257-264, 1 figure in text. August 16, 1948. (Continued on inside of back cover.) i University of Kansas Publications Museum of Natural History Vol. 4, pp. 1-466, plates 1-41, 31 figures in text December 27, 1951 AMERICAN WEASELS BY E. RAYMOND HALL . cm. Z88L MAR 1 7 1952 iMAR 1 University of Kansas Law^rence 1951 University of Kansas Publications, Museum of Natural History Editors: E. Raymond Hall, Chairman, A. Byron Leonard, Edward H. Taylor, Robert W. Wilson Vol. 4, pp. 1-466, plates 1-41, 31 figures in text December 27, 1951 University of Kansas Lawrence, Kansas PRINTED BY FERD VOILAND. JR.. STATE PRINTER TOPEKA. KANSAS 1951 23-3758 AMERICAN WEASELS (3) Plate 1. Ha*n&r**^ 30 20 10, _ ' 120 T 100 (1 / ■ I — ^. - • .•'OREGON. Tillamook. M.f.altifrontalis. 38AW, Coll. of S. J. Jewett. • CALIFORNIA, Point Arena. M.f.munda, 19722, M V. ^- \ A — 8 ■< (CALIFORNIA, West Riversi( 1. < 7 ' ' ' ■ •ARIZONA Sofford. M.f.neomexicana, 204365, U.S. N^M. ;ide. M. f. la h rostra, 7143", M. V. Z.-^'T— „. r "v >^, .-30 r C \ ^!"«iTAMAULIPAS, Motomoros. M. f. frenata, 1 16552 , U. S. N.M V^; y-- > / r -A^ • MICHOACAN Los Reyes. M.f. leucopana, 125972, U.S.N.M. GUATEMALA, G Sto. Elena. M. f. goldmani, 15953, FM NICARAGUA, fi/latogaipa. M.f.nicaraguae. 28322, A.mTnh'« PANAMA, Mt. Pirre. M.fpanamensis,\7Q970,\J.S.H.f o Q c •-5 1- a < o H Materials, Acknowledgments and Methods 21 this time because they are in breeding condition. Perhaps the ex- planation in part is to be found in the lesser weight of the female (approximately half of the male's weight) which, as indicated above, permits her to step on the pan of a steel trap without spring- ing it whereas the heavier male does spring the trap and as a con- sequence is caught. Hamilton (1933:299-300), who mentions this selective factor, found an equal number of males and females in the three newly born litters that came under his observation. I suppose that in nature there are approximately equal numbers of male and female weasels and further suppose that the selective factors which cause more males tlian females to be caught are the greater distances traveled by the males and their greater weight. MATERIALS, ACKNOWLEDGMENTS AND METHODS At a late stage in the preparation of this manuscript a total of 5,457 specimens had been examined. For the most part these were conventional study-specimens; that is to say, they were stuflFed skins with the skulls separate and each was accompanied by the cus- tomary data as to locality of capture, date of capture, name of col- lector, external measurements and sex recorded on the labels by the collectors. Skulls unaccompanied by skins, nevertheless, com- prised a large share of the total and a small proportion was made up of skins unaccompanied by skulls, mounted specimens, skeletons, and entire animals presei"ved in liquid. It was the recognition of this need for specimens from extensive areas from which no specimens previously had been collected that influenced me, appro.xi- mately a year after the study was begun, to allot for it a long span of time. The procedure adopted, in general, was to study the weasels of one species from a given geographic area in so far as the material warranted, then lay this aside until additional critical material could be obtained, and finally, some months or a year later, complete the account. In this fashion the manuscript of the American weasels received my attention in each of the past twenty-five years (September, 1926 to date of pubhcation). This is a confession of fact rather than a recommendation of procedure. This type of procedure unduly delays the diffusion of knowledge and for a variety of reasons justifiably annoys other students of the subject. Nevertheless, many gaps have been filled that otherwise would have remained open. Although specimens to solve several problems still remain to be collected and studied, it seems that a point of diminishing returns has now been reached, which, in fairness to all concerned, calls for pubhcation of the results so far obtained. For assistance in the entire undertaking, I am more indebted to Miss Annie M. Alexander than to any other one person; she provided the means by which specimens from critical areas were obtained, made it possible to examine the European collections, and assisted in other ways. The late Professor Joseph 22 University of Kansas Publs., Mus. Nat. Hist. Crinnell and Mr. Charles D. Bunker, among others, gave truly valuable en- couragement and assistance. Collections containing weasels which were examined in the study here re- ported upon were as follows: Acad. Nat. Sciences of Philadelphia American Mus. Nat. History Baylor University Berlin Zoological Museum Boston Society of Natural History Brigham Young University British Museum of Natural History California Academy of Sciences Carnegie Museum Charleston Museum Coe College Collection of J. Arnold Collection of Stanley C. Arthur Collection of Rollin H. Baker Collection of William Bebb Collection of R. H. Coleman Collection of Ian McTaggart-Cowan Collection of Stuart Griddle Collection of John Gushing Collection of Walter W. Dalquest Collection of William B. Davis Collection of J. M. Edson Collection of Ralph Ellis Collection of John Fitzgerald, Jr. Collection of Mr. Green Collection of Ross Hardy Collection of Donald V. Hemphill Collection of L. M. Huey Collection of R. W. Jackson Collection of Stanley G. Jewett Collection of E. J. Koestner Collection of J. E. Law Collection of A. H. Miller Collection of Lloye H. Miller Collection of R. D. Moore Collection of J. A. Munro Collection of O. J. Murie Collection of Robert T. Orr Collection of Arthur Peake Collection of Kenneth Racey Collection of William B. Richardson Coll. Rocky Mt. Spotted Fever Lab. Collection of Victor B. Scheffcr Collection of William T. Shaw Collection of O. P. Silliman Collection of W. E. Snyder Collection of Frank Stephens Collection of T. C. Stephens Collection of D. D. Stone Collection of Myron H. Swenk Collection of Joe and Dean Thiriot Collection of John Tyler Collection of Jack C vonBloeker Collection of Alex Walker Collection of Edward R. Warren Colorado Museum of Natural History Charles R. Conner Museum Cornell University Donald R. Dickey Collection Field Museum of Natural History Florida State Museum Fresno State Junior College Humboldt State Teachers College Illinois Natural History Survey Iowa State College Iowa Wesleyan College Kansas State Agric. College Leland Stanford Junior University Leningrad Academy of Science Los Angeles Mus. Hist. Art and Sci. Louisiana State University Mt. Rainier Nat'l Park Collection Museum of Comparative Zoology Mus. Polonais d'Hist. Nat., Warsaw Mus. Vert. Zool., Univ. California Museum of Zoology, Univ. Michigan National Museum of Canada Naturhistorika Ricksmuseum, Sweden Neuchatel University Museum New York State Museum Ohio State Museum Oklalioma Agric. and Mech. College Ottawa University, Kansas Paris Museum Provincial Museum of British Columbia Royal Ontario Museum of Zoology San Diego Society of Natural History State Hist, and Nat. Hist. Soc. Colo. State Normal School, Cheney, Wash. Texas Co-operative Research Collection United States National Museum University of Arkansas Univ. California Mus. Palaeo. University of Id.iho Univ. Kansas Mus. Nat. History University of Minnesota University of Notre Dame University of Oklahoma University of Oregon University of South Dakota University of Utah Univ. Washington Museum of Zoology University of Wisconsin Univ. Zool. Mus., Copenhagen The largest single collection is in the United States National Museum, where the specimens of the National Museum proper and the United States Biological Surveys Collection, together, provide essential materials including Materials, Acknowledgments and Methods 23 a large share of the holotypes. Specimens in all of the North American col- lections including Canada and Mexico have been made available, by loan, and in 1937 materials were examined in the principal collections of northern and central Europe. After the materials in North American collections were as- sembled, special effort, with considerable success, was made in each of several winters, to obtain specimens from areas not previously represented in collec- tions. To the many persons who were in charge of the collections consulted, to those who at my request sought critical specimens, and to those who assisted in various stages of assembling data and in preparation of the manuscript, I am grateful indeed. Likewise, I am deeply appreciative of the grants-in-aid received from the Carnegie Institution of Washington, the University of Cali- fornia Chapter of Sigma Xi, the John Simon Guggenheim Memorial Founda- tion and the Kansas University Endowment Association. I am mindful also of an obligation to those who appropriated funds, by legislative action, for re- search use by The University of California and The University of Kansas. For assistance with the illustrations I am indebted to the late Major Allan Brooks for Plate 1, to Mrs. Mary Bios for figures 25-31, to Miss Ann Murray for figures 11-13, to Mr. W. C. Matthews for all the photographs, to Mrs. Freda L. Abemathy for figures 2-9, 18-22, 24, and for retouching all the photographs except the following which were retouched by Mrs. Virginia Unruh: figs, d of plates 2, 3, 4, 9, 10, 11, 16, 17; figs, i of plates 5, 6, 7; figs, h, j, k of plate 7; figs. / and g of plates 12 and 13; and figs, c and d of plate 14. To Mrs. Unruh I am further indebted for figures 1, 16, 17 and 23 and for much temiinal assistance with preparing most of the illustrations for the engraver. The methods of study, after specimens were assembled, included first comparisons of specimens of like age and sex from each of several localities to ascertain the constant features by which full species were distinguishable, one from the other. For example, it was found tliat in every individual from Trout Lake, Washington, of the species here designated Mustela erminea, the postglenoidal length of the skull amounted to more than 47 per cent of the condylobasal length whereas it was less than 47 per cent in all in- dividuals here designated as Mustela frenata, from the same locality. Testing of specimens from other localities by means of this and other selected characters permitted the outlining of the geographic ranges of the full "species-groups." By comparing specimens of other nominal species and by examining specimens from localities geo- graphically intermediate between the nominal species, I found inter- gradation and therefore arranged the nominal species as subspecies of a single species. Intergradation here is understood to be the re- sult of crossbreeding in nature between two kinds of animals in the area where the geographic ranges of the two kinds meet. Presence of intergradation between two kinds of weasels was basis for ac- cording them subspecific rank. Absence of intergradation in nature 24 University of Kansas Publs., Mus. Nat. Hist. at every place where the geographic ranges of two kinds met or overlapped, and absence of intergradation by way of some other kind, or chain of kinds, was basis for according each of the two kinds full specific rank. By thus applying the test of intergradation, or lack of it, I found that there were four full species of weasels, of the subgenus Miistela, in all of the Americas. Next, the specimens of one species were arranged in trays in a geographic sequence. The specimens from any one locality were segregated by sex and under one sex from one place were arranged from oldest to youngest, that is to say by age. The four series with the largest numbers of individuals of a given age were selected. Seventeen cranial measurements and three external measurements were recorded for each individual of each of these four series. For each measurement, the coefficient of variation, standard deviation and probable error were computed. The four samples subjected to such analysis were a series of adult males, one of adult females, one of subadult males and one of subadult females. Also, studies of each sex were made to ascertain seasonal changes in pelage. After data were obtained on ontogenetic ( age ) variation, secondary sexual variation, seasonal variation, and degree of individual vari- ation by studying specimens in the manner described above, tests were made for subspecific (geographic) variation by comparing series of specimens of like sex, age and season, from different locali- ties. For each one of several geographically variable features noted, a map was prepared for animals of each sex. When all the data thus obtained were codified, subspecific ranges were, in a sense automatically, obtained. On the resulting map showing geographic ranges of subspecies for a species, a type locality was accurately plotted for each name that had been applied to the species, and names then were applied in accordance with the international rules of zoological nomenclature. VARIATION Variation with Age The kind of variation which results from increasing age has been dealt with extensively for the skull (of the Old World Mustela erminea) by Hensel (1881) and for the external features and to some extent for the skull by Hamilton ( 1933) in the North American forms M. erminea cicognanii and M. frenata noveboracensis. The young of both erminea and frenata are hairless and blind at birth. In M. frenata noveboracensis, the eyes open on approxi- mately the 37th day. When 2 to 4 months old, the tail is pointed Variation 25 at the tip. This is because the terminal hair of the tail, including the black tip, is short and lies flat on the tail. In subadults and adults the hair on the terminal part of the tail is as long as that on the basal part, and the tail appears to be of uniform diameter all the way out to the end. In the western subspecies of M. frenata, and in its tropical sub- species, animals so young as to have pointed tails commonly have the underparts of the body more intensely colored than do adults. The young may have salmon-colored instead of yellowish fur on the underparts. Otherwise, in animals that have attained approximately adult proportions — which appears to be at approximately 6 months of age in males — there are no variations which are ascribable to in- creasing age in the color-pattern or pelage that cause the syste- matist to confuse species or subspecies. Of the several parts of the skull in juvenal animals, the braincase and width of the posterior part of the palate are most nearly of the size attained in the adult, the facial part of the skull at birth is the least developed, and the interorbital region is, in relation to its ulti- mate adult size, intermediate in stage of development. The per- manent teeth are acquired when the animal is approximately eleven weeks old. Four age groups, based on characters of the dentition and skull, have been recognized. They are: Juvenile. — One or more deciduous (milk) teeth present. Birth to three months of age. Young. — Sutures widely open between the maxillae and nasals and between the premaxillae and nasals. Three to seven and a half months of age. Subadult. — Sutures between maxillae and nasals visible but indistinct. Seven and a half to ten months of age. Adult. — Bones of rostrum coalesced with no traces of sutures visible to the naked eye. More than ten months old. The skull as a whole increases in size until the animal is two-thirds of the way through the stage designated as young. After this time the width of the rostrum, as measured across the hamular processes of the lacrimals, increases until approximately a third of the way through adulthood. The interorbital breadth decreases from late subadulthood to adulthood and even in adults there appears to be a slight decrease in this part of the skull with increasing age. The average zoologist will readily distinguish skulls of juveniles and young from adults but usually fails to distinguish subadults from adults. Nevertheless, subadults must be distinguished from adults if geographic variation is to be measured accurately. The 26 University of Kansas Publs., Mus. Nat. Hist. reason for this is that such differences in the form (not size) of the skull as result from increasing age equal and often exceed the differ- ences of a geographic sort which serve for distinguishing subspecies that have adjoining geographic ranges. All sutures in the skull, except those between the tympanic bulla and the braincase, and those on the dorsal face of the rostrum, are obliterated while the ani- mal is a subadult. Most kinds of mammals retain sutures throughout life or until the animals are well into adulthood. Therefore, skulls of weasels offer fewer features for estimating age than do those of most mammals and the skulls of weasels that are subadults or older are more difficult to classify accurately as to age than are the skulls of most other mammals. More reliance on shape of entire skull and less reliance on extent and shape of any individual bone is necessary in estimating the age of a weasel. Wright (1947:344) shows that the weight of the baculum (os penis) is a certain means of differ- entiating adults from males of lesser age. When approximately eleven months old, Mustela frenata oribasus of western Montana molts from the white winter coat into the brown summer coat. At that time spermatogenesis starts for the first time and the weight of the baculum increases from less than 30 milligrams to more than 52 milligrams. In the autumn and early winter, most of the specimens are sub- adults. Ordinarily the few adults obtained in these seasons can easily be segregated from the subadults because ontogenetic de- velopment in the twelve additional months of life of each of the older animals has obliterated the sutures on the rostrum, heightened ( vertically ) and lengthened ( anteriorly ) the sagittal crest, widened the rostrum, and produced still other changes in form that are re- vealed by direct comparison of specimens of the two ages. Secondary Sexual Variation The secondary sexual variation, which has been detected, is in size of the animal, relative length of the tail and shape of the skull. The female is the smaller. In the small Mustela rixosa and appar- ently in Mustela africana the secondary sexual difference in size is relatively slight. In Mustela frenata and Mustela erminea, males are approximately twice as heavy as females, the degree of differ- ence very definitely depending upon the subspecies. For example, in M. e. richardsonii the recorded weights are 175 and 69 grams as opposed to 81 and 54 grams in M. e. cicognanii. In general, within one species the greatest difference in size of males and females is in those subspecies in which the animals are of large size. The secondary sexual variation in size is much more than the individual Variation 27 variation in either sex. The same is not true of secondary sexual difference in length of the tail (relative to the length of the head and body), which in eighteen subspecies of M. erminea is from 1 to 7 per cent longer in males than in females. In two subspecies, A/, e. haidarum and M. e. ohjmpica, the tail is a fraction of a per cent the longer in females if we may rely upon the few specimens for which collectors' measurements are available. In both M. erminea and M. frenata the skull of the female is ap- proximately 45 per cent lighter than that of the male, or put in the opposite way, the skull of the male is 83 per cent heavier than the skull of the female. The difference in this respect varies greatly depending on the subspecies. For example, the skull of the male is 127 per cent heavier than that of the female in M. e. richardsonii but only 33 per cent heavier in M. e. anguinae. In Mustela frenata, the subspecies noveboracensis shows most sexual dimorphism in weight of skull (3.6 and 1.7 grams) and olivacea the least (5.3 and 3.8 grams). In general, the difference in this respect is less in sub- species the individuals of which are of small size. Therefore, as might be expected, the secondary sexual variation in weight of the skull is less in M. rixosa, individuals of which are of small size, than in M . erminea or than in M. frenata, in general of larger size. Nevertheless, in M. africana, in which the individuals are of large size, there appears to be less sexual dimorphism in weight of the skull than in M. frenata or than in M. erminea, al- though it should be remarked that there are too few data for M. africana to allow of forming a trustworthy conclusion concerning the amount of secondary sexual variation in that species. The secondary sexual variation in shape of the skull consists of a slenderness in the female. In relation to the basilar length the spread of the zygomatic arches is more in males and, except in the one subspecies M. /. altifrontalis, the rostrum is broader. Also the interorbital region is relatively broader in males of most subspecies. In most subspecies of both M. frenata and M. erminea the tympanic bullae are relatively (to the basilar length) longer in females. The maximum sexual dimorphism occurs in M. erminea arctica and the minimum dimorphism in M. e. haidarum, M. e. anguinae and M. e. muricus. Taking into account all of the subspecies of each of the North American species, the shape of the skull differs most in M. erminea and least in M. frenata. In the latter species the greatest difference in shape of the skull, as was true also of its weight, is in the subspecies M. f. noveboracensis. In these two subspecies, M. /. noveboracensis and M. e. arctica, in addition to the secondary sexual variation already mentioned in the skull, females have the braincase 28 University of Kansas Publs., Mus. Nat. Hist. smoother and more rounded, the postorbital-, mastoid-, and lacrimal- processes relatively smaller, and the ventral face of the tympanic bulla at its anterior margin more nearly flush with the floor of the braincase. In the weasels, subgenus Mustela, the disparity in size of the two sexes is almost or quite as much as in any other fissiped carnivore. It is because of this large degree of difference that the skulls of the two sexes are described separately in the following systematic ac- counts. The need for such treatment was recognized by Reinhold Hensel (1881:127) more than sixty years ago when he wrote in the introduction to his "Craniologische Studien", of Mustela, as follows: ". . . die Geschlechtsdifferenzen am Schadel vieler Sauge- thiere ... so gross sind, dass man diese wie schadel ver- schiedener species behandeln muss, wahrend in anderen Ordnungen (Rosores, Edentaten) die Schadel solche Unterschiede nichtzeigen." In the past, failure to appreciate the large amount of secondary sexual variation has resulted in erroneous deductions as regards characters of certain geographic races and has been the cause of some nomenclatural confusion, as for example, in Mustela frenata macrura, where the female was named as a separate species ( Mus- tela jelskii ) . Individual Variation Individual variation is here considered to be the variation in one species which can occur between offspring of a single pair of par- ents, after variation ascribable to differences in age, sex, and season is excluded. Individual variation, therefore, is a term here used in a composite sense; it includes variations which probably repre- sent different genetic strains within certain populations and varia- tions induced within one generation by environmental factors. In skulls of weasels, the individual variation in size is more than it is in relative proportions. Hensel (op. cit.) has stressed that weasels, like other carnivores, produced "dwarfed" individuals more than do herbivorous mammals. I cannot vouch for the accuracy of this view, but can say that individual variation is not greater than in some other fissiped carnivores. Impressions to the contrary prob- ably result largely from failure to recognize age-variation. When skulls of a large series from any one locality are arranged first h\ sex, and under each sex according to probable age on the basis of extension anteriorly of the sagittal crest and of degree of postorbital constriction, individual variation is seen to be less than a cursory examination, even of only one sex, would suggest. Variation 29 Study of a large series of one age of one sex of one species from one locality shows that some parts, of the skull for example, vary more than other parts. In illustration, among 22 male topotypes of Mustela frenata washingtoni the least interorbital breadth varied 25 per cent (9.0 mm. to 12 mm.) whereas the length of the tooth- rows varied only 13.3 per cent ( 15.6 mm. to 18.0 mm. ). In color the individual variation definitely is more in areas of intergradation be- tween subspecies than in other areas. Details of one such instance of intergradation are given in the account of Mustela frenata spadix. Statements to the effect that there is much individual variation in the color of weasels, were made mostly fifty years or so ago by writers who had but few specimens from widely separated localities. Where marked climatic differences exist between localities only a few miles apart, marked differences occur in coloration of the weasels from the different localities. Much of what formerly was mistaken for individual variation now proves to be geographic variation. Individual variation actually is of slight amount in com- parison with that in mammals generally. Differences in size and relative proportions of parts usually are correlated with geographic differences in color. The color does fade slightly in the period between molts. Also as a result of the seasonal color change, in autumn along the upper margin of the Austral Life-zone, some in- dividuals become white whereas others become white on only the underparts, the upper parts changing only to lighter brown. Prob- ably it would be correct to say that this variation was a combination of seasonal and individual variation rather than either one alone. As might be supposed, individual variation is not the same in all species or subspecies. For example, p2 is always absent in Mustela africana and always present in certain subspecies of M. frenata. In some other subspecies of M. frenata, p2 is absent approximately as often as present. In the writer's experience, when only a few specimens are available for comparison, individual variation is more difficult to distinguish from specific and subspecific (geographic) variation than is age-variation or secondary sexual variation. Among the larger series of specimens examined, only one instance of what might be called a mutation in the old sense of a large, sudden change, was detected. That was the loss of the second lower molar in many (less than a third) of the specimens from Newfoundland. The six instances of abnormal coloration described on pages 41 to 43, might be regarded as mutations of large magni- tude but no evidence was found of repetition of an abnormality in 30 University of Kansas Publs., Mus. Nat, Hist. any one population. Otherwise, in every instance where plotted, the manifestations of a variation arranged themselves about the mean in such a way as to form a smooth, unimodal curve. Seasonal Variation When subspecific and specific variations are the objectives of study, seasonal variation must be understood, in order to be ex- cluded from consideration, in the same way that variations ascrib- able to age, sex and individualism must be understood in order to be excluded from consideration. In weasels, change in color of the pelage is the seasonal variation most important for the systematist to understand. Other seasonal variations in the pelage are hairiness versus nakedness of the pads of the feet, length of the pelage on the body, and possibly the density of the pelage on the body. In the northern half of North America, roughly speaking, seasonal change in color is so pronounced (white in winter and brown in summer) as to be easily recognized. South of this area, in the Austral and Sonoran life-zones, the color of the winter pelage differs only slightly from that of the summer pelage. In these more southern latitudes the winter pelage in almost all subspecies is of lighter color than the summer pelage and has a smoky suffusion. With material of the two seasons in hand for comparison, close attention to the variation will permit the systematist to recognize the difference in shade of brown as seasonal variation and not geographic or specific variation. Farther south still, in the Tropical Life-zone, seasonal difference in color was not detected in the material studied. Seasonal change in color is discussed in the section immediately following. Variation in Coloration, and Molt In all American weasels (subgenus Mustela) the color, at least in summer, is brown with more or less white or whitish on the underparts. In one species, Mustela africana, there is a longi- tudinal stripe of brown on the middle of the light-colored under- parts; this stripe is absent in each of the other three American species. Two species, M. erminea and M. frenata, always have a black tip on the tail. Of the other two species, M. africana lacks the black tip and M. rixosa may or may not have a few black hairs in the tip of its tail. White or light yellowish facial mark- ings occur in subspecies of M. frenata from the southwestern United States to Central America. Subspecies having the most extensive light-colored facial markings have the remainder of the upper part of the head black. In weasels without light facial mark- Variation 31 ings the upper parts of the head all are brown. In the two species, M. erminea and M. frenata, the extent to which the light color of the underparts extends down the insides of the legs and out on the underside of the tail, or the absence of light color on these parts, is a matter of geographic variation. The same can be said for M. rixosa except that first its tail is unicolored and second in- dividual variation as well as geographic variation accounts for the color pattern on the underparts and legs in animals from the southeastern part of the range of the species. The most remarkable feature of the coloration of weasels is the winter whitening. This occurs in the northern part of North America in each of the three species of weasels found on that continent. The black tip of the tail in M. erminea and M. frenata remains black in winter. If an individual of M. rixosa has black hairs on the tip of its tail in summer, there are thought to be black hairs there also in winter. Otherwise the winter pelage is all white in northern areas in each of the three species. In this white winter coat the animal is known as ermine. The underlying cause seems to be protective coloration. At any rate, weasels are always white in winter if they are from areas where snow lies on the ground all winter, every winter, or al- most every winter; and they are always brown if from areas where there is never, or rarely, snow in winter. The changes in color are effected by molt, one in autumn and one in spring. Animals that are brown in winter undergo the same two molts as do those that are white in winter. The capacity to acquire a white coat or a brown coat in winter is an hereditary matter just as one man grows red hair and another grows black hair. In the weasels, however, all individuals in the north turn white in winter and if one that was born there is kept through successive winters in the warmer south where there is no snow, he will still turn white each winter. A weasel born in a southern area, where all are brown in winter, molts into a brown (not white) winter coat even when kept in a cold, snowy, northern area where native weasels of the same species all turn white. Obviously, therefore, neither snow nor temperature is an immediate cause and, as we have said, the color in winter is a matter of heredity. The time of the molt, we now know, is determined by the amount of light. When nights grow longer and days shorter, a point is reached at which the lesser light received through the eyes causes the pituitary gland to cease producing a gonadotropic hormone. Directly or in- 32 Univei^ity of Kansas Publs., Mus. Nat. Hist. directly, the lack of this hormone stimulates molt and, probably enzyme action, or the lack of it, causes the melanoblasts of the cells in the hair follicle to be without pigment. Hence the hair grown from a follicle under such conditions lacks pigment (mel- anin) and is white. In spring, as the days grow longer and the nights shorter, the increasing amount of light received day by day through the eyes stimulates the pituitary gland to produce the gonadotropic hormone which directly or indirectly, stimulates molt and, probably by enzyme action, the melanoblasts are caused to be present in cells of the hair follicle and the melanoblasts pro- vide granules of melanin pigment which are incorporated in cells of the growing hair. These granules of pigment give the hair its color. Evidence in support of this hypothesis is given below. Along the Pacific Coast from British Columbia southward, M. erminea ( see fig. 25 on page 95 ) is brown in winter. This is an area where snow rarely falls and the temperature in winter ordinarily is above freezing. In the remaining part of the American range of this species the temperature in winter is below freezing much of the time and snow remains throughout the winter or for long periods. In this colder part of the animal's range, only white coats occur in winter. M. frenata likewise has a white coat in winter in the part of its geographic range where snow and freezing tem- peratm-es prevail throughout most of the winter and a brown coat in warmer, snowless areas to the southward and along the Pacific Coast. The third species, M, rixosa, exhibits a corresponding cor- relation between coat color and climate. On the Asiatic conti- nent, several species, including M. erminea, provide parallel correlations and nowhere are there any exceptions for the sub- genus Mustela. These data are an important part of the material on which we have based the induction that the underlying cause of seasonal change in color is a need for protective coloration. As regards molt, most naturalists who have written upon the sub- ject regard it as responsible for the change from the white winter coat to the brown summer coat. However, the change from brown summer coat to white winter coat has been thought by several writers to be effected by change in coloration of the individual hairs. Among those holding this opinion there may be cited Bell (1874: 197) in reference to Mustela erminea, and Coues (1877:123) in reference to American specimens to which he applied the same name. More lately Hadwen (1929) has taken this same view, and Gunn (1932) also discusses the possibility of the hairs changing Variation 33 color. Bachman (1839:228-232), MacgiUivary (1843?:158), Au- dubon and Bachman (1851 (vol. 2):62), Schwalbe (1893:538), Pearson et at. (1913:447), Miller (1930, 1931A), Hamilton (1933: 300) and Rothschild (1942), among others, have been inchned to the opinion, or positively affirm, that the color change in autumn is the result of a molt. The papers cited above contain, in turn, references to many other printed accounts deahng with this ques- tion. To my mind, it has not so far been demonstrated tliat the change in color of weasels in autumn is accomplished without a molt. Also so far as I am aware, no explanation has been given of how the pigment may disappear from the hair of weasels. Metchni- koflf's (1901:156) idea that the senile whitening of the hair in man is accomplished by phagocytes which remove the pigment granules would hardly seem to explain the relatively sudden and complete autumnal change occurring in weasels. Anyhow, Danforth (1925: 108 ) , and some other students have thought that the action of these phagocytes was at most a factor of sHght importance in the whiten- ing of hair. Whatever be the complete answer to the question of how the weasel changes color in autumn, at least one specimen of long-tailed weasel, which is in process of color change in autumn, presents clear evidence of molt of the overhairs. This specimen of M. f. longicauda is no. 188408, U. S. Nat. Mus., taken on November 12, 1897, at Rapid City, South Dakota. Other specimens of M. erminea which were taken in autumn similarly show molt to be in progress. For these and other reasons, I am inclined to the opinion that die autumnal change in color, like the one in spring, is effected by molt. During the period of the autumnal color change, Noback (1935:27) had a captive M. /. noveboracensis and, each morning, found clumps of brown hair on the floor of its cage; tliis was strong indication that molt was responsible for the color change in this instance. However, I freely admit that the evidence does not prove that the change from brown to white can be accomplished onhj by molt; in the present state of knowledge it would be unscientific to deny that the change were possible of accomplishment by other means. Also, it is true that the fifteen specimens before me of Mustela frenata, subspecies included, in process of change from brown to white, with the exception of the one from Raf)id City, South Dakota, if taken individually, do not, in macroscopic exami- nation, show definite molt lines or other absolutely convincing evi- dence of molt. However, these same specimens, insofar as examined 2—3758 34 University of Kansas Publs., Mus. Nat. Hist. microscopically, do show overhairs all white, or overhairs pig- mented throughout. The lighter color of the proximal parts of the overhairs in itself should not be accepted as evidence of color change, for in the fresh summer pelage, the same condition exists. Also, careful macroscopic examination suffices to show that in the transitional pelage of autumn, the brown overhairs generally are longer than the intermixed white overhairs. Whether the underfur behaves in exactly the same way as the overhair, I have not myself definitely ascertained, but I assume that the underfur is molted twice each year, at least in the northern populations of Mtistela frenata and in the other species of more northern distribution. Schwalbe's (1893) work, including section- ing of the skin and study of the hair follicles, led him to conclude that the underfur was molted twice each year in Mtistela erminea. In Mustela frenata noveboracensis, M. f. nevadensis, and M. f. nigriatiris, measurements taken on adult males show the overhairs to be longer in the winter pelage than in the summer pelage of speci- mens from the same locality. For example, in M. /. nigriatiris irony Berkeley, California, the overhairs of the summer coat (July and August) average 8 millimeters in length on the hinder back and 7 mm. on the belly, but average 9.5 mm. and 8 mm. respectively in January-taken specimens possessing the full winter coat. At Ann Arbor, Michigan, in the summer coat, the longest hairs on the hinder back average approximately 12 mm., and those on the belly, 9.5 mm., against 13 mm. and 9.5 mm. respectively in winter. Al- though general observations initially led me to believe that the black, terminal hairs of the tip of the tail are longer in the winter pelage than in the summer pelage, actual measurements fail to show a difference in length. The change from one coat to the other in the long-tailed weasel has been described among others by Miller (1930, 1931A), Hamil- ton (1933) and Glover (1942) on the basis of captive specimens. In a general way, the progress of the molt in their specimens agrees with that which I have been able to make out from examination of skins taken in the wild. There is, however, this difference: Their specimens show a more spotted pattern when in process of hair- change than do specimens taken in the wild. Probably the more or less unnatural conditions under which these captive animals lived modified the normal progress of molt. In wild-taken specimens of the species Mustela frenata, sub- species included, the spring molt begins on the middorsal line and proceeds laterally, producing, at almost any given time, a relatively Variation 35 sharp molt line separating the white winter hair from the incoming brown summer coat. However, in autumn the change takes place first on the belly, then on the sides, and finally makes its appearance over all the upper parts at about the same time, with the result that the upper parts have a salt-and-pepper appearance without at this time any sharply defined molt Hnes. In general, the molt pattern can be said to be reversed in the two seasons; in spring, it begins on the back and in autumn, on the belly. The difiFerence in spring and autumn color pattern is better illustrated on plate 39 than by additional description. Swanson and Fryklund (1935:123) have observed that the "spring molt proceeds differently" than the fall one in Mustela rixosa, and Barrett-Hamilton (1903:309) in com- menting on the European hare (and the stoat?) remarks, "In spring the moult, and with it the brown colour, progresses in exactly the opposite order . . ." as compared with the white color of au- tumn, which that particular writer thought resulted from removal of pigment from the hairs rather than from molt. The tail, excepting the black tip, lags in the molt in many in- stances, with the result that, especially in spring, it may retain a few white hairs as late as does the belly. In autumn it is less tardy and so far as I have observed, becomes white at about the same time that the general area of the back changes color. On the tail, the black tip itself, as clearly shown in more than a score of specimens, is molted at approximately the same time in autumn as is the pelage of the body. However, the long black hairs, which appear in, say, November, appear to increase in length until January. In spring, the long black hairs of the tip of the tail seem not to be shed at the same time as the rest of the winter pelage, but remain approxi- mately six weeks longer and then are replaced by long black hairs of the summer coat. At any rate, this is the picture presented by a half dozen specimens of M. /. nevadensis and M. f. longicauda which do show a spring molt to be in progress on the black tip of the tail. Schwalbe similarly (1893:536-537) has suggested that the black tip of the tail in Mustela erminea in spring is not molted until about two months after the pelage on the rest of the body is changed. Schwalbe {he. eit.) thinks also that in M. erminea studied by him, the black tip of the tail in autumn is replaced ap- proximately one month in advance of the pelage on the rest of the body. As indicated above, my specimens of Mustela frenata, sub- species longicauda and nevadensis, do not show this discrepancy in autumn. I have considered the possibility that the black tip of the tail, in some species of Mustela, is molted only once while the re- 36 University of Kansas Publs., Mus. Nat. Hist. mainder of the coat was undergoing two molts. My inconclusive data lend but little support to this possibility. The difFerence in pattern of color between specimens taken in autumn and spring is known to some fur-trappers of my acquaint- ance who have suggested that molt occurs in spring, whereas the individual hairs change color in autumn. Reference to plate 39 wall show how gross comparisons might lead one to this erroneous ex- planation of the color change. As to time of molt: In eight subspecies of Mustela frenata, namely, noveboracensis, occisor, primulina, spadix, longicauda, ari- zonensis, nevadensis and effera, material is available to indicate that the autumnal molt begins in October and is completed in Novem- ber, and that the spring molt occurs in March or April. A con- densed list of specimens providing basis for this statement is as follows : M. f. noveboracensis: 26 specimens in transitional pelage taken in auttunn and 14 taken in spring; M. /. occisor: One topotype has acquired one-fifth of the winter pelage on October 22, 1896; M. /. primulina: 2 in November, one in March, and 2 in April are in process of change; M. /. spadix: 6 autumnal specimens and one in April show pelage change; M. /. longicauda: 7 autumnal specimens and one in April show pelage change; M. f. arizonensis: 12 speci- mens in autumn and 3 in spring are in process of molt; M. /. effera: One No- vember-taken male has acquired four-fifths of the winter coat and another taken on April 21 at Fort Rock, Oregon, is half finished with the spring molt. It may be added that no marked difference in time of either au- tumnal or spring molt is apparent as between the more northern and more southern localities from which the mentioned specimens come. With more complete material I would expect to find a difference in this regard. The material of the other, more southern, subspecies of Mustela frenata has not been adequate to show the time of molting or the number of molts which occur in one year. Animals in the northern part of the range of Mustela frenata acquire a white winter coat, whereas those in the southern part acquire a brown winter coat, and in an intervening area the winter coat may be either browoi or white. By plotting on a map the locaH- ties of capture of all specimens examined in the winter coat, it was possible to outline this intervening area as shown in figure 10 on page 37. However, Dearborn (1932:36) shows that in Michigan some animals have a brown coat in winter at places farther north than figure 10 shows to be the case. Hamilton's (1933-306) map for New York shows the same to be true in that state. Accordingly, the boundaries of the area shown in figure 10, in which both brown Variation 37 ec M Fig. 10. Map showing the region (in black) where both the brown and white winter pelage is found in the long-tailed weasel, Mustela frenata. and white long-tailed weasels occur in winter, are known to be only approximate; with full information available the belt would be represented as wider. Hamilton (1933:302) has pointed out that "Where half of the weasels remain brown, these brown winter specimens are always 38 University of Kansas Publs., Mus. Nat. Hist. males." The results of my own examination of specimens not studied by Hamilton, in a general way provide confirmatory data. More exactly, my examination reveals that at the most northern localities where brown specimens occur, only males are in this coat. In explanation, it may be said that in plotting on a map locali- ties of capture of specimens in the winter coat, thirteen places were found where both sexes were represented and where both brown and white winter coats were found. With the two sexes, it is theoretically possible to have nine diflFerent combinations of coat color. With males all brown, there might occur females (1) all brown, (2) all white, or (3) some brown and some white. In ad- dition to these three combinations, we might have three more by finding the mentioned types of female coat color repeated where all males are white, and three more, or nine in all, by substituting a population of males some of which were brown and some of which were white. Seven of these possible combinations actually were found. The two combinations not found were all white males with all brown females, and all white males with females both brown and white. In the three instances where the males all were brown and the females all were white, the localities of capture were in the northern part of the variable area. This indicates that where the brown winter coat occurs at northern localities, the brown indi- viduals are all males. Farther south, of course, the females, too, acquire the brown winter coat. Stated in another way, there is a broad belt across North Amer- ica from the Atlantic to the Pacific in which males of Mustela frenata at any one locality may be either brown or white in winter. Inside this broad belt there is a narrower one, approximately half as wdde, in which females at any one locality may be either brown or white. In support of the idea that color of the winter coat is an hereditary matter and that it is not dependent on temperature, the following evidence derived from my transplanting specimens of Mustela frenata supports the idea that color of the winter pelage is de- pendent on heredity and not on temperature or snowfall. A male captured on June 24, 1937, in the brown summer coat in Salt Lake City, Utah, was received by me at Berkeley, California, five days later and kept in captivity almost six months. On No- vember 17, 1937, half the pelage was white and on December 27, 1937, when next examined, the animal was in the full, white, winter coat as it was on January 25, 1938, when it died. Native weasels all turn white in winter in Salt Lake City, but in Berkeley native weasels always are brown in winter. Variation 39 A juvenile or young animal, a male, captured in May, 1936, at Lafayette, Contra Costa County, California, was kept there until August 13, 1936, when transferred to Calneva at the north end of Lake Tahoe, California. The weasel was kept at Calneva until its death on December 23, 1937. In both the winter of 1936-'37 and in that of 1937-'38, the winter coat was brown as in animals from its place of origin ( Contra Costa County ) and unlike weasels of the Tahoe region nearly all of which turn white in winter. Two females, each approximately two months old, captured on May 1, 1936, at James Landing, 4 miles northwest of San Pablo, Contra Costa County, California, were kept in Berkeley, California, until August 13, 1936, when they were transferred to the mouth of Blackwood Creek, on tlie west side of Lake Tahoe, California. On October 25, 1936, both weasels escaped. On December 25, 1936, the headless body of one of these was found approximately 300 yards soutli of the mouth of Blackwood Creek. The animal had been dead at most a few days when found and was in the brown winter coat. At the place of its origin all weasels are brown in wanter but at the mouth of Blackwood Creek only 2 of 60 weasels caught there in the winter coat were brown; the other 58 were white. The headless weasel was identified, as one of the two formerly in captivity, by means of certain short toes, the ends of which had been clipped off when the animal was a captive. No trace of the second female was found. A female of unknown age, in white winter pelage, captured 4 miles southeast of Tahoe City, California, and kept there until April 3, 1937, on which date it was brought to Berkeley, California, molted to brown in the spring. The first signs of the brown coat were noted on April 14. On May 24 or 25 she gave birth to 4 young which lived less than ten days. In the following winter this animal acquired a white coat. As previously noted, weasels native to the Berkeley area, where this female was kept, have brown coats in winter. The weasels were in every instance kept in cages out-of-doors. The sides of the cages were open to the elements. A nest box in each cage provided shelter. All were of the species Mustela frenata. The significant results, it seemed to me, were that the winter coat was the kind found in the area where the weasel originated instead of the kind found in weasels native to the areas in which the specimens were held in captivity. 40 University of Kansas Publs., Mus. Nat. Hist. That the time of molt is determined by the amount of light has clearly been shown by Bissonnette (1944:223) for American weasels of the two species Mustela erminea and M. frenata. In his words {op. cit.:2A6) "Reducing the daily periods of light induced molting and regrowth of new fur. ... In the Bonaparte weasels [Mus- tela erminea], white replaced brown. . . . Increasing daily light-periods caused molting and change to dark brown. . . . Incomplete molts in both directions ( toward white or toward brown ) were produced as a result of early reversal of increase or decrease of daily light-time. . . . That this stimulus is received through the eyes and acts through the anterior pituitary gland is indicated by Bissonnette's [1935:159] studies on ferrets, a nearly related animal. That the thyroids and sex-glands are not essential is at least suggested ... by Lyman's (1942) study on the varying hare [Lepus americanus]." It can be added that Lyman (1943: 451) demonstrated in Lepus americanus that the effect of light is received through the eyes. He demonstrated this by masking the animals. To Wright (1942B:109) who studied the two American weasels, M. erminea and M. frenata, it seemed likely that the pitui- tary produced or released gonadotropic hormone at about the time of the spring molt and that this molt and the spring changes in the reproductive tracts of the weasels might be caused by a stimulus from a common source. Later, Wright (1950:130) in- jected a gonadotropic hormone into long-tailed weasels which had recently acquired their white winter pelage and thereby caused them to lose the white pelage and acquire the brown pelage. It is Lyman (1943:450) who says, in relation to Lepus americanus, "When in the physiologically white condition, the melanoblasts of the regenerating guard- and pile-hair folUcles con- tain no melanin-forming enzyme (dopaoxidase), which may be the reason for the lack of pigment." Schwalbe (1893) by sectioning the skin and microscopically examining the hair-follicles of M. erminea learned that the basal cells producing hairs lacked pigment granules in autumn when the European ermine (M. erminea) was acquiring its white winter coat and that tlie cells contained granules of pigment in spring when, as we know, the granules are incorpo- rated in the growing hair and give it its color. The above material, then, is basis for the account on pages 31 and 32 of what causes the weasel of northern areas to have a white coat in winter. The discerning student will instantly perceive that although some parts of the account on pages 31 and 32 are pre- cisely accurate, other parts are the result of inferences which need Variation 41 to be proved. More careful work of the kind that Schwalbe (1893) and Wright ( 1942B ) did is needed. The account on pages 31 and 32 is merely the best that can be given with the information now available. Many writers have commented on the yellowish color, sometimes with a greenish tinge, found on the fur of weasels in the white winter coat. The stain is more often found on the tail and hinder- parts of the body than elsewhere. Possibly, partly on this account, some have ascribed tliis color to the smearing of the fur with urine. Still others have thought it resulted from the smearing of the fur with secretions from the anal scent glands. Schumacher ( 1928 ) takes this point of view, and while it may be that he has not proved his point, still his conclusions fit the known facts and seem sound to me. Schumacher points out that the same soiling of the fur is present in summer as well as in winter, but that on the summer pelage the stain can be detected only on the Light-colored under- parts. It is from this point of view that he criticizes the systematic worth of white versus yellowish-white underparts in the summer pelage of geographic races of Miistela erminea and Mustela nivalis. Although in the long-tailed weasels (Mustela frenata) the under- parts of all the races are pigmented with some form of red, orange or yellow, it seems probable to me that the additional color resulting from the soiling effect of this glandular secretion explains the greater variation, found at a single locality, in the color of underparts than of upper parts in the summer pelage. I have neither seen nor heard of a black weasel in any part of the New World or of the Old World. I have found only one albino among American specimens. It is an adult female, no. 121424, American Museum of Natural History, of Mustela erminea richard- sonii, taken on August 30, 1935, at Hot Springs, Northwest Territory. This place, I am told by G. G. Goodwin who obtained the animal, is on the "Nahanni River where the rugged mountain ridges rise abruptly from the low mud flat lands, latitude 61, longitude 125." The shortness and coarseness of the hair corresponds to that of the summer pelage and not winter pelage. The pelage is everywhere white, even the tip of the tail. True, all except the nape and top and sides of the head has a faint yellowish-green tinge which has been supposed to result from staining by secretion of the anal scent glands but there is no pigment in the hair as in erythristic specimens. From the Old World, Farurick (1873:17) has recorded what he re- gards as an albino of Mustela vulgaris since it had no black hairs on the tip of its tail. Flintoff (1935:228, 229) records what may 42 University of Kansas Publs., Mus. Nat. Hist. have been an albino Mustela vulgaris from Yorkshire and an albino M. crminea from an unstated locality. Jackel (1873:459) mentions specimens of Mustela erininea and Mustela vulgaris, which were partly "albinistic" or "erythristic." Among the American specimens of M. erminea I have not recorded any which appeared to be either partly or wholly erytliristic or only partly albinistic. Among the 1550 skins of M. frenata which were in summer pelage or browoa winter pelage, five, described below, show marked abnormalities in color. Two of these five are partly albinistic. One is an adult male, no. 223880, U. S. Nat. Mus., from Billy's Island, Okefinokee Swamp, Georgia, which has the nose as well as the area between the eyes white. Also there is a tuft of white hairs at the anterodorsal margin of each ear, scattering white hairs suggesting a postorbital bar on each side of the head, and a patch of white hairs on the middorsal line behind the ears. Markings of this kind are not abnormal in M. f. periinsulae, the subspecies adjoining on the south, except for the white nose which clearly is an instance of partial albinism. The second specimen is a subadult male, of M. /. noveboracensis, no. 177679, U. S. Nat. Mus., in process of acquiring the brown winter coat, taken on November 27, 1911, at Gaylordsville, Connecticut. It has white markings on the nose, on the right side of the neck, on the right hind foot and right forefoot, and on the tip of the tail. The white area of the nose on the left side extends back to the eye, but on the right side barely encircles the nose-pad. On the right side of the neck, all that area between the foreleg and ear is white from the mid-dorsal line (including 7 or 8 millimeters to the left of the mid-dorsal line) down to the throat, which is white as it is also in normal individuals. The toes of the right hind foot are more extensively white than in normal specimens of noveboracensis, and all of the right forefoot as well as the wrist is white. The tail is of striking appearance because of its tricolor pattern. The proxi- mal part is of the normal brown color. The black terminal part commences proximally at the usual place, but the distal 11 milli- meters of the fleshy part of the tail bear only pure white hairs pro- ducing a terminal white pencil 35 millimeters long. The three otlier specimens abnormally colored are erythristic individuals. An adult male of M. /. latirostra, no. 7574, coll. D. R. Dickey, taken on April 14, 1918, at Covina, Los Angeles County, California, has the color of the upper parts greatly restricted, and, in addition, has spots and blotches of the color of the underparts distributed over the back and rump. A spot of this same color Variation 43 occurs above each ear. Incidentally, this and other subspecies of Mustela jrenata from the Pacific Coast of North America obviously have the factor for erythrism operating over a larger part of the body than it does in M. erminea or than in M. f. noveboracensis, where the underparts sometimes are white. In M. f. latirostra and in other subspecies from the Pacific Coast the light color of the underparts always is tinged with this reddish color. Another erythristic specimen is a young male of M. f. nevadensis, no. 23493, U. S. Nat. Mus., taken on August 6, 1890, at Birch Creek, Idaho. It has all of each foreleg, the axillary regions, and a saddle- shaped area over the shoulders of the same buff-yellow color as the underparts. The third erythristic specimen is a subadult female, of M. f. ore- gonensis, no. 47149, Mus. Vert. Zool., taken on December 20, 1930, at Carlotta, Humboldt County, California. This specimen appears to be white and initially was thought to be merely an individual in the white winter coat. Closer examination, however, shows that it has a light wash of ochraceous or faint reddish color. Also, other specimens taken in winter at Carlotta show that weasels there do not acquire a white winter coat. The only normally brown area is approximately three millimeters in diameter at the anterodorsal margin of the pinna of the right ear. The tip of the tail is black as in a normal specimen. The specimen in question is actually pure white only on top of the head from a short distance behind the ears on over the forehead nearly to the eyes, and on the inside of the ears. In a normally colored animal this area is the dark area of the head. In this freak, the other parts of the head, which, in indi- viduals of normal coloration are the white or light orange facial markings, have the reddish cast of the remainder of the body, al- though the color is less intense than on the back. The collector noted that the specimen had eyes of normal color. A possible explanation for the coloration of this specimen is that this species has three factors for color, one for the black tail tip, one for the reddish color, and a third, missing in the specimen in question, for the blackish brown. For some more exact knowledge concerning this erythristic type of coloration, we are indebted to Pitt (1921:99), who describes a pop- ulation of polecats, Mustela putoritis, in Cardiganshire, England, in which this erythristic variation is maintained in a state of nature. In ferrets, Mustela furo, Pitt {op. cit.-.WA) notes that ", . . ery- thrism is certainly dependent on a Mendelian factor, being dom- inant to albinism and recessive to the black-brown coloration. Both 44 University of Kansas Publs., Mus. Nat. Hist, in the ferret and polecat, erythrism seems to be correlated with in- creased size, and certainly in the ferret is usually accompanied by a quick temper and general increase in vitaHty." Variations of Taxonomic Worth Variations of taxonomic worth usually are referred to as charac- ters. For example, shortness of the tympanic bulla is a character, and the opposite condition, long tympanic bulla, is another char- acter. Specific variations, that is to say specific characters, are pro- vided by the color-pattern, length of tail, number of premolar teeth, shape of the tympanic bullae, and length of the braincase in rela- tion to the length of the tooth-bearing parts of the skull. Subspecific characters are provided by color-pattern, color itself, size as meas- ured by weight of the animal, and its hnear measurements, size of the skull, and size and shape of parts of the skull. The characters distinguishing subspecies from one another are not of a different nature from those distinguishing species from one another. Given any one of the above structural features, say, dorsal outline of the skull, several characters may be provided by it. For example, weasels of tlie species Mustela frenata have the dorsal outline of the skull convex in southern Louisiana, straight in Missouri and concave in North Dakota, thus providing three characters. This is geo- graphic variation. These variations, characters in zoological par- lance, when plotted on maps, reveal the geographic occurrence of, say, the convex shape of the skull. Tn combination with other characters, for example, dark color and short tail, basis is provided for recognizing a subspecies, in this instance Mustela frenata arthuri of Louisiana. Because the change from convex to flat skull takes place geographically at about the same place (in eastern Texas) as does the change from short tail to long tail, and the change from dark color to light color, it is easy to draw a line there marking the western geographic limit of occurrence of the M. f. arthuri. This same line marks also the eastern margin of the geographic range of the subspecies Mustela frenata frenata, the subspecies next ad- jacent to the westward. On this line and for several miles to either side of it weasels show varying combinations of these three char- acters or an intermediate condition as regards one or more of the characters, or both. For example, from a locality in eastern Texas a weasel may have (1) a facial pattern exactly intermediate be- tween that of the unicolored face of arthuri and that of the bi- colored face of frenata, (2) the long tail of frenata and (3) the con- vex skull of arthuri. In the sum of its characters this specimen is Variation 45 exactly intermediate between typical arthtiri and typical frenata. Another specimen from the same place may differ from the first specimen only in having the tail slightly shorter. The total "score" for the two specimens is, therefore, by a very slight margin in favor of arthuri. Let us suppose that we obtain a third specimen from the same place and that it has the face marked like that of artliuri but the tail fully as long, and the skull as lacking in dorsal convexity, as in frenata. Now tlie score is definitely for frenata. For convenience of handling, the population is referred to frenata, providing that the average of specimens from a nearby locality to the westward is not in favor of arthuri. In event the average of specimens from a lo- cality next adjacent to the westward is in favor of M. f. arthuri, the total evidence from the two localities may be weighed together and appropriate decision as to subspecific status of weasels from the area is made according to what tlie average is for the area as a whole. The three individual animals of an intermediate sort are ordi- narily termed intergrades. This implies that their characters are the result of mixed parentage — perhaps a female of M. /. arthuri and a male of M. /. frenata but probably each parent itself was an intergrade and the offspring, of which we examined three, owe their characters to reproductive processes operating in obedience to Mendelian laws of inheritance. The two kinds of animals, Mustela frenata arthuri and Mustela frenata frenata, are identified as subspecies because of the inter- gradation between them. If at this and all other places where the geographic ranges of arthuri and frenata met there was no crossbreeding (no intergrades), the two kinds would be treated as distinct species. Intergradation, and the lack of it, are accepted as the criteria of subspecies and species, respectively. These criteria suffice for animals, in this instance weasels, which have a continuous geographic distribution. Some kinds of weasels are confined to islands, as for example the islands off the coast of Alaska and British Columbia. Because weasels are land animals, crossbreeding in nature between the weasels of two islands is, of course, impossible. A modified test (used in the study here reported upon) in deciding on specific versus subspecific status in these instances can be made as follows: On tlie adjacent main- land, ascertain the degree of difference between two subspecies whose geographic ranges meet (for example, M. e. richardsonii and M. e. alascensis). Next ascertain the degree of difference between the insular kind of animal and the kind on the mainland. If the 46 University of Kansas Publs., Mus. Nat. Hist. degree of difference is greater when the insular kind is compared than when only the kinds of the mainland are compared, tlie insular kind is to be regarded as a species. If the degree of difference is no greater between the insular kind and the mainland kind than it is between the two adjacent mainland kinds, the insular kind is to be regarded as a subspecies. In short, for insular kinds, the cri- terion is degree of difference, with the limitation of geographic adjaceny, rather than intergradation. The geographic variation ( subspecific characters ) found could be spoken of as two kinds: First, there is the variation which is ex- pressed in a general trend for a long distance, producing, in general, a cline of even slope; and second, that of inconstant trend in any one direction. In his "The Rabbits of North America" Nelson (1909:34-35) has commented on the latter type of variation as fol- lows: "While studying series of specimens from all parts of the vast range occupied by the geographic races of such species as Sylvilagus -floridanus and S. auduboni, I have been impressed with evidences of fluctuation of both external and skull characters. These fluctu- ations are somewhat wavelike in character and rise to central points of extreme development and tlien sink away to intermediate borders beyond which new waves rise. Where the waves of differentiation are pronounced they mark recognizable geographic races. Within the area covered by the larger or geographically broader waves of differentiation (recognized as of subspecific value), smaller waves of differentiation are included, which may represent local variations in intensity of characters of the subspecies, or these characters may diminish and the variation tend in other directions, sometimes even closely reproducing the characters of another subspecies occupying a distinct area." In Mustela frenata, much of the geographic varia- tion at first inspection appears to be of this nature. Closer scrutiny, however, reveals that the repetition, at geographic intervals, of several features of color and structure are closely correlated with environmental features which are repeated only at these same places. In Mustela erminea, much of the variation is of the first kind, namely, that which can be expressed as long clines of relatively even slope. As several authors have said, zoological classification based on this kind of variation is like dividing the spectrum and depends largely upon the standards set, for, theoretically, the possibilities of subdivision are unlimited. Actually, however, none of the clines has an even slope and the possibilities for subdivision therefore are limited. Also, when several features are used, instead of only Variation 47 one feature, the classification is more satisfactory even if the basis is more complex. Some features of structure which provide subspecific characters are mentioned below. Total length, of males, ranges from 598 to 360 mm. in M. frenata and from 336 to 228 mm. in M. erminca. There is no cline of sus- tained slope in M. frenata but in M. erminea there is a progressive decrease in total length from north to south. Length of tail varies from as little as a half to as much as seven- tenths of the length of the head and body in M. frenata, the sub- species neomexicana having the long tail and the two subspecies arthiiri and primuUna having short tails. The geographic ranges of primttlina and neomexicana are contiguous. In M. erminea there is likewise no variation of a clinal nature in length of tail and fur- thermore the variation is much less than in M. frenata. In length of hind foot, which in males varies from 49 mm. in northern populations of M. erminea to 28 mm. in southern popula- tions, the same cline is seen as in the total length of animals of this species. In M. frenata, however, there are several decreases and increases along any straight line which can be drawn through the geographic range of the species. The range of variation in males is 41 mm. (M. /. arizonensis) to 59 mm. (M. /. macrophonius) . Weight of the entire animal is an excellent measure of size but weights are unavailable for many subspecies. In M. frenata, the two subspecies texensis and macrophonius probably are the heaviest and effera, arizonensis and helleri probably are the lightest. Geo- graphically the variation in weight behaves in approximately the same way as does the measurement of total length. In M. erminea the variation in weight of males is from 206 grams in northern animals to 58 grams in southernmost populations, there being a relatively constant gradient geographically. Degree of hairiness of tlie foot-soles in M. frenata clearly is linked with the temperature; in regions of high average temperature the hairiness is least and in regions of low average temperature it is most. The decrease in hairiness is accomplished in two ways, namely, smaller breadth and decreased length of individual hairs and decrease in number of hairs on a given area of dermal surface. This correlation holds throughout the entire north to south range of the species. Corresponding differences are found on the same latitude where topographic diversity in an east to west direction produces northern conditions at high altitudes and southern condi- tions at low altitudes. The conclusion seems unavoidable that 48 University of Kansas Publs., Mus. Nat. Hist. 12 13 Figs. 11-15. Dorsal views of adult skulls of each sex of five subspecies of the ermine, Mustela erminea, to show secondary sexual variation and geographic variation in size of the skull. Males on the left and females on the right. AU XI. Note especially the geographic variation in decreasing size of the skull from north to south in each sex, and that the secondary sexual variation in size of skull is less in ermines with small skulls than in those with large skulls. Variation 49 140 130 120 110 100 90 80 f Me richardsonii, Ft Franklin, Mackenzie. d",i33847, A M N h. \ \ • - \ %^ ^. '^ "^ ^ M e. richardsonii^ Ft Resolution, Mackenzie. ?, 129703, U.SIM.VI ! / i ' I ip M. e. invicfa, Jasper Park, Al be rtad',3593,?, 10797, CN / / / -^ 14 / ( ■-- I ! vj§ M e gulosa, Trout Lake, Washington. t?',82i77, ?,77370,uSNivi I / / :/ I \ 15' T 1--. ^ ^ M e muricus. Baker Creek, Nevada. Tionym for the weasel of the lower Amazon area. 1898. neomexicanus (Putorius frenatus) Barber and Cockerell [= Mustela frenata neomexicana], Proc. Acad. Nat. Sci. Philadelphia, p. 188, May 3, 1898, apphes to the long-tailed weasel of New Mexico, Arizona, Durango and adjoin- ing areas. 1898. haidarum (Putorius) Preble [= Mustela erminea haidarum], Proc, Biol. Soc. Washington, 12:169, August 10, 1898, apphes to the ermine of the Queen Charlotte Islands, British Columbia. History of Classification 77 1899. notius {Putorius noveboracensis) Bangs [=Mustela frenata nove- boracensis], Proc. New England Zool. Club, 1:53, June 9, 1899, was applied to tlie long-tailed weasel of the Carolinas until 1936 since wliich time it has been regarded as a synonym of noveboracensis. 1899. occisor (Putorius) Bangs [=Mustela frenata occisor], Proc, New England Zool. Club, 1:54, June 9, 1899, applies to the long-tailed weasel of central and northern Maine. Until 1936, occisor was ordinarily used as the name of a full species but since then has been arranged as a subspecific name under Mustela frenata. 1899. mundus (Putorius xanthogenijs) Bangs [= Mustela frenata munda], Proc. New England Zool. Club, 1:56, June 9, 1899, is now applied, and gener- ally has been since 1899, to the long-tailed weasel of the coastal district of Cali- fornia north of San Francisco Bay. 1899. muricus (Putorius (Arctogale)) Bangs [= Mustela erminea muricus], Proc. New England Zool. Club, 1:71, July 31, 1899, applies to the dimunitive ermine, often erroneously designated least weasel, of the western United States. 1899. oribasus (Putorius (Arctogale) hmgicauda) Bangs [= Mustela frenata oribasus], Proc. New England Zool. Club, 1:81, December 27, 1899, applies to the long-tailed weasel of the Rocky Moimtains northward from Yellowstone National Park. 1900. eskimo (Putorius rixosus) Stone [= Mustela rixosa eskimo], Proc. Acad. Nat. Sci. Philadelphia, 1900:44, March 24, 1900, is appUed to the least weasel of Alaska and adjacent parts of boreal North America. 1901. allegheniensis (Putorius) Rhoads [= Mustela rixosa allegheniensis], Proc. Acad. Nat. Sci. Philadelphia, 1900:75, March 25, 1901, appUes to the least weasel of the eastern United States. 1902. perdus (Putorius tropicalis) Merriam [= Mustela frenata perda], Proc. Biol. Soc. Washington, 15:67, March 22, 1902, appHes to the long-tailed weasel of the Lower Tropical Life-zone from southern Veracruz into Guatemala. 1903. microtis (Putorius) Allen [—Mustela erminea richardsonii]. Bull. Amer. Mus. Nat. Hist., 19:563, October 10, 1903, is a name applied to an individual ermine of small size from Shesley, British Columbia, which Allen thought was specifically distinct from the ermine of the Hudsonian Life-zone and adjacent territory. Now the name is arranged as a synonym of richardsonii. 1904. audax (Putorius) Barrett-Hamilton [= Mustela erminea arctica], Ann. and Mag. Nat. Hist., ser. 7, 13:392, May, 1904. In the original description the type locality. Discovery Bay, was erroneously stated to be in Greenland and the name audax until 1945 was applied to the kind of weasel occurring in northern Greenland whereas tlie type specimen was taken instead in northern EUesmere Island and because the weasel there is subspecifically indistinguish- able from ermines from farther west, audax is a synonym of Putorius arcticus. 1904. imperii (Putorius arcticus) Barrett-Hamilton [= Mustela erminea richardsonii], Ann. and Mag. Nat. Hist., ser. 7, 13:392, May, 1904, based on an animal from Fort Simpson, Mackenzie, Canada, proves to be inseparable from richardsonii which has priority. 1904. polaris (Putorius arcticus) Barrett-Hamilton [= Mustela erminea polaris], Ann. and Mag. Nat. Hist., ser. 7, 13:393, May, 1904, is the name used 78 University of Kansas Publs., Mus. Nat. Hist. for the ermine of eastern Greenland and since 1945 has been used for the weasel of Greenland as a whole. 1905. macrophonius (Putorius) Elliott [= Mustela frenata macrophonius], Proc. Biol. Soc. Washington, 18:235, December 9, 1905, applies to the long- tailed weasel of the mountains along the eastern border of Veracruz. 1906. leptus (Putorius streatori) Merriam [= Mustela erminea murica], Proc. Biol. Soc. Washington, 16:76, May 29, 1903, until 1945 was applied to the diminutive ermine of the Rocky Mountains from Wyoming south to northern New Mexico but proves to be a synonym of muricus with type locality in the Sierra Nevada of California. 1908. angustidens (Putorius cicognanii) Brown [=z Mustela erminea angus- tidens], Mem. Amer. Mus. Nat. Hist., 9(pt. 4):181, pi. 17, is applied to an extinct subspecies known from fossil remains of Pleistocene age from northern Arkansas. 1908. gracilis (Putorius) Brown [= MusfeZa frenata gracilis], Mem. Amer. Mus. Nat. Hist., 9(pt. 4): 182, 1908, apphes to a Pleistocene weasel known from a single skull from northern Arkansas. 1912. costaricensis (Mustela) Goldman [^Mustela frenata costaricensis], Proc. Biol. Soc. Washington, 25:9, January 23, 1912, apphes to the long-tailed weasel of Costa Rica. 1913. primulina (Mustela) Jackson [^Mustela frenata primulina], Proc. Biol. Soc. Washington, 26:123, May 21, 1913, appUes to the long-tailed weasel of the central part of the United States in eastern Kansas and adjoining areas. 1913. campestris (Mustela) Jackson [= Mustela rixosa campestris], Proc. Biol. Soc. Washington, 26:124, May 21, 1913, applies to the least weasel of the Great Plains region. 1913. olivacea (Mustela peninsulae) HoweU [:= Mustela frenata olivacea], Proc. Biol. Soc. Washington, 26:139, May 21, 1913, applies to the long-tailed weasel of the southeastern United States excepting most of Florida. 1914. meridana (Mustela) HoUister [= Mustela frenata 7neridana], Proc. Biol. Soc. Washington, 27:143, July 10, 1914, applies to the long-tailed weasel of northern South America. 1916. nicaraguae (Mustela tropicalis) Allen [= Mustela frenata nicarguae]. Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916, appUes to the long-tailed weasel of Nicaragua. 1927. arthuri (Mustela noveboracensis) Hall [z= Mustela frenata arthuri], Proc. Biol. Soc. Wasliington, 40:193, December 2, 1927, applies to the long- tailed weasel of Louisiana and adjoining areas. 1932. semplei (Mustela arctica) Sutton and Hamilton [= Mustela erminea semplei], Ann. Carnegie Mus., 21(2) :79, February 13, 1932, originally was appUed to the ermine of Southampton Island but after 1945 was appHed also to the ermine of Baffin Island, Melville Peninsula and the west side of Hudsons Bay as far south as Eskimo Point. 1932. panamensis (Mustela frenata) Hall, Proc. Biol. Soc. Washington, 45:139, September 9, 1932, applies to the long-tailed weasel of Panamd. History of Classification 79 1932. anguinae {Mustela cicognanii) Hall [^Mustela erminea anguinae], Univ. California Publ. Zool., 38:417, November 8, 1932, applies to the ermine of Vancouver Island, British Columbia. 1935. labiata (Mustela arctica) Degerb0l [^Mustela erminea semplei]. Kept. 5th Thule Exped., 1921-1924, vol. 2, no. 4, p. 25, 1935. When Degerb0l wrote his description and proposed this name he was unaware that Sutton and Hamilton had three years before based a new name on weasels from Southampton Island. Because the two names apply to the same subspecies, Degerb0rs name, labiata, must fall as a synonym of semplei which has priority. 1935. helleri (Mustela frenata) Hall, Proc. Biol. Soc. Washington, 48:143, August 22, 1935, applies to the long-tailed weasel of eastern Peru. 1936. nevadensis (Mustela frenata) Hall, Carnegie Inst. Washington, publ. no. 473, p. 91, November 20, 1945, applies to the long-tailed weasel of the western United States. For many years, animals of this subspecies were re- ferred to longicauda and from 1891 until 1936 to arizonensis. 1936. effera (Mustela frenata) Hall, Carnegie Inst. Washington, publ. no. 473, p. 93, November 20, 1945, apphes to the long-tailed weasel of the Blue Mountains region. From 1891 until 1936 this animal was referred to under the name arizonensis. 1936. altifrontalis ( Mustela frenata ) Hall, Carnegie Inst. Washington, publ. no. 473, p. 94, November 20, 1936, applies to the long-tailed weasel of the humid coastal district from Puget Sound southward into Oregon. 1936. nigriauris (Mustela frenata) Hall, Carnegie Inst. Washington, pubL no. 473, p. 95, November 20, 1936, applies to the long-tailed weasel of the coastal district of California from San Francisco Bay southward to Point Con- cepcion. Previous to 1936, xanthogenys was the name applied to this race of weasel. 1936. latirostra (Mustela frenata) Hall, Carnegie Inst. Washington, pubL no. 473, p. 96, November 20, 1936, applies to the long-tailed weasel of southern California which previously had borne the name xanthogenys. 1936. pulchra (Mustela frenata) Hall, Carnegie Inst. Washington, publ. no. 473, p. 98, November 20, 1936, is applied to the long-tailed weasel of the southern end of the San Joaquin Valley of California. 1936. inyoensis (Mustela frenata) Hall, Carnegie Inst. Washington, publ. no. 473, p. 99, November 20, 1936, is applied to the long-tailed weasel of Owens Valley, California. 1936. texensis (Mustela frenata) Hall, Carnegie Inst. Washington, publ. no. 473, p. 99, November 20, 1936, applies to the long-tailed weasel of central Texas which previously had been assigned to the subspecies frenata. 1936. perotae (Mustela frenata) Hall, Carnegie Inst. Washington, publ. no. 473, p. 100, November 20, 1936, applies to long-tailed weasel of the mountains along the Puebla-Mexico boundary. 1938. boliviensis (Mustela frenata) Hall, Proc. Biol. Soc. Washington, 51:67, May 18, 1938, applies to the southernmost known long-tailed weasel which is in the Lake Titicaca region in Peru and Bolivia. 80 University of Kansas Publs., Mus. Nat. Hist. 1944. salva (Mustela erminea) Hall, Proc. Biol. Soc. Washington, 57:35, June 28, 1944, applies to the ermine of Admiralty Island, southeastern Alaska. 1944. initis {Mustela erminea) Hall, Proc. Biol. Soc. Washington, 57:37, June 28, 1944, applies to the ermine of Baranof and Chichagof islands, south- eastern Alaska. 1944. celenda (Mustela erminea) Hall, Proc. Biol. Soc. Washington, 57:38, June 28, 1944, applies to the ermine of Prince of Wales, Dall and Long is- lands, Alaska. 1944. seclusa (Mustela erminea) Hall, Proc. Biol. Soc. Washington, 57:39, June 28, 1944, applies to the ermine of Suemez Island, southeastern Alaska. 1945. invicta (Mustela erminea) Hall, Jour. Mamm., 26:75, February 27, 1945, applies to the ermine of the Rocky Mountains for several hundred miles both north and south of the United States-Canadian boundary. 1945. fallenda (Mustela erminea) Hall, Jour. Mamm., 26:79, February 27, 1945, appUes to the ermine of the coastal mainland in southern British Columbia and northern Washington. 1945. olympica (Mustela erminea) Hall, Jour. Mamm., 26:81, February 27, 1945, applies to the diminutive ermine of the Olympic Peninsula, state of Washington. 1945. gulosa (Mustela erminea) Hall, ]oui. Mamm., 26:84, February 27, 1945, applies to the diminutive ennine of the Cascades in Washington. 1945. bangsi (Mustela erminea) Hall, Jour. Mamm., 26:176, July 19, 1945, is the name apphed today to the ermine of the western Great Lakes region. In 1925 when this study was begun, the American weasels (subgenus Mus- tela proper) were arranged as belonging to 47 kinds (including subspecies) of 29 full species. In the present account a total of 68 kinds, belonging to 4 full species are recognized in the subgenus Mustela. The increase in nxmiber of subspecies and the decrease in number of species are the nomenclatural results ordinarily obtained in this decade from a systematic study of a genus of American mammals. MUSTELA 81 CHECK-LIST OF AMERICAN SPECIES AND SUBSPECIES OF THE GENUS MUSTELA Subgenus MUSTELA Linnaeus PAGE Mustela erminea 87 Mustela erminea arctica (Merriam) 96 Mustela erminea polaris (Barrett-Hamilton) 103 Mustela erminea semplei Sutton and Hamilton 105 Mustela erminea kadiacensis ( Merriam ) 108 Mustela erminea richardsonii Bonaparte 110 Mustela erminea cicognanii Bonaparte 118 Mustela erminea hangsi Hall 124 Mustela erminea invicta Hall .- 128 Mustela erminea alascensis ( Merriam) 131 Mustela erminea salva Hall 135 Mustela erminea initis Hall 136 Mustela erminea celenda Hall 139 Mustela erminea seclusa Hall 141 Mustela erminea haidarum (Preble) 142 Mustela erminea anguinae Hall 145 Mustela erminea fallenda Hall 148 Mustela erminea olympica Hall 153 Mustela erminea streatori ( Merriam) 155 Mustela erminea gulosa Hall 159 Mustela erminea muricus ( Bangs) 161 MuMela erminea angustidens ( Brown) 165 Mustela rixosa 168 Mustela rixosa eskimo Stone 181 Mustela rixosa rixosa Bangs 184 Mustela rixosa allegheniensis Rhoads 187 Mustela rixosa campestris Jackson 190 Mustela frenata 193 Mustela frenata noveboracensis (Emmons) 222 Mustela frenata occisor ( Bangs ) 230 Mustela frenata primulina ( Jackson ) 232 Mustela frenata arthuri Hall 241 Mustela frenata olivacea Howell 244 Mustela frenata peninsulae Rhoads 250 Mustela frenata spadix ( Bangs ) 252 Mustela frenata longicauda Bonaparte 262 Mustela frenata oribasus ( Bangs ) 270 Mustela frenata alleni ( Merriam ) 274 Mustela frenata arizonensis ( Meams) 276 Mustela frenata nevadensis Hall 280 Mustela frenata effera Hall 291 Mustela frenata washingtoni (Merriam) 294 Mustela frenata saturata ( Merriam ) 297 Mustela frenata altifrontalis Hall 300 Mustela frenata oregonensis (Merriam) 304 Mustela frenata munda ( Bangs ) 3C9 82 University of Kansas Publs., Mus. Nat. Hist. PACE Mustela frenata xanthogenys Gray 315 Mustela frenata nigriauris Hall 319 Mustela frenata latirostra Hall 323 Mustela frenata pulchra Hall 328 Mustela frenata inyoensis Hall 331 Mustela frenata neomexicana (Barber and Cockerell) 333 Mustela frenata texensis Hall 338 Mustela frenata frenata Lichtenstein 341 Mustela frenata leucoparia ( Merriam ) 347 Mustela frenata perotae Hall 351 Mustela frenata goldmani ( Merriam ) 355 Mustela frenata macrophonius ( Elliot ) 360 Mustela frenata tropicalis ( Merriam ) 363 Mustela frenata perda ( Merriam) 366 Mustela frenata nicaraguae Allen 370 Mustela frenata costaricensis Goldman 372 Mustela frenata panamensis Hall 375 Mustela frenata meridana Hollister 379 Mustela frenata affinis Gray 384 Mustela frenata aureoventris Gray 387 Mustela frenata helleri Hall 391 Mustela frenata agilis Tschudi 393 Mustela frenata macrura Taczanowski 398 Mustela frenata holiviensis Hall 402 Mustela frenata gracilis ( Brown ) 404 Subgenus Gramnogale Cabrera Mustela africana 406 Mustela africana africana Desmarest 409 Mustela africana stolzmanni Taczanowski 413 Subgenus Putorius Cuvier ( Black-footed Ferret — not treated in present work ) Mustela nigripes ( Audubon and Bachman ) Subgenus Lutreola Wagner (Minks — not treated in present work) Mustela vison Mustela vison vison Schreber Mustela vison mink Pcale and Beauvois Mustela vison lutensis (Bangs) Mustela vison evergladensis Hamilton Mustela vison vulgivaga ( Bangs ) Mustela vison letifera Hollister Mustela vison lacustris (Preble) Mustela vison energumenos (Bangs) Mustela vison evagor Hall Mustela vison aestuarina Grinnell Mustela vison nesolestes (Heller) Mustela vison melampelus (Elliot) Mustela vison ingens (Osgood) Mustela macrodon (Prentiss) iJ MUSTELA 83 ARTIFICIAL KEY TO AMERICAN SPECIES OF THE GENUS MUSTELA PAGB A Length of upper tooth-rows less than 20 mm. in males and 17.8 mm, in females. B Postglenoid length of skull more than 47 per cent of condylo- basal length. C Tail without a black pencil and with at most a few black hairs at extreme tip; in both sexes mastoid breadth or- dinarily exceeds breadth of braincase, Mustela rixosa, least weasel, p. 168 C Tail with a black pencil; in females mastoid breadth or- dinarily exceeded by breadth of braincase, Mustela erminea, ermine, p. 87 B' Postglenoid length of skull less than 47 per cent of condylobasal length. D Tail with distinct black tip; midventral line white, yellowish, orange, not same color as upper parts; p2 present; thenar pad on forefoot absent, Mustela frenata, long-tailed weasel, p. 193 D' Tail without black tip; midventral line same color as upper parts; p2 absent; thenar pad on forefoot present, Mustela africana, tropical weasel, p. 406 A' Length of upper tooth-rows more than 20 mm. in males and 17.8 mm. in females. E Abdomen all white; face with blackish mask; ml lacking even a trace of a metaconid; distance between upper canines more than width of basioccipital as measured between foramina sit- uated midway along medial sides of tympanic bullae, Mustela nigripes, black-footed ferret. E' Abdomen dark brov^Ti, like back; face uniformly brown without blackish mask; ml with incipient metaconid; distance between upper canines less than width of basioccipital as measured between foramina situated midway along medial sides of tym- panic bullae, Mustela vison, mink, American mink. DIAGNOSIS OF THE GENUS Genus Mustela Linnaeus Weasels, Ferrets, Polecats, Minks Genotype. — Mustela erminea Linnaeus. Diagnosis. — Legs short; body relatively long; adults 190 mm. to 700 mm. in total length; skull ranging in basilar length from 16 to 70 mm.; facial angle slight; tympanic bullae greatly inflated (moder- ately in Lutreola), cancellous, and with paroccipital processes closely appressed to bullae; palate behind upper molars; dental r , I 3 C 1 P 2-3 M 1 . . r xt-i 1 i formula: —'—'—'—'—' -rr'' ~' ~Z' mner moiety of Ml larger than i3clp3-2m2 ^ ° outer; P4 with simple deuterocone; in ml trigonid longer than 84 University of Kansas Publs., Mus. Nat. Hist. talonid, metaconid absent (incipiently developed in Lutreola), and talonid trenchant. For many years prior to 1911, the name Mustela was appHed to martens, and Putoritis was regarded as the first available generic name for the weasels. In 1911 Thomas (1911:139) showed that M. erminea ( Mustela of Gesner ) by tautonymy was the type of Mustela and subsequently the generic name Mustela has been used for the true weasels which include the American weasels to which we now apply the specific names erminea, rixosa and frenata. The mink, Mustela (Lutreola) vison, and the black-footed ferret, Mustela (Putorius) nigripes, since 1911 also have been referred by most American authors to the genus Mustela, the names Lutreola and Putorius being regarded by these authors as having no more than subgeneric status. European writers, on the other hand, accord greater taxonomic weight to the zoological differences between fer- rets and weasels and, therefore, accord full generic rank to Putorius. Consequently, for the black-footed ferret, Europeans today write Putorius nigripes and Americans write Mustela nigripes. For the same reasons, the name of the mink is written by some European zoologists Lutreola vison and by American zoologists Mustela vison. EXPLANATION OF SYSTEMATIC TREATMENT For each full species there will be found under the account of it the follow- ing information: Type, statement of geographic range, selected characters for ready recognition, other characters of the species, a summary of geographic variation, and information on habits, in the order mentioned. For each subspecies, information is presented in the following order: earliest available zoological name, synonyms, type, geographic range, zoological char- acters for ready recognition, description (mentioning size, certain external features including color, the skull and teeth) historical material when war- ranted, remarks which may elaborate on points made in preceding paragraphs, and other information thought to be useful, and finally a Ust of specimens e.xamined. In explanation of certain of these categories it should be said that in the synonymy no attempt is made to hst every published reference to the subspecies concerned. It is aimed, however, to include at least one citation to each name- combination that has been applied, to the subspecies concerned, along with other especially important references. Mere records of occurrence are not re- garded as especially important and citations to them ordinarily are omitted in the synonymy. No comma is placed between the zoological name and the name of the author who coined and first used the name in accordance with the rules of zoological nomenclature. Otherwise a comma is interposed between the zoological name and the name of tlie user (author). When the accepted (earliest available) name of a subspecies at the head of any one of the follow- ing accounts is combined with a generic name different from that with which MUSTELA 85 it originally was placed, the authority for the name is set in parentheses. The same rule is followed with the name of a full species when it is written without any subspecific name following. Parentheses in such situations, therefore, de- note that for the terminal part of the scientific name there has been a change in generic name with which the terminal part of the scientific name is here associated. In the paragraph headed "characters for ready recognition", only a few characters, namely, those regarded as most useful for identification when the student has limited time, are mentioned. Other features useful for distinguish- ing the kind of animal in question from its near relatives are to be found in the description and comparisons. In the description, external measurements, unless otherwise indicated, are those recorded by tlie collector on the label attached to the skin. Total length is the distance from the tip of the pad on the nose to the tip of the fleshy part of the tail when the relaxed animal is laid out straight, not stretched. This measurement does not include the hairs that project beyond the end of the fleshy part of the tail. Length of tail is the distance from the base of the tail, when it is bent at right angles to the long axis of the body, to the tip of tlie fleshy part of the tail excluding the hairs that project beyond the fleshy part of the tail. Length of tail and length of tail-vertebrae are synonymous. Length of hind foot is measured from the proximal end of the calcaneum to the tip of the longest claw. Capitalized color terms, unless otherwise indicated, refer to Ridgway's (1912) Color Standards and Color Nomenclature. Some use is made of color terms taken from Oberthiir and Dauthenay (1905) because those authors show a much larger number of shades between dark brown and black than does Ridgway (1912). The colors of the upper parts of most weasels are some shade or other of dark brown. Color terms that do not have the initial letter capitalized do not refer to any one standard and consequently are used in a general sense. Relative extents of the color of the upper parts and underparts are computed from measurements of the circumference of the body at the place where the color of the underparts is narrowest. Ordinarily this place is in the lumbar region rather than in the thoracic region. An explanation of how cranial measurements were taken is given on page 417. In designating teeth, capital letters are used for teeth in the upper jaw and lower case letters are used for teeth in the lower jaw. For example: 12 denotes the second incisor tooth in the upper jaw and i2 denotes the second in- cisor tooth in the lower jaw; CI and cl refer to the canine tooth of the upper jaw and lower jaw, respectively; P3 and p3 refer to the third premolar of the upper jaw and lower jaw, respectively, bearing in mind that the first (anterior) premolar is absent in the lower jaw and upper jaw of weasels (see fig. 31 on page 416), as also, in some kinds of weasels, is the second premolar; Ml and ml refer to the first molar of the upper jaw and lower jaw respectively. In describing the skull and teeth the two sexes are treated separately because difi^erences in shape as well as size are the rule. Unless otherwise indicated, the skulls on which descriptions are based are of adults. Weights of skulls include the weight of the lower jaws. In general, every second sub- species is described. For a subspecies geographically next adjacent to the 86 University of Kansas Publs., Mus. Nat. Hist. one described, only the diflFerences between the two are enumerated. Fhi* method of description indicates also likenesses and is more economical of words than some other methods of description. Also, by use of this method, cross reference is reduced to one other subspecies. Following this formal descrip- tion, there is a comparison of the cranial and dental characters with those of geographically adjacent subspecies. In the paragraph headed "Remarks" the two words "character" and "structure" frequently appear. The word structure here is used to mean some part of an animal, as for example, a hair, a muscle, a bone, or an internal organ. A structure is not a system, as for example, the digestive system or osseus system. A character is some weight, linear dimension, volume, shape, color, or other perceptible attribute of a structure, of a system, or of an entire organism. In recording the localities of capture of specimens examined, effort has been made to be exactly as precise as the locality data on the labels of the specimens permit. The word "County" is written out in full when the name of the county is written on the label of each specimen listed from that county. When one specimen, or more, here assigned to a given county lacks the name of the county on the label, then the abbreviation "Co." is used. The sur- prising frequency with which the same place name is repeated in a given state or province makes it desirable for the collector to write the name of the county, or corresponding minor political subdivision, on labels of study speci- mens at the time they are prepared. MUSTELA ERMINEA 87 SYSTEMATIC ACCOUNTS OF SPECIES AND SUBSPECIES MUSTELA ERMINEA Linnaeus Emiine ( Synonymy under subspecies ) Type. — Mustela erminea Linnaeus, Systema Naturae, 10th ed., p. 46, 1758. Range. — From tlie British Isles and Atlantic Coast of Europe across Eiurasia and North America including Greenland, from the northernmost land, south, in North America, to the lower margin of the Canadian Life-zone; geographi- cally south to Connecticut, New York, Pennsylvania, northeastern Ohio, south- em Michigan, Wisconsin, northern Iowa, Minnesota, North Dakota, in the Rocky Mountains to northern New Mexico, in the Sierra Nevada to Mono County, Cahfomia, and on the Pacific Coast to the Golden Gate. Characters for ready recognition. — Differs from Mustela rixosa in presence of black pencil on tail, tail-vertebrae more than a fourth of length of head and body, and in regions where the two species occur togetlier, basilar length of skull more than 32.5 in males and more than 31.0 in females; from Mustela frenata, in regions where the two species occur together, by tail less than 44 per cent of length of head and body and by postglenoidal length of skull more than 46 per cent of condylobasal length in males and more than 48 per cent in females. Characters of the species. — Size medium to small (total length 225 to 340 mm. in males and 190 to 290 mm. in females); tail 30 to 45 per cent of length of head and body, with distinct black pencil; caudal vertebrae 16 to 19; skull with long braincase and short precranial portion; postglenoidal length, when expressed as a percentage of the condylobasal length, more than 48 in females and ordinarily more than 46 in males; upper parts brown; under- parts whitish, ordinarily continuous from chin to inguinal region but in subspecies in the humid region along the Pacific Coast inter- rupted in some individuals by brown of upper parts encircling body in the abdominal region. The soles of the feet in each of the sub- species are densely haired in winter and have only a relatively small area of the foot-pads exposed in summer, the intervening areas being well haired even at that season. The uniformity throughout the species as regards hairiness of the foot-soles and also the char- acter of the vibrissae makes it unnecessary to describe these features in the accounts of the subspecies of erminea. Geographic variation. — In the Old World 16 or more subspecies are currently recognized and there are 20 in North America. The features in which geographic variation is especially prominent are: 88 UNivERSiTi' OF Kansas Publs., Mus. Nat. Hist. First, size, as expressed by external measurements and weight, second, color pattern, depending on the extent, in relation to one another, of the dark-colored upper parts and light-colored under- parts, and third, breadth and depth of the rostral region of the skull. Except in size, the variation in the skull is less than in M. frenata. Likewise in tone and shade of upper parts and hue or tint of underparts, erminea is less variable than frenata and has the face all of one color without the contrasting color-pattern of the face and head seen in many subspecies of frenata. M. erminea exceeds frenata as regards variation of the size of the area occupied by tlie light-colored underparts. At one extreme is the subspecies arctica in which the area of the light color extends well up on the sides of the body, down the insides of the legs, over the feet and far out on tlie lower side of the tail whereas at the other extreme are the races streatori and olympica in which the light-colored underparts are restricted to two areas, one on the chin, throat and chest, and the other on the inguinal region. These areas may or may not be connected by a thin line of white color along the midline of the underparts. In size of animal, erminea probably exhibits the maxi- mum variation among American species of weasels; an average- sized male of the race arctica weighs 4 times as much as one of the race muricus, and in the species frenata I doubt that the difference is quite as great between individuals of the smallest race, effera, on tlie one hand, and either of the largest races, texensis or macro- phonius, on the other hand although actual weights are not avail- able for these races of frenata. As elsewhere indicated, the small- sized individuals of M. erminea are of the southern races and the large-sized individuals are of the northern races. This decrease in size southward occurs both in Asia and in America. Natural history — habitat and numbers. — Along the International Boundary east of the Turtle Mountains, Soper (1946:136) found this species present only in timbered areas and absent from many untimbered areas. Of the same species to the westward he com- ments "so far as I know at present, there is no evidence to show that any short-tailed weasels inhabit a broad strip of treeless terri- tory immediately north of the International Boundary in Canada from southwestern Alberta to southeastern Saskatchewan". The same author (1942) reports that in the general area of Wood Buf- falo Park, Northwest Territory, south of Great Slave Lake, the ermine is uncommon on pine-grown sand ridge and rolling upland and common in lower spruce-aspen parklands, stream-side conifer- ous belts, and grassy, semi-wooded swamplands. MUSTELA ERMINEA 89 Nine ermines per square mile is the number that Soper (1919:46- 47) estimated at Edmonton on the basis of the numbers that he trapped there in the winters of 1912-13 and 1913-14 and on the basis of the tracks of remaining ermines. From corresponding data he estimated the population in the winter of 1913 on the Hay River, north of Jasper Park, to be nine per square mile. In each of these instances he estimated ten weasels per square mile but he inclined to the view that one-tenth of the animals involved in his counts were long-tailed weasels (Miistela frenata). Osgood (1909B:30) and his field companion in the period July 31 to September 3, 1903, took a series of 42 specimens within a radius of 500 yards of their camp at the head of Seward Creek, Alaska, all caught in four traps, in one month. Of the 42 specimens, 28 are males and 14 are fe- males. Fluctuations of a multiannual nature are marked in this species. Bailey (1929:156) observes that in Sherburne County, Minnesota, when meadow mice are abundant for two or three years these weasels become abundant but that when the mice are scarce the weasels also become scarce. Manning (1943:56), on Southampton Island, noted "that the maximum and minimum points of the weasel cycle are much more sharply marked than those of the fox cycle and the increase and decrease are more rapid." How far an ermine will travel in a given length of time has sel- dom been recorded but Hamilton (1933:293), on March 20, 1932, "followed the track of a small weasel, presumably a male cicognanii, for four miles in the fresh snow", and Ingles (1942) observed a diminutive ermine of the subspecies M. e. mnricus, at Woods Lake, California, 286 yards from its den. Behavior As regards locomotion, Soper (1919:46), in reference to Mustela cicognanii, presumably in Ontario, Canada, writes that in the bounding gait the hind feet register almost, if not exactly, in the front-foot impressions, with the right front and hind feet lagging shghtly behind. "The distance normally is about 19 inches, repre- senting a regular rate of travel. ... In traversing open spaces they resort to long, graceful leaps upwards of six feet in length. . . . I measured a record ... of 8 feet, 2 inches." Of M. e. arctica. Dice (1921:22) writes that when it runs "the tail is carried off the ground usually at an angle of about 45 degrees." Seton (1929 (2):598) states that "At Carberry [Manitoba] I have often seen this energetic little creature seeking for Mice in the deep. 90 University of Kansas Publs., Mus. Nat. Hist. soft snow. Its actions are much like those of an Otter pursuing sal- mon. Sometimes it gallops along a log, or over an icy part of the drift; then plunges out of sight in a soft place, to reappear many yards away. . . ." Little is recorded concerning swimming but on this score Seton (1929 (2) :602) does quote J. W. Curran, who in July, 1899, at Lake Couchiching, Ontario, watched an ermine pursue a chipmunk into the water and for 100 yards before giving up the chase and wheeling around and making for shore. In swimming "The Weasel, I think, showed more of his body, and seemed to exert himself more" than the chipmunk. As to voice. Dice (1921:22), at Tanana, Alaska, heard the ermine, when excited, bark somewhat like a mink but not so loud and Seton (1929 (2):606) quotes Manley Hardy to the effect that the species has a purring note. Sense of smell was used by an M. e. muricus that Dixon (1931:72) watched as the ermine followed a three-fourths-grown pika. Con- cerning the ermine at Carberry, Manitoba, Seton (1929 (2):598- 599) writes that "The smell of blood must be as far-reaching as it is attractive to these sanguinary little creatures. I have frequently hung new-killed Rabbits and partridges temporarily in trees, and, after an absence, in some cases of a few minutes only, have found an Ermine mauling the game, though there was no sign of such a visitor when the cache was made." Enemies George Measham, of Winnipeg, found sign in the snow indicating that a great snowy owl had killed an ermine and T. Mcllwraith shot a bald eagle at Hamilton Bay which had the bleached skull of a weasel ( probably of this species ) clinging to the tliroat ( Seton, 1929 (2):603). A. B. Howell (1943:98) hkens mustehd mammals to domestic cats in their manner of crossing roads and thinks that mustelids loiter at the side of the road until the stimulus of the approaching car causes them to make a dash whereupon they are caught by the wheels and killed. Three of four weasels seen to cross the road were killed, one even having apparently crossed the road before turning back and being killed under the car. One weasel killed was Mustela erminea cicognanii. Dalquest (1948:190) in writing of this species in the state of Washington, says "I have seen only one abroad in the daytime. It dashed from a roadside thicket . . . and was crushed beneath the wheels of a car." MUSTELA ERMINEA 91 Food The killing of prey is described by Hamilton (1933:332) as fol- lows: "A rapid dash, and the bird or mouse is grabbed over the back of the skull, the fore legs encircle the animal as though hugging it, and the hind legs are brought up to scratch wildly at the captive. . . . If [the prey is] a large animal, as a rat, the weasel usually Hes on its side, while the diminishing struggles of the rodent con- tinue, but if a mouse or a small bird [is the object of attack], the weasel is apt to crouch over its prey. Little time is lost over the first [mouse] ... if two mice are present [;] a strong bite through the brain case . . . [is] sufficient. If only one animal is present, the weasel dawdles over its kill some time after life has departed." Hamilton's (1933:333) study of the contents of the digestive tracts of bodies of ermines obtained from fur trappers and fur buy- ers in New York enabled him to publish the following "Frequency Indices of Mammal Genera in Fall and Winter Food of 191 Mustela cicognanii": Microtus, 35.7 per cent; mammals undetermined to genus but principally mice, 16.3; Blarina, 15.1; Peromyscus, 11.4; Sylvilagus, 9.0; Sorex, 4.9; Rattus, 4.4; Tamtas, 3.6. Close corre- spondence is shown by the following data of Aldous and Manweiler (1942) for the ermine from Lake of the Woods, Minnesota: mice, 58.7 per cent by number and 54.5 by volume; shrews 22.5 and 21.8 per cent; birds, 2.7 and 5.0 per cent. Of the mice in stomachs, 40 per cent were microtines, 15 per cent were Peromyscus and 45 per cent were unidentified as to kind. Fragments of a small fish were found in one stomach. Summed up, the dominant winter foods were mice and shrews. Trapping of the mammal populations was done to see what the available food was and it was found that the small mammals were eaten in direct ratio to their relative abun- dance. Snowshoe rabbits and red squirrels were not eaten. The Minnesotan data were from 60 stomachs and 53 intestinal tracts re- covered from 129 weasels trapped by use of scent ( not bait ) mostly from January 1 to February 7, 1939, although a few were trapped in 1938. Analyses of contents from stomachs gave approximately the same results as those from intestines. In 1939 at Lake of the Woods, weasels were concentrated where food was abundant but no such concentration was noted in the following winter. Big short-tailed shrew (Blarina brevicauda). — In New York State, the ermine preys on Blarina as shown by Hamilton's (1933:330) seeing one being carried by a male ermine on May 6, 1931, and another being carried by a female on May 13, 1932. The same 92 University of Kansas Publs., Mus. Nat. Hist. author (1928:249) found the remains of a Blarina in a small female from Malone, New York. Kirk (1921) observed, however, that the ermine ( M. e. cicognanii ) avoided the shrew, Blarina, caught in a trap and that Blarina avoided the weasel caught in a trap. Chipmunk (genus Tamias). — Remains were found in a male ermine in New York on May 14, 1932 (Hamilton, 1933:330), and Seton (1929 (2):602) records a chipmunk at Lake Couchiching, Ontario, that was pursued into the water by an ermine. Deer mice (genus Peromyscus) . — As shown by Hamilton (1933: 33) and Aldous and Manweiler (1942), Peromyscus was second only to microtines in numerical abundance among the food items of ermines in New York and Minnesota. Peromyscus and micro- tine rodents were brought to a den of the dimunitive M. e. muricus in early August, in Fresno County, California, according to Ingles (1942). He observed that an Alpine chipmunk was active under and around the tree and that juncos reared young 40 feet from the den but that the chipmunk and juncos were unmolested by the er- mines. Lemming (genus Lemmus). — One was recovered from a female ermine ( with milk in her glands ) at Laurier Pass, British Columbia (Sheldon, 1932:201). Red-backed mouse (genus Clethrionomys) . — Criddle and Criddle ( 1925:146) record that on "May 31, 1921. — Saw a Bonaparte's weasel capture a Red-backed Vole after a long hunt during which the pursuer never once lost track of its victim." Meadow mice (genus Microtiis). — As shown by the data of Hamilton (1933:333) and Aldous and Manweiler (1942) recorded above, Microtus is the item of first importance in the diet of the ermine in New York and Minnesota. Criddle and Criddle (1925: 146) write concerning the vicinity of Treesbank, Manitoba, that "October, 1918. — Following a severe outbreak of mice in 1916-17, Bonaparte's weasel increased enormously and very soon reduced the rodents to comparative rarity. This resulted in a scarcity of food for the weasels, which in their turn became greatly reduced in numbers." Old World rat {Ratt us). —Bishop (1923) found two headless rats near a nest of this species in Albany, New York. Pika (Ochotona). — Dixon (1931:72) at Milner Pass, Colorado, on July 20, 1931, saw an ermine, of the subspecies muricus, follow- ing a three-fourths grown pika by scent and outrunning the pika. The pikas worked a relay system and the weasel abandoned the trail when the fourth pika became the object of the chase. MUSTELA ERMINEA 93 Cottontail (genus Sylvilagus). — Hamilton (1933:33), as noted above, found remains of cottontail in the digestive tracts of ermine that had been trapped for fur in winter. Possibly these remains were bait that had been placed at traps. Snowshoe rabbit (Lepus americanus). — Morse (1939:210) in a study of predation on hares and grouse in the period of notable deci- mation of these two game species in 1935-1936 in the Cloquet Valley State Forest, in St. Louis County, Minnesota, found that "weasel predation on hares appeared to be of very low incidence or alto- gether lacking." Wild birds (Class Aves). — Aldous and Manvv^eiler (1942), as noted above, found that the remains of birds constituted five per cent by volume of the food of the ermine in winter in Minnesota. Chicken (genus GaZ/w5 ) .— Criddle and Criddle (1925:145), who published relatively extensive data on the tiiree species of weasels of Manitoba, write that: "We have no record of Bonaparte's weasel killing poultry, and we doubt whether it ever does so." However, Soper (1919:46) investigated the excited cackling of a hen brooding chicks at night and found a solitary ermine that had killed three chicks and that had tiie remainder under very active scrutiny. Leopard frog (Rana pipiens). — One frog was found in a male ermine on November 20, 1931, in New York by Hamilton (1933: 300). Fish (Class Pisces). — Aldous and Manweiler (1942) found frag- ments of a small fish in one of 60 stomachs of ermine from Minne- sota. Earthworm (Phylum Annelida). — Osgood (1936:64), presum- ably at Rutland, Vermont, observed a pair of weasels from 2:15 P. M. to 5:00 P. M., in a barn and saw the female in that time make many trips for food for her young. Only earthworms were brought. Fifty traps in an adjacent, swampy field caught only one bull frog and no mice indicating that mice had been eliminated from the foraging territory of the ermine. In handling food, Dice (1921:22) noted that the Alaskan ermine did not use the feet but only the mouth. Reproduction Litters of 4, 4, 7, 7, and 8, yielding an average of 6 young per litter have been recorded from the northeastern United States by Hamilton (1933:327). He {op d^ : 321-325) described animals one day old from New York State as being flesh-colored, having the long neck of tlie adult and a fine growth of white hair two milli- 94 University of Kansas Publs,, Mus. Nat. Hist. meters in length, on the dorsal surface of the neck, that foreshadows the mane or pompadour that is prominent from the 14th to the 21st day of hfe. Six animals, when one day old averaged 1.7 grams in weight, which was three per cent of the weight of an adult female and one and one half per cent of the weight of an adult male. At two weeks of age the heavy brown mane stood out in marked contrast to the rest of the scantily, white-furred animal. The eyes opened on the thirty-fifth day of life. .^^-^-'^. Fig. 24. Mustela erminea richard- sonii, adult female. Catalogue Number 14866, U. S. Nat. Mus., Fort Chimo, Ungava. X ^■ Ventral view of body of a pregnant female to show details of mastology. Note the five pairs of mammae char- acteristic of weasels, and the uneven arrangement of mammae of the two sides which is also common among weasels. MUSTELA ERMINEA 95 1 B+M M.e.orctico 2 ^SM M.e.poloris 3 ^H M.e. sempici 4 I I M.e.kodlacensis 5 llllllll M.e.richardsonii 6 ^S M.e.cicoqnanii 7 l'-'--Vv-v1 M.e.banqsi 8 9 10 I I 12 13 M.e.invicta M.e.alascensis M.e.salva M.e.initis M.e.celendo M.e. seclusa 4 15! I6l 17 18 19 20 M.e.haidarum 1 M.e.anquinoe I M.e.fallenda J M.e. olympico M.e. strcatori M.e. quiosa M.e. muricus Fig. 25. Map showing geographic ranges of the subspecies of Mustela erminea in the New World. For rearing their young, ermines live in burrows. Bishop ( 1923), in Albany, New York, found a burrow occupied by four young and a pair of adults. The burrow had many galleries and contained a nest constructed of rat fur, fine grass and fragments of leaves. 96 University of Kansas Publs., Mus. Nat. Hist. At Woods Lake, Fresno County, California, in early August, Ingles observed (1942) some young and at least one adult at their den which was in a burrow beneath a hollow tree. The ermines used the hollow root and the hollow tree as well as the burrow beneath. Seton (1929 (2):591) quotes S. Eldon Percival, of Barretts Rapids, Ontario, as finding the living quarters of an ermine in unthreshed grain stacked in a barn and says (op. cit. :590) that John Burroughs dug out a nest, composed of leaves and the fur of mice and moles, two or three handfuls in bulk, from a cavity the size of a hat, arched over with a fine network of tree roots. Four instances in which the male as well as the female was present at a den containing young are cited by Hamilton ( 1933:328) and he gives some evidence, although not at all conclusive, that "adults customarily pair, or at least run together, at times other than the breeding season". No other writers remark on this matter. I doubt that adult ermines are associated in pairs for most of the year but such may be the case. Mustela erminea arctica ( Merriam ) Ermine Plates 2, 3, 4, 9, 10, 11 and 41 Ptitorius arcticus Merriam, N. Amer. Fauna, 11:15, pi. 2, figs. 1, la, and pi. 5, figs. 6, 6a, June 30, 1896. Putorius (Gale) erminea, Coues, Fur-bearing animals, p. 109, 1877 (part). Ptitorius richardsonii. Bangs, Proc. Biol. Soc. Washington, 10:16, pi. 1, figs. 3, 3a, pi. 2, figs. 3, 3a, and pi. 3, figs. 6, 6a, February 25, 1896 (part). Putorius cicognanii alascensis, Osgood, N. Amer. Fauna, 19:43, October 6, 1900. Putorius kadiacensis, Osgood, N. Amer. Fauna, 21:69, September 26, 1901. Putorius aiidax Barrett-Hamilton, Ann. and Mag. Nat. Hist., 13(ser. 7): 392, May, 1904, type from Discovery Bay, EUesmere Island. Putorius alascensis, Heller, Univ. California Publ. Zool., 5:345, March 5, 1910. Mustela arctica arctica. Miller, U. S. Nat. Mus. Bull., 79:97, December 31, 1912; Dice, Joum. Mamm., 2:22, February 10, 1921. Mustela arctica. Hall, Univ. California Publ. Zool., 30:420, March 19, 1929. Mustela erminea arctica, Ognev, The mammals of U.S.S.R. and adjacent countries, 3:31, 1935; Hall, Proc. California Acad. Sci., 23:559, August 22, 1944; Hall, Joum. Mamm., 26:179, July 19, 1945. Type.— Male, adult, skull and skin; no. 14062 '23010, U. S. Nat. Mus.; Point Barrow, Alaska; July 16, 1883; obtained by John Murdock, original no. 1672. The skull has a fracture, on the dorsal surface, extending from the anterior nares to the interorbital constriction and another fracture on the left margin of the nasal bone. The middle of the left zygomatic arch is broken away. Otherwise the skull is complete. Right incisor one, above and below, are missing. Otherwise the teeth are present and entire. The skin is in the brown summer pelage, well made, in a good state of preservation, and shows no ob- vious signs of fading. MUSTELA EKMINEA 97 Range. — Arctic regions of Alaska and western Canada from the Pacific Ocean to Smith Sound; from the northern limit of land south approximately to a line from Skagway through Ft. Goodhope, north shore of Great Bear Lake, south shore of Clinton Golden Lake, north shore of Baker Lake, west end of Wagner Bay to south end of Committee Bay. See figure 25 on page 95. Characters for ready recognition. — Differs from M. e. polaris in darker upper parts (Raw Umber rather than Buckthorn Brown) and less intensely colored underparts that are Sulphur Yellow, Colonial Buff or Primrose Yellow rather than Buff Yellow; from M. e. semplei, in males, in that hind foot more than 44 and basilar length more than 41 and in that females average larger, the skulls of females being only about 11 per cent heavier; from M. e. kadiacensis in hind foot more than 33 in females, zygomatic breadth amounting to more, rather than less, than distance between last upper molar and jugular foramen irrespective of sex; from M. e. richardsonii, alascensis, salva and initis, both sexes so far as known, by proximal two-thirds of under side of tail colored same as underparts rather than same as upper parts, and by zygomatic breadth amounting to more, rather than less, than distance between last upper molar and jugular foramen. Description. — Size. — Male: Six adults from Tanana, Alaska, yield average and extreme measurements as follows: Total length, 336 (310-350); length of tail, 93 (84-105); length of hind foot, 49 (45-51). Female: Five adults, one each from Alatna River, mountains near Eagle, Kamarkak in Alaska, Arctic Red River and Baillie Island in Canada, yield average and extreme measiurements as follows: Total length, 285 (272-304); length of tail, 77 (68-95); length of hind foot, 39 (34-43). Weight of 5 subadult males from Tanana is 206 (163-248) grams; adults would be heavier. Color. — Winter pelage all white except tip of tail. Summer pelage with upper parts uniform in color and Raw Umber or darker (16n) of Ridgway and about tones 2 to 3 of Chocolate of Oberthiir and Dauthenay, pi. 343, but in autumn some specimens have more light red than tones 2 or 3. Underparts Sulphur Yellow, Colonial Buff, or Primrose Yellow, often white on chin and insides of forelegs; color of underparts extends narrowly over upper lips, distally on posterior sides of forelegs onto antipabnar surface of forefeet, onto proximal two-thirds or three-fourths of underside of tail as length of tail is measured along tail-vertebrae, on medial sides of hind legs to a point between knee and ankle but reappears on antiplantar faces of toes and in some individuals is narrowly continuous onto toes; rim of ear in some specimens with short, white or pale hairs giving ears distinct whitish border; least width of color of under- parts averaging, in adult males from Alaska, 65 (46-93) per cent of greatest width of color of upper parts. Black tip of tail in 5 males in winter pelage from Tanana averaging 84 (70-93) mm. which is 91 (75-107) per cent of length of tail-vertebrae. Skull. — Male (based on 5 adult topotypes): See measurements and plates 2-4. As described in Mustela erminea richardsonii except that: Weight, 3.5 (3.1-3.9) grams; basilar length 42.5 (41.8-43.3); length of tooth-rows more than length of tympanic bulla; breadth of rostrum measured across lacrimal processes averaging more than a third of basilar length; interorbital breadth 4—3758 98 University of Kansas Publs., Mus. Nat. Hist. more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more than distance between last upper molar and jugular foramen. Female (based on 2 adult topotypes and 2 adults and 4 subadults from central Alaska): See measurements and plates 9-11. As described in Mustela erminea richardsonii except that: Weight, 1.5 (1.2-2.0) grams; basilar length, 35.7 (34.5-37.0); length of tooth-rows more than length of tympanic bulla; breadth of rostrum more than 30 per cent of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more than distance between last upper molar and jugular foramen (except in specimens from Ellesmere Island where two distances are approximately equal). Cranial differences from Mustela erminea kaneii (which occurs on the Asiatic side of Bering Strait), in both males and females, are: larger size relatively as well as actually, broader except in mastoidal region where relatively (to basilar length) the width is less; preorbital part of skull broader as well as longer. From kadiacensis differences in the skull of the male are: size less; 13 per cent heavier, relatively (to basilar length) narrower across interorbital region and zygomatic arches; tympanic bullae relatively as well as actually narrower. Judging by the single avail- able adult female of kadiacensis, the skull of female arctica is larger in all parts measured, a fourth heavier, has tympanic bullae of al- most twice the volume and the interorbital and preorbital regions, relative to the braincase, are much reduced in whatever plane measured. Differences from richardsonii, additional to those noted above in the formal description of the skull, between the males, are: larger in all parts measured except length of tympanic bulla which is about the same; 42 per cent heavier; relative to basilar length, skull broader with preorbital part longer as well as broader; tympanic bullae more inflated posteriorly. The same differences prevail between females except that the skull is 36 per cent heavier and in arctica the length of the bulla is actually more ( although relative to the basilar length less ) and its greater inflation posteriorly is hardly perceptible. Dif- ferences from alascensis, additional to those indicated in the formal descriptions of the skulls of the two, in males, are: larger in every part measured; 95 per cent heavier; relative to the basilar length, skull broader with preorbital part longer as well as broader; meas- ured at a point opposite the foramen lacerum anterius, the width of the pterygoid space is more, rather than less, than 40 per cent of its length. Excepting this difference in width of interpterygoid space, the same differences prevail between females, those of arctica being 56 per cent heavier. MUSTELA ERMINEA 99 Comparison with semplei is made in the account of that sub- species. Skull indistinguishable from that of polaris. Remarks. — The person who studies specimens of this subspecies finds labels inscribed with the names of naturalists well known to all readers of literature on the Arctic. Sir John Franklin, R. Mc- Farlane, R. Kennicott, E. W. Nelson and R. M. Anderson are names which appear commonly. Of Alaskan specimens prepared accord- ing to modern methods, a large share was obtained by O. J. Murie and L. R. Dice. The ermine was observed in the far north by early explorers and was mentioned in the literature, almost always under the name then used for the ermine of northern Europe and Asia. In 1896 Bangs misapplied to it the name richardsonii but Merriam in the same year corrected the application of this name and proposed as new for this weasel the name arctica, the name in use today. For almost 50 years after Merriam and Bangs wrote about it, arctica was treated, nominally at least, as a species distinct from its other relatives in both the Old- and New-World. The subspecific status of arctica was emphasized in 1944 (555) by the present writer in reporting in detail upon the specimens, of Mustela erminea, from Eastern Asia which were made available on loan by Professor B. S. Vinogradov and the late Anatol I. Argyropulo of the Leningrad Academy of Sciences. Specimens of Mustela erminea kaneii from the Asiatic side of Bering Strait and Mustela erminea arctica from the American side are distinguishable by slight cranial characters but in coloration and external measurements I can detect no differ- ences. Merriam's (1896:16) mention of more golden-colored upper parts and darker underparts in American specimens than in erminea was the result of his comparison of Alaskan and northern European specimens. When Old World specimens from eastern Siberia, in- stead of from Europe, are used the differences mentioned by Mer- riam do not apply. Incidentally, many Siberian specimens have the white border, on the ear, which Merriam ( loc. cit. ) noted as a distinguishing feature of arctica. When Merriam named arctica he said (1896:15, 16) "Putorius arcticus . . . has heretofore been confounded with erminea or richardsoni. . . . It is inter- esting to find in this country an Arctic circumpolar weasel which, though specifically distinct, is strictly the American representative of the Old World erminea." Bearing in mind that Merriam's concept of species and subspecies (see Merriam, 1919:6) differed from that of nearly all modern systematists it is clear from his statement quoted 100 University of Kansas Publs., Mus. Nat. Hist. above that he correctly understood the zoological relationship obtaining between the ermines of the Old and New Worlds. Ognev (1935:31) seems to have been the first to use the name combination Mustela erminea arctica for Alaskan specimens. Thereby he expresses the view adopted here, namely that the American ermine is subspecifically but not specifically distinct from the Old World animal. Whether actual intergradation ( crossbreed- ing ) ever takes place across the narrow Bering Strait I do not know. I doubt that crossbreeding occurs but considering the Diomedes (islands), tliat might serve as a half way stopping point, and re- membering Mr. Charles Brower's oral statement to me that he had seen tracks of ermine as far as 10 miles from the northern shore of Alaska out on the ice, the possibility must be granted of an oc- casional individual crossing from one side to the other of Bering Strait on the ice in winter or of being carried across when the ice broke up and drifted. If transfers of this kind occurred often one would expect ermines to occur also on Saint Lawrence Island where apparently they do not. The one skin (U. S. Nat. Mus. no. 259046) seen as labeled from there, my friend, Otto William Geist ascer- tained was imported as a skin with other furs from Siberia. Ognev (op. cit.) who used the name combination Mustela er- minea arctica for Alaskan specimens, applied it also to animals from Kamchatka. At the same time he recognized the animal from the eastern mainland of Siberia (as opposed to the peninsula of Kam- chatka) under the name Mustela erminea orientalis Ognev 1928. Hall (1944:556) applied the earlier proposed name Putorius kaneii Baird 1857, to the animal on the eastern mainland of Asia and pro- posed the new name Mustela erminea digna for the ermine of Kam- chatka. In comparing material of these two Asiatic races with topo- types and other specimens of M. e. arctica from Alaska, it seemed to me that the degree of relationship, one with the other, was about the same. M. e. digna has a slightly larger preorbital region than M. e. kaneii, and the skull is longer. In both of these particulars digna approaches closer to arctica. M. e. kaneii has longer tympanic bullae and a wider skull than digna and tlierein approaches more towards arctica than toward digna. As nearly as I can make out, digna and kaneii show a nearly equal degree of resemblance to arctica. Also the degree of difference between digna and kaneii is about the same as between either one of them and arctica. In view of the above considerations the ermines of the New and Old worlds are here regarded as only subspecifically distinct. In the original description of Putorius audax (here regarded as MUSTELA ERMINEA 101 inseparable from Ptitorius arcticus Merriam) Barrett-Hamilton er- roneously designated the type locality as "Discovery Bay, North Greenland" whereas he should have written Grinnell Land [= El- lesmere Island of modern terminology] in place of Greenland. As reference to Nares ( 1877 and 1878 ) will readily reveal, Discovery Bay is near 65° W and 81° 40' N, across Robeson Channel, to the west, from Greenland. The label on the type specimen and the specimen register in the British Museum of Natural History each designates the locality for this specimen, the type of audax, as Discovery Bay without mention of Greenland. The published accounts of Feilden (1878) and Nares (1877 and 1878) state that specimens of ermine were obtained at Discovery Bay. Probably H. C. Hart is the collector of the specimen; he was the naturalist attached to H, M. S. Discovery which wintered at Discovery Bay while H, W. Feilden was the naturalist attached to H. M. S. Alert which wintered a few miles southeast of Cape Sheridan, also on the eastern coast of Ellesmere Island. It is true that from these ships a trip was made into Greenland and an ermine (only one individual it seems) was obtained tliere, but this individual was the type specimen of Mtistela erminea polaris, in the account of which race something of the history of this specimen is given. With the material available — and it is not entirely adequate — I can detect no features by which animals from the type locality of audax can be distinguished from typical arctica which latter name has priority. Intergradation with richardsonii probably occurs completely across the continent. Intergrades here referred to arctica include those from Fort Goodhope. The one defective specimen from Lake Lebarge, Yukon, is not certainly identified as arctica and how far west of Teslin Lake tlie boundary-line between arctica and richard- sonii should be drawn remains to be ascertained. The one specimen available from Hinchenbrook Island, no. 912 Mus. Vert. ZooL, an adult female, is doubtfully referred to arctica because the damaged tympanic bullae appear to be no larger than in alascensis, and the size of the skull is more as in alascensis although intermediate be- tween that race and arctica. Shape of the skull is more as in arctica. Possibly more nearly adequate material would show the existence on Hinchenbrook Island of an insular race differing in about the same degree from arctica of the mainland as does the insular kadiacensis. Nevertheless, the males from farther south at Cape Yakataga are in all respects arctica and this argues against near 102 University of Kansas Publs., Mus. Nat. Hist. relationship to alascensis of the animal on Hinchenbrook Island. The three animals seen from Yakutat Bay are so young as not to display clearly the cranial characters of the subspecies but the extension of the color of the underparts onto the underside of the tail in them and also in the skin without corresponding skull from Glacier Bay, Alaska, is as in arctica, the race to which they are referred, and gives substantial basis for showing the geographic range of arctica as extending this far south along the Pacific Coast. Specimens examined. — Total number, 281, arranged alphabetically by Dis- tricts and from north to south in each District. Unless otherwise indicated, specimens are in the collection of the United States National Museum. Alaska. Point Barrow, 22 {V, l^ 4', 7\ 6°); Flaxman Island, 3; Collinson Point, 1^ Sahrochet River, 1^ Hulahula River, 1^ 69°20' & 141°, 1; Rampart House, 1; Yukon River, mouth of Porcupine River, 18; Alatna River, 30 mi. from mouth, 1; Koyakuk Riv., 16 mi. below Bettles, 4; Shelton, 1'; Kruzamepa, 1'; Tanana, 6; Boulder Creek, Chena River, 3; Fort Reliance, 4; Yukon River, 20 miles above Circle, 2; Mts. near Eagle, 42 (1*); Snake River, Nome, 1'; Nulato, 3; No(e)wikakat Riv., 1; Kantishna, 3; Fairbanks, 5 ( 1 20 mi. E and 1 33 mi. E); Richardson, 1; N. Fk. Kuskokwim R. at base of Mt. Sischo, 1; N. Fk. Kuskokwim R. at Junction with McKinley Fk., 1; Nenana Riv., mouth of Maurice Cr., 1; Ober Cr., trib. of Jarvis Cr., Delta Riv. region, 1; head of Savage Riv., near Jennie Cr., 1; Wonder Lake, 1°; Bear Cr., 3; Unlakleet, 3; St. Michaels, 11; 125 mi. E and a little N of Knik, Cook Inlet, on S side Matanuska Range, 1*; Hope, Cook Inlet, 1; lak Lake, 1*; head of Behring Riv., 1; Bethel, 2; Kenai Lake, 8; Kenai Peninsula, 13 (2'); He[i]nchenbrook Island, 1200 ft., T; Sunshine Point, KaUekh River, Yakataga Dist., 1"; Cape Yakataga, 3"; Yakutat Bay, 3°; Seward, 7; Seldovia, 22 (4'); Homer, 1^ Cape Elizabeth, 18; Akchookuk Lake, 1; Lake Weelooluk, 1; Kokwok Riv., 80 mi. up, 4; Nushagak, 1; Nushagak Riv., 1; Kolukuk, 1; Egooshik River at mouth, 1; Glacier Bay, 1; Becharof Lake, between Portage Bay and Becharof Lake, 1; Ugashik Riv., 4; Chignik, 7; East base Frosty Peak, 1; Pavlov Bay, 1°; Mt. Pavlof, 1"; Unimak Island, 2 (1^). District of Franklin. Cape Sheridan, 1'; Discovery Bay, Ellesmere Island, 1" (type specimen of Putorius audax Barrett-Hamilton); Axel Heiberg Island, 1"; Bache Peninsula, Ellesmere Island, 1°; Bedford Pims Island, 4'; Craig Har- bor, 2"; Cape Kellett, Banks Island, 1'; Franklin Isthmus, 1"; King William Island, 2". District of Keewatin. Ualiak, Ogden Bay, 2". District of Mackenzie. Baillie Island, 1"; FrankUn Bay, 1; Langton Bay, arm of Frankhn Bay, 15 mi. S of, 1'; Cockbum Point, 69°N, 115°W, 2"; Dolphin and Union Strait, 1°; Bernard Harbor, 2°; Kent Peninsula, 4'°; Horton Riv., near Fort Anderson, 1; Fort Anderson, 6; Anderson River, 3; Barry Island, Bathurst Inlet, V; Fort McPherson, 1; Peels River, 2; Arctic Red River, 8=; Fort Good Hope, 6; Clinton Golden, F. Yukon. Kamarkak, 1"; Herschel Island, T; Lapierres House, 2; Forty Mile, L. T. Coal Cr., 4°; head of Coal Cr., 1; Macmillan River, Forks, 1; 20 mi. W. Ft. Selkirk, 1; Slims River, near Kluane, 1'; head of Lake Lebarge, 1. 1. American Mus. of Nat. History 7. Carnegie Museum 2. Museum of Comparative Zoology 8. Leland Stanford Junior University 3. Acad. Nat. Sciences of Philadelphia 9. University of Oklahoma. 4. Field Museum of Natural History 10. Univ. Zool. Mus., Copenhagen 5. Mus. Vert. Zocil., Univ. California 11. British Museum of Natural History 6. Canadian National Museum 12. California Academy of Sciences MUSTELA ERMINEA 103 Mustela erminea polaris (Barrett-Hamilton) Ermine Putorius orcticus polaris Barrett-Hamilton, Ann. and Mag. Nat. Hist., 13 (ser. 7):393, May, 1904. Mustela erminea, Manniche, Meddelelser om Gr0nland, 45:80-85, 1 fig., 1910. Mustela arctica polaris, Miller, U. S. Nat. Mus. Bull., 79:97, December 31, 1912. Mustela erminea polaris. Hall, Joum. Mamm., 26:179, July 19, 1945. Type. — Probably female, skin only; no. 78. 6. 19. 11, Brit. Mus. Nat. Hist.; Gap Valley, Di miles northeast Cape Brevoort, 82° N, 59° 20' W, Northwestern Greenland; June 15 or 16, 1876; obtained by Lewis A. Beaumont. The skin is in full, fresh summer pelage, fairly well stuffed except for the tail which is unstuffed; the whole is in a good state of preservation. Range. — North coast, and east coast as far south as Turner Sound (between 69 and 70 degrees) of Greenland. See figure 25 on page 95. Characters for ready recognition. — Differs from M. e. arctica in lighter upper parts (near[/] Buckthorn Brown rather than Raw Umber or darker) and more intensely-colored underparts that are Buff Yellow rather than Sulphur Yellow, Colonial Buff, or Primrose Yellow; from M. e. semplei in color in same fashion as from arctica and in larger size of skull. Description. — Size. — Male: One subadult and two adults (one ad. from Sc0resby Sound and other two from Ymer Island) measure as follows, the average being given first: Total length, 318 (301, 320, 315); length of tail, 72 (69, 70, 73); length of hind foot, 46.5 (44, 46, 47). Female: No measurements taken in the flesh available but hind foot, measuring 33.5 in the dried state and therefore approximately 35 in hfe. Color. — As described in Mustela erminea arctica except that upper parts in summer near(/) Buckthorn Brown and tone 4 of Dark Fawn of plate 307 to tone 1 of Raw Umber of plate 301 of Oberthiir and Dauthenay. Underparts Buff- Yellow. Least width of color of underparts averaging, in 3 males, 66 (57-72) per cent of greatest width of color of upper parts. Black tip of tail in same males averaging 71 (70-72) mm. which is 99 (99-104) per cent of length of tail-vertebrae. The fighter-colored upper parts and more intensely yellow underparts are the distinguishing features of the subspecies polaris in comparison with other races of American M. erminea. Skull. — Male (based on 5 adults from eastern Greenland): See measure- ments. As described in Mustela erminea richardsonii except that: Weight more (not recorded); basilar length, 41.3 (39.0-42.4); length of tooth-rows more than length of tympanic bulla; breadth of rostrum measured across lacrimal processes averaging more than a third of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more than distance between last upper molar and jugular foramen. Female (based on 2 adults. Turner Sund and Kap Hoeg): See measure- ments. As described in Mustela erminea arctica except that basilar length 36.8 (35.9, 37.8), and length of tooth-rows not more tlian length of tympanic 104 University of Kansas Publs., Mus. Nat. Hist, bulla. Skulls of females not in hand when this comparison is written; only the recorded measurements are available. To me the skull of polaris is indistinguishable from that of arctica. Therefore the comparisons made of the skull of arctica with those of other subspecies will apply also for polaris. RemarJcs. — In view of the heretofore erroneous assignment of the type locality of Mustela erminea audax to Greenland, pains were taken to verify the statement by Barrett-Hamilton (1904:393) rela- tive to the type specimen of polaris. Taking pains thus seemed the more worthwhile because in the specimen register at the British Museum of Natural History, there is written to the right of cata- logue numbers 78 - 6 = 19 nos. 1 - 11, "Discovery Bay Presented by Mr. Hart Arctic Collection." This refers to no. 78.6.19.1. There are no ditto marks below but by implication this data applies also to nos. 1-11, which include the holotype of polaris. A label at- tached to the specimen does however give the locality as "Hall Land" "N Greenland" and another label has on it "Ermine, pro- cured by Mr. Beaumont Greenland Lat 89° Long W 59-20." The 89° is obviously a mistake ( on the label or in my transcription of it ) for 82°. Reference to Nares (1877:385) reveals that Lieutenant Lewis A. Beaumont, under date of June 15 and 16, 1876, wrote in his field journal as follows: "I shot an ermine." In the daily accounts of his journey from Discovery Bay on Grinnell Land [^ Ellesmere Is- land], across Robeson Channel and along the north coast of Green- land to the west base of Mount Farragut near 50° 30' W he men- tions the ermine only this once. For several other kinds of animals, Beaumont mentions individuals seen or shot, often with the nota- tion that this is the second, or third seen. This mention of a kind of animal whenever seen was in accordance with orders. On page 39 of the Discovery Report {op. cit., 1877) in "General orders to sledging parties" by Captain G. S. Nares, Commanding the Expedi- tion, we find ". . . note daily: IV State the animals seen and those shot." Reference to the map facing page 358 of the ( op. cit. ) report reveals that on the 15th and 16th, camps were made by Beaumont in Gap Valley, each 7% miles northeast of Cape Brevoort, one camp on either side of the 82° line, and separated from each other by a distance of only 2/4 air hue miles or 4/2 miles march ac- cording to his journal. These several data, then, are the bases for designating the type locality of M. e. polaris, in the way that I have stated it at the begin- ning of this account of the subspecies. MUSTELA ERMINEA 105 The light-colored upper parts and more intensely yellow iinder- parts well differentiate this subspecies from arctica or scmplei. Intergradation is suggested by a skin, no. 1462, Copenhagen Zoo- logical Museum, from Axel Heibergs Land, the color of the under- parts of which agrees with that of specimens from Greenland. Also the color of the upper parts is decidedly nearer that of animals from Greenland than to that of specimens from Ponds Inlet, Tulican ar.d Gifford River. No other specimens west or south of Greenland suggest intergradation. In Greenland itself, one adult, a female from Turner Sund, East Greenland, has the underparts no more yellowish than in some specimens from Melville Peninsula. This female is darker on the back than any one of the other 10 specimens from Greenland in summer pelage examined at the same time, but even so is not so dark colored as animals from Baffinland or other islands to the west of Greenland. The final summation of information about this subspecies would have been more precise if I had been able to have actually in hand, at the time of writing, specimens preserved in the Copenhagen Zoological Museum. The war made it impractical to secure the loan of these as previously planned. Even so, the measurements and notes on color that I obtained from this material, in 1937, in Copenhagen, suffice to prove that the subspecies polaris is well set off in color from the other American subspecies of Mtistela erminea. The best material of this subspecies is in the University Zoo- logical Museum at Copenhagen, Denmark. Specimens examined. — Total number, 35, arranged by locality from the western end of the north coast of Greenland, eastward and then southward down the east coast. Unless otherwise indicated, specimens are in the Uni- versitetets Zoologisk Museum, K0bnhavn, Danmark. Gap Valley, 7% mi. NE Cape Brevoort, 82 N, 59 20' W, 1 ( British Mus. ) ; Dragon Point, 1; Danmarks Havn (Fjeldene ved Baadskjeret, 1; hlle Fjeld, 1; Lyservig, 1; harefjeldets, 4; Rypefjeldet, 1; Baadskjeret, 1; Danmarkshavn, 3) 12; Christians Havn, 1 (not found on map); Shannon Island, 4; Germania Havn, 2; Claveringoen, 1; Carls Havn, 1; Myggbukta, 2 (British Mus.); Ymer[s] Island, 2 (Mus. Comp. Zool. ); Kap Hoegh, lamesonsland, 1 (Berhn Zool. Mus.); Scoresby Sund, 3; Turner Sund, 4. Mustela erminea semplei Sutton and Hamilton Ermine Plates 2, 3, 4, 9, 10 and 11 Mustela arctica semplei Sutton and Hamilton, Ann. Carnegie Mus., 21:79, February 13, 1932. Mustela arctica labiata Degerbol, Kept. 5th Thule Exped., 2 (no. 4):25, 1935, type from Malugsitaq, Melville Peninsula, Canada. Mustela erminea semplei. Hall, Journ. Mamm., 26:179, luly 19, 1945. Type. — Male, subadult, skull and skin; no. 6470, Carnegie Mus.; Coral Inlet, 106 University of Kansas Publs., Mus. Nat. Hist. South Bay, Southampton Island, Canada; October 8, 1929; obtained by George Miksch Sutton, original no. 3M. The skull has two holes in it: one is immediately above the left canine, and the other ( 2 X 5.5 mm. ) is 3 millimeters to the left of the median line at the juncture of the frontal and parietal bones. From this last mentioned hole a fracture extends back halfway to the lambdoidal crest. The tip of the left upper canine is broken oflF. Otherwise the skull is complete, and the teeth all are present and entire. The skin is well made and in fresh white winter pelage except for a trace of the old brown summer pelage on the back, on the tail, on the anterior borders of the ears, and in a spot 11 mm. long and 8 mm. wide on the nose. Range. — Baffin and Southampton islands, Melville Peninsula and west side of Hudsons Bay as far south as Eskimo Point. See figure 25 on page 95. Characters for ready recognition. — DifFers from A/, e. arctica, in that, in males, hind foot less than 44 and basilar length less than 41 and in that females average smaller, their skulls being only about 10 per cent lighter; from M. e. polaris in darker upper parts ( Raw Umber rather than Buckthorn Brown ) and less-intensely-colored underparts that are Sulphur Yellow, Colonial Buff or Primose Yellow rather than Buff Yellow, and in lesser size in the same fashion as from arctica; from M. e. richardsonii, of both sexes, in that proximal two- thirds of under side of tail colored same as underparts rather than same as upper parts and by least interorbital breadth amounting to more, instead of less, than distance between glenoid fossa and posterior border of external auditory meatus. Description. — Size. — Male: Ten adults and subadults, from Southampton Island, yield average and extreme measurements as follows: Total length, 282 (267-318); length of tail, 77 (59-87); length of hind foot, 40 (38-43). Female: Four subadults from Southampton Island yield average and ex- treme measurements as follows: Total length, 271 (256-288); length of tail, 71 (69-74); length of hind foot, 35 (33-38). Color. — As described in M. e. arctica except that least width of color of underparts averaging, in 7 males, 59 (45-81) per cent of greatest width of color of upper parts. Black tip of tail in 19 male topotypes averaging 72 ( 64-83) mm. which is 91 (75-122) per cent of length of tail-vertebrae. Skull. — Male (based on 2 adults and 10 subadults from Southampton Is- land ) : See measurements and plates 2-4. As described in Mustela erminea richardsonii except that: Weight, 2.0 (in one subadult) grams; basilar length, 37.5 (35.7-39.9); length of tooth-rows more than length of tympanic bulla; breadth of rostrum more than a third of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more than distance between last upper molar and jugular foramen. Female (based on 1 adult and 4 subadults from Southampton Island): See measurements and plates 9-11. As described in Mustela erminea richardsonii except that: Weight, 1.35 (in one adult) grams; basilar length, 34.2; breadth of rostrum more than 30 per cent of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breath more or less than (approximately same as) distance between last upper molar and jugular foramen. MUSTELA ERMINEA 107 In comparison with richardsonii, the skulls of males averaged smaller in every measurement taken except breadth of rostrum and interorbital breadth which are more, and zygomatic breadth and length of inner lobe of Ml which are approximately the same; skull about 20 per cent lighter; in relation to basilar length, preorbital region longer and broader in every part measured. Female aver- ages larger, in every part measured; 23 per cent heavier; in relation to basilar length, every other measurement more. It is noteworthy that the skull of the male is smaller and the skull of the female larger than in richardsonii. Differences from arctica are: Size less, in each sex; males about 40 per cent and females 10 per cent lighter; in males, skull more rounded in outline as viewed from above because zygomatic arches arise less abruptly from skull; in males tympanic bullae do not project so far ventrally from squamosal floor of braincase; with these exceptions, skull of semplei can be said to be a smaller edition of that of arctica. From polaris, semplei difFers, cranially, in the same way as from arctica. Remarks. — There is a slight increase in size of ermines toward the north which probably is the result of intergradation between semplei and arctica. Specimens from the northern part of Baffin Island are larger than those from farther south. Specimens from the main- land west of Southampton Island may owe their smaller (than in arctica) size to intergradation with richardsonii almost as much as to intergradation with semplei. Degerbc^l's name Mustela arctica lahiata was applied to speci- mens, which to me are indistinguishable from topotypes of Mustela arctica semplei, which latter name has three years priority. Deger- b'al Ontario Museum of Zoology 7. Collection of Edward R. Warren 4. United States National Museum 8. Carnegie Museum MUSTELA ERMINEA 109 The white, winter skin is only moderately well stufiFed but in a good state of preservation. The spring coat is appearing along the back. This coat is visible at only two places unless the hair be parted when the new brown pelage, which is coming in, can be seen all along the midline of the back. Range. — Kodiak Island, Alaska. See figure 25 on page 95. Characters for ready recognition. — Differs from M. e. arctica in hind foot less than 33 in females and in zygomatic breadth amounting to less, instead of more, than distance between last upper molar and jugular foramen irrespective of sex. Description. — Size. — Male: One adult and 3 subadults yield average and extreme measurements as follows: Total length, 341 (318-360); length of tail, 93 (86-102); length of hind foot, 47 (44-49). Female: An adult measures: Total length, 258; length of tail, 70; length of hind foot, 31. Color. — As described in M. e. arctica, except that least width of color of underparts averaging 54 (40-83) per cent of greatest width of color of upper parts. Black tip of tail in 3 males in summer pelage averaging 80 (70-90) mm. which is 85 (69-96) per cent of length of tail-vertebrae. Skull. — Male (based on 2 adults): See measurements and plates 2-4. As described in Mustela erminea richardsonii except that: Weight 3.1 grams; basilar length, 42.6 (42.1-43.2); length of tooth-rows more than length of tympanic bulla; breadth of rostrum measured across lacrimal processes aver- aging more than a third of basilar length; interorbital breadth more than dis- tance between glenoid fossa and posterior border of external auditory meatus. Female (based on one adult, no. 98042) : See measurements and plates 9-11. As described in Mustela erminea richardsonii except that: Weight, 1.2 grams; basilar lengdi, 33.0; length of tooth-rows more than length of tympanic bulla. Comparison with arctica has been made in the account of that subspecies. Although richardsonii and kadiacensis are described as having the zygomatic breadth less than the distance between the last upper molar and jugular foramen, the zygomatic breadth is con- siderably more in kadiacensis than in richardsonii; consequently the two dimensions are more nearly equal than in richardsonii. Except for being slightly narrower, the skull of kadiacensis is only a slightly smaller edition of that of arctica. Remarks. — When naming the weasel from the mainland of Alaska as new, under the name Pittorius arcticus, Merriam ( 1896:16) wrote: "A small form of arcticus occurs on Kadiak Island. . . , It is probably worthy of recognition as subspecies kadiacensis." The in- formality of this description possibly was in part due to the de- scriber's recognition of the fact that the degree of difference be- tween arcticus and the insular kadiacensis was slight. Specimens collected after Merriam proposed the name for the weasel of Kodiak Island show the animal there to be less different from arctica of the adjacent mainland than he thought; small size is the most pro- 110 University of Kansas Publs., Mus. Nat. Hist. nounced distinction of kadiacensis and Merriam's male type speci- men is smaller than any of the five additional males saved from Kodiak Island since that time. Even so the differences fully w^ar- rant subspecific recognition, in my opinion, although kadiacensis is not a strongly differentiated race. More adult females are needed to ascertain the norm of form and size for that sex. If the one female known is typical, the difference from arctica is more pro- nounced in females than in males. The lesser size of kadiacensis can hardly be credited entirely to the effect of insularity, for animals from the southern part of the mainland, on Kenai Peninsula for example, are smaller than those from central and northern Alaska and provide evidence of intergradation of a sort between kadiacensis and arctica. Specimens examined. — Total number, 9, all from Kodiak Island, Alaska, and unless otherwise indicated in the U. S. National Museum. Karluk, 1 (Stanford Univ.); Kodiak, 7; Kodiak Island, 1 (Field Mus. Nat. Hist.). Mustela erminea richardsonii Bonaparte Ermine Plates 2, 3, 4, 9, 10 and 11 Mustela richardsonii Bonaparte, Charlesworth's Mag. Nat. Hist., 2:38, 1838. Putorius cicognanii, Baird, Mamm. N. Amer., p. 161, 1858 (part). Putorius richardsonii, Baird, Mamm. N. Amer., p. 164, 1858 (part-Halifax, N. S.). Putorius (Gale) erminea, Coues, Fur-bearing animals, p. 109, 1877 (part). Putorius richardsoni. Bangs, Proc. Biol. Soc. Washington, 10:16, February 25, 1896. Putorius cicognani richardsoni, Merriam, N. Amer. Fauna, 11:11, June 30, 1896. Putorius (Arctogale) cicognanii cicognanii, Bangs, Proc. New England Zobl. Club, 1:18, February 28, 1899. Putorius microtis Allen, Bull. Amer. Mus. Nat. Hist., 19:563, October 10, 1903. Type from Shesley, British Columbia. Putorius arcticus imperii Barrett-Hamilton, Ann. and Mag. Nat. Hist., 13(ser. 7):392, May, 1904. Type from Fort Simpson, Mackenzie, Canada. Putorius cicognanii richardsoni, Preble, N. Amer. Fauna, 27:231, October 26, 1908. Mustela microtis. Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912. Mustela cicognanii mortigena Bangs, Bull. Mus. Comp. Zool., 54:511, July, 1913. Type from Bay St. George, Newfoundland. Mustela cicognanii, Sheldon, Joum. Mamm., 13:201, August 9, 1932. Mustela cicognanii richardsonii, Miller, U. S. Nat. Mus. Bull., 79:95, De- cember 31, 1912; Hall, Univ. California Publ. Zool, 40:368, November 5, 1934. Mustela cicognanii cicognanii. Hall, Canadian Field- Nat., 52:108, October, 1938. Mustela erminea richardsonii. Hall, Journ. Mamm., 26:77, February 27, 1945; Hall, Journ. Mamm., 26:180, July 19, 1945. Type. — Male, age unknown, skin; no. 43.3.3.4, British Museum of Natural History; probably from Fort Frankhn, Canada; presented to British Museum on or before March 3, 1843; may be the type. MUSTELA ERMINEA 111 In September, 1937, when I searched in the British Museum for the skull, I found no trace of it nor mention of it in catalogues. The skin is in white, winter pelage, mounted on a pedestal. See under remarks for Mustela e. cicognanii for reasons for and reasons against regarding this specimen as the holotype. Range. — Hudsonian and Canadian life-zones of the greater part of Canada from the Atlantic to the Pacific. See figure 25 on page 95. Characters for ready recognition. — Differs from M. e. arctica, polaris, sem- plei and haidarum, in both sexes, by proximal two-thirds of under side of tail colored same as upper parts rather than same as underparts, and interorbital breadth less, rather than not less, than distance between glenoid fossa and posterior border of external auditory meatus; from M. e. hangsi, in that, in both sexes, least width of color of underparts averages two-fifths rather than about a third of greatest width of color of upper parts, and in that skulls of males are a fourth heavier, basilar length averaging more than 40; from M. e. cicognanii, in both sexes, in that least width of color of underparts averages two-fifths instead of less than a third of greatest width of color of upper parts, in females by 20 per cent heavier skull (1.1 versus 0.92), in males by skull more, rather than less, than 1.9 grams, and basilar length more, instead of less, than 38; from M. e. invicta, in males, by skull more, instead of less, than 1.9 grams; mastoid breadth more, instead of less, than 19.9 mm.; depth of skull at anterior margin of braincase more, instead of less, than 12.4 mm.; in females, by same measurement of depth more, instead of less, than 10.1, and weight of skull averaging more, instead of less, than one gram; from M. e. fallenda in both sexes upper lips white rather than brown, in males, hind foot more than 41, basilar length more than 38.3, in females hind foot more than 29, basilar length more than 31.4, and breadth of rostrum amounting to less, instead of more, than 30 per cent of basilar length; from M. e. alascensis in males in that black tip of tail more than 43, total length more than 320, tympanic bullae more than 14 and longer than tooth-row rather than less than 14 mm. and sometimes shorter than tooth-row, females not individually distinguishable. Description. — Size. — Male: Four adults (Fort Franklin, Fort Simpson, Mts. W Fort Nelson, and Govt. Hay Camp, Wood Buffalo Park) yield average and respective measurements as follows: Total length, 331 average (340, 325, 330, 328); length of tail, 93 (102, 91, 93, 87); length of hind foot, 45 (48, 43, 45, 44). Weight of 4 adults from the Belcher Islands is 175 ( 135-180) grams. Of 10 subadults from Belcher Islands it is 119 (92-137) grams. Female: Three adults from Great Slave Lake (Willow River, Fairchild Point, and Fort Resolution) yield average and respective measurements as follows: Total length, 252 (237, 238, 282); length of tail, 69 (63, 60, 85); length of hind foot, 32 (31, 32, 34). Corresponding, average measurements for three adults from Glacier Lake are 240, 60, 32 and for 3 adults from the Athabasca Delta, 243, 65, 30. Weight of 8 subadults from the Belcher Is- lands is 69 (64-78) grams. Weight of adults would be more. Color. — Winter pelage all white except tip of tail. Summer pelage vidth upperparts unifonn in color and darker (16n) than Raw Umber, and about tones 3 to 4 of Chocolate of Oberthiir and Dauthenay, pi. 343. Underparts Sulphur Yellow, Colonial Buff, or Primrose Yellow, often nearly white on chin and insides of forelegs; color of underparts extends narrov^ly over upper lips, 112 Unrtersity of Kansas Publs., Mus. Nat. Hist. distally on posterior sides of forelegs onto antipalmar faces of toes and some- times over most of antipalmar surfaces of forefeet, on medial sides of hind legs to a point between knee and ankle but reappears on antiplantar faces of toes and in some individuals is narrowly continuous onto toes. Least width of color of underparts averaging, in a series of 12 males from the Athabaska Lake Region, 40 (25-54) per cent of greatest width of color of upper parts. Black tip of tail averaging 56 (45-63) mm. in 5 adult males from same region and thus 60 (48-70) per cent of length of tail-vertebrae. From arctica, polaris, semplei and kadiacensis, richardsonii differs in: Color darker; ventral side of tail same color as upper parts; Hght-colored underparts a fifth narrower; black tip of tail by actual measurement a fifth shorter and averaging less than two-thirds rather than more than four-fifths of length of tail-vertebrae. From cicognanii, richardsonii differs in that the underparts are a fourth wider and in some specimens more brightly colored. The width of the underparts is likewise a fourth more than in hangsi. In invicta the under- parts are not so brightly colored as in some specimens of richardsonii. From fallenda, richardsonii differs in that the upper parts often are lighter colored, upper lips white rather than colored like upper parts, and underparts as wide again. In comparison witli alascensis, the black tip of the tail averages three- fifths rather than a half of length of tail- vertebrae. Skull. — Male (based on 6 adults from 3 miles south of Big Island, Great Slave Lake): See measurements and plates 2-4; weight, 2.5 (2.1-2.9) grams; basilar length, 40.9 (39.6-43.7); length of tooth-rows less than length of tym- panic bulla; breadth of rostrum measured across lacrimal processes less than a third of basilar length; interorbital breadth less than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth less than distance between last upper molar and jugular foramen. Female (based on 4 adults: from Willow River, 1; Fort Resolution, 1; Athabasca Delta, 2; and 2 subadults, one from 3 mi. S Big Island and one from 15 mi. above Smith Landing): See measurements and plates 9-11; weight, 1.1 (0.9-1.4) grams; basilar length, 33.1 (31.5-34.2); length of tooth- rows less than length of tv'mpanic bulla; breadth of rostrum less than 30 per cent of basilar length; interorbital breadth less than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth less than distance between last upper molar and jugular foramen. The skull of the female averages 56 per cent fighter than that of the male. Comparison of the skull with that of arctica, polaris, semplei, kadiacensis, haidarum, cicognanii, hangsi, invicta, fallenda, and alascensis is made in the accounts of those subspecies. Remarks. — M. e. richardsonii has the most extensive geographic range of any American race of erminea, is centrally located with re- spect to the other races, is more abundantly represented by study specimens in zoological collections than any other race, and is a sort of average for the species as a whole in most structural features. Therefore richardsonii is used as a standard of comparison and ac- cordingly is more fully described than any one of the other races each of which by reference to richardsonii is described in compara- MUSTELA ERMINEA 113 tive fashion. This comparative description has the virtue of more clearly indicating differences between subspecies and also makes for brevity. John Richardson, Bernard R. Ross, and names of their com- panions, as written on the labels of the older specimens recall to the student's mind early explorations of the north country. Edward A. Preble obtained important specimens at several places and in recent years J. Kenneth Doutt and G. G. Goodwin have made the reviser's work easier by preparing specimens in series from areas not pre- viously well represented. The nomenclatural history of this subspecies begins with refer- ences in the literature that identify the animal as the Old World species, Miistela erminea — an identification which the study here reported upon shows to have been correct in the specific, although not in the subspecific, sense. Richardson, for example, in his "Fauna Boreah-Americana" published in 1829 so identified the animal. In 1838, Bonaparte, basing his description on Richardson's account of 1829, proposed the new name richardsonii. Richardson himself, the following year in the "Zoology of Beechey's Voyage", accepted Bonaparte's name and it has been applied to the animal in the central part of the northern timber-belt of North America ever since, except as authors used the name Mustela erminea in the belief that richardsonii was not distinct from erminea. The north and south boundaries of the range assigned to richard- sonii varied according to the notions of the particular writer who was employing the name. Until Merriam in 1896 named arctica as distinct, animals from the far north were generally included under the name richardsonii along with populations to which the latter name now is applied. Because richardsonii grades gradually into the smaller cicognanii of more southern occurrence the boundary between the two has been set farther north by one writer and farther south by another, depending probably upon what the writer felt was the half-way point in size. This point of course depended upon the samples selected as typical of richardsonii on the north and cicognanii on the south. Because Bangs, in 1896, took as rep- resentative of richardsonii the far northern and hence large-sized animals ( now separated as M. e. arctica ) , his half-way point in size between them and the small cicognanii of New England naturally fell farther north than it would have had he used as representative of richardsonii specimens from places south of the range of arctica. In 1903 J. A. Allen proposed the name Putorius microtis for a specimen from Shesley, northwestern British Columbia, a place ap- 114 University of Kansas Publs., Mus. Nat. Hist. proximately 50 miles northwest of Telegraph Creek. Considering the great disparity in size between this one specimen and the other larger specimens of normal size, from the general region, available to Allen at that time, it is not surprising tliat he thought two full species were represented. In 1943 when G. G. Goodwin called to my attention two males, as small as the type of microtis and taken by him approximately 300 miles east of Shesley, in the valley be- tween the Musqwa and Prophet rivers, I for a second time ex- amined all available specimens and data with the possibility in mind that microtis was a species or subspecies distinct from M. e. richardsonii, but again concluded that only one subspecies was in- volved because no character except size was found to distinguish the large from the small individuals of a given sex and there are, preserved from northern British Columbia, individuals of inter- mediate size. Putorius microtis Allen seems to have been based on an individual of M. e. richardsonii near the lower limit of size for that subspecies and microtis is regarded as a synonym. Barrett-Hamilton in 1904 named the animal at "Fort Simpson, British Columbia" Putorius arcticus imperii. Preble (1908:232) pointed out that Fort Simpson on the Mackenzie undoubtedly was the place intended, and arranged imperii as a synonym of M. e. richardsonii. The type specimen of imperii was stated to have been received from B. M. Ross who is known to have collected specimens, including specimens of this species ( now in U. S. Nat. Mus.), at Fort Simpson on the Mackenzie. I know of no Fort Simpson in British Columbia. If, as seems improbable. Port Simp- son, British Columbia, was the place that Barrett-Hamilton intended to designate (where so far as I know Ross did not collect), the name imperii still would seem to be a synonym of richardsonii be- cause richardsonii seems to be the race of weasel at Port Simpson. In proposing the name Putorius arcticus imperii, Barrett-Hamilton stressed that the weasel, which he was naming, was a subspecies of P. arcticus, gave characters which applied perfectly to richard- sonii but made no reference to richardsonii. Barrett-Hamilton did not refer to richardsonii possibly because he relied on Merriam's classification of 1896 wherein richardsonii is treated as a species dis- tinct from arctica. Merriam, it will be remembered, held that slight degree of morphological difference rather than intergradation was the criterion for subspecies. Although I have no record of having examined the type specimen of imperii I have but little hesitancy in treating it as a synonym, and would have no hesitancy at all in so doing if the type was certainly known to have been obtained at Fort Simpson on the Mackenzie. MUSTELA ERMINEA 115 The name Mtistela cicognanii mortigena Bangs, 1913, proposed for the ermine of Newfoundland, is placed as a synonym of rich- ardsonii only after repeated, detailed comparisons. In advance of study I supposed that the isolation of the ermine, in Newfoundland, had contributed to its differentiation, which, however, the original describer, Bangs, indicated was slight. Bangs was a careful worker and I am confident that the differences he described really existed between his specimens. Material more nearly adequate than he had from the mainland, shows the males, so far as my measure- ments and comparisons go, to be in nowise diflFerent from those in Newfoundland. Females in Newfoundland may have, on the aver- age, slightly longer hind feet than on the opposite mainland but I am not certain that they do and even if there is a slight difference in this regard as suggested by available data, I think it insufficient basis, alone, for according subspecific status to the insular animal. The name richardsonii was based by Bonaparte on Richardson's description which in turn was drawn from a specimen taken at Fort Franklin, that thus becomes the type locality. It is fortunate that Preble, in 1903, succeeded in taking specimens there because the place is near the belt of intergradation between arctica and richardsonii. Of Preble's two adult males (see Preble, 1908:232) I have examined no. 133847, which is in transitional pelage and therefore gives no clue in so far as coloration is concerned, as to affinities with arctica versus richardsonii. Specimens in the summer pelage are much to be desired from Fort Franklin. Regardless of what their coloration may be, specimen no. 133847, in external measurements and most certainly in cranial features is of the race to the south and not the race that Merriam named arctica. Because all specimens from localities to the south of Fort Franklin likewise differ from arctica of the barren grounds, considerable additional confidence is felt in allocating the name richardsonii to the animal which ranges from Fort Franklin southward rather than to the one, here designated arctica, that occurs to the northward of Fort Franklin. Although in most structural features richardsonii is a sort of average for the American races of the species, it is the extreme in high degree of sexual dimorphism. The difference in size be- tween the males and females is greater than in any other race except possibly M. e. kadiacensis in which so little is known of the female that the difference between the two sexes cannot be accurately judged. It will aid in understanding the high degree of secondary sexual difference in richardsonii to visualize two 116 University of Kansas Publs., Mus. Nat, Hist. kinds of weasels distributed over the northern half of the continent, thinking now of the geographic area in America occupied by the whole species Mustela erminea of which the subspecies richardsonii is only a part. One of the two kinds of weasel is the male ermine and the other the female. The decrease in size of the male, as measured by the weight of the skull, is in the ratio of 7 in the north to 2 in the south. This decrease is gradual whereas the corresponding decrease from 3 to 1 in the female is not gradual; half of the decrease in the female occurs in the short north to south distance comprised in the belt of intergradation, along the northern boundary of richardsonii, between it and arctica. As a result richardsonii is composed of females with medium sized skulls and males with relatively large skulls, the ratio by weight being approximately 5 to 2. The disproportion in races of ermines both to the north and to the south is less. Actually in the north (arctica) the approximate ratio by weight is 21-^:1; in richardsonii, 2%:1; in the south (muricus), 1%:1. Indicated in still another way in richardsonii the skull of the female is 56 percent lighter than that of the male and the skull of the male is 127 percent heavier than that of the female. Intergradation with races whose ranges border on that of richardsonii is complete. On the northern boundary of the range of richardsonii along the western shore of Hudsons Bay for perhaps a hundred miles north of Eskimo Point, there are intergrades with arctica. As judged by their lesser size, individuals of this population are influenced by the sempl ei-stock. Otherwise, intergradation on the northern boundary, with arctica, is abrupt whereas intergradation at the south, between richardsonii and cicognanii, is gradual. Intergradation is similarly gradual between richardsonii on the one hand and bangsi and invicta on the other. By speaking of the intergradation as abrupt, it is in- tended, in this instance, to indicate that in a relatively narrow belt, between the geographic ranges of arctica and richardsonii, ermines intermediate in color-pattern, shape of skull, and size, bridge the gap between the ermine of the tundra (arctica) and that in the forest belt (richardsonii). It may be added that the degree of diflFerence between the two subspecies just mentioned is approximately twice as much as between richardsonii and cicog- nanii. The intergradation between cicognanii and richardsonii is gradual. By gradual it is meant that the change from one kind to the other is achieved in a wider area where ermines from locality A do not diflFer appreciably from those taken at, say, locality B, 50 miles farther south, although ermines from A and those from a third MUSTELA ERMINEA 117 locality, C, say, 130 miles south, clearly show differences indicative of geographic variation. Specimens examined. — Total number, 1035, as follows. Arranged alpha- betically by provinces and districts and from north to south in each province or district. Unless otherwise indicated, specimens are in the United States National Musevun. Alberta. 15 mi. above Smith Landing, 2; Fort Smith, 2 (1'); Smith Landing, 2; LaButte, Fitzgerald, 1'; Egg Lake, 15 mi. NW Ft. Chippewyan, 4 (2'); Lobstick Island, near Ft. Chippewyan, 1; Athabasca Delta, 9 mi. above mouth of main branch, 1; Athabasca Delta, Long Creek, 1 mi. W of main branch, 2; Ft. Chippewyan, 1; Peace Point, 1'; 18 mi. below Peace Point, 1; Em- barass River, 7 (4'); Athabasca River, 1'; Ft. McMurray, 1; Athabasca River, Middle Rapid, 2; 60 mi. above Grand Rapids, 1; Boiler Rapid, 1; Entrance, 3'; St. Albert, 2. British Columbia. Fort Halket, 1; Shesley, 1'; Dorothy Lake, Mts. W of Ft. Nelson, 4000 ft., 3'; valley between Musqwa and Prophet rivers, 3800 ft., SW of Ft. Nelson, 2^ Sikanni Chief Riv., 1; Telegraph Creek, 7 (6"); head of Bad River, 2350 ft., on lake, 1; Sbc Mile, 5*; Tuchodi Lake, 2°; Iskoot River, 2°; Level Mtn., 1'; head of Tatletuey Lake, 12 mi. W Thudade Lake, 2; Robb Lake District, 5"; Ft. Grahame, 12 (2'); Sustut Mts., on trib. Sustu Riv., 25 mi. SE Thudade Lake, 2; Laurier Pass, 1; Omineca Mts., 1"; Point Creek and Clearwater River, 2; Kispiox Valley, 23 mi. N Hazehon, 5'; Hazelton, 3'; NW arm Tacla Lake, 7; N end Babine Lake, 1; Pt. Simpson, 1; Metlakatla, 1; Stuart Lake, 27; S Fk. Sahnon Riv., 1'; mouth Sahnon Riv., 1'; Vanderhoof, 4'; Wistaria P. O., near Bums Lake, 1'; Kruger Lake, 9'; Indianpoint Lake, 23'; Quesnel, 1; Ahbau Lake, 3'; Isaacs Lake, 6*; Beaver Pass, 56'; Lightning Creek, 54'; LaFontaine, 16'; Barkerville, 1'; Barkerville District, 34'; Swift River, 27'; Cunningham Creek, 34'; Itcha Mts., 1'; Anahim Lake, 1'; Chezacut Lake, 8'; Kleena Kleene, 18'; 158 mi. House (Cariboo on labels), 3'; Rivers Inlet, 6 (5°; 1'); Horse Lake, 4=^ Kingcome Inlet, 8'; Loughborough Inlet, T; McGiUi- vary Creek, 1; Camel Back, Pemberton Meadows, V; Arrow Rapids, mainland opposite Stuart Island, 1'; Butte Inlet, 9'; Green Lake, 1'; Mt. Whistler, r°; Alta Lake, 2 (F; 1"); Mons, 1^ Keewatin. Foot of Baker Lake, 1. Labrador. Okak, 3=; Nain, 22 (11'; 11'); Hopedale, 24'; Kippokak Bay, 7'; Ailik, 1; Makkovik, 26^ Labrador, 55° N, 3; Hamilton Inlet, 2^ NW River Post, interior Labrador, 5"; Cartwright, 5; Paradise, 12; Sandwich Bay ( Muddy Bay, 6; North River, 6), 12; Battle Harbor, 1"; St. Marys River, 3''; Black Bay, 16 ( 15''; 1^'); Lanceau Loup, 17 (1^). Mackenzie. Ft. Frankhn, 1'; Ft. Rae, 12; Fairchild Point, 6"; Fort Simpson, 10 (2'); Hot Springs (61°, 125°), 1'; Willow River, near Ft. Providence, 1; 35 mi. N Big Island, 7; Big Island, 9; 3 mi. S Big Island, 7; Ft. Resolution, 9; 100 mi. N Ft. Smith, 2; 75 mi. NW Ft. Smith, 1; Ft. Liard, 2; Sucker Creek, 4'; Govt. Hay Camp, Wood Buffalo Park, 2\ Manitoba. Egg Is., Rabbit Point, 1; Ft. Churcliill, 1; Ft. York, W Hudsons Bay 57° N, 1^^ Oxford House, 11; Gypsumville, 1"; Lake St. Martin. New Brunswick. Restigouche Co.: Bird Bait, north Camp, 6 mi. NE Nictau Lake, 2''; Red Brook, Tobique River, 1". Victoria Co.: Trousers Lake, 3*. Glouchester Co.: Youghall, 1'; Miramichi Road, 15 mi. from Bathurst, 13". York Co.: Scotch Lake, 2. Newfoundland. Nicholsville, 3=; Bay St. George, 48 (26=; 2'=; 1"); Cod- roy, 9 (7=; 2»). Nova Scotia. Victoria County: Cape North, 2\ Inverness County: Fizzle- ton, 3\ Richmond County: St. Peters, 1\ Pictou County: Glengary, 1". Guysborough County: East Roman Valley, 5'. Kings Co.: Wolfville, 5 (3', 2'); near Wolfville, l\ Halifax Co.: Hammond Plains, 1. Annapolis Co.: AnnapoUs Royal, 1. Digby Co.: Digby, 3. No locality more definite than Nova Scotia, 3. 118 University of Kansas Publs., Mus. Nat. Hist, Ontario. Severn River, 1'; R. C. Mission, Yellow Creek, near mouth of Albany, 2'°; Ft. Albany, 4; Charlton Island, 1; Moose Factory, 10 (7"; 3'); Abitibi, 1". Quebec. Fort Chimo, 10'; Ungava Forks, 1; Belcher Islands, Hudsons Bay (Tukarak Island, 29; Eskimo Harbor, 2; InnetalUng Island, 1; S tip Gibson Peninsula, 2; Flaherty Island, 1), 35'*; Cairn Island, Richmond Gulf, 2"; Manitounuk Sound, 4"; about 15 mi. S Great Whale River, 1"; Ft. George, 1"; Charlton Island, 1"; Waswonaby Post, 1'; Mistassinnay Post, 3'; Godbout, 36; Mt. Albert, 7 (4'^ 3'); St. Anne River, 1500 ft., 1'; Ste. Anne des Monts, 3'; "Federal Mine", 1'; Berry Mountain Camp, 1'; Berry Mountain Brook, 1'; Cascapedia River (Middle Camp, 2; Tracadie, 2; Square Forks, 1), 5'. Saskatchewan. Poplar Point, Athabasca Lake, l""; Fair Point, Athabasca Lake, 1"; Emma Lake, 1*; Harper Lake, 2'; Livelong, 3"°; Fairholm, 2*; Touch- wood Hills, 2'^ Indian Head, 1'". Yukon. Hoole Canyon, 1; Teslin Lake (30 mi. N of , 1; Lake itself, 1; "near" the lake, 1; Mts. "near", 2; Snowden Mts., 2; Teslin Post, 2; Eagle Bay, 1; Morley Bay, 2; NisutUn River, 1; NisutUn Flats, 2; Wolf River, 1; Wolf Lake, 5), 2r. Mustela erminea cicognanii Bonaparte Ermine Plates 2, 3, 4, 9, 10 and 11 Mustela cigognanii [sic] Bonaparte, Charlesworth's Mag. Nat. Hist., 2:37, 1838. Putoritts vulgaris, Emmons, Quadrupeds of Massachusetts, p. 44, 1840. Mustela pusilla DeKay, Zool. of New York, Pt. 1, Mammalia, p. 34, pi. 14, fig. 1, 1842. Type from New York State. Putorius pusillus, Audubon and Bachman, Vivip. Quadrupeds of N. Amer., 2:100, pi. 64, 1851 (pi. 1846) and erroneously labled Mustela fusea, as pointed out on p. 102 of te.\t. Putorius cicognanii, Baird, Mamm. N. Amer., p. 161, 1858. Putorius richardsoni cicognani. Bangs, Proc. Biol. Soc. Washington, 10; 18, fie;s. 4, 4a of pis. 1 and 2, and pi. 3, figs. 2, 2a, February 25, 1896 (part). Putorius cicognani, Merriam, N. Amer. Fauna, 11:10, pi. 2, figs. 3, 3a, 4, 4a and pi. 5, figs. 2, 2a, June 30, 1896. Mustela cicognanii cicognanii. Miller, U. S. Nat. Mus. Bull., 79:95, Decem- ber 31, 1912; Bishop, Joum. Mamm., 4:26, February 9, 1923. Mustela cicognanii, Jackson, Joum. Mamm., 3:15, February 8, 1922. Mustela erminea cicognanii. Hall, Joum. Mamm., 26:77, February 27, 1945; Hall, Joum. Mamm., 26:180, July 19, 1945. Type. — No type specimen designated; type locality, eastern United States. The restriction of the type locahty from the general region of northeastem North America, as given by Merriam (1896:10) to the less inclusive area of the eastern United States as earlier given by Bangs (1896:18) is supported 1. Canadian National Museum 12. 2. Museum of Comparative Zoology 13. 3. American Mus. Nat. History 14. 4. Mus. Vert. Zool., Univ. California 15. 5. Collection of Alex Walker 17. 6. Provincial Museum of Brit. Columbia 18. 7. Collection of Kenneth Racey 19. 8. Field Museum of Natmal History 9. Univ. Washington Museum of Zoology 20. 10. Collection of Ian McTaggart-Cowan 21. 11. Acad. Nat. Sciences of Philadelphia British Museum of Natural History Museum of Zoology, Univ. Michigan Carnegie Museum Royal Ontario Museum of Zoology Donald R. Dickey Collection Berlin Zoological Museum Natm-historika Ricksmuseum, Sweden Collection of William T. Shav/ Collection of J. A. Munro MUSTELA ERMINEA 119 by Bonaparte's remarks in connection with the proposal of the name cicog- TUinii. He says (1838:37-38) "During my stay in the United States, I only saw a small species of Mustela, very common throughout the Union . . .". This animal constituted basis for the name cicognanii which name, he points out, is bestowed in order that the Americans "... should have con- stantly under their eye, this very common little animal, as the perpetual memorial . . ." to the Italian Governmental representative ". . . who, for upwards of fourteen years had served, in diplomatic and commercial concerns, . . . two countries, ... so diflFerent ... as the Roman and the United States . . . ." Clearly he had in mind principally, if not exclusively, the animal of tlie United States. Range. — Transition and higher life-zones of northeastern United States south to Connecticut, central Pennsylvania and extreme northeastern Ohio; in Quebec and Ontario westward from the latitude of central Maine to Lake Nipigon and Lake of the Woods. See figure 25 on page 95. Characters for ready recognition. — DiflFers from M. e. richardsonii of both sexes, in that least width of color of underparts averages less than a third rather than two-fifths of greatest width of color of upper parts, in males skull less, instead of more, than 1.9 grams and basilar length less than 38, in females by 16 per cent hghter skull (0.92 versus 1.1 grams); from M. e. hangsi, in males hind foot less instead of more than 40, hnear measurements of skull averaging 11 per cent less (depth of skull at plane of molars 10.0 versus 11.4), in females averaging smaller, hind foot 30 versus 32 and depth of skull at plane of molars 8.6 versus 9.1. Description. — Size. — Male. Seven adults and subadults from New York and Pennsylvania, yield average and extreme measurements as follows: Total length, 266 (240-295); length of tail, 74 (66-80); length of hind foot, 36 (33-39). Hamilton (1933:294) gives the weight of 31 adults from New York as 81 (66-105) grams. Female: Twelve adults and subadults from Maine and the area south to central Pennsylvania, yield average and extreme measurements as follows: Total length, 243 (225-260); length of tail, 63 (55-72); length of hind foot, 29.8 (26-32). Hamilton (1933:294) gives the weight of 15 adults from New York as 54 (45-71) grams. Color. — As described in Mustela erminea richardsonii except that under- parts in summer Marguerite Yellow or even more whitish; least width of color of underparts averaging, in adult males from New York and Pennsylvania, 29 (27-32) per cent of greatest width of color of upper parts. Black tip of tail in same series averaging 42 (30-51) mm. which is 57 per cent of length of tail- vertebrae. Skull. — Male ( illustrated by 4 adults in table of cranial measurements, which see): See plates 2-4. As described in Mustela erminea richardsonii except that: Weight, 1.5 (1.2-1.7) grams; basilar length, 35.7 (33.8-37.6). Female (illustrated by adult and subadults recorded in table of cranial measurements, which see): See plates 9-11. As described in Mustela erminea richardsonii except that: Weight of 2 subadults, 0.92 (0.86-0.98) grams; basilar length, 32.4 (31.4-33.3). 120 University of Kansas Publs., Mus. Nat. Hist. The skull of the male, in linear measurements, is approximately 13 (12-16) per cent smaller and 40 per cent lighter than in M, e. richardsonii. In relation to the basilar length, the skull averages slightly narrower, slightly shallower as measured in the vertical plane touching the posterior borders of the last upper molars, and the preorbital part is shghtly longer. In skulls of females of cicognanii, linear measurements average 3(0-6) per cent less, the weight is 16 per cent less and tlie teeth are 5 per cent shorter. In relation to the basilar length, measurements of the skull are approximately the same or slightly less in cicognanii. In comparison with bangsi, the male sex in linear measurements of the skull and teeth averages 11 per cent less than in bangsi from Aitkin, Minn., and 6 per cent less than in bangsi from Elk River, but in relation to the basilar length the preorbital region is larger. The weight is approximately a fourth less. In females the measure- ments average less, some being the same, and in relation to the basilar length, the bullae are shorter and the skull is shallower. The weight is about the same. Remarks. — In January, 1838, in Charlesworth's Magazine of Nat- ural History, C. L. Bonaparte proposed for three kinds of American weasels the names Mustela cicognanii, Mtistela richardsonii and Mustela longicauda. In this paper Bonaparte indicates that he previously had written (for his Iconografia della Fauna Italica . . .) an account of Mus- tela cicognanii using this same name. Fasciola XXII of the Iconogr. d. Fauna Italica, presenting his account of Mustela, like the Eng- lish paper was published in the year 1838. In his article in Charles- worth's Magazine, Bonaparte refers to his book published [used the past tense] in Rome but whether it actually appeared first I am unable to determine and hence am uncertain which of the two constitutes the original description. Reference to the Italian account suggests as basis for the name M. cicognanii, (1) specimens possibly seen in tlie United States by Bonaparte, or (2) Godman's published account of the animal. In the English publication, however, Bonaparte actually says that (1) he saw the small species in the Union [= United States]. Also, he (2) mentions his earlier written Italian account, (3) men- tions that "all the [American?] naturalists" used the name M. vul- garis for this animal, (4) incidentally mentions Godman's account, and (5) in naming two other American species cites accounts of them by Richardson. Also, Bonaparte in this English article makes MUSTELA ERMINEA 121 clear that when he wrote [not necessarily published] his Italian paper he did not know of the existence of two of the three American species. In the register of mammals at the British Museum of Natural History, tliere appears: 43.3.3.3 Mustek longicauda Bonap N amer presented by Dr. J. Richardson 4 Mustek Richardsonii Bonap " 5 " Cicognanii Bonap " To the right of these entries there appears, in three lines, the notation: "The three specimens examined by Prince Canino on which he established the three species" Every part of each of the above entries is in the hand writing of J. E. Gray, in charge of the collections from 1824 to 1840 and associated with them as Keeper until 1875, The three specimens are in good condition considering their age. The catalogue or register number shows, among other things, that they were entered in the register on March 3, 1843, Questions which might occur to anyone are: (1) Was there a type specimen of Mustela Cicognanii Bonaparte? If so is it no. 43.3.3.5? (2) If there was no type specimen was there a type locality? If so what is it? Among other things that may have bearing on these questions, are these: Bonaparte in Charlesworth's Magazine appears to base the two names Mustela Richardsonii and Mustela longicauda on Richardson's published account of Mustela erminea. At any rate immediately following each of the two names, Bonaparte writes "Nob. (M, erminea Rich, F, Bor, Amer,)," Bonaparte's other, first newly proposed name, Mustela Cicognanii, in Charlesworth's Magazine has following it only "Nob, North America", although in a paragraph above he did point out that this was the animal which all naturalists, at the time he was in America, considered as M. vulgaris. Turning to Richardson's account (Fauna Boreali Americana, , . , Quadrupeds, pp. 45-47, 1829) one finds that he recognized two species, M, vulgaris and M, erminea. Of the first he gives measurements "of an old female killed at Carlton House," Of the second species he distinguishes two varieties, the first represented by a specimen, of which he gives measurements, 'Tailed at Fort Franklin, Great Bear Lake" and, the second variety "of a larger size, having a longer tail and longer fore-claws" he indicates the 122 University of Kansas Publs., Mus. Nat. Hist. size of by giving measurements of a specimen taken "in the neigh- borhood of Carlton House." The last variety is clearly the basis of Bonaparte's M . longicauda. The specimen from which Richardson took his measurements I have been unable to locate [no. 43.3.3.3 in the British Museum, appears to be another specimen, although of the same subspecies and provided by Richardson]. The first variety of Richardson's Mustela erminea, clearly is the basis of Bonaparte's M. Richardsonii. The specimen from which Richardson took his measurements may well be no. 43.3.3.4 now preserved in the British Museum of Natural History, but I could not be certain about this. Richardson's M. vidgaris is accompanied by measurements of a female which I have ascertained to my full satisfaction is the identical specimen now bearing catalogue number 43.3.3.5 said by Gray to be the specimen on which Bonaparte based his name Mustela cicognanii. Gray probably saw his guest, Bonaparte, at work on these weasels and Gray's own written indication perhaps should be accepted at its face value. I found only 4 Richardson specimens of North American weasel in the British Museum in 1937 and it is con- ceivable that Bonaparte, 100 years before, actually had at hand only one specimen each of two kinds and 2 specimens of the third. This I think is not an important consideration, though, for Gray says just which specimens did serve as basis for Bonaparte's names and there is only one specimen for each name according to Gray. But I wonder if a type specimen can be made in this way? That is to say, after a name is published in a manner which makes it available, and if two or more specimens of the kind of animal in- volved, were, or may have been, available to the describer, can a person, even the author, himself, make a type specimen by saying that one particular specimen is beyond doubt the specimen on which a given name was established even though no particular specimen was designated in the original description? I incline to the view that a specimen so designated would at most be only a lectotype, unless it were a "cotype. However, if a holotype can be made by action such as Gray took, then (1) is no. 43.3.3.3 the type specimen of Mustela longicauda Bonaparte and, (2) is no. 43.3.3.4 the type specimen of Mustela Richardsonii Bonaparte? Incidentally, Mustela longicauda Bonaparte whether based on no. 43.3.3.3 or on Richardson's account will continue in its present MUSTELA ERMINEA 123 application. The same is true of Mustela richardsonii. If the basis of Mustela cicognanii Bonaparte [the diagnosis in the Iconografia d. Fauna Itahca . . . makes it clear that the name applies to the short-tailed species] was a weasel from the eastern United States or a description of a weasel or weasels from there, the name will continue in its present application. If, instead, the name is based on no. 43.3.3.3 (from Carlton House, Saskatchewan) or on Richard- son's account of M. vulgaris, the name will apply to a diflFerent subspecies (now called richardsonii and richardsonii will fall as a synonym of cicognanii) and the ermine of the eastern United States will take the next available name. Bonaparte probably named (in manuscript at least) cicognanii before he ever saw the specimen in the British Museum. This is indicated by his state- ment in Charlesworth's Magazine (1838:37) that "I have now [ItaHcs mine] found two [other] American species . . .". Whereas the names richardsonii and longicauda are based on Richardson, the name cicognanii, even if it dates from the account in Charles- worth's Magazine, appears to have a composite basis composed at the very least of (1) animals seen by Bonaparte in the United States, and (2) those called vulgaris by some other authors. Con- ceivably the specimen no. 43.3.3.3 in the British Museum, was part of the basis. From the nature of the case it can be argued that there could be no type and that if someone should bring to light a specimen in, say, Philadelphia, bearing the notation "this is the specimen seen in the United States by Bonaparte" it would im- mediately become as important as the one in London. Any Ameri- can weasel or weasels (then alive or preserved in a zoological col- lection) that Bonaparte saw in the United States probably were of the eastern United States. Bangs (1896:18-21), for one, previous to the present consideration of the name cicognanii, restricted it to the ermine of the eastern United States. Consequently, the name cicognanii, in the present account is applied to the ermine of the eastern United States. In my opinion there was and is no type. Almost certainly there was no type if the Fauna Italica appeared before the account in Charlesworth's Magazine did. Specimens examined. — Total number, 172, arranged alphabetically by prov- inces and states, then (except where indication is given to the contrary) by counties from north to south within each state or province. Unless otherwise indicated, specimens are in the U. S. National Museum. Connecticut. Windham Co.: S. Woodstock, Woodstock Lake, V. Hartford Co.: Windsor, 1'. New London Co.: Liberty Hill, 3'. Maine. Aroostook Co.: Quimby, 1^ Ashland 2\ Piscataquis Co.: table- land on top of Mt. Katahdin, 1; Chimney Pond, 3; T. 5, R. 13, 3^; "vicinity of Chesnucook," 1"; T. 4, R. 13, 1'; Moosehead Lake, T; Grenville, 10"; Barnard, 124 University of Kansas Publs., Mus. Nat. Hist. 3 (1"). Penobscot Co.: South Twin Lake, 1'; Lincoln, 11 (7*, 2°, 2'). Franklin Co.: Seven Pond Township, 7'. Oxford Co.: Umbago Lake, T; Upton, 4'; Bethel, 1'. Hancock Co.: Bucksport, 17'; Naskeag, 1. Lincoln Co.: Booth Bay, 1=. Massachusetts. Middlesex County: Wilmington, 2; Budington, 6 (1'); Worcester Co.: Cambridge, 5 ( 1', 3'); Sterling, V. Plymouth Co.: Middle- boro, 7 (1'). New Hampshire. Carroll Co.: Ossipee, 5. Rockingham Co.: Greenland, 1'. Cheshire Co.: Dublin, 1. New York. St. Lawrence County: Ogdensburg, 1'°. Franklin Co.: Malone, 1". Lewis Co.: Locust Grove, 1. Warren Co.: Lake George, 1. Mont- gomery Co.: Amsterdam, 1. Albany Co.: Albany, 1". Renesselaer County: Berlin, 2'; Schoharie, V. Thompkins Co.: Cascadilla Creek, Ithaca, 1". Allegany Co.: Ford Brook, Wellsville, 1". Ontario Co.: Phelps, 1'. Cattaraugus Co.: Cattaraugus, 1*. Ontario (locahties locally north to south, then west to east). Thunder Bay Dist.: Grand Bay, Lake Nipigon, 5'; Macdiamiid, 2"; Oscar, 2°; 20 mi. SW Fort Wilhams, 1'; Michipicoten Island, 3°. Algoma Dist.: Michipicoten, 1; Franz, 1"; Pancake Bay, 2". Parry Sound Dist.: French River, Georgia Bay, 1^; Seguin Falls, Tvi^). Montieth, 1". Sudbury Dist.: Casselmans, Rathbun Twp., 1*. Nipissing Dist.: Smoky Falls, near Kapuskasing, 4'; Franks Bay, Lake Nipissing, 1\ Haliburton Co.: Gooderham, 1''. Simcoe County: OrilUa, 1'; no locaHty more definite than county, T". Carelton Co.: Britannia, 5 mi. W Ottawa, 1"; Ottawa, 1"; Constant Bay, NE? of Ottawa, 1". Wellington Co.: Mt. Forest, 2"; Guelph, 1". Addington Co.: Buckshot Lake, Abinger Twp., P. Fontenac Co.: Clearlake, Arden, 1". Pennsylvania (by counties from west to east). Crawford Co.: North She- nango Township, Pymatuning Swamp, 2'*; Linesville (3 mi. NW, 1; 3)2 mi. W, 2; 3 mi. W, 1; 2 mi. SW, 1; 7)i mi. SW, 1) &\ Potter County: Cherry Springs Farm, Abbott Township, 1; 3 mi. S Inez, South Fork Simamahoning Creek, 1". Sullivan County: Lopez, 1". Lackawana Co.: Scranton, l\ Wayne County: Waymart, 1. Quebec (west to east). Labelle County: Kamika [z= Kiamika] Lake, 2"; Lacoste, 2"; Trout Lake, probably in this county, 2". Megantic Co.: Black Lake, 1". Rhode Island. Newport Co.: Middletown, 2". Vermont. Lamoille Co.: Mt. Mansfield, 1. Windsor Co.: Barnard, V. Mustela erminea bangsi Hall Ermine Plates 2, 3, 4, 9, 10 and 11 Mustela erminea bangsi Hall, Journ. Mamm., 26:176, July 19, 1945. [Putorius] cicognani, Meams, Bull. Amer. Mus. Nat. Hist., 3:235, June 5, 1891. Putorius richardsoni cicognani. Bangs, Proc. Biol. Soc. Washington, 10:18, February 25, 1896 (part). Putorius cicognanii, Cory, NIamm. Illinois and Wisconsin, p. 375, 1912. Mustela cicognanii, Aldous and Manweiler, Journ. Mamm., 23:250, August 13, 1942. 1. American Mus. Nat. History 9. University of Wisconsin 2. Boston Society of Natural History 10. Mus. Vert. Zobl., Univ. California 3. Museum of Comparative Zoology II. Canadian National Museum 4. Acad. Nat. Sciences of Philadelphia 12. Field Museum of Natural History 5. Collection of Alex Walker 13. Collection of Kenneth Racey 6. Collection of W. E. Snyder 14. Carnegie Museum 7. Museum of Zoology, Univ. Michigan 15. Cornell University 8. Royal Ontario Mtiseum of Zoology 16. Nevi' York State Museum MUSTELA ERMINEA 125 Mustela cicognanii cicognanii, Bailey, N. Amer. Fauna, 49:169, January 8, 1927; Leraas, Joum. Mamm., 23:344, August 13, 1942. Type. — Male, subadult, skull and skin; no. 11541, D. R. Dickey Coll.; Elk River, Sherburne County, Minnesota; November 1, 1925; obtained by Bernard Bailey, original no. A 606. The skull is complete and the teeth all are present and entire. The skin is well made and in a good state of preservation. Range. — Southern Manitoba, northeastern North Dakota, the whole of Mirmesota, Wisconsin and Michigan and northern Iowa. See figure 25 on page 95. Characters for ready recognition. — Differs from M. e. richardsonii, in that, in both sexes, least wadth of color of underparts averages about a third, in- stead of two-fifths, of greatest width of color of upper parts, and in that skulls of males are a fifth or more lighter, basilar length averaging less than 40; from M. e. cicognanii, in tliat hind foot more than 40 in males, averaging 32 versus 30 in females, and in larger skull, depth of skull at plane of molars being 11.4 versus 10.0 in males and 9.1 versus 8.6 in females. Description. — Size. — Male: Twelve adult and subadult males from Aitkin, Minnesota, yield average and extreme measurements as follows: Total length, 316 (291-341); length of tail, 87 (70-101); length of hind foot, 43 (40- 44). Two adidts from Aitkin each weigh 170 grams. Four adult and subadult females from Elk River and Fort Snelling, Minnesota, yield average and extreme measurements as follows: Total length, 249 (240-260); length of tail, 61 (55-65); length of hind foot, 32 (30-33). Color. — As described in Mustela erminea richardsonii except that, least width of color of underparts averaging, in males from Minnesota, 32 ( 19- 51) per cent of greatest widtli of color of upper parts. Black tip of tail in 12 male topotypes in white winter pelage averaging 52 (45-58) mm. which is 60 (53-66) per cent of length of tail-vertebrae. Skidl. — Male (based on adults from Aitkin): See measurements and plates 2-4. As described in Mustela erminea richardsonii except that: Weight of 2 adults from Aitkin, 2.2, 2.3 grams (9 subadults from T. 61 N, R. 26 W, average 1.95 grams); basilar length, 39.7 (38.5-40.7); length of tooth-rows rarely more (usually less) than length of tympanic bulla. Female (based on adults from Minnesota as listed in table of cranial measurements, which see): See plates 9-11. As described in Mustela erminea richardsonii except that: Weight, of a subadult from T. 61 N, R. 26 W, 0.91 grams; basilar length, 32.8 (31.8-33.6); breadth of rostrum rarely equal to as much as 30 per cent of basilar length. From richardsonii, topotypes of bangsi differ in that cranial measurements in males are approximately 7 (5-9) per cent less, linear measurements of teeth are 10 (9-11) per cent less and the skull is a fifth lighter. In relation to basilar length the tympanic bullae of bangsi are longer. Skulls of females are individually in- distinguishable, those of bangsi averaging approximately 1 per cent less in linear measurements. Comparison with the smaller cicognanii is made in the account of that subspecies. 126 University of Kansas Publs., Mus. Nat. Hist. Remarks. — Before the subspecific name bangsi was proposed, individuals of this subspecies ordinarily were recorded in the hterature as Mustela cicognanii. The best single lot of material is in the zoological collection of the University of Wisconsin. The late naturalist Albert Lano preserved a large share of the material from Minnesota. The large series from Elk River of that same state was mostly collected by Bernard Bailey although his Aunt, Anna (Bailey) Mills, and her brother the late Vernon Bailey, at an earlier time saved some specimens from Elk River. The name bangsi was proposed in recognition of the superior work done on American weasels by the late Outram Bangs, From the range of M. e. invicta in the Rocky Mountains, that of bangsi is separated by the Great Plains from a large part of which region the species is unknown. M. e. bangsi differs from invicta in greater degree of sexual dimorphism in size, and in each sex by larger size, narrower light-colored underparts, and deeper brain- case as measured at the anterior margin of the basioccipital. In bangsi the braincase is deeper relative to the length of the skull as well as, of course, actually deeper. Of the two subspecies whose ranges do meet that of bangsi, it more closely resembles richardsonii than cicognanii. From rich- ardsonii, especially from southeastern populations of the same in which the skull is of the same size as in bangsi, the latter differs in longer hind feet. This is an average difference and by one inter- pretation the animals here referred to bangsi might be lumped with some of the populations from the southeastern part of the range of richardsonii and the whole lot treated as intergrades between richardsonii and cicognanii. Nevertheless, the animals here re- ferred to bangsi are not geographically intermediate between rich- ardsonii and cicognanii and this consideration had much to do with the decision to recognize as a separate subspecies the animals here named bangsi. Within the range of the subspecies there is some geographic variation; the hind feet of animals from Iowa average slightly shorter than those of animals from Minnesota and Wisconsin but are nowhere nearly so short as in cicognanii at the same latitude in the eastern United States. It is noteworthy that the few specimens seen from Isle Royal have the long hind feet of bangsi and not the short hind feet of cicognanii which occurs all along the northern mainland. Because an oft cited record of occurrence even though erroneous, has a way of being repeated in later works, attention is here called MUSTELA ERMINEA 127 to the alleged occurrence of this ermine in northwestern Ohio at New Bremen. Henninger (1921:239) published the original ac- count of the supposed occurrence but as I pointed out in 1937 (p. 304), the specimen concerned proved upon examination to be a female of Mustela frenata noveboracensis. Henninger was mis- led probably by the short tail; the end of the tail had been lost and healed over before the animal's death. The present study has revealed that M. erminea everywhere east of the Cascade Moun- tains assumes a white winter coat. Had this been known when Henninger obtained his specimen he probably would not have wrongly identified the animal from New Bremen which was in the brown, winter pelage. Specimens examined. — Total number, 222, arranged alphabetically by prov- inces and states and, arranged from north to south, by counties in each state. Unless otherwise indicated, specimens are in the University of Wisconsin Museum of Zoology. Iowa. Dickinson County: W side Lake Okobogie, V. Winnebago County: Lake Mills, 7'. Worth County: Northwood, 1'. Clay County: "Dewey's Pasture, near Ruthven", 1'. Manitoba. Aweme, 4*; Red River Settlement, 1°. Michigan. Isle Royal: Tobin Harbor, P; Bell Isle, 1^ Washington Harbor, 3'. Ontonagon Co.: Ontonagon, 2 ( 1', 1'); T 51N, R 43W, sl7, Porcupine Mts., 1'. Gogebic County: Little Girls Point, 5'; Ironwood, V. Iron County: no locahty more definite than county, 1^. Luce Co.: Taquahniemenaw River Falls, V. Chippewa Co.: Sault Ste. Marie, 2\ Emmet County: Wilderness State Park, 2\ Cheboygan County: Univ. Mich. Biol. Station, 1'. Washtenaw Co.: Ann Arbor, 1^. Minnesota. Kittson County: no locahty more definite than county, 1'. Roseau County: Deer Township, 1°; Falun Township, 2°. Marshall Co.?: Moose River, 5°; Warren, definitely in Marshall County, 1°. Cook County: Grand Marais, 3 (2', V). St. Louis County: 2 mi E Babbit, 14"; Bumtside [z= Bumside] Lake, 1°. Itasca County: T 61N, R 26W, 23. Clay Co.: Moor- head, 3". Aitkin Co.: Aidcin, 13 (11", 1", 1"). OUer Tail Co.: Arthur, 3"; Ten Mile Lake, 1'; Parkers Prairie, 2". Chisago Co.: North Branch, 1". Sherburne Co.: Elk River, 42 ( 16^ 5^ 20', 1^°). Hennepen Co.: Lake Minnetonka, 1"; MinneapoHs, 1°; Fort Snelling, 5 (4", 1"). North Dakota. Pembina Co.: Walhalla, V. Nelson Co.: Stump Lake, 1°. Eddy Co.: Brantford, 2\ Wisconsin. Douglas County: T. 44N, R. 13W, 1; Gordon, 1. Bayfield Co.: Brinks Camp, Washburn, 1^ "near Cable", 1. Ashland County: Bear Lake, 2. Iron County: Fisher Lake, 4; Mercer, 5. Vilas Co.: Mamie Lake, 16°; Ox Bow Lake, 1''. Oneida Co.: Tomahawk Lake, 1". Langlade County: T. 34N, R. HE, 3. Rush Co.: Ladysmith, 1. Dunn County: Colfax, 2. Door County: Mink River, ElHson Bay, 1'. Dodge Co.: Fox Lake, 1"; Beaver Dam, 12". 1. Collection of T. C. Stephens 9. Univ. Kansas Mus. Nat. History 2. Iowa State College 10. Carnegie Museum 3. Museum of Zoology, Univ. Michigan 11. Field Museum of Natural History 4. Collection of Stuart Griddle 12. British Museum of Natural History 5. United States National Museum 13. Berlin Zoological Museum 6. Collection of Alex Walker 14. Museum of Comparative Zoology 7. Donald R. Dickey Collection 15. Mus. Vert. Zool., Univ. California 8. American Mus. Nat. History 16. Collection of W. E. Snyder 128 University of Kansas Publs., Mus. Nat. Hist. Mustela erminea invicta Hall Ermine Plates 2, 3, 4, 9, 10, 11 and 41 Mustela erminea invicta Hall, Joum. Mamm., 26:75, February 27, 1945; Hall, Joum. Mamm., 26:180, July 19, 1945. Putorius cicognanii, Preble, N. Amer. Fauna, 27:230, October 26, 1908. Type. — Male, subadult, skuU and skin; no. 101122, Mus. Vert. Zool.; Bene- wah, Benewah County, Idaho; October 24, 1926; obtained by William T. Shaw. The skull has a hole in the right squamosal bone on the floor of the braincase, and lacks the hamular process of the left pterygoid. The postmolar part of the right lower jaw is missing. The teeth all are present and entire. The skin is in white, winter pelage, well made, and in a good state of preservation. Range. — Central Rocky Mountain region from Jasper Park south over Al- berta, southeastern British Columbia, Washington east of the Cascades, and north and central Idaho and northwestern Montana. See figure 25 on page 95. Characters for ready recognition. — Differs from M. e. richardsonii, in males, by skull Hghter than 1.9 grams, mastoid breadth less than 19.9, depth of skull at anterior margin of basioccipital less than 12.4, in females by corresponding measurement of depth less than 10.1, and weight of skull less than one gram; from M. e. fallenda, in both sexes, by upper lips white ( not brown ) in males by skull averaging longer (37.0 versus 35.7), in females by breadth of rostrum less, instead of more, than 30 percent of basilar length; from M. e. streatori, gulosa, and muricus by hind foot more than 36 and basilar length more than 35 in males and by hind foot more than 29.5 and basilar length more than 30.5 in females; further distinguished from streatori by white (not brown) upper lips and from gulosa by black tip of tail more than half length of tail-vertebrae. Description. — Size. — Male: Ten adults and subadults from central Idaho County yield average and extreme measurements as follows: Total length, 291 (272-328); length of tail, 86 (75-100); length of hind foot, 39.9 (38-44). Female: Five adults and subadults from the same locality yield average and extreme measurements as follows: Total length, 255 (245-270); length of tail, 71 (68-76); length of hind foot, 32.3 (32-33). Color. — As described in Mustela erminea richardsonii except that under- parts in summer Marguerite Yellow or more whitish; least width of color of underparts averaging, in four females from Idaho and Montana, 38 ( 33-43 ) per cent of greatest width of color of upper parts. Black tip of tail in same speci- mens 38 (31-42) mm. which is 57 (52-65) per cent of length of tail-vertebrae. Skull. — Male ( 5 adults from Idaho County ) : See measurements and plates 2-4. As described in Mustela erminea richardsonii e.xcept that: Weight, 1.5 (1.4-1.7) grams; basilar length, 37.0 (35.8-39.8). Female (illustrated by adult and 4 subadults in table of cranial measure- ments, which see): See plates 9-11. As described in Mustela erminea richard- sonii except that: Weight, 0.72 (0.7-0.9) grams; basilar length, 32.2 (31.6- 32.8). From fallenda, invicta differs in that the skull of the male has a relatively narrower rostrum and relatively shallower braincase. Fe- males show the same differences but the degree of difference is about MUSTELA ERMINEA 129 as great again as in males. The teetli are almost exactly the same size in the two subspecies. The weight is the same in males but in females invicta is 18 per cent heavier. From streatori, invicta differs in that males average larger in every measurement taken except that the anteroposterior diameter of the inner moiety of Ml is less; 36 per cent heavier; linear measurements of the skull are about 5 per cent larger and those of the teeth, with the one exception noted, about 6 per cent larger; relative to the basilar length the tympanic bullae are longer and the rostrum is relatively narrower. In females, measurements of the skull average 8 per cent more and those of the teeth 7 per cent more except that, as in males, the inner lobe of Ml is actually shorter. Females of invicta are 12 per cent heavier; relative to tlie basilar length the skull is narrower throughout and the tooth-rows are shorter than in streatori. From gulosa, invicta differs in that males average larger (about 12 per cent) in every measurement taken, excepting the antero- posterior diameter of Ml which is the same; 50 per cent heavier; relative to the basilar lengdi the length of the tooth-rows and inter- orbital breadth are less. In females the inner lobe of Ml is smaller but every other measurement taken of the skull and teeth is more, invicta averaging about 8 per cent larger and 22 per cent heavier; relative to the basilar length, the tooth-rows are shorter and the skull is narrower interorbitally, through the rostrum and across the zygomata. From murica, invicta of corresponding sex differs in being larger in every measurement taken; males average 17 per cent larger in cranial measurements, 13 per cent larger in dental measurements and are 83 per cent heavier; corresponding percentages for females are 11, 9 and 20. Exception must again be made for the antero- posterior diameter of the inner lobe of the last upper molar which is less in females, and only slightly more in males. In males of invicta the tympanic bullae are longer in relation to the basilar length. From the geographically remote cicognanii, skulls of both males and females of invicta are to me individually indistinguishable. There is, nevertheless, an average difference not apparent to the eye between skulls of males. If the length of the tooth-rows be taken as a standard (100 per cent), the rostrum, of invicta, as measured across the lacrimal processes is broader (89 rather than 84 per cent ) but the width across the fourth upper premolars is less, 94 rather tlian 97 per cent of the length of die tooth-rows. 5—3758 130 University of Kansas Publs., Mus. Nat. Hist. Since the skull of invicta closely resembles that of cicognanii, it follows that invicta differs from richardsonii and bangsi in about the manner described in the account of cicognanii. Remarks. — Animals of this subspecies in advance of the present study generally were recorded in the literature under the name Mustela cicognanii. The difficulty in distinguishing individual specimens of invicta on morphological grounds from those of the geographically remote M. e. cicognanii should not be taken to in- dicate that the populations do not differ appreciably. Actually they differ in several characters although in no one of these is the degree of difference sufficient to allow of using it alone as a certain means of diagnosis. In invicta, as compared with cicognanii, the light-colored underparts are wider in relation to the dark-colored upper parts and the tail is longer by 4 per cent relative to the head and body. Given a population of each of the two subspecies, in which the skull is of the same mass, the hind feet are longer in invicta, there is more sexual dimorphism in size, and the anterior part of the skull differs in some particulars as just described in the comparison of the skull of invicta with other forms. Nevertheless, each of these differences is of an average sort. Therefore, and because over-all size is about the same in the two subspecies con- cerned, one or a few specimens from, say, central Idaho, can be distinguished from animals from western Pennsylvania only with difficulty, if at all. The close resemblance of skulls of invicta and cicognanii may be a function of their living at approximately the same latitudinal position in a climate that has marked seasonal variation. Intergradation with richardsonii is complete and gradual; in one sense invicta is but little more than a small richardsonii. Inter- gradation with fallenda is shown by several specimens. These two races differ in large degree in color, and in size and shape of the skull of females. Although the geographic area where intergrada- tion in color occurs is fairly wide, the area in which intergradation in cranial characters in females occurs, appears, from the in- adequate material available, to be much narrower. Intergradation occurs freely in Washington with streatori but with muricus so far as known only in the Bitterroot and nearby mountains of northwestern Montana. The Snake River Plains and low country along much of the Columbia River appears to be uninhabited by weasels of the species erminea and hence there is opportunity for intergradation only in the mentioned area of Montana. MUSTELA ERMINEA 131 Specimens examined. — Total number, 177, as follows. Arranged alpha- betically by provinces and states then by localities from north to south in each province and by counties from north to south in each state. Unless otherwise indicated, specimens are in the United States National Museum. Alberta. Jasper House, 4'; Shovel Pass, 2'; Jasper Park, 10'; head of Smoky River, 9; Henry House, 2 (1'); Blindman River, 1'; forks of Blindman River and Red Deer River, 2 ( 1», 1*); "near Red Deer, Red Deer River", 1'; Red Deer River, 2 ( 1% 1'); Red Deer, 2'; Rosebud, 2'; Prairie, 3000 ft., 1; Didsbury, Litde Red Deer River, 1; Canadian Nat'l Park, 1°; Canmore, 1; Banff, 1'; High River, 1°; "Waterton Lake Park" in Alberta, 6\ British Columbia. Grand Forks of Fraser River, 1; Canoe River, 1'; Field, 1; Glacier, 1'; E side Beaverfoot Range, 4000-4500 ft., 6 mi. SE Fraser Creek, 8'; Wentworth Lake, 1*; Revelstoke, 2 (1', 1'); Spillimach[e]en River, 2'; Sicamous, 2; Albert River, 7000 ft., 1'; Lumby, Creighton Valley, 1'; Okanagan, 4 (2\ V, V); Kettle River Lake, Gold Range, 4000 ft., 1; Crows Nest Station, 1'; Yale District, 3; Fort Hope, 1; Chilliwack Lake, 1'; Skagit, 2 (1', 1'); Skagit Valley, 1'; Skagit Summit, V; Lightning Lakes, 2 mi. N International Boundary, 3; Osoyoos-Bridesville Summit, 2; Westbridge, V; Rossland, 5'; Creston, mouth Goat Creek, 3'; Yahk, 4\ Idaho. Bonner Co.: Coolin, 4. Benewah County: Benewah, V. Idaho County: "Pete Kings, Locksa River", 1"; 2 mi. SSE Selway Falls, 1900 ft., 1"; 4 mi. SW Selway Falls, 5800 ft., 3"; Newsome Cr., 12 mi. above jet. with 5 Fk. Clearwater River, 2'; Iron Mt., to 14 mi. W thereof, 24'; Pilot Cr., % to 2J2 mi. above Newsome Cr., 4'; Sawmill Cr., VA mi. W Newsome, 1'; between Selway River and S. Fk. Clearwater R., 4'. Montana. Teton County (of old arrangement of counties): Many Glacier, 4900 ft., 1'; Duck Lake, 6 mi. NE St. Marys Lake, 1; St. Marys, Glacier Park, 1''; Lower St. Marys Lake, 1'. Flathead County: Stanton Lake, 5. County in question: Bitter Root Mts., 1. Ravalli County: Tin Cup District, 1'; Bass Creek, 6800 ft., NW of Stevensville, 1; Capitan Peak, 7000 ft., 1; Darby, 2'; Girds Creek, 1'; Charlos Heights, 2'. Washington. Whatcom County: Twin Lakes, Winchester Mts., 3 (1"); ChiUiwack River, 2600 ft., 2; Cooper Creek, near head, 4500 ft., Hannegan Pass, 1; Cooper Cr., 4300 ft., Hannegan Pass, 1"; Beaver Creek (2500 ft., and at McMillan Ranch, 1700 ft.), 2; Barron, Bonite Mine, 5000 ft., 1. Okanogan County: Tungsten Mine, 6800 ft., Bauerman Ridge, 4; Hidden Lakes, 4100 ft., 1; West Fork Pasayter River, 4700 ft., 1. Stevens County: Orin, 1". Fend Oreille County: lone, 2". Chelan County: Lake Chelan, 1". Mustela erminea alascensis (Merriam) Ermine Plates 2, 3, 4, 9, 10 and 11 Putorius richardsoni alascensis Merriam, N. Amer. Fauna, 11:12, pi. 2, figs. 2, 2a, June 30, 1896. Putorius cicognanii alascensis, Miller, U. S. Nat. Mus. Bull., 79:96, Decem- ber 31, 1912; Swarth, Univ. California Publ. Zool., 7:140, January 12, 1911. Mustela erminea alascensis. Hall, Proc. Biol. Soc. Washington, 57:36, June 28, 1944; Hall, Joum. Mamm., 26:180, July 19, 1945. 1. Canadian National Museum 9. Univ. Washington Museum of Zoology 2. American Mus. Nat. History 10. Collection of William T. Shaw 3. Field Museum of Natural History 11. University of Idaho 4. Museum of Comparative Zoology 12. California Academy of Sciences 5. Royal Ontario Museum of Zoology 13. Museum of Zoology, Univ. Michigan 6. Cornell University 14. Charles R. Conner Museum 7. Mus. Vert. Zool., Univ. California 15. Collection of Walter W. Dalquest 8. Collection of Kenneth Racey 16. Collection of Stanley G. Jevvfett 132 University of Kansas Publs., Mus. Nat. Hist. Type. — Male, adult, skull and skin; no. 74423, U. S. Nat. Mus., Biol. Surv. Coll.; Juneau, Alaska; August 22, 1895; obtained by Clark P. Streator, original no. 4806. The skull shows malformation of the frontal sinuses due to parasites and lacks osseous tissue where the parasitic infestation was localized. The left exoccipital condyle and adjacent region is less developed than the right and the posterior part of the skull is bent slightly to the left. Otherwise the skull is unbroken. The teeth all are present and entire. The skin is in the brown summer coat, fairly well made and in a good state of preservation. A few white hairs persist where the proximal line of the black hair of the tip of the tail meets the distal line of the brown hair. Range. — Mainland of southeastern Alaska from Lynn Canal south to include Mitkof, Zarembo, Wrangel and Revillagigedo islands. See figures 25, 26 on pages 95 and 134. Characters for ready recognition. — Differs from M. e. arctica and haidarum, in both sexes, by proximal two-thirds of under side of tail colored same as upper parts rather than same as underparts and interorbital breadth less, in- stead of more, than distance between glenoid fossa and posterior border of external auditory meatus; from A/, e. salva, in males, by overall depth of brain- case including tympanic bullae less than 89 per cent of orbitonasal length, fe- males not individually distinguishable but averaging shallower through the braincase; from M. e. initis, celenda and seclusa by interorbital breadth less than distance between glenoid fossa and posterior border of external auditory meatus (females of initis, celenda and seclusa unknown); further from initis by total length less than 317 and black tip of tail less than 57 per cent of length of tail-vertebrae; further from celenda by chest white, not mostly covered by brown patch. Description. — Size. — Male: Eight adults from Windham, Alaska, yield aver- age and extreme measurements as follows: Total length, 298 (288—315); length of tail, 88 (84-94); length of hind foot, 41.3 (37-44). Female: Two adults from Juneau and Helm Bay measure, respectively, as follows: Total length, 258, 258; length of tail, , 76; length of hind foot, 32, 34. Color. — As described in Mustela crminea richardsonii except that least width of color of underparts averaging, in five females, 42 (35-53) per cent of greatest width of color of upper parts. Black tip of tail in same specimens averaging 36 (30-40) mm. which is 49 (48-53) per cent of length of tail- vertebrae. Skull. — Male ( based on 8 adults from Windham ) : See measurements and plates, 2-4. As described in Mustela erminea ricliardsonii except that: Weight, 1.8 (1.5-2.6) grams; basilar length, 37.5 (36.5-38.9); length of tooth-rows more or less than (about same as) length of tympanic bulla. Female (based on 5 adults, from locafities listed in the table of cranial measurements): See measurements and plates 9-11. As described in Mus- tela erminea richardsonii except that: Weight, 0.96 (0.7-1.1) grams; basilar length, 32.7 (31.9-33.2); breadth of rostrum more or less than (about equal to) 30 per cent of basilar length. From richardsonii, alascensis differs in that the skull of the male averages smaller in every measurement taken and is 28 per cent MUSTELA ERMINEA 133 lighter. Relative to the basilar length, the orbitonasal length is more and the braincase is shallower as measured at the anterior end of the basioccipital. The four adult females seen of alascensis are more variable than those of richardsonii and average smaller in some measurements and larger in others but give no proof of any consistent difference. From haidarum, alascensis differs in that the rostrum and entire preorbital part of the skull is actually as well as relatively much smaller in both sexes. In males of alascensis the length of the skull, and other cranial measurements of length, is more. In males, the mastoid breadth and zygomatic breadth are about the same as in haidarum^, as also is the weight. Ml is larger but ml and P4 are smaller. In females the anteroposterior extent of the inner moiety of Ml and length of tympanic bulla are about the same in the two subspecies but all other cranial and dental measurements in alascensis are less. It is 29 per cent lighter. The difference in the preorbital region is of about the same degree as in the males. Comparisons of the skuU with those of arctica, salva, initis, ce- lenda, and seclusa are made in the accounts of those subspecies. Remarks. — The relatively few specimens known of this race seem always to have been referred to in the literature by the name alascensis and the nomenclatural history is therefore simple. The original materials were obtained by the collector Clark P. Streator and the additional series of skeletons, one with skin, from Windham were procured by Stanton Price, a resident there. The subspecies is well differentiated from both arctica and rich- ardsonii. Although actual intergrades are lacking between alas- censis and the two races just mentioned I have no doubt that intergradation occurs with richardsonii and think it probably does also with arctica. The assignment of the three females from Mitkof Island, Zarembo Island, and Loring on Revillagigedo Island, is tentative because each is so young as not to show diagnostic cranial charac- ters. The two other specimens from Revillagigedo Island (Carroll Inlet), labeled as males, are in white winter pelage. Only one, no. 136358, a subadult, is accompanied by a skull. The small size of each specimen, and its cranial characters which are inter- mediate between those of males and females of alascensis of the adjacent mainland, indicate the existence of a distinct race of weasel on Revillagigedo Island. On the chance that the one specimen with a skull is a dwarf, or is wrongly sexed as seems improbable, the population is tentatively referred to alascensis. 134 University of Kansas Publs., Mus. Nat. Hist. :" = ": = g I ::::::_j ? I specimens • examined 3 M.e.arctica 4 M. e . richardsonii JM.e. olascensis 5 ^m M. e.inifis M. e.salva 6 k-:-:':":":-:-:-! M . e . relgnda 7 iililiiliij M. e .seclusa Fig. 26. Map showing known occurrences and probable geographic ranges of the subspecies of Mustela erminea in southeastern Alaska. MUSTELA ERMINEA 135 Specimens examined. — Total number, 24, arranged by localities from north to south. Unless otherwise indicated, specimens are in the Museum of Verte- brate Zoology, University of California. Alaska. Juneau, 5'; Talcu River, 1; Windham, 9; Mitkof Island, 1; St. John Harbor, Zarembo Island, 1; Wrangel, 1'; Helm Bay, Cleveland Peninsula, 1; Cleveland Peninsula, 2'; Revillagigedo Island, Carroll Inlet, 2'; Loring, 1'. Mustela erminea salva Hall Ennine Plates 2, 3, 4, 9, 10 and 11 Mustela erminea salva Hall, Proc. Biol. Soc. Washington, 57:35, June 28, 1944; Hall, Joum. Mamm., 26:180, July 19, 1945. Type. — Male, adult, skull only; no. 74641, Mus. Vert. Zool.; Mole Harbor, Admiralty Island, Alaska, December 27, 1936; obtained by A. Hasselborg. The skull (plates 2-4) shoM^s malformation of the frontal sinuses owing to parasites and lacks osseous tissue where the parasitic infestation was localized. The skull is unbroken. The teeth all are present and entire. Range. — Admiralty Island, Alaska. See figures 25, 26 on pages 95, 134. Characters for ready recognition (known only from skulls). — Differs from males of M. e. alascensis in overall depth of braincase which is more than 89 per cent of orbitonasal length; from M. e. initis, in males, in that orbitonasal length and mastoid breadth total less than 35 mm., weight of skull and lower jaws less than 2.1 grams; from M. e. celenda, in males, in that breadth of rostrvun measured across lacrimal processes less than a third of basilar length. Description- — Size. — Male: An adult from Gambier Bay measures: Total length, 320; length of tail, 95; length of hind foot, 45 (41 in dry skin). Female: A subadult from Hawk Inlet, measures: Total length, 250; length of tail, 70; length of hind foot, 33. Color. — As described in Mustela erminea richardsonii except that least width of color of underparts in four individuals 40 (38-43) per cent of greatest width of color of upper parts. Black tip of tail, in two individuals for which external measurements are given, amounting to 50 and 40 mm. respectively which is 53 and 57 per cent of length of tail-vertebrae. Skull. — Male (type and 4 adult topotypes): See measurements and plates 2-4. As described in Mustela erminea richardsonii except that: Weight, 1.7 (1.5-1.9) grams; basilar length, 37.8 (36.4-39.5, extremes are in subadults); length of tooth-rows more or less (usually more) than length of tympanic bulla; interorbital breadth rarely more than distance between glenoid fossa and posterior border of external auditory meatus. Female (2 ad. and 1 ad.-sad. topotypes): See measurements, and plates 9-11. As described in Mustela erminea richardsonii except that: Weight, 0.9 (0.8-1.0) grams; basilar length, 33.0 (32.0-33.6); length of tooth rows approximately same as length of tympanic bulla; breadth of rostrum approxi- mately 30 per cent of basilar length. From alascensis, salva differs in that males have the preorbital region slightly wider in relation to the length of the tympanic bulla; 1. United States National Museum. 136 University of Kansas Publs., Mus. Nat. Hist. also the braincase is smaller, actually as well as in comparison with the preorbital part of the skull. The tympanic bullae do not project so far below the squamosals and the braincase itself is shallower, in adults averaging only 11.5 mm. as against 12.5 mm. The overall depth of the braincase, including the tympanic bullae, when divided into the orbitonasal length gives an average of 93 ( 90-97 ) per cent whereas in alascensis the figure is only 85 (78-88) per cent. On this basis alone, everyone of the adult skulls of the two races can be distinguished. The females and subadult males show the same tendency to reduction in depth of braincase but not every individual among them can be surely distinguished. By weight the skull of salva of corresponding sex is only about 6 per cent smaller. Com- parisons with initis and celenda are made in the accounts of those subspecies. Remarks. — Most of the specimens seen were collected by Allen E. Hasselborg, resident on Admiralty Island. On the basis of skulls- few skins, and measurements taken in the flesh, are available — saloa more closely resembles alascensis than does any other subspecies so far known from southeastern Alaska. The race on Admiralty Island is only slightly differentiated from alascensis of the adjacent mainland. Specimens examined. — Total number, 26, all from Admiralty Island, Alaska, arranged in general by localities from north to south, and unless otherwise indicated in the Museum of Vertebrate Zoology, University of California. Alaska. Admiralty Island: Hawk Inlet, 2; Seymour Canal, 4; Mole Harbor, 18 (skulls only); Gambier Bay, 1; no locality more definite than Admiralty Island, 4 (1 in U. S. Nat. Mus.). Mustela erminea initis Hall Ermine Plates 4, 5 and 6 Mustela erminea initis Hall, Proc. Biol. Soc. Washington, 57:37, June 28, 1944; Hall, Joum. Mamm., 26:180, July 19, 1945. Type. — Male, adult, skull and skin; no. 289, Mus. Vert. Zool.; Saook Bay, Baranof Island, Alaska; October 9, 1907; obtained by A. Hasselborg, original no. 4. The top of the skull is fractured on the left side from the anterior nares posteriorly through the postorbital process to the posterior root of the zygomatic arch. On the left lower jaw the canine and three incisors are missing; otherwise tlie teeth all are present and entire. The skin is in process of molt, approximately nine-tenths of the incoming white pelage being in place. The skin is well made and in a good state of preservation. Range. — Chichagof and Baranof islands, Alaska. See figures 25, 26 on pages 95, 134. MUSTELA ERMINEA 137 Characters for ready recognition (only males known). — Differs from M. e. arctica, in that proximal two-thirds of under side of tail colored same as upper parts rather than same as underparts, zygomatic breadth less than distance between last upper molar and jugular foramen; from M. e. salva in that orbitonasal length and mastoid breadth total more than 35 mm., weight of skull and lower jaws more than 2.1 grams; from M. e. alascensis, by total length more than 317, black tip of tail more than 57 per cent of length of tail-vertebrae, interorbital breadth more than 10.3 and equal to, instead of less than, distance between glenoid fossa and posterior border of external auditory meatus; from M. e. celenda by chest white (not mostly covered by brown patch), breadth of rostrum measured across lacrimal processes less than a third of basilar length; from M. e. seclusa in zygomatic breadth more than distance between last upper molar and jugular foramen. Description. — Size. — Male: The type and an adult topotype measure, respectively, as follows: Total length, 330, 320; length of tail, 95, 95; length of hind foot, 45, 45. Female: No external measurements available. Color — As described in Mustela erminea richardsonii except that least width of color of underparts averages, in two young female topotypes, 50 (49, 50) per cent of greatest width of color of upper parts. Black tip of tail in three young female topotypes averaging 54 ( 52-55 ) mm. which is 67 ( 63-69 ) per cent of length of tail-vertebrae. Skull. — Male ( illustrated by type and 1 ad. topotype ) : See measurements and plates 4-6. As described in Mustela erminea richardsonii except that: Weight, 2.3 and 2.5 grams; basilar length, 39.6, and 40.5; interorbital breadth equal to distance between glenoid fossa and posterior border of external auditory meatus. Female: No adults available. From salva, initis differs in that skulls of males average larger in every measurement taken, being 41 per cent heavier. Relative to the basilar length, the interorbital and preorbital parts of the skull are larger; the relatively greater interorbital and mastoid breadths are particularly noticeable. Although the depth of the braincase, including the tympanic bullae, is both relatively as well as actually more than in salva, the depth is relatively less than in alascensis which otherwise differs from initis in about the same way that salva differs from initis. Whereas the interorbital breadth in initis is about equal to the distance between the glenoid fossa and the posterior border of the external auditory meatus, the interorbital breadth is uniformly less than this distance in both salva and alascensis. In comparison with seclusa the teeth are of the same size but all measurements of the skull are larger. The skull of initis is 25 per cent heavier. In relation to the basilar length, the interorbital and preorbital parts of the skull are much less in initis. The preorbital and interorbital regions in initis are relatively smaller in comparison 138 University of Kansas Publs., Mus. Nat. Hist. also with arctica. The one measurement of interorbital breadth in initis is greater in relation to the basilar length than in kadiacensis but the rostral region, and all that part of the skull anterior to the braincase, is relatively smaller in initis. Remarks. — The two adult males, nos. 286 and 289 from Saook Bay, provide convincing evidence of the existence of a distinct race of weasel on Baranof Island. Three other young specimens, almost subadult, from the same place are labeled as males although the basilar lengths of these skulls are only 35.5, 35.9 and 37.3 milh- meters as against 39.6 and 40.5 in the two adult males. The dif- ference in size is too great to be age-variation. The fact that 3 are definitely of one category and 2 of the other makes it doubtful that individual variation accounts for the differences. The small size of these 3 specimens and the fact that in each the anterior margin of the tympanic bulla is flush with the squamosal rather than pro- truded from the braincase, suggests that the three are females. If they are females, the amount of secondary sexual variation is rather less than would be expected by analogy with the amount obtaining in alascensis on the mainland and in salva on Admiralty Island. Another possibility that I can not disprove is that two stocks of weasels persist on Baranof Island, the two larger specimens being descendants of the stock which first became established on the is- land and the three smaller specimens being descendants of an indi- vidual ermine, or of ermines, that were rafted or otherwise trans- ported to the island at a considerably later date. Assuming for the moment that there are two stocks, it must be admitted that each one differs from any stock known from elsewhere. Therefore, each stock would be presumed to have been long resident on the island. But — two stocks as closely related as the two in question would not be expected to persist for long in an area as small as that of Baranof Island because competition would give one the ascendency. There- fore, the first suggestion, namely that the three smaller animals are really females, seems the more probable. The feasible way to clear up the present uncertainty is, of course, to obtain additional speci- mens, carefully labeled as to sex. Yet another reason why addi- tional collecting is desirable in this area is to ascertain whether there is subspecific differentiation between the ermines of Baranof and Chichagof islands. The one specimen available from the latter island, although in general like the three smaller animals from Baranof Island, differs in the fuller (less scooped out) medial side of the tympanic bulla and to a slight degree in each of some other MUSTELA EKMINEA 139 features. This specimen from Chichagof Island is labeled as a male also. Specimeris examined. — Total number, 6, arranged by localities from north to south, and in the Museum of Vertebrate Zoology, University of California. Alaska. Chichagof Island, Freshwater Bay, 1. Baranof Island, Saook Bay, 5. Mustela erminea celenda Hall Ermine Plates 5, 6 and 7 Mustela erminea celenda Hall, Proc. Biol. Soc. Washington, 57:38, June 28, 1944; Hall, Joum. Mamm., 26:181, July 19, 1945. Type. — Male, adult, skull and skin; no. 130987, U. S. Nat. Mus., Biol. Surv'. Coll.; Kasaan Bay, Prince of Wales Island, Alaska; June 16, 1903; obtained by Cyrus Catt; original no. 4407X. The skull has a piece 1.5 mm. long broken out of the left zygomatic arch. P2 is absent on both sides. The right II, and the left II and 12 are missing. The skin, in summer pelage, is fairly well-made. A scrotal pouch attests to the correctness of the sex recorded on the label. The rostral part of the skull is smaller than in average-sized males of corresponding age. Range. — Prince of Wales, Dall, and Long islands, Alaska. See figures 25, 26 on pages 95, 134. Characters for ready recognition (only males known). — Differs from M. e. alascensis and initis in chest mostly covered by brown patch, not white, and breadth of rostrum measured across lacrimal processes more than a tliird of basilar length, which cranial character serves to distinguish also salva; from M. e. seclusa in zygomatic breadth less than distance between last upper molar and jugular foramen; from M. e. haidarum in chest white (not mostly covered by brown patch), proximal two-thirds of underside of tail colored Uke upper parts ratlier than underparts, basilar length more than 38.2 mm. Description. — Size. — Male: Seven adults and subadults from Prince of Wales Island, yield average and extreme measurements as follows: Total length, 286 (277-304); length of tail, 77 (74-85); length of hind foot, 36 (35.5-40.5). Female: No specimen available. Color. — As described in Mustela erminea richardsonii except that upper parts about tone 3 of dark Chocolate Brown of Oberthiir and Dauthenay, pi. 342; lower throat and chest covered by a large patch of same color as upper parts; color of iinderparts extending to toes but in interrupted fashion on both fore- and hind-feet; least width of color of underparts averaging, in four males from Prince of Wales Island, 41 (38-49) per cent of greatest width of color of upper parts. Black tip of tail averaging, in 8 males in vmiter pelage, 65 (59-78) mm. which is 84 (69-92) per cent of length of tail-vertebrae. From its geographic neighbors alascensis and initis, celenda differs in darker color of upper parts, presence rather than absence of patch of dark color on lower throat and chest, and longer black tip on tail. From haidarum, celenda differs in darker color of upper parts, presence rather than absence 140 University of Kansas Publs., Mus. Nat. Hist. of patch of dark color on lower throat and chest, narrower hght-colored under parts, black tip of tail averaging less rather than more than nine-tenths of length of tail-vertebrae and ventral face of tail colored like upper parts rather than Uke underparts. Skull. — Male (illustrated by 5 adults): See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 2.3 (2.2- 2.5) grams; basilar length, 39.5 (38.9-40.7) mm.; length of tooth-rows more than length of tympanic bulla; breadth of rostrum measured across lacrimal processes more than a third of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more or less than (about equal to) distance be- tween last upper molar and jugular foramen. Female. — Complete skull of adult unavailable. Differences from richardsonii are indicated in the formal de- scription just given. Additional to differences therein noted, celenda differs from initis in larger interorbital and preorbital parts of the skull although dimensions of other parts of the skull and the teeth are about the same or even less. From salva, celenda differs in larger average size in every measurement taken, except for the inner moiety of Ml which is about the same. The skull of celenda is 35 per cent heavier. In relation to the basilar length the skull of celenda is wider, especially in the interorbital and preorbital regions. In comparison with alascensis the tympanic bullae are of approximately the same length; otherwise essentially the same differences obtain as are noted in comparison with salva and the zygomatic breadth is relatively more in celenda. From seclusa, in which the teeth are of comparable size, celenda differs in that every cranial measurement is more and the skull is 28 per cent heavier. Because the skull of celenda is so much longer, its dimen- sions in other planes are less in relation to the length than in seclusa. M. e. celenda is larger in every part measured than haidarum, 21 per cent heavier, and in relation to the basilar length the interorbital, and preorbital, parts of the skull are smaller, the braincase is shallower, and the skull is relatively wider across the zygomata and mastoid processes. In comparison with kadia- censis, differences are: 26 per cent lighter, skull shorter; in re- lation to the basilar length, braincase shallower as measured at the anterior end of the basioccipital, tooth-rows shorter but orbi- tonasal length more. In comparison with arctica all parts measured of the teeth and skull of celenda are smaller and its skull is 34 per cent lighter. In relation to the basilar length, the interorbital breadth of celenda is only slightly less but its skull is narrower MUSTELA ERMINEA 141 across the rostrum and zygomata, the tooth-rows are shorter, and the braincase is shallower. Remarks. — The late George Willett in the course of his work in Alaska collected most of the known specimens of this strongly differentiated subspecies. In both coloration and cranial characters the distinguishing features are so well marked that the zoologist could with reason accord full specific rank to celenda. Neverthe- less it obviously is an ermine. Also, races from other islands of southeastern Alaska tend to bridge the gap, as regards cranial features, between celenda and the mainland ermine. The specimen from Dall Island agrees in all respects with topotypes. The speci- men from Howkan on Long Island is in white winter pelage and the skull has suffered shrinkage from some chemical solution; the reference of this specimen to celenda is tentative. Specimens examined. — Total number, 25, as follows: Arranged by locali- ties from north to south. Unless otherwise indicated, in U. S. National Museum. Alaska. Prince of Wales Island: Craig, 18 (10 in Mus. Vert. Zool., and 8 in Los Angeles Mus. Hist. Art and Sci. ) ; Kasaan Bay, 2; no locality more def- inite than the Island itself, 3; Dall Island, Otter Harbor, 1 (Los Angeles Mus. Hist. Art and Sci.). Long Island, Howkan, 1 (Field Mus. Nat. Hist.). Mustela erminea seclusa Hall Ermine Plates 5, 6 and 7 Mustela erminea seclusa Hall, Proc. Biol. Soc. Washington, 57:39, June 28, 1944; Hall, Joum. Mamm., 26:181, July 19, 1945. Type. — Male, adult, skull alone; no. 31232, Mus. Vert. Zool.; Port Santa Cruz, Suemez Island, Alaska; March 24, 1920; obtained by George Willett. The skull (plates 5-7) is complete and unbroken. Of tlie upper incisors only right 13 is present. Otherwise the teeth are present and unbroken. Range. — Known only from the type locality. See figures 25, 26 on pages 95, 134. Characters for ready recognition (only the male known). — DiflFers from M. e. celenda in basilar length less than 38.2, from M. e. salva, initis and haidarum in zygomatic breadth more than distance between last upper molar and jugular foramen. Description. — Size and Color. — No external measurements or skins available. Skull. — Male: See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 1.8 grams; basilar length, 34.3; length of tooth-rows about the same as length of tympanic bulla; breadth of rostrum measured across lacrimal processes more than a third of basilar length; interorbital breadth more tlian distance between glenoid fossa and posterior margin of external auditory meatus; z\'gomatic breadth more than distance between last upper molar and jugular foramen. Female. — Skull not available. 142 University of Kansas Publs., Mus. Nat. Hist. From alascensis and salva, seclusa differs in larger teeth, shorter skull, much larger preorbital and interorbital regions, actually as well as in relation to basilar length. Excepting the teeth, which are of about the same size, the same general differences obtain in comparison with initis which, however, is 29 per cent heavier. From celenda, seclusa differs in smaller skull in all parts measured, being 22 per cent lighter. The teeth are about the same size. In relation to its length the skull of seclusa is much broader and deeper. From haidarum, seclusa differs in: teeth larger; skull shorter and more convex in dorsal outline along median longitudinal axis; in relation to basilar length, skull broader, deeper and braincase relatively shorter. Remarks. — The characters shown in the one available skull are far outside the limits of individual variation for other known sub- species. Other specimens are much to be desired to ascertain what the "average" individual is like and to learn the characters of the female. Specimen examined. — One, the holotype. Mustela erminea haidarum (Preble) Ermine Plates 5, 6, 7, 11, 12 and 13 Putorius haidnrum Preble, Proc. Biol. Soc. Washington, 12:169, August 10, 1898. Mustela haidarum, Miller, U. S. Nat. Mus. Bull., 79:97, December 31, 1912. Mustela erminea haidarum. Hall, Proc. Biol. Soc. Washington, 57:38, June 28, 1944; Hall, Joum. Mamm., 26:181, July 19, 1945. Type. — Male, adult, skull, skeleton and skin; no. 94430, U. S. Nat. Mus., Biol. Surv. Coll.; Massett, Queen Charlotte Islands, British Columbia; March 17, 1898; obtained by J. H. Keen; original no. 1800x. The skull is unbroken and complete except for osseous tissue destroyed in the region of each postorbital process; this is the result of infestation of the frontal sinuses by parasites. The skeleton is complete down to the distal ends of the tibiae; the more distal bones are in the skin. The first, right, upper incisor is missing. Otherwise the teeth all are present and entire. The skin is in the white, winter pelage but the new under fur is visible along the back and on the head although mostly covered with white hair. Range. — Queen Charlotte Islands. See figure 25, page 95. Characters for ready recognition. — DifiFers from M. e. celenda in chest white (not mostly covered by brown patch), proximal two-thirds of under side of tail colored Uke underparts instead of upper parts, in males basilar length less than 38.2; from M. e. seclusa, in male, in zygomatic breadtli less than distance between last upper molar and jugular foramen; from M. e. richardsonii and alascensis, in both sexes, in proximal two-thirds of under MUSTELA ERMINEA 143 side of tail colored like underparts instead of upper parts, interorbital breadth not less than distance from glenoid fossa to posterior margin of external auditory meatus; from M. e. anguinae and fallenda, in both sexes, in light- colored underparts more than half the width of dark-colored upper parts, proximal two-thirds of under surface of tail colored like underparts instead of upper parts, interorbital breadth equal to or more than distance between glenoid fossa and posterior margin of external auditory meatus. Description. — Size. — Male: Two adults, U. S. N. M., no. 100622, from Cumsheva Inlet, and Amer. Mus. N. H., no. 37411, and the type, measure, respectively, as follows: Total length, 283, 290, 275; length of tail, 70, 75, 60; length of hind foot, 39, 40, 37. Female: Corresponding measurements of an adult, no. 100624, and a young individual, no. 100623, each from Cumsheva Inlet, are: 252, 250; 63, 61; 31, 32. Color. — As described in Mustela erminea richardsonii except that under- parts not Sulphur Yellow but ranging from near (e) Colonial Buff through Marguerite Yellow to almost pure white; color of underparts extends distally on posterior sides of forelegs and onto toes but in many specimens interrupted at wrist by color of upper parts; color of underparts extends onto proximal three-fourths of under side of tail as length of tail is measured along tail- vertebrae; least width of color of underparts averaging, in 5 males, 79 (66-130) per cent of greatest width of color of upper parts. Black tip of tail in same males averaging 62 (60-70) mm. which is 92 (83-115) per cent of length of tail-vertebrae. The close correspondence in color-pattern of this weasel with the Arctic races, arctica, polaris, semplei and kadiacensis is noteworthy, and distinguishes it from weasels on the adjacent mainland and adjoining islands to the north and south. The color of the upper parts is darker than in the four Arctic races named. Skull. — Male ( 7 adults ) : See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 1.9 (1.7-2.0) grams; basilar length, 36.7 (35.6-37.5); length of tooth-rows more than length of tympanic bullae; breadth of rostrum measured across lacrimal processes more than a third of basilar length; interorbital breadth more than distance between glenoid fossa and posterior margin of external auditory meatus; zygomatic breadth barely less than distance between last upper molar and jugular foramen. Female (2 adults): See measurements and plates 11-13. As described in Mustela erminea ricliardsonii except that: Weight, 1.3 and 1.4 grams; basilar length, 34.2; length of tooth-rows more or less than (about equal to) length of tympanic bulla; breadth of rostrum more than 30 per cent of basilar length; interorbital breadth not less than distance between glenoid fossa and posterior margin of external auditory meatus. From richardsonii, haidarum differs in that skull of the male is actually larger in its anterior part (breadth of rostrum, interorbital breadth and orbitonasal length ) but all measurements of other parts average less. In relation to the basilar length, the tympanic bulla is shorter but all other measurements are more. In the skull of the fe- 144 University of Kansas Publs., Mus. Nat. Hist. male, which is 23 per cent heavier, the v^idth of the tympanic bulla and anteroposterior extent of the inner lobe of Ml are the same; in all other measurements the female of haidarum is larger, and in relation to the basilar length all measurements are more except the depth of the skull at the anterior margin of the basioccipital and the width of the tympanic bulla, which are less. By actual weight the skull of the male is 25 per cent lighter and the skull of the female 24 per cent heavier than in richardsonii. From fallenda and anguinae, haidarum differs in that measurements of the skulls of both sexes either average more, or are uniformly more, with two exceptions. These are the lesser length and breadth of the tympanic bulla, in comparison with males of fallenda, and the dimensions of Ml which are about the same in all three races concerned. The pre- and inter-orbital parts are larger in relation to the remainder of the skull. The postorbital breadth is actually a third more than in fallenda. In relation to the basilar length, the tympanic bulla is shorter and the braincase deeper than in males of anguinae. The skull of the male is 27 per cent heavier than that of fallenda and 58 per cent heavier than that of anguinae. The skull of the female is 59 and 50 per cent heavier than those of fallenda and anguinae, respectively. Comparison of the skull with those of alascensis, celenda and seclusa has been made in the accounts of those subspecies. Remarks. — The available specimens of this ermine were obtained by J. H. Keen in 1898, Wilfred H. Osgood and E. A. Lewis in 1900, W. W. Brown in 1914, J. A. Munro in 1917 and 1918, and Allan Brooks in 1920. M. e. haidarum has more claim to full specific status than any other race of ermine because the diagnostic structural features are numerous and individually of relatively great degree. Indeed, individual variation appears not to bridge the gap between any population of haidarum and other subspecies and strong reasons could be advanced for according haidarum the status of a full species. It differs from the subspecies of erminea on the adjoining mainland and adjoining islands to the north and south and agrees with the Arctic races (arctica, polaris, semplei and kadiacensis) in great extent of the color of the underparts, extension of this color onto the underneath side of the tail, long black tip of the tail and general form of the skull including the relatively heavy preorbital region. The color although darker than in the Arctic subspecies, is lighter than in the insular races ifnmediately to the north and south. In combination, the features MUSTELA ERMINEA 145 mentioned could be taken as indication that haidarum is a relict population from a former glacial period. Assuming that it is a relict population, the color may have become slightly darker since that period but the main response appears to have been a decrease in size for this is a much smaller animal than the Arctic ermines. The size is about what would be expected if one were to judge by the slightly larger ermines on the islands of southeastern Alaska to the north and the smaller ermine on Vancouver Island to the south. The ermines of the islands of southeastern Alaska, excepting possibly the incompletely known seclusa, have fewer characters of the Arctic races and more characters of the races of the adjoining mainland. Therefore, a possible inference is that the distinctive characters of ermines of the Alaskan islands developed with the aid of isolation from stocks which reached the islands after the glacial period. M. e. haidarum may have found its way to the Queen Charlotte Islands in the glacial period. Specimens examined. — Total number, 17, as follows. Arranged by locality from north to south. Unless otlierwise indicated, specimens are in the U. S. National Museum. British Columbia. Queen Charlotte Islands. Masset, 7 (4', 1', 1'); Skid- gate, 1; Graham Island, 5 (2', 1°, 1'); Cumsheva Inlet, 3; no locahty more definite than Queen Charlotte Islands, V. Mustela erminea anguinae Hall Ermine Plates 5, 6, 7, 11, 12 and 13 Mustela cicognanii anguinae Hall, Univ. CaUfomia Publ. Zool., 38:417, November 8, 1932. Puforius cicognanii, Baird, Mamm. N. Amer., p. 161, 1858 (part). Pulorius streatori, Swarth, Univ. California Publ. Zool., 10:102, February 13, 1912. Mustela erminea anguinae Hall, Journ. Mamm., 26:79, February 27, 1945; Hall, Journ. Jvlamm., 26:181, July 19, 1945. Type. — Male, adult, complete skeleton (no skin); no. 12482, Mus. Vert. Zool., French Creek, Vancouver Island, British Columbia; found as a des- iccated carcass on May 1, 1910; obtained by Harry S. Swarth. Range. — Vancouver Island, British Columbia. See figures 25, 27 pages 95, 149. Characters for ready recognition. — DifiFers from M. e. haidarum, in both sexes, in Hght-colored underparts less than half the wddth of dark-colored upper parts, proximal two-thirds of under surface of tail colored like upper 1. Mus. Vert. Zool., Univ. California 4. Univ. Washington Museum of Zoology 2. American Mus. Nat. History 5. Canadian National Museuem 3. Donald R. Dickey Collection 146 University of Kansas Publs., Mus. Nat. Hist. parts instead of underparts, interorbital breadth less than distance between glenoid fossa and posterior margin of external auditory meatus; from M. e. fallenda, in both sexes, anterior margin of tympanic bullae flush with squa- mosal rather than projecting from floor of braincase, in males by sagittal crest absent, in females by total length more than 238 and tooth-rows about same length as, instead of longer than, tympanic bulla; from M. e. streatori, in male, by sagittal crest absent and hind foot ordinarily more than 33.5, in female by hind foot more than 27.5, basilar length more than 30.2; from M. e. olympica, in males, by greater average size, hind foot ordinarily more than 33.4 and interorbital breadth ordinarily more than 8.5, in females by larger size, total length more than 235, tail more than 65, hind foot more than 27.5, basilar length more than 30.2. Description. — Size. — Male: Sixteen adults and subadults yield average and extreme measurements as follows: Total length, 272 (261-284) mm.; length of tail, 81 (74-86); length of hind foot, 35.0 (33.5-36). Female: Five adults and subadults have corresponding measurements as follows: 247 (241-257); 69 (66-73); 30.0 (28.0-32.0). Color. — As described in Mustela erminea streatori except that: occasionally white in winter; upper parts about tone 2 of Dark Chocolate of Oberthiir and Dauthenay; least width of color of underparts averaging, in 7 adult males, 6 (0-15) per cent of greatest width of color of upper parts. Black tip of tail in same series averaging 37 (26-46) mm. which is 46 (32-54) per cent of length of tail-vertebrae. Skull. — Male (based on 13 adults): See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 1.2 (1.0- 1.3) grams; basilar length, 34.0 (32.5-35.6); length of tooth-rows more or less (usually less) than length of tympanic bulla. Female (based on 5 adults): See measurements and plates 11-13. As described in Mustela erminea richardsonii except that: Weight 0.9 (0.77- 1.06) grams; basilar length, 31.5 (30.9-31.8) grams; length of tooth-rows more or less than (approximately same as) length of tympanic bulla; breadth of rostrum more than 30 per cent of basilar length. The sexual dimorphism in the skull is slight, the skull of the male being only a third heavier than that of the female. In fal- lenda of the adjacent mainland to the east the male is three- fourths heavier than the female. In comparison with fallenda, males are smaller, averaging less in every cranial and dental measurement taken and by weight are a fifth lighter; sagittal crest absent rather than present; tympanic bullae flush with squa- mosal rather than projecting below floor of braincase; in relation to basilar length, tympanic bullae smaller, braincase deeper and broader, skull wider interorbitally and across zygomata. Females are larger than in fallenda, and with one exception average larger in every cranial and dental measurement taken, being 6 per cent heavier. The one exception mentioned is the lesser actual length of the tympanic bulla in anguinae, in which the length of the tooth- J MUSTELA ERMINEA 147 rows is about the same as, rather than less than, the length of the tympanic bulla. The postorbital breadth is greater than in fallenda and the anterior edges of the tympanic bullae are flush with the squamosals rather than projecting below the floor of brain- case. In relation to the skull as a whole the preorbital and in- terorbital parts are larger. In comparison with streatori, skulls of males are of about the same size, anguinae being only 9 per cent heavier. The length of the tooth-rows is ordinarily less than, rather than about equal to, the length of the tympanic bulla; sagittal crest wanting rather than present since in anguinae the temporal muscles meet usually only at the posterior end of the braincase instead of all along the mid- line on its top; tympanic bullae narrower and more nearly flush with squamosal (less protruded from braincase). Relative to the basilar length, the zygomatic breadth is more, the tympanic bullae are narrower, and the braincase is deeper at the anterior end of the basioccipital. The female is 41 per cent heavier than streatori, there being no overlap in most cranial and dental measurements. Ml, however, is approximately the same size in each subspecies. The tooth-rows and tympanic bulla are of almost equal length whereas in streatori the length of the tooth-rows is less than that of the bulla. Difterences from olympica, in males, are: Ml shorter; all other measurements of teeth and parts of skull averaging larger; skull 20 per cent heavier; tooth-rows averaging shorter than tympanic bulla rather than about the same; relative to basilar length, brain- case deeper at anterior end of basioccipital and tooth-rows shorter. The skull of the female is 64 per cent heavier, larger in every measurement taken without overlap; temporal ridges meeting, rather than separated, at lambdoidal crest; length of tooth-rows about equal to, rather than shorter than, tympanic bulla; in re- lation to basilar length, skull deeper, orbitonasal length more, mastoid and zygomatic breadths more, and tympanic bullae shorter. Remarks. — References in the literature to this insular race mostly were under the name streatori until 1932 when in the course of the present study the name anguinae was proposed. A few specimens have been taken by nearly every student of small mammals who has collected on Vancouver Island. Arthur Peake and Herbert Laing have probably collected more specimens than any other two zoologists. M. c. anguinae is noteworthy for the slight secondary sexual 148 University of Kansas Publs., Mus. Nat, Hist. variation in size; the disparity between the two sexes is less than in any other American subspecies of erminea. By hnear measure- ment the body of the female is only 7 per cent shorter than in the male (178 versus 191 mm.). Linear measurements and weights of the skulls of the two sexes are further indicative of this approxima- tion in size. By weight the skull of the female is only a fourth lighter than that of the male, or, stated in another way, the male's skuU is only a third heavier (1.2 versus 0.9 grams). No geographic variation has been detected between lots of specimens from different parts of Vancouver Island. The one specimen available from Salt Spring Island presents no obvious differences from selected individuals from Vancouver Island. The winter pelage is more often brown than white. Of 17 specimens seen in winter pelage or in transition pelage, only 6 are white. These 6 are from Comox, Stamp River, Hilliers, Jeune Landing and Port Alice. Of the 34 specimens in brown pelage, 7 have the dark color of the upper parts meeting on the abdomen. Six of the 34 have brown color on the pectoral region. In two, this is a separate patch but in the other four the dark color is a continuation of the upper parts and extends in front of each foreleg over part of the pectoral region, but the two extensions, one from either side, do not meet on the underparts. The color of the lips was recorded in 22 individuals: one had both the upper- and lower-lips white; 7 had the upper lips brown and the lower lips white; in 14 both the upper- and lower-Hps were brown. Specimens examined. — Total number, 40, listed by localities from north to south as follows. Unless otherwise indicated, specimens are in the National Musemn of Canada. British Columbia. Vancouver Island: Cape Scott, 4; Shushartie, 1; Quatsino, 1'; Jeune Landing, 1'; Port Alice, 5'; Marble Creek, Quatsino Sound, 1'; Port Hardy, 5; Sayward, 2; Bear Lake, 4; Bear River, 1; Comox, 4 (3*); Stamp River, Albemi, 1°; Errington, 1'; French Creek, 1'; HilUers, 1'; Craigs Crossing, 1'; Nanaimo, 2'; Cowichan Lake, 1'; Duncan, 2\ Salt Spring Island, 1*. Mustela erminea fallenda Hall Ermine Plates 5, 6, 7, 11, 12 and 13 Mustela erminea fallenda Hall, Joum. Mamm., 26:79, February 27, 1945; Hall, Joum. Mamm., 26:181, July 19, 1945. Putorius streatori Merriam, N. Amer. Fauna, 11:13, June 30, 1896 (part- Sumas). 1. Mus. Vert. Zool., Univ. California 4. Provincial Museum of British Columbia 2. Collection of Arthur Peake 5. Collection of Kenneth Racey 3. Collection of J. A. Munro MUSTELA ERMINEA 149 123 O type locality M. e, r ic hard son ii specimens examined 2 tifiiSjiiiild M.e.invicfa M.e.anquinae M.e . fallendo lim^giM qM M.e.oly mpi c a M.e.streaton M.e .quioso M.e.mu r I c us Fig. 27. Map showing known occurrences and probable geographic ranges of the subspecies of Mustela erminea in Washington and parts of British Colum- bia and Oregon. 150 University of Kansas Publs., Mus. Nat. Hist. Type. — Male, adult, skull and skin; no. 7096, Nat. Mus. Canada; Hunting- don, British Columbia; May 21, 1927; obtained by C. H. Young, original no. 317. The brown summer sldn is well made. The skull (plates 5-7) is complete. Right p2 has the crown broken away; otherwise the teeth all are present and entire. Range. — On mainland in immediate vicinity of coast from probably op- posite Texada Island, British Columbia, south to Lake Whatcom, Washington, and east to Mount Baker Range on International boundary. See figures 25, 27 on pages 95, 149. Characters for ready recognition. — DiflFers from M. e. haidarum, in both sexes, in hght-colored underparts less than half the width of dark-coloroed upper parts, proximal two-thirds of under surface of tail colored like upper parts instead of underparts, interorbital breadth less than distance between glenoid fossa and posterior margin of external auditory meatus; from M. e. richardsonii in both sexes, by upper hps brown rather than white, in males hind foot less than 41 and basilar length less than 38.3, in females hind foot less than 29, basilar length less than 31.4 and breadth of rostrum more, instead of less, than 30 per cent of basilar length; from M. e. invicta, in both sexes, by upper lips brown (not white); in males by skull averaging shorter (basilar length 35.7 versus 37.0); in females by breadth of rostrum more, instead of less, than 30 per cent of basilar length; from M. e. anguinae, in both sexes, by anterior margin of tympanic bulla projecting from floor of braincase rather than flush with squamosal ( the difi^erence is slight in females ) , in males by sagittal crest present, in females by total length less than 238 and tooth -rows longer than, instead of about same length as, tympanic bulla; from M. e. streatori, in both sexes, by black tip of tail more than half of length of tail-vertebrae, in males hind foot more than 33.7, tympanic bulla longer than, instead of about same length as, upper tooth-rows; weight of skull more than 1/4 grams, in females weight of skull more than 0.7 grams, length of lateral side of P4, 4 mm. or more; from M . e. olympica, in males, length of hind foot more than 33, black tip of tail more than 36.5 mm., weight of skull more than 1.2 grams, basilar length more than 33.5, in females length of hind foot more than 25.5, weight of skull more than 0.66 grams, basilar length more than 28.4; from M. e. gulosa, in both sexes, by anterior margin of tympanic bulla projecting below floor of braincase rather than flush with squamosal (the difference is slight in females), in males hind foot more than 33.5, weight of skull more than IM grams, basilar length more than 33.9, in females by total length more than 222, hind foot longer than 26, weight of skull more than 0.7 grams, basilar length more than 29. Description. — Size. — Male: Seven adult topotypes yield average and ex- treme measurements as follows: Total length, 278 (249-305); length of tail, 77 (69-81); length of hind foot, 36.5 (34-40). A male topotype of un- known age weighed 113 grams. Female: Two adult topotypes, with actual measurements in parentheses, average as follows: Total length, 232 (228-236); length of tail, 60 (57- 62); length of hind foot, 27 (27-27). An adult from Morovitz Guard Station, Wash., weighed 54 grams. Color. — Winter pelage rarely white, brown pelage indistinguishable from MUSTELA ERMINEA 151 summer pelage except for slightly more smoky tinge in winter in specimens from some localities; otherwise as described in Mustela erminea streatori ex- cept that least width of color of underparts averaging, in seven adult topo- types, 18 (0-37) per cent of greatest width of color of upper parts. Black tip of tail averaging, in same series, 45 (38-52) mm. which is 58 (53-65) per cent of length of tail-vertebrae. In comparison with richardsonii and invicta, fallenda diflFers in darker color of upper parts and their extension at the expense of the hght-colored vmderparts which are narrower by a half. In correlation with this restriction in area of the light-colored underparts, the upper lips are brown instead of white. In comparison with anguinae, olympica and streatori, the longer black tip on the tail is the principal difference in color. From gulosa, fallenda differs in slightly darker color of upper parts and in narrow underparts, the width of the same being only about a fifth instead of a third of the width of the dark-colored upper parts. Skxdl. — Male ( based on 7 adults ) : See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 1.5 (1.3- 1.7) grams; basilar length, 35.7 (34.3-38.2). Female (based on 6 ads.): See measurements and plates 11-13. As de- scribed in Mustela erminea richardsonii except that: Weight, 0.85 (0.73-1.0) grams; basilar length, 30.6 (29.4-31.7); breadth of rostrum more than 30 per cent of basilar length. In comparison with richardsonii, skulls of males differ as follows: averaging smaller in every measurement taken with no overlap in several dimensions; 40 per cent lighter; in relation to basilar length, rostrum (orbitonasal length) longer and skull slightly broader in- terorbitally. Females average smaller in every cranial and dental measurement taken with no overlap in basilar length, length of tooth-rows and length of tympanic bulla; 22 per cent lighter; breadth of rostrum more, rather than less, than 30 per cent of basilar length; in relation to basilar length, pre- and inter-orbital parts of skull larger, and mastoid breadth more. Differences from males of olympica are: size larger with no over- lap in most measurements; 50 per cent heavier; tympanic bullae longer than upper tooth-rows rather than of about equal length; in relation to basilar length, rostrum shorter, braincase wider and deeper, zygomata more expanded. Females are larger with no overlap in most measurements; 35 per cent heavier; in relation to basilar length, pre- and inter-orbital regions narrower, braincase deeper and wider across mastoids. Differences from streatori, in males, are: skull averaging larger in every cranial and dental measurement taken; 36 per cent heavier; tympanic bulla longer than, instead of about same length as, upper tooth-rows. In females the inner lobe of Ml is shorter antero- posteriorly; otherwise all measurements of fallenda average larger 152 University of Kansas Publs., Mus. Nat. Hist. and it is 33 per cent heavier; rostrum and interorbital region broader in relation to remainder of skull. In comparison with gulosa, skulls of males differ as follows: averaging larger in every measurement taken with no overlap in several dimensions; 50 per cent heavier; tympanic bullae wdth anterior margins projecting slightly below squamosals rather than flush with same; length of bulla more than, rather than about same as, that of upper tooth-rows. Considering the great difference in size, the relative proportions are remarkably alike. In females, length of inner lobe of Ml about the same; otherwise averaging larger in every measurement taken; 44 per cent lighter; relative to basilar length, tooth-rows longer, skull wider across zygomata and mastoids, rostrum and interorbital regions slightly narrower, skull shallower in plane of last upper molars. Comparisons with haidariim, invicta and ongui7iae are made in accounts of those subspecies. Remarks. — Until the name fallenda was proposed in the course of the present study, most of the specimens of this race were assigned to streatori. Intergradation v/ith streatori is complete as it is also with invicta and richardsonii, in other words with each of the subspecies whose ranges meet that of fallenda. In color and in size the difference is least between streatori and fallenda. As between fallenda and invicta the size is not greatly different and the intergradation in color is gradual. Between fallenda and richardsonii intergradation is somewhat different and to fully appreciate its nature we should remember that the color of fallenda resembles that of the saturate coastal races, streatori, anguinae and olympica although the black tip of the tail is longer. In this latter feature and in several cranial details, as well as in greater degree of secondary sexual variation in size, fallenda resembles richardsonii. Because the two differ more than do most subspecies of ermine whose ranges meet, some of the intergrades at first inspection appear to be widely different from either parent stock. For example, specimens from Alta Lake, British Columbia, may give this impression because the combina- tion of large size and dark color suggests a kind of ermine different from either fallenda or richardsonii. In no instance, however, has there been found in these intergrades any character other than those occurring in one or the other of the two parent races. Along the coast in the north part of the geographic range assigned to fallenda, some specimens nearly typical of richardsonii have been taken so near to the place where fairly typical fallenda was MUSTELA ERMINEA 153 obtained that I have doubted whether there is intergradation in the usual fashion in this area; more specimens will have to be obtained from this coastal area to resolve the doubt one way or the other. The winter pelage is brown in all specimens at most localities. The only white pelage seen was in each of tliree specimens from Glacier, Whatcom County, Washington. A fourth specimen from there is in brown winter pelage. At any one locality there is much variation in the degree to which the dark color of the upper parts encroaches on die area that in most other races is light-colored. An extreme degree of encroachment is shown by a specimen taken on December 1, 1935, by R. A. Cummings, at Vancouver, British Columbia, in which the light color occurs only in three restricted areas, the chin, the throat and the lower breast; otherwise the coat is brown. There are other specimens, for instance from the type locality, which differ mainly in having an additional white spot in the inguinal region. The opposite extreme, in a specimen also from the tyjDe locality, is where the least width of the light-colored underparts on the abdominal region is a third of the circumference of the body. The two extremes are connected by a dozen inter- mediate stages. Of 64 specimens in which the color of the lips was carefully examined, one, from Vancouver, has both the upper- and lower-lips brown; 9 have both the upper- and lower-lips white; and 54 have the upper lips brown and the lower lips white. Specimens examined. — Total number, 72, arranged by localities from north to south. Unless otherwise indicated, specimens are in the National Museum of Canada. British Columbia. Horseshoe Lake, Stillwater, 2; Vancouver, 1'; Point Grey, V; Port Moody, .5^'; ChilUwack, 8 (2', 4% P); Sumas, 19 ( 18\ 1*); Thurstons Ranch, 2; Cultus Lake, 2; Mt. Baker Range, 5'; Lihumption Park, 1; Hunting- don, 14; Tami Hy Creek, 1. Washington, Whatcom Co.: Semiahmoo, 1"; New Whatcom, 1°; Lake Whatcom, 2'; 5 mi. W Glacier, 1^ Glacier (3 at 900 ft.), 4-'; E Side Easton Glacier, Mt. Baker, V; Morovitz Guard Station, 831 ft., 1". Mustela erminea olympica Hall Ermine Plates 5, 6, 7, 12, 13 and 14 Mustela erminea olympica Hall, Joum. Mamm., 26:81, February 27, 1945; Hall, Journ. Mamm., 26:181, July 19, 1945. Mustela rixosa, Svihla and Svihla, Mmrelet, 13:24, January, 1932. Mustela rixosa rixosa, Svihla and Svihla, Murrelet, 14:39, May, 1933. 1. Mus. Vert. Zool., Univ. California 5. Lcland Stanford Junior University 2. United States National Museum 6. Collection of William T. Shaw 3. Museum of Comparative Zoology 7. Collection of Kenneth Racey 4. Field Museum of Natural History 8. Collection of Walter W. Dalquest 154 University of Kansas Publs., Mus. Nat. Hist. Type.— Male, adult, skull and skin; no. 90738, U. S. Nat. Mus., Biol. Surv. Coll.; near head of Soleduck River, 4500 ft., Olympic Mountains, Clallam County, Washington; April 28, 1897; obtained by Vernon Bailey, original no. 6213. The skin is well prepared and in good condition. The skull (plates 5-7) is unbroken and the teeth all are present and entire. Range. — Olympic Peninsula, Wasliington, south to Olympia. See figures 25, 27 on pages 95, 149. Characters for ready recognition. — DiflFers from M. e. anguinae, in males, by lesser average size, hind foot ordinarily less than 33.4, and interorbital breadth ordinarily less than 8.5, in females by smaller size, total length less than 235, tail less than 65, hind foot less than 27.5, basilar length less than 30.2; from M. e. fallenda, in males, by length of hind foot less than 33, black tip of tail less than 36.5, weight of skull less than 1.2 giams, basilar length less than 33.5, in females length of hind foot less than 25.5, weight of skull less than 0.6 grams, basilar length less than 28.4; from M. e. streatori by smaller size, in males hind foot less than 33.0, basilar length ordinarily less than 32.5, in females by hind foot ordinarily not longer than 24, by breadth of rostrum less than 8.6, depth of braincase at posterior border of upper molars less than 7.6. Description. — Size. — Male: Twelve individuals of adult proportions yield average and extreme measurements as follows: Total length, 243 (205-269); length of tail, 65 (60-74); length of hind foot, 31 (29-32). Female: Corresponding measurements of six females are: 196 (188-208), 52 (45-60?), 23.4 (22.7-24.0). An adult weighs 30 grams. Color. — As described in Mustela erminea streatori except that least width of color of underparts averaging, in 12 males of adult proportions, 5 (0-11) per cent of greatest width of color of upper parts. Black tip of tail averaging, in same series, 26 (20-35) mm., which is 40 (31-58) per cent (average the same as in streatori ) of length of tail-vertebrae. Skull. — Male (based on 5 adults): See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 1.0 (0.9-1.1) grams; basilar length, 31.8 (30.6-32.5); length of tooth-rows more or less than ( about equal to ) length of tympanic bulla. Female (illustrated by 3 adults): See measurements and plates 12-14. As described in Mustela erminea richardsonii except that: Weight, 0.55 (0.52- 0.58) grams; basilar length, 27.1 (26.7-27.5); breadth of rostrum more than 30 per cent of basilar length. In comparison with streatori, skulls of corresponding sex average smaller in every measurement taken with no overlap in most of those of females. Exception is to be made for the inner lobe of Ml in males where the size is the same. By weight males are smaller by 10 per cent and females by 14 per cent. In relation to other parts of the skull the tympanic bullae are narrower and in fe- males they are shorter as well. Comparison with anguinae and fal- lenda has been made in the accounts of those subspecies. MUSTELA ERMINEA 155 Remarks. — The smaller size, especially of females, is the principal feature distinguishing this race from streatori. On the basis of available data the female of olympica is smaller than that of any other race and hence is the smallest adult weasel of the species erminea, in either the Old World or in America. Intergradation with streatori is indicated by specimens from the southern end of Puget Sound. These specimens are intermediate in size between typical examples of the two races concerned. The color of the upper parts is uniform and the color pattern varies less than in geographically adjoining races. The white color of the underparts is restricted to a thin line on the abdominal re- gion, but widens out posteriorly in the inguinal region and an- teriorly over the pectoral region, throat, chin and lower lips. The upper Hps are brown. The brown of the upper parts extends around in front of each foreleg, the two brown areas not quite meet- ing on the lower throat. The above description applies to each of the 19 specimens examined with regard to these details. Every specimen seen in the winter coat was brown, not white. Specimens examined. — Total number, 20, arranged by counties from north to south. Unless otherwise indicated, specimens are in the U. S. National Museum, Washington. Clallam Co.: Clallam Bay, 2 ( 1\ 1'); Elwah, 2"; Johnsons Ranch, 1*; Happy Lake, 1*; Boulder Lake, 2*; near head of Soleduc River, 4500 ft., 1; 12 mi. S Port Angeles, V. Jefferson Co.: Hayes Cr., 2000 ft., Elwah River, 2; head N Fork Quinault River, 4000 ft., 1; Duckabush, 3; N Fork Skokomish River, 1. Mason Co.: Lake Cushman, 2°; 4 mi. S Olympia, 1. Mustela erminea streatori (Merriam) Ermine Plates 5, 6, 7, 12, 13 and 14 Putorius streatori Merriam, N. Amer. Fauna, 11:13, pi. 2, figs. 5, 5a, 6, 6a, June 30, 1896. Putorius cicognanii, Baird, Mamm. N. Amer., p. 161, 1858 (part imless no. 2395 was a female of M. frenata). Putorius pusillus, Baird, Mamm. N. Amer., p. 159, 1858 (part). Putorius (Gale) vulgaris, Coues, Fur-bearing animals, p. 102, 1877 (part). Mustela streatori streatori. Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912; Grinnell, Univ. CaUfomia Publ. Zool., 40:101, September 26, 1933. Mustela cicognanii streatori. Hall, Murrelet, 12:22, January, 1931; Hall, Univ. California Publ. Zool., 38:417, November 8, 1932. Mustela erminea streatori. Hall, Joum. Mamm., 26:77, February 27, 1945; Hall, Joum. Mamm., 26:181, July 19, 1945. Mustela rixosa. Beer, Joum. Mamm., 29:296, August 31, 1948. 1. Mus. Vert. Zool., Univ. California 4. Field Museum of Natiu-al History 2. Univ. Washington Museum of Zoology 5. Museum of Zoology, Univ. Michigan 3. Charles R. Conner Museum 156 University of Kansas Publs., Mus. Nat. Hist. Type. — Male, adult, skull and sldn; no. 76646, U. S. Nat. Mus., Biol. Surv. Coll.; Mount Vernon, Skagit Valley, Skagit County, Washington; Feb- ruary 29, 1896; obtained by D. R. Luckey, original no. 3. The skull is unbroken and the teeth all are present and entire. The skin, in brown winter pelage, is stuffed and in good condition. Range. — Western Washington along eastern side of Puget Sound, western Oregon from the Cascades to the coast, and northwestern California south in the humid coastal district nearly to the Golden Gate. See figures 25, 27 on pages 95, 149. Characters for ready recognition. — Differs from M. e. anguinae, in male, by sagittal crest present and hind foot ordinarily less than 33.5, in female by hind foot less than 27.5, basilar length less than 30.2; from M. e. fallenda, in both sexes, by black tip of tail less than half of length of tail-vertebrae, in males hind foot less than 33.7, tympanic bulla about same length as, in- stead of longer than, upper tooth-rows; weight of skull less than 1/4 grams, in female weight of skull less than 0.7 grams, length of lateral side of P4 less than 4 mm.; from M. e. ohjmpica, by larger size, in males hind foot more than 33.0, basilar length ordinarily more than 32.5, in females by hind foot ordinarily longer than 24, by breadth of rostrum more than 8.6, depth of braincase at posterior border of upper molars more than 7.6; from M. e. giilosa and muricus, in both sexes, by upper Ups brown (not white), light color of underparts ex-tending down hind leg no farther than knee, depth of skull at posterior border of upper molars more than 7.7 in females and ordinarily more than 9.6 in males, further from muricus by tail more than 62 in males and more than 49 in females; from M. e. invicta by upper Ups wliite (not brown), in males hind foot more than 36 and basilar length more than 35, in females hind foot more than 29.5 and basilar length more than 30.5. Description. — Size. — Male: Twelve adults from Blaine and Tillamook, Oregon, yield average and exlieme measurements as follows: Total length, 255 (245-275); length of tail, 72 (64-80); length of hind foot, 31.5 (30.0- 33.5). Female: Seven adults from Blaine and Tillamook, Oregon, yield average and extreme measurements as follows: Total length, 214 (193-230); length of tail, 55 (50-63); length of hind foot, 25 (24-27). Color. — Winter and summer pelages indistinguishable; upper parts uni- form and ranging from Raw Umber to slightly darker (16n), and about tones 1 to 3 of Dark Chocolate of Oberthiir and Dauthenay, pi. 342; underparts white, in summer rarely with a faint buffy suffusion in pectoral region; color of underparts extends from chin, and often lower Hps, posteriorly to inguinal region, distally on posterior sides of forelegs onto antipalmar faces of toes (sometimes interrupted at and above wrist) and on medial sides of hind legs hardly to knee. Least width of color of underparts averaging, in twelve adults from Blaine and Tillamook, 10 (0-47) per cent of greatest width of color of upper parts. Black tip of tail, in same series, averaging 28 (24-33) mm. which is 40 (34-47) per cent of length of tail-vertebrae. Skull. — Male (based on 12 adults): See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 1.1 (1.0- I MUSTELA ERMINEA 157 1.2) grams; basilar length, 33.2 (32.5-33.8); length of tooth-rows more or less than (about same as) length of tympanic bulla. Female (based on 7 adults): See measurements and plates 12-14. As described in Mustela erminea richardsonii except that: Weight, 0.64 (0.60- 0.67) grams; basilar length, 28.5 (27.6-29.5); breadth of rostrum more than 30 per cent of basilar length. Comparison with anguinae, fallenda, olympica, gulosa and muricus is made in accounts of those subspecies. Remarks. — This weasel is rare in collections and the best mate- rial of it was obtained by Alex Walker in Tillamook County, Ore- gon, where he resides. The almost ideal series of 30 specimens showed the range of secondary sexual, age, and individual varia- tion expectable in the small ermines of the Pacific Coast of the United States and was the means of allowing satisfactory decision on questions of classification in the related subspecies in which in- dividuals are of comparable size. Intergradation with each of the geographically adjoining sub- species, olympica, fallenda, invicta, gulosa and muricus is shown by specimens examined. With the last mentioned subspecies, in- tergradation is shown by two specimens from as far south as Siski- you County, California, assigned to muricus. The application of the name streatori is difficult because it was based on a specimen from a place where two cHnes cross. The north-south cline is one of size which decreases to the south. The east- west cline is one of intensity of color, the westernmost ( coastal ) population being the most intensely colored. The type locality of streatori is at the place where two lines perpendicular to one an- other, and representing the two clines, cross. This intersection is near the place where the ranges of several subspecies meet. The nomenclatural question is, to which one of 6 subspecies should the name streatori apply. Specimens from barely within the geographic boundaries of four of these subspecies so closely resemble topo- types of streatori that a student with material at his disposal from only the area about Puget Sound naturally would apply the name streatori to all of his specimens, and knowing even of the arrange- ment adopted in the present account the student will have difficulty in identifying his specimens according to it. Not only will the student find the arrangement difficult, but probably unsatisfactory if he thinks of streatori as being the kind of animal represented by topotypes. I conceive of topotypes of streatori as being nontypical of the subspecies; they are intergrades with fallenda. My aim was initially to work out the geographic ranges of subspecies and only 158 University of Kansas Publs., Mus. Nat. Hist. subsequently to apply names, according to which type localities fell within the previously determined geographic ranges. By this procedure no greater weight was given to a holotype and to topo- types than to specimens from any other locahty. Of the 40 specimens seen in winter pelage, only one is white. It is from Darrington in the Cascade Mountains of Washington. The 39 others are brown and I doubt that the white pelage ever occurs in the low coastal territory included within the geographic range of streatori. This subspecies resembles anguinae and olympica in the great extension of area of the dark-colored upper parts at the expense of the area of the light-colored underparts. The usual ar- rangement is one where the brown of the two sides nearly meets on the midventral line leaving a sizable, inguinal area of light color connected by a thin line to the sizable area of hght color on the pectoral region. The light color of the pectoral area ordinarily is continuous with the light-colored area of the throat and chin but the dark color of the upper parts extends around in front of each foreleg. These extensions of dark color meet on the chest in only 2 of the 56 specimens examined in this regard. Across the abdo- men the dark color is continuous in 4 of the 56 specimens. The lower lips are brown instead of white in only 3 individuals and in 2 of these the lip of one side is brown and its opposite is white. The variation in color-pattern is less than in anguinae or than in fallenda. Specimens examined. — Total number, 63, arranged alphabetically by states, then by counties from north to south in each state. Unless otherwise indicated, specimens are in the U. S. National Museum. California. Humboldt County: 10 mi. NE Carlotta, 1\ Mendocino County: Russian Gulch Slate Park, 1\ Sonoma County: Mouth of Gualala River, V. Oregon. Clatsop Co.: Astoria, 1. Tillamook County: Tillamook, 16 (14^ 1"); Blaine, 12 {1\ 2', 1*, 2"). Washington Co.: Beaverton, 1°; Forest Grove, 1'. Clackamas Co.: Oregon City, 1*. Lincoln Co.: Newport, 1. Linn County: Sico, 1*. Lane Co.: Vida Fish Hatchery, 2"; McKenzie Bridge, 1°; Mercer, 1". Klamath County: Deschutes River, 6 mi. E Crescent Lake, 1*. Douglas Co.: Gardiner, 1'. Curry Co.: Port Orford, 1; Gold Beach, 2'. Washington. Skagit Co.: N end Whidby Island opposite Deception Pass, 1; Hamilton, 4; Mt. Vernon, 3. Snohomish County: Oso, 550 ft., 1; Darrington, 600 ft., 1. Pacijic County: Wallicut River, 2 mi. E llwaco, 1'. Wahkiakum Co.: 4 mi. E. Skamokawa, 3\ Coivlitz County: 4 mi. E mouth Kalama River, 2'; 6 mi. E mouth Kalama River, 1\ Skamania Co.: 15 mi. N Govt. Springs, 1300 ft., 1. 1. Mus. Vert. ZooL, Univ. California 6. Field Museum of Natural History 2. Collection of Alex Walker 7. Collection of O. J. Murie 3. Donald R. Dickey Collection 8. Collection of Stanley G. Jewett 4. Univ. Kansas Mus. Nat. History 9. University of Oregon 5. Museum of Zoology, Univ. Michigan 10. Museum of Comparative Zoology MUSTELA ERMINEA 159 Mustela erminea gulosa Hall Ermine Plates 5, 6, 7, 12, 13 and 14 Mustela erminea gulosa Hall, Joum. Mamm., 26:84, February 27, 1945; Hall, Joum. Mamm., 26:181, July 19, 1945. Putorius streatori Merriam, N. Amer. Fauna, 11:14, June 30, 1896. Type. — Male, subadult, skull and skin; no. 81998, U. S. Nat. Mus., Biol. Surv. Coll.; Trout Lake, Klickitat County, Washington; February 3, 1897; obtained by P. Schmid, original no. 147. The skin is in brown winter pelage, and appears to have been made up from a skin split along the mid-ventral line from the anus to the forelegs. It probably was dried by a trapper, is well made, and lacks a patch of hair on the left flank but otherwise is in good condition. The skull lacks the central part of the left zygomatic arch and the posterior two-tliirds of the right one. The right m2 is represented only by an abortive stump or the broken root, and il and i2 on each side are absent; otherwise, the teeth all are present and entire. Range. — Cascades of Washington from northeastern King County south to Mount Adams. See figures 25, 27 on pages 95, 149. Characters for ready recognition. — Differs from M. e. invicta and fallenda, in both sexes, by anterior margin of tympanic bulla flush with squamosal rather than projecting below floor of braincase (difference shght in females), in males hind foot less than 33.5, weight of skull less than Di grams, basilar length less than 33.9, in females by total length less than 222, hind foot shorter than 26, weight of skull less than 0.7 grams, basilar length less than 29; from M. e. muricus, in both sexes, by upper parts darker, tone 4 of Chocolate or darker (see description of color) least width of hght-colored underparts averaging one-third instead of approximately two-thirds of greatest width of dark-colored upper parts, in males, on the average, tail more than 65, weight of skull more tlian 0.90 grams, basilar length more than 30.8 mm.; from M. e. streatori, in both sexes, by upper hps white (not brown), hght color of underparts extend- ing down hind legs below knee, depth of skull at posterior border of upper molars less than 7.7 in females and ordinarily less than 9.6 in males. Description. — Size. — Male: One adult and four subadults from Mount Rainier yield average and extreme measurements as follows: Total length, 253 (238-266); length of tail, 75 (70-83); length of hind foot, 31.5 (30-33). Corresponding measurements of 9 subadults from Trout Lake are: 257 (233- 282); length of tail, 76 (56-83); length of hind foot, 30.2 (26-33). Female: Of adults, 2 from Mount Rainier and 2 from Trout Lake measure as follows: Total length, 202, 203, 216, 210; length of tail, 54, 52, 57, 51; length of hind foot, 24, 24, 25, 24. The averages for these females are 208, 54, 24.3. Color. — As described in Mustela erminea richardsonii except that color sometimes brown in winter (with more smoky tinge than summer coat); upper parts ranging from tone 2 through tones 3 and 4 of Dark Chocolate (pi. 342) into tone 4 of Chocolate (pi. 343) of Oberthiir and Dauthenay; underparts (always white in winter) in summer Sulphur Yellow or more whitish; least 160 University of Kansas Publs., Mus. Nat. Hist. width of color of underparts averaging, in 5 males from Mount Rainier, 31 (18-45) per cent of greatest width of color of upper parts. Black tip of tail, in same series, averaging 34 (29-40) mm., wliich is 45 (41-50) per cent of length of tail-vertebrae. Skull. — Male (based on 2 ad. and 13 sad.): See measurements and plates 5-7. As described in Mustela erminea richardsonii except that: Weight, 1.0 (0.95-1.16) grams; basilar length, 32.3 (30.9-33.4); length of tooth-rows more or less than (about equal to) length of tympanic bulla. Female (illustrated by 5 adults): See measurements and plates 12-14. As described in Mustela erminea richardsonii except that: Weight, 0.59 (0.53- 0.65) grams; basilar length, 28.1 (27.8-28.4); breadth of rostrinn ordinarily more than 30 per cent of basilar length. In comparison with streatori, skulls of males and females aver- age smaller in every cranial measurement taken. Teeth of about same size and males 9 per cent, and females 8 per cent, hghter. In relation to basilar length, skull of female shallov^er, tympanic bul- lae slightly shorter and, on the average, zygomata less expanded. In comparison with muricus, males average larger in every meas- urement taken; 23 per cent heavier; in relation to other dimensions, braincase shallower at anterior end of basioccipital. Females are of about equal size; in relation to other dimensions, braincase shal- lower and mastoid and zygomatic breadths less. Comparisons with invicta and fallenda have been made in the accounts of those subspecies. Remarks. — This is not a strongly marked race and in most of the characters used for differentiating it from other races it resembles either streatori to the west or muricus to the southeast. Neverthe- less, there is a geographic area, the southern Cascades of Washing- ton, tliroughout which individual characters are combined in es- sentially the same way and there are a few features, for instance, smaller skull of the female, in which gulosa differs from either of its close relatives. In view of these circumstances and because the animals can not well be included in the subspecies streatori or muricus, gulosa is recognized as distinct. The races gulosa and olympica are what might be termed weakly differentiated sub- species in contrast to the stiongly differentiated subspecies streatori and muricus. Of the 21 specimens in winter pelage, 17 are white and four are brown. The brown winter coat is distinctly paler, with more of a smoky tinge, than the brown summer pelage. The light- colored underparts are narrower than in the subspecies immediately to the east but are wider than in the coastal forms to the west. The dark color of the upper parts extends onto the chest in front MUSTELA ERMINEA 161 of the forelegs, as in the coastal forms, in only one of the 13 speci- mens in summer pelage and in it on one side only. The black tip of the tail is short as in the coastal fonns. One specimen is in transitional pelage. It has acquired approximately half of the white winter pelage and was taken on October 12, 1897, at Kee- chelus Lake. Specimens examined. — Total number, 38, arranged by counties from north to south. Unless otherwise indicated, specimens are in the U. S. National Museum. Washington. King County. 2 mi. E Skykomish, 2\ Kittitas Co.: Keechelus Lake, 3 (1'); Martin, 1'; Easton, 3. Fierce Co.: James Lake, 4370 ft., Mt. Rainier, 1; Glacier Basin, 5935 ft., Mt. Rainier, 1; Meslers Ranch, 2000 ft., 1 mi. W Rainier Park, 1. Lewis Co.: Mt. Rainier Nat'l Park, 5 (1 each from: Paradise Park, 5400 ft.; Reflection Lakes, 4900 ft.; Ohanapecosh [Hot] Springs, 2000 ft.; Tahoma Creek, '; Bear Prairie); also in Mt. Rainier Nat'l Park, Long- mire, 3 (!', 1*). Skamania Co.: Mt. St. Helens, 6000 ft., 1. Klickitat Co.: Trout Lake, 18. Mustela enninea muricus (Bangs) Ermine Plates 7, 8, 12, 13, 14 and 41 Putorius (Arctogale) muricus Bangs, Proc. New England Zool. Club, 1:71, July 31, 1899. Putorius streatori leptus Merriam, Proc. Biol. Soc. Washington, 16:76, May 29, 1903. Type from Silverton, San Juan County, Colorado. Putorius muricus, Stephens, Cahfomia Mammals, p. 248, 1906. Putorius cicognani, Taylor, Univ. CaUfomia Publ. Zool., 7:298, June 24, 1911. Mustela streatori leptus, Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912; Bailey, N. Amer. Fauna, 35:48, September 5, 1913; Dixon, Joum. Mamm., 12:72, February 12, 1931; Whitlow and Hall, Univ. California Publ. Zool., 40:246, September 30, 1933. Mustela muricus. Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912; Kellogg, Univ. California Publ. Zool., 12:358, January 27, 1916. Mustela cicognanii lepta. Dice, Joum. Mamm., 1:12, November 28, 1919; Hall, Mamm. Nevada, p. 184, July 1, 1946. Mustela rixosa, Seton, Joum. Mamm., 14:70, February 14, 1933. Mustela cicognanii leptus, Miller, Joum. Mamm., 14:368, November 13, 1933; Bailey, N. Amer. Fauna, 55:293, August 29, 1936. Mustela erminea murica. Hall, Joum. Mamm., 26:84, February 27, 1945; Hall, Joum. Mamm., 26:181, July 19, 1945. Type. — Male, young, skull and skin; no. 9146, collection of E. A. and O. Bangs in Mus. Comp. Zool.; Echo, 7500 ft.. El Dorado County, Cahfomia; July 15, 1897; obtained by W. W. Price and E. M. Nutting. The skull has a fracture along the sagittal suture and fractures on the left side of the braincase but these have been glued, and no part of the skull is missing except in the region of the right P4 which part has been shot away. On the left side m2 never developed. Excepting this tooth and the right P4, 1. Collection of Walter W. Dalquest 3. Mt. Rainier Nat'l Park Collection 2. Acad. Nat. Sciences of Philadelphia 4. Univ. Washuigton Museum of Zoology 6—3758 162 University of Kansas Publs., Mus. Nat. Hist. all the teeth are present and entire. The skin is well made but has the soles of the hind feet turned up. Range. — Near 5300 feet (Denver) to 11000 feet (Santa Fe Baldy); typi- cally boreal but taken in Upper Sonoran Life-zone in winter at Denver; from central and southwestern Montana, southern Idaho, and Blue Mountains of southeastern Washington southward east of the Cascade Divide through the Salmon River Mountains and Sierra Nevada at least into Fresno Coimty of California, in the Great Basin to central Nevada, in the Rocky Mountains into northern New Mexico; eastward to the Black Hills. See figure 25 on page 95. Characters for ready recognition. — Differs from M. e. invicta by hind foot less than 36 and basilar length less than 35 in males and by hind foot less than 29.5 and basilar length less than 30.5 in females; from M. e. gulosa, in both sexes, by upper parts lighter, tone 2 of Chocolate or lighter (see de- scription of color), least wddth of hght-colored underparts averaging about two-thirds instead of one-third of greatest width of dark-colored upper parts, in males, on the average, tail less than 65, weight of skull less than 0.90 grams, basilar length less than 30.8 grams; from M. e. streatori, in both sexes, by upper lips white (not brown), hght color of underparts extending down hind leg below knee, depth of skull at posterior border of upper molars less than 7.7 in females and ordinarily less than 9.6 in males, tail less than 62 in males and less than 49 in females. Description. — Size. — Male: An adult from Black Butte, Cahfomia, meas- ures: Total length, 227; length of tail, 55; length of hind foot, 27. Cor- responding measurements of another from Wheeler Peak, Nevada, are: 220, 56, 26. Two subadults from Colorado, one from Crested Butte and another from Coventry, measure, respectively, as follows: 238, 227; 66, 60; 30, 30. An adult from Wheeler Peak, Nevada, weighs 57.7 grams and another from 2 mi. W Black Butte, Cahf., 54.5 grams. Female: Two adults from Teton County, Wyoming, measure: Total length, 205, 200; length of tail 52, — ; length of hind foot, 23, 23.7. A subadult from 9/2 mi. E Pocatello, Idaho, measures: 197, 50, 25. An adult from Wheeler Peak, Nevada, has corresponding measurements of 190, 42, 23, and weighs 33.8 grams. Color. — As described in Mustela erminea richardsonii except that upper parts tone 2 or lighter of Chocolate of plate 343 of Oberthiir and Dauthenay; underparts white. Pale Buff or with faint wash of Sulphur Yellow; least width of color of underparts in male from Black Butte and one from Wheeler Peak, amounting to 65 and 59 per cent of greatest width of color of upper parts. Black tip of tail, respectively, 28 and 33 mm., which amounts to 51 and 59 per cent of length of tail-vertebrae. In two adult females, one from Teton County, Wyoming, and one from Wheeler Peak, Nevada, the least width of the underparts amounts to 55 and 60 per cent of the greatest width of color of upper parts. Black tip of tail, respectively, 23 and 19 mm., which amounts to 44 and 45 per cent of length of tail-vertebrae. From the other subspecies of small-sized weasels of more northwestern occurrence, namely anguinae, fallenda, olympica, streatori and gulosa, muricus differs in lighter color of upper parts, wider light-colored underparts and rel- atively longer black tip of tail. MUSTELA ERMINEA 163 Skull. — Male (illustrated by 5 adults in table of measurements, which see): See plate 7. As described in Mustela erminea richardsonii except that: Weight, 0.78 (Wheeler Peak) and 0.85 (Black Butte) grams; basilar length, 30.6 (29.8-31.2); length of tooth-rows more or less than (approximately equal to) length of tympanic bulla. Female ( illustrated by 6 adults in table of measurements, which see ) : See plates 12-14. As described in Mustela erminea richardsonii except that: Weight, 0.60 (0.575-0.645); basilar length, 28.0 (27.3-29.4); breadth of rostrum approximately 30 per cent of basilar length. In comparison with streatori, males average smaller in every meas- urement taken with no overlap in most dimensions; 25 per cent lighter; anterior margin of tympanic bulla more nearly flush with squamosal, that is to say less protruded from braincase; in re- lation to other dimensions of skull, braincase shallower anteriorly (at plane of last molars) and deeper posteriorly (at anterior end of basioccipital ) . Females average smaller in every measurement taken except mastoid and zygomatic breadths which are actually more; 6 per cent lighter; in relation to other parts of skull, pre- orbital and interorbital parts slightly smaller; in relation to length of skull, braincase shallower. Comparison with invicta and gulosa is made in the accounts of those subspecies. Remarks. — The smallest males of the entire species are of this subspecies and the females of it are barely larger than those of olympica and gulosa and hence are among the three smallest. The material now available consists only of one or a few specimens from each of several widely separated localities. If as many speci- mens per unit area were available as there are of the species M. erminea from southern British Columbia, geographic variation warranting the division of muricus into more than one subspecies might be revealed. Evidence pointing in this direction is com- prised in the pale color and small size of the pair of adults from Wheeler Peak on the eastern border of Nevada; the suggestion is that there is a distinct pale race of small individuals in the isolated spots of boreal life-zone in the mountains of the desert. The color and size of the specimens from the Toyabe Mountains, and that from the Pine Forest Mountains, both places also in Nevada, nevertheless, lend no support to this suggestion. Comparison of specimens from the Rocky Mountains of Colorado with those from the Sierra Nevada of California gives no basis for recognizing more than one subspecies. Therefore, Putorius streatori leptus Merriam with type locality at Silverton, San Juan County, Colorado, falls as a synonym of the earlier named Tutorius ( Arctogale ) muricus Bangs with type locality at Echo, El Dorado County, California. Further- 164 University of Kansas Publs., Mus. Nat. Hist. more, specimens from northern New Mexico, the southernmost known area of occurrence for the subspecies (and for the species), are as large as specimens from far north in the range of the sub- species, say, in northwestern Wyoming; there is therefore no evi- dence of progressive decrease in size to the southward as in advance of study I supposed existed in muricus. This erroneous supposi- tion was held because I knew that there was a decrease in size to the southward in the species as a whole and also in each of the subspecies richardsonii and invicta directly to the north of muricus. Intergradation with invicta is shown by specimens from south- western Montana. Where the margins of the geographic ranges of invicta and muricus approach one another elsewhere, low-lying territory, zonally unsuited to the existence of the species, occurs along the Snake and Columbia rivers, and precludes any chance of intergradation except around the head of the Snake River Plains. Two specimens, here referred to muricus, from Siskiyou County, California, in both color and cranial characters, are intergrades with streatori and might be referred with almost equal propriety to streatori. Specimens examined. — Total number, 52, arranged alphabetically by states, then by counties from north to south within each state. Unless otherwise in- dicated, specimens are in the Museum of Vertebrate Zoology, University of California at Berkeley. California. Siskiijou County: head of Rush Creek, 6400 ft., 1; Castle Lake, 5434 ft., 1. Teha7na County: 2 mi. W Black Butte, 6800 ft., 1. Placer County: ridge W of Tahoe Pines, Lake Tahoe, 1; Blackwood Creek, 6250 ft., near Tahoe Pines, 1. El Dorado Co.: Fallen Leaf Lake, 6500 ft, 1'; Echo, F. Tuolumne Co.: Ten Lakes, 9200 ft., Yosemite Park, 1. Mariposa Co.: Vogelsang Lake, 10350 ft., Yosemite Park, 1. Mono County: Mammoth, 1'. Colorado. Rio Blanco Co.: Marvine, L Boulder Co.: Camp Albion, 10600 ft., 1'; Boulder, V. Denver Co.: Denver, 1\ Park County: Jefferson, 1'. Gunnison Co.: near Placita in Gunnison County, 1'; Crested Butte, 9000 ft., 3 ( r, 2'). El Paso County: Turkey Creek, SW Colorado Springs, 6000 ft., 1'. Chaffee Co.: Arp[b]ourville, 1"; Plancock, 1. Montrose County: Coventry, 6800 ft., 1". San Juan Co.: Silverton, 1°; in San Juan County above timber- hne, r. Idaho. Bannock County: West Fork of Rapid Creek, 9)2 mi. E Pocatello, 1. Montana. Meagher Co.: Camas Creek, Big Belt Mts., 4 mi. S Ft. Logan, 1'. Beaverhead Co.: Donovan, 1°. County in question: Yellowstone Park, l^ Nevada. Humboldt Co.: Alder Creek, 6000 ft., Pine Forest Mts., 1. Ormsby County: 'A mi S Marietta Lake, 8150 ft., 1. Nye Co.: South Twin River, Toy- abe Mts., l^ White Pine County: Baker Creek (8500 ft., 8675 ft., 11100 ft.), 3. New Mexico. Taos Co.: Twining, 10700 ft., 1°. Sandoval County: 9 mi. E Cuba, 9000 ft., 1. Santa Fe Co.: Saddle S of Santa Fe Baldy, 11000 ft., Santa Fe Range, 1". Oregon. Wasco Co.: Mill Creek, 20 mi. W Warmsprings, T. Klamath Co.: Fort Klamath, 1". MUSTELA ERMINEA 165 South Dakota. Peiminsfon County. 4 mi. SE Hill City, 5300 ft., 2"; Pfan- der's Ranch, 3 mi. SSE Hill City, 5300 ft., 1"; Palmer Gulch, 3 mi. SE Hill City, 5300 ft., 1"; Spring Creek, 2 mi. W Oreville, 5500 ft., 1". Custer County: a mi. E Sylvan Lake, 6250 ft., 1". Washington, Columbia County: Butte Creek, 1; Stayawhile Spring, 5150 ft., 1. Wyoming. Crook County: 5 mi. NW Sundance, 5900 ft., 1". Teton County: Whetstone Creek, 2"; M mi. E Moran, 6700 ft., 1". Sublette County: Ji mi. NE Pinedale, 7500 ft., 1". Albany County: 30 mi. N and 10 mi. E Laramie, 6560 ft., 1"; 26 mi. N and 4Ji mi. E Laramie, 6960 ft., 1". Carbon County: 8 mi. N and 19M mi. E Savery, 8800 ft., 2". Mustela erminea? angustidens (Brown) Plates 7, 12, 13 and 14 Putorius cicognanii angustidens Brown, Mem. Amer. Mus. Nat. Hist., 9 (pt. 4):181, pi. 17, 1908. Mustela cicognanii angustidens. Hay, Iowa Geol. Surv. Bull., 23:32, 1914; Hay, Carnegie Inst. Washington, Pub. no. 322A:252, October 15, 1924; Hay, ibid.. Pub. no. 390 (vol. 2): 528, 1930; Hall, ibid.. Pub. no. 473:111, 112, November 20, 1936. Type. — Female, adult, skull and lower jaws lacking zygomata, right P2 and incisors, no. 12432, Amer. Mus. Nat. Hist.; from Conard Fissure, four miles west of Willcockson, Newton County, Arkansas; obtained sometime in the period 1903 to 1905 inclusive (see plates 8, 14). Range. — Known only from the Pleistocene deposit in Conard Fissure, at the type locality in northern Arkansas. Description. — Skull. — Male (based on nos. 12437, 12441 and 12444): See measurements and plates 7 and 8; weight, unknown; basilar length, 38.1 (36.6-39.2); length of tooth-rows more tlian length of tyinpanlc bulla; breadth of rostrum measured across lacrimal processes less than a third of basilar length; interorbital breadth ordinarily equal to distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth probably averaging approximately the same as distance between last upper molar and jugular foramen. Female (based on nos. 11766 and 12435): See measurements and plates 8, 12-14; weight, unknown; basilar length, 34.0 (32.5-35.1); length of tooth- rows more than length of t>'mpanic bulla; breadth of rostrum about equal to (more or less than) 30 per cent of basilar length; interorbital breadth less than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth probably less than distance between last upper molar and jugular foramen. Comparison of the cranial description given above with those of the Ameri- can races of erminea from the far north vdll show that many characters are held in common — more than vvdth more southern subspecies of erminea. 1. Collection of Lloye H. Miller 7. Collection of Jack C. vonBloeker 2. Museum of Comparative Zoology 8. Collection of Edward R. Warren 3. Donald R. Dickey Collection 9. San Diego Society of Natural History 4. Field Museum of Natural History 10. Acad. Nat. Sciences of Philadelphia 5. United States National Museuin 11. Museum of Zoology, Univ. Michigan 6. Colorado Museum of Natural History 12. Univ. Kansas Mus. Nat. History 166 University of Kansas Publs., Mus. Nat, Hist. Remarks. — The ten specimens studied by the writer fall into two groups of six larger individuals and four smaller. Upon comparing these with each sex of the three species of American Recent weasels, frenata, erminea and rixosa, it is seen that size, and to some degree shape, rule out of consideration both sexes of rixosa and also males of frenata. Thus we are left with females of frenata and males and females of erminea. So far as size is concerned, it can be assumed that the larger specimens are females of frenata and that the smaller are males of erminea. This assumption has in its favor also, the fact that the postglenoidal length of the skull accords with that in Recent specimens. The difference in this regard in Recent animals is that the postglenoidal length of the skull, expressed as a per- centage of the total ( condylobasal ) length of the skull, amounts to: in frenata in erminea (J ordinarily less than 46 $ ordinarily more than 46 5 less than 47 5 more than 48 In the fossils the percentage for the larger skulls is 46; for the smaller skulls it is 48. It may be that the ten fossil skulls are six female frenata and four male erminea but I think not. In the first place a skull of different shape, seemingly of the frenata stock, is known from the deposit and it is almost certain that two subspecies of the same species would not occur at the same place at the same time. It is possible, of course, that parts of the deposits were laid down at times so far apart that a shift in geographic range of two subspecies had oc- curred. This one skull, seemingly of the frenata stock, is the type of Piitoritis gracilis Brown (see p. 404) and was regarded as the only known specimen of gracilis. Regardless of the specific identity of this one specimen named gracilis, the chances of obtaining other- wise from a deposit, like that in Conard Fissure, six females of frenata and four males of erminea without a male frenata or a fe- male of erminea coming to light are so slight as strongly to incline me to the view that the six larger specimens are males of the same species to which the 4 smaller specimens belong. By either this interpretation, or the one initially considered (of female frenata and male erminea), the animals from the fissure are at least sub- specifically distinct from any American Recent weasel. Further- more, by this latter interpretation each sex of this weasel, angiis- tidens, is intermediate between the frenata and erminea stocks in the feature of postglenoidal length which feature, at any place MUSTELA ERMINEA 167 where the two Recent species occur together, serves to distinguish one from the other. In the northernmost subspecies of erminea (arctica for example) the postglenoidal length in some males is no longer than in males of frenata. Considering general size, an- gustidens agrees better with erminea than with frenata and this circumstance has influenced me to place angtistidens as a sub- species of erminea. Today, erminea is not known to occur nearer Conard Fissure than northern Iowa, more than 400 miles to the northward. In comparison with the race there, bangsi, males of angustidens are of approximately the same size but in the shorter distance between the glenoid fossa and anterior margin of the tympanic bulla, and also in the lesser postglenoidal length of the skull, angustidens re- sembles the northernmost American subspecies of erminea. Fe- males of angustidens differ more from any living weasel than the males do. The females are much larger than those of bangsi, and among living American races of erminea most closely resemble intergrades between arctica and richardsonii which intergrades are found approximately 1700 miles to the north of Conard Fissure. In females, the preorbital part of the skull in M. e. arctica is broader and in M. e. richardsonii narrower than in angustidens. If it seems strange that females of angustidens resemble one sub- species whereas males, in size, resemble another subspecies almost a thousand miles distant, it should be remembered that the degree of sexual dimorphism varies much from one subspecies to another in the Recent animals. An example is furnished by Mustela er- minea fallenda and Mustela erminea invicta. The assemblage of mammals from Conard Fissure includes sev- eral species of boreal predilections which, like Mustela erminea, now occur only much farther north than Arkansas. At one time the edge of the sheet of ice was only about 200 miles north of Arkansas. It may be significant that the cranial characters of the female ermine from the Fissure, and qualitative cranial characters of males from there, are most nearly approximated among Recent weasels by those which live along the southern edge of the frozen tundra. In view of what has been said, the possibility should be consid- ered that the distinctive cranial features of angustidens may be the result of evolutionary change in time as well as of geographic varia- tion resulting from horizontal placement. 168 University of Kansas Publs., Mus. Nat, Hist. MUSTELA RIXOSA Bangs Least Weasel (Synonymy under subspecies) Type. — Putorius rixosus Bangs, Proc. Biol. Soc. VVasliington, 10:21, Feb- ruary 25, 1896. Range. — From Norway and Switzerland eastward through Sibera and all the way across North America, but unknown from Iceland, Greenland and the Arctic islands west of Greenland; in North America, from the Arctic Life- zone south to Central British Columbia, Montana and into parts of the Upper Austral Life-zone as in the eastern half of the continent. The southern extension of range in the Appalachians (to North Carolina) is not duplicated in the Rocky Mountains of western North America probably because the region there suitable for rixosa south of Central British Colunlbia and Montana is occupied by the almost equally small Mustela erminea muricus and related subspecies which seem to fill the ecological role that rixosa plays where it occurs. The small size of females of M. erminea cicognanii in New England may similarly account for the absence of rixosa there. Characters for ready recognition. — Differs from both Mustela er- minea and Mustela frenata by tail a fourth or less of length of head and body and without a black tip ( at most a few black hairs at ex- treme tip in rixosa), and from M. frenata and from M. erminea in regions where it and rixosa occur together, by basilar length of skull less than 32.5 in males and less than 31.0 in females. Characters of the species. — Size small: Total length less than 250 in males and 225 in females; tail a fourth or less of length of head and body, and without a black pencil and at most with a few black hairs at extreme tip; caudal vertebrae 11 to 16, normally 15 in M. r. rixosa, and 11 in one M. r. eskimo examined; skull with long braincase and short precranial portion, thus essentially same shape as in M. erminea but the largest males of M. rixosa always with a lesser basilar length that even the smallest females of M. erminea or M. frenata of the same geographic area. In fact no specimens of M. frenata have skulls so small as the largest M. rixosa, and skulls of equal size of M. erminea and M. rixosa, for example, M. erminea muricus of Colorado and M. rixosa eskimo of Alaska, differ in that when the skulls are viewed from directly above those of rixosa have the mastoid processes more prominent, or the braincase is higher in relation to its width or both differences together prevail. Stated in another way, comparison of skulls of equal size of rixosa and erminea shows that in the latter the brain- case is more nearlv flat and is wider above and in front of the MUSTELA RIXOSA 169 mastoid processes; therefore, the greatest breadth of the braincase equals or exceeds the mastoid breadth, whereas the reverse is ordinarily true of rixosa. Geographic variation. — In the Old World four subspecies are currently recognized (see Allen, 1933:316) and the same number is here recognized in North America. Length of the tail, length of head and body and hind foot, breadth of the rostral part of the skull in relation to its length, and position on the side of the head of the line of demarcation between the dark color of the upper parts and the white underparts, are the features in which geographic variation has been detected. The general impression is that the amount of geographic variation is much less than in Mustela frenata and only slightly less than in Mustela erminea of the same geo- graphic area. Nomenclature. — It is exceptional for a species which occurs in both the Old- and New-World to take its specific name from New World material, especially if the name was proposed as recently as 1896; most circumboreal species take their names from descriptions of European specimens. Although the least weasel, Mustela rixosa (Bangs) 1896, seems now to be an exception, it may yet turn out that the first available name was based on European material. Zim- mermann (1943) shows that the least weasel actually was named on the basis of European material long before 1896 and concludes that the name Putorius minutus Pomel, 1853, based on a specimen from France, is the first available name. Because Putorius nowdays is relegated to subgeneric rank under the generic name Mustela, we have for consideration the name- combination Mustela minuta (Pomel). Unfortunately for Zimmer- mann's conclusion, Mustela minuta Pomel is not available because it is preoccupied by Mustela minuta Gervais [=z Palaeogale minuta (Gervais), 1848-1852 — see Simpson, 1946: 2, 12], a name applied to another species of small mustelid from the Oligocene or lower Miocene deposits of Europe. Some other early names thought by Zimmermann (1943:290) to have been based on the dwarf weasel of Europe are judged to be nomina nuda and therefore are to be ignored. The name Mustela minor Nilsson 1820 was thought by Miller (1912:402) to be a renaming, and hence a synonym, of Mustela nivalis Linnaeus. If that is the case the name does not apply to the dwarf weasel. If the name Mustela minor Nilsson was instead based on the dwarf weasel, the name might still be unavailable, de- 170 University of Kansas Publs., Mus. Nat. Hist. pending on rulings on secondary homonyms, because the name might be preoccupied by [Lutra] minor Erxleben 1777 which is a synonym of [Mustela] lutreola Linnaeus 1766. Two names seem- ingly available for weasels, and in use for them today, which might replace rixosa as the name of the species, are, first, Mustela boc- camela Bechstein, 1801, of Sardinia [= Mustela nivalis hoccamela of Miller, 1912, 405] and second, Putorius numidicus Pucheran, 1855, of Morocco and Algeria [= Mustela numidica of Allen, G. M., 1939, 183]. As they stand in the current literature, Mustela numidica is a species distinct from the dwarf weasel and the other name, Mustela nivalis hoccamela, is an insular subspecies of the mouse weasel. Zimmermann (1943:292), however, implies that M. numidica may belong to the dwarf weasel group when he says "Ob auch iberica Barr.-Ham. als Unterart zu 7ninuta Pom. zu stellen ist, soil hier nicht untersucht werden, ebensowenig die von Ca- brera vermutete Zugehorigkeit der grossen nordafrikanischen M. numidica Puch. zur Hberica-Gruppe ". The answer to this prob- lem requires a taxonomic, rather than a nomenclatural, decision. Whether either M. numidica or M. hoccamela are conspecific with the dwarf weasel I cannot at this time ascertain for want of ade- quate specimens. Because these two names, M. hoccamela, and M. numidica, are assigned to kinds of weasels which are currently regarded as specifically distinct from the dwarf weasel, and because all the other names which certainly have been assigned to Old World populations of the dwarf weasel before 1896, so far as I know, are nomina nuda or are preoccupied, the next available name, Mustela rixosa (Bangs, 1896), is here employed. Remarks. — This species may have a wider geographic range in northeastern North America than is now known. Strong (1930:7) writes that the Naskapi Indians of the interior country of Labrador between Hamilton Inlet and Ungava Bay "have only one name for weasel, me-tah-kwut, but they say there are three kinds in their territory, a large, an intermediate, and a very small weasel. The latter suggests the least weasel . . . which has not been re- corded from northern Labrador." In the northern part of the range of the species, the winter pelage is white and the summer pelage is brown. In the southern part of the range, that is in the range of the subspecies allegheni- ensis, the winter pelage is either brown or white and the time of the molt into winer pelage is irregular; each of eleven individuals from Pennsylvania, Michigan and Ohio, taken in December, Jan- MUSTELA RIXOSA 171 uary, February and March is mostly white but retains some con- siderable part of the brown pelage of the previous coat on top of the head and usually also along the midline of the entire dorsum. These eleven animals include individuals of each sex. Of each sex, some are adults and some are subadults. Therefore, the delayed or incomplete fall molt, at present, cannot be correlated with either sex or with any particular age. No wild-taken specimens of M . erminea or of M. frenata of the same region show this delayed or incomplete molt. Possibly this delay or incompleteness of molt is the result of the same cause that lies behind the birth of some M. rixosa in midwinter. As listed below, several litters of young have been found in midwinter. In fact it appears that in the United States, young may be born in every month of the year although, accord- ing to existing information, more litters are produced in spring and in winter than in summer and autumn. Many juveniles and young of aUegheniensis examined in study collections clearly were born in spring but about as many seem to have been born in mid- winter as at any other time (in the light of present knowledge) and this is in contrast to what we know of the two other species of American weasels since their young, so far as known, are born in spring. One instance is worthy of detailed comment. An adult female, no. 783 Ohio State Museum, taken on January 31, 1931, at Vinton, Meigs County, Ohio, bears the following notation on the attached label "nest plowed out of ground. Very small yng escaped — marked like parent. 5 was nursing." The enlarged mammae on the dried skin substantiate the statement that the female was nursing young. She has a brown mask continuous from one ear through the eye, across the forehead and through the other eye to the opposite ear. On each side of the body a stripe of brown 5 to 10 mm. wide extends from the upper part of the foreleg back to the thigh and base of the tail, uniting there with its opposite and covering the tail. There are a few spots of brown on the shoulders, and nmip and one on the middle of the back. Other- wise the specimen is white. One implication of the statement on the label that the young which escaped were marked like the parent (presumably this female parent) is that this female is a partial albino. I am more inclined, however, to the view that there was an unseasonable activity of the particular glands of in- ternal secretion the hormones of which promote embryonic growth 172 University of Kansas Publs., Mus. Nat. Hist. and that these glands, or others controlled by them, were in some way responsible for an abnormal progress of molt, or for a re- versal of molt in that one molt began before the previous molt had been completed. Excepting this one specimen, no. 783 from Vinton, Ohio, all of those in transitional pelage indicate that the direction of the molt pattern is the same as in M. frenata and M. erminea. That is to say, the autumnal molt begins on the midventral line and the molt in spring begins on the mid-dorsal line. Furthermore, the normal progress of each molt appears to follow the same pattern that has been described above for Mustela frenata. A possible explanation of unseasonal molt in the southeastern area of occurrence of the species Mustela rixosa, and a possible explanation of the abnormal molt of the female from Vinton, Ohio, is that the species has only relatively recently invaded the area, and has had insufficient time to adjust the physiology of its molting mechanism to the longer periods of daylight that obtain later in autumn and earher in spring than farther north. In the other two species of American weasels, the change in length of periods of light, it will be recalled, is known to indirectly control both molt and some changes in the sexual cycle. Wright (1942B:109) has shown that molt in spring precedes by one or two months the birth of young in M. frenata, that the two phenomena are correlated in a way that is statistically significant, and recognizes that progres- sively longer periods of dayhght may be the causal stimulus. The suggestion made above that M. rixosa does not hve in New England or in the Rocky Mountains of the western United States because each of the two areas already is inhabited by weasels of almost equally small size, is in Hne with the idea that rixosa is a recent immigrant to America, or more precisely that rixosa arrived later than erminea. Natural History.— Habitat and Numbers.— Soper (1946:136) recounts that near the junction of the Antler and Souris rivers, Mani- toba, this species occurs "both in the river valleys and on the upper prairies", and later (1948:55), with reference to the Grand Prairie of the Peace River region of Alberta, writes that the least weasel "inhabits both parklands and mixed wood forest environments." At most times, wherever found, the least weasel is regarded as rare. Not only mammalogists regard it as rare and as a desirable catch, but Indians likewise value it, probably because of its rarity. For example, Osgood (1901:69-70), who caught a female least MUSTELA RIXOSA 173 weasel at Tyonek, Alaska, writes tliat: "The natives regard the cap- ture of one of these rare animals as a piece of great good fortune. One old Indian who frequently visited our cabin told us that his brother who had caught one when a small boy had in consequence become a Ijig chief; and he assured me that since I had caught one I must surely be destined to become a man of great wealth and power." Swenk's (1926:313-330) account of the species in Clay County, Nebraska, shows, however, that the animal was far more abundant in 1916 and 1917 than subsequently and inferentially than it was before 1916. Clearest proof of multiannual fluctuation is provided by P. O. Fryklund's (Swanson and Fryklund, 1935:120-126) receipt of weasels from Roseau County, Minnesota. From 1895 to 1932 he had approximately equal opportunity to receive least weasels each year. Those which came to his attention were distributed by years as follows: 1895-1927, 7 individuals in all; winter of 1927-28, 3 individuals; winter of 1928-29, 59 individuals; 1929-1930, 84 in- dividuals; 1930-1935, 3 individuals. "These records indicate a very definite increase in the abundance of least weasels in the Roseau region [in] the two years from the autumn of 1928 to the spring of 1930. Mr. Fryklund has handled 166 least weasels in his 40 years in Roseau County, and of these, 143 were taken in the two years mentioned." The maximum home range of the least weasel is two acres and a weasel seldom travels farther than ten rods from its burrow ac- cording to Polderboer (1942:146) who, in the period December 20, 1939, to January 2, 1940, studied four least weasels and one long- tailed weasel on a 144 acre farm in Butler County, Iowa. Behavior Of the voice, Llewellyn (1942:441) records that his captive speci- men taken in Virginia uttered a shrill shriek when seizing prey or when teased. When excessively annoyed the weasel also emitted musk. The sense of smell is used in hunting as was witnessed by George L. Fordyce; he observed a least weasel following the scent of a Peromyscus and saw the least weasel overtake and kill the mouse (Seton, 1929 (2):637). At a nest in a clover stack, in Manitoba, Criddle (1947:69), on December 27, 1946, found the least weasel "to have been rather remiss in its sanitary habits as its pile of dung was almost, or quite, touching the nest and only just to the side of its entrance." There were 117 voids. 174 University of Kansas Publs., Mus. Nat. Hist. Enemies The great-horned owl, barn owl and long-tailed weasel are to be counted as enemies since Nelson (1934:252) found the fur, skull and other fragments of the skeleton of a least weasel in one of 26 pellets of the great-horned owl in Wisconsin; Handley (1949:431) found the skull and other skeletal remains of a least weasel in one of 22 pellets of the barn owl in Virginia; and Polderboer, Kuhn and Hendrickson (1941), in Iowa, found the remains of a least weasel in the den and scats of a Mustela frenata. A domestic cat in Michi- gan killed a least weasel (Dearborn, 1932B:277). Food Mice are killed by the least weasel biting into the back of the head and neck according to Allen (1940:460) who reported upon the growth of five young, from Michigan, that he had in captivity. He further states that a weasel was able to kill a mouse in 30 sec- onds. One large Microtus introduced into the cage slept with a weasel for several days and ate parts of the mice that the weasel killed but then the weasel killed this mouse! Llewellyn (1942: 440-441), in writing of a captive from Virginia, says: "When a live mouse was placed in the cage, the weasel sprang upon it almost instantly. Grasping the mouse by the back of the head, the weasel bit its victim through the skull several times in rapid succession and held on witli its sharp teeth. The sound of the teeth piercing the bone was distinctly audible at a distance of several feet. Dur- ing this interval the weasel hugged the mouse closely with its fore legs and pressed it firmly to its belly through a kicking motion of the hind legs. The hold on the back of the head was not relinquished until the mouse was dead. The killing took only a few seconds. Upon releasing the mouse the weasel usually came to the front of the cage and inspected the observer for an interval of several sec- onds after which it returned to its prey and began its meal at once. Sometimes the blood would be licked from the wound in the back of the head or perhaps an ear would be chewed a bit and the blood licked off, but never did the weasel 'cut the throat' of its prey and 'suck the blood.' "The weasel ate the head and brain first, beginning at the back of the head and working forward. Just before reaching the nose the process was reversed and eating then proceeded from the base of the skull toward the tail of the mouse. The tip of the nose, maxilla with teeth, and the tail seemed to be the parts least pre- MUSTELA RIXOSA 175 ferred; they were not eaten when an abundance of food was pres- ent. At no time did the weasel place its front feet on the mouse in an attempt to hold it. A second or third mouse was killed im- mediately upon being placed in the cage even though the first one had not been consumed. The weasel, however, usually returned to the partially eaten mouse and finished it before starting on a new one. Upon completing a meal, especially if the meal had been particularly bloody, the weasel rubbed its chin on the bottom of the cage, scooting along and appearing more snake-like than normal. Whenever I attempted to remove a mouse, or partially eaten one, from the cage, the weasel hung to the mouse tenaciously, and often allowed itself to be Hfted up in this manner. "In the six days that the weasel was kept in captivity it was fed 10 house mice having a total weight of 118 grams. As no food was given on one day, the amount of food eaten is probably shghtly below the actual capacity of the animal. Since the weasel weighed only about 32 grams, the average amount of food eaten a day was slightly in excess of one-half the weight of the animal." Polderboer (1942:146-147) found in three dens, in Iowa, bits of Reithrodontomys (harvest mice) and Peromyscus maniculatus (deer mouse), and in the digestive tract of one least weasel there was a bone fragment and a few hairs of a deer mouse. In the ac- count, given beyond, of a nest, Griddle (1947:69) records the Penn- sylvania meadow mouse (Microttis pennsylvanicus druminondi) and the Gapper red-backed vole (Clethrionomys gapperi) as prey at Treesbank, Manitoba. The same author, concerning the same place, earlier (1926:199-200) wrote that in 1922 the meadow mouse, Microtus minor, "went into winter quarters in great numbers and its homes were well stocked with provisions. ... all went well until the middle of February, 1923. Then, within a few days, each was taken possession of by a least weasel ( Mustela rixosa ) and the inhabitants were quickly destroyed. One dwelHng was occupied by one of these weasels for about two weeks during which time I observed that it had dragged several mice over the snow to its temporary home. This residence was examined in April, and in it were discovered six dead Microtus minor, one Evotomys, the head of another, and at least six or eight remnants of small rodents in- cluding Microtus drummondi, these last remains being chiefly in- dicated by the hair-lined nest of the weasel. "The homes of 27 other vole communities examined at this time were all found to have been entered by weasels, the inhabit- 176 University of Kansas Publs., Mus. Nat. Hist. ants having been killed and partly eaten. Moreover, the weasels had made the homes temporary centers from which they raided other rodent habitations in the vicinity. Thus from being an abmi- dant animal this vole was reduced to insignificance in the course of a few weeks, while all other kinds of mice had suffered severely from the same enemy." An instance of predation on Peromyscus, reveahng some of the methods of capturing prey, is recounted by Seton (1929 (2):636- 637 ) who quotes a letter to him from George L. Fordyce, of Youngs- town, Ohio, as follows: "While out in the field this morning (Dec. 26), walking along the bank of a ravine at the edge of our golf course, I saw a Field-mouse run out of the bushes into the rough grass that is just outside of the fair-green of the course. In another instant, what I thought at first to be a white Mouse came out at the same place. The Mouse ran into a wheel track, and disappeared under the grass, coming out about 6 feet from where it went in. The white animal followed through the same course, and when it came out, I saw that it was a small Weasel, very little larger than the Mouse, and that it was following the trail of the Mouse by scent. "For a time the Mouse ran in circles, and zigzagged about, often . within 4 or 5 feet of the Weasel; but the latter seemed so intent on the trail, that it did not notice the Mouse to one side. Af- ter a time the latter started toward the open golf course; and when the Weasel reached the point where the trail was straight, it sighted the prey, made a sudden dash forward, and, although 25 feet be- hind, overtook the Mouse while it was going 3 or 4 feet. "For a few seconds, they seemed to fight, until the Weasel got the Mouse by the throat, and started for the bushes, dragging the body. When it came to within about three feet of me, I moved a little to see what it would do. It dropped its victim, and ran into the ravine. The Mouse had a drop of bright red blood in the center of its white throat. I waited near by for 15 or 20 minutes, thinking the Weasel might come back, but it did not show up again; even an hour later, tlie Mouse had not been disturbed." There are two suggestions, but no proof that I know of, in the hterature that the least weasel eats insects. Abbott (1884:27-32- 1st ed., 1884) gives considerable information on the food (some insects included ) of the 'little weasel" which he describes ( op. cit.; 27) as having "a little pointed tail of a uniform brown color". Al- though this suggests Mustela rixosa, Abbott mentions on the next MUSTELA RIXOSA 177 page (page 28) that a specimen of the smaller weasel measured six and a half inches from the tip of the snout to the base of the tail and that the tail itself measured two and a fourth inches to the tip of the last caudal vertebra. These measurements indicate that Miistela erminea was involved. Because of the uncertainty as to the species of Mustela involved, Abbott's interesting data on food, nest and behavior are not recorded in the present work. Seton (1929 (2):636) says that of several least weasels brought to D. Nicholson at Morden, Manitoba, most of them decayed so quickly that they could not be saved as specimens. To Seton this indicated that insects were an important part of the food of the weasels. In summary: Least weasels are known to eat harvest mice, deer mice, meadow mice and red-backed mice; it is suspected that they eat also insects. Reproduction Polderboer (1948:296) has taken six specimens in "northeastern Iowa [in] . . . January and December — all males in winter pelage. None of these males showed signs of sexual activity; in all, the testes were retracted and diminutive in size. ... A male least weasel in brown pelage was taken November 17, 1945, at Marion, Iowa. This specimen had large testes that had de- scended into the scrotum. The testes, when removed, were about the size of medium-sized garden peas. Microscopic examination of the testes and the vasa deferentia showed mature sperms to be present. . . ." On July 1, 1917, in Clay County, Nebraska, a nest with four young was found (Swenk, 1926:321). On July 29, 1939, an adult and five young were plowed out of the ground in Allegan County, Michigan; one of the two young males weighed 40.5 grams two days after capture (Allen, 1940:459-460). On August 12, 1932, ten young with the mother, were found in Roseau County, Minnesota (Swanson and Fryklund, 1935:125). September appears to have been the month of birth of a specimen, no. 8472 in the Carnegie Museum, taken on November 24 in Pittsburgh, Pennsylvania. In October, a young least weasel is recorded from Pennsylvania (Winecoff, 1930:313). Early December was the time of birth of a specimen, approximately 10 weeks old, no. 88077, University of Michigan, taken on February 21 in Allegan County. On Decem- ber 25, 1927, in Washington County, Pennsylvania, "five full-sized, though young . . . animals were caught under the same pile of com fodder" (Sutton, 1929:253). The first week of January 178 University of Kansas Publs., Mus. Nat. Hist. seems to have been the time of birth of a juvenile, no. 88080, Uni- versity of Michigan, taken in Livingston County, Michigan, on March 27, 1943, since the specimen is approximately seven weeks old. On January 15, 1929, in Washington County, Pennsylvania, four young with the eyes yet unopened were obtained from a nest (Sutton, 1929:254). On January 25, 1928, young, the eyes of which may not yet have been open, were taken from a den in Washington County, Pennsylvania, by WinecofiF (1930:313), who records other young having been taken in the same month as well as in Febru- ary. On March 10, a female from North Portal, Saskatchewan, gave buth to four young (Dunk, 1946:392). On April 18, 1916, four young, half grown, were taken in Nebraska (Swenk, 1926:317). On April 2, 1929, three young were found in Roseau County, Min- nesota, according to Swanson and Fryklund (1935:125) who re- mark that: "The Pennsylvania and Minnesota records show that least weasels may be bom any time from July to early February in the northern states." Now, with all of the above records available, it turns out that November, May and June are the only months in which young least weasels have not been reported. Of course some of the young, for which the ages were not specified, were bom in preceding months. Even so, the data now available suggests that, in the United States, young least weasels may be born in every month of the year. The number per htter is 3, 4, 4, 4, 5, 5, and 10, yielding an average of 5. The rate of growth of the young has not been studied enough to allow of judging if it diflFers significantly from that of other species of the genus. Allen (1940:459-460), however, tells us that of the three young females and two young males captured on July 29, 1939, in Allegan County, Michigan, one male that was killed on July 31, 1939, weighed 40.5 grams. The male remaining alive in- creased from 46 grams (August 5) to 62.5 grams on September 20, having eaten 63 mice while in captivity. The females in the period of August 5 to September 4 increased in weight as follows: 41 up to 49 grams; 44 to 50 grams; and 47 to 58 grams, having eaten, by September 26, 60, 64 and 65 mice. Concerning a nest in which young were found, Sutton (1929: 254) writes that on January 15, 1929, near Burgettstown, Washing- ton County, Pennsylvania, an animal was seen to enter a small hole in a creek bank. After the observer dug in a distance of approxi- mately six inches an adult, female least weasel was seen and obtained. Back of the animal, the hole, which turned sharply MUSTELA RIXOSA 179 downward, was full of water. The weasel first seen was a female nursing young. A chamber, to the side of the hole, filled with dead grass, comprised a nest containing four young with the eyes yet unopened. Several nests occupied by adult least weasels or by least weasels that were old enough to shift for themselves have been found. Polderboer ( 1942:145-147) in the winter of 1939-40, on a 144 acre farm in Butler County, Iowa, found four least weasels living, singly, in burrows dug by moles and pocket gophers. The nests therein made by mice were used by the least weasels. Winecoff (1930:312-313) mentions one den in Pennsylvania that contained the remains of only mice, "and not a hint of a feather." Above, in the account of food of the least weasel. Griddle's (1926:199-200) ac- count of the havoc wrought by least weasels among the meadow mice (Microtus ochrogaster minor) has been given. In this ac- count he mentions the fur-lined nests of the weasels that had appropriated the homes of the Microtus. Griddle's (1947:69) later account of a nest at Treesbank, Manitoba, is as follows: "A Nest of the Least Weasel. — When a least weasel finds its way into a locality that has a large number of mice in it, it selects for its home one of their nests that has been made in a well concealed place. This it immediately starts to improve by lining it with hair plucked from its victims before eating them; and as long as sufficient numbers of mice remain in the district the weasel con- tinues adding their hair to the nest, so that the thickness of its walls give one a good idea of the length of time it has been in use. The nest is not only used for sleeping in, as most of the food is consumed in it. Frozen mice are taken in to be thawed out and the weasel carries those it has recently killed in to prevent them getting frozen, or perhaps to have them warm for its next meal. "On January 27, 1946, my son Percy called my attention to a nest that he had just uncovered in a clover stack that we were using. This nest had originally been made by a Drummond's vole, Microtus pennsylvanicus drummondii, and taken from it by the least weasel, Mustela rixosa, the tracks of which had been noticed about the stack yard since the first snow in early November. "The nest had evidently been in use for at least three months and the continual additions made to its walls had been so great that they were nearly an inch thick of hair matted together so closely that it appeared to be felt. The hair alone weighed nearly 22 gm., so that with this for protection the weasel must have been warm and comfortable through the severest winter weather. 180 University of Kansas Publs., Mus. Nat. Hist. m^^m M . r . c i I< i m o 2 t^Vi'-V'vAv'-y/.VI M . r . r I X o s a • specimens examined llllillillll III! M.r. compestri leqheniensis Fig. 28. Map showing occurrences and probable geographic ranges of the sub- species of Mustela rixosa in North America. MUSTELA RIXOSA 181 "In the nest were two red-backed mice, Clethrionomys gapperi, one of which had the base of its skull eaten out. No hair had been removed from either of them, but a Microtus lying in a side tunnel some feet away had the long hair plucked from its back and sides. In and close about the nest were found forty-three front parts of mice skulls which had evidently been discarded be- cause of the sharp teeth in the maxillaries. Seven full stomachs and eleven hind feet of adult Microtus with parts of leg bones were disclosed in, or under, the weasel's bed and a few small bits of skin with hair attached were scattered among the plucked hair of the nest. "This weasel seems to have been rather remiss in its sanitary habits as its pile of dung was almost, or quite, touching the nest and only just to the side of its entrance. It was composed of 117 voids all of which contained much hair and broken bone, "Six other mouse nests found in the same stack, or others ad- joining it, had been thinly lined with hair. One of these had two mice in it, a red-backed with its brain eaten out and a Microtus with some hair plucked from its neck. Another nest contained the front part of a skull with teeth and the hind feet and tail of a red-back. Besides the mice found in the nests seven others were discovered tucked away in side tunnels. One of these mice had most of the hair plucked from its back. Whether all these mice and nests belonged to the same weasel or not I am unable to say, but it is usual for them to have several nests in the area surrounding the one that is used as their headquarters or home." Mustela rixosa eskimo (Stone) Least Weasel Plates 14 and 15 Putorius rixosus eskimo Stone, Proc. Acad. Nat. Sci. Philadelphia, 1900:44, March 24, 1900. Putorius (Gale) vulgaris, Coues, Fur-bearing animals, p. 102, 1877 (part). Putorius rixosus Bangs, Proc. Biol. Soc. Washington, 10:21, February 25, 1896 (part); Merriam, N. Amer. Fauna, 11:14, June 30, 1896 (part). Mustela rixosa eskimo. Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912; Swenk, Joum. Mamm., 7:327, November 23, 1926; Hall, Univ. Cahfomia Publ. Zool., 30:421, March 19, 1929. Type. — Female, age in question, no. 848 in Acad. Nat. Sci. Philadelphia; Point Barrow, Alaska; July 25, 1898; obtained by E. A. Mcllhenny. Type not seen by me. Range. — Alaska and Yukon Territory. See figure 28 on page 180. Characters for ready recognition. — Differs from M. r. pygmaea of eastern Asia in longer tail, averaging 11 rather than 16 per cent of length of head 182 University of Kansas Publs., Mus. Nat. Hist. and body, and in study skins reaching only to heel instead of to point be- tween heel and toes; from M. r. rixosa in shorter tail averaging 16 rather than 19 per cent of length of head and body and not extending beyond outstretched hind feet in study skins; white of underparts extending dorsally as a reentrant angle from upper lip to behind eye, rather than deUmited dorsally by a boundary between white and brown color that extends straight across cheeks from upper lip to side of body well below eye and ear; breadth of rostrum measured across lacrimal processes more, instead of less, than 85.5 per cent of orbitonasal length; from M. erminea of same region by basilar length of skull less than 32; tail less than 50 and lacking black pencil. Description. — Size. — Male: The original describer hsts measurements of topotypes as follows: Total length, 204, 230; length of tail, 28, 31; length of hind foot, 20, 22. Allowing 5 per cent for shrinkage, the hind feet of 5 topotypes yield an average measurement of 23 for the hind foot. Female: Measurements of two topotypes are: Total length, 184, 180; length of tail, 25, 25; length of hind foot, 24, 18. In four other topotypes the hind feet, allowing 5 per cent for shrinkage, yield an average of 21. Color. — Winter pelage all white, rarely with few white hairs in tip of tail but no black pencil; summer pelage with upper parts about Raw Umber and tone 3 of Chocolate pi. 343 of Oberthiir and Dauthenay; underparts white, ex-tending over upper lip, insides of limbs and over all four feet. Line of demarcation between underparts and upper parts extends from upper Up posterodorsally to behind eye down to base of ear, up behind ear for a third or more of its height, and back along side of body. Tail unicolor all around and same color as upper parts. Least width of color of under- parts averaging 83 per cent of greatest width of color of upper parts. Skull. — Based on topotypes; see measurements and plates 14 and 15; weight, 0.82 (0.74-0.93) grams in males, and 0.80 and 0.84 in two females; basilar length, 29.5 (27.6-30.1) in males and 27.8 (27.1-28.8) in females; other- wise as described in M. e. richardsonii. Remarks. — Among the earliest specimens preserved was one by Edward W. Nelson in the course of his explorations of the Upper Yukon, and one in 1874 by L. T. Turner from St. Michaels, Alaska. Bangs, in 1896 (p. 22) mentioned the occurrence of the species in Alaska, but it was not until 1900 (p. 44) that Stone named the subspecies, and then principally on the basis of specimens obtained two years before by E. A. Mcllhenny. The large size, broad skull, light color and short tail are the distinguishing subspecific characters of the race eskimo, and the three characters first mentioned are distinguishing features also of the subspecies of Mustela erminea, namely arctica, which in- habits the same region. Possibly eskimo also will be found on Banks Island and the other Arctic islands between Alaska and Greenland, as is M. e. arctica; at the present time no specimens MUSTELA RIXOSA 183 of Mustela rixosa are known from these islands although some race of rixosa would be expected to occur there. Animals from southern Alaska average slightly smaller than those from northern Alaska, and this decrease in size toward the south probably represents intergradation with M. r. rixosa. Further evi- dence of intergradation is furnished by the short tail of the specimen from 15 miles east of Atlin; in other particulars this specimen agrees with the subspecies rixosa to which it is here referred. Neverthe- less, the short tail, and color pattern, namely reentrant angle of white behind the eye, is to be seen in all Alaskan specimens ex- amined in the brown pelage, even in no. 107591, from Tyoonok on Cook Inlet, which Osgood (1901:69) and Swenk (1926:323) thought might not differ from the subspecies M. r. rixosa. Each of four male topotypes, hardly subadult in age, probably of a single litter, is much larger than any other specimen seen from Point Barrow. The basilar length, for example, is 31.9 as against 29.5, and the weight of the skull (with lower jaws) is as much as 1.5 grams, as against 0.93 in the heaviest of the other males. Initial examination of materials from Point Barrow raised the suspicion that two distinct species were represented — rixosa and a larger one possibly allied to M. nivalis of the Old World. Nevertheless, further study almost completely allayed the suspicion because the only difference discernable is one of size, and it is supposed that additional specimens will bridge the gap in size and show that M. r. eskimo at Point Barrow averages larger than the adult speci- mens now available indicate. The four large males of subadult age are nos. 42814-42816 and 42818 of the American Museum of Natural History. Of the fourteen adult and subadult skulls examined, two display lesions resulting from infestation of the frontal sinuses by nematode parasites. None of the young skulls show such infestation. Specimens examined. — Total number, 42 as follows. Arranged alphabeti- cally by Territory and District and unless otherwise indicated in the United States National Museum. Alaska. Barrow and Point Barrow, 19 {S\ T, 2\ V); Wainwright, 1'; Mts. back of Icy Cape, V; west of Beechy Point, 1'; west edge of Colville River Delta, 1'; Koyukuk River, 16 mi. above Beetles, 1; upper Yukon, 1; Fort Yukon, 1; Stephens Village, 1; Wales, 1"; McDonald Creek, tributary of Salcha Slough, 1; near head of Toklat River, 1; head of Kantishna River, 1; St. Michael, 4 (2'); Tyoonok [= Tyonek], 1; Bethel, 3; vie. Bristol Bay, 1. Yukon. La Pierres House, 1; Klotassin River, tributary of White River, 1. 1. American Mus. Nat. History 4. Collection of W. E. Snyder 2. Mus. Vert. Zool., Univ. California 5. Colorado Museum of Nat\u-al History 3. Acad. Nat. Sciences of Philadelphia 6. Canadian National Museum 184 University of Kansas Publs., Mus. Nat. Hist. Mustela rixosa rixosa (Bangs) Least Weasel Plates 14 and 15 Putorius rixosus Bangs, Proc. Biol. Soc. Washington, 10:21, pi. 1, fig. 6, pi. 2, fig. 6, pi. 3, fig. 4, February 25, 1896; Merriam, N. Arner. Fauna, 11:14, pi. 2, figs. 7, 7a, June 30, 1896. Putorius pusillus, Baird, Mamm. N. Amer., p. 159, 1858. Putorius (Gale) vulgaris, Coues, Fur-bearing animals, p. 102, 1877. Miustela]. rixosa, Thomas, Proc. Zobl. Soc. London, p. 168, March, 1911. Mustela rixosa rixosa, Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912; Swenk, Joum. Mamm., 7:327, November 23, 1926. Type. — Female, adult, skin and skull; no. 642 Bangs Coll. in Mus. Comp. Zool.; Osier, Saskatchewan; July 15, 1893; obtained by W. C. Colt; original no. 79 according to describer. The skull lacks the basioccipital, basisphenoid, and left zygomatic arch. The "crowns" of the lower canines are missing; otherwise the teeth are present and entire. The skin is fairly well made, with soles of hind feet up, in good condition and in summer pelage. Range. — From northern British Columbia and Great Slave Lake south on the west side of the Rocky Mountains to Ootsa Lake, British Columbia, and on the east side of the Rocky Mountains, south to central Montana, North Dakota and Minnesota; eastward in Canada, entirely north of St. Lawrence River, to Atlantic Ocean. See figure 28 on page 180. Characters for ready recognition. — Difi^ers from M. r. eskimo in longer tail averaging 19 rather than 16 per cent of length of head and body and extending beyond outstretched hind feet in study skins, rather than to a point short of tips of toes; boundary between brown upper parts and white underparts extending straight across cheeks from upper fip to side of body well below eye and ear, rather than with reentrant angle from upper lip carrying white upward to point behind eye, and with breadth of rostrum less, instead of more, than 85.5 per cent of orbitonasal length; from M. r. campestris by smaller size: hind foot less than 25 in males and ordinarily less than 22 in females; in males total length less than 216 and tail averaging less than 34, and in females total length averaging less than 182 and tail averaging less than 29; color said to average darker; from M. r. allegheniensis by three average differences, namely lighter color, longer tympanic bullae and larger size of males; from M. frenata and M. erminea of same region by basilar length of skull less than 32; tail less than 50, and lacking black pencil. Description. — Size. — Male: Six adults and subadults from Shaunavon, Saskatchewan, yield average and extreme measurements as follows: Total length, 202 (188-208); length of tail, 32.5 (31.5-34.0); length of hind foot, 22.8 (21-24). Female: One adult and 3 subadults from the same area yield average and extreme measurements as follows: Total length, 172 (162-190.5); length of tail, 27.4 (24-34); length of hind foot, 19.6 (17.5-22). Color. — Winter pelage all white, rarely brown; as described in M. r. eskimo except that fine of demarcation on side of head betwen upper parts and under- parts passes almost straight back without the dorsally directed reentrant area MUSTELA RIXOSA 185 of white behind the eye and ear; least width of color of underparts averaging 52 per cent of greatest width of color of upper parts. Skull (Based on those from Shaunavon, Sask.) — Sec measurements and plates 14 and 15; weight, 0.88 (0.70-0.98) grams in males and 0.55 (0.54-0.56) in females; basilar length, 29.5 (28.4-30.4) in males and 26.1 (24.7-27.0) in females; otherwise as described in M. e. richardsonii. Remarks. — As early as 1858 (p. 159) Baird recognized an indi- vidual of this race from Pembina, Minnesota, as pertaining to a distinct species. Although he used for it tlie specific name pusilltis originally proposed by DeKay for a small weasel from the state of New York, Baird wisely noted that the specimen he described "may be different from the New York species. . . ." After pre- paring this account, Baird included a second specimen, from Fort Steilacoom, Washington Territory, which he thought might be the same, but the differences that he was careful to point out, in tlie light of later knowledge, show it to be of the species Mustela er- minea. Only a few other naturalists followed Baird in distinguish- ing the least weasel as a separate species until Bangs in 1896 (p. 21 ) clearly differentiated it and proposed for it the name Putoritts rix- osus, which continues in use today and applies to the species. The accumulation at the National Museum of Canada, through the energy of Dr. R. M. Anderson, of a good series of specimens from Saskatchewan in the general vicinity of the type locality allows for the first time an adequate conception of the amount of secondary sexual variation and individual variation and permits recognition of subspecific characters to differentiate between M. r. rixosa and the subspecies eskimo and campestris. In comparison with the subspecies allegheniensis the basis for segregation is less clear and will remain somewhat in doubt until additional adults of allegheni- ensis from, say, Pennsylvania, become available with accurate ex- ternal measurements taken in the flesh and especially with com- plete skulls. Intergradation with the subspecies eskimo is suggested by the short tail of the specimen from fifteen miles east of Atlin, British Columbia; in other particulars that specimen, a skin-alone, agrees with the subspecies rixosa. Intergradation with campestris is indi- cated by increased size of some specimens from North Dakota, and is suggested with allegheniensis by the color of specimens from Wisconsin and Illinois. Three specimens from Winona County, in southeastern Minnesota, unfortunately are skulls-alone without external measurements. Also, two of these skulls are of young animals. The one adult, unsexed, is from Crystal Springs. 186 University of Kansas Publs., Mus. Nat. Hist. Selected cranial measurements are: basilar length, 28.5; length of tympanic bulla, 10.9. These measurements accord with those of males of the subspecies rixosa to which the specimens from Winona County, therefore, are here assigned. The possibilities have not been excluded, however, that the adult is an unusually large female of the subspecies campestris or a male of allegheniensis that has tympanic bullae longer than average for that subspecies. Some hesitation is felt in assigning the specimens, 8 in all, from eastern Canada to the subspecies rixosa. The skin-alone from Eagle River and the skin, with part of the skull, from St. Michael Bay, are in transitional pelage and are of no help in appraising subspecific characters. The one adult specimen which does have a complete skull is from an island south of the Comb Hills. This animal in all respects agrees with selected individuals of M. r. rixosa from Saskatchewan, but each of the five other skins in sum- mer pelage has spots of dark browai color on the breast. Only about one specimen in three of rixosa from Saskatchewan is simi- larly marked. Furthermore, on some of the specimens from eastern Canada the spots are larger than on any of the animals from farther west. The greater frequency of brown spots on the breast, the larger average size of these spots, and the darker average coloration of the upper parts are suggestive of geographic variation, the existence of which has to be proved by additional and more complete specimens from eastern Canada. For the time being, specimens from there are tentatively assigned to the race rixosa. Of 56 subadult and adult skulls only 3 ( 1 North Dakota; 1 Cal- gary, Alberta; and 1 Island S Comb Hills, Queb.) display lesions resulting from infestation of the frontal sinuses by nematode para- sites. None of the young skulls shows such infestation. Specimens examined. — Total number, 87 as follows. Arranged alpha- betically by provinces and states and within each from north to south. Unless otherwise indicated, specimens are in the United States National Museum. Alberta. Miette River, 1'; 5 mi. NW Camrose, 1'; Camrose, 2 (1\ T); "near Camrose", 2'; Fks. Blindman and Red Deer rivers, 1^ Innisfail, 1*; Vet- eran, V; Diddsbury [= Didsbury], 1; Calgary, 2 (1°, 1"); Shepard, 1\ British Columbia. Clarks Ranch, Halfway River, Peace River Dist., 1*; 15 mi. E Atlin, 1"; Wistaria, P. O., 3 {2\ V); Ootsa Lake, 1". Labrador. Davis Inlet, 1^; 30 mi. upriver and 20 mi. toward Groswater Mts., Eagle River, 1; St. Michael Bay, 1. Mackenzie. Old Fort Rehance, V; Fort Resolution, 2; Fort Smith, 1. Manitoba. Gypsumville, 1*; Lake St. Martin Reserve, 1*. Minnesota. Roseau County: Cedarbend, 2'; Grimstad, 1'; America, 2 (1', 1'°); Malung, 1^°; Norland, 1"; Falun, 3 ( 1', 1", 1"); Palmville, 1"; Spruce, 1"; Stokes, 1'". No locahty more definite than Marshall County, 1'. Clay Co.: Moorhead, V. Winona County: "near" Whitman, 1"; Altura, 1"; Crystal Springs, 1". MUSTELA RIXOSA 187 Montana. Sun River Valley, 1; Wibaux in Wibaux County, 1. North Dakota. Walsh Co.: Grafton, 15 (3', 1°, 5", 2\ 1", V\ 1"). Mc- Henry County: 4 and 4M mi. N Upham, 2. Wells Co.: 1". Morton Co.: Mandan, 1*. Ontario. Algoma Dist.: Tatnall, near Oba, 1*. Moose Factory, 1". Quebec. Island S of Comb Hills, James Bay, 1". Saguenay County: Na- tashkwan, 1. Saskatchewan. Osier, 1"; "near Regina," 1'; Dollard, 2'; Shaunavon (and "near" and 1 ™. NE), 9'; Klintowel P. O. (about 15 mi. N of Eastend), 1'; Eastend and "near" Eastend, 2\ Mustela rixosa allegheniensis (Rhoads) Least Weasel Plates 14, 15 and 41 Putorius allegheniensis Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1900: 751, March 25, 1901. Putorius rixosus allegheniensis, Cory, Mamm. Illinois and Wisconsin, p. 378, 1912. Mustela allegheniensis. Miller, U. S. Nat. Mus. Bull., 79:96, December 31, 1912. Mustela rixosa allegheniensis, Swenk, Joum. Mamm., 7:328, November 23, 1926. Type. — Probably male adult, skin and skull, no. 6195, Acad. Nat. Sci. Philadelphia; near Beallsville, Washington Co., Pa.; about 1885 or 1886; obtained by Robert Hawkins. Type not seen by me. Range. — Wisconsin, northern Illinois, northern Indiana, Michigan, Ohio, Pennsylvania east to Dauphin County and south in the mountains to north- western North Carolina. See figure 28 on page 180. Characters for ready recognition. — Distinguished from M. r. rixosa by three average differences, namely, darker color, shorter tympanic bullae, and smaller size of males; from M. r. campestris in smaller size: hind foot less than 25 in males and less than 22 in females; in males total length less than 216 and tail averaging less than 34, color averaging darker; from M. frenata and M. erminea of same region by basilar length less than 31, tail less than 45, and lacking black pencil. Description. — Size. — Male: An adult or subadult from Fair Oaks, Pa., a subadult from Finleyville, Pa., and an adult from Huttonsville, W. Va., measure, respectively as follows: Total length, 206, 194, 191 (average 197); length of tail, 37, 32, 28 (32); length of hind foot, 23 in each. An adult from Roanoke, Indiana, weighs 40.6 grams. 1. Canadian National Museum 10. 2. Collection of Kenneth Racey 11. 3. Field Museum of Natural History 12. 4. Royal Ontario Museum of Zoology 13. 5. Univ. Kansas Mus. Nat. History 14. 6. American Mus. Nat. History 15. 7. Collection of Alex Walker 16. 8. California Academy of Sciences 17. 9. Provincial Museum of British Columbia 18. Mus. Vert. Zool., Univ. California Collection of Ralph Ellis Collection of O. J. Murie Acad. Nat. Sciences of Philadelphia Museum of Zoology, Univ. Michigan Museum of Comparative Zoology Carnegie Museum University of Minnesota Collection of Mr. Green 188 University of Kansas Publs., Mus. Nat. Hist. Female: Two young from Leasuresville, Pa., and Middle Paxton Twp., Pa., measure, respectively, as follows: Total length, 188, 172; length of tail, 33, 30; length of hind foot, 20.5, 21. An adult from Monroeville, Ohio, weighs 40.5 grams and a young individual from Middle Paxton Twp., Pa., 39.3 grams, and a subadult from Swan Creek Exp. Station, Allegan Co., Mich., weighs 49 grams. Color. — Winter pelage either all white, or brown as in summer; upper parts about Raw Umber, or tone 2 of Carbo Brown of pi. 342 of Oberthiir and Dauthenay. Underparts white at least on thoracic region; approxi- mately three-fourths of specimens with brown rictal spot at angle of mouth or with this area covered by brown upper parts which extend down on each side and meet on the vmderparts in about one specimen out of three; upper lips and hind feet ordinarily brown; toes of forefeet ordinarily white (see under remarks for details of color pattern). Least width of color of under- parts in the specimens in which the dark color of the upper parts does not encircle the body averages 60 per cent of greatest width of color of upper parts, or including all specimens the percentage is 42. Skull (based on specimens from Pa. Hsted in table of cranial measurements, which see and plates 14 and 15). — Basilar length 29.7 and 28.6 in male and 28.0 in female; weights unavailable; otherwise as described in M. e. rich- ardsonii. The length of the tympanic bullae seems to be actually less, and less in relation to the basilar length, than in other American subspecies of M. rixosa. Remarks. — Robert Kennicott's mention in 1859 (p. 245) of what seems to be this subspecies is the earhest reference to it that I can identify in the hterature. He used the specific name pusillus and it was not until 1900 that Samuel N. Rhoads proposed the name Putorius allegheniensis. Since 1900, several records of occurrence have been published which have made the geographic range of this race better known. An adequate number of specimens has been gathered only from Ohio and from western Pennsylvania. Many from Ohio are with- out accurate external measurements taken in the flesh. The ma- jority of the specimens from Pennsylvania owe their preservation to the willingness of local officials, who pay bounties on weasels, to save the skins of Mustela rixosa. These specimens ordinarily comprise the skin with locality but because the feet, external measurements in the flesh, and skulls are unavailable, the material is far from adequate and to give an accurate notion of the usual or average cranial characters of allegheniensis in Pennsylvania, skulls from there are especially desirable. A smaller percentage of the specimens from Ohio than from Penn- sylvania have the brown color of the upper parts meeting on the underparts. Also, more of the specimens from Ohio are lighter MUSTELA RIXOSA 189 colored and this suggests intergradation with the subspecies cam- pestris and rixosa to the westward. From Pennsylvania 23 animals in brown pelage are available. In 5 there is a rictal spot at the angle of the mouth; in 5 the area is white and in 13 the brown color of the upper parts is continuous over the area in question. Only 2 of 23 have the upper lips white. Eight have the color of the upper parts meeting on the venter thus restricting the white of the underparts to the chin, throat, and pec- toral region, and 6 of these have a white area in the inguinal region as well. The toes of the forefeet are white in 3 of 4 animals suit- able for examination in this regard and the hind feet are marked with white in 3 of the 8 animals which have the hind feet preserved. Mustela rixosa in Pennsylvania parallels the species Mustela frenata in that in this relatively humid area of the northeastern United States the color of the upper parts is darker and the area of the dark-colored upper parts is increased at the expense of the area of the light-colored underparts. Also Mustela erminea in this same region ( range of the subspecies Mustela cicognanii ) shows the same tendency to darker color of upper parts and their extension in area at the expense of the area of the light-colored underparts, or was men- tioned above. It is difficult to account for the seeming absence of the species from New England and all that part of Canada and the United States south of the St. Lawrence River and northeastward from Pennsylvania. The size of females of M. erminea cicognanii in that territory is so little more than in rixosa that the latter possibly can- not successfully compete with the erminea stock which may already occupy the ecologic niche to which rixosa is adapted. It will be remembered that in western North America in territory seemingly climatically suitable for rixosa it occurs no farther southward than the line below which M. erminea has become reduced to a size comparable with that of M. rixosa. Of 41 subadult and adult skulls assigned to this subspecies 24 have obvious lesions in the frontal sinuses evidently resulting from infestation by nematodes. More in detail, none of the specimens from Illinois (3 individuals), Pennsylvania (3 barely subadult), or West Virginia (2) displays lesions. From Wisconsin, Indiana, Vir- ginia and North Carolina there is one specimen each and each specimen displays lesions. From Ohio, 17 of 23 specimens dis- play lesions. From Michigan 3 of 8 specimens display lesions; 2 adults and one subadult have lesions and 5 subadults do not have lesions. 190 University of Kansas Publs., Mus. Nat. Hist. Specimens examined. — Total number, 102 as follows: Arranged alpha- betically by states and within each state by counties from north to south. Un- less otherwise indicated, specimens are in the United States National Museum. Indiana. Huntington Co.: Roanoke, 1. Wells County: Harrison Town- ship, 1'. Illinois. Lake Co.: Deeriield, 3'; no locahty more definite than county, 1'. Cook Co.: Northfield, 1^ La Grange, 1'. Michigan. Tuscola County: 8 mi. N Caro, 1\ Santilac County: Deckerville, V. Allegan County: Swan Creek Exp. Station, 1*; Swan Creek Farm, 1'; T. 2N, R. 14W, 1'; Allegan, 1\ Livingston County: George Reserve, 1*; J2 mi. N Unadilla, 1. Oakland County: Rochester, V. Macomb County: Romeo, V. Wa.shtenaw County: 5 mi. SW Ann Arbor, 1'. Branch County: vie. Coldwater, 1\ North Carolina, "near Marshall," 1. Ohio. Northern part of state, 1*. Williams Co.: Stryker, l^ Lucas Co.: Monclova, V. Erie Co.: Sandusky, 2'; marsh near Sandusky, 1'; Berlin Heights, 1'; no locality more definite than county, 1°. Wood Co.: 10 mi. NE Bowfing Green, V; Bowling Green, 4'; 3 mi. E Bowling Green 1'; Plain Township, 1°; Portage Townsliip, l^ Loraine County: Welfington, 1*. Huron County: west of Monroeville, V. Summit County: Ira, 3*. Portage County: SufBeld, l\ Hancock County: Vanburen, 1^; Findlay, 1*; 9 mi. S Findlay, 1^ no locality more definite than county, 7 (2^ 2*, 3°). Mahoning Co.: Ellsworth, 1. Craw- ford County: "near Crestline", 1*. Delaware Co.: Sunbury, 1°; Lewis Center, 1*; no locality more definite than county, 1*. Licking Co.: Johnstown, 1°. Fairfield County: Baltimore, 1*; Violet Township, 1*. Meigs [= Gallia?] County: Vinton, 1*. Pennsylvania. Erie County: McKeen Twp. 1. Crawford County: Spring- boro, 1°; Pymatuning Swamp, between Hartstown and Shermansville, Sads- bury Twp., 3'. Mercer County: Shenango Twp., 1. Lawrence County: Little Beaver Twp., 1. Butler County: Leasuresville, 1'; Clearfield Twp., 1; Valencia, 1'. Armstrong County: Ford City, Burrell Twp., 1. Indiana County: Smicks- burg, 1; N. Mahoning Twp., 2; White Twp., 1. Allegheny Co.: South Hills, Pittsburgh, 1'; "near Pittsburgh", 1'; Fair Oaks, T. Westmoreland County: Bolivar, 1. Dauphin County: Middle Paxton Twp., 1. Washington County: Finleyville, 1; Rea, 5; Beallsville, 1°; Claysville, 1. Green Co.: Deep Valley, 1; Waynesburg, 1; Jefferson, 1; Cumberland Twp., 1. Fayette County: Acme, 1'; Somerset County, 1. Lancaster County, 1. West Virginia. Randolph Co.: Huttonsville, 1. Wisconsin. Sauk County: Sumpter Twp., 1". Dodge Co.: Beaver Dam, 1*. Dane Co.: Madison, 1; McFarland (= MacFarland), 1. Mustela rixosa campestris Jackson Least Weasel Plates 14 and 15 Mustela campestris Jackson, Proc. Biol. Soc. Washington, 26:124, May 21, 1913. P[utorius]. pusillus, Aughey, Sketches of the physical geography and geology of Nebraska, p. 119, 1880, Omaha. Mustela rixosa campestris, Swenk, Joum. Mamm., 7:329, Nov. 23, 1926. Type. — Female, adult, skin and skull; no. 171490, U. S. Nat. Mus., Biol. Surv. Coll.; Beemer, Cuming County, Nebraska; April 18, 1911; obtained by G. Sharp; x catalogue no. 8440. 1. Museum of Zoology, Univ. Michigan 5. American Mus. Nat. History 2. Field Museum of Natural History 6. Acad. Nat. Sciences of Philadelphia 3. Collection of E. J. Koestner 7. Carnegie Museum 4. Ohio State Museum 8. Collection of W. E. Snyder MUSTELA RIXOSA 191 The skull is unbroken. On tlie left side, CI and P2 are missing; the other teeth are present and entire. The skin is excellently made and in a good state of preservation. Range. — South Dakota, Nebraska and Iowa. See figure 28 on page 180. Characters for ready recognition. — Differs from M. r. rixosa and M. r. allegheniensis in larger size: Hind foot more than 25 in males and ordinarily more than 22 in females; in males total length more than 216 and tail aver- aging more than 34; color possibly slightly paler than in M. r. rixosa and averaging paler than in M. r. allegheniensis; from M. frenata and M. erminea of the same region by basilar length less than 32; tail less than 50, and lacking black pencil. Description. — Size. — Male: Four adults from Nebraska yield average and extreme measurements as follows: Total length, 231 (225-237); length of tail, 36 (32-39); length of hind foot, 29 (28-31). Female: Six adults from Nebraska yield average and extreme measurements as follows: Total length, 192 (184-225); length of tail, 35 (28-40); length of hind foot, 23 (20.5-26). Color. — Winter pelage ordinarily white; as described in M. r. eskimo ex- cept possibly paler and certainly with line of demarcation on side of head between upper parts and underparts passing almost straight back without the dorsally directed reentrant angles of white behind the eye and ear; least width of color of underparts in four specimens from Nebraska averaging 80 (49-89) per cent of greatest width of color of upper parts, but in a fifth animal in summer pelage the brown color of the upper parts encircles the body. Skull. — See measurements in table and plate 15; weight 1.1 grams (male from Brown Co., S. D. ); basilar length, 30.7 in male from Clay Co., Neb., and 28.8 in female from same county; otherwise as described in M. e. richardsonii. Remarks. — In his revisionary treatment of the American races of Mustela rixosa, Myron H. Swenk (1926:313) credits Samuel Aughey with recording this animal, M. r. campestris, from Nebraska, as early as 1880, under the name Putorius pusillus. In 1908, Swenk recorded the animal from the same state under the name rixosus and in 1913 the race campestris was formally named by H. H. T. Jackson. On the testimony of a friend who had previously obtained several specimens for him, Swenk (1926:321) records the least weasel from Oshkosh, Garden County, Nebraska, which is a marginal record of occurrence to the southwest for M. r. campestris. At an early stage in the study of American weasels the writer examined the specimens from Nebraska saved by Mr. Myron H. Swenk and recorded measurements of them. However, at the time of writing this account the specimens were not available for exami- nation and the account of coloration is accordingly incomplete. The large size, particularly the large external measurements, comprises the principal distinguishing character of this subspecies of the least weasel. 192 University of Kansas Publs., Mus. Nat. Hist. Of the four adults examined from Iowa and South Dakota one exhibits lesions such as result from infestation of the frontal sinuses by nematodes. Specimens examined. — Total number, 21 as follows. Arranged alphabeti- cally by states and by counties, from north to south in each state. Unless other- wise indicated, specimens are in the United States National Museum. Iowa. Howard Co.: Chester, 1\ Palo Alto Co.: Emmetsburg, P. Kassuth County: Algona, 1". Clayton Co.: National, 1. Storey County: Nevada, 1". Wapello Co.: Ottumwa, V. Henry Co.: Mount Pleasant, 1'. Nebraska. Holt County: Page, 1*. Madison County: Norfolk 1*. Cuming Co.: Beemer, 1. Hamilton County: Chapman, 1*. Clay County: Inland to 1 mi. east thereof, 7*. South Dakota. Brown County: shore of Sand Lake, S.15 T.126N, R.62W, 1. Day County: Waubay Migratory Waterfowl Refuge, 1. McCook County: Salem, T. 1. Coe College 4. Collection of Myron H. Swenk 2. Iowa State College 5. University of South Dakota 3. Iowa Wesleyan College MUSTELA FRENATA 193 MU STELA FRENATA Lichtenstein Long-tailed Weasel (Synonymy under subspecies) Type. — Mustela frenata Lichtenstein, DarstcUung neuer oder wenig be- kannter Saugethiere, pi. 42 and corresponding text unpaged. 1832. Range. — From southern Canada southward over all of the United States, Mexico, Central America, Venezuela, and the republics of western South America to southern Peru and extreme northern Bolivia. All the life-zones from Alpine Arctic to Tropical are inhabited. In the extremely desert region of southeastern California and western Arizona the species is scarce or possibly absent although recovery of a skull (see under account of M. f. neomexicana) from near the center of this region at Potholes on the Colorado River, and a reported occurrence in the mountains of Baja California, Mexico, indicate that a few individuals of the species live in favorable habitat even in this desert region. Characters for ready recognition. — Differs from Mustela erminea, in regions where the two species occur together, by tail more than 44 per cent of length of head and body and by postglenoidal length of skull less than 46 per cent of condylobasal length in males and less than 48 per cent in females (see under characters of the spe- cies); from Mustela rixosa by presence of black pencil on tail, caudal vertebrae more than a fourth (%-%) of length of head and body, basilar length of skull more than 34 mm.; from Mustela africana by absence of thenar pad on forefoot, underparts without longitudinal, median, abdominal stripe of same color as upper parts, upper lips narrowly (rather than broadly) edged with color of underparts, longest facial vibrissae extending to or behind posterior margin of ear; presence of p2; more inflated (see pis. 23 and 30) tympanic bullae. Characters of the species. — Size large: Total length 300 to 550 mm.; tail two-fifths to seven-tenths of length of head and body, with distinct black pencil at end; caudal vertebrae 19 to 23; skull with long precranial portion; postglenoidal length, expressed as a per- centage of the condylobasal length, less than 47 in females and ordi- narily less than 46 in males; upper parts brown; light-colored under- parts, in summer pelage, tinged with buffy or yellowish and continu- ous from chin to inguinal region; some subspecies (southwestern United States, Mexico, Central America, and Florida) with white or yellowish facial markings which do not occur in any other Ameri- can species of the genus Mustela. 7—3758 194 University of Kansas PxmLS., Mus. Nat. Hist. Geographic variation.— Forty-two subspecies are recognized, and the species is geographically more variable than any of the other 3 American species. Color, color-pattern especially on the head, relative proportions of the tail, hind feet, body including the head, and shape and size of the skull are the principal features in which geographic variation has been noted. The variation in the skull extends to the basicranial region ( shape and size of tympanic bullae and related structures), interorbital region and preorbital region. Natural History.— Rahitat and Numbers.— As has already been remarked, the long-tailed weasel is absent from the extreme desert of the southwestern United States and northwestern Mexico. Pos- sibly the absence of water to drink is the limiting factor. In south- em Nevada the finding of weasels only in places that were well watered, even though small rodents suitable as food for weasels were even more abundant in the surrounding desert, supports this possibility that the absence of water to drink is the limiting factor. Also at Berkeley, California, in early December of 1927 in the can- yon at the head of Dwight Way and in the autumn and winter of 1928 in Strawberry Canyon on the campus of tlie University of California, I trapped extensively for this species in different habitats and obtained, in all, four individuals no one of which was farther than 10 feet from water. The lesser cruising range of the individual weasel than of, say, the coyote, probably explains why, in. an arid region, for example Pahranagat Valley, Nevada, only the meadow mice and their riparian associates are preyed upon by the long- tailed weasel whereas the coyote preys upon these riparian rodents and also upon the kangaroo rats and other rodents which are so abundant in adjoining habitats that are devoid of water. In areas where water is available every few hundred yards, no particular habitat seems to be avoided in summer providing there is food for the long-tailed weasel. In winter (January and March) there obviously was a choice of habitat, possibly occasioned by more abundant food or more satisfactory shelter, or both, in Centre County, Pennsylvania, where Glover (1943B) found the popula- tion density in the chestnut-oak habitat to be one weasel per 6.5 acres in areas of tree cuttings and slash and one weasel per 13.3 acres in the open forest. In the scrub oak-pitch pine forest type the population was one weasel per 26.4 acres in tree cuttings and slash and one weasel per 38.2 acres in tlie open forest. No weasel was found in an area of 9.6 acres comprising a wood lot, the edge of the forest, abandoned fence rows and an abandoned orchard. The two types of forest in which he did find weasels, 25 in all, MUSTELA FRENATA 195 comprised 381.6 acres. Glover's (op. cit.) data is the only precise information known to me on actual numbers of long-tailed weasels in a given area of any considerable size. Fluctuations which I elsewhere (1946:57) have designated as multiannual fluctuations occur in this species but seemingly not in the degree that they do in Mustela erminea. This difference be- tween the two species is to be expected because M. frenata does not range so far northward toward the polar regions as does M. er- minea and populations of most kinds of animals in the polar, at least in the arctic, regions are subject to more extreme and more regular fluctuations than are kinds of animals in temperate or tropical regions. Indication of the means by which decrease in the weasel population is brought about is afforded by Osgood's (1935: 156) observations around Rutland, Vermont. In the late winter of 1934, tracks indicated that weasels left their usual haunts and hunted cross lots, vainly trying to find food. Testing of the small mammal population in the spring and summer of 1934 showed that it was at low ebb. In the fall of 1934 mice and shrews were abundant again but weasels seemed to be entirely absent. The decrease in the population of weasels lagged behind the decrease in the population of the herbivorous prey as did the subsequent in- crease; this, of course, is the normal relation of carnivorous species of mammals and their prey, at least in and above the Transition Life-zone. The average distance away from the central den which four weasels (sex unspecified) traveled in a single night at Ames, Iowa, was 312 feet; the maximum distance was 642 feet. These data were obtained in the winter of 1939 by Polderboer, Kuhn and Hendrickson (1941:115) who studied the tracks in the snow. In Manitoba, Griddle and Griddle (1925:143) noted that a female which lived in their basement often wandered more than half a mile away in search of food. In Michigan, Quick (1944:75) found the maximum distance traveled in one day ( = night? ) by a large male to be 3.43 miles although two miles was the average distance traveled by this individual. In 1942, from January 4 to March 4, in Gentre Gounty, Pennsylvania, Glover (1943B) studied tracks of 11 males and 10 females, in newly fallen snow, and ascertained that the distance traveled in a single night averaged 704 ( 60-2535 ) feet for the male and 346 ( 20-1420 ) feet for the female. The weasels in the open timber traveled farther per trip than those in the brushland and dense stands of trees. 196 University of Kansas Publs., Mus. Nat. Hist. Behavior An adult female (now the holotype of Mustela frenata neva- densis) seen running across a field, and, I think, unaware of my presence, at every bound bent her back up so far that she re- minded me of a measuring worm. For part of the time when run- ning, the tail was held ofiF the ground straight out behind, and then, for a while, inclined upward at an angle of about 45°. An- other weasel that I saw in the daytime, and that I think was un- aware of my presence, was bounding along among the Baccharis bushes on the south-facing slope of Dwight Way Canyon, Berkeley, California. This individual, at each bound, arched the back up so high as to remind me, again, of a measuring worm. The long-tailed weasel is a land mammal and unlike its close relative, the mink, is seldom seen in the water. That it can swim, however, is attested by the capture of one while it was swimming across the Rio Ramos in Mexico (Davis, 1944:381). Also, Green (1936), in May, in Gratiot County, Michigan, saw a weasel, nm- ning with a Peromyscus in its mouth. The weasel dropped tlie mouse, entered the water and swam to a hole among stones. More instances of climbing, than of swimming, have been re- ported in the literature for the long-tailed weasel. Seton (1929 (2):625) quotes William M. GraflBus of Pennsylvania as having seen a weasel closely pursue a red squirrel nearly to the topmost branch of a large hemlock. When the squirrel loosed its hold and dropped into a stream, the weasel descended to the ground and caught and killed the squirrel when it emerged from the water. Pearce (1937:483), in central New York State, on July 29, 1931, watched a weasel chase a chipmunk up a black cherry tree ten inches in diameter, and noted that the first rush carried the weasel "straight up the trunk for approximately 10 feet, where it hesitated momentarily before continuing. Then, instead of climbing verti- cally, it made progress by traveling in short ascending spirals around the trunk, scarcely making 3 feet in height for each circuit of the tree. Upon reaching the limb by which the chipmunk escaped, the weasel followed out along this in the same spiral manner. This limb had a diameter of about 4 inches at its base and extended upward at an angle of perhaps 20 degrees above the horizontal. ... it made its way head first almost down to the ground, using the same spiral mode of progress, but at a leisurely pace . . . While traveling down the side limb it ap- peared practically to wrap its sinuous body around the limb." A male long-tailed weasel, from Colorado, which I kept captive MUSTELA FRENATA 197 was often fed freshly killed mice. These I thrust through one of the small openings in the wire mesh. The weasel quickly learned to seize any part of a mouse tlius introduced and his tugging aided in getting the mouse into the cage. Occasionally a mouse too large to be got through the mesh had to be withdrawn. In such an in- stance, if the weasel had already had hold of the mouse, he would screech frightfully. I have heard no other vocal sounds from a weasel except a kind of purring. The sense of smell apparently is well developed; at any rate it is keen enough to allow the weasel to follow the trail of an in- tended victim by the scent left by the latter. Murie's (1935:321- 322) account, for example, of a weasel pursuing a snowshoe rabbit gives clear evidence that the weasel relied on scent in following the rabbit. A captive male weasel obtained at Gainesville, Florida, stamped his hind feet when annoyed (Moore, 1945:259). A male from Colorado that I kept for months in a cage at La- fayette, California, was several times found in a sleep so deep that he was awakened with difficulty. Seton (1929 (2):629-630) writes: "In my small menagerie, I have had half-a-dozen Weasels of the New York species. Their sleeping dens are arranged so as to be easily and silently opened. Several times I have lifted the lid to find the weasel in a deep sleep — a sleep so profound that I had to poke him vigorously with a stick before he awoke, looked up, and rushed forth with a little puflF of wrath, and a little puff of smell." Feces and urine were ordinarily deposited in one particular place by each of the captive weasels that I have observed. Hamilton (1933:294) records that a large male M. /. noveboracensis, in a week, averaged 10 evacuations every twenty-four hours, that urina- tion immediately precedes defecation, and describes the feces as black or brown, long and narrow and often spiral-shaped owing "to the matted fur of some rodent that had been eaten". Quick (1944:77) writes, concerning four winter dens in Michigan, that "The latrines of weasels were in the entries of used dens and scats could be collected there by the handful." Polderboer, Kuhn and Hendrickson (1941:116) in the spring of 1939 at Ames, Iowa, gathered scats "from latrines found at the entrances of burrows and from latrine chambers found within burrows." Scats were found by them in the linings of some nests. Courage of a high order might be credited to the long-tailed weasel because individuals have attacked animals much larger than the weasels. Actually, however, in few if any of these instances was the motive for attack known. That a hawk was attacked is 198 University of Kansas Publs., Mus. Nat. Hist. suggested by Soper's (1919:45) account of Mitstela frenata nove- boracensis wherein he repeats a story told to him of a hawk observed in unsteady flight, and obviously in distress, which when it plum- meted to earth was with a weasel which escaped from the observer. Charles Tatham, Jr., of Cambridge, Massachusetts, according to Seton (1929 (2): 630, 631) observed one that attacked his dog. Persons and long-tailed weasels have figured in some rather strange encounters. For example, Oehler (1944:198) recounts that in the autumn of 1940 at Cincinnati, Ohio, an animal, mis- takenly thought to be a chipmunk, was seen to dash into a hollow log whereupon pounding on tlie log brought out the weasel which bit and clung to tlie hand of one man whose companion was bitten when he attempted to free the man that was bitten first. Seton (1929 (2): 631) writes that on the night of September 5, 1897, on Roosevelt's old ranch, near Medora, North Dakota, a man turned over his saddle (which was lying on the ground) to dis- lodge what was thought to be a pack-rat. The animal was a long-tailed weasel which attacked him. It ran up his legs a num- ber of times aiming at his throat before being killed by a dog. Criddle and Criddle (1925:146) wrote: "August 20, 1919.— A longicauda in the Insectary ran at me this morning apparently with a view to intimidating. It uttered a shrill cry while making the attack, but retreated after advancing within two feet." The same authors (op. cit.: 147) further write that a "Long-tailed Weasel was caught in a trap set for gophers, and, on being released by Miss M. Criddle, at once turned upon its liberator and bit savagely at her boot. It then moved a short distance away to a tub of water, where it drank thirstily, merely glancing at the observer from time to time while doing so, and then ran ojff out of sight. "Mr. T. Criddle records a similar experience. After liberating a large weasel from a trap, it immediately rushed at him and persisted in its attack with such ferocity that it was three times picked up and thrown, on each occasion to a greater distance, be- fore it finally abandoned its offensive. "We have no record of a weasel making an unprovoked attack upon anyone." Wight (1932: 164) in Michigan, detected a weasel attacking a hen. The weasel fled at Wight's approach but returned and at- tacked him several times. Finally the weasel went around Wight to reach the hen. In Wight's words "There was no evidence of in- furiation, but rather a well directed offense at the one object, re- gardless of its size, which stood between the weasel and an oppor- MUSTELA FRENATA 199 tunity to satisfy its desire to kill, which was probably based upon the uncontrollable urge of hunger pangs." Weasels of each of the three North American species have been successfully kept in captivity. A type of cage satisfactory for keeping the animals in the laboratory is described by Bissonnette and Bailey (1940:761-763). Some of the captives used their teeth to break glass water-containers and to gnaw slivers of wood from the cages. Ingested slivers of wood and bits of broken glass caused the deaths of some of the captives. Weasels kept by me all were of the species Mustela frenata. They thrived on a meat diet but I was always careful to give them, every few days, if not each day, some small rodents entire, thinking that the bits of bone and fur ingested might, in some way unknown to me, keep the di- gestive tract in better condition than would flesh devoid of hair and bone. Three young weasels approximately the size of mice, in the Okefinokee Swamp of Georgia, were obtained by a hunter who, according to Harper (1927:303), raised them by feeding "milk for a few days, and then fresh meat." Litters of young born in captivity have been successfully raised by the mothers (Hamilton, 1933) and success in getting the animals to breed in captivity and to rear their young is recorded by Wright ( 1948A ) . He has found, however, that the majority of his captive adult males show no interest in mating when placed with females in heat. He, there- fore, uses only selected males and when a female in heat is to be bred, he places one of his responsive males with her one day, an- other of his responsive males with her the second day and thus alternates a couple of males for three or four days. Even so, slightly fewer than half of the females which were thus bred pro- duced young. A weasel in the white winter coat was used by Audubon and Bachman (1856:177, Quarto edit.) to drive rabbits out of their bur- rows in the same fashion that ferrets commonly are used. Although these naturalists refer to their animal as an ermine it probably was Mustela frenata noveboracensis, the long-tailed weasel. The ani- mal's teeth (probably canines) were blunted and a long cord tied on its neck. With the aid of this weasel 12 rabbits were caught in one morning and more than 50 in four weeks. Enemies Little is recorded concerning enemies of weasels and it may be that other vertebrates are not an important factor in removing the annual increase. Errington (1935:195-198), in Iowa, found four. 200 University of Kansas Publs., Mus. Nat. Hist. putrid weasels about dens of red foxes, Vulpes fulvus. No remains of weasels were found in the feces of the foxes and it appears that the foxes do not eat the weasels. The label on an adult female specimen of M. /. spadix from Boone County, Iowa, bears the date May 10, 1938, and the annotation, by T. G. Scott, "fox-killed." Bailey (1931:328) recounts that "Weller saw a coyote carrying one in its mouth" at an elevation of 11,500 feet in the Pecos Mountains of New Mexico. The type specimen, a young female, of M. /. peninsulae from Hudsons, Florida, according to Rhoads (1894:155) "... was caught in the woods by a cat." Barber and Cock- erell (1898:189) mention one that was killed by a dog in Mesilla Park, New Mexico. Moore (1945:258) records the death of a weasel in Florida. Circumstantial evidence indicated that it was killed by the bite of a water moccasin. In the Biological Surveys Collection of mammals in the United States National Museum, the label with the skull of an adult male weasel, No. 160663, from Banning, California, carries the information that the skull was taken from the stomach of a Crotalus (rattlesnake). In reporting on a study of owl predation in Delaware County, Pennsylvania, Pearson and Pearson (1947:143) mention that "weasels are found throughout the county but . . . were never eaten by the owls." The Uinta spermophile at some places and times probably is a prey sought by the long-tailed weasel but Warren (1924:265) records Citelhis armatus repeatedly chasing weasels in August, at Camp Roosevelt, Yellowstone National Park, and how the ground squirrels at one time ignored the weasel even when it came within a few inches of a squirrel. Warren (1932:71), on August 2, 1931, at Grand Mesa, Colorado, obtained a large male weasel with two porcupine quills in it; one was near the mouth and another "in the skull." Osgood (1935:156) writes that near Rutland, Vermont, a male weasel "taken in April, was heavily parasitized and had several short porcupine quills em- bedded in its neck, head, and shoulders." The remainder of Os- good's account implies that the weasel may have turned to porcu- pine because the normal food for weasels was scarce at the time. Porcupine quills, then, are a hazard for weasels although it is un- likely that the porcupine is ever to be classed as an enemy of the weasel. An accident of another sort, which must at the very least have been annoying to the weasel that suflFered it, was recorded by Soper (1921:37). The animal had a stick lodged crosswise be- tween the fourth upper premolar teeth. MUSTELA FRENATA 201 The recorded actions of several kinds of animals which are too small to be dangerous to the weasel suggest tliat they recognize that the weasel is a danger to them. Borell and Ellis (1934:21) mention that a weasel in Nevada caused a great disturbance among the chipmunks. Long (1938:250) heard pikas give evidence of terror hy a peculiar cry when a weasel was in a rock slide occupied by the pikas. Seton (1929 (2):629) writes "On June 14, 1915, as I prowled around the south side of the lake on my homeland at Greenwich, Conn., my attention was called to a pair of song spar- rows and a male towhee that were noisily mobbing a Weasel, twit- tering around and darting at him, as though they knew full well his evil ways. The weasel paid little heed, but soon dived from sight in a stone wall." No account has been found of an American weasel or ermine rolling, tumbling and frolicking in a manner that aroused the curi- osity of birds to a degree which permitted the weasel to come witliin leaping distance of the birds. Accounts of such behavior are on record for the English stoat (ermine). Food and Hunting Weasels are active both in the daytime and at night. Whether the time of activity varies with the season, with the locality, with the sex or with other conditions, I do not know. Adult, live, free- living, actively moving weasels that I recall having seen all were observed in the daytime: two were in Alameda County, California, two were in White Pine County, Nevada, one was in Scotts Bluff County, Nebraska, and one was in Laramie County, Wyoming. I recall ten adults, from the same three states, and one from Wash- ington State, that got into my traps; two of these certainly got in the traps in the night; one certainly got in the trap in the daytime; the other eight were found in traps which may have caught the weasels either in the night or in the daytime. Soper (1946:136) in speaking of M. f. longicauda north of the International Boundary in Canada remarks that it has the "habit to some extent of hunting at all times of day." Criddle and Criddle (1925:144) in writing of Mustela frenata longicauda in Manitoba record that "The shrill cry of a rabbit [Lepiis americanus] in the dark is nearly always due to the weasel's attack. Indeed, we have often watched the latter at work during the twilight hours. First would come the almost noiseless run of the small rabbit with its characteristic dodging and this would be followed by the appearance of the agile foe which, at times, would leap high over obstacles and at others 202 University of Kansas Publs., Mus. Nat. Hist. move swiftly beneath them. Then there would follow intermittent cries of the rabbit as the weasel secured a temporary hold of its quarry, for be it noted that this hunter apparently bites anywhere to begin with and it is probable that the blood made to flow acts as an aid to tracking as well as weakening the prey. Several similar close encounters might occur before the rabbit would be finally over- come, but weasels are very persistent when they once get into con- tact with their victims and it is therefore very seldom that the latter escape. In killing, they either penetrate the brain with their teeth, or dislodge the vertebrae behind the head." These and more than two score other observations which record the time when weasels were seen make it clear that some were active at night and that some were active in the daytime. As to the routes traveled while the weasels are hunting, Quick (1944:77) says of four individuals that he studied in Washtenaw County, Michigan: "The weasels appeared to prefer hunting cer- tain coverts with noticeable regularity, but rarely cruised the same area on two consecutive nights." The killing technique of fifteen captive Mustela frenata nove- boracensis was studied by Glover (1943A). For the weasels he released 19 mice, 3 brown rats, 6 cottontails and 4 ring-necked pheasants. Most of the mice were killed by a bite on the back of the head, with the body and legs of the weasel hugging the back of the victim. "The weasel shoved the prey in close to the stomach with the hind legs, and the kill was made in a reclining semi-curled-up position". On each of the rats (Rattus) an initial grip was secured at the base of the ear. When the rat rested, a new hold was taken by the weasel. Finally the weasel secured a hold at the base of the skull and near the ear, and a hght crushing sound followed. Four of the six cottontails were killed by bites on top of the head and ear; two cottontails succumbed from neck wounds. In three instances, neither of two weasels could be in- duced to make a determined attack on the cottontails or to kill them. At times the cottontails proved to be able opponents for weasels by striking out with their front feet and by kicking with their strong hind legs. In killing the pheasants the teeth of the upper jaw of the weasel pierced the top of the braincase and the teeth of the lower jaw entered the region of the auditory process. The forelegs hugged the neck of the pheasant, the body of the weasel was extended in a riding position on the back of the bird and no amount of kicking or rolling dislodged the weasel. MUSTELA FRENATA 203 Polderboer, Kuhn and Hendrickson (1941) describe a cotton- tail cached by a weasel as having the muscles of the neck severed from the region behind the right mastoid process and noted "that hemorrhage in the region of the right jugular vein had occurred." Concerning the methods of killing mammals smaller than cotton- tails, the accounts by Nichols and Nichols (1935:297-299) and that by Svihla (1931) corroborate Glover's (1943A) account, as do also the accounts of Miller (1931B:164) and Moore (1945:257). The latter says that his captive male, from Gainesville, Florida, customarily bit its rodent prey at the base of the skull and used the feet to manipulate the live prey. Miller ( loc. cit. ) emphasized that his male weasel (M. /. longicauda) grasped where it could, used its snakelike body to coil over the prey and shifted the grip of its teeth to the nape of the neck or back of the skull. The captives that I have had [one from Salt Lake Gity, Utah; three from Gontra Gosta Gounty, Galifornia; and the same individual reported upon by Miller (1931:150)] customarily employed the techniques of killing small rodents that were described by Glover and Miller {loci cit.). Allen (1938:225-229) experimented with the ability of four dif- ferent males of M. /. noveboracensis from Michigan to kill adult cottontails. The method used was to place the weasel in a cage of quarter-inch hardware cloth approximately three feet long, two feet wide, and two feet high. The bottom of the box was covered with several inches of straw. One cottontail was offered to each weasel. In two instances the weasel attacked and bit the cottontail, was struck by the hind feet of the cottontail, retired from the attack and died a few hours later as a result of the blows of the cottontail's hind feet. In the other two instances the weasel rendered the cottontail helpless by severing the neck muscles from the skull. Subsequently an incision made by the weasel, in each of the two instances gave access to blood on which the weasel fed until it was full, in one instance by licking "blood as a cat laps milk". One rabbit was subdued in 10 minutes and the other in 15 minutes. Allen (op. cit.) points out that cottontails form a considerable portion of the weasel's food and thinks that they are killed in burrows more easily than they were in the cage. In writing of the three species of weasels, including Mustela frenata, found at Treesbank and vicinity, Manitoba, Norman Griddle and Stuart Griddle (1925:143, 144), in my opinion, correctly ex- plain the killing of more prey than weasels need. "The fact that 204 University of Kansas Publs., Mus. Nat. Hist. weasels frequently kill many more animals than they require for immediate use has been universally interpreted as a lust for killing — a supposition which we believe to be quite erroneous. It is true that weasels often kill more than they need, but the surplus is not necessarily wasted because the animals always store it for future use, in much the same way as do badgers, minks or skunks, and with the same object in view as squirrels have in gathering nuts. We have observed many such stores, but as far as our observations go, the habit of killing in excess occurs much more prominently in the late summer and autumn months than in the spring. Indeed, we have no records of excessive spring slaughter and this indicates that the supposedly blood-thirsty habit of weasels is no more a lust for killing than is the woodsman's foresight in providing his larder with meat for the winter months. It should be noted in this connection that members of the weasel family, when undisturbed, do not leave their victims scattered about, but carefully store them away, and in many instances the bodies are buried with earth or taken under ground to preserve them. We suspect that this instinct for preserving food for future use accounts for most of the excessive killing by carnivorous animals instead of this latter indicating an aimless desire for slaughter which would unnecessarily deplete the food supply of the future. This instinct, however, does not seem to be as definite as that of some rodents, and there is no doubt that much of the stored meat decays before it can be utilized." Griddle and Griddle (1925:146) note that a weasel in the vicinity of Treesbank was carrying a rat [Rattus] and that "Two small punctures in the throat were the only evidence of the manner in which its death had been brought about." Gonsiderable information has been recorded concerning the food of Mustela frenata and a little information is on record as to kinds of foods not taken that could have been taken. For example, Ingles (1939:253, 254) on May 14, 1938, near Shasta Gity, Gali- fornia, noted that nestlings of russet-backed thrushes were ignored by an adult weasel and four young weasels which were feeding instead on meadow mice and a mole. Howard (1935:322, 323) records that a weasel in Michigan which carried bits of meat from beef bones on a porch ignored a red squirrel which drew on the same food supply but which retreated to the end of the porch when the weasel appeared. Quick (1944) records that in the winter of 1940 on a 640 acre area in Washtenaw Gounty, Michigan, MUSTELA FRENATA 205 four resident weasels did not kill any of the 10 rabbits or several pheasants but subsisted on smaller animals. Glover (1943A) thought that M. frenata kills only a few adult cottontails in the wild. To judge from these observations, M. frenata chooses small mammals as prey in greater measure than it does birds or larger mammals. Records of prey taken, attacked or pursued by MusteJa frenata include the following: Broad-footed mole (Scapanus latimanus). — One was fed on by an adult M. frenata and four young, on May 14, 1939, "near Shasta City," California (Ingles, 1939:253, 254). Dusky shrew (Sorex cinereus. — A female weasel, at Majestic, Long Island, N. Y., was shot when carrying a Sorex cinereus that had a small hole in the top of its head (Nichols and Nichols, 1935: 297-299). Big short-tailed slirew (Blarina hrevicauda). — One was taken from the stomach of a weasel (Hamilton, 1928:249). Townsend ground squirrel (Citellus townsendii) . — Alcorn saw a weasel five miles west of Fallon, Nevada, carrying a squirrel (Hall, 1946:192). Richardson ground squirrel (Citellus richardsonii) . — The at- tempted capture of one of these squirrels in Saskatchewan is re- corded by Seton (1929 (2):625). Belding ground squirrel (Citellus beldingi). — Grinnell, Dixon and Linsdale (1937:233) recount that at Tuolumne Meadows, California, a weasel killed a ground squirrel of this species. Thirteen-lined ground squirrel (Citellus tridecemlineotus) . — Er- rington (1936:406, 407) found a den in Palo Alto County, Iowa, on June 22, 1934, where he collected 32 fecal pellets. Sixteen samples contained thirteen-lined ground squirrels, 9 contained rabbits, 9 contained mice (7 Microtus, 1 Feromyscus and 1 uni- dentified); red-winged blackbirds and unidentified fringillids were represented as also were ground beetles, grasshoppers and other insects. One red-winged blackbird lay near the entrance of the den. Franklin ground squirrel (Citellus franklinii). — Sowls (1948:126) records that at Delta, Manitoba, a weasel was observed killing one of these squirrels and that "the weasel had taken the squirrel from its hibernating burrow as evidenced by tracks in the snow." On July 19, 1917, in the vicinity of Treesbank, Manitoba, T. Criddle saw a weasel attacking one of these ground squirrels which was 206 Univeesity of Kansas Fuels., Mus. Nat. Hist. in mortal terror and squeaking continuously. Eventually the squir- rel was tlirown on its back "and would have been speedily killed but for an interruption" (Griddle and Griddle, 1925:146). Golden-mantled ground squirrel {Citellus lateralis). — On August 15, 1941, along the Kaweah River in Sequoia National Park, Boyer (1943:99, 100) saw a weasel chasing a Citellus lateralis; three or four times the weasel grasped the back of the neck of the squirrel which each time threw off the weasel until the two, weasel after the squirrel, plunged into the river. The squirrel, bleeding at the base of the skull, was rescued and entered a hole; the weasel got out of the water and under a rotting log. Follett (1937:365) at 2 p. m. in Plumas Gounty, Galifornia, saw a weasel have hold of the lower jaw of a golden-mantled ground squirrel near its throat. Alcorn watched a weasel chase a golden-mantled ground squirrel in Nevada (Hall, 1946:192) and Grinnell and Dixon (1919:681) record that on August 4, 1911, near Monache Meadows in eastern Tulare Gounty, Galifornia, a weasel pursued, captured and killed a golden-mantled ground squirrel. Eastern chipmunk (Tamias striatus). — Pearce (1937:483) in central New York State, on July 29, 1931, saw a chipmunk scamper up a tree pursued by a weasel. Ghipmunk (subgenus Neotamias). — Stanford (1931:363) on No- vember 11, 1931, at Fish Lake, Utah, saw a weasel pursuing a chipmunk. On August 5, 1910, "near Independence Lake," Nevada Gounty, Galifornia, Louise Kellogg recorded that a weasel seized and ran off with a chipmunk (Grinnell, Dixon and Linsdale, 1937: 233). Allen (1938:228) observed that a chipmunk (whether Tamias striatus or T. minimus not specified) was killed in 30 sec- onds whereas 10 to 15 minutes were required by the caged, male Mustela frenata noveboracensis to kill a cottontail. Red squirrel (Tamiasciurus). — Seton (1929 (2):625) records the capture of one in Pennsylvania, and Grinnell, Dixon and Linsdale (1937:232), at Gisco, Galifornia, saw one closely pursued by a weasel. Flying squirrel (Glaucomys). — Burroughs (1900:77, 78) records remains of one of these squirrels along with the remains of other animals in a food cache of a Mustela but his account does not make clear whether Mustela frenata or Mustela erminea was the species of weasel involved. Northern pocket gopher (Thomomys). — In "July. 1939, near Stillwater [Nevada], Alcorn pursued . . . [a] weasel and caused MUSTELA FRENATA 207 it to drop ... a pocket gopher [Thomomtjs bottae] which was about two-thirds grown" (Hall, 1946:192). Grinnell, Dixon and Linsdale (1937:233) write that "at least twice, weasels in the [Yosemite] Valley were seen carrying pocket gophers." Relative to Thomomtjs talpoides in the vicinity of Treesbank, Manitoba, Griddle and Griddle (1925:146) record that on September 11, 1918, an individual of Mustela frenata longicauda took seven pocket gophers dead .... It seized the rodents by the middle of their back and held them high while carrying them away. They were stored in a gopher burrow some two hundred yards distant. On February 17, 1921, "Game across the marks of a weasel carting some object over the snow. An investigation revealed a recently- killed pocket gopher with its captor still in possession." Griddle (1930:279), at Aweme, Manitoba, "frequently observed this weasel [M. f. longicauda] . . . carrying a pocket gopher to its larder, and twice it has been encountered in mid winter with freshly killed gophers in its possession". The evidence already presented that weasels levy heavily on pocket gophers is strengthened by the many references in the literature to weasels having been caught in traps set for pocket gophers in the burrows of those rodents and by the many statements, not quoted here, that living quarters of weasels are in burrows made originally by pocket gophers. For example, the present writer, in an account of the Mammals of Nevada (Hall, 1946:191, 192), has said of the long-tailed weasel, Mustela frenata nevadensis, that "All the three dens that were excavated . . . were originally burrows of pocket gophers . . . . Although we have found weasels in many situations in Nevada, . . . they most often were obtained from the burrows of pocket gophers." Excluding the weasels taken by Alcorn, more specimens of the remaining lot were caught in traps set in the bur- rows of pocket gophers than by all other means combined. All of the 22 weasels taken by Alcorn [within a radius of 10 miles of Fallon] were obtained in gopher traps. Mexican pocket gopher (Cratogeomys). — At Ghalchicomula, 8000 feet, Puebla, Nelson (1918:470 and letter dated March 9, 1928) saw a weasel fastened to a pocket gopher. Nelson obtained the pocket gopher and found that its neck muscles were torn loose from the skull. Grasshopper mouse (Onychomys) . — Barber and Gockerell (1898: 189) found remains of this mouse in the stomach of a weasel at Mesilla Park, New Mexico. 208 University of Kansas Publs., Mus. Nat. Hist, White-footed mice (Peromyscus). — Green (1936) saw a weasel in Gratiot County, Michigan, in May, carrying a Peromyscus. Quick (1944:76), in winter, in Michigan, found one dead, probably killed by a weasel. From Washtenaw County, Michigan, Quick ( 1944:77) examined 294 scats of free-living weasels and found Peromyscus in 189 scats, Microtus in 83, small birds in 20, red squirrel in 3, and hair of weasels in small quantities (probably from the animals which deposited the scats) in 36. He concludes (op. cit., 78) that the winter food was 65 to 70 per cent Peromyscus, 23 to 33 per cent Microtus, and 2 to 7 per cent small birds. Wood rats (Neotoma). — A female long-tailed weasel weighing 250 grams was taken one mile north of Kent, Texas, while eating a Neotoma albigula (Davis and Robertson, 1944:263). A wood rat house under observation by Vestal (1937:364) in Contra Costa County, California, was invaded by one weasel which ate two adult wood rats (Neotoma fuscipes) and one young. In the same area he saw a weasel in a wood rat nest some months later ( Vestal, 1938:5). Three miles east of Reno, Nevada, on May 13, 1936, W. B. Richardson watched a long-tailed weasel carrying a half-grown round-tailed wood rat (Neotoma lepida) across a rock slide (Hall, 1946, 192). Harper (1927:303) records three wood rats [Neotoina floridana] and two cotton rats [Sigmodon hispidus] found dead in the den of a female weasel and her three young in the Okefinokee Swamp of Georgia. Another female and three young approximately half grown were found in the swamp in a hollow pine log. Con- tents of the den as described to Harper were nearly a peck of wood rats, whole and in pieces; remains of several kinds of birds including robins and quail, and a piece of joint snake (Ophisaurus ventralis). Meadow mice (Microtus). — Polderboer, Kuhn and Hendrickson (1941), in 1939, at Ames, Iowa, identified "A total of 118 items . . . in 97 winter scats and 48 in the 38 spring scats." Their combined data are as follows: Frequency Percentage Meadow mouse 71 42 85 Harvest mouse 36 21 75 Deer mouse 17 10 23 Meams cottontail 14 8 42 Short-tailed shrew 9 5 42 House mouse 3 1 86 Tree sparrow 2 1 02 Grasshopper 1 .60 Shaw pocket gopher 1 .60 Least weasel 9 5 40 Unidentified material 3 1 . 85 MUSTELA FRENATA 209 Polderboer, Kuhn and Hendrickson divide their data into two categories, winter and spring. Items recorded in winter but not in spring are house mouse, tree sparrow, and grasshopper. J terns recorded only in spring were pocket gopher and least weasel. The samples of cottontail and least weasel all were from the scats of one large male weasel. Of a total of 14 pheasants, 24 quail and 35 cottontails on the 160 acres involved in the study only two cotton- tails appear to have been killed by the weasels — really by one weasel of four which lived on the area. Food items taken from the nests (3) and adjacent caches of food in the dens, were as follows: meadow mouse, 30; short-tailed shrew, 4; pocket gopher, 2; deer mouse, 2; least weasel, 1; tree sparrow, 1. The authors remark tliat the abundance of several prey species does not cause the weasels to ignore the slirews which are said to be distasteful to carnivores. Two horned larks, apparently killed by weasels, were found on the 160 acre area studied; the horned larks were not in caches of food, nor were remains of horned larks found in scats. Dearborn (1932:34, 37) for Michigan, on the basis of contents of (37?) intestinal tracts and "feces collected partly in winter and partly in summer" found that, by frequency of occurrence, mammals comprised 83 per cent of the food, birds 10 per cent and insects 7 per cent. Frequency indices for the genera of mammals in per- centages of food items of all kinds were as follows: Microtus, 31 per cent; Peromyscus, 24 per cent; Sylvilagus, 14 per cent; Sorex, 7 per cent; Blarina, 5 per cent; Scalopus, 2 per cent. Griddle and Griddle (1925:146), for the vicinity of Treesbank, Manitoba, record that on October 3, 1913, a weasel was seen to take a field mouse down a hole. They add {op. cit.: 147) that "Once while ploughing, we observed a Long-tailed Weasel carrying a field mouse. . . ." Ingles (1939:253, 254), in June, 1938, near Mt. Shasta City, Galifornia, found an adult and four young weasels which fed on several Microtus montanus montanus. Green ( 1936 ) in May, in Gratiot Gounty, Michigan, in the vicinity of a nest in v/hich there were four young weasels, found "several" dead Micro- tus. Hamilton (1933:330) records that in New York State a male weasel, on April 5, 1932, at Ithaca, had eaten a Microtus and that in May, 1927, a female weasel was seen carrying a Microtus in its mouth. Hamilton's (1933:333) study of the contents of the digestive tracts of bodies of weasels obtained from fur trappers and fur buyers enabled him to publish the following "Frequency Indices 210 University of Kansas Publs., Mus. Nat. Hist. of Mammal Genera in Fall and Winter Food of 163 Mustela nove- boracensis": Microtus, 33.6 per cent; Sylvilagus, 17.3; mammals undetermined to genus but principally mice, 17.1; Peromijsctts, 11.3; Rattus, 9.1; Blarina, 5.9; Sciurus, 2.7; Tamias, 1.0; Condylura, 0.8; Ondatra, 0.8. Grinnell, Dixon and Linsdale (1937:233, 234) quote W. Fry con- cerning a weasel which reared six young at Giant Forest, California, in 1919, as follows: "This parent weasel, after the birth of her young, remained at the premises for a period of thirty-seven days; during which time, from actual count, the following numbers of mammal species fell victim to her: mice [genera not specified] 78; gophers 27; moles 2; chipmunks 34; wood rats 3; ground squirrels 4. This is a total of 148 animals for the . . . thirty-seven days . . . not a bird was captured during the period." Rats (Rattus).— Criddle and Griddle (1925:146), on the farm at Treesbank, Manitoba, record a long-tailed weasel, on July 2, 1918, running away from the farm buildings carrying a rat; July 11, 1919, "Two longicaudas . . . have been seen running off with rats on several occasions."; July 11, 1920, "There are two large weasels about the buildings [;]. . . . Each has been noted with rats and this afternoon one of them was seen running into the woods carrying a rat, followed by two excited swallows." The authors {op. cit. :147) add "In the fall of 1924, Mr. A. Gooper, a prominent poultryman of Treesbank, observed a large weasel carrying a freshly killed rat which it stored below ground and then returned towards the poultry-house, causing no little apprehension to the owner. Within a short time, however, the weasel reappeared with another rat which it hid as before. In this way several rodents were ac- counted for during the afternoon, and Mr. Gooper assures us that the weasel Tcept up the good work for some days'." Hamilton (1933:330) in New York State in May, 1927, saw a male weasel in possession of a rat. Big jumping mouse (Zapus major). — In the Warner Mountains of Galifornia, on Parker Greek, H. G. Bryant frightened a weasel that dropped a freshly killed jumping mouse (Grinnell, Dixon and Linsdale, 1937:232). Snowshoe rabbit (Lepiis americanus) . — Adolph Murie (1935: 321-322) writes that: "Four miles north of Funkley, Minnesota, early on the morning of November 13, 1921, . . . watched from the top of a 30-foot spruce a weasel . . . hunting a varying hare. . . . The ground was covered with six inches of fresh snow . . . both animals . . . [had] their [white] winter pelage. MUSTELA FRENATA 211 "My attention was first attracted to the hare as it came hopping steadily but unhurriedly from the north. Directly in front of me, about 75 feet from the tree I had climbed, the hare crisscrossed back and forth at various angles over an open area about 20 feet in diameter. After producing a maze of tracks, the hare 'froze' near one edge of tlie pattern. In a few minutes the weasel appeared, all his faculties focused on the warm trail. Expertly he followed its convolutions, passing at times within a few feet of the watching hare. Not until the weasel had followed every turn of the trail to within three feet of its termination did the hare skip off. It came out to the road almost directly below me, turned at right angles northward and was soon out of sight. At the road the weasel lost the trail, . . . and then ran parallel with it, once more in hot pursuit. "Ten minutes later the hare emerged from the north as before, came on directly to the tracked-up area, and continuing its strata- gem, leisurely hopped about to leave its zigzag trail. Then it sat down quietly to wait. . . . The weasel ['s] . . . nose led him through tlie network with Uttle trouble. He was almost upon the hare before it jumped off and followed the same path [as] . . . before. . . "The hare had to show his big heels [a third time] ... as the weasel approached him. This time the weasel failed to follow. . . . After examining a few brush heaps he vanished into the woods behind me." Seton (1929 (4):723, 724) writes tliat in December of 1886 in the sandhills nortlieast of Carberry, Manitoba, he saw a weasel chasing a snowshoe rabbit which took refuge near his feet under the sleigh and so escaped the weasel. Thurber (1940:356) men- tions a month-old varying hare that was rescued from a weasel and of approximately the same size as the weasel. Griddle and Griddle (1925:146) for the vicinity of Treesbank, Manitoba, record "August 21, 1921. — Heard cries of a small rabbit at dusk to-night, which investigation showed was being attacked by a large weasel. The rabbit was later carried to the weasel's store chamber below ground." They record furtlier (op. cit., 146, 147): "November 8, 1924.— Shot a bush rabbit and left it lying. Two hours later [it] . . . was found to have been dragged be- neath a brush pile and partly eaten. Innumerable weasel tracks left no doubt as to the identity of the thief." In describing a weasel that wintered in a nest in a threshing machine, the same autliors {op. cit. :143) say that no bird remains were found in the pile of 212 University of Kansas Publs., Mus. Nat. Hist. approximately three pounds of droppings adjacent to the nest. In a store chamber some 140 yards away from the nest, two bush rabbits ( Lepus americanus ) had been dragged to the entrance and numerous smaller rodents were taken below ground. The rabbits were buried beneath the snow and eaten as necessity arose. Narrow selectivity on the part of the weasel in choosing food is almost always shown in instances where the food of weasels has been studied. For example, the weasel which lived in the threshing machine ate rodents and rabbits and not poultry although the weasel had ready access to the poultry building. The weasel which lived in the bag of feathers in the basement of Stuart Griddle's house ignored grouse, approximately 20 in number, in favor of other non-avian food. Cottontail (Sylvilagus) . — Polderboer, Kuhn and Hendrickson ( 1941 ) mention that one of 4 weasels which they studied on a 160 acre area at Ames, Iowa, in 1939, had a cache of food in a pocket gopher burrow 10 rods distant from tlie weasel's den. The cache contained only two cottontails, one partly eaten. Leopold (1937) records seeing a Mustcla (probably a long-tailed weasel but pos- sibly an ermine) kill a third-grown cottontail by biting it at the base of the skull. Leopold describes the blood sucking or licking, suggesting that he shared the popular misconception that weasels suck blood. The supposition that weasels suck blood has been refuted by many observers, for example by Svihla ( 1931 ) . My own observation of captives makes me think that weasels do not suck blood. Seton (1929 (2):626) quotes B. H. Warren as seeing a weasel dragging a freshly killed, still warm, rabbit that contained nine embryos almost ready for birth. A young rabbit was seen being carried by a weasel in Hidalgo County, Texas, in March, 1935 (Mulaik, 1938:104). An instance of a cottontail being chased in June in South Carolina is recorded by Hamilton (1933:330), Addy (1939:372, 373), in Virginia, on August 14, 1939, shot a large weasel which was pursuing a Sylvilagus that was only a foot and a half ahead of the weasel. The rabbit stopped when a shot was fired and permitted itself to be stroked and petted. Tracking showed that the weasel had chased the rabbit for a half mile. On November 20, 1942, at Lake James, Indiana, a weasel was seen by Grosjean (1942:443) attacking a "young rabbit" in the throat of which the weasel had made five large holes from which there was no obvious bleeding. Seton (1929 (4): 798) recounts that in 1910 at Base Lake, Michigan, F, C. Hicks saw a cottontail with a weasel hanging to its legs rush to the cottage. When only four feet from MUSTELA FRENATA 213 Hicks the weasel loosed its hold and the cottontail escaped under the cottage. Burroughs (1939:253) on May 14, 1939, in Saginaw County, Michigan, records that a young cottontail weighing be- tween 200 and 250 grams was carried from the nest and killed. Burroughs was attracted first by the "hissing scream" of the weasel, strode toward the sound, flushed an adult cottontail, and discovered the empty nest from which the weasel had taken the young cot- tontail. Brush rabbit {Sylvilagus bachmani). — Vestal (1937:364) in Con- tra Costa County, California, found a brush rabbit that appeared to have been killed by a weasel. Reptiles. — Grinnell, Dixon and Linsdale (1937:234) recount that in July, 1889, in Wilson Canyon, near Pasadena, California, a weasel killed a red racer by severing the backbone of the snake. In April, 1935, in Hidalgo County, Texas, a half grown bull snake ( Pituophis sayi sayi) was regurgitated by a young weasel. Russell (1930:504, 505) has recorded finding in California a male weasel and a king snake (Lampropeltis getulus boylii) three feet five inches long in mortal combat. The weasel killed the snake but the weasel, inca- pacitated by the conflict, was easily picked up by hand and was also saved as a specimen. Wild birds. — In the spring of 1940, in Washtenaw County, Michi- gan, one bobwhite, of 10 bobwhite living on a 640 acre area, was killed by one of four weasels that lived on the area. No other quail was killed there. The one unfortunate bird was killed in the mouth of an abandoned den where the quail roosted (Quick, 1944:76). A male weasel, subspecies M. /. effera, was seen by Booth (1946:439) attempting to enter the nesting hole of a pair of Slickers, Colaptes. One song sparrow (Melospiza melodia), and one slate-colored junco (Junco hyemalis) were recorded by Quick (1944:76) as killed by weasel in Michigan. Chicken (genus Gallus). — Quick (1944:78) writes that in one year (1938-1939) weasels were reported to have killed 1.03 per cent of all chickens in one township of Washtenaw County, Michigan, and that of the total damage to all kinds of poultry, 59 per cent was done by weasels. Weasels entered 19 per cent of the chicken coops on the study area. Farmers killed 68 per cent of the weasels seen in barn yards. Spring and summer were the seasons in which most of the weasels were observed in barn yards. Internal evidence in Quick's {op. cit.) account leads me to suspect that some losses of poultry were charged to weasels when Rattus was actually re- sponsible. 214 University of Kansas Publs., Mus. Nat. Hist. Griddle and Griddle (1925:146), quote a neighbor in the vicinity of Treesbank, Manitoba, as recording that on October 29, 1917, "A weasel last night made its way into our fowl-house, the door being inadvertently left open. The weasel killed eleven fowl, some of which were dragged into the yard. All the largest fowls were selected, tlie pullets remaining untouched though they were in the majority. Next night the weasel dug a hole beneath the building and killed a hen and two cocks, returning for another during the day, making a total of fourteen in all." Griddle and Griddle (1925: 146) remark that the weasel proved to be a large one, probably an old male. The same authors (op. cit. -.147) record that at their farm at Treesbank, Manitoba, on January 31, 1925, "A Long-tailed Weasel killed three hens last night, and rather severely bit a cock about the neck. This, or another weasel, had been around the farm-yard for some time ( The specimen was a large male ) . . . . "In the fall of 1924, Mr. A. Gooper, a prominent poultryman of Treesbank, observed a large weasel carrying a freshly killed rat which it stored below ground and then returned towards the poul- try-house, causing no little apprehension to the owner. Within a short time, however, the weasel reappeared with another rat which it hid as before. In this way several rodents were accounted for during the afternoon, and Mr. Gooper assures us that the weasel 'kept up the good work for same days'. "Being a farmer of many years' standing, Mr. Gooper has naturally lost some poultry through the agency of weasels, but while he re- marks that 'there are good as well as bad actors among weasels', he has the practical good sense to recognize the value of an efficient ratter even though it be a weasel. "Our sister, Maida Griddle, writes under date of March 4, 1925: " 'There is another weasel ( longicauda ) in the fowl-house, a well-behaved one this time. It came and took a piece of meat out of my hand quite nicely, which it carried down a hole and then came and sniffed all over my mitt to see if there was any more. I thought it had been killed when I visited the farm buildings next day as there was a strong smell of musk on the cat and in the fowl house, but the weasel was there as cheeky as ever. It got hold of my skirt twice and tried to pull me down its hole. I think it wanted the cloth for a bed, as it was taking straw and other material down the burrow. The poultry were very frightened at first, but they are getting used to the weasel's presence now'." In commenting on the economic role of the long-tailed weasel in Manitoba, Griddle and Griddle (1925:145) write as follows: MUSTELA FRENATA 215 "Supply and demand are prominent factors in governing our weasels' food habits. The two smaller species, as we have already pointed out are so dependent upon mice for a living that they increase or diminish with the fluctuation of these creatures. The Long-tailed Weasel, however, is not so easily checked by the temporary disappearance of any particular kind of game. If mice are scarce it devotes greater attention to gophers or bush rabbits and if these are not in sufBcienct numbers to satisfy its appetite, the animal raids a poultry house as a last resource. In nine years out of ten, this weasel will find sufficient food about the fields and woods, but on the tenth it may be obliged to temporarily turn to domestic animals. It is at such times that the weasel is seen and its deeds recorded. A thousand mice may have been killed in the meantime, but the destruction of half a dozen hens is alone used as evidence of the weasel's economic standing. "In the last twenty years we have permitted weasels to frequent the farm buildings at will and the poultry house has been no exception. In that time rats and mice suffered severely from the weasels, while the total number of poultry taken were six. Many times that number, however, have been killed by rats. "When we review our experiences of the past, we are astonished to find what few poultry have been killed by weasels. Our own losses in forty-two years have not exceeded fifteen birds and even these were usually eatable. There have been reports of losses from time to time from neighbors, but on looking into details we find that there are very few farmers who have experienced more than three separate occasions of weasel depredation and the total loss per farmer in the last thirty years does not, we are sure, ex- ceed ten birds. This is surely a remarkably small payment to weasels in general for the great good done by them in killing rodents. "We wish to point out, too, that only the exceptional weasel becomes a poultry killer. In most cases apparently it is a fully- grown male that does the killing. There are exceptions, of course, but when we see a large weasel actively engaged in rodent hunting within a few feet of a brood of newly hatched chickens and not even looking at them, we must at least pause to ask if this animal is the enemy that we were taught to believe it to be." A suggestion that weasels sometimes obtain the prey killed by hawks is offered by Griddle and Griddle (1925:147) who write: "Hawks are not always the aggressors, as is shown by an incident 216 University of Kansas Publs., Mus. Nat. Hist. reported by Mr. H. L. Seamans, of Lethbridge, Alberta. Mr. Seamans noted a large buzzard suddenly fly straight upwards from a fence post, and then alight upon another one some distance away. A little while afterward this bird once more arose in the same manner as before, and presently repeated the performance again. An investigation then followed and revealed that a Long- tailed Weasel was following the hawk from post to post. "We should hardly exjDect a weasel to attempt to capture a bird of the above type. On the other hand, it is possible that these animals might be able to startle a hawk sufficiently to cause it to drop its prey, which would thus provide food for the weasel." The following frequency index is compiled from the foregoing data on prey of Mtistela frenata. Moles (family Talpidae), 5 Shrews (family Soricidae), 26 Pigmy weasel {Mtistela rixosa), 1 Ground squirrels (genus Citellus), 23 Chipmunks (genus Tatnias), 38 Tree squirrel (possibly all Tamiasciurus), 8 Flying squirrel (genus Glaucomys), 1 Pocket gophers (family Geomyidae), 34 Mice (order Rodentia), 96 Harvest mice (genus Reithrodontomys), 36 Grasshopper mouse (genus Onychomys), 1 Deer mice (genus Peromyscus), 235 Cotton rat (genus Sigmodon), 2 Wood rats (genus Neotoma), 14 Meadow mice (genus Microtus), 248 Muskrat (genus Ondatra), 1 Old World rats (genus Rattus), 19 House mouse (genus Mus), 1 Jumping mouse (genus Zapus), 5 Varying hare (Lepus americanus), 5 Rabbits (genus Sylvilagus), 48 small birds, 32 cliickens, 17 lizard, 1 snakes, 4 insects, 3 More significant than the above compilation, of course, are the results of careful studies of the food of the long-tailed weasel in re- stricted areas. Examples of such studies are those of Polderboer, Kuhn and Hendrickson (1941) and Hamilton (1933:333). According to Hamilton's (1933:332) observations on captive weasels, "There seems to be little relative difference in the amount they eat, regardless of their activities. "In general, more food is taken in summer than in winter. Usually about a third their weight every 24 hours is eaten, but a growing young weasel will consume much more. A young male noveboracensis, weighing 145 grams, consumed an entire chipmunk, fur and bones, weighing 85 grams, in 24 hours. A day later it ate all of a partly grown rat, 105 grams, in the same length of time." Moore (1945:253) records that a captive male that he obtained at Gainesville, Florida, consumed, on the average, between 63 and 70 grams of flesh and blood per day. The weasel itself weighed approximately 320 grams. Sanderson (1949:413), concerning seven young weasels from Manitoba, that he raised in captivity, writes: "From the fifth to MUSTELA FRENATA 217 the seventh week of age, they consumed approximately 22 pjr cent of their body weight per day; from the eighth to the tenth week (just before reaching mature size) they consumed approxi- mately 24 per cent; but after reaching maturity they consumed only 18 per cent. When given all the food they would take in one day, they ate as much as 40 per cent of their body weight." Griddle and Griddle (1925:143, 146) say that weasels drinking at a bird trough "held their mouths very close to the water and as far as we could see, lapped the liquid up with rapid movements of the tongue. As a rule, after drinking, they would merely spring to the ground and vanish amid a bunch of scolding birds, but occasion- ally we have seen an animal slowly drag itself through the water and follow this performance by some rapid gambols, or a quick run, a method of drying which most of us have practiced in our youth". According to Hamilton's (1933:332) observations on captives, "Weasels are great drinkers, and while they take but little at a time, about 25 c. c. is drunk by a large animal during a day . . . ." Reproduction Philip L. Wright's several papers (1942A, 1942B, 1947, 194SA, and 1948B) reporting on his detailed studies of Mustela frenata (subspecies orihasus and longicauda) in captivity have yielded a large share of the precise information that we have concerning breeding and reproduction in this species. He has found that a single litter, of up to 9 young is born in the spring, usually in April. At three months of age the females "are full grown". The young males remain sexually immature during the first summer but the young females, as well as the females which are more than a year old, come into heat in the midsummer and are bred by the adult males. After a long period of quiescence lasting for several montlis, the embryos resulting from these matings become active in early spring and develop to full term in less than 27 days after they be- come implanted. The adult males are sexually active from April into August, when the testes are at maximal size and are conspicuous in the scrotum. A gradual regression takes place starting in August and extending into September. By October the testes may be fully regressed and the molt to white may start in this month. The white winter weasel, of either sex, is sexually inactive. The testes of the sexually active male in early spring and late summer are seven to eight times the size of the fully regressed testes. Females which had borne and suckled young were first found to be in oestrus 65 to 104 days after birth of the young. Lactation lasts for approxi- 218 University of Kansas Publs., Mus. Nat. Hist. mately 5 weeks. In 18 litters the length of the gestation period varied from 220 to 337 days with an average of 279 days. The female in heat has the vulva much swollen and she will remain in this condition for several weeks if not bred. Wright (1948A) de- scribes the actual mating as beginning with a scuffle after which the male grabs the female by the scruff of the neck with his teeth and holds her until she becomes subdued when he clasps her lower abdomen with his front feet and arches his back over her posterior regions. The two animals remain locked in this position usually for two hours and sometimes for longer than three hours. If the animals are left together, copulation may take place again on the same day or upon succeeding days. Hamilton (1933:316-321) writes of a freshly born M. /. novebora- censis that it ". . . was pink and much wrinkled. The wet- ness .... did not entirely obscure a few sparse, rather long, white hairs . . . over its back and head. It had the pro- nounced and extraordinarily long neck of the adult." At one day of age the average weight of six individuals in the litter was 3.1 grams, which is 3 per cent of the weight of the adult female and 1/2 per cent of the weight of an adult male. At two weeks of age "The silky white hair . . . obscures the general flesh color of the skin, evident a week earlier. The hair on the back of the head and neck, also over the shoulders, is slightly longer than that of the back . . ." but there is no crest or mane or pompadour at this or any other age such as characterizes the juvenal ermine. When 21 days old one young male 'liurried from the nest chamber and commenced to eat some meat." At three and a half weeks "They are all eating small pieces of meat. . . . The canine teeth have made their appearance in both the upper and lower jaw, but just a hint of the incisors show. Some of the cheek teeth are through, as the meat appears to be thoroughly masticated by the little ones." On the 36th and 37th days the eyes opened. Sanderson (1949:415) found that a litter of seven young of Mus- tela frenata longicauda, from Manitoba, raised in captivity, "reached the peak of their growth" at approximately ten weeks of age. Several nests have been found. In Manitoba, Sanderson (1949: 412) excavated a burrow at the mouth of which he had trapped the adult female and in which he found eight young approximately five weeks old. The "burrow was about three inches in diameter, with two chambers at a depth of twelve inches. One of these was empty, the other contained the young. The two surface-openings were but two feet apart and the entire burrow was no more than MUSTELA FRENATA 219 three feet long. . . . The meager nest material consisted en- tirely of finely chopped grass. There was no mouse hair present, no accumulation of fecal material, and no storehouse containing food." Charles O. Handley has written me that on January 25, 1929, on the Sinkola Plantation, Thomas County, Georgia, he investigated the living quarters of a family of five weasels, four of which had been shot five days before by a hunter. According to the hunter each of the four which had been killed was approximately two- thirds the size of one which escaped into a hole in the ground. Handley found that the weasels had been using as headquarters a burrow in the trunk of an old uprooted oak as well as a nearby gopher burrow. The burrow in the oak was approximately ten feet long and had been excavated in the rotten wood. In a dis- tance of fifty feet along the gopher tunnel there were several used openings with pathways leading away from each. On February 6, Handley, with the help of a friend, trapped a large male weasel near this place. Criddle and Criddle (1925:143) describe a female which, one winter, slept in a bag of feathers in a basement of a house occupied by one of the authors; another weasel in winter made its head- quarters in a threshing machine. The nest of the latter "was somewhat roughly constructed and consisted of a convenient bunch of straw and chaff under the cylinder." Harper (1927:303) in the Okefinokee Swamp of Georgia dis- lodged a weasel from the house of a wood rat and was told of a den found in the swamp "in the trunk of a hollow cypress tree" from which a mother weasel and three young "about the size of mice" were obtained. "The bed contained, I suppose, a bushel or more of rabbit hair, rat hair, and squirrel hair. It looked like it must have been used as a den for several years, although there was no stink that I could detect except the musk from the old Weasel." Another female and tliree young approximately half grown were found in a hollow pine log. Between January 6 and April 12, 1940, on 640 acres of land, in Washtenaw County, Michigan, four weasels were studied and each weasel used only one den in this period (Quick, 1944:78). Criddle (1930:279) remarks that M. /. longicauda at Aweme, Mani- toba, often makes its temporary headquarters in the burrows of pocket gophers (Thomomys) . A female and three young weasels were found by Shaw ( 1921 : 167 ) using a nest of a mountain beaver in the burrow of that animal. Green (1936), in May, in Gratiot 220 University of Kansas Publs., Mus. Nat. Hist. County, Michigan, saw a weasel enter a hole under a decayed log and investigated finding four young weasels in a nest mostly of Microtus fur. In the early part (winter and spring) of 1939, at Ames, Iowa, Polderboer, Kuhn and Hendrickson (1941) studied four weasels Hving in four separate dens on 160 acres typical of Iowa farmland and excavated three of the dens. One den was in a weed patch in an old mole run. The nest chamber, approximately nine inches in diameter and six inches below the surface of the ground "was filled with grasses packed in a layer-like formation. In the center of this mass was a nest hollow lined with patches of mouse and shrew fur. Beneath this layer of fur and at the sides of the nest were skins, various bones, and skulls of partially eaten mice and shrews . . . scats [were in the nest] ... At intervals, layers of clean grass had been laid over the filth of the former bed, thus giving the nest a stratified appearance." A second den, of a lairge male, was in a field of sweet clover two feet high in the former burrow of a FrankHn's ground squirrel. The nest cell, seven inches in diameter and nine inches below the surface of the ground, "was lined with grasses mixed witli much rabbit and mouse fur. Some scats, and bones and fur of mice and shrews were matted together in layers at the bottom of tlie nest." When this den was abandoned the male weasel occupied, for a month, another bur- row, 20 rods distant, of a Franklin ground squirrel, in the field of sweet clover. The nest cell measured 11 by nine inches and was 11 inches below the surface of the ground. "Two nest layers were present. The first, composed chiefly of coarse straw and grass, had apparently been occupied at some time by a spotted skunk. . . . On top of the skunk nest was the weasel nest composed of fine grasses, mouse fur, and skeletal remains of mice." Relation of the Sexes to each other and to the young Quick (1944:75) writes that on March 28, in Michigan, he found the tracks of a male and those of a smaller animal, supposedly a female, meeting. The two "then led along the fence for about 18 chains and both entered the den of the male .... Only the tracks of the smaller weasel left the den on the same date. Observa- tion on April 12 showed that the large male still occupied the den." I am at a loss to explain this behavior since breeding would not be expected to occur in late March and since I suppose that MUSTELA FRENATA 221 the male and female do not live together except in the breeding season. Consequently, I wonder if the sign was wrongly read. Hamilton (1933:328), however, writes that M. /. novehoracensis Fig. 29. Map showing the geographic ranges of the subspecies of Mustela frenata and Mustela africana. 1. M. f. novehoracensis 12. M. f. nevadensis 23. M. /. inyoensis 34. ^f. f. costariccnsis 2. M. f. occisor 13. M. f. effera 24. M. f. neomexicana 35. M. f. panarnensis 3. M. f. primulina 14. M. f. washingtoni 25. M. f. texensis 36. M.f. meridana 4. M. f. arthuri ^5. M. f. saturata 26. M. f. frenata 87. M. f. affinis 5. M. f. olivacca 16. M. f. altifrontalis 27. M. f. Icucoparia 38. M. f. aureoventris 6. M. f. peninsulae 17. M. f. orcgoncnsis 28. M. f. perotae 39. M. f. helleri 7. M. f. spadix 18. M. f. munda 29. M. f. macrophonius 40. A/. /. macrura 8. M. f. longicauda 19. M. f. xanthogenys 30. M. f. goldmani 41. M. f. agilis 9. M. f. oribasus 20. M. f. nigriauris 31. M. f. tropicalis 42. M. f. boliviensis 0. ^f. f. allcni 21. M. f. latirostra 32. M. f. perda 43. M. a. africana 1. M. f. arizonensis 22. M. f. pulchra 33. M. f. nicaraguae 44. M. a. stolzmanni 222 University of Kansas Publs., Mus. Nat. Hist. is to "be found in pairs when caring for the young. During mid- May, 1927, I several times saw a male of this species carrying food to a den of young ones." Green (1936), in May in Gratiot County, Michigan, remarks that while he was uncovering and examining a nest of four young weasels, two adults ran about excitedly and one removed a young weasel. In instances where several nearly full-grown young have been obtained from one den it has been my experience (Hall, 1946:191) that the only adult trapped there was the female; no adult male was found or in the one instance when found he was living alone in a den 200 yards away from the den of the female and her young. Data are too few to warrant a definite conclusion about the extent to which the male aids in rearing the young, but I have wondered if he might not do so when the young were less than half grown and then live alone when they were more than half grown. Mustela frenata noveboracensis (Emmons) Long-tailed Weasel Plates 16, 17, 18, 31, 32 and 33 Putorius Noveboracensis Emmons, Quadrupeds of Massachusetts, p. 45, 1840. Mustela fusca DeKay, Zool. of New York, Pt. 1, Mammalia, p. 34, 1842. Putorius fiiscus Audubon and Bachman, joum. Acad. Nat. Sci. Philadelpliia, 8 (Pt. 2):288, 1842; Audubon and Bachman, Vivip. quadrupeds of N. Amer., 3:234, pi. 148, 1853 (pi. 1848). Putorius noveboracensis, DeKay, Zool. of New York, Pt. 1, Mammalia, p. 34, 1842; Baird, Manmi. N. Amer., p. 166, 1858; Merriam, N. Amer. Fauna, 11:16, pi. 4, figs. 1, la, 2, 2a, pi. 5, figs. 3, 3a, text figs. 4-6, 30, June 30, 1896; Bangs, Proc. Biol. Soc. Washington, 10:13, pi. 1, figs. 2, 2a, pi. 2, figs. 2, 2a, and pi. 3, figs. 3, 3a, February 25, 1896; Cory, Mamm. Illinois and Wisconsin, p. 366, plates, 1912. Putorius erminea, Audubon and Bachman, Vivip. quadrupeds of N. Amer., 2:56, pi. 59, 1851. Putorius agilis Audubon and Bachman, Vivip. quadrupeds of N. Amer., 3:184, pi. 140, 1853. Putorius richardsonii, Baird, Mamm. N. Amer., p. 164, 1858 (part). Putorius (Gale) erminea, Coues, Fur-bearing animals, p. 109, 1877 (part). Putorius noveboracensis notius Bangs, Proc. New England Zool. Club, 1:53, June 9, 1899. Type from Weaverville, Buncombe County, North Carolina. Mustela noveboracensis noveboracensis. Miller, U. S. Nat. Mus. Bull., 79:97, December 12, 1912; Soper, Joum. Mamm., 4:251, November 1, 1923. Mustela cicognanii, Henninger, Joum. Mamm., 2:239, November 29, 1921; Seton, Lives of game animals, 2:584, 1929 (part, Ohio); Hamilton, Amer. Midland Nat., 14:290, July, 1933 (part, Ohio); Lyon, Amer. Midland Nat, 17:109, January, 1936 (part, Ohio). Mustela noveboracensis, Jackson, Joum. Mamm., 3:15, February 8, 1922. Mustela frenata noveboracensis. Hall, Carnegie Instit. Washington Publ. 473: 104, November 20, 1936; Hall, Amer. Midland Nat., 18:304, March, 1937. Type — Wilhamstown, Berkshire County, Massachusetts. Type specunen not known to be in existence. MUSTELA FRENATA 223 Range. — Altitudinally, sea level to highest parts of mountains of eastern United States; Canadian Life-zone of Ontario and Quebec southward through eastern United States in Canadian, Transition and Upper Austral life-zones to and including upper edge of Lower Austral Life-zone in the Carolinas and northern parts of Georgia, Alabama, and Mississippi; westward from the At- lantic Coast to St. Croix and Mississippi rivers. See figure 29 on page 22L Characters for ready recognition. — DiflFers: From M. f. olivacea, in males, by width of tympanic bulla which is less than rather than more than 8.5 mm., and in adult females by total length which is less than rather than more than 345 mm. and by mastoid breadth which is less than rather than more than dis- tance between articular faces of exoccipital condyle and glenoid fossa; from M. /. occisor by a number of average differences including smaller size, rela- tively shorter tail and relatively narrower skull (see measurements); from M. f. spadix by least width of color of underparts amounting to less than 41 per cent of greatest width of color of upperparts, absence of color of under- parts on ankles and feet, adults with hind foot less than 50 in males and 40 in females, orbitonasal length less than 15.5 in males and 13.5 in females, length of tooth-rows less than 18.0 in males and 15.7 in females, mastoid breadth less than 25.5 in males and 22.0 in females; from M. /. primulina in males by interorbital breadth averaging more than 24 per cent of basilar length, orbitonasal length averaging more than 34 per cent of basilar length or 64 per cent of mastoid breadth, tympanic bullae less inflated anteromedially, than posteromedially, and in females by orbitonasal length amounting to more than two-thirds of mastoid breadth, by zygomatic breadth averaging less than 21, and by anterolateral margin of tympanic bullae not projecting below squa- mosal; from M. /. arthuri in males, by zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla and by convex dorsal outline of skull in longitudinal axis. The length of tlie hind foot averages more than the basal length in males whereas the reverse is true in females. The tail, relative to the length of the body, is longer in males than in females. The average differences in external Description. — Size. — Male and Female: LoCAIiITT Number of specimens averaged Total length Length of tail Per cent of body- length Lencth of hind foot Massachusetts Sad. cT 413 (390-432) 146 (127-159) 54% 46.0(41.0-48.0) 4 ad. 9 311 (298-321) 104 ( 95-114) 50% 33.9 (31.5-37.0) Liberty Hill, Conn 10 ad. cf 411 (379-438) 141 (124-155) 52% 47.1 (43.0-51.5) 6 ad. and sad. 9 318 (303-338) 105 ( 80-123) 49% 33.0 (31.7-36.0) Beaver Dam, Wise 10 ad. cf 407 (372-431) 130 (113-143) 47% 46.0 (42.0-50.0) 4 ad. 9 326 (303 338) 99 ( 86-108) 43% 35.6 (34.5-38.0) Washtenaw Co., Mich.. 10 ad. c? 371 (3.50-405) 130 (115-140) 54% 45.0 (40.0-50.0) 10 ad. 9 306 (290-335) 97 ( 90-120) 46% 34.0 (30.0-40.0) 224 University of Kansas Publs., Mus. Nat. Hist. measurements of the two sexes in Massachusetts, are: total length, 104; length of tail, 42; length of hind foot, 12.1. In Michigan, where the males are smaller, corresponding diflFerences are only, 65, 33, and 11. Weight of 19 adult males from New York (Hamilton, 1933:294), 225 (196-267) grams and in 13 adult females, 102 (72-126) grams. Weights of 2 adults from Michigan are: $ 258; 9 101 grams. Externals. — Longest facial vibrissae black, brown, or white (often all three colors in same specimen ) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 19. Color. — Upperparts, in summer, Vandyke Brown or darker than tone 4 of Burnt Umber of Oberthiir and Dauthenay, pi. 304. Sometimes approaching tone 2 of Warm Sepia of Oberthiir and Dauthenay, pi. 305. Underparts, in summer, ranging from white through Napthalene Yellow (Peterboro, N. Y. ), Pale Orange Yellow (eastern Mass.), near Primuline Yellow (unusual speci- men from Leelanau Co., Mich.) to near (c) Deep Chrome (no. 19053, U.S. Nat. Mus., Roan Mts., N. C. ). In winter, all white except tip of tail, or upper parts near ( 12" 1 ) Rood's Brown and tone 2 of Raw Umber of Oberthiir and Dauthenay, pi. 301, with underparts white or sometimes tinged with yellowish. Tip of tail at all times black. Upper parts of unifonn color except for occa- sional slight darkening of nose. Color of underparts extends distally on pos- terior sides of forelegs to foot and sometimes over upper sides of toes and on medial sides of hind fimbs only to knees. Least width of color of underparts averaging, in a series of twenty-two males, mostly in full winter pelage, from Liberty Hill, Connecticut, 21 (11-40) per cent of greatest width of color of underparts. In eleven females from the same place, corresponding percentages are 20 (14-29). Black tip of tail in same series of males, most of which are in full winter pelage, 70 (60-75) mm. long; thus longer than hind foot and averaging 50 per cent of length of tail-vertebrae. Skull and teeth. — Male ( based on ten adults from Massachusetts ) : See measurements and plates 16-18; weight, 3.6 (3.3-4.4) grams; basilar length, 44.6 (43.3-46.0); zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more or less than length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less (usually more) than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 6 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa behind or directly below posterior fourth of ml. Female ( based on five adults from Mass. ) : See measurements and plates 31-33; weight, 1.7 (1.2-2.1) grams; basilar length, 36.5 (35.2-38.1); zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital MUSTELA FRENATA 225 breadth more or less than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more or less (usually less) than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 4 to 5/2 upper incisors; height of tympanic bulla less than distance from its an- terior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum. The skull of the female averages 53 per cent hghter than that of the aver- age male. Comparisons of tlie skull with tliose of M. /. oJivacea, M. f. spadix, M. f. primulina, and M. /. arthuri, are made in the accounts of those subspecies. As compared with that of M. /. occisor the skull of adult male noveboracensis, is of smaller average size with relatively (to basilar length of Hensel) lesser mastoid and zygomatic breadths. In addition to the zygomatic arches of noveboracensis being less widely bowed outward they seem to be more rounded posteriorly. Comparisons of subadult females indicate that these differences exist in the females as well as in the adult males. Remarks. — The earliest of the post-Linnean references to this weasel mostly were under the specific name erminea in the be- lief that the American animal was the same as the larger of the two common species of weasel in the Old World. The name noveboracensis, now in use for this subspecies, was applied in 1840 and since that time the males usually have borne that name; the females, because they are smaller, were more frequently con- fused with some other species. Audubon and Bachman in 1853 even proposed the name agilis for the female in the mistaken behef that it was a species distinct from the male. After 1896, when Bangs correctly classified the weasels of the eastern United States, the males have been correctly identified and the females, except by a few authors, likewise have been correctly named. Because many early American naturalists did their first collecting of mam- mals in the geographic range of noveboracensis, the person who examines labels of specimens of this subspecies can find data written in the hand of Spencer Fullerton Baird, Theodore Roosevelt, and other naturalists famous for their work as scientists or accomplish- ments otherwise. The material is more nearly adequate than is that of many other subspecies and the number of specimens is exceeded — and only slightly — by that of the subspecies nevadensis, which like noveboracensis has a relatively large geographic range. Intergradation with Mustela frenata spadix is indicated by sub- adult males from western Wisconsin, namely, one from Gordon, 8—3758 226 University of Kansas Publs., Mus. Nat. Hist. three from Colfax and one from Meridean. Linear measurements of the teeth of these specimens are exactly intermediate between those of spadix from Elk River, Minnesota, to the west, and nove- boracensis from, say, Beaver Dam, Wisconsin, to the east. The specimens from western Wisconsin show approach to spadix also in that the length of the tooth-rows and breadth of the rostrum are shghtly greater than in noveboracensis from farther east, say, Beaver Dam, Wisconsin, Indeed, animals from as far east as Beaver Dam itself might be thought of as showing some approach to spadix. Although, along the eastern seaboard, the upper lips, with rare exceptions, are the same color as the underparts, farther west, in Michigan and Wisconsin, the lips more often than not are white. Animals from Beaver Dam have slightly shorter black tips on the tails, broader extent of the light color of the underparts and females average slightly larger than typical noveboracensis, say, those from Massachusetts. Each of these differences reflects characters found better developed in the spadix-longicauda stock to the west. Toward the southern part of its range where noveboracensis meets M. /. olivacea there is a marked increase in yellowness of the underparts. This coloration of the underparts, since it is not so well marked in the northern part of the range of noveboracensis, might be regarded as showing intergradation with olivacea and primulina, each of which has far more intensely colored under- parts than does noveboracensis. Excepting this increase of yellow on the underparts, however, there are few if any characters of noveboracensis which undergo marked change as approach to the range of olivacea is made. Indeed, the characters of noveboracensis remain constant to within a relatively short distance of the geo- graphic range of olivacea. Notwithstanding the state of affairs described above, intergrada- tion seems to take place. Three specimens referred to novebora- censis but which at the same time are regarded as intergrades with olivacea are as follows: No. 28.300, Charleston Museum, from five miles east of York, South Carolina, is an adult female with a badly crushed skull. In external measurements the specimen agrees with noveboracensis. The underparts, as regards color and width, are intermediate. The general proportions of the skull and tympanic bullae agree with those of noveboracensis but the skull is larger than in any female of true noveboracensis and ap- MUSTELA FRENATA 227 proaches that of olivacea. The same can be said of a young female, no. 80, Ohio State Museum, from Roswell, Georgia. Another female, no. 171559, U. S. Nat. Mus., from Lookout Mountain, 1500 ft., Fort Payne, Alabama, is barely subadult. The external measurements are nearer those of olivacea. The color and narrowness of the underparts are typical of noveboracensis. The proportions and especially size of the skull show approach to olivacea, though they are nearer to noveboracensis when all features are taken into account. In the northern part of its range individuals of noveboracensis attain larger size than farther south. This ten- dency reaches its extreme, in males at least, in M. f. occisor of Maine. Specimens of noveboracensis from the Adirondacks of New York average larger (see cranial measurements on page 418) than those from farther south, and thus approach occisor in size as well as in geographic position. Also, occasional individuals which strongly show characters of occisor are found even farther south than the Adirondacks of New York. This is true of no. 96518, U. S. Nat. Mus., J ad., from Lunenburg, Massachusetts. The animal has a large skull of relatively great width much as in occisor, although its external measurements, relative length of tail and long, terminal, black brush place it with noveboracensis rather than with occisor. Of a pair of specimens from Ossipee, New Hampshire, the male, no. 77108, U. S. Nat. Mus., has a long (175 mm.) tail, and short (60 mm.) black pencil as in occisor, although otherwise it is referable to noveboracensis. Still another specimen, a sub- adult male, no. 4193, Mus. Comp. Zool., from Upton, Maine, has a longer (51 mm.) hind foot than noveboracensis although it other- wise agrees with that subspecies. As remarked by Bangs (1899: 55), other than fully adult specimens from the range of occisor are "troublesome", and would not be selected as distinct from noveboracensis if placed in a series of that subspecies, say, from New York State. In view of the facts that several specimens from intermediate localities combine the characters of noveboracensis and occisor, that noveboracensis in the northern part of its range averages larger than it does farther south and thus approaches occisor in size, and that occasional large specimens resembling occisor in several, but not all, features sometimes crop up in the northern part of the range of noveboracensis, it appears that nove- boracensis and occisor intergrade. Therefore they are treated as two subspecies of the single species, Mustela frenata. 228 University of Kansas Publs., Mus. Nat. Hist. Intergradation with M. /. primulina has been commented on in the discussion of that subspecies. Female, no. 159980, U. S. Nat. Mus., from Golconda, IlHnois, has many characters of primtdina but two young males from there agree better with noveboracensis. Examination of 283 adult and subadult skulls for malformation of the frontal sinuses revealed only ten that were not obviously malformed. Two were from New York, one from Massachusetts, one from Pennsylvania, and six from the 52 specimens from Michi- gan and Wisconsin. In addition, skulls of many young and even juveniles were malformed. Specimens examined. — Total number, 555, arranged alphabetically by states and provinces and, unless otherwise noted, from north to south by counties in each state. Except as otherwise noted specimens are in the United States National Museum. Alabama. DeKalb Co.: Fort Payne, 1. Connecticut. Litchfield Co.: Riverton, V; Gaylordsville, 1. Hartford Co.: East Hartford, 4 (3'); Glastonbury, 2'; South Glastonbury, 4\ Windham Co.: Plainfield, 2 (P). Fairfield Co.: Greenwich, 2'. New London Co.: Liberty Hill, 35 (33*, 2"). District of Columbia. Washington, 3; near Washington, 1; Eastern Branch, 1; Congress Heights, 1; Bennings, 1; no definite locaUty, 1. Georgia. Towns Co.: Young Harris, 1. Cherokee Co.: Canton, 1. Cohh Co.: Roswell, 1". Indiana. St. Joseph County: Notre Dame, 2°. Porter Co.: Hebron, 1. Miami Co.: Denver, 5 (4*, 1=°). Wells Co.: Bluffton, 1. Howard Co.: Russiaville, 1. Jay County: Salamonia, 1'. Boone County, 1'. Knox Co.: Bicknell, 3. Illinois. Lake Co.: Camp Logan, 3'; Fort Sheridan, T. Cook Co.: W Nortlifield, 2; Flossmoor, 1°; no locaUty more definite than county, 1. Du Page Co.: Bloomingdale Spg., 1°. Carroll Co.: Savanna, T. McLean County: Normal, 1". Champaign County: Harwood Township, 1"°. Pike Co.?: Milton Spring, 1°. Pope Co.: Golconda, 3. Kentucky. Woodford Co.: Midway,!. Hancock Co.: Hawesville, L Maine. Oxford Co.: Upton, 1*; Bethel, 1". Maryland. Howard Co.: Long Comer, 1; Hanover, 1. Montgomery Co.: Gaithersburg, 1; Garret Park, 1; Chevy Chase, 1; Bethesda, 1. Prince Georges Co.: Laurel, 18; Plummer Island, 3; Oxon Hill, L Talbot Co.: Easton, 1. Dorchester County: Cambridge, 5'. Massachusetts. Middlesex Co.: Wilmington, 6; Burlington, 6; Lexington, 1*; Wayland, 2*. Berkshire Co.: New Marlboro, 1'. Worcester Co.: Lunenburg, 2; Lancaster, 1'; Princeton, 2*. Norfolk Co.: So. Weymouth, 1*. Plymouth Co.: Wareham, 5^ Michigan, Marquette Co.: Michigamme, 1. Charlevoix County: Thumb Lake, 1°; M mi. N Thumb Lake, V. Leelanau County: Leland, 3°; Duck Lake, 2 mi. S Leland, 1°; Lost Pond, 8)2 mi. S Leland, T. Osceola County: Le Rov, 2". Huron County: Rush Lake, 1°. Saginaw Co.: East Saginaw, 1. Oak- land Co.: Royal Oak, 4°; South Lyon, 1°. Livingston County: Portage Lake, !•. Washtenaw Co.: Portage Lake, 6°; Waterloo, 2'; Lima, 1"; Ann Arbor, 11"; 3 mi. E Ann Arbor, V; 2 mi. SE Ann Arbor, 1°; 2 mi. S Ann Arbor, 1*; 3 mi. S Ann Arbor, 1"; Dixboro, 1'; Pittsfield, 3 (2°); Saline, 1'; near Sahne, 2*; 1 mi. S Saline, 2"; York, 2°; Manchester, 2'. Lenawee Co.: Morenci, 1*. Cass County: Marcellus Township, T. Berrien County: Harbert, 1'; Warren Wood Preserve, 1'; Warren Woods, 1". MUSTELA FKENATA 229 New Hampshire. Grafton Co.: Franconia, 1'. Carroll Co.: South Chatham, 4 (3'); Ossipee, 2; Intervale, 1'. Merrimack Co.: Webster, 2'. New Jersey. Morris Co.: Morristown, 1. Essex Co.: West Orange, 1*. Mercer Co.: Princeton, 1". Ocean Co.: Point Pleasant, V. Camden Co.: Haddonfield, 1'°. Cumberland County: Millville, 2". New York. St. Lawrence County: Ogdensburg, 1". Clinton Co.: Rouses Point, 1'". County?: Adirondacks, 12. Essex Co.: Elizabethtown, 1; Schroon Lake, 1; no locality more definite than county, 1. Lewis? Co.: Locust Grove, 4; Lyons Falls, 1. Warren County: Lake George, 6; Caldwell, 1. Hamilton County: Beaver Brook, /2 mi. above mouth Indian Lake, V\ Oswego Co.: Scriba, 2"; Palermo, 1". Monroe Co.: Penfield, 3. Madison Co.: Peterboro, 6 (2'). Schoharie Co.: Schoharie, 1'. Rensselaer Co.: East Shodack, 1". Tompkins Co.: Taughannock Falls, 2"; Ithaca, 4 (3"); Glenside, Ithaca, 1"; 6 mi. Creek, Itliaca, 1". Green County: Lanesville, V. Orange Co.: Pop- lopen's Pond, 1'; Highland Falls, 1'. Putnam County, 1". Westchester Co.: Sing Sing, 4; Armonk, 1^ Hastings, 3 (2', 1"). Nassau Co.: Flushing Meadows, 1'; Flushing, 1"; near Flushing, 1'; Oyster Bay, 2. Long Island: Cold Spring Harbor, 1; Bridgehampton, V. County in question: Severance, 3; Lake Grove (Long Island?), 1. North Carolina (east to west by counties). Wake Co.: Raleigh, 4 ( 1', 1', 2'). Mitchell Co.: Magnetic City, foot of Roan Mountain, 6; Roan Mt., 1; Roan Mt., 6000 ft., 3. Buncombe Co.?: Valley of Black Mts., 1'. Madison County, 2". Ohio. Trutnbull Co.: Warren, 1". Seneca Co.: Tiffin, 1". Summit Co.: Ira, 2'. Crawford Co.: Gafion, l"'. Ashland County: Loudonville, 1°. Au- glaize Co.: New Bremen, S"'. Franklin Co.: 3 mi. N Columbus, 1''; Minerva Park, Columbus, 5''. Fairfield County: Sec. 32, Pleasant Twp., 1"; Lancaster, 1"'. Clinton County: Reesville, 1; J2 mi. S and J2 mi. W Wilmington, 2". Pike Co.: Waverly, l^^ Ontario. Sudbury District: Metagama, 2"*. Carleton Co.: Ottawa, 2". Muskoka County: Lake of Bays, 1; Bracebridge, 1. Haliburton Co.: Gooder- ham, r. Simcoe Co.: OriUia, 4 (2', 2\) Prince Edward Co.: Bloomfield, 1". York Co.: Toronto, 1'. Waterloo Co.: Branchton, 3"; Preston, 2"; no locality save county, V. Welland Co.: Ridgway, 1'. Elgin Co.: St. Thomas, 1". Essex Co.: Kingsville, 1"; Point Pelee, 1". Pennsylvania (east to west by counties). Crawford Co.: Pymatuning Swamp, 3J2 mi. W Linesville, T'; Meadville, 2'\ Beaver County: Beaver, 1"; Raccoon Creek, 1"^. Butler Co.: Mars, 1'^ Leasuresville, 4". Allegheny Co.: Allegheny, 1. Warren County: Bear Lake, 2''. Westmoreland Co.: Kingston, 1"; Laughlinstown, 2". Somerset Co.: Confluence, 1''; Tub Mill Run, 2 mi. N Springs, 1'". Jefferson Co.: Siegel, 1". Clearfield ? Co.: Penfield,!". Cam- bria County: Cresson, V\ Fulton County: Well's Tannery, 1". Clinton County: near Round Island, 2". Cumberland Co.: Carfisle, 1. Snyder Co.: 5 mi. S Selinsgrove, 1. Northumberland Co.: Pottsgrove, 1. Union Co.: Mif- flinburg, 1. Sullivan County: Lopez, 7 (4", 3"). Chester Co.: Westtown, !"■; Valley Forge, V; W Bradford Township, 1'°; no locality more definite than county, 3. Philadelphia Co.: Hohnsburg, 2'°. Bucks County: V. Pike Co.: Milford, 1. Rhode Island. Providence Co.: Chepachet, 1. Washington Co.: Lake Warden, 2. Quebec. Megantic County: Black Lake 1". County in question: Meach Lake, 1". South Carolina. York County: 5 mi. E York, 1". Laurens County: Lau- rens, 1'". Tennessee. Campbell Co.: Highcliff, 1. Carter ? Co.: Roan Mts., 1". Hamilton Co.: Walden Ridge, near Soddy, 3. Vermont. Windsor Co. : Hartland, 1'. Virginia. Shenandoah Co.: Toms Brook, 1. Arlington Co.: Arfington, 1; Ballston, 1; Alexandria, 1. Fairfax Co.: Falls Church, 3; Mt. Vernon, 2; no 230 University of Kansas Publs., Mus. Nat. Hist. locality more definite than county, 1. Prince William Co.: Occoquan, 1. Essex County: Montague, 1. Prince George County, 1. Norfolk Co.: Wal- laceton, 1. Grayson County: Mt. Rogers, 3. County in question: Dismal Swamp, 1; Massanutten Mt., 1. West Virginia. Hardy County, 1. Pendleton Co.: radius of 2 mi. Smoke Hole, 1". Greenbriar Co.: White Sulphur, 2'. Wisconsin. Douglas County: Gordon, 1"°. Vilas County: Mamie Lake, 4. Dunn County: Colfax, 4'°; Meridean, 1'°. Door County: state game farm, 17'"; no locality more definite than county, 1'". Dodge County: Rolling Prairie, 1"; Beaver Dam, 52"'. Dane County: Wingra Lake, 1'°. Waukesha County: Pewaukee, 2'". Racine County: Waterford Township, 2'". Rock County: Mil- ton, 1'°; Bowers Lake, 1'°. Walworth Co.: Lane's Mill, 8 mi. N Elkhom, 7 {!'', 6'"-); Delavan, 7. Mustela frenata occisor (Bangs) Long-tailed Weasel Plates 16, 17, 18, 31, 32 and 33 Putorius occisor Bangs, Proc. New England Zool. Club, 1:54, June 9, 1899. Mustela occisor. Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912. Mustela frenata occisor. Hall, Carnegie Instit. Washington Publ. 473:104, November 20, 1936. Type. — Male, adult, skull and skin; no. 9102, coll. of E. A. and O. Bangs in Mus. Comp. Zool.; Bucksport, Hancock County, Maine; January 15, 1899; ob- tained by Alvah G. Door but measured and sexed by O. Bangs. The skin is well made and in good condition. It is in full, white winter- dress with black-tipped tail. The skull has the posterior half of the left zygo- matic arch broken away; otherwise the skull is unbroken and complete. Left 13 and right P3 are missing. The teeth otherwise all are present and entire. Range. — Maine; possibly north locally to south side of St. Lawrence River in Quebec and possibly occurring in western New Brunswick. Zonal range Canadian and probably Transition. See figure 29 on page 221. Characters for ready recognition. — DiflFers from M. /. noveboracensis by a number of average differences including larger size, relatively longer tail and relatively wider skull (see p. 225, and measurements on pp. 418, 419). Description. — Size. — Male: Five adults yield average and extreme measure- ments as follows: Total length, 443 (430-465); length of tail, 163 (154-175); length of hind foot, 50 ( 47-54 ) . Tail averages 58 per cent as long as head and body. Length of hind foot averages more than basal length. Female: Measurements of two subadult female topotypes are as follows: Total length, 346, 318; length of tail, 116, 110; length of hind foot, 39, 35.5. 1. Boston Society of Natural History 14. Collection of Edward R. Warren 2. Collection of Alex Walker 15. Charleston Museum 3. American Mus. Nat. History 16. Univ. Kansas Mus. Nat. History 4. Museum of Comparative Zoology 17. National Museum of Canada 5. University of Notre Dame 18. Carnegie Museum 6. Field Museum of Natural History 19. Collection of R. H. Coleman 7. San Diego Society of Natural History 20. University of Wisconsin 8. Collection of R. W. Jackson 21. Collection of W. E. Snyder 9. Museumi of Zoology, Univ. Michigan 22. Collection of William Bebb 10. Acad. Nat. Sciences of Philadelphia 23. Ohio State Museum 11. Mus. Vert. Zool., Univ. California 24. Royal Ontario Museum of Zoology 12. New York State Museum 25. British Museum of Natiu-al History 13. Cornell University 26. Berlin Zoological Museum MUSTELA FRENATA 231 Tail amounts to 50 per cent and 54 per cent of body-length respectively. Length of hind foot more or less than (about equal to) basal length. The average diflFerences in external measurements of the two sexes are: Total length, 111; length of tail, 50; length of hind foot, 12.5. Externals. — As described in Musfela frenata noveboracensis. Color. — As described in Mustela frenata noveboracensis except that black tail-tip in series of 10 males in full v^^inter pelage 60 (45-80) mm. long; thus averaging 39 per cent of length of tail vertebrae. Skull and teeth. — Male (based on 3 adults): See measurements and plates 16-18. As described in Mustela frenata noveboracensis except that: Weight, 4.2 (4.1-4.3) grams; basilar length, 45.7 (44.9-46.9); zygomatic breadth more or less than (about equal to) distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate rarely less than length of P4; anterior margin of masseteric fossa behind or directly below posterior half of m2. Female (based on 2 subadults): See measurements and plates 31-33. As described in Mustela frenata noveboracensis except tliat: Weight, 2.0 ( 1.9-2.1) grams; basilar length, 37.3, 38.2. Comparison of the skull with that of M. f. noveboracensis is made in the account of that subspecies. Remarks. — Excepting a specimen in the Academy of Natural Sciences of Philadelphia, obtained in 1893, and two in the Boston Society of Natural History, obtained in 1925, I have seen no ma- terial of this subspecies in addition to that examined by Bangs at the time he prepared the original description in 1899. Anderson (1945:56, 57) records a specimen, Canadian National Museum Catalogue Number 18426, from Kamouraska County, Que- bec, as of this subspecies and thinks that occisor occurs north of Maine "locally to south side of lower St. Lawrence River in Quebec; probably also in western New Brunswick." So far as the available material of occisor permits one to judge^ it is distinguished from noveboracensis by a combination of char- acters no one of which invariably can be relied upon as diagnostic. Employing adult males, average differences indicate that M. f. occisor is larger in each of the external and cranial measurements; tail relatively longer; black tip of tail relatively shorter; mastoid and zygomatic breadth relatively greater and zygomatic arches more nearly square posteriorly. Considering the large number of specimens of noveboracensis which are available in comparison with the few of occisor it is not surprising that some noveboracensis should be found which exceed in size those of occisor. This is the case as regards the basilar length of a very old male, no. 96518, U. S. Nat. Mus., from Lunenburg, Massachusetts. Also, the skull is actually broader than any of those 232 University of Kansas Publs., Mus. Nat. Hist. of occisor. However, this specimen is much older than any occisor examined. In a female, no. 4260, Mus. Comp. ZooL, from Liberty Hill, Connecticut, the skull is longer (but nairower) than in either of the two available females of occisor. The average differences pointed out above which characterize this extreme northern population of noveboracensis-like weasel in com- parison with true noveboracensis without much question are geo- graphic variations. Whether or not tliese variations are of a degree sufficient to warrant nomenclatural recognition is debatable. With equally scanty material from other regions I have not named varia- tions seemingly as great as those shown by occisor. The paucity of material of occisor is a handicap in making a decision in this in- stance. Each of the adult and subadult specimens, except the one from Perry, shows malformation resulting from the infestation of the frontal sinuses with parasites. Specimens examined. — Total number, 18, listed by counties from west to east and unless otherwise indicated in the Museum of Comparative Zoology. Maine. Oxford Co.: South Andover, 2 (Boston Soc. Nat. Hist.); Umbagog Lake, 1. Franklin Co.: Seven Pd. Township, 1. Piscataquis Co.: Grenville, [= Greenville?], 1. Hancock Co.: Bucksport, 10. Washington County: 3rd Mopang Lake, 1 (Acad. Nat. Sci. Phila. ); Perry, 1 (Boston Soc. Nat. Hist.). County in question: Moosehead Lake, 1. Mustela frenata primulina Jackson Long-tailed Weasel Plates 16, 17, 18, 31, 32 and 33 Mustela primulina Jackson, Proc. Biol. Soc. Washington, 26:123, May 21, 1913. Putorius noveboracensis, Baird, Mamm. N. Amer., p. 166, 1858 (part). Mustela longicauda longicauda. Dice, Joum. Mamm., 4:108, May 9, 1923. Mustela longicauda primulina, Linsdale, Joum. Mamm., 9:141, May 9, 1928. Mustela frenata primulina. Hall, Carnegie Instit. Washington Publ. 473:104, November 20, 1936. Mustela frenata, Leopold and Hall, Joum. Mamm., 26:143, July 19, 1945. Type.— Male?, young, skull and skin; no. 168006, U. S. Nat. Mus., Biol. Surv. Coll.; 5 miles northeast of Avilla, Jasper County, Missouri; May 11, 1905; obtained by Hartley H. T. Jackson; original no. 7869X. The skin lacks the distal part of the tail — the part which bears the black tip. Otherwise the skin is complete and well preserved. The teeth of the permanent dentition all are present and entire. The lower jaws are complete and unbroken. The skull is broken transversely through the interorbital region, transversely through the braincase and longitudinally through the basioccipital. Both zygomatic arches are gone. The type is judged to be a male rather than a female as stated by the original describer, Jackson (1913:123), whose measurements of hind foot, interorbital constriction, maxillary tooth-row, and mandibular tooth-row agree with those of males and are larger than those of any female seen of this subspecies. MUSTELA FRENATA 233 Range. — Upper and Lower Austral life-zones west of the Mississippi River in Missouri and Arkansas, the soiithcastcni half of Iowa, eastern half of Kansas and Oklahoma, northern Louisiana and northeastern Texas. Southern and southwestern Umits of range undetemiined. See figure 29 on page 221. Characters for ready recognition. — DiflFers from M. f. novehoracensis in males by interorbital breadth averaging less than 24 per cent of basilar length, or- bitonasal length averaging less than 34 per cent of basilar length or 64 per cent of mastoid breadth, tjmpanic bullae as much inflated anteromedially as postero- medially, and in females by orbitonasal length amounting to less than two- thirds of mastoid breadth, by zygomatic breadth averaging more than 21 mm., and by anterolateral margin of tympanic bulla projecting below squamosal; from M. f. spadix by least width of color of under parts amounting to less tlian 40 per cent of greatest width of color of upper parts, by absence of color of underparts on hind leg below knee, and by smaller size (hind foot less than 50 in males and 40 in females; orbitonasal length less than 15.5 in males and 13.5 in females; length of tooth-rows less than 18 in males and 15.7 in females; mastoid breadth less than 25.5 in males and 22 in females); from M. f. longicauda by Brussels Brown rather than near (h) Clay Color of upper parts, least width of underparts less than 40 per cent of greatest wadth of color of upper parts, absence of color of underparts on hind leg below knee, zygo- matic breadth less than 28.8 in males and 24.1 in females; from M. /. neo- mexicana by Brussels Brown rather than Buckthorn Brown color of upper parts, in absence of white frontal spot and broad white bands on sides of head, in anterolaterally rounded, rather than "square", tympanic bullae and in zygomatic breadth of less than 30 in males and 24 in females; from M. /. frenaia and M. f. texensis by absence of white facial markings and smaller size ( basilar lengtli of adult males less than 47, tail-length less than 155 in males, and hind foot less than 40 in females); from M. f. arthuri by less evenly spreading zygomatic arches (see pis. 16, 17 and 18), greater inflation of the tympanic bullae anteromedially and more nearly straight (less convex) dorsal outline of skuU. Description. — Size. — Male: Eighteen adults and subadults from Douglas County, Kansas, yield average and extreme measurements as follows: Total length, 397 (371-440); length of tail, 133 (120-147); length of hind foot, 43 (40-47). Tail averages 50 per cent as long as head and body. Length of hind foot averages less than basal length. Corresponding measurements, originally taken in inches and fractions thereof, of 9 adults and subadults from Boone County, Arkansas, are as follows: 413 (384-438); 138 (127-155); 41 (38-44). Female: Six adults and subadults from Douglas County, Kansas, yield average and extreme measurements as follows: Total length, 339 (317-355); length of tail, 107 (95-115); length of hind foot, 35 (34-37). Tail averages 46 per cent of length of head and body. Length of hind foot less than basal length. Corresponding measurements, originally taken in inches and fractions thereof, of 5 adults and subadults from Boone County, Arkansas, are as follows: 355 (332-397); 113 (101-127); 33 (29-38). The average difi^erences in external measurements of the two sexes, in Douglas County, Kansas, are: Total length, 58; length of tail, 26; length of hind foot, 8. An adult male from Boone Co., Iowa, weighed 293 grams. 234 Univ'ersity of Kansas Publs., Mus. Nat. Hist. Externals. — Longest facial vibrissae black or dark brown (often both colors in same specimen) and extending beyond ear; carpal vibrissae colored either like underparts or upper parts and extending to apical pad of fifth digit; hairi- ness of foot-soles as shown in figure 20, Color. — Upper parts, in summer, Brussels Brown to near ( 14 n ) Brussels Brown, or tones 2 to 4 of Raw Umber of Oberthiir and Dauthenay, pi. 301. Chin and rarely upper Hps white. Remainder of underparts Picric Yellow to PrimuUne Yellow. In winter, color essentially the same except for smoke-gray effect in upper parts and more whitish in underparts. Tip of tail at all times black. Upper parts of uniform color except for occasional darkening of nose and mid-dorsal region. Color of underparts extends distally on posterior sides of forelegs onto antipalmar faces of toes, on medial sides of hind legs only to a point between knee and ankle. Least width of color of imderparts averaging, in a series of 21 males from Lawrence, Kansas, 23 (9-35) per cent of greatest width of color of upper parts. Black tip of tail in same series, most of which are in full winter pelage, 52 (40-70) mm. long; thus longer than hind foot and averaging 39 per cent of length of tail-vertebrae. Skull and teeth. — Male (based on ten adults from Douglas County, Kansas): See measurements and plates 16-18; weight, 3.7 (3.3-4.2) grams; basilar length, 44.8 (43.8-46.0); zygomatic breadth more or less (less in 80 per cent) than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior end of tympanic bulla (less in 70 per cent); mastoid breadth more or less (less in 80 per cent) than postpalatal length; postorbital breadth less than length of upper premolars and, except in one specimen, more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less (less in 70 per cent) than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic buUa; least width of palate less than outside length of P4 (except in one specimen); anterior margin of tympanic bulla as far posterior to foramen ovale as widtli of 2/2 to 5 upper incisors; height of tympanic bulla more (except in one speci- men) than distance from its anterior margin to foramen ovale; length of tympanic bulla more tlian length of lower molar and premolar tooth-row and longer (except in one specimen) than rostrum; anterior margin of masseteric fossa behind or just below posterior eighth of ml. Female ( based on 5 adults and subadults from Douglas County, Kansas ) : See measurements and plates 31-33; weight, 2.2 (2.0-2.4) grams; basilar length, 38.9 (37.6-40.7); zygomatic breadth less than distance between condylar fora- men and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and except in one specimen, more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than greatest length of P4; t>Tnpanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrLim. The skull of the female averages 41 per cent hghter than that of the male. Compared with tlie skull of M. /. noveboracensis from Massa- chusetts, that of the male of primulina, in dorsal view, is seen to be MUSTELA FRENATA 235 shorter anteriorly to the postorbital processes and to have a more marked postorbital constriction. In lateral view the dorsal outline of the skull of primnlina is less concave in the postorbital region. In ventral view tlie skull of primtilina is seen to be wider across the mastoid processes and zygomatic arches but the most pronounced difference is in the tympanic bullae. In noveboracensis each bulla is scooped out on the anterior part of the medial face and appears to be narrower anteriorly than posteriorly whereas in primulina the anterior part of the medial face is not scooped out but is moderately inflated and the bulla appears to be of uniform breadth anteriorly and posteriorly. By actual measurement the breadth of the bulla averages 59 per cent of its length in primulina but only 50 per cent in noveboracensis. Other respects in which the skull of the male of primulina differs from that of noveboracensis are as follows: Linear measurements of teeth more; relative to the basilar length, the length of the tooth-rows averages more, whereas the interorbital breadth and orbitonasal length are less. When skulls of females are compared, each of the differences mentioned above is found to apply, except that the degree of dif- ference is in some parts greater, for example, in the tympanic bullae. In primulina, the bulla is in general like tliat of the male novebora- censis, whereas in the female noveboracensis it is less inflated, es- pecially anteromedially, shorter, relatively narrower, and in ventral view projects little or none below the squamosal floor of the brain- case. The breadth of the bulla averages 51 per cent of its length in primulina but only 47 per cent in noveboracensis. The bullae pro- ject below the basioccipital on the average, for a distance of 2.9 millimeters in female primulina and only 2.3 millimeters in female noveboracensis. In primulina the temporal ridges are well devel- oped and fuse to form a low sagittal crest, but in noveboracensis the ridges are absent. Also, in primulina the mastoid processes project farther laterally beyond the braincase. The skull of female nove- boracensis is much lighter than that of primulina. Average weights of the two are 1.7 and 2.2 grams. The skulls of females of primulina and noveboracensis differ more than do the skulls of males. Compared with the skull of spadix, that of the male, and the female, of primulina averages smaller in every part measured. Ex- pressed in percentages of the basilar length, the two depth measure- ments of the skulls are not significantly different, but, excluding the measurements of the bullae and teeth, the other cranial measure- ments are less. The main difference in relative proportions is in the tympanic bullae which average only a half millimeter shorter in males of primulina and one and one-tenth millimeters shorter in 236 University of Kansas Publs., Mus, Nat. Hist. females. The bullae are, therefore, relative to the basilar length, longer in primulina. The skull of primulina, then, differs from that of spadix mainly in smaller size and relatively longer tympanic bullae, especially in males. Compared with the skull of M. f. longicaiida, that of both sexes of primulina averages smaller in every part measured, except in males where the length of the tympanic bulla, and breadth and length of Ml are the same or slightly larger in primulina. Relative to the basilar length, the length of the tympanic bullae, and in females only, the depth measurements are greater in primulina but all the others, in both sexes, are less. These ratios reflect the relative nar- rowness of the skull of primulina. Upon direct comparison the narrowness is especially noticable in the interorbital region, mas- toid region, tympanic bullae, and across the zygomata. Compared with the skull of M. f. neomexicana that of both sexes of primulina averages smaller in every part measured. Excepting the measurements of the teeth, most of the other measurements are constantly larger. Relative to the basilar length, the length of tooth- rows and length of tympanic bulla are more, but excepting the depth measurements, the others are less. Still other differences are, in primulina, less well-developed sagittal crest, anterolateral comer of bulla rounded rather than "square", and in males a transversely con- vex rather than flat interorbital region. Compared with M. /. frenata and M. /. texensis, the skulls of males of primulina differ in being smaller in every part measured but relative to the basilar length, have longer tooth-rows, a lesser zygo- matic breadth and are less constricted interorbitally. Compared with the skull of M . /. olivacea, those of both sexes of primulina average smaller in every part measured, have shallower ( dorsoventrally ) tympanic bullae, a lower sagittal crest and slightly weaker postorbital processes on the frontals. Relative to the basilar length, the several cranial measurements are about the same. Comparison of the skull with that of M . /. arthuri has been made in the account of that subspecies. Remarks. — The first specimens of this race known to have been preserved in study collections are one in the United States National Museum, taken at Bridge, Carroll County, Missouri, many years ago by J. Burbage, and less than a dozen specimens preserved before 1900 from eastern Kansas in the University of Kansas Museum of Natural History. In 1913 Hartley H. T. Jackson bestowed a name on this animal on the basis of two specimens taken by him in south- western Missouri. Later, through the efforts of Charles D. Bunker, MUSTELA FRENATA 237 and his associates at the University of Kansas, nearly 100 specimens were saved from eastern Kansas, principally from Douglas County. In the course of the present study, Lav/rence V. Compton obtained a topotype for the California Museum of Vertebrate Zoology, and with the assistance of Mr. B. G. Roberts, a good series of specimens from Boone County, Arkansas, was preserved in the same museum. In the early years of the 20th Century, the late B. H. Bailey at Coe College, Iowa, collected specimens from that state. The specimens from these several sources suffice to give a relatively clear idea of the characters of this subspecies. Mustela frenata primulina is closely related to M. f. novebora- censis, from which, on the average, it differs in the lighter color of the upper parts of the summer coat, in the more intense coloring of the underparts, and in certain cranial features pointed out above. In the southern part of its range, however, noveboracensis has the underparts only a little less intensely colored with yellow than primulina. Also, the skull of the one topotype from 7^2 miles south- east of Carthage, a subadult male in brown winter pelage, is almost exactly intermediate between that of noveboracensis from Massa- chusetts and primulina of Douglas County, Kansas, and Boone County, Arkansas. M. /. primulijia often has the underparts white in winter, as does this topotype which agrees with the average of noveboracensis in small size of teeth and narrowness across the mastoid processes and zygomatic arches. However, it agrees with primulina in shape and relative size of the rostrum. It is almost exactly intermediate in shape and width of the tympanic bullae. Three other males, but no females, all in winter pelage, are avail- able from eastern Missouri. Of the two from Silex, Lincoln County, one is nearer noveboracensis and the other nearer primulina on the basis of cranial characters. The third specimen, from four miles south of Lesterville, so far as I can determine by examination of individual cranial characters and tabulation of results, is exactly intermediate. Final decision on the proper allocation of specimens from the parts of Missouri represented can best be made when skulls of females are available. From the fact that the skull of the female referred to noveboracensis from Golconda, Illinois, shows almost as many characters of primulina as of noveboracensis, it is judged that females from as far west as Silex and Lesterville, Missouri, will show even more characters of primulina and so be referable to that form. If this supposition be correct, the present reference of the almost exactly intermediate males, from eastern Missouri, will stand; otherwise, it may not. 238 University of Kansas Publs., Mus. Nat. Hist. Additional intergrades with noveboracensis are available from eastern Iowa. Of five specimens from Hillsboro, Iowa, two males and a female have tympanic bullae like those of primulina but the other two males have bullae like those of noveboracensis. The fe- male is smaller than primulina and in this small size and in general configuration of the skull, viewed dorsally, is more nearly like nove- boracensis. As a whole, the population averages almost exactly in- termediate. The same is true of 3 males and one female from Mus- catine. The subadult male from Keosaqua, to my eye, resembles noveboracensis in the greater length of the skull anteriorly to the postorbital processes, and in the relative narrowness across the mas- toidal region, but otherwise is more like that of primulina. Two males and one female from Tipton, although in each instance var- iously intermediate, are as a whole nearer primulina. No. 2865, Coe College, male adult, from Cedar Rapids, has characters of the three races, spadix, noveboracensis and primulina. In the skull, the width suggests spadix, the narrow mastoid region, noveboracensis, and the tympanic bullae are as in spadix or primulina. One male, no. 12, Coe College, from Dubuque, is as narrow across the mastoid region as is noveboracensis although the bullae are well inflated as in primulina. The skull, without corresponding skin, of a female, no. 140a, Iowa State College, from Green's Island, also resembles noveboracensis in narrowness of the mastoidal region, and in small size of skull, but in larger teeth, broader tympanic bullae, and sagittal crest is referable to primulina. Of two females from Vin- ton, one adult is typical of primidina but the other, a subadult, is practically indistinguishable from female noveboracensis, from Ann Arbor, Michigan. Three males from Vinton agree well with prim- ulina except that the interorbital region is wider than average and thereby suggests spadix or noveboracensis. An adult female from New Hartford also is typical of primulina except for the broader interorbital region. Three males from Fayette are typical of primu- lina. Other specimens from Iowa are intergrades with spadix, or if not with spadix, with the animal of northwestern Iowa which in some ways combines the characters of longicauda and spadix. For ex- ample, no. 2665, Coe College, an adult male from Davenport, has the anterior part of the skull (all that is preserved) heavily ridged as in spadix and in addition, the underparts are marked with the shade of reddish displayed by topotypes of spadix and with some yellowish as seen in longicauda. The color pattern, however, is as in primulina. A young male, no. C-51, Iowa State College, from Kelley, Story County, has anteriorly truncate bullae as are more MUSTELA FRENATA 239 frequently found in tlie longicauda-spadix stock of northwestern Iowa, than in primulina. In other respects, the animal, in so far as can be judged from the broken skull, agrees with primulina as it certainly does in color, color pattern, and external measurements. An adult male, no. 499a, Iowa State College, from 2 miles east of Ledges St. Park, in Boone County, in short body, size of teeth, and size of skull, in so far as the broken parts can be measured, resembles primulina more closely than it does any other subspecies. The long tail, long hind foot, wide extent of the light-colored under- parts, and extension of the color of the underparts onto the hind feet are more as in spadix. Other intergrades with spadix from Iowa are mentioned in tiie account of spadix. The specimen from Swartz, Louisiana, suggests intergradation with arthuri in that tlie anteromedial part of the tympanic bulla is less inflated than in typical primulina. Intergrades with longicauda are available from Riley and Pratt counties, Kansas. No. 7182, Univ. Kans., subadult male in winter pelage, from near Winkler, has a skull of larger size as in longicauda with which race it seems to agree in large size of body, tail and hind foot, although the collector's measurements are lacking. Color pat- tern and relative proportions of the skull throughout are as in primulina. The young male, no. 3495, Univ. Kans., from Pratt, Kansas, agrees in external measurements and large size of skull with longicauda, but has the color and color pattern precisely as in primulina. The teeth are smaller as in primulina. Immaturity pre- vents judging of its relationships on the basis of relative proportions of the skull. The two specimens, skins only, available from Oklahoma, are pro- visionally referred to primulina. These are remarkable for the re- striction of the color of the underparts and for the intensity of the yellow coloration of the underparts. The specimen from Norman has the color of the underparts entirely absent from the hind legs and not extending posteriorly to the penis. On the chest and lower throat, large spots of color of the upper parts are present and the yellow area of the underparts on the belly is narrower than in any other specimen of primulina examined. The specimen from 8 miles northwest of Stillwater has the color of the underparts only a little less restricted although this color does extend over the inguinal region almost to the knees. The skin of the posterior part of the body of a weasel is available from 10 miles south of Sulphur Springs, Texas. It, likewise, is only provisionally referred to primulina. The coloration is about as in the specimens from Oklahoma but the dis- tribution of the color of the underparts cannot be made out. 240 University of Kansas Publs., Mus. Nat. Hist. The dark color of the upper parts occurs far westward in animals which otherwise display characters of longicauda. Among these intergrades, the larger size of longicauda generally is combined with this dark color. This geographic behavior of the dark color of the upper parts is analogous to the condition described in M. /. spadix. Stated in another way, the dark color of the upper parts is the character, of the eastern animal, last to disappear as one goes westward across the Mississippi Valley toward the range of longi- cauda which is a subspecies of markedly different size, shape of skull, and coloration. Only two of 29 specimens from Kansas show infestation of the frontal sinuses. All four of the specimens from Missouri have the frontal sinuses malformed as do 9 of the 14 from Arkansas examined in this respect. An adult female from Boone County, Iowa, bears the date May 9, 1938, and the annotation by T. G. Scott, "Fox-killed." Specimens examined. — Total number, 131, arranged alphabetically by states and from north to south by counties in each state. Except as otherwise indi- cated, specimens are in the University of Kansas, Museum of Natural History. Arkansas. Boone County: 3 mi. E Bergman, 4'; 3 mi. SE Bergman, V; 3 mi. S Bergman, 1^ 3 mi. SW Bergman, 1'; 4 mi. SE Bergman, 2'; 5 mi. SE Berg- man, 1'; 4/'2 mi. SE Bergman, 3'; 5 mi. SE Bergman, 1'; 5 mi. S Bergman, 2'; 5 mi. SW Bergman, 2\ Washington Co.: Fayetteville, 1". Crawford County: 10 mi. S Winslow, 1. Sebastian Co.: Fort Smith, V. Iowa. Fayette County: Fayette, 3*. Dubuque County: Dubuque, V; Green's Island, V. Butler County: New Hartford, 1*. Hardin County: Union, 1°. Benton County: Vinton, 5*. Linn County: Cedar Rapids, 1\ Boone County: Worth Township, Sec. 21, 1°; 2 mi. E Ledges St. Park, 1'. Story County: Kelley, 1°. Cedar County: Tipton, 3*. Scott County: Davenport, 2*. Muscatine County: Muscatine, 4\ Henry County: Hillsboro, 5^ Van Buren County: Keosaqua, 1°; no locality more definite than county, 1°. Taylor County, 1. Kansas. Riley County: near Winkler, 1. Pottawatomie County: Onaga, 1'. Atchison County: Doniphan Lake, 1; 5 mi. NE Muscotah, 1; no locality more definite than county, 1. Douglas County: Lawrence, 8; 6 mi. NW Lawrence, 1; 1/2 mi. W Lawrence, 1; 6 mi. S Lawrence, 1; 7 to 7/2 mi. SW Lawrence, 14; 10 mi. W Lawrence, 1; Chnton, 4; Baldwin, 1; no locality more than county, 29 (2*). Woodso7i County: 1/2 mi. S Neosho Falls, 1". Greenwood County: 8 mi. SW Toronto, 2. Pratt County: Pratt, 1. Louisiana. Quachita Parish: Swartz, 1". Missouri. Carroll County: Bridge Creek, P. Lincoln County: Silex, 1'; 1 mi. E Silex, V. Reynolds County: 4 mi. S Lesterville, 1\ Jasper County: 5 mi. NE Avilla, 1'; 7% mi. SE Carthage, V. Oklahoma. Payne Co.: 8 mi. NW Stillwater, 1'. Cleveland Co.: Nor- man, v. Texas. Hopkins Co.: 10 mi. S Sulphur Springs, 1". 1. Mus. Vert. Zool., Univ. California 7. Ottawa University, Kansas 2. University of Arkansas 8. Donald R. Dickey Collection 3. United States National Museum 9. Oklahoma Agric. and Mech. College 4. Coe College 10. University of Oklahoma 5. Iowa State College 11. Collection of RoUin H. Baker 6. Collection of W. E. Snyder 12. Louisiana State University MUSTELA FRENATA 241 Mustela frenata arthuri Hall Long-tailed Weasel Plates 16, 17 and 18 Mustela novehoracensis arthuri Hall, Proc. Biol. Soc. Washington, 40: 193, December 2, 1927. Mustela frenata arthuri Hall, Carnegie Instit. Washington Publ. 473:105, November 20, 1936. Type. — Male, subadult, skviU and skin; no. 37515, Mus. Vert. Zool.; Remy, St. James Parish, Louisiana; December 15, 1926; obtained by Stanley C. Arthur. The skin is stuffed and well preserved. The skull (plates 16-18) is un- broken. The teeth all are present and entire. The presence of a well-developed scrotal pouch shows the specimen to be a male. Contrary to what was stated in the original description the specimen was taken in 1926 and not in 1925. Range. — Lower Austral Life-zone of southeastern Texas, Louisiana, and into Mississippi. See figure 29 on page 221. Characters for ready recognition (of males). — Differs from M. f. olivacea in smaller size ( adult males with hind foot and basilar length less than 45 ) , depth of skull at anterior margin of basioccipital, ignoring sagittal crest, amounting to more than 63 per cent of mastoid breadth, and greater convexity of dorsal out- line of skull in longitudinal axis (see pis. 16-18); from M. /. novehoracensis, in males, by zygomatic breadth not less than distance between anterior palatine foramen and anterior margin of tympanic bulla and by convex dorsal outline of skull in longitudinal axis; from M. f. primulina by evenly spreading zygomatic arches, lesser inflation of tympanic bullae anteromedially than posteromedially, and convex dorsal outline of skull in longitudinal axis; from M. f. texensis and M. f. frenata by absence of white facial markings and postorbital breadth more than distance between posterior borders of P4 and P2. Description. — Size. — Male: The type, a subadult male, measures (inches and quarter fractions thereof, transposed into millimeters) as follows: Total length, 390; length of tail, 113; length of hind foot, 44. Tail is 41 per cent as long as head and body. Length of hind foot less than basal length. Typical female unknown. Externals. — Longest facial vibrissae black, or dark brown (both colors in the type) and extending beyond ear; carpal vibrissae same color as underparts and extending to within 3.5 millimeters of apical pad of fifth digit. Hairiness of foot-soles in type slightly less than shown in figure 20. Color. — Upper parts in summer tone 4 of Burnt Umber of Oberthiir and Dauthenay, pi. 304; underparts as described in M. f. olivacea. In winter, upper parts ( based on type ) near ( 1 ) Brussels Brown or grayer than tone 4 of Burnt Umber of Oberthiir and Dauthenay, pi. 304, darker on top of head from nose to a line connecting posterior margins of ears. Chin and posterior third of each upper lip white. Remainder of underparts white with wash of Warm Buff. Tip of tail black. Color of underparts extends distally on posterior sides of forelegs over toes but represented on antipalmar faces of feet by only a tew scattered hairs. Color of underparts extends distally on medial sides of hind limbs only to knees. Least width of color of underi^arts amounting to 15 per cent of greatest width of color of upper parts. Black tip of tail 50 mm. long; thus longer than hind foot and 44 per cent as long as tail-vertebrae. 242 University of ICansas Publs., Mus. Nat. Hist. Skull and teeth. — Male (based on type and 2 subadults): See measure- ments and plates 16-18. As described in M. /. noveboracensis except that: Weight, 4.0 (3.7-4.3) grams; basilar length, 43.5 (43.3-43.6); zygomatic breadth not less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than length of upper premolars; interorbital breadth more than distance between foramen opticimi and anterior margin of tympanic bulla; least width of palate more or less than length of P4; t>Tnpanic bulla longer than rostrum. Female: Typical skull unknown. The skull from 12 miles east of Eagle Lake, Texas, lacks the convexity in the dorsal longitudinal axis and the skull agrees with those of larger individuals of primulina except that the anteromedial faces of the tympanic bullae are less inflated, and the mastoid and zygomatic breadths are greater than in any female seen of primulina. Probably this greater breadth is the result of intergradation with M. f. frenata to the west- ward. Compared with the skull of M. /. olivacea that of arthuri diflFers as follows: Averaging smaller in every part measured; basilar length 5 mm. less; by weight a fourth hghter; relative to basilar length, interorbital breadth greater and zygomatic and especially mastoid breadth less; dorsal outline of skull more convex in longitudinal axis; tympanic bullae narrower and less inflated especially on anteromedial faces. Compared with the skull of noveboracensis that of arthuri has the zygomatic breadth equal to or exceeding the distance from the anterior palatine foramen to the anterior margin of the tyinpanic bulla, whereas the zygomatic breadth is less than this distance in noveboracensis. Also, in arthuri, the rostrum is relatively shorter, the braincase is more inflated anteri- orly, the zygomatic arches are more uniformly spreading, and the dorsal outline of the skull is distinctly convex, both transversely and longitudinally, whereas it is transversely more nearly flat in noveboracensis and longitudinally is concave in the interorbital region. Compared with M. /. primulina, arthuri has narrower bullae, which are much less inflated on their anteromedial faces, a less marked postorbital constriction, a braincase which is narrower across the mastoid region and broader anteriorly, and a skull, which, in longitudinal axis, has the dorsal outline markedly more convex. Compared with the skull of M. /. texensis that of arthuri is smaller in every part measured; length one-fifth less; one-half as heavy; postorbital constriction less marked; braincase relatively narrower posteriorly and tympanic bullae less inflated especially anteromedi- MUSTELA FRENATA 243 ally. Compared with the skull of M. f. frenata that of arthuri is smaller in every part measured; basilar length 6 mm. less; a third lighter; postorbital constriction less marked; relative to the basilar length the rostrum is broader, longer and deeper; the zygomatic expanse and breadth of the braincase across the mastoids is less; the dorsal profile of the skull is more convex in longitudinal axis; zygomata evenly spreading rather than abruptly protruding from skull posteriorly; tympanic bullae less inflated anteromedially. Remarks. — In 1926, Stanley C. Arthur, then Director of the Division of Wild Life, for the Louisiana State Department of Conservation, obtained specimens of this weasel. Some of them were mounted and the remainder were placed in the collections of the United States National Museum and the Museum of Verte- brate Zoology. In 1938 to 1940 George H. Lowery saved specimens from Baton Rouge, which showed the color of the summer pelage and revealed that the size of males was more than was indicated by the original materials. In 1940 and 1941 Rollin H. Baker ob- tained specimens from eastern Texas which greatly extended the known geographic range. In addition to the localities represented by specimens examined, Arthur (1928:117) has recorded specimens from Greensburg, St. Helena Parish; Braithwaite, Plaquemines Parish; Geismar, Assump- tion Parish; Laurel Hill, West Feliciana Parish; French Settlement, Livingston Parish; and Kentwood, Tangipahoa Parish. All these localities lie within the eastern half of southern Louisiana. A skin-only, no. 38902, Mus. Vert. Zool., obtained from a fur buyer by Stanley C. Arthur, was taken in Mississippi "south of Jackson". Possibly it is of the subspecies arthuri. Intergradation with Af. f. oUvacea is indicated by a specimen from Mobile County, Alabama, commented on in the account of olivacea. Intergradation with primulina is indicated by the shape of the anteromedial part of the bullae of the specimen from Swartz, Louisiana, that is referred to primulina. The lack of specimens from the northern two-thirds of Mississippi and from western Tennessee, prevents any definite statement as to the limits of range of arthuri in those areas. In comparison with animals from the type locality, the slightly larger size of the adult male from Baton Rouge, and the still larger size of the adult male of primulina from Swartz, Louisiana, suggests that the olivacea "influence" may 244 University of Kansas Publs., Mus. Nat. Hist. extend farther west in the latitude of northern Louisiana than any- where else. None of the skulls examined shows malformation of the frontal sinuses such as results from infestation by parasites in some races. Arthur (1928:115) speaks of the ". . . cut-over swamp land, where the tupelo and cypress have been removed, . . ." as constituting suitable habitat for this animal. Specimens examined. — Total number, 13, as follows: Texas. Colorado Co.: 12 mi. N Eagle Lake, V; 5 mi. W Eagle Lake,l'; 3 mi. S Garwood, 1*. Louisiana. East Baton Rouge Parish: Baton Rouge, 4°. Livingston Parish: Springville, 1\ Saint James Parish: Convent, 1"'; Remy, 2 ( P, 1'). Assump- tion Parish: near Lake Verret, 1'. Mississippi. Harrison Co.: Saucier, 1°. Mustela frenata olivacea Howell Long-tailed Weasel Plates 16, 17, 18, 31, 32 and 33 Mustela peninsulae olivacea Howell, Proc. Biol. Soc. Washington, 26:139, May 21, 1913. Mustela frenata olivacea, Hall, Carnegie Instit. Washington Publ. 473:104, November 20, 1936. Ttjpe. — Male, adult, skull and skin; no. 180802, U. S. Nat. Mus., Biol. Sur- veys Coll.; Autaugaville, Autauga County, Alabama; December 22, 1912; obtained by L. S. Golsan. The skull (plates 16-18), although cracked at two places in the interorbital region, is in one piece and not warped out of shape. The teeth all are present and entire. The skin is exceptionally well-made and in perfect condition except for the ex-treme tip of the tail which is broken off. Range. — Lower and Upper Austral life-zones in eastern Mississippi, Alabama, Georgia, South Carolina, and northern Florida. See figure 29 on page 221. Characters for ready recognition. — Differs from M. /. peninstdae in finer, softer pelage and shorter (less than 15.8 in ad. $ ) tympanic bullae; from M. /. novchoracensis, in adult males by wider tympanic bulla which is more than rather than less than 8.5, in adult females by total length which is more than rather than less than 345, and by mastoid breadth which is more than rather than less than distance between articular faces of exoccipital condyle and glenoid fossa; from M. /. arthuri in larger size (adult males with hind foot and basilar length each more than 45); depth of skull at anterior margin of basioccipital, ignoring sagittal crest, amounting to less than 63 per cent of mastoid breadth, and lesser convexity of dorsal outline of skull in longitudinal axis (See pis. 16-18). 1. Mus. Vert. Zool., Univ. California 4. Collection of Rollin H. Baker 2. United States National Museum 5. Louisiana State University 3. Collection of Stanley C. Arthur Catalogue Se; no. 47165 $ 47166 $ 47167 $ 41023 $ 41025 $ 223880 $ 198 S Average 7 $ 49385 9 41024 9 iCngth of tail Length of hind foot 160 48 147 47 145 45 140 47 142 47 145 49 140 48 146 47 124 45 [not typical] 125 41 MUSTELA FRENATA 245 Description. — Size. — Male and female: External measurements of adults are available as follows: Locality Total length Box Springs, Talbot Co., Georgia 454 Box Springs, Talbot Co., Georgia 435 Box Springs, Talbot Co., Georgia 422 Thomas Co., Georgia 443 Grady Co., Georgia 395 Okefinokee Swamp, Georgia 416 Okefinokee Swamp, Georgia 425 427 Gainesville, Alachua Co., Flordia 396 Thomas Co., Georgia 380 51527 9 Talbot Co., Georgia 376 128 43 The length of the hind foot averages less than the basal length in both males and females. The tail averages 52 per cent as long as the head and body in males and 51 per cent in females. Average differences in measure- ments of the two sexes are: Total length, 49; length of tail, 19; length of hind foot, 5. An adult male, no. 41023, and an adult female, no. 41024, each taken in February, 1929, on the Sinkola Plantation, Thomas County, Georgia, weighed 15 ounces (425 grams) and 7 ounces (198 grams) respectively according to Charles O. Handley. Externals. — As described in Mustela frenata noveboracensis, except that hairiness of foot-soles slightly less than shovra in figure 19. Color. — Upperparts, in summer, near tone 4 of Burnt Umber of Oberthiir and Dauthenay, pi. 304. In winter hghter, between tones 3 and 4 of Raw Umber of Oberthiir and Dauthenay, pi. 301. Dark spot at each angle of mouth present or absent. Underparts ranging from Massicot Yellow to Cream Buff except on chin and upper hps which are white. Tip of tail black. Upper parts of uniform color. Color of underparts extends distally on posterior sides of forelegs over antipalmar faces of toes and on medial sides of hind limbs to ankles. Least width of color of underparts averaging, in a series of five males from Talbot Co., Georgia, 29 (extremes 24-34) per cent of greatest width of color of upper parts. Black tip of tail in same series, averaging 65 (extremes 60-70) mm. long, thus longer than hind foot and averaging 43 per cent of length of tail-vertebrae. The spot at the angle of the moutli is absent in one-third of the specimens examined. The upper lips are white in specimens from the southern part of the range of olivacea but in the northern part of the range of the subspecies the upper lips are dark colored as in noveboracensis. Skull and teeth. — Male (based on 5 adults from Talbot Co., Georgia): See measurements and plates 16-18; weight, 5.3 (5.0-6.4) grams; basilar length, 48.3 (45.8-50.1); zygomatic breadth more or less (usually less) than distance between condylar foramen and Ml and more or less (usually more) than distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less than (averaging about equal to) postpalatal 246 University of Kansas Publs., Mus. Nat. Hist. length; postorbital breadth less than length of upper premolars and more or less than ( about equal to ) width of basioccipital meaured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than (about equal to) distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than greatest length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla not less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum (one exception); anterior margin of masseteric fossa below posterior half of m2. Female (based on 2 adults from Thomas Co., Ga., and one from Talbot Co., Ga. ): See measurements and plates 31-33; weight, 3.8 (3.5-4.0) grams; basilar length, 43.4 (42.7-44.0); zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less ( usually more ) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including 13) upper incisors; height of tympanic bulla not less (usually more) than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum. The skull of the female averages 28 per cent lighter than that of the male. Compared with the skull of M. f. peninsulae, of which only one good skull, and that a female, is available, that of M. /. olivacea averages smaller and has relatively and actually smaller and less in- flated bullae. As compared with the skull of M . /. noveboracensis, that of olivacea in the case of males is larger in every part measured and relative to the basilar length is broader across the zygomatic arches and mastoids. However, the rostrum and interorbital region are relatively narrower. The orbitonasal length is relatively less. The tympanic bullae are broader and more inflated. The same dif- ferences hold as between females of noveboracensis and olivacea. Indeed, the females of these two races differ more than do the males. Additional, selected differential cranial characters in the females are, in olivacea, as follows: Weight averaging 3.8 grams rather than 1.7 grams; braincase with, rather than without, sagittal crest; anterior border of tympanic bulla separated from foramen ovale by breadth of less than, rather than breadth of more than, 4 upper incisors (in- cluding 13 ) ; height of tympanic bulla not less than, rather than less than, distance from its anterior margin to foramen ovale; squamosal bone, between anterior margin of tympanic bulla and foramen ovale, ventrally concave rather than ventrally convex. Comparisons of the skulls with those of M. /. arthuri and M. f. primulina are made in the accounts of those subspecies. MUSTELA FRENATA 247 Remarks. — Excepting two young specimens from South Carolina in the Charleston Museum, no specimens of this race of large weasel seem to have been preserved until Arthur H. Howell, in the course of his study of the mammals of Alabama, procured specimens on which his name, olivacea, was based. Later, Francis Harper ob- tained three instructive specimens from Okefinokee Swamp. Really adequate material, for the localities represented, owes its preserva- tion to the alertness of Charles O. Handley, when he resided at Thomasville, Georgia, and to Hallie E. Fuller of Geneva, Talbot Co., Georgia. The distinctness of M. f. olivacea from M. f. peninsidae is not satis- factorily established due to inadequate material of peninsidae. Dif- ferences shown by the specimens seen indicate that, as compared with olivacea, peninsulae is larger, has transversely wider light- colored underparts which possess more yellow, and a larger skull with more inflated tympanic bullae. In each of these characters, olivacea is intermediate between noveboracensis on the north and peninsulae on the south. The question arises, tlierefore, whether the animals here recognized under the name olivacea really constitute a recognizable subspecies or instead are only representatives of a subspecies which reaches its extreme development in Florida. In the latter event, the name peninsidae would apply to all. Examina- tion of more material from Florida, especially from the southern half of Florida, will be necessary to answer this question. This large weasel of tlie southeastern United States is remarkably difiFerent from noveboracensis. Indeed, were it not for actual inter- grades such as the two from Fort Payne, Alabama, and York, South Carolina, which are described in the account of M. f. noveboracensis, and the six specimens from northwestern Alabama, which are re- ferred to olivacea, the systematist, I believe, would have little or no hesitancy in designating the two as distinct species, especially on the basis of differences to be seen in the skull. Not only are the two forms structurally more different than usually is the case but between two geographically, adjacent subspecies of the same species of mammal, but the belt where intergradation oc- curs appears to be narrow. Nevertheless, when material of the two races is laid out in geographic order, and examined in mass, certain features are seen to undergo gradual change as a person's eye travels from specimens from, say, the center of the range of nove- boracensis to specimens from southern localities adjoining the ter- ritory occupied by olivacea. One of these features subject to gradual change is the color of the underparts. Beginning at the 248 University of Kansas Publs., Mus. Nat. Hist. Adirondacks of New York where a large number of the specimens have white underparts, the underparts become more intensely yel- lowish southward through the range of noveboracensis into that of olivacea. Indeed, this progressive trend seems to continue right on southward through the range of olivacea into that of peninsidae. Turning in the opposite direction we find that the least width of the underparts decreases gradually northward toward the range of noveboracensis. There is, likewise, a decrease to the northward in length of the skull and relative, as well as actual, narrowing of the braincase and tympanic bullae. However, in least width of color of underparts and the mentioned cranial features, the trend stops relatively abruptly at the southern boundary of the geographic range of noveboracensis and does not continue on, northward, into the range of noveboracensis as is the case with the change in in- tensity of yellowness of the underparts. Two males, in the United States National Museum, Biological Surveys Collection, from near Leighton, Alabama, no. 178386 from the Tennessee River nine miles north [of Leighton?] and no. 180240 from La Grange Mountain, although clearly referable to olivacea on the basis of cranial characters, show some approach to nove- boracensis in lesser size of the skull and agree with noveboracensis in the narrowness of the color of the underparts. Also, these speci- mens, hke others from the northern part of the range of olivacea, for instance, no 31.227, Charleston Museum, from Mayesville, South Carolina, have the color of the underparts extended only part way out on the hind limb toward the foot. In specimens of olivacea from the southern part of its range the color of the underparts is extended onto the hind feet and this trend reaches its extreme in peninsidae, specimens of which have the feet and larger parts of the limbs marked with the light color of the underparts. An adult female, no. 32.32, Charleston Museum, although typical of olivacea in most respects, is nevertheless an intergrade. The teeth are as small as in some specimens of noveboracensis. The size of the skull is only slightly nearer that of olivacea than it is to that of noveboracensis. The proportions of the skull, however, are dis- tinctly those of olivacea. Five other specimens, from northwestern Alabama, namely two from eight miles north of Nauvoo, two from Shoal Creek, and one from White Creek, also show intergradation between noveborancesis and olivacea. The remarks concerning color and color pattern of the specimens from Leighton apply equally well to the five from northwestern Alabama. In cranial characters, no. 51658 from Shoal MUSTELA FRENATA 249 Creek is referable to oUvacea, as also is no. 51677 from the same place, providing it is a female rather than a male as sexed by the collector. No. 57146 from White Creek also is referable to oUvacea although the skull shows some approach to that of noveboracensis. Of the two males from near Nauvoo, no. 51652 is to me indistinguish- able from noveboracensis, but no. 51653 does have some characters of oUvacea, although on the whole, the latter, too, seems to be a little nearer noveboracensis than oUvacea. However, because the mean of these seven specimens from northwestern Alabama is nearer oUvacea than noveboracensis the former name may be ap- plied. Another specimen from "Souinlonie" Creek, Clark County, Mis- sissippi, has the coloration and rostral configuration of primuUna, narrow mastoidal breadth and smaller teeth of noveboracensis and skull of large size with "full" braincase as in oUvacea. No. 235364, U. S. Nat. Mus., from the Mobile River at the "L. and N. RR. Cross- ing", Mobile County, Alabama, although definitely oUvacea, shows approach to arthuri in that the dorsal outline of the skull is longi- tudinally more convex and the tympanic bullae are less inflated than in oUvacea and in that the color of the underparts is almost exactly as in the type specimen of arthuri. The young specimen labeled as from "Silver Springs", Florida, has large tympanic bullae (17 mm. long) and several characters that show its relationship to peninsulae as that race is now understood. Because the sex is un- known the identification as oUvacea is tentative and is made on the assumption that the specimen is a male. If it is instead a female, the animal is referable to peninsulae. An adult, female specimen in the Charleston Museum, no. 27.239.1, taken at St. Matthews, South Carolina, on December 8, 1927, contained four embryos which averaged 19 mm. in length and 47.75 centigrams in weight. Another adult female, in the Charles- ton Museum, no. 32.32, taken on February 21, 1932, at the same place, has prominent mammae, and the collector has noted that two were slightly active. Sixteen of twenty-nine adults examined show infestation of the frontal sinuses by parasites. However, in none is the malformation of the frontal region so great as frequently occurs in M. /. nove- boracensis. Specimens examined. — Total number, 52, arranged alphabetically by states and from north to south by counties in each state. Except as otherwise indi- cated specimens are in the University of California Museum of Vertebrate Zoology. 250 University of Kansas Publs., Mus. Nat. Hist. Alabama. Lawrence County: White Creek, 1; Little Sand Mt., Shoal Creek, 2. Winston County: VA mi. N Nauvoo, 1; 8 mi. N Nauvoo, 1. Lauderdale Co.: near Leighton, 9 mi. N Tennessee River, l\ Colbert Co.: Leighton, 1'. Autauga Co.: Autaugaville, 1\ Dale Co.: Midland City, 1\ Mobile Co.: Mobile River, 12 mi. NE Mobile, 1\ Florida. Alachua Co.: Gainesville, 4'. Marion Co.: "Silver Springs," 1. Georgia. Spalding County, 1. Lainar County, 1. Talbot County: south- west part of county, 1; Box Springs, near Geneva, 3; Upatoie Creek, 1 mi. SW Box Springs, 2; 3 mi. SE Geneva, 1; 4 mi. W Geneva, 1; 5 mi. W Geneva, 1; 2 mi. E Geneva, 1. Chattahoochee County, 2. Grady County: Beachton, 3'; locality no more definite than county, 4. Thomas County: Sinkola Planta- tion, 2; locality no more definite than county, 2. Charlton County: Vi mi. E Chesser's Island, Okefinokee Svv^amp, 1'. County in question: Billy's Island, Okefinokee Swamp, 1'; Okefinokee Swamp, V. Mississippi. Clark County: Souinlonie Creek, 1. South Carolina. DarZingfon Co.: Society Hill, 1'. Sumter County: Mayes- ville, 1'. Calhoun County: St. Matthews, 2*. Georgetown County: Sampit, r. Charleston County: Rantowles, 1'; 8 mi. N Charleston, 1*. Beaufort Co.: Yemassee, V. Mustek frenata peninsulae ( Rhoads ) Long-tailed Weasel Plates 16, 17 and 18 Putorius peninsulae Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1894:152, June 19, 1894; Bangs, Proc. Biol. Soc. Washington, 10:10, February 25, 1896. Mustela peninsulae, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, Mustela p. peninsulae, Bailey, Bailey Mus. and Library Nat. Hist., l(no. 5):1, December 1, 1930. Mustela frenata peninsulae. Hall, Carnegie Instit. Washington Publ. 473: 105, September 20, 1936. Type.— Female, young, part skull and skin; no. 8515, Acad. Nat. Sci. Philadelphia; Hudson's, Pasco County [14 miles north of Tarpon Springs], Florida; before 1895; obtained by W. S. Dickinson. The skull has been cut vertically in two at the plane of the glenoid fossae. These fossae and all the cranium posterior to them are missing. In addition to the part of the cranium anterior to the glenoid fossae, tlie lower jaws are preserved complete. The teeth all are present and entire. The prominent sutures on the rostrum and palate show the specimen to be young and its small size leaves but little doubt that the animal was a female. The fight facial markings are more extensive than in any of the referred specimens. In the type these fight facial markings consist of a median isolated spot immediately in front of the ears, a larger one on the nose, with an interrupted bar on each side extending posteroventrally in front of and anterior to the eye, a wider bar, on each side, extending anterodorsally between the ear and eye and finally an isolated spot at the anterior border of each ear. The skin is stuffed and in fair condition except that the vertebrae remain in the tail. Range. — Austral and probably Tropical life-zones of Florida south of latitude 29°. See figure 29 on page 221. 1. United States National Museum 4. Charleston Museum 2. Florida State Museum 5. American Mus. Nat. History 3. Cornell University MUSTELA FRENATA 251 Characters for ready recognition. — Differs from M. f. olivacea in coarser pelage and larger tympanic bullae. Description. — Size. — Male: No external measurements available. Female: The type a young animal and no. 2379, an adult from Tarpon Springs, measure respectively as follows: Total length, 375, 378; length of tail, 100, 130; lengtli of hind foot, 40, 44.5. Externals. — As described in Mustela frenata noveboracensis except that hairiness of foot-soles as shown in figure 20. Color. — Upper parts (in winter) near tone 3 of Burnt Umber of Oberthur and Dauthenay, pi. 304. Dark spot at each angle of mouth present or absent. Tip of tail black. Underparts Reed Yellow except on chin and usually on legs where white. Upper lips white entirely around. Upper parts of uni- form color. Color of underparts extends distally on legs over both sides of feet and on front legs over wrists. Proximal part of tail slightly hghter below than above. Least width of color of underparts, in seven specimens, averaging 41 (extremes 31-52) per cent of greatest width of color of upper parts. Black tip of tail, in each of two females, 45 mm. long; thus slightly longer than hind foot and amounting to 36 per cent of length of tail- vertebrae. The spot at the angle of the mouth is absent in four of the ten specimens and is present on botli sides in the other six. Skull and teeth. — Male (based on an adult from Apopka and the anterior part of an adult from Enterprise): See measurements and plates 16-18. As described in Mustela frenata olivacea except that: Weight, 7.0 grams; basilar length, 49.8. Female (based on an adult from Tarpon Springs, Florida): See measure- ments. As described in Mustela frenata olivacea except that: Weight, 4.7 grams; basilar length, 44.2; zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla. In comparison with M. /. olivacea^ the insuflBcient material of M. /. penimuJae suggests that its skull averages larger and has relatively as well as actually larger and more inflated tympanic bullae. Remarks. — The first published mention of this weasel seems to have been the original description which appeared in 1894. This description was based on a single specimen sent to Samuel N. Rhoads by W. S. Dickinson, who, in the following year, procured another specimen at Tarpon Springs. So far as known only eight other specimens, as listed under "Specimens examined", have found their way into collections of study specimens. H. H. Bailey (1930:1) credits the range of this subspecies as extending south "to the shores of Florida Bay and the Gulf of Mexico, where ever high ground occurs." Evidence of intergradation between M. f. peninsulae and M. f. olivacea is provided by specimens of olivacea from Gainesville, 252 University of Kansas Publs., Mus. Nat. Hist. Florida, and the Okefinokee Swamp, Georgia. These specimens, on the average, have the color of the underparts wider, the skull larger, and the tympanic bullae relatively larger than do specimens of olivacea from farther north. In these features, approach to M. f. peninsulae is shown. Light facial markings occur in this subspecies. They are similar to those possessed by weasels which occur at the same latitude and under corresponding climatic conditions on the Pacific Coast. The type specimen and one from Tarpon Springs have white facial markings. Two of the three specimens from Apopka also show white facial markings, although in reduced amount. One of the four specimens of M. /. olivacea from Gainesville, Florida, has well- developed light ( white ) facial markings. Also of the four specimens of M. /. olivacea examined from Okefinokee Swamp, Georgia, one has prominent white facial markings. However, in it the pattern is so unusual as to suggest that it is an instance of partial albinism rather than an outcropping of a racial tendency, or a pattern of coloration induced by climatic factors. None of the eight available skulls show any infestation of the frontal sinuses by parasites. Specimens examined. — Total number, 10, arranged by counties from west to east. Florida. Pasco County: Hudson's, 1'. Pinellas Co.: Tarpon Springs, l^ Hernando Countij, 1'. Polk Co.: Aubumdale, 1"; no locality more definite than county, 1". Orange County: Apopka, 3'. Volusia Co.: Enterprise, 1*. Seminole Co.: Osceola, 1°. Mustela frenata spadix (Bangs) Long-tailed Weasel Plates 16, 17, 18, 31, 32 and 33 Putorius longicauda spadix Bangs, Proc. Biol. Soc. Washington, 10:8, Feb- ruary 25, 1896; Merriam, N. Amer. Fauna, 11:21, figs. 10, 11, June 30, 1896; Cory, Mamm. Illinois and Wisconsin, p. 374, 1912. Mustela longicauda spadix. Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912; Bailey, Journ. Mamm., 10:156, May 9, 1929. Mustela longicauda, Johnson, Journ. Mamm., 11:439, November 11, 1930. Mustela noveboracensis, Murie, Journ. Mamm., 16:321, November 15, 1935. Mustela frenata spadix. Hall, Carnegie Instit. Washington Publ. 473:105, November 20, 1936. Type. — Male, young, skull and skin; no. 3265/1786, Amer. Mus. Nat. Hist.; Fort Snelling, Hennepin County, Minnesota; June 25, 1889; obtained by Ed- gar A. Mearns; original no. 812. The skull is complete although there are fractures on the top of the brain- 1. Acad. Nat. Sciences of Philadephia 4. Field Museum of Natural History 2. United States National Museum 5. American Mus. Nat. History 3. Florida State Museum MUSTELA FRENATA 233 case, on the right side of the braincase and at the middle of the right zygo- matic arch. The teeth all are present and entire. The skin, although overstiifiFed, is complete, well preserved, and in summer pelage. Range. — Upper Austral and Transition life-zones of Minnesota, northern and western Iowa, southeastern North Dakota, eastern part of South Dakota, and northeastern Nebraska. See figure 29 on page 221. Characters for ready recognition. — Differs from M. f. noveboracensis and M. /. primulina in that specimens of all ages have least width of color of underparts amounting to more than 41 per cent of greatest width of color of upper parts, and have light color of underparts extended onto hind foot rather than stopped short of ankle; adults with hind feet more than 50 in males and 40 in females; orbitonasal length more than 15.5 in males and 13.5 in females; length of tooth-rows more than 18.0 in males and 15.7 in females; mastoid breadth more than 25.5 in males and 22.0 in females. From M. f. longicauda by color darker than near (h) Clay Color, in males by a flattened occiput in which the depth of the skull, exclusive of the sagittal crest and taken at the anterior border of the basioccipital, amounts to less than 58 per cent of the mastoid breadth. Description. — Size. — Male: Three adults from Elk River, Minnesota, yield average and extreme measurements as follows: Total length, 458(444-467); length of tail, 154 (140-165); length of hind foot, 55(52-59). Tail averages 51 per cent as long as head and body. Length of hind foot averages more than basal length. Corresponding measurements of tliree subadults from Madison, Minnesota, are as follows: 453(438-469); 157(152-165); 50 (47- 51 ). Tail averages 53 per cent as long as head and body. Female: Tliree adults from Elk River, Minnesota, yield average and ex- treme measurements as follows: Total length, 387(380-391); length of tail, 131(121-138); length of hind foot, 44(43-46). Tail averages 51 per cent as long as head and body. Length of hind foot more or less than (approxi- mately equal to) basal length. Corresponding measurements of two adults and one subadult from Madison, Minnesota, are as follows: 385(379-396); 137(119-159); 42(38-44). Tail averages 55 per cent as long as head and body. The average differences in external measurements of the two sexes from Elk River, are: Total length, 71; length of tail, 23; length of hind foot, 11. At Madison, corresponding differences are 68, 20, and 8. Two adult females from Elk River, Minnesota, weigh 205 and 210 grams. Externals. — Longest facial vibrissae black, brown, or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot- soles ( in summer pelage ) as shown in figure 19. Color. — Winter pelage all white except tip of tail. In southern part of range sometimes assumes a brown winter coat. Summer pelage with upper parts ranging from near (16') Cinnamon Brown to Vandyke Brown. Chin and upper hps white. Remainder of underparts ranging from near (a) Olive Ocher to Ochraceous Buff and Pale Orange Yellow. Tip of tail at all times black. Upper parts of uniform color except for occasional slight darkening of nose. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and ankles, on medial sides of hind limbs 254 University of Kansas Publs., Mus. Nat. Hist. to ankle, over antiplantar faces of toes and distomedial fourth of each tarsiis, and over proximal fifth to third of under side of tail. Least width of color of underparts averaging (in 3 specimens from Elk River) 54(47-59) per cent of greatest width of color of upper parts. Black tip of tail averaging same length as hind foot and 28 per cent of length of tail-vertebrae. Save for Uie greater width of the light-colored underparts and relatively short black tip of the tail, both featm-es of M. f. longicauda, spadix is variously intermediate, depending on locafity, as between noveboracensis and longicauda. Skull and teeth. — Male (based on 3 adults from Elk River, Minn.): See measurements and plates 16-18. As described in Mustela frenata longicauda except that: Weight, 5.6(5.0-6.5); basilar length, 49.0(48.7-49.2); zygomatic breadth sometimes less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more or less (about equal to) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadtli more or less tlian distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as widtli of 4 to 5 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; lengtli of typanic bulla less than lengdi of rostrum; anterior margin of masseteric fossa below talonid of ml. Female ( based on 4 adults from Elk River, Minn. ) : See measurements and plates 31-33. As described in Mustela frenata longicauda except that: Weight, 3.5 (3.3-4.0) grams; basilar length, 42.9 (42.3-43.2); least widdi of palate more or less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors. The skull of die female averages 33 per cent fighter than that of the male. Skulls of adult males of spadix from Elk River, Minnesota, as compared with those of longicauda from Alberta, are larger in every part measured. Relative to tlie basilar length these skulls of spadix are broader across the mastoid region, narrower across the zygomata, deeper through the plane of the postorbital processes, shallower through the braincase and have relatively shorter tym- panic bullae. Whereas the tympanic bullae of longicauda are, on the average, approximately as long as the rostrum (orbitonasal length), in spadix the rostrum is longer than the bulla. Viewed posteriorly, the braincase of spadix is seen to be much shallower and wider than that of longicauda. Indeed, the depth of the brain- case, measured at the anterior end of the basioccipital, amounts to only 56 per cent of tlie mastoid breadth in spadix as against 61 per cent in longicauda. The longer, waistlike, postorbital constriction, relatively smaller braincase, and especially the relatively narrower zygomatic expanse in spadix imparts to its skull a more slender ap- pearance than has the skull of longicauda. These differences are not shown by the skulls of females. To be sure, spadix, in most of its cranial measurements, averages slightly larger, has a relatively MUSTELA FRENATA 255 shallower braincase and is relatively deeper through the postorbital processes, but these diflFerences are so slight that inclusion of one more specimen, of slightly different proportions, in the average might cause the average measurements to read as they do in longi- catida. Compared with noveboracensis, from Massachusetts, adult skulls of spadix, taking sex into account, are larger in every part measured and are relatively as well as actually wider and deeper throughout. Also, in spadix: Sagittal and lambdoidal crests higher, especially in females; anterior margin of tympanic bulla projecting up sharply from squamosal; occiput more flattened in posterior view; tooth- rows relatively and actually longer but orbitonasal length relatively shorter; postorbital processes more robust; zygomatic arches widely bowed outward rather than evenly rounded; canines larger; squa- mosal less swollen ventrally, especially in females. Between noveboracensis and spadix, the differential cranial characters are greater in number and degree between females than between males. Comparison of the skull with that of M. f. primulina is made in dis- cussion of that subspecies. RemarJis. — Edgar A. Mearns in 1889 and the early nineties took several specimens of this weasel and it was principally on tliese that Bangs in 1896 (p. 8) based his description. The best material, how- ever, is that from Elk River, Minnesota, collected in later years by Bernard Bailey, and supplemented by one specimen taken in 1885 by Vernon Bailey and another by his sister Anna Bailey in 1891 at the same place. Mustela frenata spadix has just one structural feature of a "unique" kind which serves to differentiate it from the geograph- ically adjoining subspecies. This feature is large size. The other diagnostic characters ascribed to spadix are of an intermediate sort — intermediate as between two extremes, one found to the westward in longicauda and the other to the eastward in noveboracensis. For example, the dark-colored upper parts are merely darker than in longicauda and merely lighter than in noveboracensis. The color is not "different"; it is only "intermediate". Furthermore, each of tlie characters ascribed to spadix, including large size itself, undergoes change from one part of its geographic range to another; the char- acters are not constant over a wide area. Indeed, excepting the large size which remains relatively uniform over the northern two- thirds of the range, no two locahties have been found from which the specimens can be said really to agree in characters. 256 University of Kansas Publs., Mus. Nat. Hist. By way of illustration, the coloration of the upper parts may be cited. Near the range of noveboracensis the average coloration of individuals from one locality is only a little Hghter than in novebora- censis. Farther westward the average coloration is a little lighter and farther west\vard yet, toward the range of the extremely light colored longicaitda, the average coloration is lighter still. Although all these animals are darker than longicaitda and lighter than noveboracensis, those from the three places do not agree among themselves. Because of the lack of more than one character of a "unique" kind and because of the inconstancy, geographically, of other characters, and for that matter, lack of constancy geograph- ically in combination of characters, the writer regards spadix as a barely recognizable subspecies. Examination of the specimens of spadix shows that the individual variation in a single species is greater in a region of intergradation than it is some distance inside the borders of the geographic range of a well-marked subspecies. This is illustrated by three specimens of M. /. spadix in fresh summer pelage from the single locality, Elk River, Minnesota. In these, the color of the upper parts varies from a little darker than Cinnamon Brown to Vandyke Brown. At any one locality well within the range of longicauda, or noveboracensis, there is nowhere nearly so much variation in color, even in much larger series of specimens. Study of the specimens here assigned to spadix reveals that some features regarded as of diagnostic value for one or the other of the two races, longicauda and noveboracensis, behave differently. For example, the dark coloration of the upper parts, which is character- istic of noveboracensis, manifests itself far westward within the range of spadix whereas the wider extent of the light-colored underparts, which is characteristic of longicauda, and the Olive Ocher, rather than Pale Orange Yellow, color of these underparts, are seen in varying degree all the way across the range of spadix. Thus, these animals are colored above like noveboracensis and be- low like longicauda, but not vice versa. In these animals, then, the longicauda type of underparts is dominant, in one sense of the word, over the noveboracensis type of underparts, and the noveboracensis type of upper parts is dominant over the longicauda type of upper parts. Each of these features is subject to actual intergradation and does not always behave as a "unit character", that is to say, one which is either present or absent. However, the noveboracensis type of upper parts is carried much farther west before being di- luted than is the noveboracensis type of underparts. Indeed, within MUSTELA FRENATA 257 the range of novcboracensis itself, the broad extent of tlie longicauda type of underparts is manifest. This is, of course, near the western margin of the range of novehoracensis. The large size of males of spadix, as exemplified by specimens from Elk River (see measurements on p. 421), seems to be retained across the northern part of the range here assigned to the subspecies. This larger size than is found in longicauda from Alberta, is shown also by some specimens from eastern North Dakota which are as- signed to longicauda. However, the average of these Dakotan specimens, all characters considered, is nearer to my concept of longicauda. Inspection of the cranial measurements of spadix shows also that in addition to its large size it is distinguishable from any one of the geographically adjoining races by its relatively (to basilar length) greater, as well as actually greater, mastoidal breadth. This might be included widi size as a unique character distinguishing spadix from longicauda and novehoracensis. However, it is not clear whether or not this greater mastoidal breadth is more than a function of the large size. Excepting the greater mastoidal breadth and generally larger size of the skull, the cranial features distinguishing males of spadix from longicauda are features in which spadix shows approach to nove- horacensis. This is true, in spadix, of the relatively longer ( in com- parison with longicauda) rostrum, relatively lesser zygomatic breadth, relatively shallower braincase measured at the anterior end of the basioccipital, and relatively deeper skull as measured at the posterior borders of the last upper molars. This same approach to novehoracensis already has been pointed out with respect to color of the upper parts and is evident also in the relative shortness of the tail which averages only 51 per cent of the length of the head and body rather than 55 per cent as in longicauda. Because the longicauda type of animal previously has been re- garded as specifically distinct from the novehoracensis type of animal, comment is offered below on selected specimens, referred to spadix, which are regarded as intergrades with novehoracensis or with other subspecies. No. 8722, Univ. Wisconsin, adult male, in the white winter coat, from north central Itasca County, Minnesota, obviously has char- acters of M. /. spadix or longicauda that occur to the west and M. f. novehoracensis of the east. Selected outstanding characters of longicauda are its long tail, anteriorly truncate tympanic bullae and large teeth. Characters indicating its affinities with novehoracensis 9—3758 258 University of Kansas Publs., Mus. Nat. Hist. are smaller size of skull, general narrowness of skull, and relatively low tympanic bullae. The skull is intermediate as regards several individual structural features. For example, although long and narrow and in this feature more nearly approaching noveboracensis, the skull is wider than usual in that subspecies and thus approaches that of longicauda or spadix. The hind foot, in the dried state, measures 47 millimeters. This large hind foot, obviously long tail (the specimen lacks external measurements), and anteriorly trun- cate bullae constitute basis for here referring the specimen to spadix. However, the seemingly small size of the body and the nar- row skull clearly show relationship to noveboracensis. Specimens, referred to spadix, from northern Iowa, are instruc- tive as showing what happens where the ranges of noveboracensis, primulina, spadix, and perhaps longicauda, meet. No. 47167, Univ. Mich. Mus. Zool., a nearly adult female, taken on November 22, 1915, at Island, Clay County, and in process of assuming a brown winter pelage, retains enough of the dark summer pelage to show that the color was slightly lighter than average for spadix. The color pattern, white lips, and extension of light color of the under- parts onto the feet, agrees with spadix or longicauda as does also the long tooth-row. The over-all length of the skull is intermediate between that of spadix and primulina. The proportions of the an- terior part of the skull and of the tympanic bullae resemble those found in primulina. A subadult male skull only, no. 123846, Ameri- can Museum of Natural History, from Webb, Clay County, shows approach to primulina in the narrowness of the rostrum. A young male from Ruthven, Iowa, no. 48340, Univ. Michigan, has a large skull approaching in size that of spadix, has the longi- cauda-spadix type of light-colored underparts and color pattern, and is slightly darker above than true longicauda. Another sub- adult male in the white winter coat from Palo Alto County, no. 35756, Univ. Michigan, has a large skull, which shows approach to primulina in its narrowness anteriorly and in some other features. Although the tail is of moderate length, the body is large as in spadix or longicauda, and the length of the hind foot suggests spadix or longicauda. A subadult male, no. 425a, Iowa State College, from Manson, Iowa, in brown winter pelage, agrees with primulina in the restric- tion of the area of the light color of the underparts and in less ex- panded zygomatic arches. The teeth are intermediate in size be- tween those of noveboracensis and primulina on the one hand and those of spadix and longicauda on the other. In other respects it agrees with, or is more nearly like, spadix. MUSTELA FRENATA 259 An adult female, no. 426a, Iowa State College, from Bamum, in the brown winter coat, agrees with primulina except that the or- bitonasal length of the skull is more as in spadix and the presence of some light color on the lower part of tlie hind legs suggests spadix. The skull only, no. 440a, Iowa State College, labeled merely Webster County, Iowa, is almost a dupHcate of no. 426a. A subadult male, no. 427a, Iowa State College, from Moorland, Iowa, only about six miles southeast of Bamum, likewise is indis- tinguishable from primulina except for having a white winter coat and in being relatively broad in the mastoidal region. Neverthe- less, both of these animals are here referred to spadix because the average of specimens from this general area is nearer that of spadix. No. 497a, Iowa State College, an adult female in white winter pelage, from Ames, approaches primulina in the narrow rostrum and smaller teeth but otherwise approaches or even agrees with spadix. Two adult males, without external measurements, from Pilot Mound, Iowa, have skulls quite like males of longicauda from Al- berta. The only approach noted to eastern forms is the restricted color of the underparts on no. 2856, Coe College, which has a brown winter coat. The color of the underparts is not extended so far out on the feet as in longicauda. Also the tympanic bullae of this specimen are a trifle narrower. The other male, no. 2652, is in the white winter coat. The one female from the same place, no. 2660, Coe College, in brown winter pelage, has a skull notably unlike that of longicauda or spadix; the skull is narrower and prac- tically indistinguishable from that of the largest female skull of primulina available from Lawrence, Kansas, save that the tooth- row is much longer. The color pattern also agrees with that of primulina or novehoracensis in that the color of the underparts extends only as far as the knee on the hind legs and is narrow on the belly. Nevertheless, another adult female, no. 120a from Amaqua Township, some 6 miles southwest of Pilot Mound, is in aU respects typical of spadix. This is the more remarkable be- cause another comparable specimen from less than 20 miles to the southwest in Worth Township is equally typical of primulina. Two young females from Chester, Iowa, nos. 2656 and 2874/ 2873, Coe College, have skulls larger than those of corresponding age of primulina or novehoracensis. The color is as in spadix. The color pattern of the underparts also is as in spadix or longicauda except that the width of the area of light color on the belly is re- stricted somewhat although not so much as in novehoracensis or 260 University of Kansas Publs., Mus. Nat. Hist. primulina. Of four males from the same place, also in the collec- tion of Coe College, no. A2874 is a white skin only and does not provide diagnostic characters. The three other males, each in summer pelage, are marked and colored as are the two females from the same place except that male no. 2861 has the color of the underparts so much attenuated on the hind legs that it barely, un- interruptedly, extends to the feet. No. 2658 is young, or perhaps barely subadult. The skull is large and referable to spadix. The two adults, nos. 2861 and 2657, differ cranially from typical (Elk River, Minn.) spadix only in being slightly narrower across the mastoids and in having the bullae a little narrower. In these de- partures they show some approach to primulina and to novebora- censis. Another male, subadult, no. 2867, Coe College, from De- corah, which has acquired half of the white winter coat, agrees with the males from Chester except that the preorbital part of the skull is shortened much as in some specimens of primuliria. From Lansing, in extreme northeastern Iowa, a large subadult male, no. 2864, Coe College, of 453 mm. in total length and half through with acquiring the white winter coat, agrees with the males previously described from Chester except in having the palate nar- rower as in noveboracensis. The adult female available from Lansing, no. 2863/2862, Coe College, in white winter pelage except for the top of the head, although a large skin, has a skull smaller than that of any spadix or longicauda and of about the same size as that of no. 3838, Univ. Kansas Mus. Nat. Hist., of primulina, from Lawrence, Kansas, except that the skull of no. 2863/2862 is much narrower across the mastoids. This specimen, then, shows approach to noveboracensis in narrowness of the mastoidal region, to primulina in other respects and to spadix. Many of these instructive specimens from Iowa, made available to the present writer by Mr. W. F. Kubichek, were brought to- gether at the Coe College Museum by the late B. H. Bailey. Most of them were obtained from trappers who did not supply the con- ventional external measurements taken in the flesh. Even though these are lacking, the specimens clearly show that actual inter- gradation occurs where the ranges of M. /. longicauda, spadix, noveboracensis and primulina meet. The dark color of the upper parts, restriction of the color of the underparts on the ankles with the result that the color reaches the toes in interrupted fashion, and large skull, of no. 18912 of the Museum of the University of South Dakota, from Roberts County, South Dakota, clearly place this specimen with spadix, rather than MUSTELA FRENATA 261 with longicauda. Likewise, male, subadult, no. 11376, Univ. South Dakota, from Clay County, South Dakota, is referable to spadix. Although without external measurements, the specimen obviously is large. The patch of summer pelage on its head and neck is darker than the summer pelage of longicauda, and the orbitonasal length is greater than the length of the tympanic bullae; all these features are characters of spadix. The adult male from Fort Sisse- ton, South Dakota, no. 188407, United States National Museum, figured by Merriam (1896, p. 20, figs. 7-9), is almost exactly inter- mediate between longicauda and spadix, although here referred to the latter. Five specimens, nos. 147375, 147432, 147762, 148720 and 148721, U. S. Nat. Mus., including 3 skulls only from Beemer, Cuming County, Nebraska, are intergrades between M. /. longicauda, M. f. primulina and M. /. spadix. One skin is in white winter pelage and the other, a female, is in summer pelage which in coloration and colpr pattern agrees with that of spadix. External measure- ments of the male agree with those of longicauda. Measurements of the female agree with those of spadix except that the tail is shorter as in primulina. The skulls are as long as in longicauda but are more slender than in either lotigicauda or spadix although nearer the latter in this respect. In dorsal aspect, the skulls espe- cially posteriorly to the orbital region, resemble primulina. All points considered, the animals seem best referred to s])adix. Although the degree of development of certain morphological features has been settled upon as indicative of the race spadix, some doubt remains as to where the western boundary of its range should be shown. This results from the fact that color has been taken into account as one diagnostic feature and this feature is lacking in the white winter specimens which, from the following places, are all that are available: Kittson County, Minnesota; Moorhead, Minnesota; Casselton and Valley City in North Dakota; Araiour, South Dakota and Clay County, South Dakota. In sum- mary, more specimens in the summer coat will be required to establish definitely the boundary between the ranges of longicauda and spadix. Surber (1932:49) has referred to additional specimens of this weasel in the University of Minnesota Museum as from Winona, Hennepin and Isanti counties of that state. At Elk River, Minnesota, B. Bailey (1929:156) found this species to be about half as abundant as Mustela cicognanii and that it is "more often found in the open timber and about the dry ridges 262 University of Kansas Publs., Mus. Nat. Hist. and fields". Of seventeen adult or subadult skulls of this race from Minnesota, ten have obvious marks of infestation of the frontal sinuses. In no skull, however, has the infestation resulted in so much malformation, as occurs in noveboracensis. Specimens examined. — Total number, 76, arranged alphabetically by states and from north to south by counties in each state. Iowa. Lijon County: Granite, 1''. Howard County. Chester, 6\ Winne- shiek County: Decorah, 1'; 8 mi. NE Ossian, l\ Allamakee County: Lansing, 2\ Clay County: Island, 1=; Webb, 1'. Palo Alto County: Ruthven, 1'; no locahty more definite than county, 1". Callioun County: Manson, V^. Web- ster County: Bamum, 1'*; Moorland, 1"; no locality more definite than county, V\ Boone County: Pilot Mound, 3'; Amaqua Township, Sec. 19, 1". Story County: Ames, 1". Minnesota. Kittson County, V. Roseau Co.: 2)2 mi. SW Roseau, Jadis Township, 1^". Itasca County: T61N, R26W, 1*. Clay Co.: Mcorhead, -2\ Atkin Co.: Atkin, 1'. Otter Tail County: Lake Lizzie, 1°; Parkers Prairie, 1', Grant Co.: 3 mi. NW Barrett, 1=. Benton (now Mille Lacs?) County: Prince- ton, r. Sherburne Co.: Elk River, 13 {&, A'\ 3', 1"). Hennepin Co.: Fort Snelling, 6 (5', 1'). Carver? Co.: Chaska, 1''. Lac qui Parle Co.: Madison, 5 (3', 2''); no locahty more definite than county, 2 ( 1", 1"). Yellow Medicine Co.: Wood Lake, l^ Blue Earth County: Rapidan, 1". County in question: Moores Lake, l^ Nebraska. Cuming Co.: Beemer, 5*. North Dakota. Cass Co.: Fargo, 1'; Casselton, 1'. Dickey Co.: Oakes, 1'. South Dakota. Roberts County, 1". Marshall Co.: Fort Sisseton, 1'. Douglas Co.: Annour, 1". Clay County, 1". Mustela frenata longicauda Bonaparte Long-tailed Weasel Plates 16, 17, 18, 31, 32 and 33 Mustela longicauda. Bonaparte, Charlesworth's Mag. Nat. Hist., 2:38, 1858. Putorius longicauda, Baird, Mamm. N. Amer., p. 169, 1858; Coues, Fur- bearing animals, p. 136, 1877; Bangs, Proc. Biol. Soc. Washington, 10:7, figs. 1, la of pis. 1, 2 and 3, February 25, 1896; Merriam, N. Amer. Fauna, 11:19, pi. 3, figs. 3, 3a, 4, 4a, pi. 5, figs. 1, la, text figs. 7-9, June 30, 1896. Mustela longicauda longicauda, Bailey, N. Amer. Fauna, 49:166, January 8, 1927. Mustela frenata longicauda. Hall, Carnegie Instit. Washington Publ. 473: 105, November 20, 1936; Hall, Canadian Field-Nat., 52:108, October, 1938. Mustela frenata, Sowls, Joum. Mamm., 29:126, May 14, 1948. Type. — Possibly not in existence. No. 43.3.3.3 [from Carlton House, Sas- katchewan] in the British Museum of Natural History has been regarded by 1. Coe College 10. Collection of Alex Walker 2. Museum of Zoology, Univ. Michigan 11. Mus. Vert. ZooL, Univ. California 3. American Mus. Nat. History 12. Field Museum of Natural History 4. University of Wisconsin 13. Acad. Nat. Sciences of Philadelphia 5. Carnegie Museum 14. Leland Stanford Junior University 6. Collection of W. E. Snyder 13. Museum of Comparative Zoology 7. Colorado Museum of Natural History 16. University of South Dakota 8. United States National Museum 17. Illinois Natural History Survey 9. Donald R. Dickey Collection 18. Iowa State College MUSTELA FRENATA 263 several zoologists as the type. It is a subadult female, skull and sldn, from North America. See the account of M. erminea cicognanii for reasons for and reasons against regarding this specimen as the holotype. No. 43.3.3.3 from the collection of Dr. John Richardson is in the white winter coat and now ( Sept. 24, 1937 ) is prepared as a study skin. Evidences of its once having been mounted are: holes in the soles of the hind feet for supporting-wires, large straight wire in the tail, folds in the skin of the now backward-projecting hind feet, and unevenness of the skin on the back resulting from straightening out the specimen. The tip of the tail and some skin from the middle of the belly are missing. Otherwise the skin is intact. The skull is that of an animal in its first year, lacks the zygomatic arch on each side, but otherwise is complete and unbroken. The teeth all are present and entire except that p2 on the right side is missing from its alveolus. Range. — Transition and Upper Sonoran life-zones of the Great Plains, south- ward from central Alberta, Saskatchewan and southern Manitoba through east- em Montana, the Dakotas and Nebraska into southeastern Wyoming, north- eastern Colorado and western Kansas. See figure 29 on page 221. Characters for ready recognition. — DiflFers from M. /. primulina in near ( h ) Clay Color rather than Brussels Brown of upper parts, least viddth of color of underparts more than 40 per cent of greatest width of color of upper parts, color of underparts extended onto hind foot rather than stopped short of ankle, zygomatic breadth more than 28.8 in adult males and more than 24.1 in adult females; from M. f. spadix in fighter color being near ( h ) Clay Color, in males by deeper occiput in which the depth of the skull, exclusive of the sagittal crest and taken at the anterior border of the basioccipital amounts to more than 59 per cent of the mastoid breadth; from M. f. oribasus in near (h) Clay Color rather than near ( 14n ) Brussels Brown color of the upper parts and in males by deeper occiput in which the depth of the skull, exclusive of the sagittal crest and taken at the anterior border of the basioccipital, amounts to more than 59 per cent of the mastoid breadth; from M. f. alleni in larger size, adult males having a total length of more than 400 millimeters, hind foot more than 45, basilar length more than 43.5, and females having a total length of more than 375 and basilar length not less than 40.0; from M. f. nevadensis in near (h) Clay Color rather than near ( 14n to 1 ) Brussels Brown of upper parts, basilar length more than 40 in females and averaging more than 45 in males; from M. /. neomexicana by near (h) Clay Color rather than Buckthorn Brown color of upper parts, absence of white and Argus Brown facial markings, and length of tooth-rows amounting to more than 37 per cent of basilar length. Description. — Size. — Male: Five adults from Alberta yield average and extreme measurements as follows: Total length, 438 (418-473); length of tail, 158 (140-193); length of hind foot, 50 (46-54). Tail averages 56 per cent as long as head and body. Length of hind foot averaging more than basal length. Corresponding measurements of five adults and subadults from North Dakota are as follows: 465 (445-516); 164 (150-179); 51 (50-54). Tail averages 55 per cent as long as head and body. Female: Six adults (Alberta, 4; Saskatchewan, 1; Manitoba, 1) yield aver- age and extreme measurements as follows: Total length, 401 (383-425); length of tail, 145 (141-159); length of hind foot, 43 (41-44). Tail averages 57 per cent as long as head and body. Length of hind foot more or less than (approximately equal to) basal length. 264 University of Kansas Publs., Mus. Nat. Hist. The average differences in external measurements of the two sexes are: Total length, 37; length of tail, 13; length of hind foot, 7. General compari- sons indicate that the Alberta-taken males may not attain so large a size as those from some other areas. Thus the differences in external measurements might be some greater elsewhere, say, in North Dakota. Externals. — Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and e.xtending to apical pad of fifth digit; hairiness of foot-soles ( in summer pelage ) only slightly greater than shown in figure 20. Color. — Winter pelage all white except tip of tail. Summer pelage with upper parts near (h) Clay Color or near tone 3 and 4 of Snuff Brown of Oberthiir and Dauthenay, pi. 303. Chin and upper lips white. Remainder of underparts ranging from near (a) Olive Ocher to near (16') Ochraceous Buff. Upper parts of uniform color except for occasional darkening of head in front of ears. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind limbs to ankles over antiplantar faces of toes and distomedial third of each tarsus, and over proximal fourth to third of under side of tail. Least width of color of underparts averaging, in a series of 10 males from Alberta, 58 (45-60) per cent of greatest width of color of upper parts. Corresponding figures for 10 females from the same place are 57 (50-74). Black tip of tail in same series of males, most of which are in full summer pelage, averaging 43 ( 35-60 ) mm. long. Thus, averaging shorter than hind foot and 27 per cent of length of tail-vertebrae. As compared with M. /. neomexicana, longicauda lacks the white facial markings, black ears, black forehead and nose, but otherwise is similarly colored. As compared with M. f. nevadensis, M. f. orihasus and M. /. spadix, each of color pattern similar to longicauda, selected differences of longicauda are its much lighter color, especially of the upper parts, with less conspicuous darkening on the nose. From M. f. primulina, longicauda differs in lighter color of upper parts, reddish rather than yellowish imderparts, and light- rather than dark-colored hind feet. Skull and teeth. — Male ( based on 5 adults from Alberta ) : See measure- ments and plates 16-18; weight, 4.7 (4.6-4.9) grams; basilar length, 46.0 (44.7-46.8); zygomatic breadth more than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tj'mpanic bulla; mastoid breadth more than postpalatal length; postorbital breadth less than length of upper premolars and more than width of basioccipi- tal measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum more or less (usually less) than length of tympanic bulla; least width of palate less than greatest length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including 13) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth- row and longer or shorter than rostrum; anterior margin of masseteric fossa below talonid of ml or anterior half of m2. Female (based on 5 adults: Alberta, 3; N. D., 1; Sask., 1.): See measure- ments and plates 31-33; weight, 3.1 (2.8-3.5) grams; basilar length, 42.3 MUSTELA FRENATA 265 (40.0-43.7); zygomatic breadth more or less (approximately equal to) than distance between condylar foramen and Ml or that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate not more than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including 13) upper incisors; height of tympanic bulla not less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-rows and longer or shorter than rostrum. The skull of the female averages 34 per cent Hghter than that of the male. Comparisons of the skull with those of M. f. primulina, M. f. spadix, M. f. oribasus, M. f. alleni, M. f. nevadensis, and M. f. neomexicana are made in accounts of those subspecies. Remarks. — Richardson's (1829:47) account on which Bonaparte may be said to have based his name, records measurements in inches and hues which I transpose into millimeters as follows: Total length, 440 mm.; length of head and body, 305; length of tail (vertebrae), 135; length of tail (including fur), 164 mm. Specimen no. 43.3.3.3 in the British Museum, which has by some persons been regarded as the type, yields measurements as follows: Total length, 408 (which allows for 15 mm. loss of the fleshy part of the end of the tail); length of head and body, 272; length of tail (vertebrae), 136 (r= 121 -f 15); length of tail (including fur), 162 (142 + 20 mm. that appears to have been lost). Richardson's specimen would appear to have been of unusual proportions and to have been larger than no. 43.3.3.3. Some reasons for and reasons against regarding this specimen as the holotype are given in the account of M. erminea cicognaiiii. The name longicauda was applied to practically all long-tailed weasels of the western United States at one time but as one after another of the geographic variants in the mountainous regions were designated as separable, the name longicauda came to be restricted to the light-colored, relatively large, animal of the Great Plains. The intergradation of longicauda with spadix and oribasus has been commented on in the discussions of those subspecies. The larger size and darker color of specimens referred to longicauda from Devils Lake and Grafton, North Dakota, are features in- dicative of intergradation there with spadix. Two young females from Waterton Lake Park, Alberta, by their darker than average color, suggest intergradation with oribasus, as, for that matter, does the specimen from Waterton Lake [= Chief Mountain Lake, 266 University of Kansas Publs., Mus. Nat. Hist. in Montana] itself, which, however, is even darker than the two specimens taken on the Canadian side of the hne and hence is referred to orihasus. An adult female, no. 175586, U. S. Nat. Mus., from Moose Pass, Alberta, examined after the above was written, is larger than any other female seen of longicauda and in this re- spect may show approach to orihasus, which in the northern part of its range is of large size as judged by males from the Bowron Lake region. One male, no. 8564, Nat. Mus. Canada, from Max Lake, Turtle Mountain, Manitoba, presents puzzling characters. The external measurements of 465, 170, and 57, are in keeping with the great length of the skull which has a basilar length of 48.8. The tooth- rows are 19.3 in length and the mastoid breadth, 25.4. The relative narrowness indicated by the mastoid breadth is maintained through- out the skull. The only other specimens relating to the Turtle Mountains that have been seen are two male, skins without measure- ments or corresponding skulls, nos. 38902 and 38903, Amer. Mus. Nat. Hist., labeled as from either "Stump Lake or Turtle Mts.", North Dakota. One of these, no. 38902, is much darker than the other. Possibly it is from the Turtle Mountains and the other, lighter-colored one, is from Stump Lake. Study of additional speci- mens from the Turtle Mountains might show the existence there of a distinct race. Four specimens, in the collection of Myron Swenk, from Inland, Clay County, Nebraska, are instructive as showing how inter- gradation occurs between primulina and longicauda. A subadult male, no. 10, is intermediate in external measurements and in color but in each instance is nearer primulina. The same is true of the least width of the color of the underparts. The color of the underparts extends uninterruptedly over the hind legs to the toes as in longicauda, but is absent from the underside of the tail as in primulina. In the skull, the basilar length, breadth of bulla, and size of teeth are nearer longicauda, as are also the ratios to the basilar length of the length of tooth-rows, breadth of the rostrum, length of the tympanic bulla, and depth of the braincase at the anterior margin of the basioccipital. Ratios to the basilar length of the interorbital breadth, mastoid breadth, zygomatic breadth, and depth of the skull at the posterior borders of the upper molars are nearer to those of primulina. The relatively long rostrum, as represented by the orbitonasal length, is nearest to that of spadix. A young, almost subadult, female, no. 7, agrees with primulina in color, color pattern, and length of hind foot. The other external MUSTELA FRENATA 267 measurements are intermediate, but nearer those of primulina. Size of skull and teeth are as in longicauda. Relative proportions of parts of the skull are not diagnostic in specimens as young as this female. An adult female, skull only, no. 8, agrees with, or approaches nearer to, longicauda in size of skull and teeth and in relative proportion of every part studied. A juvenile, skull only, of questionable sex, no. 9, provides no diagnostic characters. On the basis of color, these specimens from Inland are distinctly nearer primulina. On the basis of cranial characters they are distinctly nearer longicauda. External measurements are intermediate and are a little nearer those of primulina. By placing the most weight on the cranial characters, the animals may be referred to longi- cauda. The same may be said of 2 skins, one skin with a skull, from Hastings, Nebraska. In each skin the color-pattern is as in primulina; in one the under side of the tail is nevertheless lighter- colored more as in longicauda and the skull, adult male 121651 American Museum of Natural History, approaches nearer to primu- lina in narrowness but has the large teeth of longicauda. Intergradaticn with neomexicana is suggested by one specimen, no. 7936, Univ. Kans., from Thomas County, Kansas, which has well-developed white facial markings. The specimen, no. 180, Kansas Agric. College, from Glasco, is mounted, of large size, in white winter pelage, and lacks external measurements. On the basis of its obvious large size, and a hind foot measurement of 49 millimeters obtained from the mounted skin, the animal is provisionally referred to longicauda rather than to primulina. Putorius culhertsoni is a name now credited to Coues (1877: 136). Although Coues probably intended only to indicate that Baird wrote this name on the labels of two specimens in the mammal collection of the Smithsonian Institution, Coues gave an "indication" of the application of the name by publishing at the same time the catalogue numbers of specimens whose labels bore the name and thus, in accordance with article 21 of the Interna- tional Rules of Zoological Nomenclature, himself becomes the author of the name. Of the two specimens mentioned by Coues, only the first recorded by him, no. 4320 (with skull no. 37995, U. S. Nat. Mus.), can now be found. Fortunately, the skull of this specimen labeled (see Lyon and Osgood, 1909:218) as taken at Fort Laramie, Wyoming, is well preserved. Its only defects are a fracture in the left zygomatic arch and the absence of parts of each of the first lower molars. In 26S University of Kansas Publs., Mus. Nat. Hist. deciding on the subspecific application of the name Putorius cul- bertsoni Coues, the skull of the type must be principally rehed upon, for there is available only one other specimen, a skin only (no. 12596, U. S. Nat. Mus.), from the same place, and it, like the type, is in white winter pelage and lacks flesh measiu-ements. The ranges as now known of three subspecies of Mustela frenata approach near to Fort Laramie. These are M. /. longicauda, M. f. alleni, and M. /. nevadensis. The skull of the type of culbertsoni is not typical of any one of the three mentioned races. The small size of its teeth and relative (to basilar length) shallowness of the frontal region of the skull through the postorbital processes of the frontal are as in nevadensis. The zygomatic arches are not so greatly expanded as in some specimens of longicauda and are more like the average for nevadensis or alleni, as also is the relatively (to basilar length) long orbitonasal length. However, each of these characters is subject to variation and alone is not surely diag- nostic, especially toward the margin of the range of any one of the subspecies concerned. The same may be said of the relatively great breadth of the skull interorbitally — a feature typically found in longicauda. More important, in my estimation, is the large size of the skull; all parts measured (excepting the teeth, the depth at the posterior border of the last upper molars, the zygomatic breadth, and the depth of the tympanic bullae) equal or approach nearest to the average for males of longicauda of similar age. The small size of alleni prevents its identification with culbert- soni. The question of application lies between nevadensis and longicauda. If the long-tailed weasel at Fort Laramie is found to be referable to the race earlier named longicauda, no change in current nomenclature will be effected. If, on the other hand, the long-tailed weasel from Fort Laramie is found to be referable to nevadensis this name will have to fall before the earlier proposed name culbertsoni. There is, however, a third possibility, namely, that the long-tailed weasel of the Transition and Upper Sonoran zones of southern Wyoming and northern Colorado, as for ex- ample, at Lay, Colorado, may represent a recognizable race char- acterized by size about as in longicauda, relative proportions of skull about as in nevadensis and coloration intermediate, to which the name culbertsoni may apply. For more detailed discussion of this possibility, see remarks under M. f. nevadensis. Satisfactory application of the name Putorius culbertsoni Coues requires an adequate series of adult specimens, of both sexes in the summer coat with external measurements taken in the flesh, MUSTELA FRENATA 269 from the type locality and like material from elsewhere in southern Wyoming. On the evidence furnished by the skull of the type of culbertsoni, that name tentatively is placed in the synonomy of longicauda. Only 2 of 25 adults examined for malformation of the frontal sinuses by parasites showed evidence of disease. Specimens examined. — Total number, 138, arranged alphabetically by prov- inces and states and further by districts or counties from north to south ex- cept as otherwise indicated. Unless otherwise indicated speciinens are in the collection of the United States National Musevmi. Alberta. St. Albert, 1; S. Edmonton, 3; Islay, 4'; Battle River, south of Camrose, V; Daysland, 1'; Moose Pass, 1; Blindman River, 2 (1% 1'); Red Deer, 3 (2', 1*); Bearberry Creek near Sundre, 1'; Canad. Nat. Park, N.W. Territory, 1*; Red Deer River, Didsbury, 1; Canmore, 1; Calgary, 11 (6*, 2°, 1"); Red Deer River, 3'; Little Sandhill Creek, Red Deer River, V; Waterton Lake Park, 2'; Sweetgrass Hills, 1'; Alberta, V. Colorado. Yuma County: Wray 4 ( 1', 3' ) . Kansas. Rowlins County. 7 mi. N, 3 mi. W Beardsley, l**; 6 mi. S and 2 mi. E Atwood, 1"; 15 mi. SE Atwood, 1'. Thomas County, near Brewster, 2°; no locaUty more definite than county, 2'. Trego County, 2 ( 1' ) . Cloud Co. : Glasco, 1'°. Manitoba. Portage la Prairie, 3'; Carberry, 2 (1', 1°); Carman, 1*; Max Lake, Turtle Mt., 1\ Montana. Glacier Co.: St. Marys Lake, 1; Blackfoot, 1; Blackfoot Agency, 1. Blaine County. 6 mi. east Chinook, 1*. Pondera County. Yz mi. SE Con- rad, 1'. Toole Co.: Shelby Junction, 1. Hill Co.: Havre, 1. Fergus Co.: Moccasin Mts., 5 mi. NW Hilger, 1; 7 mi. NE Hilger, 1. Rosebud Co.: % mi. N Ingomar, 1. County in question. Milk River, 2. Nebraska. Dawes County: Chadron, 2". Cherry Co.: Kennedy, 1; no locality more definite than county, 1. Brown County: Long Pine, 1". Antelope County: Nehgh, 1". Adams Co.: Hastings, 2'. Clay County: Inland, 4". North Dakota ( arranged by counties from west to east ) . Divide Co. : Crosby, 1. Mountrail Co.: Lostwood, 1. Little Missouri River, 1. Golden Valley Co.: Sentinel Butte, 1. Billings Co.: Medora, 1*. McLean Co.: 3 mi. W Elbowoods, 1. Oliver Co.: Ft. Clark, 2. Morton Co.: Mandana [= Man- dan?], 1. S/oux Co.: 3 mi. N Cannonball, 1. Logan Co.: 6 mi. SW Napoleon, 1. Rolette County: Turtle Mts., 1"; Fish Lake, 1. Benson Co.: Ft. Totten, 3°; Sully Hill Nat. Park, 1. Ramsey Co.: Devils Lake, 2. Stump Lake or Turtle Mts., 2'. Nelson Co.: Stump Lake, 1. Grand Co.: Larimore, 1. Walsh Co.: Grafton, 11 (4", 3', 2'). Stutsman Co.: Jamestown, 1. Barnes Co.: Valley City, 1. Saskatchewan. Wingard, 5; Osier, 2'; Simpson, 1'; Touchwood Hills, 4"; South arm Last Mountain Lake, 1'; Rush Lake (Assinaboia, N.W.T.), 2\ South Dakota. Pennington Co.: Rapid City, 1. Wyoming. Goshen Co.: Fort Laramie, 2. 1. National Museum of Canada 9. Univ. Kansas Mus. Nat. History 2. Museum of Comparative Zoology 10. Kansas State Agric. College 3. American Mus. Nat. History 11. Collection of Myron H. Swenk 4. Field Museum of Natural History 12. Leland Stanford Junior University 5. Acad. Nat. Sciences of Philadelphia 13. Museum of Zoology, Univ. Michigan 6. Collection of Alex Walker 14. Royal Ontario Museum of Zoology 7. State Hist, and Nat. Hist. Soc. Colo. 15. British Museum of Natiu-al History 8. Mus. Vert. Zool., Univ. California 270 Unu'ersity of Kansas Publs., Mus. Nat. Hist. Mustela frenata oribasus (Bangs) Long-tailed Weasel Plates 16, 17, 18, 31, 32, 33 and 40 Putorius (Arctogale) longicauda oribasus Bangs, Proc. New England Zool. Club, 1:81, December 27, 1899. Putorius longicauda, Coues, Fur-bearing animals, p. 136, 1877 (part). Mustela longicauda oribasus. Miller, U. S. Nat. Mus. Bull., 79:98, Decem- ber 31, 1912. Mustela longicauda oribasa. Hall, Univ. California Publ. Zool., 40:368, November 5, 1934. Mustela frenata oribasa. Hall, Carnegie Instit. Washington Publ. 437:105, November 20, 1936. Type. — Female, adult, skull and skin; no. 9058, collection of E. A. and O. Bangs, but now in collection of Mus. Comp. Zool.; source of Kettle River, 7500 feet [the summit between middle fork of Kettle River and Cherry Creek at Pinnacles — oral information from the collector, Feb. 12, 1936], British Columbia; September 10, 1898; obtained by Allan Brooks; original no. 1368. The skull (plate 40) is complete and unbroken. The teeth all are present and entire except right F which has the anterior half broken away. The skin is complete, fairly well made, and in summer pelage. Range. — Canadian and Hudsonian life-zones from near 56 °N in the Rocky Mountains of British Columbia and Alberta and Ootsa Lake along the Fraser and Cliiicotin rivers south to Alta Lake, in the Caribou and Monashee moun- tains, probably in the SeUcirks and Rockies, and tlirough the Rockies of Mon- tana into extreme northern Wyoming. See figure 29 on page 221. Characters for ready recognition. — DiEFers from M. /. longicauda by near (14 n) Brussels Brown rather than near (h) Clay Color of upper parts and in males by relatively shallower occiput in which the depth of the skull, ex- clusive of the sagittal crest and taken at the anterior border of the basioccipital, amounts to less than 59 per cent of the mastoid breadth; from M. f. nevadensis by greater average size, see measurements. Description. — Size. — Male: Two adults from Florence, Montana, measure as follows: Total length, 440, 440; length of tail, 165, 161; length of liind foot, 47, 49. Corresponding measurements of an adult male from Quesnel, British Columbia, are: 443; 168; 55. Tail amounts to 60, 58, and 61 per cent as long as head and body. Length of hind foot averages more than basal length. Female: The type specimen, the only typical adult or subadult specimen of this sex of which external measurements are available, measures: Total length, 392, length of tail, 150, length of hind foot, 46. Tail is 63 per cent as long as head and body. Length of hind foot amounts to more than basal length. The differences in external measurements, between the one female and the average of the tliree males are: Total length, 49; lengtli of tail, 15; length of hind foot, 4. Externals. — Longest facial vibrissae brown or white (often both colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to or beyond apical pad of fifth digit; hairiness of foot-soles (in summer pelage) shghtly less than shown in figure 19. MUSTELA FRENATA 271 Color. — Upperparts, in summer, near ( 14 n ) Brussels Brown, more black- ish and less reddish than tone 4 of Burnt Umber of Oberthiir and Dauthenay, pi. 304; in type near tone 4, pi. 301 of Oberthiir and Dauthenay. Underparts, in summer, Buff Yellow or near (20 c) Amber Yellow. In winter, all white except tip of tail which is at all times black. Upper parts of uniform color except for occasional slight darkening of top of head and along mid-dorsal line of back. Color of underparts extends distally on posterior sides of fore- legs over feet, on medial sides of hind hmbs over antiplantar faces of toes and over proximal two-thirds of ventral side of tail. Least width of color of underparts amounting to 43 per cent of greatest width of color of upper parts, 75 per cent in male from 4 miles northeast of Quesnel, British Columbia, and 52 (33-66) in four males from Montana. Black tip of tail in four males from Montana averaging 50 (44-60) mm. long. Thus averaging approximately as long as hind foot and 33 per cent of length of tail-vertebrae. Color not different than in many specimens of M. f. nevadensis. Color comparison with M. f. longicauda has been made in the account of that sub- species. Skull and teeth. — Male (based on 5 adults and 2 subadults from British Columbia and 4 adults from Montana): See measurements and plates 16-18. As described in Mustela frenata longicauda except that: Weight, 5.0 (3.8- 6.0) grams; basilar length, 46.7 (43.6-48.8); postorbital breadth in one of nine instances less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tym- panic bulla; breadth of rostrimi less than length of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2/2 to 5 upper incisors; length of tympanic bulla not less than length of lower molar and premolar tooth-row and shorter than rostrvmi. Female (based on the type specimen): See measurements and plates 31- 33, 40. As described in Mustela frenata longicauda except that: Weight, 3.5 grams; basilar length, 41.6 mm.; zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than outside length of P*; tympanic bulla as far posterior to foramen ovale as width of 4/2 upper incisors; height of tympanic bulla less than dis- tance from anterior margin of tympanic bulla to foramen ovale; length of tympanic bulla less than length of rostrum. If more than one skull were avail- able of the female of orihasus it is believed that the description would agree with that of longicauda in nearly all features. The skull of the female is 30 per cent lighter than that of the average male. Comparison with longicauda reveals that, on the average, skulls of males are larger, relative to the basilar length broader across the mastoids, shallower through the braincase as measured at the an- terior end of the basioccipital exclusive of the sagittal crest, with longer rostrum. Compared with nevadensis, the skull averages larger in all measurements taken, and has a relatively broader rostrum, relatively greater mastoid breadth and a braincase which 272 University of Kansas Publs., Mus. Nat. Hist. is shallower relative to the basilar length. By weight, the skull of nevadensis is a fourth lighter, and in linear measurements 5 to 18 per cent smaller. Remarks. — Some of the specimens from Montana, which here are referred to oribasus, more than half a century ago were listed by Coues (1877:138) under the name longicauda. It was not until 1899 that this race was given a name by Bangs, who at that time (1899B:81) accurately made out the distinctive color features. Distinctive cranial characters cannot be described with assurance even now because there still are too few specimens. The type specimen, at one time examined by the present writer, has on the stuffed skin no well-developed mammae, scrotal pouch, or other visible sexual part. Probably the collector's sex mark for female is correct. As judged by the two skulls of subadult males from the Barker- ville region, individuals of this race attain larger size than do those of longicauda. On the basis of larger size than either longicauda or nevadensis, the specimens from the Rocky Mountains of Mon- tana and two from northern Wyoming are referred to this race. The short, wide, flat, tympanic bullae, relatively great mastoidal breadth, and some other features of the specimen from Donovan, Montana, point toward oribasus, whereas nearly as many more cranial fea- tures, in this instance mainly differences in size, are indicative of nevadensis to which race the specimen might almost equally well be referred. Another male from Darby, in the Bitterroot Valle>- of Montana, has a slightly longer hind foot than those from Flor- ence, but a female from Hamilton, agrees more nearly with neva- densis. The average of all the specimens from the Bitterroot Valley is a little nearer oribasus. Four skulls from Buffalo, Wyoming, here referred to nevadensis show approach to oribasus in size of skull. The specimens from Big Snowy Mountains, and the Highwood Mountains of Montana are too young clearly to show size of the adult skull, but are distinctly darker colored than longicauda of the plains country proper. Of two subadult females from Tacy, Mon- tana, the color of the one in summer pelage is distinctly nearer that of oribasus and nevadensis than it is to that of longicauda to which some approach in color might be expected. The reduced size of both of the specimens is further suggestive of nevadensis and it may be that adult specimens from these more eastern moun- tainous areas in Montana will show that nevadensis is the name proper to apply to animals of this region. Intergradation with nevadensis is suggested by specimens col- MUSTELA FRENATA 273 lected from along the upper reaches of Okanagan Lake, British Cohimbia, by Major Allan Brooks and Mr. J. A. Munro and by a series of skulls from lone, Pend Orielle County, Washington, lent me by Mr. Walter Dalquest. At each place, the average of all specimens is nearest to that of nevadensis. Specimens from near Waterton Lake show several steps in the transition from the light-colored longicauda type of coloration to the darker coloration characterizing oribasus. One taken here, at a time when the body of water referred to seems to have been known as Chief Mountain Lake, is barely dark enough to be placed with oribasus. Two other specimens from across the Canadian Border labeled as "Waterton Lake Park" are slightly lighter colored above, and on this account are placed with longicauda. The two adult males from Lillooet, British Columbia, are refer- able to oribasus although neither is quite typical. One has a satu- rated coloration suggestive of that of aJtifrontalis and the skull is shorter and broader than in other specimens of oribasus. The fe- male from Lillooet, skin alone, no. 916, Prov. Mus., B. C. is small for oribasus. The female, no. 1539, collection of Kenneth Racey, from Alta Lake, in brown winter pelage, in almost every measure- ment falls nearly midway between altifrontalis and oribasus but slightly nearer the latter. The skull from Chezacut and 3 animals from Wistaria, British Columbia, probably are females and show a greater average size than specimens from farther to the southeast. For example, the basilar length of the skull, 44.8 (44.3 to 45.1), exceeds that of the type specimen. The animals from Wistaria on Ootsa Lake furnish the northwesternmost station of occurrence of which I have record for this subspecies. The northernmost records of occurrence, at "Clearwater River, Peace R[iver]., B. C," and at Little Prairie, are furnished by a white skin without skull, no. 257450, U. S. Nat. Mus., purchased on August 2, 1932, at the place mentioned by W. H. Sheldon and Richard Borden, and a skull with white winter skin, no. 3585, Pro- vincial Museum, British Columbia, respectively. The characters distinguishing longicauda and oribasus are not shown by white win- ter skins; the skull shows some features of longicauda, and the ref- erence of tiiese specimens to oribasus rather than longicauda is tentative. Only the skull from Little Prairie shows evidence of infestation of the frontal sinuses by parasites. In the Barkerville area of British Columbia, Mr. and Mrs. Thomas T. McCabe obtained only 2 274 University of Kansas Publs., Mus. Nat, Hist. skulls of this subspecies from a total of 238 weasel skulls gathered by local trappers. The others were Mtistela erminea. Specimens examined. — Total number, 46, listed by localities from north to south and unless otherwise indicated, in the United States National Museum. British Columbia. West of Hudson Hope, 1"; Clearwater River, tributary to Peace River, 1; Little Prairie, a few miles south of Peace River and about 40 miles west of the main highway between Dawson Creek and Fort St. John, 1'; Wistaria, 3*; Four Mile Creek, 4 mi. NE Quesnel, 1^ Isaacs Lake, 3200 ft., 1^ Barkerville region, 1=; Clear River, 4800 ft., 1'; Chezacut, 1"; Lillooet 3 (2', 1'); Alta Lake, 1'°; source of Kettle River, 7500 ft., 1°; E side Beaverfoot Range, 4000 to 4500 ft. between Eraser Creek and 6 mi. SE of Eraser Creek, V; Cranbrook, 1^ head of Cross River, 10 mi. below Assiniboine Pass, 1"; camp east of "Kootanie", 1"; camp east of Kootanie River, 1". Alberta. Thoral Creek, 7000 ft., 50 mi. NE Jasper, 1'. Montana. Glacier? Co.: Chief Mt. Lake (=:Waterton Lake), 1. Flathead Co.: Columbia Falls, 1. Chouteau? Co.: Highwood Mts., 1. Fergus? Co.: Big Snowy Mts., 1. Wheatland Co.: Harlowton, 1'. Ravalli Co.: Florence, 2; Hamilton, 1"; Darby, 1'; Carlos [= Charlos] Heights, 2=; Tin Cup District, 2=; no locality more definite than county, 2". Beaverhead Co.: Donovan,!. Madi- son Co.: Sheridan, 1'. Gallatin Co.: Ranch 7-11, Eldridge, l\ Stillwater County: Tacy, 2". County in question: Gallatin Valley, 1; Yellowstone Park, 1'. Wyoming. Glen Creek, Mammoth Hot Springs, 1. Park Co.: Four Bears, V. Mustela frenata alleni (Merriam) Long-tailed Weasel Plates 18, 19, 20, 31, 32 and 33 Putorius alleni Merriam, N. Amer. Fauna, 11:24, June 30, 1896. Mustela alleni. Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912. Mustela frenata alleni, Hall, Carnegie Instit. Washington Publ. 473:106, November 20, 1936. Type. — Male, adult, skull and skin; no. 186451, U. S. Nat. Mus. (formerly 4485/5120, collection of Dr. C. Hart Merriam); Custer, South Dakota; obtained by Vernon Bailey; original no. 90. The skull is complete and unbroken. The upper incisors are missing. All the other teeth are present although the premolars, and especially the canines, are much worn, possibly as the result of the animal's efforts to free itself from a trap. The skin is fairly well made, in a good state of preservation, and entire. Range. — Canadian, Transition and Upper Sonoran life-zones of the Black Hills of South Dakota and adjacent semi-bad-land territory of Wyoming and Nebraska southward to Mitchell, Scottsbluff County. See figure 29 on page 221. Characters for ready recognition. — DifiFers from M. f. longicauda in smaller size, adult males having a total length of less than 400, hind foot less than 45, 1. Collection of Ian McTaggart-Cowan 7. American Mus. Nat. History 2. Mus. Vert. Zool., Univ. California 8. Royal Ontario Museum of Zoology 3. National Museum of Canada 9. Coll. Rocky Mt. Spotted Fever Lab. 4. Provincial Museum of British Columbia 10. Collection of Kenneth Racey 5. Museum of Comparative Zoology 11. British Museum of Natural History 6. Field Museum of Natural History 12. Museum of Zoology, Univ. Michigan MUSTELA FRENATA 275 basilar length less than 43.5, and in adult females total length less than 375, and basilar length less than 40; from M. f. nevadensis in near Clay Color rather than near (14 n to /) Brussels Brown of upper parts in summer. Description. — Size. — Male: External measurements of the type specimen are: Total length, 372; length of tail, 137; length of hind foot, 44. Tail is 58 per cent as long as head and body. Length of hind foot more than basal length. Female: No external measurements for typical adults are available. No. Ml #41 from Mitchell, Scottsbluff Co., Nebraska, an adult female which is an intergrade with the larger M. f. longicauda, measures as follows: Total length, 367; length of tail, 120; length of hind foot, 41. Externals. — Longest facial vibrissae dark brown or white and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) as shown in figure 20. Color. — ^Winter pelage unknown; probably white except, of course, tip of tail. Summer pelage as described in Mustela frenata longicauda except that: Least width of color of underparts averaging, in 3 males from Black Hills, 54 (38-62) per cent of greatest width of color of upper parts. Black tip of tail averaging 43 (40-45) mm. long. Thus, averaging approximately same length as hind foot and in type specimen amounting to 33 per cent of length of tail-vertebrae. Skidl and teeth. — Male (based on the type and no. 7440 Amer. Mus. Nat. Hist., from Hill City, S. Dak.): See measurements and plates 18-20. As de- scribed in Mustela frenata longicauda except that: Weight, 3.1 (3.0-3.2) grams; basilar length, 41.0 (40.9-41.0); mastoid breadth not less than post- palatal length; breadth of rostrum more than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 5 upper in- cisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale. Female (based on no. 7441, American Mus. Nat. Hist., from Black Hills, S. Dak. ) : See measurements and plates 31-33. As described in Mustela frenata longicauda except that: Weight, 2.0 grams; basilar length 37.6. The skull of the female is 35 per cent fighter than the average for the two males. Comparison with M. /. longicauda and M. /. nevadensis reveals that the tympanic bullae average more nearly flat and tliat the skull is smaller. Remarks. — Animals of this subspecies were described and named by Merriam in 1896 as a distinct species on the basis of two or pos- sibly three specimens from the Black Hills of South Dakota and the name seems never to have been applied to specimens from other regions. Vernon Bailey obtained only the one specimen, the type, on his trip in 1888, but two more were obtained for the American Museum of Natural History by Walter Granger in 1894. Mustela frenata alleni combines the light coloration of M. f. longi- cauda with the small size of M. f. nevadensis. Indeed, the size may average less than that of nevadensis. M. f. alleni seems to reach its extreme of small size in the Black Hills of South Dakota. Speci- 276 University of Kansas Publs., Mus. Nat. Hist. mens from Mitchell, Scottsbluff County, Nebraska, here referred to alleni are of larger size and in this respect are intermediate between the subspecies alleni and longicauda. Of the two specimens avail- able from Chadron, Nebraska, and here referred to as longicauda, the female, Ml #6, is almost exactly intermediate in size between alleni and longicauda, whereas the male, Ml #11, is as large as the average-sized longicauda. None of the nine skulls (5 adults) shows malformation resulting from the infestation of the frontal sinuses with parasites. Specimens examined. — Total number, 10, as follows. Wyoming. Crook Co.: Sundance, 1'. South Dakota. Pennington Co.: Hill City, 1=; 20 mi. N Elk Mt., 1'. County in question: Black Hills, V. Custer Co.: Custer, 2 ( 1\ 1"). Nebraska. Scottsbluff County: Mitchell, 4\ Mustela frenata arizonensis (Meams) Long-tailed Weasel Plates 19, 20, 21, 31, 32 and 33 Putorius arizonensis Mearns, Bull. Amer. Mus. Nat. Hist., 3:234, June, 1891; Merriam, N. Amer. Fauna, 11:22, fig. 12, June 30, 1896. Mustela arizonensis, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912. Mustela frenata arizonensis. Hall, Carnegie Instit. Washington Publ. 473: 106, November 20, 1936. Type. — Female, adult, skull and skin; no. 2490/1886, Amer. Mus. Nat. Hist; San Francisco Forest [then (1886?), Yavapai County], Arizona; June 20, 1886; obtained by Edgar A. Mearns. The skull (plates 31-33) is complete and unbroken save for a small punc- ture in the right squamosal. The incisors above and belovi^ and M^ and P^ on each side are missing. Four canines are preserved separately. Otherwise the teeth are in place. The skin has been taken down from a mount. Some hair has been lost from in front of the ears. Seven mammae are evident and show the animal to have been nursing young. The shghtly faded color was mentioned by Mearns in the original description. He says (1891:234): "The memorandum of the colors was made before skinning, the specimen having been subsequently preserved in a solution of alum and salt, which extracted much of the coloring matter." Range. — Transition to Hudsonian life-zones of Arizona and extreme western New Mexico, along the Colorado River, and south of the Little Colorado River, from San Francisco Mountain region along Mogollon Plateau to extreme west- ern New Mexico. See figure 29 on page 221. Characters for ready recognition. — Difi^ers from M. f. neomexicana by near (14 n) Brussels Brown rather than Buckthorn Brown color of upper parts, in absence rather than presence of white frontal spot continuous with color of underparts, in basilar length of less than 44 in males and 39.3 in females; 1. United States National Museum 3. Collection of Myron H. Swenk 2. American Mus. Nat. History MUSTELA FRENATA 277 from M. /. nevadensis in that total length averages less than 375 in males and 330 in females, basilar length averaging less than 41 in males and less than 36.7 in females. Description. — Size. — Male: No. 24679/32071, from Springerville, and no. 248993 from the Kaibab Plateau, measure respectively, as follows: Total length, 363, 367; length of tail, 140, 143; length of hind foot, 41.5, 41.0. Tail is 63, and 64 per cent as long as head and body. These males, the only spec- imens of that sex of which external measurements are available, probably are grading toward nevadensis and therefore are nontypical. Female: Three specimens, one young from Little Spring, a subadult from Deadmans Flat and the type specimen, measure respectively as follows: Total length, 323, 296, 302; length of tail, 110, 101, 109; length of hind foot, 38, 33, 36. These average, 307, 107, 36. Tail averages 53 per cent as long as head and body. Differences in external measurements of the two sexes are: Total length, 56; length of tail, 39; hind foot, 5.5. Externals. — Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot soles ( in summer pelage ) about as shown in figure 19. Color. — Winter pelage unknown. Summer pelage with upper parts near (14 n) Brussels Brown or tone 2 of Raw Umber of Oberthiir and Dauthenay, Pi. 301, darker on top of head from nose to line connecting posterior margins of ears. Tip of tail always black. Chin and upper lips white. Remainder of underparts Buff Yellow to Straw Yellow and rarely Ochraceous Buff. Color of underparts extends distally on posterior sides of forelegs over toes onto anti- palmar faces of feet and wTists, on medial sides of hind legs to ankles and over antiplantar faces of toes, medial third of tarsus, and over proximal fifth to fourth of ventral side of tail. Least width of color of underparts averaging, in 8 specimens, 44 (29-54) per cent of greatest width of color of upper parts. Black tip of tail, in four females averaging 35 (33-38) mm. long. Thus, aver- aging shorter than hind foot and 32 per cent of length of tail-vertebrae. Three of the eight specimens before me (no. 242671 from 25 mi. SE Flagstaff, not available at time of this accounting) have the dark spot near the angle of the mouth faintly indicated, whereas the other five lack the spots. The color is as in M. /. nevadensis. Skull and teeth.— Male (based on 55211, 65231, and 248993; see p. 422): See measurements and plates 19-21; weight 2.7 and 3.1 grams; basilar length, 40.4; zygomatic breadth more than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more than postpalatal length; postorbital breadth less than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; inter- orbital breadth more or less than distance between foramen opticum and an- terior margin of tympanic bulla; breadth of rostrum less than length of tym- panic bulla; least width of palate more or less than medial length of P4; an- terior margin of tympanic bulla as far posterior to foramen ovale as width of 3/2 (including 13) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than 278 University of Kansas Publs., Mus. Nat. Hist. length of lower molar-premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below talonid of ml. Female ( based on the type specimen ) : See measurements and plates 31-33; weight, 1.6 grams; basilar length, 35.5; zygomatic breadth less than distance between condylar foramen and Ml and more than distance between anterior palatine foramen and anterior margin of tympanic bulla (nearly equal in each instance); postorbital breadth less than length of upper premolars and greater (7.1-8.4) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate equal to inside length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 (including 13) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar-premolar tooth-row and greater than length of rostrum. The skull of the female averages 41 per cent Hghter than that of the male. Compared with the skull of M. /. nevadensis, that of arizonensis is smaller, less heavily ridged and has more inflated tympanic bullae and a relatively greater mastoid breadth. Comparison with the skull of M. f. neomexicana is made in the account of that subspecies. Remarks. — In 1891 Mearns ( 234-235 ) named this weasel as a full species on the basis of two individuals taken by him in 1886 and 1887. Since that time only a few additional specimens have been preserved. Only four are adults. Although this material does not permit of a definition of the subspecies as precise as could be wished, still, it clearly shows tliat the animals from the plateau region of Arizona are recognizably different from those farther north in the Sierra Nevada of CaHfornia and those of the Rocky Mountains and Great Basin region northward to the Canadian border. These more northern animals have gone by Mearns' name, arizonensis, since the date of its proposal until 1939 when the name nevadensis was pro- posed. The smaller size, especially of the skull, and the greater inflation of the tympanic bullae are the outstanding characters which dis- tinguish arizonensis from the similarly marked nevadensis. The bullae are relatively much inflated throughout but especially so on the posteromedial parts. Although the three adult males and two subadult females avail- able of this subspecies are smaller in most parts measured than any of the scores of nevadensis of similar age tliat have been measured, overlap in size probably will be found as additional specimens of arizonensis become available. A young female, no. 18513, coll. D. R. Dickey, from Little Spring, does have certain cranial measure- ments as large as are found in the minimum-sized nevadensis from farther north. MUSTELA FRENATA 279 Intergradation with the two subspecies whose geographic ranges adjoin that of arizonensis is indicated by specimens at hand. One of tliese is the adult male from 25 miles southeast of Flagstaff, which shows decided approach to neomexicana, in color and in possessing white facial markings less well developed than in neomexicana. Even better developed white facial markings, with intervening blackish coloration, are displayed by no. 148271, U. S. Nat. Mus., from 8500 feet altitude on Willow Creek, New Mexico. This sub- adult female shows approach to neomexicana also in larger size of the skull and entire animal. The great inflation of the posterior part of each of its bullae and the dark color of the upper parts are char- acters of arizonensis. The color of the underparts stops at the ankles leaving the hind feet dark colored, in which respect the specimen is unlike either neomexicana or arizonensis. If additional specimens showing the same characters as this one be found at other nearby localities they probably should be given recognition as a separate subspecies. For the present it seems best to regard the specimen merely as an intergrade. Although it might, with almost equal propriety, be referred to either neomexicana or arizo- nensis, the specimen is here placed with the latter. The subadult male from Springerville, Arizona, is of larger size than the topo- typical male of arizonensis and in this respect shows slight approach to nevadensis. The narrower mastoidal breadth and slightly less inflated tympanic bullae of the male from the Kaibab Plateau may reflect merely individual variation or may represent intergradation in these features with nevadensis. The statement made by Merriam (1896:22) that, "The type speci- men . . . is an immature female and is of unusually small size. A male obtained by him [Mearns] near the same place is of the nor- mal size, as is another male in the Department collection from Springerville, Ariz., collected by E. W. Nelson", needs correction. The female is not immature. The specimen obtained by Mearns near the same place probably refers to Amer. Mus. No. 2489, from Quaking Asp Settlement, which lacks both the skull and external measurements. As stuffed it is of small size for a male. The male from Springerville, as shown by the external and cranial measure- ments, is not of normal ( i. e. average ) size, but is smaller than the average for the other populations of similarly colored weasels re- ferred to by Merriam (op. cit.) as arizonensis but here described under the name nevadensis. None of the skulls shows signs of infestation of the frontal sinuses by parasites. 280 University of Kansas Publs., Mus. Nat. Hist. Specimens examined. — Total number, 17, arranged alphabetically by states and from north to south by counties in each state. Unless otherwise indicated specimens are in the collection of the United States National Museum. Arizona. Coconino Co.: VT Park, Kaibab Plateau, 1; Deadman Flat, 6400 ft., 1'; Little Spring, V; Government Prairie, near Parks, 1'; Coconino? Co.: San Francisco Forest (Yavapai Co., in 1886), V; 25 mi. SE Flagstaff, 1; Quak- ing Asp Settlement, V. Apache Co.: Springerville, 1; North Fork White River, White Mts., 8200 ft., 4'; head San Francisco River, Judd Ranch, Alpine, 1'; 2 mi. SE Big Lake Knoll, 8700 ft., 24 mi. S Springerville, l\ Greenlee County: S end Blue Range, 9000 ft., Prieto Plateau, 1; Beaver Creek, 7000 ft., 1". New Mexico. Grant Co.: MogoUon Mts., Willow Creek, 8500 ft., 1. Mustela frenata nevadensis Hall Long-tailed Weasel Plates 19, 20, 21, 33, 34, 35 and 39 Mustela frenata nevadensis Hall, Carnegie Instit. Washington Publ. 473:91, November 20, 1936. Putorius longicauda, Coues, Fur-bearing animals, p. 136, 1877 (part); Mer- riam, N. Amer. Fauna, 5:83, July 30, 1891. Putorius (Gale) hrasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part). Putorius arizonensis, Merriam, N. Amer. Fauna, 11:22, figs. 13, 14, June 30. 1896 (part); Stephens, Mammals of Cahfomia, p. 247, 1906. Mustela arizonensis, Grinnell and Swarth, Univ. Cahfomia Publ. Zool, 10:376, October 31, 1913; Whitlow and Hall, Univ. California Publ. Zool., 40:247, September 30, 1933. Mustela arizonensis arizonensis, Grinnell, Univ. California Publ. Zool., 40:102, September 26, 1933. Mustela frenata, Boyer, Joum. Mamm., 24:99, February 20, 1943. Type. — Female, adult, skull and skin; no. 41053, Mus. Vert. Zool; three miles east Baker, White Pine County, Nevada; May 30, 1929; obtained by E. R. Hall and W. C. Russell; original no. 2674, E. R. H. The skull (plates 33-35) is complete and unbroken. The teeth all are present and entire. The skin is fairly well made. Eight mammae are evident and show the animal to have been nursing young. Range. — Altitudinally, 700 feet at Wenatchee, Washington, to the highest parts of the mountains of the western United States; Upper Sonoran Life-zone to Arctic Alpine Life-zone; southern British Columbia in the Cascades and territory west to Monashee Mountains, and Nelson, southward in the Cascades of northern Washington, over western Wa.shington, Idaho, Utah, and Nevada to northeastern Arizona and northern New Mexico; westward from the eastern base of the Rocky Moimtains in Colorado to the western base of the Sierra Nevada and Cascades of California and to the Cascades of southern Oregon. See figures 29 and 30 on pages 221 and 314. Characters for ready recognition. — Differs from M. f. oribasus by smaller average size, see measurements; from M. f. longicauda by near (14 n to I) Brussels Brown rather than near (h) Clay Color of the upper parts, and in males by a shallower occiput in which the depth of the skull, exclusive of the sagittal crest, and taken at the anterior border of the basioccipital, amounts 1. Mus. Vert. Zocil., Univ. California 3. American Mus. Nat. History 2. Donald R. Dickoy Colhction 4. San Diego Society of Natural History MUSTELA FREN'ATA 281 to less than 59 per cent of the mastoid breadth; from M. f. alleni by near (14 n to /) Brussels Brown rather than near (h) Clay Color of upper parts in summer; from M. /. neomexicana by near ( 14 n to / ) Brussels Brown rather than Buckthorn Brown color of upper parts, in absence of white frontal spot continuous with color of underparts, in basilar length of less than 46 in males and 40 in females; from M. f. arizonensis by total length averaging more than 375 in males and 330 in females, basilar length averaging more than 41 in males and 36.7 in females; from M. f. inyoensis by absence of white facial markings; from M. f. pulchra by absence of light facial markings, near ( 14 n to I) Brussels Brown rather than near (16 /) Buckthorn Brown color of upper parts, and lesser size, hind foot less than 40 in females and basilar length averaging less than 46.0 in males; from M. f. xanthogentjs by absence of light facial markings and near (14 n to I) Brussels Brown rather than Buckthorn Brown color of upper parts; from M. /. munda by absence of white facial markings, presence of color of underparts on ventral face of proximal third of tail, and hind foot of less than 50 in males; from M. /. saturata by presence of light color of underparts on tail and ankle and in lesser average breadth across mastoid processes of skull (see measurements); from M. f. oregonensis by absence of nasofrontal white patch, presence of light color of underparts on ventral face of tail, and shorter skull, which, relative to its length in males, is deeper through the braincase; from M. /. washingtoni by presence of light color of underparts on ventral face of tail, by skull which in male relative to basilar length is shorter in the preorbital region and wider across the zygomata and mastoid processes, and in female has longer preorbital region and larger bullae (see measurements); from M. /. altifrontalis by lighter col- ored upper parts which are tones 1 to 3 of Raw Umber, pi. 301, rather than tone 4 of Brownish Drab, pi. 302, of Oberthiir and Dauthenay, by Buff- Yellow to Straw Yellow rather than near ( 14' a to 16' c ) Ochraceous-Buff color of underparts, by least width of color of underparts amounting to more than 37 per cent of greatest width of color of upper parts, by presence of color of underparts on ventral side of tail and on hind leg over ankle, and by lesser depth of skull through frontal region; from M. /. effera by larger size, males averaging 12^2 per cent larger in external measurements, 8 per cent larger in linear measurements of skull, and 22 per cent heavier in weight of skull, total length averaging 400 rather than 360, basilar length averaging 43.6 rather than 40.5. Description. — Size. — Male: Twenty-one adults from the southern half of the Sierra Nevada of California yield average and extreme measurements as follows: Total length, 400 (356-428); length of tail, 150 (125-178); length of hind foot, 46.1 (42-50). Tail averages 60 per cent as long as head and body. Length of hind foot averaging more than basal length. Corresponding measurements of twelve adults from extreme southern and southwestern Colorado are as follows: 407 (355-431); 150 (133-170); 46.0 (42-49). Female: Ten adults from the Sierra Nevada of California yield average and extreme measurements as follows: Total length, 349 (336-362); length of tail, 127 (120-133); length of hind foot, 36.3 (32-39). Tail averages 57 per cent as long as head and body. Length of hind foot less than basal length. Corresponding measurements of ten adults from the Rocky Moun- tains of central Colorado are as follows: 347 (325-375); 123 (111-141); 40 (32-43). 282 University of Kansas Publs., Mus. Nat. Hist. The average differences in external measurements of the two sexes, in the Sierras of California are: Total length, 51; length of tail, 23; length of hind foot, 9.8. Weight of 7 adult males from Cahfomia is 267 (226-345) grams. Two adult females from there weigh 148 and 115 grams and 3 from White Pine County, Nevada, 134, 122 and 124, giving an average of 129 grams. Externals. — Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) about as shown in figure 19. Color. — Upper parts, in summer, near (14 n to I) Brussels Brown or tones 1 to 3 of Raw Umber of Oberthiir and Dauthenay, pi. 301, darker on top of head from nose to fine connecting posterior margins of ears. Chin and upper lips white. Remainder of underparts Buff-Yellow to Straw Yellow and sometimes Ochraceous-Buff especially in young, and in some adults from southern Colorado. In winter, all white, except tip of tail, or upper parts near (/) Snuff Brown or fighter than Brussels Brown with a smoked effect, and underparts white. Tip of tail at all times black. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind legs to ankles, over antiplantar faces of toes, medial third of tarsus and usually over proximal tenth to three-foiuiiis of ventral side of tail. Least width of color of underparts averaging, in a series of twenty males from the southern half of the Sierra Nevada of Cah- fomia, 59 (37-76) per cent of greatest width of color of upper parts. In seven males from southern Colorado corresponding percentages are 55 (37-71). Black tip of tail in series from Sierra Nevada averaging 50 ( 40-60 ) mm. long; thus longer than hind foot and averaging 33 1/3 per cent of length of tail- vertebrae. Skull and teeth. — Male (based on 25 adults, from Sierra Nevada of Cah- fomia): See measurements and plates 19-21; weight, 3.7 (2.9-4.9) grams; basilar length, 43.6 (40.6-46.1); zygomatic breadth more than distance be- tween condylar foramen and Ml (save in four instances) and more tlian dis- tance between anterior palatine foramen and anterior margin of tympanic bulla (save in two specimens); mastoid breadth more (80 per cent of speci- mens) or less (20 per cent) than postpalatal length; postorbital breadth less than length of upper premolars and more or less than width of basioccipital measured from medial margin of one foramen lacemm posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than medial length of P4 (except in two specimens); anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostmm; anterior margin of masseteric fossa not carried farther forward than point directly below hypoconid of ml. Female (based on ten adults from Sierra Nevada of Cahfomia): See measurements and plates 33-35; weight, 2.2 (1.8-2.4) grams; basilar length, 38.2 (36.7-39.5); zygomatic breadth more (except in one specimen) than distance between condylar foramen and Ml and more (save in two speci- MUSTELA FRENATA 283 mens) than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and less than (except in one specimen) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least widtli of palate more or less tlian either outside or inside length of P4 but generally less than inside length; tympanic bulla as far posterior to foramen ovale as width of 3 to 5/2 upper incisors; height of tympanic bulla more or less (usually more) than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than length of rostrum. The skull of the female averages 41 per cent lighter than that of the average male. Compared with the skull of M. /. longicauda, that of both sexes averages smaller in every measurement taken. Males of nevaden- sis, on the average, relative to the basilar length, are narrower in the interorbital region and across the zygomata but have the orbi- tonasal length greater. Stated in another way, the rostrum of longicauda appears to be shorter and broader and the zygomata are more expanded. Females of nevadensis, on the average, relative to the basilar length are narrower across the mastoid processes and zygomata and have the braincase deeper at the anterior margin of the basioccipital. Also in nevadensis the mastoid processes do not project so far laterally beyond the braincase, the lambdoidal crest and postorbital processes are less well developed and except in the interparietal region, the temporal ridges hardly meet and they form a sagittal furrow rather than a low sagittal crest which characterizes adult females of longicauda. Each of these differences separating the females of longicauda from those of nevadensis are of the same nature, although not necessarily of the same degree, as tliose which appear in longicauda with increasing age. The differences men- tioned above are readily appreciable when series of specimens are compared. However, none of the differences is of great degree, and most parts of the skulls of the two subspecies are of similar rela- tive proportions. Even so, there is but little overlap in actual size. Comparisons with the skulls of M. /. orihasus, alleni, neomexicana, arizonensis, inyoensis, ptdchra, xanthogenys, munda, saturata, ore- gonensis, washingtoni, altifrontalis, and effera are made in the ac- counts of those subspecies. Remarks. — The populations to which the name nevadensis at present is assigned have gone by the name arizonensis since Mearns proposed this name in 1891. Before that time Coues (1877:141) had included individuals of this race under the name Puforius longi- cauda. 284 University of Kansas Publs., Mus. Nat. Hist. Among the populations here assigned to M. f. nevadensis, there is some geographic variation but it is of lesser degree than in most other species of mammals which range over the same region. Com- parison of 20 adult males from the Rocky Mountains of Colorado with 25 adult males from a place as far distant as the Sierra Nevada of California shows that the two populations closely resemble each other. The specimens from Colorado average a trifle wider across the zygomata, have a longer body and therefore relatively shorter tail, and, except in southern Colorado, a slightly longer hind foot. Comparison of ten adult females from each of the two areas reveals that those from Colorado have a markedly longer hind foot, and a tail somewhat shorter relative to the length of the body. The men- tioned differences are the only ones found among the great num- ber of points investigated, except that as remarked by Merriam ( 1896:23 ) the Sierran animal has the yellow of the underparts reach- ing farther up under the chin, the underside of the tail on the aver- age is more suffused with yellowish and the white on the upper lip is more extensive. As regards the last mentioned feature, my check of 34 skins from Colorado reveals that the white extends all the way around the upper lip in every specimen but one, whereas in 69 specimens from the Sierra Nevada the white extends all the way around the upper lip in only 39. However, as further re- marked by Merriam {loc. cit.), not only this but the other color features are inconstant in addition to being slight. When the oc- currence of the dark spots near the angles of the mouth are tabu- lated, it is found that in 33 Colorado-taken specimens they are absent in 19, faintly indicated in 13, and well developed in 1. In 62 California-taken specimens they are absent in 37, faintly indi- cated in 20, and well developed in 5. In northwestern Colorado, southern Wyoming, and possibly through the Bear River Divide into southeastern Idaho, long-tailed weasels here referred to nevadensis approach longicauda in large size and occasionally in other features, more closely than do speci- mens of nevadensis from most other places in its range. This tend- ency is thought to be significant for much of the area in question lies in or below the Transition Life-zone, the same life zones in which farther to the eastward true longicauda occurs. One specimen that illustrates this approach to longicauda is an adult male, no. 2334, collection of E. R. Warren, from 6160 feet, Lay, Routt [now Moffat] County, Colorado. In large size and, relative to the basilar length, shorter rostrum and shorter tympanic bullae, it agrees with longicauda but the darker color and, relative MUSTELA FRENATA 285 to the basilar length, narrowness of the rostrum, mterorbital region, zygomatic expanse and the shallowness through the region of the postorbital processes place it with nevadensis. Of two other speci- mens from Steamboat Springs, Routt County, a young male, no. 4010, in the collection of E. R. Warren, has a hind foot (50 mm.) as long as in longicauda; and the other, no. 138195, U. S. Nat. Mus., an adult male, agrees well enough in size and proportions with nevadensis but has the coloration typical of longicauda. From Wyoming, one subadult female, no. 177553, U. S. Nat. Mus., from Garrett, is intermediate in size and coloration but is nearer to nevadensis in these particulars, as it is in all other points consid- ered except size of the molar teeth which are as large as in longi- cauda and larger than in any female nevadensis from Colorado or California. Another female, an adult, no. 179304, U. S. Nat. Mus., from Lonetree, Wyoming, agrees with longicauda in size of skull. Indeed, ten of seventeen cranial measurements exceed the maximum for Colorado-taken nevadensis. Where differences exist in relative proportions of the skull as expressed in percentages of the basilar length, the specimen approaches nevadensis in 5 instances and longicauda in only 3. The color is intermediate but much nearer that of nevadensis with which the animal agrees also in external measurements. Ten subadults (5 of each sex) from within 12 miles of Laramie (not Fort Laramie) show greater resemblance to nevadensis but definitely approach longicauda. Average external measurements are: J , 408, 155, 44; 5 , 361, 134, 40. The two other specimens examined from this general locality, a young female, no. 2711, Mus. Vert. Zool., from Fort Bridger, and a sub- adult female, no. 188377, U. S. Nat. Mus., from Bridger Pass, show no departures from nevadensis of similar age. The specimens from scattered localities in the Transition Life- zone of northwestern Colorado and southern Wyoming are larger than nevadensis is elsewhere, and also in certain other features resemble longicauda of the plains to the eastward. Everything considered, the animals in question are much more like nevadensis than longicauda. Study of more specimens, especially from Wyo- ming, might provide grounds for recognizing as a different sub- species the animals in this large area comprising parts of Colo- rado and Wyoming from which so few specimens now are available. Possibly the name Putorius culbertsoni Coues would apply. De- cision on that point will require adequate material from the type locality. Fort Laramie. See discussion of this name under M. /. longicauda. 286 University of Kansas Publs., Mus. Nat. Hist. In southeastern Idaho males are larger than they are at most other places within the range of nevadensis. An average of 7 adults and subadults from Pegram, Montpelier, Springfield, and the vicinity of Pocatello, reveals, when compared with the average of nevadensis from Colorado and that of longicaiida from tlie Great Plains, that this population from southeastern Idaho is nearest to longicaiida in linear measurements of the orbitonasal length, mas- toid breadth, length of tympanic bullae, and as expressed in per- centage of the basilar length, length of tooth-row, breadth of ros- trum, and zygomatic breadth. In all other points of size, relative proportions and color, the animals approach nearer to, or actually agree with, nevadensis. The specimens commented upon clearly show intergradation be- tween nevadensis and longicauda. Similarly, the specimens from Scottsbluff County, Nebraska, here referred to M. /. alleni, by their larger size suggest intergradation of that subspecies with the larger nevadensis-longicauda stock although the approach is more toward longicauda than nevadensis. Between oribasiis and nevadensis, however, there is no lack of material showing intergradation. As set forth in the account of oribastis, specimens from Montana are truly intermediate structurally as well as geographically. Intergradation with washingtoni is shown by specimens from the northern part of the Cascade Range in Chelan and Okanogan counties, Washington. The adult male, U. S. Nat. Mus., no. 235183, from Bald Mountain, is referable to washingtoni on the basis of cranial characters but all the other adult and subadult specimens examined from Chelan and Okanogan counties are nearer nevaden- sis on the basis of cranial characters. Indeed, some show no ap- proach to washingtoni in cranial characters. As might be expected on geographic grounds, tlie specimen from Easton, U. S. Nat. Mus., male subadult, no. 116870, shows approach to washingtoni. This is true of the coloration of the hind limbs, small size of tlie tympanic bullae, and relatively greater length of the preorbital part of the skull. However, the greater width of the light color of the under- parts and relatively great breadth across the mastoid processes and zygomatic arches are points of agreement with nevadensis. Simi- larly, a series of 7 specimens from the Entait River, 20 miles above its mouth, in tone of color is nearer to washingtoni, as is one of the two skulls of adult males in length of the preorbital region. How- ever, in greater breadth of the skull otherwise, and in the relatively great width of the light color of the underparts, the animals are nearer to nevadensis, to which they are here referred. Some of MUSTELA FRENATA 287 these characters mentioned above in which departure is shown from typical nevadensis are characters that show approach to altifrontalis. This is especially true of the more intense coloration and restriction of the color of the underparts. Complete inter gradation with effera is shown by specimens from southern Oregon. The change from small effera to the larger nevad- ensis here is gradual; consequently in northeastern California and southern Oregon the size increases gradually to the northward. Specimens showing complete intergradation with oregonensis and saturata are wanting. However, one specimen from Crescent Lake suggests oregonensis in having near ( 18 ) apricot yellow underparts such as occur frequently in oregonensis. Also some specimens from northern California approach saturata in having the color of the underparts reduced in the extent to which it reaches out on the under side of the tail. This fact and the consideration that the two races are less diflFerent from one another than are otlier kinds which defi- nitely are known to intergrade leave no doubt but that material from the intervening localities would show complete intergradation. Intergradation between nevadensis and mimda is indicated by specimens from South Yolla Bolly Mountain, Trinity County, which are commented on at greater length in the account of M. f. munda. M. f. inyoensis is so closely related to nevadensis as to leave no doubt tliat specimens from suitable localities will show actual in- tergradation. That intergradation occurs directly with the bridled weasel of the interior valleys of California, M. /. xanthogenys, is shown by specimens from along the west-facing flank of the south- ern part of the Sierra Nevada. Probably intergradation occurs all along the Sierra Nevada on the western slope but specimens are lacking to show this. Weasels are known to occur in the foothill territory and the lesser attention given to this region by mammal collectors than to the higher parts of the mountains may explain the lack of preserved specimens. Individual specimens, here re- ferred to nevadensis, but, showing varying degrees of approach to xanthogenys are as follows: A female from Hume; a male and a female from 8000 feet elevation, Monache Meadows; a male from 9800 feet elevation on the east fork of the Kaweah River; and 7 specimens, probably one family, from one-half mile south of Min- eral King, 7850 feet. Of the specimens from 7850 feet, the adult male has no Hght facial markings and the head is only slightly darker than the back. The adult female has much restricted, light facial markings and the intervening areas are darker than in the male. The five juveniles trapped in the same burrow as the female, 288 University of Kansas Publs., Mus. Nat. Hist. each has more extensive light facial markings than the adult female although the area of this varies from only slightly more than in the female to as much as in typical specimens of xanthogenys. Also, the dark color of the head in these five specimens averages darker than in nevadensis and more as in weasels to the southwestvvard es- pecially latirostra. One of the five juveniles is lighter colored over all of the upper parts than nevadensis and is suggestive of xantho- genys in this respect. Finally, the adult male has on the underparts small spots of ochraceous orange suggestive of latirostra and some individuals of pulchra. No. 30655 42628, U. S. Nat. Mus., taken on Mount Whitney, also shows white facial markings and some other features of the valley-inhabiting xanthogenys. A suggestion of intergradation with arizonensis is furnished by specimens, referred to that race, from Springerville and the Kaibab Plateau. No speci- mens happen to be available from the region in which intergrada- tion would be expected between nevadensis and neomexicana. Since neomexicana and arizonensis intergrade it is probable that nevadensis also will be found to intergrade with neomexicana. In summary, nevadensis is judged to intergrade with each of the sub- species of Mtistela frenata whose range adjoins that of nevadensis. This subspecies is remarkably free from injury to the frontal sinuses such as result from the presence of parasites. In 98 adults from Oregon, California, Nevada, and Colorado, no malformation was noted. Only 1 of the 26 specimens from Washington was mal- formed and it was an intergrade with washingtoni. The single adult from New Mexico was diseased, as were 3 of the 6 from British Columbia, 1 of the 20 from Idaho, and 1 of the 7 from Utah. Specimens examined. — Total number, 568, arranged alphabetically by provinces and states and from north to south by counties in each state. Unless otherwise indicated specimens are in the collection of the United States Na- tional Museum. Arizona. Apache Co.: 15 mi. E Luka Chu Kai Navajo School, 8000 ft., 2. British Columbia. Monte Cr., 20 mi. E Kamloops, 1'; Sicamous, 2; Okana- gan, 18 {T, &, V, V'); Monashee Pass, 1°; Swan Lal'pes, 55 (43-81) per cent of greatest width of color of upper parts. Black tip of tail in same series of males averaging 58 (53-63) mm. long; thus averaging longer than hind foot and 33 per cent of length of tail-vertebrae. Skull and teeth. — Male (based on 6 ads., t>'pe and 5 topotypes): See measurements and plates 22-24. As described in M. f. nigriauris except that: Weight (6 ads.), 5.3(4.5-6.1) grams; basilar length, 47.6(46.0-48.6); (one skull, no. 335, with postorbital breadth more than distance between posterior borders of P4 and P2); interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2 to 3/2 (includ- ing 13) upper incisors; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than rostrum. Female: Adult skull of typical female not seen. As compared with the skull of the type specimen of inyoensis, skulls of adult males of pulchra are larger throughout, relatively broader, especially in the preorbital part of the skull, have more inflated tympanic bullae, and are less convex in dorsal outline. Comparison of the skull with that of latirostra has been made in discussion of that subspecies. Comparison of skulls of adult males of nigriauris and pulchra shows that those of pulchra average larger in every measurement taken except those of ml, Ml, P4, and deptli of skull at posterior borders of upper molars. The basilar length is only slightly more and it follows that, relative to this length, other measurements of the skull are relatively, as well as actually, larger. In no one measurement is there an entire lack of overlap, but the skulls of adult males, and probably adult females, may be dis- tinguished from those of nigriauris by the combination of the fol- lowing mentioned, average difi^erences: Tympanic bullae larger in each of three dimensions; preorbital and interorbital parts of skull broader and notably heavier; interorbital breadth greater; zygo- matic arches more expanded laterally; mastoid processes more prominent. As compared with xanthogenys, differences of similar nature, but of greater degree, distinguish pulchra. As compared with those of nevadensis (represented by specimens from Mono 330 University of Kansas Fuels., Mus. Nat. Hist. Co., Calif.), skulls of adult males of pulchra average larger in every measurement taken and no overlap exists in basilar length, orbito- nasal length, mastoid breadth, zygomatic breadth, length of tym- panic bulla, or depth of skull at either the anterior margin of the basioccipital or at the posterior margins of the upper molars. Rel- atively, the preorbital portion is about the same size in the two forms. Remarks. — The best material of this big weasel was obtained in 1910 and 1911 by John Wimmer and forwarded to the California Academy of Sciences through John R. Rowley, although in 1905, one specimen had been obtained by A. S. Bunnell for the collec- tions of the United States Bureau of Biological Survey, another by J. Grinnell for the Museum of Vertebrate Zoology in 1912, and in 1933, another by L. M. Huey, for the San Diego Society of Nat- ural History. The males from the type locality are relatively uniform in size and shape of skull. The one exception is no. 137935, U. S. Nat. Mus., slightly younger than the others. Its skull is relatively more slender than any of the others and does not display several of the differential characters. The male, no. 127566, U. S. Nat. Mus., from Alila (=: Earlimart) is intermediate in cranial features between pulchra and xanthogenys as known from specimens taken in the vicinity of Fresno. The skull of the female, no. 127565, from the same place, is too young to provide diagnostic characters. Since the skull of an adult female of topotypical pulchra is unknown, doubt attaches to the identification of the adult, female specimen, no. 115895, U. S. Nat. Mus., from Delano. It has a relatively broad skull in comparison with the adult female of xanthogenys from Los Banos. The adult female, no. 9998, San Diego Soc. Nat. Hist., from 2 mi. SW Simmler, shows approach to nigriauris in slightly reduced size. The skin alone from Coahnga, a male, taken on April 10, 1935, measures 462, 179, 47. The adult female, with crushed skull, from 4 miles east of Coalinga, measures 350, 129, 40. In size, these specimens agree better with pulchra than with xanthogenys. The skin alone from 3 miles south of Coalinga is un- sexed and without external measurements. Skulls of adults from Coa- linga are needed to permit of more positive identification of the sub- species found there. The female from 4 miles east of Coalinga, taken on February 21, 1936, is in process of molt on the under- parts, and the longer hair which is near (20') Naples Yellow con- trasts strongly with the incoming shorter hair which is near (10 c) ' MUSTELA FRENATA 331 Salmon-Orange. The skin alone, no. 16270, Mus. Vert. Zool., from Isabella, was made up from a decayed animal and is of but little use. It is referred to pulchra purely because of geographic near- ness of Isabella to the type locality of pulchra. The most that can be told from the specimen is that it is a relatively light-colored, bridled weasel. The fact that the color is slightly darker than in pulchra may or may not indicate intergradation with nevadensis. No. 54103/41042, U. S. Nat. Mus., consisting of crushed bits of skull and the skin of the head, is from Willow Spring, Kern County. This marginal locality is really in the Mojave Desert rather than in the San Joaquin Valley. The light color of the skin of the head suggests pulchra, but it is realized that a complete specimen might show the animal there to be unHke pulchra. None of the skulls shows evidence of having had the frontal sinuses infested by parasites. Specimens examined. — Total number, 18, listed by counties from north to south. Unless otherwise indicated, specimens are in the Museum of Verte- brate Zoology. California. Fresno Co.: Coalinga, 1*; 4 mi. E Coalinga, 1; 3 mi. S Coalinga, 1'. Tulare County. Alila ( = Earlimart ) , 2\ Kern County: Delano, 1'; But- tonwillow, 9 (6% 2'); Isabella, 1; Willow Spring, V. San Luis Obispo County: 2 mi. SW Simmler, 1'. Mustela (renata inyoensis Hall Long-tailed Weasel Plates 22, 23 and 24 Mustela f renata inyoensis Hall, Carnegie Instit. Washington Publ. 473:99, November 20, 1936. Putorius xanthogenys, Merriam, N. Amer. Fauna, 11:25, June 30, 1896 (part). Mustela xanthogenys xanthogenys. Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912. Type. — Male, adult, skull (with skeleton) and skin; no. 25907, Mus. Vert. Zool; Carl Walter's Ranch, 2 mi. N Independence, Inyo County, California; June 26, 1917; obtained by A. C. Shelton; original no. 3143. The skull (plates 22-24) is complete and unbroken. All teeth are present and entire. The skin is well made and in good condition. Range.— From 3700 feet (Lone Pine) to at least 4000 feet (Alvord); Lower Sonoran Life-zone of the floor of Owens Valley in Inyo County, Cali- fornia. See figures 29 and 30 on pages 221 and 314. Characters for ready recognition. — Differs from M. f. nevadensis in presence of white facial markings; from M. f. pulchra in near (I) Brussels Brown rather than near (16 /) Buckthorn Brown to near (h) Yellow Ocher color of upper parts and basilar length of less than 45 in males; from M. f. lati- 1. United States National Museum 3. San Diego Society of Natural History 2. California Academy of Sciences 4. Collection of J. Arnold 332 University of Kansas Publs., Mus. Nat. Hist. rostra in brownish rather than blackish color of inside of ear and orbitonasal length of more than 15. Description. — Size. — Male: Two adults, the type specimen and no. 25392/32805, measure, respectively, as follows: Total length, 423 and 390; length of tail, 170 and 145; length of hind foot, 42 and 44. Tail is 67 and 59 per cent as long as head and body. Length of hind foot less than basal length. Female: No. 12400, Field Mus. Nat. Hist., which is young, has the follow- ing measurements: Total length, 390; length of tail, 150; length of hind foot, 39. Tail is 63 per cent as long as head and body. Length of hind foot less than basal length. The differences in external measurements between the two sexes, as represented by the male type specimen and by the young female, are: Total length, 33, length of tail, 20; length of hind foot, 3. Externals. — Longest facial vibrissae black or dark brown and reaching beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) slightly less than shown in figure 19. Color. — Large spot between eyes, band confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, upper throat, chin, lower hps and in some specimens part or all of upper hps white; patch between eyes and bars in front of ears tinged with some shade of yellowish in one specimen; dark spot posterior to each angle of mouth present in four of five specimens; tip of tail black; remainder of upper parts, in summer, near (l) Brussels Brown or tones 1 to 2 of Raw Umber of Oberthiir and Dauthenay, pi. 301; sfightly darker brown on forehead, nose and about eyes. In winter near (/') Snuff Brown or lighter than Brussels Brown with a smoked effect. Underparts Buff-Yellow, winter and summer. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists and on medial sides of hind legs over antiplantar faces of toes. Least width of color of underparts averaging, in 5 available specimens 34 (24-42) per cent of greatest width of color of upper parts. Black tip of tail, in two adult males, averaging 53 (45 and 60) mm. Thus longer than hind foot and averaging 34 per cent of length of tail-vertebrae. Skull and teeth. — Male (based on the type): See measurements and plates 22-24. As described in M. f. nigriauris except that: Weight, 4.4 grams; basilar length, 44.7; postorbital breadth not less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; length of tympanic bulla less than length of lower molar and premolar tooth- row. Female: Adult unknown. Compared with the skull of the male of nevadensis, no single dif- ference not covered by individual variation in nevadensis has been detected. Selected differences of inyoensis in comparison with latirostra are larger size, less inflated tympanic bullae and relative narrowness of the postorbital, interorbital and preorbital parts of MUSTELA FRENATA 333 the skull. Comparison of the skull with that of M. /. pulchra is made in the account of that subspecies. Remarks. — Although two specimens of this subspecies were taken during the Death Valley Survey conducted by Dr. C. Hart Merriam, only three additional individuals are known to have been saved as study specimens since that time. M. /. inyoensis as now known may be thought of as closely similar to M. /. nevadensis except for the presence of well-developed white facial markings like those found in the weasels of the San Joaquin Valley and coastal region of California south of San Francisco Bay. The nonwhite areas of the head are almost the same color as the back and not distinctly blackish as in M. f. latirostra and M. f. nigri- auris. The one specimen in the winter coat, no. 25392/32805, U. S. Nat. Mus., from Lone Pine, is brown rather than white. The brown has the pale smoke-tinge common in the winter pelage of subspecies whose members are either brown or white in winter. The range of this subspecies is thought to include the floor and lower elevations of Owens Valley although it may occur in limited numbers south- westward along the base of the Sierra Nevada and through the mountains in places of low elevation like Walker Pass its range may meet that of pulchra. The type specimen was taken in an alfalfa field by ranch hands, who, according to A. C. Shelton (MS), stated tliat tlie species was common at the type locality. None of the five specimens shows infestation of the frontal sinuses by parasites. Specimens examined. — Total number, 5, listed by localities from north to south. California. Inyo Co.: Alvord, 4000 ft., 1 (U. S. Nat. Mus.); 2 mi. N Inde- pendence, 1 (Mus. Vert. Zoo!.); Lone Pine, 3 (2 in Field Mus. Nat. Hist, and 1 in U. S. Nat. Mus.). Mustela frenata neomexicana (Barber and Cockerell) Long-tailed Weasel Plates 1, 22, 23, 24, 34, 35 and 36 Putorius frenatus neomexicantis Barber and Cockerell, Proc. Acad. Nat. Sci. Philadelphia, 1898:188; Lantz, Trans. Kansas Acad. Sci., 19:178, 1905. Mustela frenata neomexicana. Miller, U. S. Nat. Mus. Bull., 79:100, Decem- ber 31, 1912; Bailey, Animal Life of Carlsbad Cavern, p. 97, 1928; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936. Mustela frenatus neomexicanus, Bailey, N. Amer. Fauna, 35:19, September 5, 1913. Type. — Male, adult, skull and skin; no. 10475, Mus. Comp. Zool.; Arm- strongs Lake, Mesilla Park, Dona Ana County, New Mexico; February 1, 1898; obtained by A. C. Tryson; original no. 58 of C. M. Barber. 334 University of Kansas Publs., Mus. Nat. Hist. The skull is imperfectly cleaned but unbroken. The right upper incisors, right P2 and left p3 are broken away. The skin is indifferently stuffed but in a good state of preservation except that the distal part of the tail is missing. The animal's coat is ragged, and this imperfect appearance is heightened by injury to the posterior part of the body, probably at the time of capture. Range. — From 3800 feet (type locaHty) to 9000 feet (Cloudcroft, N. Mex.); Upper Sonoran and Lower Sonoran life-zones of northern Mexico, southeastern Arizona, New Mexico and western Texas, panhandle of Oklahoma, southeastern Colorado and southwestern Kansas. See figure 29 on page 221. Characters for ready recognition. — Differs from M. f. frenata and M, f. texensis by Buckthorn Brown rather than Brussels Brown color of upper parts, mastoid breadth of adult males ordinarily more, rather than less, than post- palatal length; from M. /. leucoparia by Buckthorn Brown rather than Argus Brown color of upper parts, distance from anterior margin of tympanic bulla to foramen ovale less, rather than more, than four-fifths height of tympanic bulla; from M. f. arizonensis and M. f. nevadensis by Buckthorn Brown, rather tlian near ( 14 n ) Brussels Brown or, in winter, white color of upper parts, in presence of white frontal spot continuous with color of underparts, in basilar length of more than 46 mm. in males and 40 mm. in females; from M. /. longicauda by Bucktliom Brown rather than near (h) Clay Color of upper parts, by presence of white facial markings on Argus Brown head, and by length of tooth-rows amounting to less than 37 per cent of basilar length; from M. /. primulina by Buckthorn Brown rather than Brussels Brown color of upper parts, in presence of white frontal spot and broad white bands on side of head, in anteriorly truncate rather than anterolaterally rounded bullae and zygomatic breadth of more tlian 30 in males and 24 in females. Description. — Size. — Male: The type specimen (see Barber and Cockerell, 1898:188) measured: Total length, 500; length of tail, 205; length of hind foot, 50. Tail 70 per cent as long as head and body. Length of hind foot less than basal length. Female: No. 21779 from Tombstone, Arizona, measured: Total length, 419; length of tail, 165; length of hind foot from dried skin, 41 (probably 43 in flesh). Tail 65 per cent as long as head and body. Length of hind foot less than basal length. The average differences in external measurements of the two sexes, as known from these two individuals, are: Total lengtli, 81; length of tail, 40; length of hind foot, 7. Compared with M. /. frenata, the size, proportions of parts and difference in size of the two sexes, appears to be about the same. Externals. — Longest facial vibrissae colored like upper parts [in the type specimen some of tlie "long bristles of the upper lip" are white as pointed out by Barber and Cockerell {op. ci^: 188)] and extending beyond ear; carpal vibrissae colored like underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 20. Color. — Broad white bands on sides of head, extending anterodorsally an- terior to each ear, confluent with white spot between eyes and with color of underparts; posterior half or all of each upper hp edged with white; usually few white hairs on top of head between ears; remainder of top of head near Argus Brown of Ridgway and Chocolate, tone 4, of Oberthiir and Dauthenay; MUSTELA FRENATA 335 dark spot posterior to each angle of mouth usually absent; tip of tail black; remainder of upper parts varying, in different specimens, from Buckthorn Brown to Dresden Brown of Ridgway, and Brown Pink ( tones 3 to 4, pi. 297, of Oberthiir and Dauthenay); underparts Antimony Yellow or near (c) Warm BuflF of Ridgway, and Brown Pink (tone 1, pi. 297, of Oberthiir and Dauth- enay); color of underparts extends distally on legs over forefeet and hind feet. Least width of color of underparts averaging 46 (41-5.5) per cent of greatest width of color of upper parts; black tip of tail 35 to 45 mm. long in females; 43 to 68 mm. long in males and averaging 21 (20-36) per cent as long as tail-vertebrae. No specimen of this subspecies in the white winter coat has been seen. Animals taken in midwinter are available from Mesilla Park, Willcox, and 10 miles east of Roswell. From M. /. frenata, neomexicana differs in: upper parts and underparts much Hghter colored; white facial markings more extensive; color of under- parts more extended onto feet. From M. /. leucoparia, neomexicana differs aS follows: above and below, much hghter colored, but white facial markings less extensive and color of underparts less extended onto feet and legs. Skull and teeth. — Male (based on adults: the type; no. 131582 from Ber- ino. New Mexico; and no. 1485 from Seward Co., Kansas): See measurements and plates 22-24. As described in Mustela frenata frenata except that: Weight, 6.2 (4.9 and 7.5); basilar length, 49.3 (48 and 50.5); mastoid breadth more than postpalatal length; least width of palate less than length of P4; anterior margin of masseteric fossa directly below m2 or heel of ml. Female (based on three adults): See measurements and plates 34-36. As described in Mustela frenata frenata except that: Weight, 3.1 (2.6-3.5) grams; basilar length, 42.7 (40.8-45.5); zygomatic breadth less tlian distance between condylar foramen and Ml and more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla. The skull of the female averages 50 per cent hghter than that of the male. As compared with the skull of the male of M. /. frenata, that of neomexicana is decidedly more angular and ridged. The post- orbital constriction is narrower, the mastoid breadth greater (it is less than the postpalatal length in some subadult males), the sagit- tal crest much higher with impressions of the temporal and mas- seter muscles carried farther forward on the frontals, rostrum shorter and tympanic bullae wider and more inflated. Similar, though less marked, differences exist between the females. As compared with M. f. leucoparia and perotae, the same differences as noted above between frenata and neomexicana exist. In addi- tion the tympanic bullae are so far removed from the foramen ovale that the distance from the anterior end of each bulla to the foramen ovale, instead of being less than the height of tympanic bullae, is in leucoparia more than four-fifths this height and in perotae more than the entire height. Also, in perotae, the squamosal, anterior to each tympanic bulla, is ventrally convex rather than ventrally 336 UNnERsiTY OF Kansas Publs., Mus. Nat. Hist. concave as in neomexicana. Compared with M, /. longicatida, neo- mexicana is relatively narrower in the interorbital region, has rela- tively shorter tooth-rows, a V-shaped rather than a U-shaped inter- pterygoid space and in males has the interorbital region flat rather than convex and the sagittal crest is higher. The same differences are to be noted in comparison with nevadensis but here the differ- ence in relative length of tooth-row is less. The same differences exist also in comparison with M. f. arizonensis except that its inter- orbital breadth, relative to the rest of the skull, is about the same. Difference in size is especially marked here; even females of neo- mexicana average larger than males of arizonensis. Remarks. — When Barber and Cockerell named this subspecies in 1898, they had three specimens. Only two others are known to have been taken before this time. These are a skeleton, without corresponding skin, taken at Lozier, Texas, in 1890 by Wm. Lloyd, and no. 21779/36482, U. S. Nat. Mus., taken on April 6, 1893, by R. D. Lusk at Tombstone, Arizona. On the back of a label re- cently attached to the last mentioned specimen the name C. K. Worthen appears and probably signifies that the specimen was purchased from this dealer in vertebrate specimens. M. f. neomexicana has a large geographic range. The old male from Liberal, Seward County, Kansas, extends the known range far to the northeast. Geographically, this occurrence is logical for the southwestern desertlike conditions extend to this part of Kan- sas. Probably the subspecies occurs in southeastern Colorado and in the panhandle of Oklahoma where conditions are similar. Bailey (1905:198) lists neomexicana as a member of the mammalian fauna of Texas. As stated by him (loc. cit. :19S) this inclusion is based on geographic grounds and not on actual specimens. Strecker (1926:13) also includes neomexicana in his list of Texas mammals but writes me, under date of January 9, 1928, that "I included Mustela frenata neomexicana as a Texas mammal on the strength of its being mentioned by Bailey. . . ." On better ground, Bailey (1928:97) lists the subspecies as occurring in southeastern New Mexico at Carlsbad Cavern. However, Bailey (loc. cit.) knew of the existence of weasels just below El Paso and at Lang- try, Texas. An unsexed skeleton, no. 167891, in the United States National Museum, from Lozier, Texas, is not certainly identifiable to subspecies. If, as I think, the animal is a female, its skull is intermediate between that of frenata and neomexicana although when all features are considered it is seen to be nearest the latter. MUSTELA FRENATA 33/ The large size (basilar length of 46.5 mm.) may reflect some rela- tionship to texensis. The field notes of the collector furnished me by Dr. H. H. T. Jackson (MS), describe the color as brownish yellow above and sulphur below. The admission of this sub- species to the list of mammals of Texas is made certain by the fe- male (no. 1572, Texas Cooperative Research Collection) taken on July 28, 1940, U mi. NW Kent, Texas, by C. E. Scull. The skull alone from Durango (City of), extends the known range far to the south. This skull is typical of neomexicana. Skins from the same place would be especially interesting as show- ing the approach, if any, in color, of neomexicana to M. /. leu- coparia. Mr. D. D. Stone of Casa Grande, Arizona, writes, under date of February 2, 1927, that a weasel was seen by an acquaintance of his in a field near Chandler, Maricopa County, Arizona. Probably this was neomexicana. If so, its range extends much farther west than collected specimens show. Actual intergradation with M. /. frenata is not shown by the material at hand. The two females from Albuquerque, although typically neomexicana as regards color, have smaller, less prom- inently ridged skulls than females of neomexicana of the same age from farther south and approach M. /. nevadensis. Probably the geographic ranges of M. f. neomexicana and M. /. latirostra do not meet; the only evidence of the existence of weasels in all of the large area, comprising western Arizona and the deserts of eastern CaUfornia, which intervenes between the ranges of the two subspecies is the skull of a young individual, no. 68842, Mus. Vert. Zool., from 10 miles northeast of Bard, Imperial County, Cali- fornia. There, on December 29, 1932, Jack C. vonBloeker, Jr., retrieved the weathered skull with some of the vertebrae attached, from the top of a woodrat's nest beneath a mesquite tree near the west bank of the Colorado River. The idea that the carcass may have been washed down the river from far upstream gains no support from a comparison of the speci- men itself for the tympanic bullae are larger than in nevadensis and the skull is larger than the largest males seen of arizonensis, the two up-river races. On the basis of size the skull could be either a male of latirostra or a female of neomexicana. These two subspecies, like arizonensis and the skull in question, have much inflated bullae. However, the immaturity of the specimen con- ceals any other diagnostic cranial features, and prevents referring it certainly to either neomexicana or latirostra. In any event the 338 University of Kansas Publs., Mus. Nat. Hist. specimen provides no evidence of intergradation between the two forms last mentioned. Speculating on its identity, I should say that it might be either an intergrade between arizonensis and nevadensis, from southern Utah or northwestern Arizona, or a mem- ber of an unnamed race resident in the lower part of the valley of the Colorado River. Whereas M. /. panamensis and M. /. aureoventris are the dark- est-colored weasels and occur in regions of heavy rainfall, M. f. neomexicana is the lightest-colored American weasel and occurs in an extremely arid region where the rainfall and humidity are slight. According to Barber and Cockerell (1898:189) "The type speci- men was shot in the grass on the shore of Armstrong's Lake. . . ." Bailey (1928:97) found the tracks of one of these ani- mals "in the great pit at the west entrance to" Carlsbad Cavern and supposes they "hunt the cave walls for mice and other small game". Data on the label attached to no. 230973 states that the specimen was taken, two miles west of Willcox, Arizona, in a prairie dog town. Only two of the 23 skulls show evidence of infestation of the frontal sinuses by parasites. Specimens examined — Total number, 28, arranged alphabetically by states and from north to south by counties in each state. Unless otherwise indicated specimens are in the United States National Museum. Arizona. Graham Co.: Safford, 1. Cochise Co.: 2 mi. W Willcox, 1; Will- cox, 1; 8000 ft., Chiricahua Mts., 1; 6000 ft., Pinery Canyon, Chiricahua Mts., 1'; Tombstone, 1; Sulphur Spring Valley, l^ Durango. "Durango City," 1. Kansas. Seward Co.: Liberal, 1'. New Mexico. Bernalillo Co.: 3 mi. NW Albuquerque, 2. Lincoln Co.: 7800 ft., South Fork Eagle Creek, White Mts., 1. Chaves County: Pecos River, 10 mi. E Roswel], 8'; Dexter, 1'. Otero Co.: Cloudcroft, 9000 ft., V. Dona Ana Co.: Mesilla Park, 2 (V, 1°); Berino, 2. Texas. Culberson County: IVi mi. NW Kent, 1'. Terrel Co.: Lozier, 1. Mustela frenata texensis Hall Long-tailed Weasel Plates 22, 23 and 24 Mustela frenata texensis Hall, Carnegie Instit. Washington Publ. 473:99, November 20, 1936. Mustela frenata, Strecker, The Baylor Bull., 27:14, September, 1924. Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926 (part). 1. Collection of Lloye H. Miller 4. Museum of Comparative Zoology 2. Mus. Vert. Zobl., Univ. California 5. British Museum of Natxu-al History 3. Univ. Kansas Mus. Nat. History 6. Texas Co-operative Research Collection MUSTELA FRENATA 339 Type. — Male, adult, skull with skin of head, neck and tail; no. 14821, Amer. Mus. Nat. Hist.; Kerr County, Texas; September 17, 1897; obtained by H. P. Attwater. The skull (plates 22-24) and dentition are complete and unbroken. The preserved parts of the skin are not stuffed. Range. — Lower Sonoran and possibly Upper Sonoran life-zones of central Texas. See figure 29 on page 221. Characters for ready recognition. — Differs from Mustela frenata arthuri in possessing white facial markings and postorbital breadth less than distance be- tween posterior borders of P4 and P2; from M. /. frenata in larger size of body and skull, the basilar length of which in adult males is more than 52.5; from M. f. neomexicana in Brussels Brown rather than Buckthorn Brown color of upper parts and basilar length of skull more than 52.5. Description. — Size. — Male: Measurements taken from the dried skins of a young male, no. 15476, Mus. Comp. Zool., from Kerr County, Texas, and a subadult male, no. 2017, Baylor Univ. Mus., from 5 mi. N Waco, Texas, are, respectively, as follows: Total length, 600 and more than 510; length of tail, 200 and 225; length of hind foot, 52 and 52. Female: Skins unknown. Externals. — As described in Mustela frenata frenata. Color. — As described in Mustela frenata frenata. Skull and teeth- — Adult male: See measurements and plates 22-24. As described in Mustela frenata frenata except that: Weight, 8 grams; basilar length 54; least width of palate less than length of P4; anterior margin of mas- seteric fossa anterior to middle of m2. Female: Skull unknown. Remarks. — The type specimen, taken by the veteran collector of Texan mammals, H. P. Attwater, appears to have been the first one of these animals to find its way into the collection of any museum or naturalist. The second specimen from Kerr County was secured by, or through, the well-known commercial collector, F. B. Armstrong. Two trade skins, from Kerr County, taken on December 10, 1938, are in the Texas Cooperative Research Collec- tion, as is also the skeleton of a young animal from Fredericksburg. The two other specimens from McLennan County (both males contrary to the statement of Strecker, 1924:14), owe their preser- vation to the alertness of John K. Strecker, Curator of the Baylor University Museum, who has given a complete account of their history. The range of this subspecies is thought to include much of cen- tral Texas. The preserved parts of the skin of the type specimen show it to have been generally large. The part of the tail preserved measures 226 millimeters and the skin of the head and neck is correspond- ingly large. The skin alone, no. 427, from near Waco, Texas, has. 340 University of Kansas Publs., Mus. Nat. Hist, as now stuffed, a body 365 millimeters long. Individuals of this race attain larger size than those of any other American member of the subgenus Mustela with the possible exception of Mustela frenata macrophonius from Veracruz, Mexico. In addition to large size, texensis and macrophonius are analogous in that each has a small geographic range at the northern end of an extensive range of its similarly colored southern relative from which it differs mainly in size. Each of the two groups, goldmani and macro- phonius on the one hand and perotae, frenata and texensis on the other, has relatively uniform color, color pattern and body propor- tions over a large region but at its northern extremity develops a "giant" population, M. /. macrophonius and M. /. texensis, respec- tively. The skull of the type specimen of M. /. texensis is the larg- est one seen of any American weasel. The type specimen of M. f. macrophonius has a basilar length that is greater by one-tenth of a millimeter but in every other measurement taken the skull of M . /. texensis is the larger. Its weight, 8 grams, also shows it to be larger. The broad, white bands in front of the ears are confluent with the white patch between the eyes on both sides in two specimens and on one side only in one other specimen. A white patch be- tween the ears is present in four specimens. The dark spot at each angle of the mouth is absent on both sides in four specimens and on one side only in one other specimen. Thus out of a possible twelve cases, the broad bands in front of the ears are confluent with the spot between the eyes in five cases. Four of the six specimens have a white spot between the ears. The dark spot at each angle of the mouth is present three out of a possible twelve times. The skull of no. 2017, from five miles north of Waco, is smaller than either of the two skulls seen from Kerr County and in this respect approaches M. f. frenata. There is no actual evidence of intergradation with any other subspecies but intergradation prob- ably does take place with M. /. neomexicana and possibly with M. /. arthuri and M. f. primulina. Strecker (1924:14) remarks that of the two specimens obtained near Waco, one was taken in a trap baited for mink and the other was shot in a hen house. None of the four skulls had the frontal sinuses infested with parasites. Specimens examined. — Total number, 7, arranged by counties from north to south. Texas. McLennan Co.: 5 mi. N Waco, 1 (Baylor Univ. Mus.); Erath, 1 (Baylor Univ. Mus.). Gillespie County. Fredericksburg, 1 (Texas Cooper- ative Research Collection), Kerr Co.: 4 ( 1 in Mus. Comp. Zool.; 1 in Amer, Mus. Nat. Hist.; and 2 trade skins in Texas Cooperative Research Collection). MUSTELA FRENATA 341 Mustela frennta frenata Lichtenstein Long-tailed Weasel Plates 1, 22, 23, 24, 36, 37, 38 and 40 Mustela frenata Lichtenstein, Darstellung neuer oder wenig bekannter Siiugethier, 1832, pi. 42, and corresponding text, unpaged; Seton, Lives of game animals, 2:576, 1929. Mustela brasilietisis Sevastianoff, Mem. de I'Acad. Imp. Sci. St. Petersburg, 4:356-363, tab. 4, 1813, name on plate only, the description being in the text (not of Gmelin, 1788); Gray, Proc. Zool. Soc. London, 1865:114. Putorius frenatus, Baird Mamms. N. Amer., p. 173, 1858; Merriam, N. Amer. Fauna, 11:26, pi. 3, figs. 1, la, lb, June 30, 1896; Bailey, N. Amer. Fauna, 25:198, October 24, 1905. Putorius (Gale) brasiliensis aequatorialis Coues, Fur-bearing animals, p. 142, 1877, part ? ("merely as a substitute for Gray's [supposedly] pre- occupied name" that is, aureoventris). Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part). Putorius mexicanus Coues, Fur-bearing animals, p. 142, 1877, [nomen nudum, cited by Coues in synonymy as "Putorius mexicanus, Berlandier, MSS. ic. ined. 4 (Tamaulipas and Matamoras)"]. Putorius brasiliensis frenata, Allen, Bull. Amer. Mus. Nat. Hist., 3:219, April 17, 1891. Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 6:197, May 31, 1894; Bangs, Proc. Biol. Soc. Washington, 10:9, February 25, 1896; Allen, Bull. Amer. Mus. Nat. Hist., 8:74, April 22, 1896. Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936. Type. — Female, adult, skull and skin; no. 991, Berlin Zool. Mus., Mexico City, Mexico; June, 1829; obtained by F. Deppe. The specimen once mounted, now is remade into a study skin and lacks the distal part of the tail. The skull ( plates 36-38, 40 ) lacks the basicranial region. Range. — Altitudinally, sea level ( Brownsville, Texas ) to 7600 feet ( Tlalpam, Mexico ) ; from southern Texas as far south as Mexico City; Lower Sonoran to at least Transition life-zone. See figure 29 on page 221. Characters for ready recognition — Difi^ers from M. f. perotae in nonexten- sion of blackish over anterior fourth of neck, least width of color of underparts more than 37 per cent of greatest width of color of upper parts; height of tympanic bulla more than distance from its anterior margin to foramen ovale; from M. f. leucoparia by restricted white facial markings that cover less than half surface of head in front of ears, by nonextension of black of head onto anterior half of neck and by wdder (more than 7.8) tympanic bullae; from M. f. neomexicana by Brussels Brown rather than Buckthorn Brown color of upper parts and mastoid breadth less than postpalatal length; from M. f. texensis by smaller size of body and skull (basilar length in adult males less than 52.5); from M . /. arthuri by white facial markings and postorbital breadth less than distance between posterior borders of P4 and P2; from M. /. tropicalis by non- extension of blackish over anterior fourth of neck, least width of underparts more than 37 per cent of greatest width of upper parts, postorbital breadth of adult males less than distance between posterior borders of P4 and P2. Description. — Size. — Male: Fifteen adults and subadults from BrovvTisville, Texas, yield average and extreme measurements as follows: Total length, 485 (430-556); length of tail, 202 (165-250); length of hind foot, 48 (40-55). 342 University of Kansas Publs., Mus. Nat. Hist. Averages believed to be reliable but extremes probably are not. Tail averages 71 per cent as long as head and body. Length of hind foot less than basal length. Corresponding measurements of an adult male (topotype, no. 50826) from Tlalpam, Mexico, are: 505, 203, 53. Another adult male, from Miqui- huana, Tamaulipas, Mexico, measures: 520, 215, 52. Female: Six adults, subadults and yoimg from Brownsville, Texas, yield average and extreme measurements as follows: Total length, 420 (362- 456); length of tail, 173 (126-200); length of hind foot, 41 (40-46). Tail averages 70 per cent as long as head and body. Length of hind foot more (vtath possible exception of no. 36362/48732 U. S. Nat. Mus.) than basal length. The average differences in external measurements of the two sexes are: Total length, 65; length of tail, 29; length of liind foot, 7. Externals. — Longest facial vibrissae black and reaching beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 20. Color. — Spot between eyes, broad band, confluent with color of under- parts, on each side of head extending anterodorsally anterior to each ear, and posterior two-thirds to one-half of each upper hp, white; remainder of sides and top of head, posteriorly to line connecting posterior margins of ears, blackish; dark spot posterior to angle of mouth present on both sides in about half the specimens; tip of tail black; remainder of upper parts Brussels Brown; chin white; remainder of underparts near (16'a) Ochraceous-Buff (same color in juveniles and young), which color ex-tends distally on posterior sides of forelegs over forefeet and on medial sides of hind legs to feet and sometimes onto upper sides of toes. Least width of color of underparts averaging, in a series of seventeen males from Brownsville, Texas, 47 ( extremes 38-53) per cent of greatest width of color of upper parts. Black tip of tail, in same series, averaging 49 (extremes 40-55) mm. long, thus about equal to length of hind foot and averaging 24 per cent of length of tail-vertebrae. Skull and teeth. — Male (based on ten adults from Brownsville): See measurements and plates 22-24; weight (three adults, one topotype and two from Brownsville, Texas), 6.2 (5.3-7.2) grams; basilar length, 49.8 (48.2- 51.3); zygomatic breadth more than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoidal breadth less than postpalatal length; postorbital breadth less than length of upper premolars (less than distance between posterior borders of P4 and P2) and not greater (usually less) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostnmi less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 or 4 (including 13) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter (usually longer) than rostrum; anterior margin of masseteric fossa just behind m2. Female ( based on two adults from Brownsville, Texas ) : See measurements and plates 36-38, 40; weight, 3.4 (3.3-3.5) grams; basilar length (six, adult MUSTELA FRENATA 343 to young) 43.3 (41.3-47.3); 2ygomatic breadth more or less than distance between condylar foramen and Ml and more than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less than (about equal to) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than outside length of P4; tympanic buUa as far posterior to foramen ovale as width of 2 to 3/2 (includ- ing 13) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum. The skull of the female averages 45 per cent hghter than that of the aver- age male. Comparison of the skull with those of M. f. arthuri, tropicalis, perotae, leucoparia and neomexicana has been made in accounts of those subspecies. As compared with M. /. texensis (known only from males), the only difference detected is smaller size. Remarks. — As Merriam (1896:27) has said: "In 1813 a Russian naturalist, Sevastianoff, gave the name 'Mustela hrasiliensis to a weasel brought to St. Petersburg by Capt. A. J. Krusenstem on his return from a voyage around the world. The animal was said to have come from Brazil, but no definite locality was given." This name was long applied by many European naturalists to American weasels which had white facial markings, and several American naturalists adopted the name. However, Lichtenstein in 1832 ap- pHed the name Mustela frenata to the weasels of the vicinity of Mexico City and that name was used for bridled weasels from Mexico and the southwestern United States by most subsequent German writers and by several Americans. In 1896 Merriam (1896:27) showed that Sevastianoff's Mustela hrasiliensis, 1813, although probably the same as Mustela frenata, was preoccupied by Gmelin's Mustela hrasiliensis, 1788, applied to an otter and that Lichtenstein's name must be used as the next available one. Since that time, 1896, frenata has been the name applied to the large bridled-weasels of Texas and the high table land of Mexico south to Mexico City. It may be added that in 1937 search by the writer among the specimens and records at the Russian Academy of Sciences, in Leningrad, failed to reveal any trace of the type spec- imen of Sevastianoff's Mustela hrasiliensis. The geographic range of this subspecies is relatively large and, as might therefore be expected, specimens show geographic varia- tion. The specimens from Tlalpam, which Merriam {op. cit. :27) regards as topotypes, differ in certain respects from specimens from Texas. The skull of the adult male "topotype", no. 50826, differs 344 University of Kansas Publs., Mus. Nat. Hist. from any other adult male seen in that the basilar length, the length of the upper tooth-rows, the orbitonasal length, the ratio of the same to the basilar length, the mastoidal breadth, the zygomatic breadth, the depth of the skull at the posterior margins of the upper molars, and the length and breadth of Ml, are greater. The height of the tympanic bullae is less than the average height for these structures in more northern specimens. The specimens from Tlalpam have also larger external measurements than the average of more northern specimens. All of these features show an ap- proach to the subspecies of more southern distribution. On the other hand, the blackish of the head is not more intense or more extended posteriorly onto the neck than in specimens from Browns- ville, Texas. The skin, with skull crushed, no. 767, in the Paris Museum, from 3200 meters elevation near Toluca, does have the black color of the head extended 30 millimeters posteriorly to the ears. In this feature, and also in the extensively white face on which the white bar in front of each ear connects with the frontal spot, as well as with the color of the underparts, the specimen re- sembles leucoparia. Better material from the western part of the state of Mexico may show the range of leucoparia to extend east- ward almost or quite to Toluca. An adult male, taken on July 15 at Miquihuana, Tamaulipas, is unique in several respects. The top of its head is black, rather than blackish, and this color extends posteriorly on the top and sides of the neck almost halfway to the shoulders. All of the upper parts are much more darkly colored than in other specimens of tliis race. The least width of the color of the underparts is 63 per cent of the greatest width of the color of the upper parts; thus the color of the underparts is considerably more extensive than in any other specimen seen. The underparts are more intensely colored than in the average specimen. The mastoidal breadth is greater than in any other adult male and amounts to more than the postpalatal length. On available maps the elevation of Miquihuana is given as 1892 meters ( about 6200 feet ) . Thus the dark colors can hardly be ascribed to more tropical conditions than those under which animals from Brownsville, Texas, live. Brownsville is only a few feet above sea level and only 235 miles farther north. The diflFer- ence noted, therefore, seems to be of geographic significance. How- ever, there is from Alvarez, San Luis Potosi, approximately 115 miles south of Miquihuana, a yoimg (nearly subadult) female, no. 21968, which is as light colored as specimens from Brownsville. Texas, or Tlalpam, Mexico. The only distinctive feature of this MUSTELA KREN'AIA 345 specimen is the much greater extent of its white facial markings; they are more extensive even than in the specimen from Miqui- huana. Finally, the series from Brownsville, Texas, indicates that the animal there is smaller than frenata from the vicinity of Mexico (city). The skull is similarly proportioned except that relative to the basilar length the orbitonasal length is more. Several other measurements of the skull of the adult male from Tlalpam, as pointed out above, are actually, although not relatively, greater than in any specimen from Brovvoisville. The similarities between speci- mens from the two localities, Tlalpam and Brownsville, are strik- ing; since the two localities lie at opposite, extreme ends of the range more geographic variation would be expected. All that is known of the characters of populations from intermediate locaHties is that the one specimen from Alvarez shows no peculiarities whereas the one from Miquihuana suggests the existence there of a geographic variant. None of the specimens seen shows actual intergradation with M. f. neomexicana or with M. /. arthuri but it is supposed that frenata intergrades with each of these subspecies. The diflFerence between frenata and arthuri is greater than between frenata and neomexicana. Bailey (1905:198) records tracks of a weasel seen just below El Paso which he supposed had been made by a weasel of the neomexicana type. He also cited the taking of a weasel at Langtry which suggested to him {op. cit.) "... a continuous range from the country of frenatus up the Rio Grande to the type locahty of neomexicanus at Mesilla Valley," New Mexico. Other records of occurrence in Texas cited by Bailey, in addition to those provided by specimens examined by the writer, are San Diego, Bee- ville, and Port Lavaca. The Port Lavaca record is the easternmost one assigned to the subspecies frenata; possibly specimens from there would be referable to arthuri. The series of thirty-four specimens from Brownsville, Texas, per- mits measuring the amount of individual and age variation in several features. For instance, the material is sufficient to show that external measurements of subadults and those that fall in the upper part of the category designated as "young" may be included with the measurements of adults, because the mentioned measure- ments are not appreciably greater in adults. The series of skulls, although not providing more than six of any one age, shows the range of variation in size and proportion of certain parts and en- ables the student the better to evaluate cranial characters of nearby 346 University of Kansas Publs., Mus. Nat, Hist. races known from only a few specimens. For example, not one of the twenty skulls of males from Brownsville and immediate vicinity is as large as either of the two specimens of texensis from Kerr County. The white facial markings vary much in size and shape. In the series of thirty-four skins from Brownsville the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in three specimens and on one side only in six other specimens. These bands are confluent with the color of the underparts in all but two specimens. In one specimen the con- nection is lacking on both sides and in the other on one side only. A white patch between the ears is present in two specimens. The dark spot at each angle of the mouth is absent on both sides in eleven specimens and absent on one side only in ten others. In six other specimens from parts of Texas north of Brownsville, the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in one specimen. A white spot between the ears is present in one specimen. The dark spot at each angle of the mouth is absent on both sides in six speci- mens and on one side only in three other specimens. In eleven specimens from Mexico, the broad white bands in front of the ears are confluent with the white spot between the eyes on both sides in two specimens and on one side only in one other specimen. The white spot between the ears is present in one speci- men. The dark spot at each angle of the mouth is absent on both sides, in six specimens, and on one side only in one other specimen. Thus, in 51 specimens the broad bands (one in front of each ear) are confluent with the white patch between the eyes in nine- teen out of 100 instances, and not with the color of the underparts in three instances. A white spot between the ears is present in four specimens. The dark spot at each angle of the mouth is pres- ent 47 out of a possible 98 times. Four Juvenal specimens from Brownsville, Texas, with their dates of capture and probable age, are as follows: no. 58574, 5 , three weeks old, taken on February 15; no. 17318/24239, j , four weeks old, taken on March 16; no. 45899, 5 , forty days old, taken on May 21; no. 21778/36481, ^ , thirty days old, taken on October 20. In the order given, the dates of birth of these four juveniles would be approximately as follows: January 25, February 15, April 1, and September 20. The dates of birth of other specimens less than three months old as judged by the stage of development of the skull, and reckoning backward from the dates of capture, are as follows: MUSTELA FRENATA 347 April 1, April 30, May 25, October 12, and December 21. Thus, young appear to be brought forth at Brownsville, Texas, in the fall, winter and spring, that is to say from the latter part of September until the latter part of May. Mustela frenata frenata is either free of the parasites that infest the frontal sinuses of most weasels, or withstands their presence remarkably well, for only one skull shows a definite pathological condition of the frontal sinuses. Allen (1896:74) quotes H. P. Attwater, with respect to this species in Bexar County, Texas, as follows: "Not common, but occasionally met with in the chaparral and cactus lands, where Wood Rats, Rab- bits and Quail abound. They were frequently met with around San Antonio during the great 'Tramp Rat' [= Sigmodon hispidus texianus, see Bailey (1905:116)] invasion of 1889-90." Specimens examined. — Total number, 63, arranged by counties, and in Mexico by states, from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum. Texas. Bexar Co.: San Antonio, 2 (1*). Goliad Co.: Charco, 1. Nueces Co.: Corpus Christi, V. San Diego County (not found), 1. Hidalgo Co.: La Hacienda, 1. Duval Co.: San Diego, 2\ County in question: Lower Rio Grande, 1. Cameron Co.: Brownsville, 34 (3', 4% 3', 2\ 1°, 1% 1'); no locality more definite than county, 2. Nuevo Leon: Rio Ramis, 20 mi. NW Montemorelos, 1". Tamaulipas: Matamoros, 6; Miquihuana (now in Nuevo Leon), 1*. San Luis Potosi: Alvarez, 1*. Mexico: Region montagneuse des environs de Toluca, Nevada Toluca, 3200 M., V Distrito Federal: City of Mexico, 2 (1'); Tlalpam, 2. No locality more definite than Mexico, 4 (1', 3'). Mustela frenata leucoparia (Merriam) Long-tailed Weasel Plates 1, 24, 25, 26, 29, 30, 36, 37 and 38 Putorius frenatus leucoparia Merriam, N. Amer. Fauna, 11:28, June 30, 1896. Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 2:165, October 21, 1889. Putorius frenatus frenatus, Allen, Bull, Amer. Mus. Nat. Hist., 22:259, July 25, 1906. Mustela frenata leucoparia. Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936. Type — Male, adult, skull and skin; no. 34914/47179, U. S. Nat. Mus., Biol. Surv. Coll.; Patzcuaro, Michoacan, Mexico; July 27, 1892; obtained by E. W. Nelson; original no. 2960. 1. American Mus. Nat. History 6. Field Museum of Natural History 2. Acad. Nat. Sciences of Philadelphia 7. Berlin Zoological Museum 3. Univ. Kansas Mus. Nat. History 8. British Museum of Natiu-al History 4. Museum of Comparative Zoology 9. Paris Museum a. Donald R. Dickey Collection 10. Texas Co-operative Research Collection 348 University of Kansas Publs., Mus. Nat, Hist. The skull (plates 29 and 30) lacks most of the braincase; a fragment, con- sisting of the supraoccipital and the coalesced frontals and parietals remains. The rostrum, left zygomatic arch, palate, left pterygoid, left glenoid fossa and right postorbital process are intact. The teeth all are present and entire. The lower jaw lacks the right coronoid process and the lateral part of the articular condyle. The skin is well made and in good condition. It diflFers from an adult male topotype (36855, U. S. Nat. Mus.) and other referred specimens in having: the black of the head extended farther posteriorly on the neck, the maximum amount of white on the head, and a white stripe 50 mm. long extending down the middle of the nape from a point between the ears more than half way to the shoulders. Range. — Sonoran and Transition life-zones of mountains west of Mexico (city) in Michoacan and Nayarit. See figure 29 on page 221. Characters for ready recognition. — DiflFers from M. /. goldmani in least width of color of underparts more than 47 per cent of greatest width of color of upper parts, hind feet colored like underparts rather than like upper parts; postorbital constriction less than, rather than more than, combined length of upper premolars; from M. f. macrophonius by same details of coloration as from goldmani and by ventrally concave rather than ventraUy convex pretympanic part of squamosal; from M. f. perotae by least width of color of underparts more than 40 per cent of greatest width of color up upper parts; height of tympanic bulla more than three-fifths distance from its anterior margin to foramen ovale; from M. f. frenata by white facial markings that cover half of surface of head in front of ears, by extension of black of head onto neck half- way to shoulders and by narrower (less than 7.8) tympanic bullae; from Af. /. neomexicana by Argus Brown rather than Buckthorn Brown color of upper parts and distance from anterior margin of tympanic bulla to foramen ovale more, rather than less, than four-fifths of height of tympanic bulla. Description. — Size. — Male: Two adults and one young from Los Reyes and Patzcuaro, Michoacan, yield average and extreme measurements as fol- lows: Total length, 514 (510-523); length of tail, 206 (196-215); length of hind foot, 55 (52-58). Tail averages 67 per cent as long as head and body. Length of hind foot more than basal length. Female: One adult from Artenkiki, Jalisco, and one subadult from Patz- cuaro, Michoacan, measure, respectively, as follows: Total length, 412, 400; length of tail, 159, 159; lengtli of hind foot, 41, 42. Tail averages 64 per cent as long as head and body. Length of hind foot equal to or greater than basal length. The average diflFerences in external measurements of the two sexes are: Total length, 108; length of tail, 47; length of hind foot, 13. Mustelu frenata leucoparia has a greater total length and length of tail than either M. f. frenata or goldmani. The hind foot is longer than that of frenata and approximately the same as in goldmani. Relative to the body length, the tail averages longer than that of goldmani and shorter than that of frenata. Externals. — As described in Mustela frenata frenata. Color. — Broad white bands on sides of head, extending anterodorsally an- terior to each ear, confluent with white spot between eyes and with color of underparts; posterior third of each upper lip white; remainder of sides and top of head, and neck posteriorly to point halfway to shoulders from ears, black; MUSTELA FRENATA 349 no dark spots at angles of mouth; tip of tail black; remainder of upper parts Argus Brown; chin white and sometimes also chest, neck and medial sides of hind legs; remainder of underparts near (16') Ochraceous-Buff (near (a) Ochraceous-Buff in juvenal female), which color extends distally over all of each foreleg (except its lateral face proximally from about middle of forearm) and on medial side of hind leg and over most of upper side of each foot. Least widtli of color of underparts averaging, in eight specimens, 54 (ex- tremes 44-61) per cent of greatest width of color of upper parts; black tip of tail averaging, in four males, 52 (extremes 38-78) mm. long, thus aver- aging 25 per cent of length of tail-vertebrae. As compared with M. /. freiiata and goldmani: white facial markings more extensive; color of underparts less restricted and more extended on legs; black tip of tail relatively of about same extent as in frenata and thus much less than in goldmani; black color of head extending farther posteriorly than in frenata but not so far as in goldmani. Skull and teeth. — Male (adult): See measurements and plates 24-26, 29, 30. As described in Mustela frenata frenata except that: Weight (no. 128972) 6.3 grams; basilar length, 51.2; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla more or less than (about equal to) distance from its anterior margin to foramen ovale; anterior margin of mas- seteric fossa anywhere from slightly anterior, to slightly posterior, to m2. Female (based on no. 26153): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 44.5; zygomatic breadth less than distance between condylar foramen and Ml, or than between anterior palatine foramen and anterior margin of tympanic bulla; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla not more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row or than length of rostrum. The skull of the female is 43 per cent lighter than that of the male. Comparison of the skull with those of M. f. perotae, goldmani and neomexicana has been made in the accounts of those sub- species. As compared with that of frenata the main difference is the less inflated tympanic bulla, the height of which is approxi- mately equal to, rather than decidedly more than, distance from its anterior margin to foramen ovale. Remarks. — The first specimen known to have been preserved is the alcoholic in the British Museum of Natural History, taken in September, 1891, on the Rio Santiago in Jalisco, by D. A. C. Buller. The other known specimens of this white-faced weasel are divided between the American Museum and the United States National Museum. The two referred specimens from Jalisco were the last of several helpful ones collected in Mexico and Central America by J. H. Batty, and these two were taken less than three months before Batty 's tragic death in Chiapas (see Allen, J. A., 1906:191). 350 University of Kansas Publs., Mus. Nat, Hist. The five specimens from Michoacan were taken by Nelson or Nel- son and Goldman together. Merriam had only three of these when he named the subspecies and remarked (1896:29) that "This form is the poorest subspecies described in the present paper." Al- though the form is not strongly marked, the two additional speci- mens from Michoacan and better comparative material than Mer- riam had confirm several of the diflFerential characters ascribed to it by him and indicate the existence of still other characters. M. f. leucoparia occurs in the Sonoran and Transition life-zones. No. 27258 from Los Masos, and no. 26153 from Artenkike (see specimens examined for other spellings) approach true frenata in coloration. Each of these specimens has a few white hairs between the ears and the white patch between the eyes is confluent on one side only with the lateral white bands on the side of the head. No. 27258 from Los Masos has a dark spot at each angle of the mouth. The 7 other specimens are relatively uniform in coloration. Each has the white spot between the eyes confluent on both sides with the extensive white areas on each side of the face. None has a dark spot at either angle of the mouth. Of these 7 specimens, the type specimen and three others have white hairs forming a median line between the ears and a fifth specimen has a white spot behind each ear. M. f. leucoparia is most like M . /. frenata. Unlike frenata, leu- coparia has tympanic bullae that are less inflated, narrower and less projected, at their anterior margins, from the cranium. In these characters leucoparia is intermediate between M. /. frenata and M. /. goldmani. The latter subspecies has the least inflated, nar- rowest and least projecting tympanic bullae of the three. The black color of the head extends, on the average, farther posteriorly than in M. /. frenata but not so far as in M. /. goldmani. The gen- eral color, too, is intermediate between that of M. /. frenata and that of the much darker M. f. goldmani. The white facial mark- ings are more extensive than in either M. /. frenata or M. f. gold- mani. This applies to both the white area between the eyes and the one on each side of the head between the ear and eye. M. /. neomexicana, whose range possibly meets that of M. /. leucoparia, also has more extensive white facial markings than M. /. frenata but less extensive markings than M. /. leucoparia. On the basis of skulls alone, specimens of frenata from Tlalpam and those of leucoparia from Los Reyes can hardly be distinguished. This fact, and the circumstance that the specimens from the north- em part of the range of leucoparia closely resemble frenata in color, MUSTELA FRENATA 351 constitute sufficient evidence for regarding the two as only sub- specifically distinct. The female, no. 26153 from Artenkike, as mentioned above, approaches true frenata in coloration. On this account it is not to be regarded as typical and it was because no other skulls of adult females were available that this one was used for comparison with females of allied races. M. /. leucoparia is, then, a subspecies of the large, temperate- zone group and is unique in possessing the maximum extent of white facial markings. None of the seven skulls shows signs of having had the frontal sinuses infested with parasites. Specimens examined. — Total number, 8, all from Mexico. Localities are listed by states from north to soutli. Specimens from Michoacan are in the United States National Museum; one from Rio Santiago is in the British Museum of Natural History; all others are in the American Museum of Natural History. Nayarit: Tepic, 1. Jalisco: Rio Santiago, 1; Los Masos, 1; "Artenkike" (J. A. Allen, 1906, p. 238, writes "Artenkikil" and, on p. 259, "Artenkiki". ) , 1. Michoacan: Zamora, 1; Los Reyes, 1; Patzcuaro, 3. Mustela frenata perotae Hall Long-tailed Weasel Plates 36, 37 and 38 Mustela frenata perotae Hall, Carnegie Instit. Washington Publ. 473:100, November 20, 1936. Putorius frenatus, Merriam, N. Amer. Fauna, ll:pl. 3, fig. 2, June 30, 1896. Type. — Female, adult, skull and skin; no. 54278, U. S. Nat. Mus., Biol. Surv. Coll.; 12,500 feet, Cofre de Perote, Veracruz, Mexico; May 26, 1893; obtained by E. W. Nelson; original no. 4864. The skull (plates, 37-39) lacks the right zygomatic arch. Left p2 is missing. The skin is fairly well made and in good condition except that the extreme tip of the tail has been broken off and there are two holes in the right hind leg. The underparts show the beginning of a spring molt. Range.— From 7500 (?) feet (Perote) to 13,500 feet (Popocatepetl), Upper Sonoran, Transition and Boreal life-zones of mountains along Puebla-Mexico boundary, eastward to western central Veracruz and south into Oaxaca. See figure 29 on page 221. Characters for ready recognition — Differs from M. /. frenata, its nearest relative, in extension from head of blackish onto anterior fourth of neck; restriction of color of underparts (least width of same less than 37 per cent of greatest viddth of color of upper parts), height of tympanic bulla less than distance from its anterior margin to foramen ovale; from M. f. macrophonius and M. f. goldmani in presence of, rather than absence of, color of underparts on hind feet; upper parts (black) Brussels Brown rather than Argus Brown or darker; from M. f. tropicalis in larger size (adult female with total length more than 400, basilar length more than 40, weight of skull more than 3 352 University of Kansas Publs., Mus. Nat. Hist. grams); postorbital breadth less than combined length of upper premolars; ml more than 5.4 long; from M. f. leucoparia in white facial markings so restricted that spot between eyes is not confluent with white stripe in front of ear, or, if so, narrowly (less than 4 wide) confluent; color of upper parts extending onto antipahnar face of forefoot, least width of color of imderparts not more than 40 per cent of greatest width of color of upper parts; height of tympanic bulla not more than three-fifths distance from its anterior margin to foramen ovale. Description. — Size. — Male: A nontypical specimen from Cerro San Felipe, Oaxaca, measures: Total length, 500; length of tail, 205; length of hind foot, 52. Female: The type specimen, measures: Total length, 418; length of tail, 160; length of hind foot, 45. In this male the tail is 70, and in the female, 62 per cent as long as the head and body. In each the hind foot is longer than the basal length. The differences in external measurements between these two specimens, representing the two sexes, are: Total length, 82; length of tail, 45; length of hind foot, 7. Externals. — ^As described in Mustela frenata frenata. Color (based on type specimen). — Color and color pattern as described in Mustela frenata frenata except that: blackish of sides and top of head ex- tends one-fourth of way back to shoulders from ears; throat and breast as well as chin white; remainder of underparts near (16' c) Ochraceous-Buff; least width of color of underparts equals 36 per cent of greatest width of color of upper parts; black tip of tail equal to 28 per cent of length of tail-vertebrae. Skull and teeth. — Male (based on a referred specimen from Cerro San Felipe which certainly is nontypical ) : See measurements. As described in Mustela frenata frenata except that: Weight, 4.9 grams; basilar length, 49.2; postorbital breadth more than distance between posterior borders of P4 and P2; tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla. Female (based on type specimen, an adult): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight 3.4 grams; basilar length, 43.5; zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla one-half to three-fifths dis- tance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum. The skull of the female is 33 per cent lighter than that of the nontypical ( and smaller than average ) male from Cerro San Felipe. Comparison of the skull with that of M. /. tropicalis is made in the account of that subspecies. Compared with the skull of M. /. MUSTELA FRENATA 353 macrophonius, that of the female of perotae is more flattened, has the longitudinal dorsal outHne distinctly concave rather than flat just behind the postorbital processes, and much wider tympanic bullae. Accordingly, the basioccipital is sHghtly narrower in perotae. The more marked postorbital constriction of the type specimen of perotae possibly is due to its relatively greater age. As compared with the skull of M. /. leucoparia, that of the female of perotae has less inflated tympanic bullae, the height of each being half as great as distance from its anterior margin to foramen ovale, whereas, in leucoparia (as represented by no. 26153) the two distances are equal. As compared with that of M. /. frenata, the skull of the female of perotae differs mainly in the lesser in- flation of the tympanic bullae and their relative position. The height of each bulla is in perotae only half as much as, but in frenata more than, the distance from its anterior margin to foramen ovale. The anterior margin of the bulla is much less projected from the floor of the braincase in perotae. The squamosal anterior to each bulla is convex ventrally in perotae but flat or concave ventrally in frenata. Remarks. — The type specimen and a juvenal female from the town of Perote were taken in the spring of 1893 by E. W. Nelson. Of these two, the type specimen was mentioned and figured by Merriam (1898:30, fig. 16 [= fig. 15], pi. 3, fig. 2) as Putorius fre- natus. The referred nontypical specimen from Cerro San Fehpe, Oaxaca, was referred by Merriam {op. cit. -.29) to Putorius frenatus goldmani with the comment that it was intermediate ". . . both in coloration and cranial characters, between typical frenatus and goldmani; . . .". No other pubhshed references to this sub- species, or specimens of it, have been seen. In 1941 and 1942, W. B. Davis and associates took four specimens along the bound- ary between the states of Puebla and Mexico. Although the specimen from Cerro San Fehpe, Oaxaca, is re- ferred to Mustela frenata perotae, to the description of which it answers best, that specimen, on account of its structural charac- ters and geographic position relative to adjacent races, is in reahty an intergrade between several of the adjacent races. Some of its intermediate characters are pointed out in the discussion of M. /. goldmani. In the specimens from 45 and 55 kilometers ESE of Mexico (city) the black color of the top of the head does not extend so far behind the ears as in the holotype of M. /. perotae and in this feature the two specimens show intergradation between the two subspecies, perotae and frenata. 12—3758 354 University of Kansas Publs., Mus. Nat, Hist. The type specimen taken on May 26, is acquiring new hair on the belly and lower sides which appears to be the result of a nor- mal molt. As would be expected from its geographic position, M. /. perotae resembles M. f. frenata of northern Mexico and the high mountain forms of southern Mexico more than it does the lowland tropical forms. This is true as regards size of entire animal, proportions of its parts, and size, general angularity and major proportions of its skull. The marked postorbital constriction, convex supralacry- mal face of rostrum, width of tympanic bullae and angularity of the braincase place it nearest M. f. frenata as does also the color and color pattern. The ventrally convex squamosal anterior to each tympanic bulla and the slight degree of projection from the cranium of the anterior margin of each tympanic bulla are inter- mediate in degree between the condition in M. f. macrophonius and that in M. f. frenata. Thus M. /. perotae combines several characters of M. /. frenata on the one hand with some of M. f. macrophonius on the other and in some features, for instance in the size, shape and degree of inflation of the tympanic bullae, presents intermediate stages of development. On the eastern plain below the high mountain, Cofre de Perote, there ranges the similarly colored, smaller, tropical weasel, Mus- tela frenata tropicalis. Between M. f. perotae and M. /. tropicalis there is marked differentiation in the skulls with much less differenti- ation in coloration. The differences in typical skulls of the two sub- species are so pronounced that one would, at first glance, hardly believe it possible for direct intergradation to occur between them on the sides of this mountain. Merriam (1896:30) thought that it did not. The two skulls figured by him (op. cit.:Sl) are a topotype of M. /. tropicalis from Jico and the one which now is the type specimen of M. /. perotae. They show the great difference in size and proportions and are females of comparable ages, not of differ- ent ages as I suspected before examining the skulls. However, de- spite this marked difference in the skulls, there is some, although not conclusive, evidence of intergradation furnished by a young female from Xuchil, Veracruz. This specimen is described in con- nection with M. f. tropicalis ( see p. 366 ) . None of the seven skulls shows marked deformity of the inter- orbital region, but two of the three adults appear to have had these parts infested with nematodes. Specimens examined. — Total number, 7, all from Mexico, listed by localities from north to south. Specimens from Veracruz and Oaxaca in the United States National Museum; remainder in Texas Co-operative Research Collection. MUSTELA FRENATA 355 Mexico: Monte Rio Frio, 45 Km. ESE Mexico City, 1; 55 Km. ESE Mexico City, 1; N slope Mt. Popocatepetl, 13,555 ft., 1. Puebla: Rio Otlati, 8700 ft., 1. Veracruz: Cofre de Perote, 12,500 ft., 1; Perote, 1. Oaxaca: Cerro San Felipe, 10,000 ft., 1. Mustela frenata goldmani (Merriam) Long-tailed Weasel Plates 1, 24, 25, 26 and 30 Putorius frenatus goldmani Merriam, N. Amer. Fauna, 11:28, June 30, 1896; Elliot, Proc. Biol. Soc. Washington, 18:236, December 9, 1905. Mustela frenata goldmani. Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936. Type. — Male, adult, skull and skin; no. 77519, U. S. Nat. Mus., Biol. Surv. Coll.; Pinabete, Chiapas, Mexico; February 10, 1896; obtained by E. A. Gold- man (on attached label collectors recorded as Nelson and Goldman); original no. 9279. The skull (plates 24 and 30) has the rostrum badly injured. All the right, and part of tlie left nasal, the upper part of the right maxilla, the postorbital process and intervening area of frontals are missing. Each zygomatic arch is broken but the parts are present and attached to the skull. The frontal and interorbital regions are greatly malformed owing to parasites that infested the sinuses. Right 12 and 13, right and left i3, and the medial parts of the paraconid and protoconid of right ml are missing. The light facial markings are less extensive than in any of the referred specimens. These markings consist of a separate spot between the eyes and a white line, confluent with the color of the underparts, on each side of the head, that extends from the base of the ear to above the eye. The dark color of the underparts is repre- sented at the angles of the mouth by a spot on the left side and a similar dark area, confluent with the dark color of the face, on the right side. The large size, characters of the skull, and scrotal pouch on the skin prove the specimen to be a male as stated on the label. Range. — Two thousand five hundred feet (El Cipres, Guatemala) to 9500 feet (near Tecpam, Guatemala), Upper Tropical Life-zone of mountains and western coasts of southern Mexico, Guatemala and Salvador. See figure 29 on page 221. Characters for ready recognition (characters based on males). — Differs from M. f. nicaraguae and M. /. perda by larger size (total length of adult males more than 489), least width of color of underparts not less than 26 per cent of greatest width of color of upper parts, weight of skull of adult male more than 5 grams; from M. /. macrophonius by smaller size (total length of adult males less than 540), skull of male with basilar length less than 52.5 and weight less than 6 grams; from M. f. perotae (typical specimens of same sex not available ) by darker color of upper parts which are Argus Brovra or darker rather than Brussels Brown; nonextension of color of underparts onto hind feet; from M. /. leucoparia in least width of color of underparts not more than 37 per cent of greatest width of color of upper parts; color of underparts not extended onto hind feet; black tip of tail two-fifths rather than one-fourth as 356 University of Kansas Publs., Mus. Nat. Hist. long as tail-vertebrae; height of tympanic bulla less than four-fifths distance from its anterior margin to foramen ovale. Description. — Size. — Male: Four adults yield average and extreme meas- urements as follows: Total length, 508(500-512); length of tail, 196( 185-207); length of hind foot, 55.5 ( 54-58 ) . Tail averages 63 ( 59-67 ) per cent as long as head and body. Length of hind foot more than basal length. Female: Typical specimen unknown. Externals. — Longest facial vibrissae black and reaching beyond ear; carpal vibrissae wholly or in part of same color as upper parts and reaching as far as hypothenar pad; hairiness of foot-soles distinctly less than that shown in figure 20 on page 60. Color. — Spot between eyes, band, confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear and posterior third of each upper lip, white; remainder of sides and top of head and neck posteriorly to or slighdy behind shoulders, black; dark spots at angles of mouth usually absent; tip of tail black; remainder of upper parts Argus Brown or near (n) Argus Brown; chin, throat and breast white; remainder of under- parts near (16' c) Ochraceous-Bufi^; color of underparts extending distally on posterior sides of forelegs onto medial toes and on hind legs to points between knees and heels. Least width of color of vmderparts, in five adult males, aver- aging 28 (extremes 26-33) per cent of greatest width of color of upper parts; black tip of tail, in four adult males, averaging 40 per cent of length of tail- vertebrae. Skull and teeth. — Male ( based on five adults ) : See measurements and plates 24-26, 30; weight, 5.4 (5.3-5.5) grams; basilar length 49.9 (49.6-51.3); zygomatic breadth (except in no. 12523 from Salvador) more than or equal to distance between condylar foramen and Ml or between anterior palatine foramen and anterior margin of tympanic bulla. Mastoid breadth less than postpalatal length; postorbital breadth more or less than length of upper pre- molars and greater than width of basioccipital measured from median margin of one foramen lacerum posterior to its opposite; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of five upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of t>'mpanic bulla more than length of lower molar and premolar tooth-row and shorter than or equal to length of rostrum; anterior margin of masseteric fossa lirmiediately behind m2. Female: Typical skull unknown. Comparison of male skull with that of M. /. per da made in dis- cussion of that form. Comparison with that of M. f. nicaraguae shows similar differences, some of which are more pronounced. For example, squamosals anterior to tympanic bullae more convex ventrally and these bullae project less from braincase than in M. f. perda; thus the difference in these features is greater between goldmani and nicaraguae than between goldmani and perda. MUSTELA FRENATA 357 As compared with the skull of the male of M. /. macrophonius, each one of the skulls of the adult males of M. f. goldmani is smaller in every measurement taken, with two exceptions. The width of the tympanic bullae was more in three specimens of M. /. goldmani as was also the depth of the same in three specimens. Relative to the basilar length all but two of these measurements average less in goldmani; the exceptions are the zygomatic breadth and depth of the skull at the anterior margin of the tympanic bullae which average more. Relative to the basilar length, the orbitonasal length and depth of the skull at the posterior margin of Ml are less in each skull of goldmani. Thus, excepting the width and height of the tympanic bullae and the relative zygomatic breadth and relative depth of the braincase posteriorly, the skull of gold- mani is shorter and relatively as well as actually narrower and hghter throughout. As compared with the skull of the male of M. f. leucoparia, that of M. f. goldmani averages a trifle shorter and no skull of goldmani equals that of leucoparia in actual or relative zygomatic and mas- toid breadths or length or height of tympanic bullae. In depth, the skull of gold7nani averages actually and relatively greater. Its teeth are smaller. The squamosal anterior to each tympanic bulla is con- vex ventrally whereas it is concave ventrally in leucoparia as in frenata. Remarks. — When Merriam (1896:28) named this subspecies, he had only one specimen but he called attention to the more impor- tant diagnostic characters, which additional specimens show per- tain to the race as a whole. M. f. goldmani in typical form occurs in high mountains of the Upper Tropical Life-zone and is most closely related to M. f. frenata and M. /. macrophonius. The altitude at which the two speci- mens were taken, twenty miles southeast of Teopisca in Chiapas, is not known. Merriam (1896:28) states that the type specimen was obtained at "about 8200 feet." The specimen taken by Stirton in Salvador comes from 8000 feet and the one obtained by Barber in Guatemala from 9500 feet. The specimen from Duenas, the skin alone of a young animal, is not instructive. As regards size, goldmani is larger than the immediately adjacent subspecies from the Lower Tropical Life-zone but is smaller than M. /. leucoparia or macrophonius. As compared with M. /. frenata, goldmani is longer, has an actually as well as relatively shorter tail, and a much longer hind foot. 358 University of Kansas Publs., Mus. Nat. Hist. The most outstanding difference in externals from frenata is the naked foot soles. Molting probably takes place twice each year although actual proof of this is lacking. In number 133254 from twenty miles south- east of Teopisca, taken on May 12, the molt is well advanced. An- other specimen from the same place still retains the winter coat. In color, goldmani is much darker than frenata, has less extensive white facial markings, longer black tip on tail, more restricted color of underparts, and lacks the extension of color of the under- parts onto the hind feet. Of the adult males from the high mountains, the type specimen from Chiapas is lightest, and the one from Salvador is darkest. This progressively darker color to the southward probably is geographic variation. In total length and relative and actual length of tail, the speci- men from Salvador is the smallest of the five adult males from the higher mountains. In addition to its darker color and smaller size, no. 12523 from Salvador shows certain distinctive cranial char- acters. The zygomatic breadth is less than, rather than more than, or equal to, the distance between the condylar foramen and Ml or than that between the anterior palatine foramen and the anterior margin of the tympanic bulla. This difference appears to be cor- related with geographic position, since no. 15953 from Guatemala has the three distances about equal and therefore is intermediate in this respect between the specimen from Salvador and those from Chiapas, in which the zygomatic breadth is greater than the other two measurements. Also in the greater depth of the skull and smaller size of the teeth, this specimen from Salvador approaches the subspecies of the Lower Tropical Life-zone. It has, however, the longest, highest and widest tympanic bullae of any of the five specimens. The amount of ventral convexity of the squamosal in front of each tympanic bulla appears not to be greater than in the other specimens. As indicative of intergradation with perotae, leticoparia and pos- sibly frenata, there is the specimen from Cerro San Felipe, Oaxaca. The degree of restriction of the color of the underparts is inter- mediate between that of goldmani and leucoparia. The same is true as regards the amount of projection from the braincase of the anterior margins of the tympanic bullae. The squamosal imme- diately anterior to each tympanic bulla is flat instead of ventrally convex as in goldmani or ventrally concave as in leucoparia and frenata. In accordance with the custom adopted in this paper of MUSTELA FRENATA 359 referring every specimen to some one subspecies, this specimen from Cerro San Felipe is referred to Mustela frenata perotae, to the description of which it most nearly answers. Possibly goldmani, as here constituted, is a composite form. The specimens from the high mountains closely resemble one another. However, a specimen, no. 68541 from "Finca El Cipres", Guatemala, which place Mr. G. Goodwin tells me is at an elevation of 2500 feet, approximately 5 miles north of Retalhuleu, has a basilar length of 47.3 and is correspondingly small in other parts. This suggests the existence of a small, lowland race on the western side of the central divide corresponding to perda and tropicalis on the east- ern side. From only a few miles away, at San Sebastian, there is available, the adult skull of a still smaller animal. This skull only, no. 41026, in the Berlin Zoological Museum, has a basilar length of 46.1, zygomatic breadth of 27.4, and other cranial measurements notably smaller than those of specimens from the high mountains. A skin-only, no. 12038, collection of Donald R. Dickey, from La Cebia, altitude 2150 feet, near the city of San Salvador, seemingly represents an animal smaller than typical goldmani. This speci- men from La Cebia has the light color of the underparts extended distally on the hind legs to the tips of the toes as in M. /. tropicalis. However, the upper parts are darker and resemble those of M. f. goldmani. A fourth specimen from only 3500 feet elevation, on the south side of Volcano Tajumulco, Guatemala, no. 41768, Field Museum of Natural History, a subadult male, measures only 490 in total length and has the least color of the underparts so restricted as to amount to only 22 per cent of the greatest width of the color of the upper parts. Both these features are suggestive of the low- land races. These four specimens indicate that the lowland population on the western side of the divide is smaller than the mountain popula- tion. The juvenile from Carolina and a young male from Finca Cipres, however, both closely resemble individuals of goldmani from the higher mountains. All these animals here are referred to goldmani. More specimens may reveal an amount and a pat- tern of geographic variation in weasels of this region that will re- quire application of another subspecific name. The female, no. 68540, from Puebla agrees remarkably well with the skull of the female, no. 132528, of macrophonius. Differences displayed by the specimen from Puebla are its slightly narrower braincase and longer space between the foramen ovale and anterior 360 University of Kansas Publs., Mus. Nat. Hist. end of the tympanic bulla. Considering the far eastern location of Puebla (just north of Rio Motagua, at 89° W, according to a sketch map provided by Mr. G. G. Goodwin), this specimen might be expected to show some approach to the small lowland races. Actually, however, it displays the characters of goldmani better than does the subadult female from Volcano San Lucas, which is nearer the metropolis of goldmani, and I assume at a higher eleva- tion than Puebla. Concerning this weasel Merriam (1896:29) says: "Mr. E. W. Nelson writes me that this fine weasel is found sparingly in the forest about Pinabete, Chiapas, at an altitude of 7000 to 8000 feet (2100 to 2500 meters). The type specimen was shot in the afternoon while hunting on a heavily wooded hill slope. It was heard making long, slow leaps over the dry, crisp leaves. Coming to a log, it stood up and rested its fore feet on the log, in which position it was shot by Mr. Goldman." The specimen taken by R. A. Stirton in Salvador comes from an elevation of 8000 feet in the rain forest of the Upper Tropical Life-zone. Mr. Stirton tells me that one morning on visiting his traps set for small rodents, he found in one the partly eaten re- mains of a Heteromys. Leaving these remains as found he placed a steel trap beside them and on the following morning found the male weasel in the trap. At least three of the ten specimens had the frontal sinuses in- fested with parasites. Specimens examined. — Total number, 15, listed by localities from north to south, and imless otherwise indicated in the American Museum of Natural History. Mexico. Chiapas: 20 mi. SE Teopisca, 2'; Pinabete, 1'. Guatemala. Puebla, 1; Finca Porvenir, 3500 ft., S slope Volcan Tajumulco, 1'; Sierra [= ? Cerro] Santa Elena, 9500 ft. (near Tecpam), !■; Carolina, 1; Volcano San Lucas, 1; "Finca El Cipres", 1; "Finca Cipres", 2500 ft., 1; Finca San Isidro, San Sebastion, Dept. Retalhuleu, 1^; Dueiias, V; no locality more definite than Guatemala, 1*. Salvador. Los Esesmiles, 8000 ft., Chalatenango, 1°; La Cebia, 2150 ft., near Sun Salvador, 1°. Mustela frenata macrophonius (EUiot) Long-tailed Weasel Plates 24, 25, 26, 30, 37, 38 and 39 Futorius macrophonius EUiot, Proc. Biol. Soc. Washington, 18:235, Decem- ber 9, 1905. 1. United States National Museum 4. British Musem of Natiwal History 2. Field Museum of Natural History 5. Donald R. Dickey Collection 3. Berlin Zoological Museum MUSTELA FRENATA 361 Mustela macrophonius. Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912. Mustela frenata macrophonius, Hall, Carnegie Instit. Washington Publ. 473: 109, November 20, 1936. Type. — Male, adult, skull and sldn; no. 14063, Field Mus. Nat. Hist.; Acho- tal, Veracruz, Mexico; January 15, 1904; obtained by Edmund Heller and Charles M. Barber; original no. 3424. The skull (plates 24-26, 30) is complete and unbroken. Excepting right P2, which has been aborted or broken away, all the teeth are present. The skin is well made and in good condition. As shown by the scrotal pouch, the specimen is a male. Range. — Tropical Life-zone, probably into Boreal Ufe-zones, of mountains along eastern border of southern Veracruz. See figure 29 on page 221. Characters for ready recognition. — Differs from M. frenata frenata and M. f. perotae and M. /. leucoparia in lacking color of underparts on hind feet and in larger skull (skulls of adult males with basilar length more than 52.5); from M. /. goldmani by larger size of skull (see above) and entire animal and wider tympanic bullae; from M. f. tropicalis and M. /. perda by larger size (total length of adult males more than 510), postorbital breadth amounting to less than combined length of upper premolars. Description. — Size. — Male: External measurements of the type specimen, an adult, are: Total length, 598; length of tail, 246; length of hind foot, 59. Tail 70 per cent as long as body; length of hind foot more than basal length. Female: The skin, without field collector's measurements, of an adult fe- male from Perez, Veracruz, shows this sex to be correspondingly large. Be- cause the skin is understuffed and because the hind feet are skinned out, re- liable measurements can not be obtained from the dried skin. Externals. — As described in Mustela frenata goldmani except that all carpal vibrissae are of same color as upper parts and that hairiness of foot-sole^ is halfway between that shown in figures 20 and 21. Color. — As in darkest individuals of M. f. goldmani, thus, color of upper parts on posterior part of back near (n) Argus Brown. Color of underparts near ( 12 ) Mikado Orange in a juvenile, extending distally on posterior sides of forelegs onto inner toes and on hind legs to points between knees and heels. Least vddth of color of underparts 28 per cent of greatest width of color of upper parts. Black tip of tail 34 per cent of length of tail-vertebrae. Skull and teeth. — Male (based on type specimen): See measurements and plates 24-26, 30. As described in Mustela frenata frenata except that: Weight, 6.9 grams; basilar length, 54.1; zygomatic breadth less than distance between condylar foramen and Ml or that between anterior palatine foramen and an- terior margin of tympanic bulla; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 6 upper in- cisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; anterior margin of masseteric fossa below posterior half of m2. Female (based on no. 132528): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 43.5; zygomatic breadth less than distance between condylar foramen and Ml and more or less than (in the single specimen, equal to) that between 362 University of Kansas Publs., Mus. Nat. Hist. anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more or less than (about equal to) outside length of P4; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale. The skull of the female is 48 per cent hghter than that of the male. Comparison of the skull with that of M. /. goldmani is made in the account of that subspecies. Similar differences probably exist between males of perotae and macrophonius. As compared with skulls of males of M. f. tropicaUs and perda, the skull of the male of macrophonius is larger in every measurement taken. The post- orbital constriction is less, rather than more, than the combined length of the upper premolars. Relative to the basilar length, the following measurements are less than in any specimen of tropicalis or perda: length of tooth-rows ; orbitonasal length; depth of skull at posterior border of upper molars; and depth of skull at anterior margin of basioccipital. Remarks. — This large weasel appears to have escaped the notice of naturalists until the spring of 1903 when J. Friesser obtained an adult female and juvenal male at Perez for the collection of the United States Bureau of Biological Survey. These specimens were tentatively referred to Mustela tropicalis. In the following Janu- ary, Edmund Heller and Charles M. Barber obtained the adult male that was made the type specimen by Elhot who did not see, or if he did, did not mention, the specimens from Perez. He did, however, refer a young female from Xuchil, Veracruz, to his Pu- torius macrophonius. This young female is here referred to Mus- tela frenata tropicalis. The extent of the geographic range of this subspecies is not well known. Mustela frenata macrophonius and M. f. texensis are the largest American weasels. The basilar length in the type specimen is greater by one-tenth of a millimeter than in tlie type specimen of M. f. texensis. The other cranial measurements taken are greater in M. /. texensis. The skull of the female from Perez is one of the largest skulls examined of that sex. The juvenal male has teeth as large as those of the type specimen and the skull is the largest for its age of any seen. Although the skin of the female is under- stuffed and hence does not provide reliable measurements, it shows that the female is also large. The white bands in front of the ears are confluent with the white patch between the eyes on one side only in one specimen. It is the juvenal male. These bands are not confluent with the color of the MUSTELA FRENATA 363 underparts on either side in the female and on one side only in the adult male. None of the specimens has a white patch between the ears. The dark spot at each angle of the mouth is present only in die juvenile where it occurs on each side. Of the three specimens, the juvenile is the darkest and the adult male the lightest. The white facial markings are most extensive in the juvenal male and the least extensive in the adult female. M. /. macrophonius most closely resembles M. /. goldmani but in the relatively flattened braincase, deep constriction of the post- orbital region and general angularity of the skull approaches M. /. perotae and M. /. frenata. Only one of the three skulls, that of the female, shows evidence of infestation of the frontal sinuses by parasites, and this did not result in malformation of the interorbital region. Specimens examined. — Total number, 3, all from Mexico, listed by localities from north to south. Veracruz: Achotal, 1 (Field Mus. Nat. Hist.); Perez, 2 (U. S. Nat. Mus.;. Mustela frenata tropicalis (Merriam) Long-tailed Weasel Plates 25, 26, 27, 30, 37, 38 and 39 Putorius tropicalis Merriam, N. Amer. Fauna, 11:30, pi. 3, figs. 5, 5a, 6, 6a, text fig. 16, June 30, 1896; Merriam, Proc. Biol. Soc. Washington, 15:68, March 22, 1902. Putorius frenatus, Merriam, N. Amer. Fauna, 11:27, June 30, 1896. Mustela tropicalis tropicalis. Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916. Mustela frenata tropicalis, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936. Type — Male, adult, skull and skin; no. 54994, U. S. Nat. Mus., Biol. Surv. Coll.; Jico, Veracruz, Mexico; July 9, 1893; obtained by E, W. Nelson; original no. 5195. The skull (plates 25-27, 30) is complete. All the upper incisors, except the second and third on the left side, are missing. The right upper canine is broken. The skin is well made and in good condition. Range. — Up to 5000 feet ( as now known ) in Tropical Life-zone of Veracruz, Mexico. See figure 29 on page 221. Characters for ready recognition. — Differs from M. f. frenata and M. f. perotae in least width of color of underparts not exceeding 36 per cent of greatest width of color of upper parts and in postorbital breadth exceeding length of upper molar and premolar tooth-rows; from M. f. macrophonius and M. /. perda in least width of color of underparts averaging more than 29 per cent of greatest width of color of upper parts; and from M. f. perda by longer tympanic bullae which in males are more than 14.9; and from M. /. macro- phonius by lesser basilar length ( not more than 48 ) and in postorbital breadth exceeding length of upper molar and premolar tooth-row. 364 University of Kansas Publs., Mus. Nat. Hist. Description.— Size.— Male: The type specimen and no. 12764/11058, a subadult, from Jalapa, Veracruz, measure, respectively, as follows: Total length, 444, 442; length of tail, 175, 160; length of hind foot, 50, 47. The tail is 65 and 57 per cent as long as the head and body. The hind foot is more or less than (approximately equal to) the basilar length. Female: Merriam (1896:31) gives the measurements of a female topotype (probably no. 54993, U. S. Nat. Mus., which has no measurements written on the attached label) as: Total length, 333; length of tail, 121; length of hind foot, 37. The length of the tail amounts to 57 per cent of the length of the body. The length of the hind foot of no. 54993, U. S. Nat. Mus. is the same as the basal length. The differences in external measurements between the male and the female topotypes are: Total length, 111; length of tail, 54; length of hind foot, 13. Externals. — As described in Mustela frenata frenata except that carpal vibris- sae do not reach apical pad of fifth digit and hairiness of foot soles is less. Color. — As described in M. /. frenata except that: Blackish of head extends half way or more from ears to shoulders; upper parts near ( 14) Brussels Brown or shghtly faded tone 2 of Maroon of Oberthiir and Dauthenay, pi. 341; under- parts of Juvenal pelage near (a) Ochraceous-Buff. Least width of color of underparts averaging (in three specimens from Jico and one from Jalapa) 34 (extremes 30-37) per cent of greatest width of color of underparts. Black tip of tail, in two male topotypes, 57.5 (55 and 60) mm. long; tlius longer than hind foot and in each individual comprising 34 per cent of length of tail- vertebrae. As compared with M. f. frenata: White facial markings slightly less extensive; blackish (not black) of head extending onto neck; upper parts sfightly darker; ventral side of tail noticeably darker; color of underparts more restricted, av- eraging approximately one-third rather than nearly one-half width color of upper parts; black tip of tail one-third rather than one-fourth length of tail and much longer than hind foot. Similar differences of lesser amount exist between perotae and tropicalis. M. f. perda, macrophonius and goldmani bear the opposite relation to tropicalis. That is to say, in the latter: White facial markings shghtly more extensive; blackish of head less extended over neck; upper parts markedly hghter; color of underparts less restricted and black tip of tail shorter. Skull and teeth. — Male (based on type specimen and a subadult, no. 11058, from Jalapa): See measurements and plates 25-27, 30. As described in Mustela frenata perda except that: Weight 4.7 (4.6 and 4.7) grams; basilar length 46.7 (45.5 and 47.8); zygomatic breadth more or less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 (including 13) upper incisors; anterior margin of masseteric fossa below middle of m2 or posterior to that tooth. Female ( based on no. 54993 and no. 1060 ) : See measurements and plates 37-39. As described in Mustela frenata perda except that: Weight (of 54993) 2.2 grams; basilar length, 37.5 (36.0-39.0); zygomatic breadth more or less than distance between anterior palatine foramen and anterior margin of tym- panic bulla; least width of palate more than greatest length of P4; height of MUSTELA FRENATA 365 tympanic bulla equal to one-third to three-fourths of distance from its anterior margin to foramen ovale. The skull of the adult female is 53 per cent lighter than that of the type specimen, a male. Comparison of the skulls of males and females with those of M. /. perda, the nearest relative, has been made in the discussion of that subspecies. Some of the features that readily distinguish skulls of M. /. tropicalis from those of M. /. frenata, perotae and macrophonius are as follows: Weight less than 4.8 grams; basilar length less than 48; postorbital breadth more than length of upper M-Pm tooth-row. The skulls of male frenata, perotae and macro- phonius are much larger, heavier, and are decidedly more angular with more constricted postorbital region the least width of which is less than the length of the upper premolars. In frenata the anterior margins of the tympanic bullae are protruded much farther from the braincase. The skull of the female of M. /. tropicalis is smaller, weighing less than 3 grams; basilar length less than 41; postorbital breadth more tlian length of upper molar and premolar tooth-row. Remarks. — This subspecies was originally described by Merriam as a full species. Later he described Putorius tropicalis perdus as another subspecies. Allen (1916) placed P. t. perdus in syn- onymy but named Mustela tropicalis nicaraguae as new. In the present paper all three forms are recognized but are regarded as only subspecifically distinct from the other bridled weasels of Mexico and Central America. The limits of the geographic range of tropicalis are fairly well known on the south and west but the only specimen available from the tropical coastal region north of Jico, is a young female from a point 50 miles south of Victoria. Thus, how far north along the coast it ranges toward Matamoros, where M. /. frenata occurs, is not known. The three specimens from Jico, a young female from Jalapa and another adult collected by J. Potts and labeled as coming from Mexico City, are assumed to be typical. The latter specimen certainly came from an elevation lower than that of Mexico City be- cause M. /. frenata occurs there. Although the female from Jalapa, agrees well with specimens from Jico, a male, no. 12764/11058, from Jalapa, has a relatively broader skull, as in perda, although the tympanic bullae are short as in tropicalis. The resemblances to perda in features of coloration are: slightly darker upper parts, and the termination just below the knees of the color of the under- parts. There are three specimens labeled as from Orizaba that 366 University of Kansas Publs., Mus. Nat. Hist. indicate intergradation with perotae as does also the coloration of the Juvenal female from 5 kilometers north of Jalapa. The speci- mens labeled as from Orizaba are old, poorly-prepared skins, only two of which have partial skulls. The size and coloration of the skins suggest perotae as do also the partial skulls in some respects although the skulls show greater resemblance to those of tropicalis. The topotype, female, no. 54993, was figured by Merriam (1896, fig. 16, p. 31 ) along with that of what now is the type specimen of M. /. perotae. Merriam called attention to the great difference in size between the skulls of the two sexes of M. /. tropicalis and compared the condition to that found in noveboracensis. Although the skull of the female from Jico is fully adult, it probably is exceptionally small. The young female from Xuchil is indistinguishable in coloration from the ju venal female of M. /. perotae from Perote, but in size of skull and size of teeth is intermediate between the female of tropicalis from Jalapa and the females from Cofre de Perote. There is then, indication of intergradation with M. /. perotae as well as with M. /. perda. M. f. tropicalis differs from M. f. perotae and M. /. frenata in about the same way that M. /. perda differs from M. f. goldmani and M. /. macrophonius. M. f. tropi- calis and perda each is smaller and more intensely colored than goldmaiii and macrophonius, and inhabits the lowland to the east of their highland relative. At least five of the nine skulls have the frontal sinuses infested by parasites. Specimens examined. — Total number, 13, all from Mexico, listed by lo- calities from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum. Tamaulipas: 50 mi. S Victoria, 1°. Near? Mexico City, 1. Veracruz: Jico, 3; 5 km. N Jalapa, 1"; Jalapa, 2 (1^ 1"); Xuchil, 1'; Orizaba, 4 (2^ D. Mustela frenata perda (Merriam) Long-tailed Weasel Plates 25, 26, 27, 30, 37, 38 and 39 Putorius tropicalis perdus Merriam, Proc. Biol. Soc. Washington, 15:67, March 22, 1902. Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part). Mustela tropicalis perda. Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912. 1. American Mus. Nat. History 4. Berlin Zoological Museum 2. Museum of Comparative Zoology 5. Louisiana State University 3. Field Museum of Natural History 6. Texas Co-operative Research Collection MUSTELA FRENATA 367 Mustela tropicalis tropicalis, Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916. Mustela frenata perda, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936. Type. — Male, subadult, skull and skin; no. 100041, U. S. Nat. Mus., Biol. Surv. Coll.; Teapa, Tabasco, Mexico; March 31, 1900; obtained by E. W. Nelson and E. A. Goldman; original no., 14074. The skull (plates 25-27, 30) is unbroken and all the teeth are present and entire. The skin is well made and in good condition. Range. — Fifty feet (Chichen Itza) to 4000 feet (San Vicente) in Lower Tropical Life-zone south from southern Veracruz through southern Mexico into Guatemala. See figure 29 on page 221. Characters for ready recognition. — DifFers from M. f. nicaraguae in lesser extent of color of underparts (not more than 22 per cent of greatest extent of color of upper parts ) , black tip of tail more than 38 per cent of length of tail, and broader skull (in adult males, mastoid breadth more than 23.9 and zygo- matic breadth more than 27.4); from M. /. tropicalis in more restricted color of underparts (least width of color of underparts less than 28 per cent of greatest width of color of upper parts) and shorter tympanic bullae, which in males are less than 15; from M. /. goldmani by total length not exceeding 489, least width of color of underparts not exceeding 24 per cent of greatest width of color of upper parts, weight of adult skull less than 5 grams and basilar length less than 48.5. Description. — Size. — Male: The type specimen and another subadult from San Vicente, Chiapas, measure, respectively, as follows: Total length, 473 and 443; length of tail, 184 and 169; length of hind foot, 51 and 51.5. The tail is 62 and 64 per cent as long as the head and body. The length of the hind foot is greater than the basal length. Female: Estimates made from the dried skin of no. 218036 are: Total length, 375; length of tail, 140; length of hind foot, 40. The hind foot of no. 65422 from Catemaco also measures 40. The average differences in external measurements of the two sexes are: Total length, 83; length of tail, 37; length of hind foot, 11. Externals. — As described in Mustela frenata goldmani except that hairiness of foot soles is slightly less. Color. — As described in Mustela frenata goldmani except that: back near (n) Argus Brown or Carbon Brown, tone 3, of Oberthiir and Dauthenay, pi. 342; underparts Ochraceous-Buff. Least width of color of underparts, in four specimens, averaging 20 (extremes 18-22) per cent of greatest width of color of upper parts; black tip of tail, in two subadult males, averaging 48 (ex- tremes 46-49 ) per cent of length of tail-vertebrae. Skull and teeth. — Male (based on type specimen and subadult no. 132997 from San Vicente ) : See measurements and plates 25-27, 30; weight 4.4 grams (same for each); basilar length 45.7 (45.3 and 46.1); zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more or less than (approximately equal to) length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its oppo- 368 University of Kansas Publs., Mus. Nat. Hist. site; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum not greater than length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4/2 to 5/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below middle of m2. Female (based on two subadults, nos. 65422 and 218036): See measure- ments and plates 36-39; weight, 2.4 (2.3-2.5) grams; basilar length, 40.5 (40.4-40.6); zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than outside length of P4 and less than inside length of same; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla equal to one-third to one-half distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than (about equal to) length of rostrum. The skull of the female averages 48 per cent lighter than that of the male. Comparison of the skull of the male with that of M. /. nicaraguae has been made in the account of that subspecies. The skull of the male as compared with that of M. /. tropicalis has shorter tympanic bullae, deeper braincase at anterior margin of basioccipital, lesser zygomatic and palatal breadth and smaller P4 and ml. The skull of the female is larger in every measurement taken except those re- flecting width of the preorbital portion. This part is actually nar- rower but probably mainly because the females of perda are younger than those of tropicalis. Features in which three skulls of subadults of M. f. perda differ from the five adults of M. /. goldmani and show no overlap are: lesser basilar length, lesser weight, greater relative length of upper tooth-rows, greater rela- tive width of rostrum, greater relative length of rostrum, lesser mastoid and zygomatic breadths, lesser width, length and height of tympanic bullae; lesser outside length of P4 and greater relative depth of braincase at anterior margin of basioccipital and at poste- rior margin of Ml. Features in which per Jfl averages less are: length of tooth-rows, interorbital breadth, orbitonasal length, relative zygo- matic breadth, length of ml, outside and inside lengths of P4, width and length of Ml, and depth of skull at posterior margin of Ml. Features in which perda averages more than goldmani are: rela- tive interorbital breadth, relative mastoid breadth and depth of skull at anterior margin of basioccipital. The length of the inner MUSTELA FRENATA 369 half of Ml averages the same. As compared with goUhnani, the skull of the male of perda is shorter, otherwise generally smaller, but relatively broader except across the zygomatic arches, and relatively deeper. The anterior margins of the tympanic bullae project slightly less from the braincase and the squamosals imme- diately in front of these bullae are slightly more convex ventrally. Remarks. — Described by Merriam in 1902 as a subspecies of Putorius tropicalis, the form perda was regarded by Allen ( 1916:99) as not subspecifically distinct from P. t. tropicalis. This is the eastern, lowland subspecies of the Tropical Life-zone, corresponding to M. f. goJdmani of the higher mountains just as M. /. tropicalis corresponds to M. /. frenata and perotae of the high mountains and table land. The difference in size between perda and nicaraguae and between perda and tropicalis is slight. M. /. perda is slightly less richly colored than M. f. nicaraguae but has the color of the underparts more restricted and has a longer black tip on the tail. In these respects it is second only to M. /. panamensis among Central American weasels. Evidence of intergradation with goldmani is furnished by the specimens from Coban, Guatemala, and the nearby locality San Cristobal in Verapaz, Guatemala. Re- duced size as compared with goldmani suggests affinity with perda but the greater width of the light-colored underparts, which aver- ages 24 (extremes 18-32) per cent of the greatest width of the color of the upper parts, shows approach to goldmani. Farther north, in Chiapas, however, specimens of perda from San Cristobal and San Vicente are readily distinguishable from those of goldmani taken a few miles away at Pinabete and near Teopisca. The latter two localities are, however, several thousand feet higher than San Cristobal (Chiapas) and San Vicente. Two of the nine skulls (only 3 adult) examined for malformation of the frontal sinuses reveal infestation by parasites. Specimens examined. — Total number, 18, listed by localities from north to south, and unless otherwise indicated in the United States National Museum. Veracruz: Catemaco, 1. Tabasco: Teapa, 2 (1'). Chiapas: San Cristobal, 1; San Vicente, 1; no localit>' more definite than state, 1. Yucatan: Chichen-Itza, 1". Guatemala. Coban, 2 ( P, 1'); Finca la Providenci, S. Cristobal, Verapaz, 3'; central Guatemala, 1; no locality more definite than Guatemala, 5 (2'). 1. British Museum of Natural History 3. Berlin Zoological Museum 2. Museum of Zoology, Univ. Michigan 370 University of Kansas Publs., Mus. Nat. Hist. Mustela frenata nicaraguae Allen Long-tailed Weasel Plates 1, 25, 26, 27 and 30 Mustela tropicalis nicaraguae Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916. Putorius tropicalis, Allen, Bull. Amer. Mus. Nat. Hist., 24:661, 1908. Mustela frenata nicaraguae. Hall, Carnegie Instit. Washington Publ. 473: 109, November 20, 1936. Type. — Male, subadult, skull and skin; no. 30754, Amer. Mus. Nat. Hist., Matagalpa, Nicaragua; April 16, 1910; obtained by W. B. Richardson; original no., 712. The skull (plates 25-27, 30) of the type specimen lacks the entire right zygomatic arch. Otherwise it is complete. The teeth all are present and imbroken. The skin is complete and unfaded but only partly stuffed. Range. — Honduras and Nicaragua. Altitudinal and zonal limits unknown. See figure 29 on page 221. Characters for ready recognition. — Differs from M. /. costaricensis and M. f. goldmani in shorter black tip of tail (not more than 35 per cent of length of tail) and lesser width (usually not more than 7 mm.) of tympanic bulla; from M. f. perda in greater extent of color of imderparts (22 or more per cent of greatest width of color of upper parts), shorter black tip on tail (not more than 35 per cent as long as tail) and narrower skull, the mastoid breadth in adult males being less than 23.9 and the zygomatic breadth less than 27. Description. — Size. — Male: Average and extreme measurements of five subadults and one young (four from Matagalpa and one from San Rafel del Norte) are: Total length, 450 (420-480); length of tail, 178 (150-190); length of hind foot, 48 (46-50). Tail averages 65 (extremes 56-69) per cent as long as head and body. Length of hind foot (measurements from dried skins) more than basal length. Female: Measurements unrecorded. Externals. — As described in Mustela frenata goldmani, except that hairiness of foot soles (between tliat shown in figures 20 and 21) is less, sHghtly less even than in M. /. perda. Color. — ^As described in Mustela frenata goldmani except that: Back near (n) Argus Brown, or Carbon Brown, tone 4 of Oberthiir and Dauthenay, pi. 342. Underparts Ochraceous-Buff. Least width of color of underparts, in four males, young, subadult and adult, 24 (extremes 22-26) per cent of greatest width of color of upper parts; the corresponding per cent in one female is 32; black tip of tail, in two subadult males, averaging 29 (extremes, 28-30) per cent of length of tail-vertebrae; corresponding per cent in one female, 36. Skull and teeth. — Male (based on type specimen, one adidt topotype [?] and one subadult from San Rafel del Norte): See measurements and plates 25-27, 30. As described in Mustela frenata perda except that: Weight, 4.2 grams (estimated for adults); basilar length 45.0 (44.8-45.5); interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as viddth of four to five upper incisors; length of tympanic MUSTELA FRENATA 371 bulla not less than length of lower molar and premolar tooth-row; anterior margin of masseteric fossa below anterior margin of m2 or posterior to that tooth. Female: Skull unknown. Comparison of the skull of the male with that of M. /. costari- censis is made in the account of that subspecies. As compared with that of M. f. perda, which it most closely resembles, the skull of the male has a narrower, shorter rostrum, lesser interorbital breadth, lesser mastoid and zygomatic breadths and slightly shal- lower braincase, measured at anterior margin of basioccipital. The tympanic bullae are slightly less projected, at their anterior margins, from the braincase and the squamosal, directly anterior to each, is a little more convex ventrally. The skull of M. f. nicaraguae is, then, slightly shorter than that of M. /. perda and relatively nar- rower. Remarks. — When naming this form, Allen (1916:100) character- ized it as "Similar to M. tropicalis tropicalis but general colora- tion much darker and the white face markings somewhat reduced in area." In the sentence preceding the one quoted, Putorius tropi- calis perdus was placed as a synonym of Putorius tropicalis tropi- calis. M. f. nicaraguae and M. /. perda are nearly alike in color and color pattern but differ in cranial characters. M. /. perda and M. /. tropicalis are widely different in color and more especially in color pattern but differ only slightly in cranial characters. The aggregate difference between perda and nicaraguae is less than that between perda and tropicalis. All three are lowland forms and each is smaller than the adjacent highland forms, namely, M. /. goldmani, macrophonius, perotae and frenata. The weasels from Honduras definitely are not typical of nica- raguae as it is known from the specimens from Nicaragua itself. The specimens from the state of Tegucigalpa, Honduras, are larger. Some are darker than topotypical nicaraguae. The dorsal out- line of the skull is more nearly flat (less convex) in some. In these and several other differential features studied, the average of speci- mens from Tegucigalpa is intermediate toward goldmani, but everything considered the animals seem best placed with nica- raguae rather than with goldmani or perda, to which latter also, they show some resemblance. With better material from Nica- ragua and additional specimens from Salvador (here referred to goldmani) a restudy of all the material now referred to the three races named would be profitable. Aims of this restudy might be to determine if a highland race additional to goldrnani should be 372 University of Kansas Publs., Mus. Nat. Hist. recognized and if the lowland races perda and nicaraguae differ from one another in the way that the existing specimens indicate. In the five males from Matagalpa, the narrow white band in front of each ear is confluent with the color of the underparts on one side only in one specimen and on both sides in two specimens. None of these bands is confluent with the white patch between the eyes. A dark spot at the angle of the mouth is present on one side in one specimen. The corresponding area is dark colored in all other specimens but not separated from the color of the up- per parts. In the specimen from San Rafel del Norte the white bands are not confluent with the color of the underparts. The female from Mambacho has the mentioned bands confluent with the color of the underparts. This female approaches M. f. costa- ricensis in the dark color of the upper parts but has more extensive white facial markings than some specimens from much farther north. Like a female seen of M. /. costaricensis, this one has a "frosted" nape but the white hairs on the back of the neck are less numerous than in the female of M. /. costaricensis. M. f. nicaraguae in typical form, then, is thought of as a small, lowland, tropical subspecies only slightly differentiated from M. /. perda. By reason of its intermediate characters, it constitutes a link between the lowland forms, and the larger animals called M. f. goldmani and M. /. costaricensis. None of the four skulls from Nicaragua shows signs of infestation of the frontal sinuses by parasites. Specimens examined. — Total number, 16, listed by localities from north to south. Specimens are in the American Museum of Natural History, unless otherwise indicated. Honduras: Alto Cantoral, 2; Cerro Grande La Paz, 1. La Flor Archaga, 1'; Comayagiiela, 1'; vicinity of Tegucigalpa, 2; no locality more definite than Honduras, V. Nicaragua: San Rafel del Norte, 1; Matagalpa, 6; Ma[o]mbacho, 1. Mustela frenata costaricensis Goldman Long-tailed Weasel Plates 25, 26, 27, 28, 29 and 30 Mustela costaricensis Goldman, Proc. Biol. Soc. Washington, 25:9, January 23, 1912. Mustela brasiliensis, Gray, Ann. and Mag. Nat. Hist, 14(ser. 4):374, 1874. Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part). Putorius affinis, Merriam, N. Amer. Fauna, 11:31, June 30, 1896 (part). Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Lonnberg, Arkiv for Zoo!., 14(no. 4):16, 1921. Mustela frenata costaricensis. Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936. 1. Museum of Comparative Zoology 2. Berlin Zoological Museum MUSTELA FRENATA 373 Type.— Male, young, skull and skin; no. 13770/37149, U. S. Nat. Mus.; San Jose, Costa Rica; obtained by C. H. Van Patten. The skull (plates 28-30) is complete and unbroken. All teeth are present and unworn. The skin apparently has been remade. It lacks the distal two- tliirds of the tail. The head is somewhat shrunken. The color is possibly faded but if so only to a slight degree. Otherwise, the sldn is in good con- dition. The orange color of the underparts is so intense as to suggest that the full, adult pelage has not been acquired. No white markings are present on the face. There is no sex mark on the label attached to the skin but the size and proportions of the skull and the scrotal pouch on the skin prove that the specimen is a male. The presence of sutures on the dorsal face of the rostrum and the short, wide, and low sagittal crest show the specimen to be young. Range. — Costa Rica. Altitudinal and zonal range unknown. See figure 29 on page 221. Characters for ready recognition. — Differs from M. f. panamensis in lighter color of upper parts (tone 2 rather than tone 4 of Reddish Black of Oberthiir and Dautlienay, pi. 344) and longitudinally flat interorbital region of skull; from M. f. nicaraguae in darker color of upper parts (of Oberthiir and Dauthe- nay, tone 2 of pi. 344 rather than tone 4 of pi. 342) and greater width (more than 7) of tympanic bulla. Description. — Size. — Male: No collector's measurements available of fully grown animals. Estimated measurements of adult males: Total length, 470; length of tail, 165; length of hind foot (taken from dried skins of 3 adults), 52 (50-52). Tail estimated to average 55 per cent as long as head and body. Length of hind foot more or less than (about equal to) basal length. Female: A subadult or adult, from the Candelaria Mountains, and a sub- adult from Irazu, measure, respectively: Total length, 370, 385; length of tail, 130, 150; length of hind foot, 40, 31. Tail 59 per cent as long as head and body. Length of hind foot probably about equal to basal length. The estimated differences in external measurements of the two sexes are: Total length, 92; length of tail, 25; length of hind foot, 16 (probably average difference is less). Externals. — As described in M. /. panamensis (figure 21) except that foot soles are slightly more hairy. Color. — As described in Mustela frenata panamensis except that: back is near Reddish Black, tone 2 of Oberthiir and Dauthenay, pi. 344; chin, Hps, and throat white or whitish; remainder of underparts near (c) Ochraceous- Buff; color of underparts rarely extending distally onto toes of forefeet. Least width of color of underparts, in eleven specimens, averaging 23 (10-36) per cent of greatest width of color of upper parts; black tip of tail, in six speci- mens, averaging 36 (31-38) per cent of length of tail-vertebrae. Skull and teeth. — Male (based on 2 adults, no. 3.2.1.6. from vicinity of San Jose and no. 11408, U. S. Nat. Mus., from "Costa Rica"): See measure- ments and plates 25-30; weight, 5.9 grams; basilar length 49 +; zygomatic breadth more than distance between condylar foramen and Ml or than be- tween anterior palatine foramen and anterior margin of tympanic bulla; mas- toid breadth less than postpalatal length; postorbital breadth in undiseased skulls less than length of upper premolars (less than distance between pos- 374 University of Kansas Publs., Mus, Nat. Hist. tenor borders of P2 and P4) and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; inter- orbital breadth more or less than distance between foramen opticmn and an- terior margin of tympanic bulla; breadth of rostrum more or less (about equal to) length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors; height of tympanic bulla less than distance from its an- terior margin to foramen ovale; tympanic bulla longer or shorter than (about equal to) lower molar and premolar tooth-row and longer or shorter than (about equal to) rostrum; anterior margin of masseteric fossa directly below posterior border of m2. Female: Skull of adult unknown. Comparison of the skull of the male with that of M. f. panamensis has been made in the account of that subspecies. As compared with that of M. /. nicaraguae the skull of M. f. costaricensis is heavier and in every measurement taken is larger. The skull is generally more massive and it follows that most measurements of depth and width are greater in relation to the basilar length as well as actually greater. The individual teeth are larger and the tympanic bullae wider and at their anterior ends are more projected from the brain- case. Indeed the skull is more hke that of M. /. goldmani than like that of M. f. nicaraguae. Remarks. — The half dozen ill-prepared skins, with partial skulls inside, of this form in the United States National Museum long were referred either to Mustela brasiliensis or Mustela afinis. It was not until 1912 when Goldman studied these specimens that the distinctive characters of the Costa Rican weasel were recognized and made the basis of the name costaricensis. M. f. costaricensis is well differentiated from M. f. nicaraguae and M. /. goldmani which occur to the northward and from M. f. pana- mensis which occurs to the southward and is a large, heavy-skulled, dark-colored animal with white facial markings restricted or ab- sent. In the type specimen and the female from the Candelaria Mountains the white facial markings are only narrow facial bars or a few white hairs, but in the young male from Cervantes there is a well developed bar 6 millimeters wide on each side of the face and a separate naso-frontal spot, 10 x 12 mm. The young female from Cachi has a V-shaped frontonasal spot, on the right side of the face a white bar 5 mm. wide and 17 mm. long connected with the color of the underparts, and on the left side a white spot in front of the ear and another between the ear and eye. White facial markings were not recorded in the other specimens. The color of tlie upper parts is only a little less dark than those of M. /. panamensis. Owing MUSTELA FRENATA 375 to the numerous white hairs on the dorsal side of the neck, the nape of the female from the Candelaria Mountains has a frosted appearance not present in other specimens. M. /, costaricensis is a large animal and among its geographic neighbors is approached in size only by a specimen of panamensis from Boquete, Panama. Also the young male from Cervantes sug- gests panamensis in the less flattened interorbital region, but even so is more like costaricensis. The small size of two young males, one from Navarro and the other from the vicinity of San Jose, is suggestive of M. /. nicaraguae. However, the large size of most of the specimens and the configuration of the skull are more as in M. f. goldmani than in M. /. nicaraguae and thus suggest that the known specimens are of high mountain subspecies. The long black tip of the tail is another point of resemblance to M. f. goldmani, the high mountain subspecies to the north. Perhaps in the lowlands of Costa Rica, there are weasels of another subspecies. Of the eight skulls examined for malformation of the frontal sinuses, each of the two adults and two subadults shows signs of having the frontal sinuses infested with parasites. Specimens examined. — Total number, 14, listed by localities from north to south. Costa Rica: Irazii (Frasu or Irasu on label), 3000 M., 1'; Cervantes, 1'; San Jose, 1'; vicinity of San Jose, 2*; Azahar Cartago, 1°; Tucurrique, 1*; Cachi, 1*; El Munco [= Muneco?] (Rio Nivarro [= Navarro?]), 4000 ft, 10 mi. S Cartago, Caribbean Slope, V; Navarro, 1'; Candelaria Mts., 1'; no locality more definite than Costa Rica, 3'. Mustela frenata panamensis Hall Long-tailed Weasel Plates 1, 25, 26, 27, 28, 29 and 30 Mustela frenata panamensis Hall, Proc. Biol. Soc. Washington, 45:139, September 9, 1932; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936. Mustela brasiliensis, Alston, Biol. Cent. Amer., Mammalia, p. 78, 1879. Putorius affinis. Bangs, Bull. Mus. Comp. Zool., 39:49, April, 1902; Holli- ster, Proc. Biol. Soc. Washington, 28:143, July 10, 1914. Mustela affinis, Goldman, Proc. Biol. Soc. Washington, 25:10, January 23, 1912; HolUster, Proc. Biol. Soc. Washington, 28:143, July 10, 1914. Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Goldman, Smithsonian Miscel. Col., 69 (no. 5):161, 1920. Type. — Female, subadult, skull and skin; no. 170970, U. S. Nat. Mus., Biol. Surv. Coll.; Rio Indio, Canal Zone, near Gatiin, Panama; February 17, 1911; obtained by E. A. Goldman; original no. 20897. 1. Berlin Zoological Museum 5. Naturhistorika Ricksmuseum, Sweden 2. American Mus. Nat. History 6. Museum of Zoology, Univ. Michigan 3. United States National Museum 7. Museum of Comparative Zoology 4. British Museum of Natural History 376 University of Kansas Publs., Mus. Nat. Hist. The skull is complete and unbroken. The left lower incisor is broken oflF but all the other teeth are present and entire. The skin is well made and seems to be in faded, worn, first, adult pelage. Range. — Sea level (type locality) to 5800 feet (Boquete, see Bangs [1902: 49]); Upper Tropical and Lower Tropical life-zones of Panama. See figure 29 on page 221. Characters for ready recognition. — Difi^ers from both M. f. meridana and A/. /. costaricensis in darker tone (tone 4 of Oberthiir and Dauthenay, pi. 344) of color of upper parts and in convex dorsal outfine of skull (Compare figures of mentioned subspecies on plates 25-27). Description. — Size. — Male: Two adults from Boquete in the Museum of Comparative Zoology, nos. 10112 and 10113, measure, respectively, as follows: Total length, 480 and 400; length of tail, 170 and 143; length of hind foot, 52 and 43. Hind feet of two other adult males measure 46 on dried skins. Tail, in two specimens mentioned above, is 55 and 56 per cent as long as head and body. Length of hind foot, in each of three adults, slightly longer than basal length. Corresponding measurements of no. 178970 from Mt. Pirre are: 422, 164, 50. Tail 64 per cent (same per cent as in young male, no. 137514 from Boquete) as long as head and body, and hind foot longer than basal length. Female: An adult and a young from Chiriqui, nos. 18434 and 18435 (Acad. Nat. Sci. Philadelphia), measure, respectively: Total length, 372, 389; length of tail, 138, 144; length of hind foot, 42, 41. The type speci- men measures: Total length, 408; length of tail, 159; length of hind foot, 46.5. Tail 64 per cent as long as head and body, and hind foot longer than basal length. The average difi^erences in external measurements of the two sexes from the vicinity of Boquete are: Total length, 59; length of tail, 15; length of hind foot, 6. Externals. — Longest facial vibrissae black and extending beyond posterior border of ear; carpal vibrissae wholly, or in part, black and extending as far as hypothenar pad; hairiness of foot-soles as shown in figure 21. Color. — Usually, posterior fourth of each upper hp and sometimes few hairs in front of ear, white; sides and top of head and neck posteriorly to, or behind, shoulders, black; dark areas at angles of mouth confluent with color of upper parts; tip of tail, black; remainder of upper parts near (n) Bay of Ridgway and Reddish Black, tone 4, pi. 344 of Oberthiir and Dauthenay; chin and lips, whitish; remainder of underparts Warm Buff or near (16' c) Ochra- ceous-Buff; near (12) Salmon-Orange in juveniles and small young; color of underparts extending distally on posterior sides of forelegs to wrists, but not to soles, and on hind legs to or slightly below knees. Least width of color of underi^arts, in seven specimens, averaging 18 (extremes 11-28) per cent of greatest width of color of upper parts; black tip of tail, in five adults and subadults, averaging 45 (extremes 41-50) per cent of length of tail-vertebrae. Skull and teeth. — Male (based on three adults from Boquete): See measurements and plates 25-30; weight, 5 (4.5-5.4) grams; basilar length, 45.2 (42.8-48.3); zygomatic breadth more or less than distance between condylar foramen and Ml or than between anterior palatine foramen and ante- rior margin of tympanic bulla; mastoid breadth less than postpalatal length; MUSTELA FRENATA 377 postorbital breadth more than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum poste- rior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum approxi- mately same (more or less than) length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4/2 to SJz upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or slightly less than (approximately equal to) length of lower molar and premolar tooth-row or length of rostrum; anterior margin of masse- teric fossa directly below posterior fourth of m2. Female ( based on subadult, type specimen and one adult from Siola ) : See measurements; weight, 3.3 and 2.1 grams; basilar length, 41.3 and 39.3; zygomatic breadth more than distance between condylar foramen and Ml and more or less than ( about equal to ) that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than com- bined length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than length of P4 (less in the adult); anterior margin of tympanic bulla as far posterior to foramen ovale as width of five upper incisors; height of tympanic bulla less than (about half) distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of lower molar and premolar tooth-row or than rostrum. The skull of the one adult female from ChiriquI is 58 per cent lighter than the average of the two adult males. The skull of the male of M. /. panamensis as compared with that of M. f. meridana, is heavier and averages larger in nearly every measurement taken. Relative to basilar length, tooth-rows, orbi- tonasal length, interorbital breadth and zygomatic breadth averag- ing narrower. Mastoid breadth always narrower. Tympanic bullae longer, narrower, and usually slightly less protruded. P4 and ml larger. Dorsal outline of skull, viewed laterally, more convex. Postorbital breadth actually and relatively greater. Postorbital processes, mastoid processes, and sagittal crest not so well de- veloped. Differences between skulls of females, in so far as known, similar to those described between males. As compared with M. /. costaricensis, M. f. panamensis has a lighter skull averaging smaller in every measurement taken except interorbital breadth, which is greater. Relative to basilar length, width of rostrum, interorbital breadth and depth of skull at plane of upper molars, less. Tympanic bullae shorter, narrower, less protruded. P4, Ml, and ml larger. Dorsal outHne of skull, viewed laterally, more convex. Postorbital breadth relatively and actually greater. Postorbital processes, mastoid processes, sagittal crest and lambdoidal crest less developed. No skull of an adult female of M. f. costaricensis is available for comparison. 378 University of Kansas Publs., Mus. Nat. Hist. Remarks. — This subspecies had not been recognized by previous workers because specimens from Panama were supposed to be Mustela affinis Gray up imtil 1916, when Allen (1916:100) re- stricted the type locality of M. affinis to Bogota, Colombia. At that time Allen referred specimens from Panama to Mustela affinis cos- taricensis, and Goldman (1920:161) followed Allen. The specimens examined show much variation. Part of this is geographic variation. For instance the specimens from Boquete approach M. /. costaricensis in size more than do those from fartlier south. Too few adult females have been seen to ascertain the amount of secondary sexual variation. Bangs (1902:49) suggested that the sex of no. 10113 was wrongly recorded and that it was not really a male. If so, this would reduce the range of apparent vari- ation in size of males from Boquete by half and bring it into accord with the amount normally existing in adult males from one locality. No. 10113 is adult but the skin shows no mammae which would prove it to be a female instead of a pigmy male. Although even smaller than 10113, the type specimen is so much larger than fe- males of M. f. meridana that I have wondered if it is correctly sexed. However, the fact that it was sexed by E. A. Goldman, a collector of wide experience, lessens the possibility that a mistake was made. The color of the underparts is more restricted in panamensis than in any other subspecies of the species. Excluding the speci- men from Mt. Pirre, the least width of color of the underparts averages 16 ( extremes 6-24 ) per cent of greatest width of the color of the upper parts. This feature, together with the black color, imparts an appearance to the Panama weasel that is strikingly Hke that of a mink. M. /. panamensis is one of the two blackest weasels; M. /. aureoventris is the other. Each of these subspecies occurs in a region of heavy rainfall and there clearly is a positive correla- tion between high humidity and intensity of color. The black tip of the tail, as regards extent, here reaches the maximum attained among Central and South American weasels. The foot soles are less hairy than in any other member of the subgenus Mustela. The tympanic bullae are lower and less inflated than in any other sub- species of the species. Adequate specimens from central and southern Panama may re- veal the existence of one or more additional subspecies since ani- mals from each of the three localities now represented differ from those from the other two and some of these differences are corre- lated with geographic position. However, specimens from all three MUSTELA FRENATA 379 localities agree in several features. For example all of them have the dorsal outline of the skull highly convex, transversely, and, more especially, longitudinally. In this respect they are sharply dif- ferentiated from any other American weasel. Nevertheless, M. /. panamensis is clearly a link between the North and South Ameri- can subspecies and panamensis intergrades with the adjacent sub- species. The large size of the skull and teeth and the slightly more ventrally projected tympanic bullae of no. 10112 from Boquete approach features seen in M. /. costaricensis. The smaller size of skull and teeth of no. 178970 from Mt. Pirre are points of resem- blance to M. f. meridana. The type specimen was selected from a region where M. f. pana- mensis is thought to have its distinctive characters well developed. The specimen is not adult and, therefore, does not show as many differential characters as does a nontypical adult from Boquete. Nevertheless, the majority of the above mentioned differential char- acters are shown by the type specimen and an adult from the same place would, it is judged, show all the differential characters better than would an adult from Boquete. Of the 11 skulls examined, 6 show no signs of having had the frontal sinuses infested with parasites. Specimens examined — Total number, 19, listed by localities from north to south and unless otherwise indicated in the United States National Museum. Panama: Boquete, 10 (3\ l^ 1', 3*, 1°); Rio Garicha [e], 5300 ft., V; Siola, 1°; Chiriqui, 1°; Rio Indio, near Gatun, 1; Mt. Pirre 3 (2°); Caloverora, 1' (locality not found, possibly misspelling of Calovebora); no locality more definite than Panama, V. Mustela frenata meridana HoUister Long-tailed Weasel Plates 25, 26, 27, 37, 38 and 39 Mustela meridana HoUister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914. Putorius affinis, Robinson and Lyon, Proc. U. S. Nat. Mus., 24:147, October, 1901; Allen, BuU. Amer. Mus. Nat. Hist., 30:256, December 2, 1911. Mustela affinis, Osgood, Field Mus. Nat. Hist. Publ. 155, zool. ser. 10:61, January 10, 1912. Putorius macrurus, Allen, Bull. Amer. Mus. Nat. Hist, 31:92, April 19, 1912. Mustela affinis affinis, Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916 (part). Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916 (part). Mustela frenata meridana. Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939. 1. Museum of Comparative Zoology 4. Berlin Zoological Museum 2. California Academy of Sciences 5. British Museum of Natural History 3. American Mus. Nat. History 6. Acid. Nat. Sciences of Philadelphia 380 University of Kansas Publs., Mus. Nat. Hist, Type. — Male, subadult, skull and sldn; no. 123341, U. S. Nat. Mus., 16.30 meters elevation, Montes de Merida, near Merida, Venezuela; August 14, 1903; obtained by S. Briceno. The skull (plates 25 and 26) lacks the right exoccipital condyle and pos- terior half of the right zygomatic arch. The teeth all are present, unworn and entire. The sldn is well made and complete. Range. — Near sea level (San Julian) to 8500 feet (Montes de Culata, Merida, Venezuela), and 9000 feet (Santa Elena, Colombia). Temperate to Subtropical life-zones of Venezuela and northern and western Colombia. See figure 29 on page 221. Characters for ready recognition. — Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, presence of p2 and two roots, rather than one root, on P2; from M. fre- nata panamensis in lighter color of upper parts (tone 3 rather than tone 4, pi. 344, Reddish Black, Oberthiir and Dauthenay), flat rather than convex dorsal outline of skull immediately behind postorbital processes (see pi. 27); from M. /. affinis, in males, by lesser average breadth and length of skull and greater actual and relative size (see measurements) of facial part of skull; from M. f. aureoventris, in males, by lighter upper parts (tone 3 rather than tone 4, pi. 344, Reddish Black, Oberthiir and Dauthenay) and by smaller skull and teeth (basilar length less than 45, length of ml less than 6.3, width of Ml less than 4.8, outside length of P4 less than 5.7). Description. — Size. — Male: Average and extreme measurements of topo- types (as recorded by collectors on labels, and so uniform as to show them not to be accurate to within more than 5 mm.) are as follows: Total length, 434 (410-460); length of tail, 164 (150-180); length of hind foot, 50 (no variation in collectors' measurements). Tail averages 61 per cent as long as head and body. Length of hind foot more than basal length. Correspond- ing measurements of no. 22191, a young male from Merida, measured by Osgood or Conover, are 439, 165, 54. The adult male no. 18703, from Paramo de Tama (eastern boimdary of Venezuela) has the following measurements vmtten on the label by Osgood: 404, 150, 47. Female: Average and extreme measurements of topotypes (as recorded by collectors on labels and so uniform as to show them not to be accurate to within more than 5 mm.) are as follows: Total length, 347 (320-370); length of tail, 128 (120-130); length of hind foot, 40 (no variation in col- lectors' measurements). Tail averages 57 per cent as long as head and body. Length of hind foot more than basal length. Two females, adult no. 11034 and young no. 11033 from Cincinnati, Santa Marta, Colombia, measured by M. A. Carriker, Jr., measure, respectively, as follows: 371, 330; 140, 140; 38, 36. No. 14463, adult, from Rio Zapata, Colombia, measured (by J. H. Batty), 315, 138, 39. No. 32182, adult, from Mira Flores, Cauca, Colombia, meas- ured (by W. B. Richardson), 375, 150, 43. The average differences in external measurements of the two sexes, at Merida, are: Total length, 87; length of tail, 36; length of hind foot, 10. Externals. — Longest facial vibrissae black (few rarely white) and extending beyond ear; carpal vibrissae colored like underparts or upper parts, and not extending beyond apical pad of fifth digit; hairiness of foot soles shghtly greater than shown in figure 21. MUSTELA FRENATA 381 Color. — As described in Mustela frenata panamensis except that: Pos- terior foiirt:h of each upper hp rarely, and small spot in front of ear usually, white; black of head proper not extending back of ears and grading in- sensibly into color of upper parts; anterior half of upper parts of adults "frosted" with numerous white hairs (tick bites?), upper parts near (n) Bay or tone 2 of Reddish Black (pi. 344, Oberthiir and Dauthenay) or tone 3 in freshest, unfaded pelage. Least width of color of underparts (in ten males from Merida) 20 (17-23) per cent of greatest width of color of upper parts. Black tip of tail, in same series, 60 to 75 mm. long, thus longer than hind foot and 41 (40-44) per cent as long as tail-vertebrae. Skull and teeth. — Male (based on tj'pe specimen and seven topotypes, five adults and three subadults): See measurements and plates 25-27; weight, 4.1 (3.8-4.3) grams; basilar length, 43.6 (42.3-44.3); zygomatic breadth more than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadtli greater than length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum greater than length of tympanic bulla; least width of palate greater than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or less than (approximately equal to) alveolar length of lower molar and premolar tooth-row and shorter than rostrum; anterior margin of masseteric fossa posterior to m2 and confined to posterior third (34 per cent average, 32 minimum, 37 ma.\imum) of mandible. Female (based on four adult topotypes): See measurements and plates 37-39; weight (no. 143665), 2.3 grams; basilar length 37.2 (36.3-38.2); zygomatic breadth more or less than distance between condylar foramen and Ml or than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth (sinuses badly infested with parasites) more than length of upper premolars or vddth of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than length of P4; tympanic bulla as far posterior to foramen ovale as combined viddth of 4 to 5 upper incisors; height of tympanic bulla less than (one half to three fourths of) distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row. The skull of the female is 44 per cent Ughter than that of the average male. Comparisons of the skull with those of M. /. panamensis and affinis have been made in the accounts of those subspecies. As compared with the skull of the male of M. /. aureoventris, that of meridana averages smaller in every measurement taken. Indeed, none of the skulls of meridana equals that of aureoventris in basilar length, length of tooth-rows, length of tympanic bulla, depth of skull at anterior margin of basioccipital or at posterior 382 University of Kansas Publs., Mus. Nat. Hist. margin of upper molars, or measurements of teeth. Relative to the basilar length, most of the measurements are greater in meri- dana. Exceptions are the relative length of the tooth-rows, and the two measurements of depth of the skull which average less. Remarks. — In 1914 when Hollister named this weasel he com- pared it with M. /. affinis and most of the differential characters which he ascribed to meridana were merely "more than" or "less than" in affinis. In affinis, HoHister included specimens from Chiriqui, Panama, and the coast of Venezuela. The specimens from these three places were referred by Allen (1916:101) to Mustela affinis costaricensis, and he restricted (op. cit. :100) the type locality of Mustela affinis affinis to Bogota, Colombia, and synonymized Mustela meridana with M. a. affinis. Hollister prob- ably would not have named meridana had he had specimens from Bogota for comparison and had he regarded them as topotypes of affinis for the difference is slight. Nevertheless, within the large geographic range of M. /. meridana there is some geographic varia- tion. There is more of such variation in the color of the pelage than in shape and size of the skull. The specimen from San Juhan is darker than the average and in this respect approaches true panamensis. San Juhan is situated at a relatively low elevation on the coast of Venezuela. M. /. meridana so closely resembles M. f. affinis that the writer has no quarrel with anyone who would synonymize meridana. However, as represented by topotypes, the two races unquestion- ably are, on the average, different, and specimens from the south- eastern part of the range of affinis probably are individually distinguishable from topotypes of meridana. Variation in the skulls of the series from Merida is relatively small. This apphes to both males and females. The external measurements recorded by native collectors are not accurate to within more than five millimeters but, considering this, variation in external measurements also seems to be slight. The difference in size of the two sexes appears to be uniformly greater than in weasels from Central America. The twenty-six topotypes show that the color and color pattern are relatively uniform. AU are of nearly the same tone except juveniles or young which are, as in the case of panamensis, much brighter colored on the under- parts. Also, the young have darker-colored upper parts. The adults, without exception, have numerous white hairs scattered over the back of the head, neck and between the shoulders. I MUSTELA FRENATA 383 have no trustworthy evidence to support the suggestion that these white hairs are the results of tick bites or that they are caused by other parasites which damage the hair folUcles. The white facial markings vary relatively little in the 45 specimens carefully ex- amined in this regard. Also, the variation in color pattern of the two sides of the head is small. Indeed, within rather narrow limits, the color of the two sides of the head is the same in every speci- men except two. In these two the white spots anterior to the ears are confluent with the color of the underparts. Only one specimen, no. 21342, has a white spot between the eyes and this spot is small. Ten of the twenty-six specimens have a definite white spot or band in front of each ear. Two specimens have such a spot on one side only. The dark spots at the angles of the mouth are present on two sides in three specimens and on one side only in three others. The mentioned spots are, then, present nine out of a possible fifty-two times. When the spots are absent, dark color usually is present in the required area but is con- fluent with the color of the upper parts. A young male from San Julian, Robinson and Lyon (1901:147) state ". . . was shot . . . as it ran over some bowlders in a ravine. Its eyes shone with the same greenish hght as do the eyes of our common weasel, and it emitted the same strong odor". No. 14463, Am. Mus. Nat. Hist., from Rio Zapata, Colombia, according to data on the label, was "taken in timber belt in valley in baUc hills" and the native name is Cosonebi. Two specimens taken on the Paramo de Tama, head of Tachira River, Venezuela and Co- lombia are commented on by Osgood (1912:61) as follows: "One . . . was caught in a steel trap baited with birds and set by the side of a rushing mountain stream. . . . The other was shot in midday as it came prowling about our Tiouse' in the clearing. • • • Of the thirty-three skulls before me, twelve have the frontal sinuses malformed by parasites. These twelve include most of the adults for few of the subadults and fewer of the young show pathologic conditions in the frontal region. Note on localities. — Several of the localities in Colombia men- tioned in "Specimens examined" are described and located by Chapman (1917:640-656, pi. 41) in his "Distribution of Bird-hfe in Colombia." Place names for Colombia on labels, not found on any map, or duplicated names of which I can not certainly select one, are Rio Barrotow, Rio Oscuro, Rio Zapata, Rio Japata, Guasca 384 University of Kansas Publs., Mus. Nat. Hist. and El Baldro. Sonson may or may not be the town of that name situated some eighty miles northwest of Bogota and on the east flank of the Central Andes west of the Magdalena River on the drainage of the Cauca River. In Venezuela most of the specimens from Merida are labeled 1630 meters, Montes de Merida. San Julian is some seven miles east of La Guaira (see Robinson and Lyon, 1901:136). San Esteban is located a little way back from the coast between Puerto Cabello and Valencia. Paramo de Tama is on the Venezuelan-Colombian border near the source of the Tachira River (see Osgood, 1912:35). Mt. Duida is shovm as at 3° 30' N and 65° 40' W by Chapman (1931:13) and Mt. Auyan- tepuy as near 5° 15' N and 62° 50' W by Chapman (1937:760). Specimens examined. — Total number, 78, arranged by localities from north to south and unless otherwise indicated in the British Museum of Natural History. Venezuela: San Juhan, 1'; Caracas, 2; Galipare, Cerro del Avila, 6500 feet, 1; San Esteban, V; Merida, 45 (10', 14% 10^ 2\ 2\ T); Pdramo de Tama, 1*. Colombia: Paramo de Tama, 1*; Cincinnati, 3'; Valdiva, 3800 ft., 1; Me- delHn, 2; 7200 ft., Barro Blanco, 1'; Santa Elena, 9000 ft., 1=; Santa Elena, 1'; Sonson, 2 (1', T); Mt. Auyan-tepuy, 1'; Pueblo Rico, 5200 ft., 1^ Mira Flores, 1'; Jerico, near Cauca River, 1; Tomel, 20 mi. NE Quitichao, 1; Mt. Duida, 1'; El Tambo, Cauca, 1°; El Baldro, 1'; Rio Japata, 2'; Rio Zapata, 4500 ft., 1; Rio Oscuro, 3300 ft., 1; Rio Barrotow, 3300 ft., 1; Guasca, 1'; no locality more definite than Colombia, 1. Mustela frenata affinis Gray Long-tailed Weasel Plate 30 Mustela affinis Gray, Ann. and Mag. Nat. Hist, 14(ser. 4):375, 1874. Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part). Putorius affinis, Merriam, N. Amer. Fauna, 11:31, June 30, 1896. Mustela affinis affinis, Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916; Allen, Bull. Amer. Mus. Nat. Hist., 35:220, May 31, 1916. Mustela frenata affinis, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939. Type. — Male, adult, skull with skin; no. 54.1.11.3 (skull originally num- bered 195d, later 54.6.3.4), Brit. Mus. Nat. Hist.; Colombia [given as new Granada in original description!; purchased from Mr. S. Stevens. Type locality restricted by Allen (1916:99) to Bogota, Colombia. The skin is in a good state of preservation and has been made over into a conventional study specimen from a mount on exhibition. Exposure to light when mounted probably accounts for the faded color. The skull (plate 30) lacks the middle 9 mm. of the right zygomatic arch, occiput, basioccipital and 1. United States National Musexim 5. Collection of Alex Walker 2. American Mus. Nat. History 6. Naturhistorika Ricksmuseum, Sweden 3. Berlin Zoological Museum 7. Carnegie Museiun 4. Field Museum of Natmal History 8. Museum of Comparative Zoology MUSTELA FRENATA 385 posterior two-thirds of the left tympanic bulla. The teeth all are present and entire. Range. — Four thousand six hundred feet (Quetame) to 9154 feet (El Car- men), Tropical to Temperate life-zones of eastern Andes of Colombia. See figure 29 on page 221. Characters for ready recognition. — Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, by presence of p2 and by two roots rather than one root on P2; from A/, frenata meridana, in case of males, by, on average, greater breadth and length of skull and lesser actual and relative size (see measurements) of facial part of skull; from M. /. aureoventris by lighter-colored upper parts (tone 2 rather than tone 4, pi. 344, Reddish Black of Oberthiir and Dauthenay); from M. /. macrura by darker color (Reddish Black, tone 2, pi. 344, Ober. and Dauth., rather than Chocolate, tone 3, pi. 343, Ober. and Dauth.). Description. — Size. — Male: Measurements in life, estimated from dried skins, are: Total length, 455; length of tail, 175; length of hind foot, 52. Pro- portions of parts supposedly as described in Mustela frenata meridana. Female: Estimates from two dried skins: Total length, 365; length of tail, 135; length of hind foot, 43. Proportions of parts supposedly as described in Mustela frenata meridana. The estimated differences in external measurements of the two sexes are: Total length, 90; length of tail, 40; length of hind foot, 9. Externals. — As described in Mustela frenata meridana. Color. — As described in Mustela frenata panamensis except that: posterior fourth of each upper lip and spot in front of each ear white in approximately half of the specimens; black of head proper not extending back of ears and grading insensibly into color of upper parts; upper parts near (n) Bay, or tone 2 of Reddish Black (pi. 344, Oberthiir and Dauthenay). Least width of color of underparts ( in five males from vicinity of Bogota ) 24 ( 15-29 ) per cent of greatest width of color of vipper parts. Black tip of tail, in same series, 60 to 75 mm. long, thus longer than hind foot and averaging 38 per cent as long as tail-vertebrae. Skull and teeth. — Male (based on three adults and two subadult topotypes) : See measurements and plate 30. As described in Mustela frenata meridana except that: Weight, 4.5 grams (estimated); basilar length 45.8 ±; inter- orbital breadth not greater than distance between foramen opticum and an- terior margin of tympanic bulla ( type as in meridana where interorbital breadth is more than distance between foramen opticum and anterior margin of tympanic bulla); least width of palate not less than length of P4; masseteric fossa confined to posterior two-fifths (38 to 40 per cent; average 39 per cent) of mandible and not extended anteriorly to middle* of m2. Female: No adults examined. As compared with M. f. meridana the skull of the male is larger, to the average amount of 2.2 mm. in basilar length and 1.2 mm, in zygomatic breadth of adults; length of tooth-rows and mastoid breadth average greater but relatively less; breadth of rostrum, interorbital breadth and orbitonasal length average actually and 13—3758 386 University of Kansas Publs., Mus. Nat. Hist. relatively less. Thus the skull of affinis is longer and broader, but the facial region is actually, as well as relatively, smaller. As com- pared with the skull of the male of M. f. aureoventris, that of M. f. affinis is about the same in basilar length. However, in no speci- men of affinis are the measurements of length of tooth-rows or breadth of rostrum, actually, or relatively, as great as in aureo- ventris. The same is true of all measurements taken of Ml, P4 and ml. The specimens from the vicinity of Quito and north of there, although referred to macrura, are nearly as dark as typical affinis, approach affinis in cranial characters, and indicate inter- gradation of affinis w^ith macrura. Remarks. — Mustela affinis was named by John Edward Gray in 1874 (p. 375) on the basis of a specimen from New Granada. Al- though usually synonymized with Mustela brasiliensis by later au- thors until 1896 when Merriam (1896:31) apphed the name to weasels from Costa Rica, nearly all the South American and sev- eral of the Central American weasels have, at one time or another, had Gray's name, affinis, applied to them. Gray, in 1865 (p. 115) when giving measurements of Mustela aureoventris, probably men- tioned the specimen, that later became the holotype. In 1916 (p. 98) Allen restricted the type locality to Bogota, Colombia. Allen's action was a necessary procedure in clearing up the systematics of South American weasels and was based on good grounds. As set forth by Allen {loc. cit.), and more in detail by Chapman (1917:642), Bogota has long been the shipping point for Colombian vertebrate specimens, many of which were obtained in the moun- tains to the east. Allen (1916A:220) quotes Thomas as saying that the type specimen was purchased from Stevens at about the same time that a number of Colombian birds were purchased from the same dealer. Also, specimens from Bogota agree with Gray's de- scription of the type specimen. Mustela frenata affinis, as here defined, constitutes one of the sev- eral slight geographic variants met with, on the sides of, and be- tween, the three north and south mountain chains of Colombia. The others are lumped under the name Mustela frenata meridana. M. f. affinis, in common with specimens from the northern part of the range of macrura has large teeth. Weasels of all of the region from Quito to Bogota have large teeth. To the north there is the smaller- toothed meridana and to the south the smaller-toothed macrura grading into the still smaller-toothed agilis, and boliviensis. Two skins, without corresponding skulls, from Caqueta are lighter colored than any others of affinis; possibly the skins are faded by MUSTELA FRENATA 387 exposure to light. Since they probably come from an elevation of less than 1000 feet in the Amazonian region, they may pertain to another subspecies. Complete, unbroken, skulls of affinis are needed to ascertain the degree to which affinis and meridana differ in cranial features. The several specimens from the immediate region of Bogota show well the color and the color pattern but lack collectors' measure- ments. None of the ten skulls examined shows malformation of the frontal region due to infestation of the frontal sinuses by parasites. Pos- sibly three of the four adults were infested, although not severely. Specimens examined. — Total number, 27, arranged by localities from north to south and unless otherwise indicated in the United States National Museum. Colombia: El Cannen, 1^ W. Cundinamarca, 1'; Muzzo [= Muzo ?], 1*; Bogota, 1; Castillo, near Bogota, 1°; Fambrias, near Bogota, 1'; Bogota district, T; Choachi, 9 {l\ 2^ T); Paramo de Choachi, 2 (1', T); Laguna del Verjon (=City ot Bogota), 1'; Quetame, 2'; Fusagasuga, 1; Caqueta, 2'; no locality more definite than Colombia, 3 ( 1^ ) . Mustela frenata aureoventris Gray Long-tailed Weasel Plates 27, 28 and 29 Mustela aureoventris Gray, Proc. Zool. Soc. London, 1864:55, pi. 8, 1864; Gray, Proc. Zool. Soc. London, 1865:115, 1865. Putorius (Gale) brasiliensis var. aequatorialis Coues, Fur-bearing animals, p. 142, 1877, part ? ("merely as a substitute for Gray's [supposedly] preoccupied name", that is, aureoventris). Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916 (part). Mustela macrura, Lonnberg, Arkiv for Zool., 14 (no. 4): 11, 1921 (part ?). Mustela frenata aureoventris. Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939. Type. — Probably female, juvenile, skull with skin, no. 64.6.6.3 (formerly 1432a), British Mus. Nat. Hist.; probably Subtropical Life-zone of western Ecuador (locahty given as Quito, probably because received from that place). The skin, once exhibited as a mount, has lost some hair from the back and other parts of the body and is not suitable for remaking into a conventional study specimen. The skull lacks the occiput, basioccipital, premaxillae, upper incisors, two of the lower incisors, all of the canines, premolars 2/2 on both sides, right P3, left p3, and has the left jugal mesially defective. The premolars present are not all fully emerged. Range. — Pacific coastal regions of Ecuador and Colombia; Subtropical and Tropical hfe-zones. See figure 29 on page 221. 1. American Mus. Nat. History 4. Berlin Zoological Museum 2. Museum of Comparative Zoology 5. Paris Museum 3. British Museum of Natural History 388 University of Kansas Publs., Mus. Nat. Hist. Characters for ready recognition. — DifFers from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, by presence of p2 and by two rather than one root on P2; from Mustela frenata macrura by Reddish Black, tone 4, plate 344 rather than Chocolate, tone 3, pi. 343 (of Oberthur and Dauthenay), or slightly darker color of upper parts; from M. /. affinis and M. /. meridana by darker color (tone 4 rather than tone 2, Reddish Black of Ober. and Dauth.) of upper parts and larger size of teeth (Ml with length more than 2.4 and breadth more than 4.7; P4 with outside length more than 5.9; length of ml more than 6.2). Description. — Unless otherwise stated, information concerning this sub- species is derived from the one referred specimen available, a young male, no. 34677, Amer. Mus. Nat. Hist. Size.— Male: Total length, 470; length of tail, 160; length of hind foot, 50. Tail 51 per cent as long as head and body. Female: Not known. Externals. — Longest facial vibrissae black and reaching beyond ear. Carpal vibrissae reaching to or beyond apical pad of fifth digit; hairiness of foot soles slightly less than shown in figure 20. Color. — Sides and top of head and neck posteriorly to shoulders black; white facial markings represented by only five white hairs anterior to right ear, one anterior to left ear and three far back on forehead; dark areas at angles of mouth confluent with color of upper parts; tip of tail black; remainder of upper parts near (n) Bay or Reddish Black, tone 4 of Oberthiir and Dau- thenay, pi. 344; chin whitish; remainder of underparts Warm BuflF, deep orange in juvenile, type specimen, according to Gray (1864, pi, 8); color of under- parts extending distally on posterior sides of forelegs to wrists but not reach- ing foot soles and on hind legs to or slightly below knees. Least width of color of underparts equal to 15 per cent of greatest width of color of upper parts. Black tip of tail equal to 27 per cent of lengtli of tail-vertebrae. In color, no. 34677 is, to me as it was to Allen ( 1916:101), indistinguishable from the darkest specimens (nos. 178970 and 10112) of M. f. panamensis. Therefore, M. f. aureoventris is one of the two darkest subspecies of weasels. Skull and teeth. — Male: See measurements and plates 27-29; weight, 4.3 grams; basilar length, 45.8; z>'gomatic breadth approximately equal to dis- tance between condylar foramen and Ml and to distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more than length of upper pre- molars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less (at least in young specimen) than length of tympanic bulla; least widtli of palate seldom if ever greater than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of three (including 13) upper incisors; height of tympanic bulla not greater than dis- tance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than orbitonasal length; anterior margin of masseteric fossa below anterior half of m2. Skulls of males of M. /. aureoventris, and Mustela frenata macrura from the MUSTELA FRENATA 389 vicinity of Quito so closely resemble one another as not to be distinguished with the material now available, although the teeth of aureoventris are larger. Comparisons of the skulls of males with those of M. f. meridana and affinis, which are readily distinguishable from those of aureoventris, have been made in the accounts of those subspecies. Female: Skull of adult unknown. Remarks. — This subspecies of the Tropical Life-zone, or at least the Subtropical Life-zone, of Ecuador, in certain cranial characters resembles Mustela frenata macrura of the Temperate Life-zone. The two differ markedly in color. Nevertheless, a large number of the specimens collected in Ecuador are intermediate in color as well as in zonal distribution. The type specimen is young or a juvenile. The measurements of no. 34677 from Gualea indicate an animal similar in size to M. /. affinis. Gray (1864:55) states that the type specimen meas- ures "Length of body and head 6 inches, of tail 4/2 inches." The plate (pi. 8) accompanying Gray's original description (loc. cit.) is marked one-half natural size and represents the animal as hav- ing a head and body length of eight and one-half inches. One year later Gray (1865:115) gives the measurements of this species as "Length of body and head 12, tail 8 inches." Since he had at this time another specimen, larger than the type specimen (which specimen later, probably, became the type of Mustela affinis Gray), the larger measurements probably were taken from it. Geographically, and as regards cranial characters, Mustela frenata aureoventris is most closely related to M. f. affinis and to the north- em section of M. f. macrura, but in color to M. /. panamensis. M. f. aureoventris and M. /. panamensis are the two darkest-colored sub- species and each occurs in a region of extremely heavy rainfall. There is a skin only, no. 32620, Amer. Mus. Nat. Hist., from Munchi- que, obtained on June 1, 1911, which is appreciably darker than specimens of M. f. affinis in corresponding pelage and is intermedi- ate between M. /. affinis and M. /. aureoventris in color as it is geo- graphically. The specimen measures 495, 202, 52. The name Mustela aureoventris Gray has been regarded by most authors as preoccupied by Mustela auriventer Hodgson (1841:909). However, the writer is not of this opinion and agrees with Thomas (1920:224) that "The name aureoventris is not invalidated by the auriventer of Hodgson, as, apart from 'one-letterist' differences, its first half comes from the adjective aureus, while Hodgson's name is based on the substantive aurum, so that not only the spellings but the derivations are different." The spelling of Gray's name should be aureoventris for this is the spelling in the original de- 390 University of Kansas Publs., Mus. Nat. Hist. scription which in pagination precedes the colored plate of the animal that is labeled Mustela aureiventris. Putorius brasiliensis var. aequatorialis Coues (1877:142) is the only name known to the writer that has been proposed as a substitute for Mustela aureoven- tris Gray. Thomas (1920:224) treats Mustela macrura Taczanowski as a synonym of Mustela aureoventris Gray. Allen (1916:101) also treats the two names as applying to the same kind of weasel but regards aureoventris as preoccupied and therefore uses the name macrura. Taczanowski's original description (1874:311) and plate of Mustela macrura indicate an animal that is lighter colored than M. /. affinis. Gray's original description (1864:55) and plate of aureoventris indicate an animal that is darker colored than M. f. affinis. Indeed Gray (1865:115) in speaking of the type of aure- oventris as compared with an adult from New Granada [=z Colom- bia] that probably later became the type specimen of Mustela affinis, states: "The young from Quito is much darker than the adult; . . ." Comparison of the plates accompanying the orig- inal descriptions of aureoventris and macrura well illustrate the dif- ference stated in the written descriptions. My examination of the type specimens of M. macrura and M. /. aureoventris shows them to have been fairly accurately portrayed in the plates ac- companying the original descriptions. Accordingly the two names are used for the two kinds of animals which appear, however, to be only subspecifically distinct. Comparison of Gray's plate (1864, pi. 8) with the available specimens from South America indicates that the name aureoventris is based on an individual that is lighter colored than no. 34677 Amer. Mus. Nat. Hist., from Gualea, Ecuador, but on one which resembles no. 34677 more than it does the lighter-colored speci- mens from the Temperate Zone of Ecuador and northern Peru. Because Quito, Ecuador, is in the Temperate Life-zone and because the available specimens from this zone in Ecuador and northern Peru are distinctly lighter colored than Gray's plate representing the type of aureoventris shows this specimen to be, it is judged to have come from an altitude lower than that of Quito (9350 feet, according to Chapman, 1926:717); probably it came from the Sub- tropical Life-zone of Ecuador. Indeed Gray (1864:55) did not say that the specimen was collected or obtained at Quito but that it was ". . . received from Quito. . . ." Chapman (1926:717) has pointed out that Quito, since 1846 has been the distributing point for bird skins which specimens ". . . come from the vicin- MUSTELA FRENATA 391 ity of the city, from the 'Napo' region on the Amazonian slopes of the Andes, and from Nanegal, Gualea, and other locaHties on the Pacific side rarely below the Subtropical Zone." It is also pointed out that only some of the specimens are labeled with their approxi- mate place of capture and that even then these localities cannot be accepted as definite; they indicate mainly whether the specimen is from the eastern or western side of the Andes. The above mentioned considerations and information gained by study of the specimens cause me to think that the type is an inter- grade tending toward the lighter-colored Mustela f. macrura of the Temperate Zone although suflBciently dark to be referred to the dark subspecies represented by no. 34677 Amer. Mus. Nat. Hist., from Gualea, Ecuador. The skull of no. 34677 shows no infestation of the frontal sinuses by parasites. Specimens examined. — Total number, 3, as follows: Ecuador: Gualea, 1, Amer. Mus. Nat. Hist. Colombia: 8325 ft., Munchique, 1, Amer. Mus. Nat. Hist. In the British Museum of Natural History, the type, ( 1 ) . Mustela frenata helleri Hall Long-tailed Weasel Plates 27, 28 and 29 Mustela frenata helleri Hall, Proc. Biol. Soc. Washington, 48:143, August 22, 1935; Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939. Type. — Male, adult, skull and skin; no. 24133, Field Mus. Nat. Hist.; 3000 feet. Hacienda San Antonio, Rio Chinchao, Peru; August 22, 1922. Obtained by Edmund Heller. Original no. 6589. The skull (plates 27-29) is complete and unbroken. The teeth all are present, entire and but slightly worn. The skin is well made, unfaded, and in good condition. Range. — Three thousand feet (type locahty) to 6700 feet (Ambo), Tropical and Subtropical Ufe-zones of eastern Peru. See figure 29 on page 221. Characters for ready recognition. — Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, presence of p2 and two roots rather than one root on P2; from Mustela frenata macrura by darker color (Carbon Brown, tone 3, pi. 342 rather than Chocolate, tone 3, pi. 343, Oberthiir and Dauthenay) of upper parts. Description. — Size. — Male: Measurements of the type specimen and topo- type, no. 24132, are, respectively, as follows: Total length, 382, 418; length of tail, 152, 164; length of hind foot, 52, 48. Tail 66 and 65 per cent as long as head and body. Hind foot more than basal length. Female: Measurements of two referred females, no. 24134 from Ambo and no. 24136 from Huanuco, are, respectively, as follows: Total length. 392 University of Kansas Publs., Mus. Nat. Hist. 328 and 303; length of tail, 118 and 103; length of hind foot, 39 and 38.5. Tail 56 and 51 per cent as long as head and body. Hind foot shorter than basal length. The average differences in external measurements of the two sexes are: Total length, 85; length of tail, 49; length of hind foot, 11, Externals. — Longest facial vibrissae black and extending beyond ear; carpal vibrissae same color as upper parts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figiue 20. Color. — Rarely a few white hairs anterior to each ear; posterior fifth of each upper Lip white; top of head, posteriorly to sUghtly behind ears, black, grading into color of upper parts of body; dark spots at angles of mouth absent; tip of tail black; remainder of upper parts near (n) Argus Brown and Carbon Brown, tone 3 (pi. 342, Oberthur and Dauthenay); chin whitish; remainder of underparts Warm Buff; color of underparts extends distally on posterior sides of forelegs to wrists but not reaching foot-soles and on hind legs to slightly below knees. Least width of color of underparts 24 per cent of greatest width of color of upper parts in each of two males and 19 to 30 per cent in three females. Black tip of tail longer than hind foot and averaging 40 ( 39-42 ) per cent of length of tail-vertebrae. Skull and teeth. — Male (based on type specimen and adult no. 24132): See measurements and plates 27-29. As described in Mustela frenata macrura except that: Weight, 4.5 (4.2 and 4.8); basilar length, 44.6 (44.0-45.3); zygomatic breadth more than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bidla; breadth of rostrum more or less than (approximately equal to) length of tympanic bulla; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of rostrum; anterior margin of masseteric fossa posterior to m2 by length of that tooth. Female (based on nos. 24134 to 24136): See measurements. As described in Mustela frenata macrura except that: Weight, 1.7 (1.5-1.9) grams; basilar length, 36.5 (35.3-38.1); zygomatic breadth less than distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more or less than (approximately equal to) outside length of P4; length of tympanic bulla less than length of rostnmi. The skull of the female averages 62 per cent fighter than that of the male. The skull of the male is generally large and heavy as are the teeth. Comparison with macrura is made in the account of that subspecies. From males of affinis those of helleri differ in: skull shorter; breadth of rostrum and interorbital breadth actually and relatively greater. Remarks. — The five specimens examined of this subspecies were taken by Edmund Heller for the Field Museum of Natural History in 1922 and 1923. It is to honor his contributions to mammalogy that the subspecies is named helleri. No. 24135 is the specimen carried as a pet for some time by Mr. and Mrs. Heller and of which Mrs. Heller (1924:481) has given an account. MUSTELA FRENATA 393 This subspecies is insuflBciently known, especially as to geo- graphic range; probably it occupies a considerable range in the Tropical Life-zone along the eastern base of the Andes. The three females, two from Ambo and one from Huanuco, come from a much higher altitude than do the two males and the climate is said to be arid at Ambo and Huanuco. The skulls of the females are 62 per cent lighter and correspondingly smaller in measure- ments, than those of males. This difference is more than that found in any other South American weasel and it may be that the females are of a subspecies other than helleri. The type specimen has a broad skull with major proportions strik- ingly like those of Mustela stolzmanni. Possibly the similar chmatic conditions under which the two live have left their impress in simi- lar fashion in this part of each of the two species. The teeth, tympanic bullae, and certain other parts of the skull are, however, so dijfferently proportioned as to show that the skulls represent two species. The referred male has a much longer skull than the type specimen and die relative proportions of breadth and depth of the two skulls differ widely. Judging from large series of weasels ex- amined from localities outside the range of M. f. helleri, the two skulls probably represent almost the maximum of individual varia- tion occurring in one subspecies. The dark color is as might be expected since helleri inhabits the humid Tropical Zone. None of the five skulls shows signs of having had the frontal sinuses infested by parasites. Specimens examined. — Total number, 5, all in the Field Museum of Natural History. Peru: 3500 ft., Hacienda Buena Vista, Rio Chinchao, 1; 3000 ft.. Hacienda San Antonio, Rio Chinchao, 1; Huanuco, 1; Ambo, 2. Mustela frenata agilis Tschudi Long-tailed Weasel Plates 27, 28, 29, 39 and 40 Mustela agilis Tschudi, Fauna Peruana, p. 110, 1844; Gray, Proc. Zool. Soc. London, 1865:113, 1865; Taczanowski, Proc. Zobl. Soc. London, 1874:311, 1874; Taczanowski, Proc. Zool. Soc. London, 1881:648, 1881; Allen, Bull. Amer. Mus. Nat. Hist., 35:104; April 28, 1916; Thomas, Proc. U. S. Nat. Mus., 58-224, 1920. Mustela macrura, Allen, Bull. Amer. Mus. Nat. Hist., 35:103, April 28, 1916. Mustela frenata agilis. Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939. Type. — No type specimen, or type locality more restricted than cold, barren highlands of the Cordillera [referring to Peru] designated. Range. — High, barren Cordillera of Peru ( see Tschudi, orig. dcscr. ) ; as here 394 University of Kansas Publs., Mus. Nat. Hist. restricted, Temperate Life-zone and higher in western Andes and intermoun- tain valleys of Peru. See figure 29 on page 221. Characters for ready recognition. — Differs from Mustela frenata macrura by lighter color (Chocolate, tone 2 rather than 3, pi. 343, Oberthiir and Dauthenay) of upper parts; length of upper tooth-rows, in females, less than 13; inside length of P4 more than 4.6; from A/. /. aureoventris by smaller teeth (maxi- mum size just given for agilis); from M. f. boliviensis by hghter color, upper parts being Chocolate, tone 2, pi. 343, rather than tone 4 or darker of Carbon Brown, pi. 342 (Oberthiir and Dauthenay). Description. — Size. — Male: The stuffed skin of an adult, from Lima, meas- ures: Total length, 460; length of tail, 125; length of hind foot, 45.7. A skin alone from Huarochiri has a body, as now stuffed, 277 mm. long. The tail is missing and the bones of the hind feet have been removed. Female: The mounted specimen, no. 565, Mus. Polonais d'Hist. Nat., yields measurements, taken by me, as follows: Total length, 250; length of tail, 75; length of hind foot, 32.5. The female, no. 21147, from Macate, measures, 300, 102, 34. Externals. — Longest facial vibrissae, either dark- or hght-colored and extend- ing beyond ear; carpal vibrissae either dark- or light-colored and extending to apical pad of fifth digit; hairiness of foot soles as shown in figure 20. Color. — Tschudi's description of the color is, in substance, as follows: Head, back and tail reddish gray; base of hair gray, followed by broader grayish- yellow ring and then reddish-brown tip; nose simply dark brown or upper fips edged with white; throat, breast, belly and higher parts of inner sides of ex- tremities whitish gray, at times wholly gray, bases of hairs always gray; feet darker than body, almost chestnut brown; tail darker on tip than at base; ears externally dark brown, internally whitish. No. 565 possibly somewhat faded from exposure to hght, has all the upper parts near (14' /) Ochraceous-Tawny or Cinnamon, and tone 4 of Oberthiir and Dauthenay, plate 323; posterior half of each upper lip white; no other white facial markings present; dark spot at each angle of mouth ( one spot conflu- ent with color of upper parts); tip of tail probably black (tip missing); under- parts white, belly probably originally with slight tinge of yellow or aUied color; color of underparts extending distally on forelegs to feet and onto upper sides of toes and on hind legs to just above heels. Least width of color of xinder- parts equal to about one-fourth of greatest width of color of upper parts. No. 21147, subadult, from Macate, has a white band confluent with the underparts extending anterodorsally anterior to each ear and the posterior third of each upper Up white. Top of head near (n) Mars Brown, and Carbon Brown, tone 3 (pi. 342, Ober. and Dautli. ); tip of tail black; remainder of upper parts near (16" /) Tawny-Olive, and Chocolate (tone 2, of pi. 343 of Ober. and Dauth.) or Raw Umber (tone 3 of pi. 301 of Ober. and Dauth.); anterior half of underparts, including posterior sides of forelegs and antipalmar faces of forefeet, white; remainder of underparts tinged with Warm Buff and extended on posterior legs almost to ankles. No. 8. 1. 10. 1., male adult, from Lima, is also light colored, and as described in no. 21147, except that left side of head has a white spot rather than bar; posterior eighth of each upper hp white; white frontonasal spot present, 11 x 11 mm.; antipalmar faces of forefeet spotted with brown color of upper parts; MUSTELA FRENATA 395 color of underparts extending distally on hind legs along medial side of foot to point halfway between heel and tip of inner toe. No. 13257 from Huarochiri in color and color pattern closely resembles no. 21147. It difiFers from no. 21147 in slightly Hghter color of upper parts, en- tirely white underparts, less extension of color of underparts onto forefeet, few white hairs instead of white band in front of each ear; color of underparts more restricted. In each of the four specimens, the least width of the underparts, expressed as a percentage of the upper parts, is as follows: no. 13257, 11 per cent; no. 21147, 29 per cent; no. 565, 31 per cent; no. 8.1.10. 1., nineteen per cent. Skull and teeth. — Male (based on no. 8.1.10.1.): See measurements and plates 27-29. As described in Mustela frenata macrura except that: Weight 4.1 grams; basilar length, 42.5; zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; tympanic bullae shorter than rostrum. Female (based on no. 21147): See measurements and plates 39 and 40. As described in Mustela frenata macrura except that: Weight (no. 21147, sub- adult), 1.5 grams; basilar length, 35.2; least width of palate less than outside length of P4; tympanic bulla as far posterior to foramen ovale as combined width of five upper incisors; no. 565 answers to the same description but difiFers from no. 21147 in greater basilar length and larger tympanic bullae which are slightly more projected, at their anterior margins, from the braincase. To judge from the skull of the female from Macate and the skull of the male from Lima, the skull and teeth of agilis are smaller than in any other South American subspecies of Mustela frenata, except M. /. boliviensis. Remarks. — Tschudi almost certainly used the name Mustela agilis in a composite sense. His statement (see quoted matter below) about the marked variation in color of this species, as represented by the skins carried by the Indian women as purses, indicates that the forms here designated as Mustela macrura, M. helleri and possi- bly others additional to the one here called agilis were included by him under the name Mustela agilis. Taczanowski took account of Mustela agilis when he described other species from Peru. Allen (1916:104) and Thomas (1920:224) were not convinced that Mustela agilis and Mustela macrura were distinct species or sub- species. Search on August 28, 1937, in the Musee d'Histoire Naturelle, at Neuchatel, Switzerland, by Mr. Theodore Delachaux, assistant I there, and the writer, revealed no trace of weasels from Tschudi's collection, although some other specimens of mammals that he j figured in the "Fauna Peruana" are preserved in that Museum. Not only were the collections of specimens examined but the new cata- logue and old catalogue of mammals were vainly searched for men- 396 University of Kansas Publs., Mus. Nat. Hist. tion of weasels deposited by Tschudi. Later, at the British Mu- seum of Natural History, on page 105 of a personal notebook, of the late Mr. Oldfield Thomas, record was found of his fruitless search for the same specimens of Mustela in May, 1902, at Neuchatel. Although Tschudi certainly used the name Mustela agilis in a composite sense, as subspecies are at present understood, his de- scription most nearly apphes to the light-colored animals from western Peru — the lightest colored of any South American weasels seen. They are of approximately the same color as North American subspecies inhabiting semiarid regions, for example Mustela frenata longicauda of the Great Plains. Another, but in my opinion less weighty, justification for applying Tschudi's name agilis to these light-colored weasels of western Peru is that by one line of reasoning, Taczanowski in naming macrura ( jelskii is a synonym of it ) from farther eastward in Peru, and that Hall in naming helleri from still farther eastward, and boliviensis to the southeastward, geographically restricted the apphcation of the name agilis. Hall's action did this because he recognized geo- graphic variation and employed the subspecies concept. Taczan- owski, however, proposed his name macrura for a kind of animal which he indicated was specifically (as opposed to subspecifically) distinct from agilis and his account (1881:649) of jelskii indicates that he thought Mustela agilis Tschudi might occur in the same place as the animals which he named as new kinds. Thus, we can not credit Taczanowski with intent to restrict the name agilis geo- graphically, even though later authors may choose to rule that his naming of macrura in efi^ect did so restrict the application of Mus- tela agilis Tschudi. The equivalents in millimeters given by Allen (1916:104) for Tschudi's measurements of 9 to 10 inches entire length, and tails of 4 inches to 4 inches and 4 lines, apparently are based on the London scale in use today. If Tschudi employed the Rhine scale also of eight Hues to the inch, but one which has the foot longer by an amount of 20 millimeters, or the Leipzig scale in which the foot is 22 millimeters shorter than the London foot, the measurements recorded by Tschudi differ in one direction or the other from those computed by Allen. However, knowledge of which scale Tschudi employed would not help much, if any, in more precise application of the name agilis because he does not indicate whether his measure- ments are of male or female animals; animals of the two sexes of the same subspecies differ more in external measurements than animals of the same sex of different subspecies of Peruvian weasels. MUSTELA FRENATA 397 Specimen no. 565, in the Polish Museum of Natural History, with- out definite locality, is provisionally referred to this subspecies. The specimen is intermediate in several respects between the female from Macate and the one of macrura from Cutervo. Tschudi (1844:111-112) has given the following account: "Lebensweise und geographische Verbreitung. Das peruanische Wiesel lebt auf den kalten, oden Hochebenen der Cordillera an sonnigen Steinhaufen und Felsen gewohnlich in Gesellschaft von 8-12 Stiicken. Diese Thierchen sind so ausserordentlich behende und scheu, dass bei dem leisesten Gerausche die ganze Schaar mit Blitzesschnelle verschwindet. Es ist uns auch nie gelungen, eines derselben zu eriegen. Die Indianer aber verstehen es, dieselben lebendig einzufangen und zu zahmen. Ein sehr zahmes sahen wir bei einer uns be- freundeten Dame in Tarma; gegen alle Fremden biss es mit Wuth und liess sich nicht anfassen, wiihrend es sich von seiner Herrin Alles gefallen liess; sie ofFnete ihm den Mund und steckte ihm den Finger hinein, ohne dass es eine bose Miene dazu machte, wiihrend es bei der geringsten Bewegung, die wir machten, es zu ergreifen, grimmig auf uns lossprang. Wenn es eingeschiichtert wurde, versteckte es sich in den Busen seiner Gebieterin und kroch ihr bald nachher zum Aermel heraus. An den Wanden und Meublen kletterte es mit grosser Behendigkeit und schliipfte durch so kleine Ritzen und Locher, dass wir fast an der Moglichkeit dieses Hindurchdringens gezweifelt haben wiirden, wenn wir es nicht selbst mit angesehen batten. Wenn es unartig war, wurde es mit einer Schnur an seinem kleinen Halsbande festgebunden; dadurch vermehrte sich sein Zom, so dass es zuweilen gegen die Dame auffuhr. Mehrmals verschwand es wahrend 8-10 Tagen und kam dann plotzlich wieder zum Vorschein. Seine Nahrung bestand in Gemiise und Fleisch, besonders aber liebte es Zuckerbrod in Milch aufgeweicht; einmal machte es sich an einen Kanarienvogel, den es auch todtete. Es erhielt seine Strafe und verschwand dann fiir immer. Die Indianer soUen dieses Wiesel zum Fange der Viscacha abrichten (davon weiter unten). Sie nennen es Comadreja, auch Ardilla. ( [footnote] Ardilla ist spanisch und heisst Eichhomchen. Mit diesem Namen werden sehr verschiedene Thiere bezeichnet; ausser dem Sc. variabilis und der Gahctis agilis auch noch mehrere Nager vmd einige Didelphysarten. ) Die Indianerinnen verfertigen sich aus dem kleinen Felle Geldbeutel. Des Sonntags triflFt man unter den vielen tausend Punaindianer- innen die nach den grossen Dorfem der Sierra kommen, um ihre Einkiiufe zu machen, kaum ein halbes Dutzend, die nicht solche Borsen mit sich fiihrten, imd dann kann man auch die verschiedensten Farbennuancen, die bei dieser Species vorkommen, beobachten." None of the tliree skulls referred to this subspecies shows infes- tation of the frontal sinuses by parasites. Specimens examined. — Total number, 4. Peru: Macate, 1 (Field Mus. Nat. Hist.); Huarochiri, 1 (Mus. Comp. Zool.); Lima, 1 ( British Mus. Nat. Hist. ) ; no locality more definite than Peru, 1 ( Mus. Polonais d'Hist. Nat.). 398 University of Kansas Publs., Mus. Nat. Hist, Mustela frenata macrura Taczanowsld Long-tailed Weasel Plates 1, 27, 28, 29, 30, 37, 38, 39 and 40 Mustela macrura Taczanowski, Proc. Zobl. Soc. London, 1874:311, pi. 48, May 19, 1874; ibid., 1881:647, May 17, 1881; ibid., 835, November 15, 1881; Lonnberg, Arkiv for Zool., 8 (no. 1):21, 1913 (?); Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914; Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Lonnberg, Arkiv for Zool., 14 (no. 4):11, 1921. Putorius (Gale) braziliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877. Mustela jelskii Taczanowski, Proc. Zobl. Soc. London, 1881:647, May 17, 1881. Mustela affinis, Lonnberg, Arkiv for Zool., 8 (no. 1):21, July 12, 1913. Mustela aureoventris, Thomas, Proc. U. S. Nat. Mus., 58:224, 1920. Mustela frenata macrura. Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939. Type. — Male, adult, mounted skin, wi^h skull separate; no. 561, Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Lake Junin, central Peru; 1873; obtained by M. Jelski. The skull (plates 27-29, 30), mounted wdth the skin but removed by me for study, lacks the right jugal, the basisphenoid, the basioccipital and parts of each exoccipital bearing the exoccipital condyles. The right tympanic bulla, although detached from the skull, is preserved separately. The teeth all are present and entire. The skin is fairly-well mounted, in a good state of preservation, and shows no fading due to exposure to light. Range. — Altitudinally, 3200 (Guainche) to at least 12000 feet (Pichincha); Upper Subtropical and Temperate life-zones of central Peru and Ecuador north from tlie states of Apurimac and Cuzco, Peru, to San Antonio, northern Ecuador. See figure 29 on page 221. Characters for ready recognition. — DiflFers from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts; presence of p2 and two roots rather than one root on P2; from Mustela frenata helleri, M. f. affinis and M. f. aureoventris by Hghter color of upper parts which are Chocolate tone 3, pi. 343, Oberthiir and Dauthenay, whereas, with reference to the same color standard, the colors are: in helleri, Carbon Brown, tone 3, pi. 342; in affinis. Reddish Black, tone 2, pi. 344; in aureoven- tris. Reddish Black, tone 4, pi. 344; from M. f. agilis by darker color (Choco- late, tone 3 rather than 2, pi. 343, Obertliiir and Dauthenay) of upper parts, length of upper tooth-rows, in females, more than 13, inside length of P4 more than 4.6; from M. f. boliviensis by lighter color of upper parts which are as above rather than tone 4 of Carbon Brown, pi. 342 of Oberthiir and Dauthenay, and larger size (in males, hind foot more than 45 and ml more than 5.6). Description. — Size. — Male (measurements as recorded by Taczanowski in the original description, for two specimens, type and topotype, with correction of the length of tail of his "female" [=male]): Total length, 420, 415; length of tail, 150, 145; length of hind foot, 51, 51. An adult from Yano Mayo, Rio Tarma, was measured by Hendee as 394, 134. Hind foot re- laxed measures, 47. Tail 55 per cent as long as head and body. Length of MUSTELA FRENATA 399 hind foot more than basal length. Female (based on measurements given by Taczanowski (1881:647) of no. 564): Total length, 323; length of tail, 120; length of hind foot, 37. Tail 59 per cent as long as head and body. Length of hind foot approximately equal to basal length. Differences in external measurements of the two sexes are: Total length, 87; length of tail, 23; lengtli of hind foot, 13. Externals. — Longest facial vibrissae extending beyond ear; carpal vibrissae color of either upper parts or underparts; hairiness of foot-soles as shown in figure 20. Color. — (Based on specimens from Cutervo and south thereof). Rarely few white hairs between eyes and in front of ears; top of head posteriorly to shghtly behind eyes, near (n) Chesinut-Brown (Ridgway) and Carbon Brown, tone 2 or darker (pi. 342, Oberthiir and Dauthenay); posterior half of upper lip rarely white; dark spots at angles of mouth absent; tip of tail black; remainder of upper parts near (/) Russet (Ridgway) and Chocolate, tone 3 (pi. 343, Ober. and Dauth. ); underparts white or whitish on medial sides of forelegs, otherwise cream color with tinge of Ochraceous-Buff; color of underparts extended distally on posterior sides of forelegs to just below elbow (in type specimen) or onto forefeet (in specimen from Yano Mayo) and on medial sides of hind legs to points between knees and ankles. Least width of color of underparts averages (in six skins) 17 (14-21) per cent of greatest width of color of upper parts. Black tip of tail longer than hind toot and averaging 36 (32-49) per cent of length of tail-vertebrae. Skull and teeth. — Male (based on type specimen and no. 562): See measurements and plates 27-30; weight, not known; basilar length, 43.2 (40.8 and 45.5 ) ; zygomatic breadth more or less than distance between condylar fora- men and Ml and more than that between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less than postpalatal length; postorbital breadth more than length of upper premolars and greater than width of basioccipital, measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more than distance between fora- men opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic buUa; least width of palate more than inside length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 (including 13) upper incisors; height of tympanic bulla not more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below or behind m2. Female (based on no. 564, from Cutervo, Peru, type specimen of Mustela jelskii Taczanowski ) : See measurements and plates 37-40; weight, not known; basilar length, 38 ±; zygomatic breadth less than distance between condylar foramen and Ml and not greater than that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than alveolar length of upper premolars and (probably) more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than inside length of P4; tympanic bulla as far posterior to foramen ovale as width of at least 5/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; 400 University of Kansas Publs., Mus. Nat. Hist. length of tympanic bulla more than length of lower molar and premolar tooth- row and longer or shorter than rostnmi. As compared with that of helleri, the skull of the male of macrura from Junin southward has a lesser mastoid breadth, notably smaller teeth, and a flatter skull which averages lighter throughout. The skulls of females available indicate that the skull and teeth are larger than in agilis. Remarks. — Seven years after Taczanowski named this subspecies, he applied the name jelskii to a female taken farther north than the original examples of macrura. As indicated in synonymy, various other names have been applied to animals included by the present author in this subspecies, Mustela frenata macrura intergrades with M. f. affinis as shown by practically all the referred specimens from north of Junin. As one proceeds northward the color of the weasels becomes progres- sively darker and the teeth become larger until the conditions found in affinis are met with near the northern border of Ecuador. From the material available it appears that the light-colored upper parts found in macrura characterize weasels of, at least, the Temperate Zone, from Marcapata, Peru, to near Quito, Ecuador. West of the range of macrura there exists the still lighter-colored subspecies, M. /. agilis. Immediately adjacent on the north, east, and south, darker-colored weasels occur. So far as color is concerned, the geographic range of the subspecies M. f. macrura is not difficult to define. However, the small size of the teeth characterizes only that part of this light-colored subspecies from Junin southward including the subspecies boliviensis at the southern extremity of the range of the species. From Cutervo northward the light-colored weasels of the Temperate Zone have teeth similar in size to those of the darker, more northern affinis. To designate the slightly larger- toothed, light-colored animals from Ecuador as a subspecies dis- tinct from affinis and macrura is one solution but at present it seems best to refer all of these light-colored animals to macrura. The type specimen and topotype no. 562 differ more in the amount of inflation of the tympanic bullae than adult males of comparable ages from a given locality usually do. In other respects, the dif- ferences between the two skulls are not greater than those ordi- narily found in specimens from the same locality. No. 562 has the tympanic bullae greatly, relative to tlie other South American wea- sels, inflated posteriorly. Otherwise, the bullae agree with those of tlie type specimen. Specimens from southwestern Ecuador, average large, and in- MUSTELA FRENATA 401 elude the largest specimens of the species Mustela frenata seen from South America. A subadult male, no. 61406, in the American Mu- seum of Natural History, is the largest. Its external measurements are 482, 191, 56. The basilar length of the skull is 48.2 and the zygo- matic breadth is 30.3. Although not so large as this specimen, the corresponding measurements of specimens from Alamor, El Chiral, and even from as far away as Sigsig also are distinctly large. The skull of the female from Ollantaytambo and that of the male from Marcapata have teeth equally as small as do the specimens from Lake Junin. The skin alone, no. 194328, from Ollantaytambo has the color of the underparts extended over the entire upper sides of the forefeet. The male from Marcapata has less of this color on the forefeet and is in this respect intermediate between the specimens from Lake Junin and the one from Ollantaytambo. In size of teeth the female, type specimen of M. ielskii, from Cutervo, shows an approach to the larger-toothed weasels of tlie northern part of the range of macrura. The specimens in the Riksmuseum from the vicinity of Quito, Ecuador, have been rather fully described by Lonnberg (1921:11- 17) and need little comment here, except to say that they show, as he suggested, that the weasel of the Temperate Zone of Ecuador is an intermediate link between M. /. macrura and M. f. affinis. The adult female and juvenal male labeled as from Ambato have little left of the skulls except some of the teeth and the assignment of the specimens to the subspecies macrura is made mainly on geo- graphic grounds. These two specimens probably are part of the shipment of birds and mammals of which Chapman (1926:703) speaks as follows: "A small collection of native-made skins pur- chased by the American Museum from a commission merchant in New York City as from 'Ambato' proved to be from the eastern slope of the Andes." Another skin in the same Museum, labeled by a native collector as from "Baeza arriba" [= above Baeza] is so dark colored and has the color of the underparts so much restricted, as to suggest that it belongs to the race aureoventris. Possibly, there- fore, it was taken not at Baeza, Ecuador, which I find to the east- ward of Quito at 77° 55' W and 0° 25' S, but at some place of the same name on the Pacific Slope, unless the locality has been altogether vvTongly recorded on the label. If the specimen was taken near the Baeza above referred to, then it gives evidence of an unnamed race of Mustela on the eastern slope of the Andes, characterized by its dark color. Unfortunately the specimen is young and its skull 402 University of Kansas Publs., Mus. Nat, Hist. therefore ofiFers insuflBcient basis for the judging of its subspecific relationships. Other specimens, in the British Museum of Natural History, re- corded as taken "near Quito" and here tentatively listed under macrura, mostly, include specimens so dark colored as to lead me to think they came from country, lower than Quito, adjacent to the range of aureoventris. Nematodes taken from the right frontal sinus of no. 562 from Junin proved to belong to the superfamily Oxyuriodea according to Professor W. B. Herms and Mr. O. L. Williams, who have inde- pendently identified them. Because these worms had been dried fifty-five years in the mounted specimen and were later boiled in cleaning the skull, a more accurate determination was impossible and whether or not they pertain to the same species found in North American weasels cannot be said. Of 18 adult skulls examined for this type of infestation, 13 were found affected as judged by the evident malformation of the frontal region. Specimens examined. — Total number, 74, arranged by localities from north to south and unless otherwise indicated in the American Museum of Natural History. Ecuador: Ibarra, 6600 ft., and 7500 ft., 2'; San Antonio, 8000-8500 ft., 5 mi. N Quito, 4 (2', 2'); Nono, 10000 ft, 1; Mindo, 1=; Zambiza, 8000-8100 ft, NE Quito, 4 (2\ 2'); Carapungo, 8500 ft., NE Quito, 1^ Panecillo, 10000 ft, near Quito, 2'; Guapulo, 8800 ft., 3 mi. E of Quito, 1^ Pichincha, 10500 ft., and 12000 ft., 2 (l^ 1'); San Ignacio, 11500 ft., Pichincha, 1; Santa Rosa, 9600 ft., Rio Pita, 2; near Santa Rosa, 9000 ft., 1; Rio San Rafel, 9000 ft., 1; N side Quito, 9000 ft, 1=; Quito, 1^ near Quito, 5'; Nara Papallacta, 11000 ft., 1=; below Papallacta, 9000 ft., 1=; Chillo Valley, 1^ "Hacienda Hda", 10000 ft., Pintag, Valencia, 1; Baeza arriba, 1; Ambato, 2; San Francisco, 8000 ft., E of Ambato, 1; Chunchi, Pagma Forest, 6400 ft., 1'; Canar, 2600 M., 1'; Malletura, 7600 ft., 1; Contrayerbas, 11000 ft., 1; Sisig, 8500 ft., 3^ El Chiral, 1; Almor, 1; Guainche, 3200 ft., 1; no locaHty more definite than Ecuador, 4'; "Received from Quito", 1*; Quisaya, 6000 ft. (locahty not found), 1'; La CaroUna (lo- cality not found), 1'. Peru: La Lejia, 1; Huancabamba, 4 (2'); Cutervo, 9000 ft., 1'; Conde- chacha, 7000 ft., Rio Utcubamba, 1'; San Pedro, 8600-9400 ft., S of Chacha- poyas, 1; Celendin, 1'; Junin, 2"; Yana Mayo, Rio Tarma, 1'; Ollantaytambo, 9000 ft., 3 (1\ 2'); Ocabamba, 1'; Anta Cuzco, 3400 and 3500 M., 2*; Mar- capata, 1'. Mustela frenata boliviensis Hall Long-tailed Weasel Plates 28, 29 and 30 Mustela frenata boliviensis Hall, Proc. Biol. Soc. Washington, 51:67, March 18, 1938. Mustela frenata macrura, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939 (part). 1. British Museum of Natural History 5. Museum of Comparative Zoology 2. Naturhistorika Ricksmuseum, Sweden 6. Mus. Polonais d'Hist. Nat., Warsaw 3. Univ. Zool. Mus., Copenhagen 7. United States National Museum 4. Berlin Zoological Museum 8. Acad. Nat. Sciences of Philadelphia MUSTELA FRENATA 403 Type — Male, adult, skull and skin; no. 72587, Amer. Mus, Nat. Hist.; Nequejahuira, 8000 feet, Bolivia; May 19, 1926; obtained by G. H. H. Tate; original no. 4135 (see plates 28-30). Range. — As now known 8000 to 9500 feet in the Andes from Limbani, Peru, south to Nequejahuira, Bolivia; upper Subtropical and Temperate life-zones. See figure 29 on page 221. Characters for ready recognition. — Di£Fers from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts; presence of p2 and two roots rather than one root on P2; from Mustela frenata macrura by darker color of upper parts (tone 4 or darker of Carbon Brown, pi. 342 rather than tone 3 of Chocolate, pi. 343, Oberthiir and Dauthe- nay) and lesser size (in males hind foot less than 45 and ml less than 5.6); from Mustela frenata agilis by darker color of upper parts (as given above rather than tone 2 of Chocolate, pi. 343, of Oberthiir and Dauthenay). Description. — Size. — Male: The type and two young specimens from Limbani, Peru, measure respectively, as follows: Total length, 383, 368, 304; length of tail, 140, 132, 115; length of hind foot, 43, 44, 41. Tail 55 per cent as long as head and body. Length of hind foot approximately equal to basal length. Female : Unknown. Externals. — As described in Mustela frenata macrura. Color. — Top of head blackish posteriorly to behind ears; upper lips same color as upper parts of head; dark area at angle of mouth not separated from upper parts as a distinct spot; tip of tail black; remainder of upper parts near (n) Mars Brown of Ridgway and tone 4 or darker of Carbon Brown (pi. 342, Oberthiir and Dauthenay); underparts Cream-Colored with strong wash of Ochraceous-Buff; whitish on insides of forelegs to just below elbow; color of underparts extended distally on forelegs over ankles onto antipalmar faces of inner toes, and on hind legs to knees. Least width of color of under- parts averages 15 (11-19) per cent of greatest width of color of upper parts. Black tip of tail in type longer than hind foot and amounting to 36 per cent of length of tail-vertebrae. Skull and teeth. — Male (based on the type): See measurements and plates 28-30. As described in Mustela frenata macrura except that: Weight, 2.8 grams; basilar length, 41.6; zygomatic breadth less than distance between anterior palatine foramen and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors. Female: Skull unknown. Remarks. — Apparently the first specimens of this race to find tlieir way into a zoological collection were the two young males taken on February 17, 1904, at Limbani, by Geo. Ockenden (sic). M. f. boliviensis is smaller than any other South American weasel except possibly agilis. Better material of the two races probably will show even agilis to be larger. Early in my study of Mustela after examination of the one young specimen, from Limbani, in the United States National Museum, 404 University of Kansas Publs., Mus. Nat. Hist, an account of this race was drawn up, but the account was dis- carded for want of satisfactory material and the animal was referred to macrura. Then, in 1937, when the two other specimens were studied, the race was formally characterized as different from previ- ously recognized kinds. The collector has noted on the labels of the two young from Limbani that they were shot in tlie afternoon when running to- getiier beneath bushes. The frontal sinuses of the type are mal- formed as a result of infestation by parasites. Specimens examined — Total number, 3, as follows: Peru: Carabaya, Limbani, 2 (one in U. S. Nat. Mus, and one in Berlin Zool. Mus.). Bolivia: Nequejahuira, 1 Amer. Mus. Nat. Hist. Mustela frenata (?) gracilis (Brown) Plates 39 and 40 Putorius gracilis Brown, Mem. Amer. Mus. Nat. Hist., 9(pt. 4):182, pi. 17, 1908. Mustela gracilis. Hay, Iowa Geol. Surv. Bull., 23:32, 1914; Hay, Carnegie Instit. Washington Publ. 322A:252, October 15, 1924; Hay, Carnegie Instit. Washington Publ. 390(vol. 2):528, 1930. Mustela frenata gracilis. Hall, Carnegie Instit. Washington Publ. 473:112, November 20, 1936. Type. — Adult skull without lower jaws, probably of a female, no. 12431, Amer. Mus. Nat. Hist.; from Conard Fissure, four miles west of Willcockson, Newton County, Arkansas; obtained sometime in the period 1903 to .1905 inclusive. ( See plates 39 and 40. ) Range. — Known only from the Pleistocene deposit in Conard Fissure, at the type locality in northern Arkansas. Description. — Skull. Probably female ( based on the type ) : See measure- ments and plates 39 and 40; weight unknowTi; basilar length, 38.1; least width of palate less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of rostrum. Comparison and remarks. — The type specimen was the only indi- vidual referred by Brown (1908) to this species. The remaining material of weasels from this deposit was referred by Brown to his Ptitoriiis cicognanii angustidens. Examination of the original ma- terials convinces the writer, too, that the specimens, except no. 12431, are of the species erminea [= cicognanii of Brown]. No. 12431 itself may possibly be erminea but is far more probably of the species frenata. The uncertainty is due to the fact that an occa- sional skull alone of a subadult male erminea is extremely diflBcult certainly to distinguish from a skull alone of an adult female frenata. MUSTELA FRENATA 405 This is true among Recent specimens in the northern Mississippi Valley today; more exactly in Iowa and southern Minnesota the females of frcnata, oftentimes intergrades between the subspecies Mustela frenata longicauda, M. f. noveboracensis and M. /. prim- ulina, by only the skulls are next to indistinguishable from certain, unusually slender skulls of male erminea. At other places where the ranges of the two species meet, this difficulty is not so often encountered. Also, the type of gracilis has the skull broken in such a way that the postglenoid length in relation to the length of the skull as a whole could not be accurately determined in this particular skull. The type specimen of gracilis surely is an adult and because of its small size is thought to be a female. Of known long-tailed weasels of the species frenata, gracilis is structurally nearest to M. /. prim- ulina which occurs in the same region today and to M. /. nove- boracensis, the long-tailed weasel of the eastern United States. M. gracilis differs from noveboracensis and agrees with primulina in possessing well-marked temporal ridges which fuse to form a low sagittal crest, in having the mastoid processes projecting farther, laterally, beyond the braincase, in having the anterior ends of the tympanic bullae produced below the squamosal rather than on the same plane with the squamosal, and in having the bullae more in- flated anteromedially. M. gracilis differs from both noveboracensis ( 97 ^ and 56 $ with skulls of comparable age ) and primulina ( 64 ^ and 24 $ with skulls of comparable age ) in that the zygo- matic breadth amounts to less than 58 per cent of the basilar length. Another difference from any one of the skulls of females of primulina is the longer rostrum, which, when measured from the posterior base of the postorbital process of the frontal to the anterior end of the nasal on the same side, amounts to more than 35 per cent of the basilar length. As pointed out by Brown (1908:182) this specimen represents the extreme of slender skull among known kinds of American weasels. Selected measurements of no. 12431, the type specimen of Mustela gracilis, are as follows: Basilar length of Hensel, 38.1 mm.; length of upper tooth-rows, 14.3 to 14.4; breadth of rostrum, 11.0; interorbital breadth, 8.5; orbitonasal length, 13.6; mastoid breadth, 18.2; length of tympanic bulla, 13.0; breadth of tympanic bulla, 6.3; depth of tympanic bulla, 3.25; outside length of P4, 4.5; inside length of P4, 4.7; breadth of Ml, 3.4; length of inner moiety of Ml, 1.8; depth of skull at anterior margin oi basioccipital, 12.2; depth of skull at posterior borders of last upper molars, 11.3; distance from foramen ovale to tympanic bulla, 3.6 mm. 406 University of Kansas Publs., Mus. Nat. Hist. MUSTELA AFRICANA Desmarest Tropical Weasel (Synonymy under subspecies) Type. — Mustela africana Desmarest, Nouv. Diet. d'Hist. Nat., vol. 19, p. 376. 1818. Range. — Known from the headwaters of the Amazon in eastern Peru and from near the mouth of the same river, on its southern side in Brazil, all within the Tropical Life-zone. See figure 29 on page 221. Characters for ready recognition. — DiflFers from Mustela frenata, the only geographically adjacent species of the genus, by: presence of thenar pad on forefoot; presence of a longitudinal, median, ab- dominal stripe of same color as upper parts; upper lips being broadly edged, entirely round, with color of underparts; failure of longest facial vibrissae to reach posterior margin of ear; absence of p2; relative flatness (see pi. 29, fig. i and pi. 39, fig. h) of tympanic bullae. Characters of the species. — Size large (total length of adults approximately 500 mm.); foot-soles naked; thenar pad present on forefoot; length of claws, measured on concave sides, less than one and one-fourth times depth of claws measured at bases; longest facial vibrissae not reaching posterior margin of ear; tail relatively long haired; tail at all ages terminating in point as is characteristic of only juveniles and very young of Mustela frenata and M. erminea; tip of tail, and muzzle, only slightly darker than remainder of upper parts; upper lips broadly edged with color of underparts; pelage coarse, harsh and sparse; longitudinal, median, abdominal stripe of same color as upper parts present; skull broad and deep; brain- case large, rounded, and much inflated anteriorly; palatal region wide; tympanic bullae less inflated than in any other American species of the subgenus; angle of lower jaw reduced; dental formula I) 1 J H^) i; teeth heavy; medial lobe of Ml but sHghtly larger than lateral lobe. See plates 28, 29, 30, 39 and cranial measurements. Geographic variation. — The reddish versus chocolate color of the upper parts constitutes the only variation of a geographic nature so far detected. Remarks. — One of the most noteworthy of the several unique characters of this large, tropical weasel is the longitudinal, median, abdominal band. The species exhibits the minimum degree of development of certain features that become progressively less apparent as one proceeds southward from Central America. The MUSTELA AFRICANA 407 relative uniformity of the coloration of the upper parts (reduction in intensity of black color on the muzzle and tip of the tail) and the reduction of the tympanic bullae are two cases in point. Viewed dorsally the general outline of the skull is most nearly matched by that of the skull of Mustela frenata meridana from Venezuela or that of M. f. helleri from Peru. However, the resemblance is not close. The tympanic bullae, although unique among American weasels, are more hke those of M. /. meridana from Venezuela than like those of any other kind. The great postorbital width ( relatively less in M. africana than in several South American subspecies of Mustela frenata) and small angular process of the mandible are characters, in varying degrees, also common to all South American weasels. Structurally M. africana clearly is more nearly like other subspecies of M. frenata from South America than it is like any species or subspecies from North America. Mustela africana is the most primitive of the American weasels. The distinctive cranial and dental characters, excepting the reduc- tion in number of premolars, are of a primitive nature. For ex- ample, the relatively wide postorbital region, the large braincase that is inflated anteriorly, and the flattened, tympanic bullae, are points of resemblance to the holarctic Mustela erminea, which species is regarded as nearest the original stem form; also the mentioned characters correspond to ontogenetic stages passed through by other weasels. Mostly on these accounts, one is led to look upon M. africana as a migrant from North America. It may have become isolated from its original stock, by a water bar- rier in the Central American region, for a length of time suJQBcient to permit of a degree of differentiation to develop between it and the North American weasels which prevented crossbreeding with the frenata stock when that stock, at a later time, reached South America. This assumption is suggested only by evidence from the Recent specimens. No remains of true weasels (subgenus Mustela) have been recorded from deposits in South America older than the Recent period. The alternate possibility, that M. africana inter- grades with some race of M. frenata in western or northern South America, has been considered and regarded as highly improbable. Cabrera (1940:15) has made the distinctive structural characters of Mustela africana basis of the generic name Grammogale to in- clude the one species africana. I am incHned to accord Gram- mogale only subgeneric rank. It is possibly significant that Mustela africana is intermediate in several respects between Lyncodon and typical Mustela. The 408 University of Kansas Publs., Mus. Nat. Hist. median, longitudinal, abdominal band of the same color as the upper parts in M. africana and the relative uniformity of the col- oration of its upper parts might be considered as an intermediate stage between the dark, bicolored (black muzzle and tail tip and brown body) upper parts and light-colored underparts of the North American weasels on the one hand and the light, uni- colored upper parts and dark-colored underparts of the Patagonian weasel ( Lyncodon ) on the other hand. The number of premolars, ^-^, is also intermediate between the numbers | and | of the North American Mustela and the Patagonian Lyncodon, respectively. The more medially, as opposed to anteriorly, directed medial cusp of P4 (characteristic of approximately half of the specimens exam- ined), and the structure of the skull in general of M. africana also seem to be morphologically intermediate between those parts of Mustela and Lyncodon. On the chance that Lyncodon is closely enough related to Mustela, to be included in a group with Mustela rather than in a group with Grisonella, it is worth noting that Lyncodon lujanensis Ameghino (1889:324, 325), from the villa of Lujan and at the city of Cordoba, at each place in the Pampean [— Pleistocene] forma- tion of Argentine (see also Cabrera, 1928:263) is the first and only fossil form of this group recorded from the whole of South Amer- ica. Actually, however, Lyncodon seems to me to be as nearly related to Grisonella, if not more so, than to Mustela. If Lyncodon is more closely related to Grisonella and Grison than to Mustela, then the above remarked intermediacy in characters of M . africana has more of interest as a tendency to parallelism than it has of phylogenetic import. Appraisal of phylogenetic relationships would require appraisal of the ancestral stem forms of the Grison stock and the Mustela stock. None of either is known from deposits of the Pliocene, the period of time immediately preceding the Pleisto- cene. None of the skulls of Mustela africana seen or figured has the nasomaxillary sutures entirely obliterated and the specimens would, judged on this character alone and by analogy with North American species, be regarded as young and subadult. However, the sutures close at what seems to be a later age than in M. frenata and M. erminea. The condition of the mammae in the type specimen of M. stolzmanni and in the specimen from Moyobamba, indicate that they have borne young. North American weasels old enough to bear young lack visible traces of the nasomaxillary sutures. I have MUSTELA AFRICANA 409 examined no skulls of africana with greatly worn teeth and hence cannot say if the sutures are obliterated in advanced age. If available data be correct, this species is unique within the genus in that the two sexes are of approximately the same actual size and of the same relative proportions in the body and in the skull. There was no difference between individuals said to be of differ- ent sexes from Pard, described and figured by Goeldi (1904:61-62, pis. 1, 2). The undoubted female, type specimen of Mustela afri- cana stolzmanni, is as large as the undoubted male, no. 37475, of the same species, but of a different subspecies, from Para, Brazil. All the specimens of M. a. africana that I have handled are labeled male and those of M. a. stolzmanni female. More material may show that the female is smaller than the male, as is the case in all near relatives of M. africana. Little has been recorded concerning the habits of this species. Tate (1931:254) states that a live individual which he saw in a cage at Pard had been captured "swimming in the salt water of the estuary about half a mile away from the shore." On the label of the speci- men from Moyobamba, there appears: "caught in Willow tree." Subspecies examined. — All described forms, of which there are two. Mustela africana africana Desmarest Tropical Weasel Plates 28, 29, 30 and 41 Mustela africana Desmarest, Nouv. Diction. d'Hist. Nat., 19:376, 1818; Cabrera, Bol. Real Soc. Espari. de Hist. Nat., 13:429, November, 1913; Cabrera, Bol. Real Soc. Espaii. de Hist. Nat., 14:175, pi. 1, March, 1914. Putorius (Mustela) brasiliensis paraensis, Goeldi, Zool. Jahrb. abt. f. sys- tematik, geogr, u., Biol., 10:556, pi. 21, September 15, 1897, type from Para, Brazil, near Pard, Ward of Marco da Legoa, Brazil; Goeldi, Bol. do Mus. Paraense, 3:195 [translation of orig. descr.], August, 1901. Putorius paraensis, Goeldi, Bol. do Museu Goeldi, 4:61, pis. 1, 2, 1904. Mustela affinis paraensis, HoUister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914. Mustela paraensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:105, April 28, 1916; Tate, Joum. Mamm., 12:253, August 24, 1931. Mustela stolzmanni paraensis. Hall, Carnegie Instit. Washington Publ. 473: 111, November 20, 1936; Hall, Physis, 16:167, pi. 1, figs. 1-4, 1939. Type. — Male, adult or subadult, mounted; no. 848, Paris Museum; from the "Cabinet de Lisbonne 1808," originally from South America as determind from the characters of the animal; probably came from Brazil, and for the present assvuned to be from Pard. On August 25, 1937, the skull was in the mounted skin and the specimen was in the position shown in the figure published by Cabrera ( 1914, pi. 1 ). Except for the loss of the distal part of the tail, and fading because of exposure to hght, the specimen was in good condition. See also tmder remarks. 410 University of Kansas Publs., Mus. Nat. Hist. Range. — Known from the south side of the Amazon River, near its mouth at Para and Cameta, Rio Tocantins, in the Tropical Life-zone of Brazil. See figure 29 on page 221. Characters for ready recognition. — Differs from Mustela frenata, the only other geographically adjacent species of the genus, in presence of median, longitudinal, abdominal stripe of same color as upper parts and naked foot- soles, in absence of p2 and in reduced size of tympanic bullae ( see pis. 29 and 30 ) and from Mustela africana stolzmanni by lighter color of upper parts which although near Chestnut-Brown are in adults 10' I (darker in yg. M. C. Z., no. 30802), instead of 11' n as in M. a. stolzmanni. Description. — Size. — This is a relatively large weasel. Goeldi (1897:559) gives the total length of the type specimen of his P. b. paraensis, a female, as 520 mm. (495 in the flesh) and, by computation from his figures, the length of the tail as 200 (205 in the flesh). These measurements probably include the hairs on the tip of the tail as probably also do the measvuements given of two other specimens (see Goeldi, 1904:62). One of these specimens, a female, measured: Total length, 520; length of tail, 200. The other specimen, a male, measured: Total length, 510; length of tail, 200. The skin of no. 37475, Amer. Mus. Nat. Hist., a male, has the following measiurements written on the at- tached label: Total length, 548; length of tail, 234; length of hind foot, 56. The hairs project 20 mm. beyond the tip of the last vertebra of the tail and probably are included in the measurements of total length and length of tail. Collectors' measurements of a young male from Cameta, and a subadult labeled as male, from Para Murutucu, are respectively as follows: 500, 430; 210, 190; 50 and 54. Externals. — Foot-soles naked, except for a few scattered hairs on ventral sides of interdigital membranes; length of claws, measured on concave sides, not more than one and one-fifth times depth of claws measured at bases; carpal vibrissae not extending beyond apical pad of first digit (not beyond hypo- thenar pad except in one young specimen); longest facial vibrissae not extend- ing to posterior margin of ear; superior genal tuft not found, hairiness of foot- soles as shown in figure 22. Color. — Upper parts near (10 /) Chestnut-Brown and relatively uniform since tip of tail and muzzle are only slightly darker than remainder; underparts with longitudinal stripe of same color as upper parts extending along median line of belly from throat or breast posteriorly to within 40 to 50 millimeters of anus. Underparts otherwise near ( 20" a ) Olive-Ocher ( hps and chin whiter in one young specimen). Color of underparts extends distally on median sides of forelegs to bare foot-soles and on median side of hind legs two-thirds of distance from knee to ankle. Upper lips broadly edged with whitish, which color passes posteriorly below and not touching eye to ventral margin of concha of ear. An inverted, basally broad, V-shaped extension passes upward 4 millimeters, just posterior to the eye. Skull and teeth. — See measurements (plates 28-30). Male: (based on 3 adult and subadult topotypes and figures and descriptions published by Goeldi, 1897 and 1904.) Weight, 7.0 grams; basilar length, 45.8 (44.6-47.8); skull broad and deep; braincase large, rounded, and much inflated anteriorly; dis- tance from postorbital process to anterior, nasal notch approximately equal to breadth across exoccipital condyles; palatal region wide; tympanic bullae less MUSTELA AFRICANA 411 inflated than in any other species; mastoid bone, laterally, concave; length of upper tooth-rows in adults and subadults less than breadth of palate measured between two outer margins of fourth upper premolars; alveolar distance be- tween CI and P4 less than length of P4; teeth heavy; medial lobe of Ml only slightly larger than lateral lobe; deuterocone of P4 heavy and often incHned mesially; p2 absent ( P2 present above on both sides in only one of seven speci- mens seen or described); lower jaw heavy; masseteric fossa not extending anteriorly to posterior fourth of talonid of ml; paraconid of ml low and base of cleft between it and protoconid relatively low on tooth. Female: No skull examined but from figures pubHshed by Goeldi (1904, pi. 2), apparently as described in the male. Remarks. — Desmarest in 1818 gave a remarkably good description of this animal which he named as a new species, Mustela africana, but mistakenly indicated that the single specimen known to him came originally from Africa. Until 1913 the name was applied, wrongly, to weasels of northern Africa or to those of the Azores Islands and St. Thomas Island. In that year Cabrera (1913:429) identified the species with the one later named Piitorhis ( Mustela ) brasiUensis paroensis by Goeldi (1897:556, pi. 21) from Para, Brazil. Despite Cabrera's clear identification in 1913, and his later mention of the correct application of the name Mustela africana, it was not correctly employed by other authors, including myself who even as late as 1936 (p. Ill) instead used Goeldi's name. In 1937 Mr, Cabrera called my attention to his published account of Mustela africana and so permitted me to examine the type specimen in the Paris Museum, whither I was bound when I received Mr. Cabrera's letter. My own examination of the specimen fully confirmed the conclusions published by Cabrera (1913:429). As a matter of historical interest, however, it is worth noting that Cabrera ( 1913 ) originally supposed the type specimen to have been taken as booty of war from Portugal by the French and that Cabrera later, at the request of P. Trouessart, pointed out (1914:176) that the specimen had been acquired in exchange ("a cambio") since according to Dr. Trouessart the Museum register showed that offer had been made to Portugal to return this and other specimens but that Portugal had replied that it had nothing to reclaim. Dr. P. Rode in August, 1937, at the Paris Museum, gave it to me as his opinion that the spcimen had been an outright gift from the "Cabinet de Lisbonne" to E. Geoffroy St.-Hilaire on his trip to Portugal in 1808 when he was given also from the same cabinet several primates, all from Brazil. Of the labels attached to the pedestal on which the specimen is mounted, that of most ancient appearance is glued to the bottom of the stand and bears in a hand apparently written be- 412 University of Kansas Publs., Mus. Nat. Hist. fore Trouessart's entries on the same label, the information "Du Cabinet de Lisbonne 1808" and "J. H. S. 1809." The opened mouth of the mounted specimen permits one to determine that P2 is absent on each side above. The stuffed scrotal pouch and hair projecting downward about the preputial opening clearly show the animal to have been a male. The least faded portions of the mounted specimen, its sides, are of the same reddish color as characterizes adults from Pard and not of the darker chocolate color of specimens of M. stolzmanni from Peru. The specimen is indistinguishable from topotypes of P. paraensls of Goeldi and his name will have to fall as a synonym of Mustela africana Desmarest. Goeldi gave an extended description, with figures of the skull, head, and entire animal, when he named paraensis. As his account shows, he was unaware that Taczanowski had described a similar weasel from the headwaters of the Amazon, or for that matter that any weasel excepting Mustela affinis Gray, had been found in South America. Goeldi's later account of additional specimens (1904: 61, pis. 1, 2) gives much useful information about the animal. Photographs of several specimens and photographs and detailed measurements of several skulls are presented by him. Para, and Cameta, Brazil, places from which Mustela africana africana is known, are nearly 2000 miles from the localities in east- ern Peru and eastern Ecuador from which M. a. stolzmanni is known, and no specimens, from intermediate localities, are avail- able to show actual intergradation of the two. However, the similarity in structure of the two weasels is so great as to indicate close affinity. Furthermore, it is understood that environmental conditions at and between the two localities are similar. These con- siderations, in the light of our knowledge of actual intergradation of geographic races of weasels in other places, cause me to treat, with a feeling of assurance, M. africana [^= P. paraensis Goeldi] and M. stolzmanni Taczanowski as subspecies of a single species. M. Rodolpho Legueira Rodriguez writes me, under date of June 16, 1928, that the type specimen of Putorius (Mustela) brasiliensis paraensis Goeldi is stuffed and preserved in a "vitrine" at the Museum Goeldi (Museum Paraense) De Historia Natural e Eth- nographia. Para, Brazil. The one young specimen seen, that from Gametd, is darker col- ored than any of the four older specimens examined. It is almost exactly the Chestnut Brown of Ridgway (1912) and therefore ap- MUSTELA AFRICANA 413 preaches closely in color the adult specimens of M. a. stolzmanni. This same tendency to greater richness of color in young than in adults is seen also in Mustela frenata. Specimens examined. — Total number, 5, all from eastern Brazil, as follows: Pard, 2 (V, V); Para Murutucu, 1"; Rio Tocantins, Cameta, 1'; type speci- men, 1*. Mustela africana stolzmanni Taczanowski Tropical Weasel Plates 39 and 40 Mustela stolzmanni Taczanowski, Proc. Zool. Soc. London, 1881:835, November 15, 1881; Allen, Bull. Amer. Mus. Nat. Hist., 35:105, April 28, 1916. Mustela stolzmanni stolzmanni. Hall, Carnegie Instit. Washington Publ. 473:111, November 20, 1936; Hall, Physis, 16:167, pi. 1, figs. 5, 6. 1939. Type. — Female, adult, mounted skin, with skull separate; no. 563, Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Yurimaguas, Peru; 1880; obtained by J. Stolzmann. The skull (plate 40), mounted with the skin but removed by me for study, consists of the premaxillae, maxillae, two halves of the lower jaw and dentition. Of these parts, right m2, left coronoid process, right P4 and Ml and adjacent part of maxilla are lost. The skin is well mounted, in a good state of preserva- tion and shows no fading due to exposure to light. Inguinal mammae are dis- tinctly shown on the skin and prove that the specimen is a female. Except for a few scattered hairs on the lower throat, a spot six by eight millimeters on the medial side of the region of the olecranon of the left foreleg and another of similar size in the left axilla, the underparts are, excepting the ventral longi- tudinal, abdominal stripe, unmarked by color of the upper parts. Range. — Known from the Tropical Life-zone of eastern Ecuador and Peru from Jatun Yacu south to Valle del Perene. See figure 29 on page 221. Characters for ready recognition. — Difi^ers from Mustela frenata, the only other geographically adjacent species of the genus, in presence of median, longtitudinal, abdominal stripe of same color as upper parts and naked foot soles, in absence of p2 and in reduced size of tympanic bullae (see pis. 28, 29, 30, 39 and 40) and from Mustela africana africana by darker color of upper parts which, although near Chestnut Brown, are ll'n instead of 107 as in M. a. africana. Description. — Size. — Male : unknown. Female: Taczanowski (1881:836) gives, among others, the following meas- urements of the type specimen: Total length, 523, length of body, 260; length of tail without hair, 190 (with hair 224); length of hind foot, 54. Whether or not the measurements were taken from the animal when in the flesh I do not know. Allowing for shrinkage of hind feet and changes due to the pos- ture of the now mounted specimen, I get from it essentially the same meas- urements. Collectors' measurements of a subadult from Moyobamba and a young female from Valle del Perene, are respectively, as follows: 469, 415; 1. American Mus. Nat. History 3. Museum of Comparative Zoology 2. British Museum of Natural History 4. Paris Museum 414 University of Kansas Fuels., Mus. Nat. Hist. 184, 160; 57, 52. My own measurements of the dry hind feet on the sldns are respectively, 48 and 49. Externals. — As described in M. a. africana except that the length of tlie concave sides of the claws are approximately one and one-fourth times the depth; thus the claws are relatively longer than in M. a. africana. Color. — As described in M. a. africana with the following noted exceptions: Upper parts near (11' n) Chestnut-Brown; area of hghter ventral coloration on the throat and sides of head less strongly tinged with yellow; pelage more dense, finer and softer than in M. a. africana. Skull and teeth. — Male: Skull unknown. Female: See measurements and plate 39 and 40. As described in male of Mustela africana africana except that: Weight, 4.7 grams. As contrasted with M. a. africana, the dentition of the lower jaw is lighter; the transverse diameter of m2 is 1.2 mm. in the type and also in the specimen from Moyobamba as against 1.5 to 1.7 in three male topotypes of M. a. africana. Remarks. — After the Polish naturalist, Stolzmann, in the course of his explorations in Peru, obtained the single specimen which was made the type, no other naturalist, so far as known, visited the type locality until thirty-two years later when Wilfred H. Osgood and M. P. Anderson spent more than a month collecting at Yuri- maguas (see Osgood, 1914:147), but secured no topotypes of this little-known weasel. C. O. Schunke took the second specimen in tlie Valle del Perene in April, 1921; L. Rutter on January 25, 1924, took the third specimen, and W. Clark-Maclntyre took the fourth specimen on the Jatun Yacu. This obscure place name is shown on the map (fig. 4, page 827) published by Brown (1941) and is the stream flowing from the west to the town of Napo, Napo is situated at approximately 1° 2' S and 77° 49' W. In the female from Moyobamba there are only 3 pairs of mammae. One pair is inguinal and two pairs are on the posterior part of the abdomen. Taczanowski (1881:836) relates that this species was taken in the forest to which it appears to be restricted since the inhabitants of the village did not know of the animal. He points out also that the previously known Peruvian species [M. /. macrura and M. f. agilis] live in the treeless territory of eight to eleven thousand feet altitude whereas M. stolzmanni was found in the humid forest of the great plain of the Maynas at an elevation of 500 feet or less above sea level. The frontal sinuses of the specimens seen reveal no malformation as a result of infestation by parasites. Specimens examined. — Total number, 4, as follows: Ecuador: R. Tatun [= Jatun] = Yacu, 1, Mus. Comp. Zool. Peru: Yurimaguas, 1 in Mus. Polonais d'Hist. Naturelle, Warsaw; Moyo- bamba, 2700 ft. [6° S, 77° W], 1 in Brit. Mus. Nat. Hist.; Valle del Perene, 1200 meters, 1 in Amer. Mus. Nat. Hist. EXPLANATION OF CRANIAL MEASUREMENTS APPEARS ON PAGE 417 (415) 416 University of Kansas Publs., Mus. Nat. Hist. 11,2.3 CI P2P3 P4 Ml il.2.3 Fig. 31. Four views of the skull and a lateral view of the left lower jaw to show points between which measurements of the skull were taken. Based on M. /. primulina, from 3 mi. E Bergman, Boone County, Arkansas, obtained December 12, 1933, by B. G. Roberts; ad. 9. 62854 Mus. Vert. Zoiil. X 1 %• Cranial MEAStmEMENxs 417 EXPLANATION OF CRANIAL MEASUREMENTS Basilar length (of Hensel). — From the anteriormost border of the foramen magnum to a line connecting the posterior margins of the alveoh of the first upper incisors. F to F' on fig. 31. Condylobasal length. — Least distance from a hne connecting the posteriormost parts of the exoccipital condyles to a line connecting the anteriormost projections of the premaxillary bones. Length of tooth-rows. — Least distance between a line connecting posterior borders of upper molars and a line connecting anterior faces of middle upper incisors. G to C on fig. 3L Breadth of rostrum. — Least distance from lateral base of hamular process of lacrimal bone to corresponding point on opposite side of skull. B to B' on fig. 3L Interorbital breadth. — Least distance across top of skull between orbits (eye sockets). OtoO'onfig. 3L Orbitonasal length. — Distance on anterior part of skull from posterior margin of base of postorbital process of frontal bone to posteriormost part of anterior border of nasal bone on same side of skull. A to A' on fig. 31. Mastoid breadth. — Greatest distance across mastoid bones perpendiciJar to long axis of skull. E to E' on fig. 31. Zygomatic breadth. — Greatest distance across zygomatic arches of cranium perpendicular to long axis of skull. D to D' on fig. 31. Tympanic bulla: Length. — From posterior face to most anterior part of anterior border. H to H' on fig. 31. Breadth. — From bottom of pit immediately posterior to external auditory meatus to medial face of bulla at right angle with longitudinal axis of skull. J to J' on fig. 31. Depth. — Least distance from ventral face of basioccipital, excluding median ridge, to line touching ventralmost points of the two bullae. L to L' on fig. 31. ml, Length. — Greatest length which rarely or never is alveolar length. P4.— Lateral. — Length from posterior margin of tooth to anteriormost part of the protocone ( anterolateral cusp ) . Medial. — Length from the posterior margin of tooth to anteriormost part of the deuterocone ( anterointemal cusp). ML— Breadth. — Distance from medial edge of crown to lateral margin of crown, approximately at a right angle with longitudinal axis of the skull. Length. — Greatest diameter, anteroposteriorly, of the inner lobe or inner half of the tooth. Depth of skull at anterior margin of basioccipital. — Measured from anterior end of ventral face of basioccipital, excluding median ridge, vertically to dorsal face of parietal excluding sagittal crest. K to K' on fig. 31. Depth of skull at posterior borders of Msl. — Measured from ventral face of palatine bones at posterior edge of upper molars to dorsal face of frontals in plane of postorbital processes of frontals. S to S' on fig. 31. 14—3758 418 University of Kansas Publs., Mus. Nat. Hist. TABLE OF CRANIAL MEASUREMENTS d o o O .a O a •-3 o 1^ S ^ ^ s 3 T3 ja L^ M J3^ -^ — • R s 1^ J2 JO M » C 2 03 S c **I! o -tJ M •S 0) M M ■^ •£ a •n 1^ a a 1 pa v-5 m O Mustela frenata 45.1 16.6 13.7 10.9 15.3 47.0 17.4 14.6 11.9 15.9 43.2 15.9 13.0 10.4 14.8 44.7 16.3 13.7 11.3 15.7 47.0 16.8 14.9 12.0 16.7 43.3 15.9 13.1 10.5 14.8 44.0 16.3 13.2 10.9 15.3 46.0 17.4 14.4 12.0 16.4 41.5 15.2 12.1 9.7 14.2 44.4 16.5 13.3 10.6 15.6 46.1 17.7 14.1 11.3 16.4 40.6 15.6 12.3 9.8 13.7 43.0 15.6 13.0 10.2 14.9 45.4 16.5 13.4 11.0 15.7 39.7 14.7 11.6 9.6 13.8 37.5 13.4 10.7 8.7 12.9 39.8 14.5 11.2 9.3 13.4 36.4 12.8 9.8 7.9 12.1 36.5 13.4 10.5 8.5 12.9 38.1 14.0 11.0 8.6 13.3 35.2 13.0 10.0 8.3 12.1 37.3 13.5 10.8 8.6 13.0 37.9 14.1 11.2 8.9 13.5 36.8 13.0 10.4 8.5 12.8 37.2 13.4 10.6 8.8 13.0 37.6 13.7 11.3 9.7 13.5 37.0 12.9 10.0 8.0 12.0 36.5 13.1 10.5 8.4 12.6 37.5 13.7 11.0 8.8 13.1 35.9 12.8 9.8 7.8 12.0 Mustela frenata 45.8 16.5 13.3 11.0 16.2 44.9 16.6 14.5 11.8 15.6 46.9 17.1 14.4 11.0 15.8 45.0 16.1 14.0 11.2 15.3 45.7 16.6 14.1 11.3 15.7 37.7 14.0 11.6 9.1 13.0 38.2 13.8 11.5 9.2 13.6 Mustela frenata 44.8 16.8 13.5 10.6 14.9 46.0 17.8 14.1 11.5 15.4 43.8 16.2 12.9 10.0 14.3 average maximum minimum average maximum minimum average maximum minimum average maximum minimum average maximum minimum average maximum minimum average maximum minimum average maximum minimum average maximum minimum average maximum minimum A. N. S. P. M. C. Z. M. C. Z. M. C. Z. M. C. Z. M. C. Z. average maximum minimum average maximum minimum average maximum minimum average maximum minimum cf ad. 10 Adirondaks, New York . c? ad. 10 Massachusetts. d' ad. 10 Liberty Hill, Connecticut. cf ad. 10 Beaver Dam, Wisconsin. 1 \ cf ad. 10 Washtenaw Co., Michigan. 9 ad. 8 Adirondaks, New York . 9 ad. 5 Massachusetts. 9 ad. 6 Maryland. 9 ad. 5 Beaver Dam, Wisconsin . 9 ad. 9 Washtenaw Co., Michigan. 4137 cf ad. Washington Co., Maine. . . 7267 cf ad. Moose Head Lake, Maine . 5501 cf ad. Bucksport, Maine 9142 d^ ad. Bucksport, Maine av 9101 9 sad. Bucksport, Maine . 9122 9 sad. Bucksport, Maine . cf ad. 10 Douglas Co., Kansas. cf ad. and sad. 8 Boone Co., Arkansas. 9 ad. and sad. 11 Douglas Co., Kansas. 9 ad. and sad. 6 Arkansas. 44.7 16.5 13.2 10.5 15.0 46.5 17.1 14.0 11.8 16.3 42.5 15.6 12.6 9.7 14.1 38.9 14.4 11.3 8.6 12.9 40.7 15.3 12.0 9.2 13.4 37.6 13.8 10.8 7.9 12.4 39.3 14.1 11.4 8.7 13.1 40.1 14.6 11.9 8.8 13.7 38.8 13.7 U.O 8.5 12.6 Cranial Measurements 419 j3 ■s 1 o e o >> tS3 Tympanic Bxilla a -s c 0) s «-4 2 Depth of Skull at Ant. marRin of basioccipital |l 1 •a 2 r C j3 m J5 p r 2 0) cS >-) 1 ffl ^ a g-ao D noveboracen.sis 23.3 26.7 24.7 28.0 22.0 25.3 15.4 16.2 14.6 7.8 8.4 7.0 3.5 4.0 3.1 5.9 6.2 5.5 5.3 5.7 4.9 5.5 5.9 4.9 4.3 4.5 4.0 2.4 2.7 1.9 14.1 14.5 13.5 12.7 13.3 12.2 23.2 25.6 21.8 26.5 28.2 25.1 15.4 16.4 14.3 7.7 8.2 7.3 3.4 3.7 3.1 5.9 6.3 5.6 5.2 5.5 5.0 5.5 5.9 5.2 4.4 4.7 4.0 2.4 3.0 2.0 14.1 14.7 13.5 12.7 13.0 12.0 22.4 23.5 20.8 26.3 28.0 24.0 15.0 15.9 14.4 7.5 8.0 6.9 3.3 3.6 3.0 5.8 6.3 5.3 5.1 5.6 4.6 5.4 6.1 5.0 4.2 5.0 3.7 2.2 2.5 2.0 14.3 15.2 13.4 12.6 13.2 11.7 23.0 23.8 22.1 26.3 27.4 23.9 15.2 16.2 14.2 8.0 8.2 7.7 5.8 6.1 5.6 5.3 5.7 4.7 5.5 5.8 5.1 4.3 4.6 3.9 2.4 2.7 2.1 13.9 14.5 13.0 12.6 13.2 11.8 22.0 22.8 21.1 24.9 26.1 23.5 14.7 15.2 14.1 7.6 7.9 7.3 5.6 6.0 5.0 5.0 5.3 4.7 5.3 5.6 4.9 4.2 4.5 3.8 2.2 2.5 1.9 13.4 14.2 12.6 11.9 12.3 11.5 18.7 19.3 17.5 20.9 22 1 io^o 13.5 14.3 12.6 6.5 7.0 5.9 2.8 2.9 2.5 4.8 5.2 4.5 4.3 4.7 4.0 4.6 5.0 4.2 3.5 3.7 3.2 1.7 1.9 1.7 11.4 12.1 10.3 10.2 10.8 9.4 18.5 19.6 17.2 20.1 20.8 19.3 13.3 13.9 12.8 6.3 6.9 5.6 2.3 2.7 2.0 5.0 5.1 4.9 4.3 4.5 4.2 4.5 4.7 4.4 3.5 3.7 3.3 1.8 2.0 1.4 11.4 11.6 11.0 10.0 10.4 9.6 18.8 19.6 18.2 21.0 21.7 20.1 13.1 13.5 13.0 6.6 6.8 6.4 4 9 5.2 4.7 4.4 4.6 4.2 4.7 4.9 4.5 3.6 4.1 3.1 1.9 2.3 1.4 11.9 12.6 11.6 10.9 11.4 10.3 18.7 19.5 17.6 21.0 21.7 20.2 13.5 13.8 13.1 6.5 6.8 6.1 4.9 5.0 4.9 4.5 4.5 4.4 4.6 4.8 4.3 3.5 3.7 3.3 1.8 2.1 1.5 11.6 12.3 10.8 10.2 10.3 9.8 18.5 19.1 18.1 20.5 21.3 19.5 13.2 13.8 12.9 6.3 6.7 6.0 4.8 5.0 4.6 4.3 4.5 4.1 4.5 4.7 4.3 3.5 3.7 3.3 1.8 2.0 1.5 11.8 12.2 11.3 10.3 10.7 10.0 oceiaor 24.5 24.2 24.0 24.2 24.2 27.5 27.7 28.0 27.4 27.7 15.7 15.2 16.4 15.8 15.8 8.5 7.7 8.0 7.3 7.9 3.5 3.1 4.0 2.8 3.4 6.2 6.0 6.0 6.1 6.1 5.6 5.2 5.8 5.3 5.5 5.9 5.6 6.1 5.6 5.8 4.6 4.2 4.5 4.3 4.4 2.8 2.4 2.6 2.4 2.6 13.3 14.2 14.7 14.5 14.2 13.0 13.0 13.2 12.6 13.0 19.0 19.5 21.7 21.8 14.1 13.6 6.3 6.5 2.6 2.5 5.2 4.6 4.6 4.3 4.6 4.5 3.8 3.5 2.1 1.7 12.5 12.3 10.5 10.3 primulina 24.0 24.9 23.2 27.2 28.2 26.2 15.5 16.3 14.4 8.3 8.8 8.0 3.4 3.9 3.0 6.1 6.6 5.8 5.5 5.9 5.2 5.8 6.1 5.5 4.6 4.7 4.3 2.4 2.6 2.2 14.2 14.8 13.6 12.8 13.2 12.0 24.1 24.8 22.8 26.9 27.7 26.1 15.5 16.4 14.9 8.1 8.8 7.5 3.6 4.2 3.1 6.0 6.5 5.8 5.5 6.0 5.2 5.6 6.2 5.2 4.3 4.9 4.0 2.2 2.5 2.0 14.2 14.6 13.9 12.3 13.0 11.7 20.3 21.4 18.8 22.6 23.8 21.1 13.5 15.1 13.0 6.9 7.5 6.3 2.9 3.3 2.5 5.2 5.7 4.8 4.8 5.2 4.5 5.0 5.4 4.6 3.5 4.2 3.4 1.8 2.1 1.5 12.6 13.5 11.7 11.2 11.7 10.0 20.3 21.0 19.8 23.1 23.8 22.5 14.0 14.5 13.5 6.9 7.2 6.5 2.9 3.0 2.8 5.2 5.5 5.0 4.8 5.0 4.5 5.0 5.3 4.7 3.8 4.2 3.5 1.9 2.0 1.7 12.5 13.0 12.3 10.7 11.8 10.3 420 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial MEASxmEMENTs — Continued e 2 '*^ o v a o O o "3 O to a s OQ & ^ ja o J3 2 •a S J3 a •«^ til'c) 0 o 5 In .a 13 9 "o J3 5 i s d o o ••io a 3! 0) jQ O a h n hJ n o U. S. N. M. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. U. S. N. M. Cornell Cornell U. S. N. M. M. V. Z. M. V. Z. M. V. Z. F. S. M. A. N. S. P. A. N. S. P. M. V. Z. Walker Walker Dickey Dickey Dickey Walker Dickey U. S. N. M. A. M. N. H. N. M. C. F. M. N. H. F. M. N. H. U. S. N. M. N. M. C. U. S. N. M. A. M. N. H. M. V. Z. U. S. N. M. Mustela frenat» 246345 d" ad. Convent, Louisiana 43.4 15.4 13.7 11.5 14.5 37515 cfsad. Remy, Louisiana 43.8 16.4 12.7 10.9 15.0 38901 cf sad. Springville, Louisiana *45.0 *16.5 15.6 Mustela frenata 47165 d" ad. Talbot Co., Georgia 50.1 18.3 15.0 12.5 16.5 47144 d' ad. Talbot Co., Georgia 49.2 18.3 13.8 11.7 16.9 471C6 cf ad. Talbot Co., Georgia 48.5 17.6 14.6 11.5 16.5 47167 d' ad. Talbot Co., Georgia 45.8 17.3 14.2 11.5 16.2 47147 cf ad. Talbot Co., Georgia 48.1 18.0 14.0 10.9 16.9 av. 5 48.3 17.9 14.3 11.6 16.6 41023 d* ad. Thomas Co., Georgia 48.8 17.5 14.9 11.9 16.0 41025 d' ad. Grady Co., Georgia 44.8 16.9 13.2 10.9 15.5 40934 d'ad. Grady Co., Georgia 47.8 17.4 14.1 12.1 15.5 40935 d' ad. Grady Co., Georgia 47.4 18.0 13.6 11.7 16.2 av. 4 47.2 17.5 14.0 11.7 15.8 223880 d" ad. Okefinokee Swamp 49.0 18.8 14.2 11.6 14.6 198 d' ad. Okefinokee Swamp 47.3 18.0 14.0 11.4 16.3 652 d' ad. Okefinokee Swamp 47.0 17.0 13.8 11.8 16.0 180802 d' ad. Autaugaville, Alabama 46.5 17.3 13.4 10.7 16.3 51527 9 ad. Talbot Co., Georgia 43.5 16.4 12.5 9.8 14.5 41024 9 ad. Thomas Co., Georgia 42.7 16.0 12.0 9.6 14.7 41022 9 ad. Thomas Co., Georgia 44.0 16.1 12.8 10.6 15.3 Mustela frenata 49387 d'ad. .A.popka, Florida 49.8 18.6 15.1 12.2 15.8 9379 9 ad. Tarpon Springs, Florida 44.2 16.6 13.8 11.0 15.8 8515 9 yg. Pasco Co., Florida 15.9 11.8 9.6 14.2 Mustela frenata 53795 d'ad. Elk River, Minnesota 49.2 19.1 15.9 11.9 17.1 A23 d'ad. Elk River, Minnesota 49.0 18.7 15.9 13.5 16.5 A37 d' sad. Elk River, Minnesota 48.7 18.9 15.2 11.9 16.5 A865 d" sad. Elk River, Minnesota 46.8 17.6 15.4 16.7 A846 d" sad. Elk River, Minnesota 46.1 18.0 14.2 10.6 15.5 11548 9 ad. Elk River, Minnesota 42.3 16.4 13.1 10.7 15.0 A174 9 ad. Elk River, Minnesota 43.2 16.2 13.8 10.8 15.3 9688 9 ad. Elk River, Minnesota 43.2 16.8 12.7 10.5 15.0 188410 9 ad. Elk River, Minnesota 43.0 16.2 13.2 10.7 14.9 av. 4 42.9 16.4 13.2 10.7 15.1 Mustela frenata 15875 d'ad. Red Deer, Alberta 46.5 18.6 15.1 12.1 15.3 8060 d'ad. Sweet Grass HilLs, Alberta 45.5 18.1 15.5 12.2 15.5 7021 d'ad. Canadian Nat. Park, Alberta . . 46.8 17.8 13.8 11.4 16.0 8567 d'ad. Calgary, Alberta 44.7 17.2 13.9 11.3 14.8 75725 d'ad. St. Albert, Alberta 46.5 18.3 15.0 12.4 15.6 av. 5 46.0 17.9 14.7 11.9 15.4 6968 9 ad. Davsland, Alberta 43.7 16.8 13.1 14.9 68731 9 ad. S. Edmonton, Alberta 42.5 16.0 13.4 10.3 14.3 16044 9 ad. Blindman River, Alberta 40.0 15.1 12.2 8.8 13.2 53792 9 ad. Grafton, North Dakota 42.8 16.5 13.6 10.5 14.9 75483 9 ad. Wingard, Sask 42.3 16.9 12.9 9.8 15.6 av. 5 42.3 16.3 13.0 9.9 14.6 Cranial Measurementts 421 ■B Tympanic Bulla 2 s •s— 13^ ■s , ^ « o -■H ' \ ^.9 "3 ■3 5-R 3i J .2 g •sag «- 1 s 1 N3 & J a; 3 ■3 -C 03 a arthuri 22.3 26.6 15.4 7.7 3.3 5.4 4.9 5.1 3.7 2.0 14.8 12.4 22.3 25.5 15.5 7.5 3.7 4.9 5.7 5.8 4.2 2.3 14.2 12.4 16.0 8.3 5.8 5.5 5.7 4.6 2.5 olivacea 26.7 30.8 17.2 9.2 4.0 6.8 6.2 6.2 4.7 2.5 !'5.0 13.8 26.0 29.0 17.9 9.4 4.2 6.8 6.0 6.3 4.8 2.8 15.5 13.4 27.0 30.9 17.5 9.8 4.3 6.8 5.7 6.0 4.5 2.5 15.6 13.2 25.7 29.4 15.9 8.7 4.1 6.1 5.6 6.0 4.9 2.3 15.4 13.7 26.2 28.9 17.2 8.8 4.2 6.1 5.7 6.2 4.6 2.3 14.5 13.2 26.3 29.8 17.1 9.2 4.2 6.5 5.8 6.1 4.7 2.5 15.2 13.5 27.3 31.2 17.9 8.7 4.2 6.4 5.9 6.3 4.9 2.5 17.0 13.4 24.7 28.3 16.8 8.8 3.9 6.2 5.7 5.8 4.3 2.4 15.0 13.3 25.1 29.1 17.9 8.8 4.5 6.6 6.0 6.0 4.7 2.6 15.0 14.2 24.1 29.1 15.8 8.5 4.5 6.4 5.7 6.0 4.6 2.2 15.0 13.5 25.3 29.4 17.1 8.7 4.3 6.4 5.8 6.0 4.6 2.4 15.5 13.6 26.1 30.4 16.5 8.6 4.4 6.1 5.7 6.0 4.6 2.4 15.6 13.2 26.8 29.8 17.2 8.8 4.3 6.5 5.8 6.3 4.6 2.5 14.8 13.2 25.8 30.5 17.3 8.6 4.5 6.2 5.8 6.0 4.5 2.5 15.5 13.6 25.5 29.2 15.6 8.3 3.7 6.8 5.8 6.1 4.9 2.6 14.5 13.2 22.4 25.7 15.5 7.8 3.8 5.8 5.6 5.8 4.2 2.2 13.5 12.3 22.8 26.0 15.8 8.0 3.9 5.8 5.0 5.4 4.1 2.1 14.4 12.2 23.4 25.8 15.3 8.2 3.6 6.1 5.7 5.8 4.4 2.1 13.2 12.0 pcninsulae 27.8 31.4 17.6 10.2 4.8 6.3 5.9 6.2 4.7 2.4 15.3 14.6 23.7 27.1 16.4 8.4 4.3 6.3 5.7 5.7 4.1 1.9 13.3 12.9 6.4 5.7 5.9 4.6 2.0 12.3 spadix 28.0 31.7 15.9 9.5 3.6 6.9 6.5 6.7 5.4 2.7 15.5 14.0 28.1 32.1 16.4 9.7 4.0 6.8 6.2 6.7 4.9 2.0 15.9 15.0 27.0 29.9 15.0 9.2 3.8 6.6 6.1 6.5 5.0 3,0 15.0 13.8 26.1 15.2 8.4 3.3 6.2 5.5 5.9 4.3 2.3 24.9 28.2 14.8 8.5 3.7 6.6 5.9 6.6 4.8 2.9 is'.i i2!8 22.9 27.0 13.8 7.7 3.1 6.6 5.6 5.6 4.4 2.1 13.0 12.3 24.5 26.8 14.5 8.6 3.7 5.8 5 3 5.5 4.5 2.1 13.9 13.2 23.3 25.6 15.2 7.6 3.2 5.9 5.4 5.7 4.1 2.0 14.0 12.2 23.3 26.0 14.7 8.1 3.3 5.7 5 3 5.6 4.3 2.4 13.8 12.8 23.4 26.4 14.6 8.0 3.3 6.0 5.4 5 6 4.3 2.2 13.7 12.6 longicaudi a 25.3 30.8 15.2 8.9 3.4 6.9 6.3 6.5 4.9 2.6 15.0 14.6 31.0 6.9 6.3 6.6 5.0 2.5 13,8 25.7 30.4 i'^.i ■8.7 3.9 6.3 5.7 5.9 4.4 1.9 15^5 13.4 24.8 29.7 15.5 8.7 3.6 6.0 5.4 5.6 4.3 2.4 15.0 13.4 25.0 29.4 15,2 8.3 3.4 6.6 6.2 6.4 4.6 2.0 16.0 13.6 25.2 30.3 15.4 8.6 3.6 6.5 6.0 6.2 4 6 2.3 15.4 13.8 24.0 26.1 14.7 8.3 3.1 5.7 5.2 5.4 4.3 2.0 13.3 12.2 24.1 26.7 14.5 8.8 3.3 5.6 5.4 5.5 4.3 2.3 14.0 12.5 22.5 24.5 13.4 8.2 2.8 5.6 5.0 5.0 4.0 2.0 12.8 11.5 23.2 26.1 15.3 8.0 2.8 6.1 5.4 5.6 4.3 2.4 14.0 12.3 23.3 25.9 14.8 8.3 3.2 6.4 5.9 6.1 4.8 2.4 14.3 12.1 23.4 25.9 14.6 8.3 3.0 5.9 5.4 5 5 4 3 2,2 13.7 12.1 422 University of Kansas Publs., Mus. Nat. Hist, Table of Cranial Measurements — Continued I S a o O o S a ■a O 09 tt> CO ■.a OS o o .a as m T3 03 a _a _o £ O M. V. Z. M. V. Z. Cowan N. M. C. N. M. C. M. C. Z. M. V. Z. 44568 43817 443 2676 2695 9058 62791 U. S. N. M. 186451 A. M. N. H, A. M. N. H. 7441 7440 913 6 M. V. Z. 55211 M. V. Z. 65231 U. S. N. M. 248993 A. M. N. H. average maximum minimum average maximum minimum average maximum minimum average maximum minimum 2490 cf ad. cfad. 9 ad. 9 ad. average maximum \ d' ad. minimum J U. S. N. M. 212423 average maximum minimum average maximum minimum d' ad. 9 ad. U. O. 3709 U. S. N. M. 65930 M. V. Z. 13778 M. V. Z. 13779 (f sad. cf sad. c? ad. cf ad. cf ad. 9 ad. 9 ad. cf ad. cf ad. 9 ad. cf ad. cf ad. cf ad. 9 ad. 10 10 14 6 9 ad. Barkerville reg., B. C 48. 8 Isaacs Lake, British Columbia 48.8 Quesnel, British Columbia 46.6 Lillooet, British Columbia 47.5 Lillooet, British Columbia 45.0 Source Kettle River, B. C 41.7 Beaverfoot Range, B. C 42. 0 Custer, South Dakota 40.9 Hill City, South Dakota 41.0 Black Hills, South Dakota 37 . 6 near Parks, Arizona 40 . 4 Alpine, Arizona 39.6 Kaibab Plat., Arizona 40. 4 S. F. Forest, Arizona 35.5 25 Sierra Nevada, California. . . S and SW Colorado. California . Colorado . NE Oregon . 43.6 46.1 40.6 43.7 44.6 41.6 38.2 39.5 36.7 38.5 39.7 36.1 40.5 41.8 39.3 Vale, Oregon 37. 4 22 Mt. Adams, Washmgton . 11 Mt. Adams, Washington . 43.7 47.7 40.0 37.7 39.0 37.1 cf ad. Mt. Ashland, Oregon 45.8 cf ad. Siskiyou, Oregon 42.6 cf ad. Jackson Lake, California 45.5 cf ad. Jackson Lake, California 43.8 av. 4 44 . 4 19.1 19.5 17.7 18.5 17.5 16.4 16.4 15.0 Mustela frenata 17.2 13.0 17.7 15.8 12.0 17.3 14.5 10.1 15.9 14.3 10.3 15.9 15.9 11.8 15.9 12.8 13.5 10.5 10.2 14.8 15.1 Mustela frenata 13.8 11.0 14.1 15.6 13.7 11.0 13.9 14.1 12.2 9.1 13.2 15.5 15.1 15.6 Mustela frenata 12.5 .... 14.2 12.2 9.7 13.7 12.9 9.8 14.5 13.8 10.8 8.6 12.9 Mustela frenata 16.6 13.7 10.7 15.1 17.6 14.9 12.0 16.2 15.2 12.5 9.9 14.0 16.5 13.9 10.6 15.0 17.3 14.8 11.1 16.4 16.0 12.8 9.9 13.8 14.7 11.8 9.2 13.4 15.1 12.4 9.9 14.2 13.9 11.0 8.6 12.6 14.8 12.2 9.3 13.4 15.4 13.1 10.2 14.4 14.0 11.1 8.5 12.5 Mustela frenata 15.2 12.3 9.6 13.7 16.4 12.7 10.0 14.4 14.4 11.9 9.2 13.1 9.2 .... Mustela frenata 16.7 13.7 10.5 15.5 18.0 15.4 12.0 16.5 15.6 12.5 9.0 14.5 14.3 11.5 8.8 12.8 14.9 12.0 9.3 13.2 13.3 10.8 8.2 12.1 Mustela frenata 17.2 14.1 11.1 15.8 15.9 14.0 10.9 14.4 18.0 13.2 10.0 16.2 16.9 12.7 9.9 15.3 17.0 13.5 10.2 15.4 M. V. Z. 52144 9 ad. S. Fork Mt., California. . .38.2 14.7 10.8 8.1 13.5 Cranial Measurements 423 I ■ T>Tnpanic Bulla S ■S — ^^ •4^ 1 to o 1-Z tJ a N /■ ^ ^ r ' N ^ CS •:; o 0^ X> 3 .M w -^ Ef-s JS^ i o S 2.2_ T3 "S s J3 a j2 Q "3 13 § Depth 0 Ant. I basioc Depth 0 poster of Ms oribasus 28.5 32.2 17.4 9.8 3.2 6.8 6.2 6.4 5.3 2.8 15.5 14.6 28.4 31.7 15.9 9.7 4.0 6.9 6.3 6.4 5.3 2.9 15.5 13.7 26.4 30.7 15.8 9.2 3.9 6.2 5.8 6.1 4.6 2.5 15.0 13.1 25.5 30.1 15.2 9.1 3.8 6.4 5.9 6.3 4.6 2.5 15.0 13.1 25.6 31.0 15.2 8.9 3.0 6.2 5.6 6.1 4.7 2.9 15.0 13.4 24.9 26.7 14.3 8.2 2.8 6.1 5.5 5.7 4.5 2.5 14.0 12.6 24.4 27.0 14.3 8.1 3.1 5.9 5.5 5.6 4.3 2.2 14.0 12.3 alien! 22.0 27.7 13.7 7.9 3.0 5.7 4.9 5.3 3.9 2.5 14.3 12.3 23.3 25.7 13.6 8.3 2.3 5.1 4.8 5.2 4.2 1.9 13.7 12.0 22.3 23.1 13.8 7.3 3.1 5.5 4.9 5.0 3.6 1.8 12.2 10.5 arizonensis 23.3 26.0 14.3 8.9 3.1 5.7 5.2 5.5 4.0 2.2 13.5 12.1 22.4 25.6 13.7 8.3 3.2 5.4 4.9 5.1 4.0 2.1 14.0 11.4 22.9 26.3 14.1 8.4 3.2 5.8 5.5 5.5 4.0 1.7 14.8 12.0 19.9 21.7 13.3 7.7 2.9 4.8 4.6 4.7 3.6 1.6 12.3 10.3 nevadensis 23.9 28.0 15.0 8.4 3.4 6.9 5.4 5.6 4.3 2.2 14.4 12.5 26.1 31.4 15.9 9.0 4.0 6.4 5.8 6.1 4.8 2.7 15.2 14.8 22.1 25.0 14.4 7.8 2.9 5.5 4.9 5.1 3.9 1.8 13.7 11.6 24.2 27.9 15.1 8.6 3.5 6.1 5.5 5.7 4.5 2.1 14.5 12.6 25.3 29.3 15.8 9.2 4.0 6.8 5.9 6.0 4.8 2.6 15.6 13.0 23.1 26.5 14.4 8.2 3.0 5.7 5.2 5.3 4.0 1.9 13.9 12.1 20.9 23.1 13.4 7.3 2.9 5.2 4.8 5.0 3.8 1.7 12.3 11.0 21.8 23.4 14.1 7.9 3.2 5.6 5.2 5.4 4.1 2.0 13.0 11.8 20.1 22.4 12.7 6.8 . 2.6 4.9 4.5 4.7 3.6 1.5 11.4 10.0 20.6 23.1 13.4 7.6 2.8 5.4 4.9 5.1 3.9 1.9 12.9 11.1 22.1 24.6 13.9 8.1 3.2 5.6 5.3 5.4 4.1 2.3 13.8 11.8 19.8 22.0 12.9 7.0 2.5 5.1 4.3 4.6 3.4 1.7 12.0 10.6 effera 22.1 25.6 14.1 7.8 3.3 5.5 5.0 5.3 3.9 2.0 13.5 11.8 23.3 27.3 15.0 8.4 3.5 5.9 5.3 5.8 4.1 2.6 14.3 12.2 20.5 25.0 12.3 7.2 3.2 5.0 4.7 5.0 3.6 1.6 12.6 11.4 19.5 22.0 13.1 6.9 3.0 5.4 4.9 5.0 3.7 1.8 12.3 Washington! 23.4 27.0 14.6 8.0 3.1 5.9 5.4 5.6 4.3 2.2 14.4 12.6 26.4 29.6 15.8 8.7 3.4 6.5 6.0 0.1 4.8 2.6 15.8 13.7 22.1 24.6 13.5 7.6 2.7 5.4 4.9 5.0 4.0 1.7 13.1 11.5 20.2 22.5 12.9 7.2 2.8 5.1 4.7 4.8 3.9 1.9 12.2 10.4 21.1 24.5 13.6 7.7 3.0 5.6 4.9 5.0 4.2 2.1 13.0 11.0 19.4 21.3 12.3 6.8 2.3 4.7 4.4 4.2 3.4 1.7 11.2 9.8 Baturata 26.0 27.9 15.1 8.9 3.9 6.1 5.5 5.9 4.3 2.3 14.2 12.6 24.5 27.7 14.7 8.5 4.0 5.7 5.0 5.5 4.1 1.9 14.8 12.4 24.7 27.4 14.6 8.7 3.2 6.3 5.9 6.2 4.6 2.5 14.6 12.8 24.2 26.9 15.1 8.6 3.3 5.6 5.2 5.5 4.0 2.2 14.2 12.3 24.9 27.5 14.9 8.7 3.6 5.9 5.4 5.8 4.3 2.2 14.5 12.5 19.8 21.8 12.4 7.2 2.7 5.0 4.6 4.9 3.7 1.8 12.2 10.4 424 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial Measurements — Continued § ••3 I a 3 C 3 O c 03 ■a o o 5o li o 5 O A C o A -*^ a) v n u O —I O average maximum minimum Walker Walker Walker Walker 6" ad. 392 185 89 45 M. V. Z. 11747 C. A. C. 3907 K. M. N. H. 9595 U. S. N. M. 32019 M. V. Z. V.O. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. M. V. Z. F. M. N. H. M. C. Z. M. C. Z. M. V. Z. M. V. Z. U. S. N. M. F. M. N. H. U. S. N. M. Walker A. N. S. P. Wisconsin U. Snyder M. V. Z. Stanford U. F. M. N. H. Stanford U. Stanford U. F. M. N. H. Stanford U. M. V. Z. M. V. Z. IT. S. N. M. 34325 1413 19720 19722 19718 19714 19715 19716 9598 8632 5459 40302 19723 135010 9597 91764 1440 11863 4232 2626 79640 863 6559 1651 487 7031 236 5851 30327 43574 Mustela frenata f 45.8 17.5 14.2 11.2 15.9 10 Tillamook Co.. Oregon { 48.0 18.9 15.0 12.0 16.8 I 42.4 16.1 13.2 10.0 14.8 9 ad. Blaine, Oregon 39.7 15.1 12.8 10.1 13.7 9 ad. Blaine, Oregon 37.8 13.9 11.9 9.5 13.4 $ ad. Blaine, Oregon 38.3 14.3 11.3 9.5 12.7 ■ 9 ad. Tillamook, Oregon 37.8 14.1 11.0 8.6 12.5 t Mustela frenata cf ad. Eureka. California 44.0 16.8 13.2 10.7 15.3 cf ad. Requa, California 41.8 16.0 13.5 10.9 14.0 cf ad. Gold Beach, Oregon 43.0 16.9 13.1 10.5 15.5 d" sad. Grant.s Pass, Oregon 42.9 16.3 13.7 10.5 15.7 9 ad. Carlotta, California 37.8 14.7 10.8 8.8 13.0 9 ad. 13 mi. S Grants Pass, Oregon . 39.4 14.7 12.0 9.7 13.9 Mustela frenata , cf ad. Point Arena, California 49.0 18.5 15.2 11.7 16.8 cf ad. Point Arena, California 48.5 18.9 14.4 10.5 17.2 cf ad. Gualala, California 45.7 17.2 13.7 10.5 15.5 av. 3 47.7 18.2 14.4 10.9 16.5 cf ad. 6 mi. W Inverness, California. . 48.2 18.4 13.9 11.3 17.2 cf ad. 6 mi. W Inverness, California. . 48.2 18.9 14.5 11.4 16.6 cf ad. 5 mi. W Inverness, California. . 46.5 17.5 13.8 II. 0 15.9 av. 3 47.6 18.3 14.1 11.2 16.6 c? ad. Nicasio, California 46.5 18.5 14.0 10.5 15.4 cf ad. Nicasio, California 44.2 17.8 14.0 11.4 16.9 o' ad. Point Reves. California 39.7 15.5 13.3 9.0 13.4 d' ad. 4 mi. N Vallejo, California 45.8 17.7 13.0 10.7 15.9 av. 4 44.1 17.4 13.6 10.4 15.4 9 ad. Point Arena, California 42.3 16.0 12.3 12.2 14.0 9 ad. Point Reyes, California 38.7 15.3 11.0 8.7 14.5 9 ad. Point Reyes, California 39.5 15.2 11.1 8.2 13.1 9 ad. Point Reyes, California 38.7 15.4 12.0 9.7 12.7 Mustela frenata cf ad. 5 mi. W Fresno, California . 43.9 16.5 13.8 10.4 14.1 cf ad. Fresno, California 43.4 16.8 12.9 10.0 14.8 cf ad. Selma, California 43.7 16.2 13.3 9.9 14.5 9 ad. Selma. California 39.4 15.0 12.0 8.9 13.2 9 ad. Tegner School, California 43.4 16.5 12.6 9.3 15.1 Mustela frenata cf ad. Palo Alto, California 48.1 18.7 15.0 11.2 17.5 cf ad. Palo Alto. California 48.0 18.7 13.9 11.0 16.2 d' ad. Menlo Park, California 47.1 17.8 13.3 9.8 14.9 d' ad. P.ilo Alto, California 46.5 18.3 13.4 11.0 16.1 d' ad. Palo Alto, California 46.1 18.1 14.5 10.7 15.1 d' ad. Menlo Park. Cahfornia 46.1 17.8 13.5 10.4 15.5 av. 6 47.0 18.2 13.9 10.7 15.9 9 ad. Hayward, California 40.7 15.3 11.8 8.8 13.0 9 ad. Palo Alto, California 41.2 16.2 11.1 8.6 14.3 9 ad. Morro, California 42.2 16.1 12.2 8.9 14.2 Cranial Measurements 425 ■s Tympanic Bulla i ♦*«^ -J -•A 1 kull a •gin a ital -■s ' .0 •a S 3 i tS3 it a ■s 1 u CQ a Q 1 Length ml g ■3 CQ 0 Depth of S Ant. mai basioccip \ Depth of S ' posterior of Msl altifrontal 8 25.1 29.2 15.6 8.5 3.5 6.1 5.5 5.7 4.5 2.5 15.4 13.6 26.0 31.6 16.5 9.0 3.6 6.5 6.0 6.0 4.9 2.8 16.2 14.0 23.9 26.0 15.0 7.7 3.2 5.6 5.0 5.2 4.0 2.1 14.5 12.4 21.8 24.0 13.8 7.7 5.2 5.0 5.0 3.7 1.9 13.5 11.8 20.8 22.7 13.4 7.3 '2^8 4.9 4.5 4.7 3.2 1.5 12.6 10.7 19.8 22.7 13.2 7.0 3.1 4.4 4.7 3.8 1.9 13.4 11.3 20.1 23.2 13.0 7.3 4^9 4.7 4.8 3.8 2.0 13.5 11.0 oregonensis 24.3 28.0 14.6 8.3 3.4 6.0 5.4 5.8 4.3 2.2 13.6 12.7 22.9 26.6 14.2 7.6 3.1 5.8 5.3 5.4 4.0 2.2 13.4 12.3 23.4 26.2 15.1 8.2 3.3 5.8 5.2 5.5 4.4 2.0 14.7 12.5 23.2 26.5 15.2 8.8 4.0 5.5 4.8 5.2 4.3 2.1 14.0 12.5 20.7 23.2 13.4 7.5 2.2 5.3 5.0 5.5 4.0 1.8 13.7 11.0 21.5 24.0 14.2 7.6 2.7 4.8 4.4 4.7 3.7 1.9 12.6 10.8 munda 27.5 33.2 16.2 9.3 3.6 6.5 5.6 6.0 4.8 2.6 14.2 14.2 26.8 32.+ 16.0 8.5 3.8 6.5 5.6 6.2 5.0 2.9 14.7 13.8 26.4 31.7 15.5 8.5 3.1 6.5 5.7 5.8 4.9 2.3 13.4 14.2 26.9 32.7 15.9 8.8 3.5 6.5 5.7 6.0 4.9 2.6 14.1 14.1 26.2 30.5 15.7 8.5 3.5 6.5 5.6 6.0 4.8 2.6 14.2 14.2 26.5 30.0 16.3 8.7 3.5 6.3 5.6 6.2 5.0 2.9 14.7 13.8 25.2 30.3 15.4 7.9 3.5 6.1 5.7 5.8 4.9 2.3 13.4 14.2 26.0 30.3 15.8 8.4 3.5 6.3 5.7 6.0 4.9 2.6 14.1 14.1 25.2 30.5 15.0 8.8 3.5 6.9 5.9 6.3 5.2 2.7 15.2 13.0 24.4 27.6 15.7 8.2 3.7 6.4 5.7 5.9 5.0 2.6 12.6 13.7 23.1 26.5 13.7 7.7 3.2 6.1 5.1 5.5 4.4 2.5 13.6 12.1 24.4 26.9 15.1 8.0 2.8 6.3 5.7 6.2 4.8 2.3 14.5 13.0 24.3 27.9 14.9 8.2 3.3 6.4 5.6 6.0 4.9 2.5 14.0 13.0 23.6 25.5 14.5 8.3 2.9 5.0 5.2 4.0 1.9 13.4 11.3 21.0 23.7 13.0 7.5 3.0 '5.1 4.9 5.4 3.9 2.3 12.9 10.8 20.2 22.8 12.7 6.8 2.7 5.3 4.9 5.0 3.9 1.9 14.7 10.8 21.7 24.7 13.7 7.4 2.9 5.5 5.0 5.3 4.2 2.0 13.2 11.7 xanthogenys 23.8 28.5 14.7 7.9 3.2 5.7 5.2 5.3 4.2 2.1 14.8 12.6 24.0 27.5 14.5 7.5 5.8 5.3 5.5 4.4 2.0 13.7 12.7 23.7 27.1 15.2 8.5 3^7 5.8 5.3 5.4 4.2 2.2 13.8 12.3 21.3 24.3 13.7 7.2 5.3 4.0 13.0 U.4 22.8 24.9 15.0 7.5 3.6 5.6 5^5 5^9 4.5 2^0 13.3 12.2 nigriauria 32.9 15.5 6.5 6.4 6.7 5.2 2.4 14.3 27! 6 29.6 15.7 '8.7 '3.4 6.2 5.8 6.0 4.8 2.7 i4;6 14.1 25.6 30.0 15.0 8.0 3.3 6.5 6.0 6.5 4.6 2.4 14.6 13.8 25.2 31.1 15.1 8.0 2.9 6.3 5.7 6.0 4.9 2.5 14.5 14.3 26.0 29.5 14.9 8.3 2.4 6.3 6.0 6.1 4.6 2.6 15.0 13.2 29.6 15.6 8.5 6.0 5.6 5.7 4.8 2.5 14.0 26!6 30.5 15.3 8.3 3;6 6.3 5.9 6.2 4.8 2.5 iiis 14.0 21.8 24.9 14.2 7.9 3.0 5.2 4.9 4.9 4.1 1.8 12.4 11.0 21.7 24.8 13.4 7.8 3.0 5.3 5.0 5.4 4.1 2.1 12.6 11.0 22.7 24.3 15.1 8.1 2.9 5.5 5.4 5.6 4.2 2.0 13.2 11.4 426 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial Measurements — Continued a o o .a B 3 d 3 "3 •♦J d o a ■73 a OS CQ o £ a 3 •4^ -a k. OJ ! -fi^ o s ^ ^ 11 o d 0 [mi £ i a S a m I-! m O M. V. Z. U. S. N. M. U. S. N. M. M. V. Z. Stephens S. D. M. Stephens Stephens S. D. M. S. D. M. C. A. S. C. A. S. M. V. Z. U. S. N. M. C. A. S. C. A. S. M. V. Z. .^257 52701 52702 3258 20 7098 22 19 6748 7194 335 337 16668 137935 336 338 25907 d' ad. 2 mi. N Independence, Calif .. . 44.7 17.3 A. M. N. H. 14821 cf ad. Kerr Co., Texas 54.0 19.2 M. C. Z. 15476 cf yg. Kerr Co., Texas 53.3 18.9 Baylor U. 2017 cf sad. 5 mi. N Waco, Texas 52.0 18.3 Kansas U. 1485 cf ad. Liberal, Kansas 50.5 17.9 U. S. N. M. 131582 c? ad. Berino, New Mexico 47.7 17.5 U. S. N. M. 36482 9 ad. Tombstone, Arizona 45.5 16.5 U. S. N. M. 230973 9 ad. Willcox, Arizona 42.5 15. 1 U. S. N. M. 225629 9 ad. Albuquerque, New Mexico 40.8 15.0 M C.Z. 240 cf ad. Brownsville, Texas 49.4 17.4 A. N. S. P. 724 d" ad. Brownsville, Texas 50. 1 17.9 U S. N. M. 58684 cf ad. Bownsville, Texas 48.2 17.3 U S. N. M. 63857 cf ad. Brownsville, Texas 48.6 18.0 U. S. N. M. 44976 c? ad. Brownsville, Texas 50.9 18.0 av. 5 49.4 17.7 A. M. N. H. 24405 9 ad. Brownsville, Texas 47.3 16.2 U. S. N. M. 58685 9 sad. Brownsville, Texas 41.3 15.0 U. S. N. M. Illll 9 sad. Brownsville, Texas 42.5 15.4 av. 3 43. 7 15.5 B. Z. M. 991 9 ad. Mexico 15.5 B. Z. M. 992 9 ad. M6xico 13.9 M C.Z. 20841 cf ad. Miquihana, Nuevo Leon 50.2 18.0 U. S. N. M. 50826 cf ad. Tlalpam, D. F 51.3 18.3 U. S. N. M. 125972 cf ad. Los Reyes, Michoacan 51.2 17.5 U. S. N. M. IfrfJ cf ad. Patzcuaro, Michoacan 18.7 A. M. N. H. 26153 9 ad. Artenkiki, Jalisco 44.5 16.0 12.8 9.7 15.5 12.9 9.9 14.0 12.4 9.4 13.3 Mustela frenata 14.9 11.6 15.4 15.5 12.3 15.5 14.2 11.0 15.3 13.7 11.1 16.5 16.9 14.6 11.5 15.9 12.5 10.0 14.6 11.8 9.5 12.8 15.3 15.1 12.1 12.1 14.3 12.7 Mustela frenata cf ad. San Diego. California 44.4 17.8 13.4 10.2 15.0 cfad. El Vido, California 43.7 16.9 12.7 9.9 14.3 cf ad. El Cajon, California 43.2 16.7 13.9 10.0 15.0 cfad. San Diego, California 42.5 16.6 14.1 12.1 14.7 cfad. San Jacinto Plain, California. . . 41.9 16.9 13.1 10.6 14.0 cfad. Jamacha, California 47.0 18.5 15.6 11.3 16.8 av. 6 43.8 17.2 13.8 10.7 15.0 9 ad. Santa Ysabe!. California 40.1 15.5 11.7 9.3 13.1 9 ad. Ballena. California 40.0 15.0 12.3 9.3 13.5 9 ad. Santa Ysabel. California 42.0 16.1 12.9 9.4 14.2 9 ad. Jamacha, Cahfornia 39.8 15.0 11.8 9.1 13.7 av. 4 40.5 15.4 12.2 9.3 13.6 Mustela frenata cfad. Buttonwillow, California 48.6 18.8 15.2 11.6 17.2 r cfad. Buttonwillow, California 48.6 18.8 15.0 11.8 17.2 cfad. Buttonwillow, California 48.1 18.8 14.8 12.0 17.1 cfad. Buttonwillow, California 47.2 18.2 14.0 10.3 16.0 cfad. Buttonwillow, California 47.0 18.3 14.5 11.5 16.5 cfad. Buttonwillow, California 46.0 17.4 14.7 11.3 16.1 av. 6 47.6 18.4 14.7 11.4 16.7 Mustela frenata 13.3 10.8 15.9 Mustela frenata 16.0 12.6 17.2 16.3 12.8 18.3 16.5 13.5 16.7 Mustela frenata 15.4 12.4 16.4 15.0 11.1 15.9 12.5 10.0 14.3 12.3 9.8 13.9 12.2 13.8 10.7 8.9 12.9 16.3 17.5 Mustela frenata 15.0 12.0 16.0 10.0 16.8 14.4 Cranial Measurements 427 ■3 Tympanic '. BuUa s 1 1 i -♦J »„ i2 ■^ « ■s 1 . , kuUa ■gin 01 ital rt-d t ^ ^~ \ / — > "5° Jo ■■s ji s J3 0 C 01 5 3 >> ■3 a 2 j3 0 J3 to J ■3 1 ■3 -a 2 m a Q Depth poste of M latirostra 23.5 27.6 15.6 8.4 3.9 6.0 5.5 5.5 4.3 2.2 14.0 12.8 24.0 14.7 8.7 4.0 5.7 5.6 5.6 4.2 2.0 14.2 13.4 24.0 27^2 15.1 8.2 4.2 5.8 5.3 5.5 4.2 2.5 14.0 12.5 24.0 28.7 15.1 8.1 3.9 5.9 5.6 5.5 4.4 2.5 14.3 13.5 24.0 28.2 14.7 8.0 3.2 6.0 5.5 5.7 4.2 2.1 14.0 12.9 24.8 29.2 16.7 8.9 3.8 6.0 5.8 6.2 4.8 2.2 14.0 13.2 24.1 28.2 15.3 8.4 3.8 5.9 5.6 5.7 4.4 2.3 14.1 13.1 21.9 24.5 14.2 8.0 3.0 5.3 4.8 5.1 4.1 2.0 14.0 11.3 21.6 23.5 13.7 8.1 3.2 5.1 4.9 5.0 4.0 2.1 12.0 11.2 22.0 24.5 14.0 7.8 3.0 5.8 5.3 5.7 4.5 2.0 13.4 11.5 20.2 23.7 13.7 7.0 2.9 5.2 4.8 4.9 3.9 1.9 12.6 11.5 21.4 24.1 13.9 7.7 3.0 5.4 5.0 5.2 4.1 2.0 13.0 11.4 pulchra 27.4 31.4 16.7 9.1 3.6 6.6 5.9 6.4 5.1 2.7 16.2 13.7 27.1 32.7 16.6 9.0 3.5 6.6 6.3 6.3 5.0 2.4 16.5 14.3 27.7 31.2 16.4 9.2 3.6 6.4 5.7 5.9 5.1 2.8 15.2 14.0 26.1 29.5 15.6 8.5 3.3 6.0 5.6 5.9 4.1 2.0 15.0 12.9 27.0 29.5 15.5 9.1 3.2 6.3 5.6 5.9 5.0 2.3 15.0 13.0 26.3 32.6 15.9 8.9 3.7 6.1 5.7 5.8 4.6 2.2 15.0 13.3 26.9 31.1 16.1 9.0 3.5 6.3 5.8 6.0 4.8 2.4 15.5 13.5 Inyoensis 25.3 29.5 15.9 9.0 3.7 6.0 5.7 5.9 4.6 2.4 15.0 13.2 texensis 28.6 35.1 17.5 8.0 3.5 6.9 6.5 6.7 5.0 2.5 16.7 15.1 28.2 34.8 18.0 8.3 4.0 6.4 6.3 6.5 4.8 2.4 16.0 15,0 29.2 18.0 9.1 6.6 6.3 6.4 4.9 2.6 16.3 15.6 neomezicana 28.3 35.0 17.0 9.0 4.0 6.3 5.9 6.0 4.9 2.4 15.5 14.3 26.3 31.2 16.5 7.9 3.4 6.4 5.7 6.3 4.7 2.5 15.8 13.5 22.3 26.6 15.2 7.5 3.3 5.9 5.6 5.9 4.5 2.2 13.8 12.0 22.6 26.5 14.5 7.0 3.1 5.5 5.3 5.6 4.1 1.8 14.0 12.0 21.6 24.5 14.5 7.5 3.0 5.2 4.7 5.0 3.9 1.8 13.0 11.0 frenata 25.9 33.0 16.7 7.9 4.3 5.9 5.9 6.1 4.5 2.1 15.0 13.8 27.0 32.2 16.5 8.5 4.1 6.5 6.2 6.3 4.8 2.7 16.0 14.0 27.2 31.0 16.6 8.3 4.2 6.3 5.9 6.1 4.8 2.7 15.5 13.5 26.0 31.0 16.0 8.2 4,8 6.5 5.7 6.1 4.8 2.6 15.0 13.6 26.9 16.6 7.9 3.4 6.5 5.6 6.1 4.7 2.3 16.0 13.4 26.6 31^8 16.3 8.2 4.2 6.4 5.9 6.2 4.7 2.5 15.5 13.7 5.9 5.4 5.4 4.1 2.1 12.2 22^7 27!6 i4;6 '&.9 '3.3 5.5 5.2 5.4 4.1 2.0 i4;6 11.7 23.8 26.7 14.3 7.5 3.2 5.9 5.5 5.7 4.2 2.0 13.5 11.8 23.3 26.9 14.2 7.2 3.3 5.8 5.4 5.5 4.1 2,0 13.8 11.9 22.8 27.0 13.7 5.7 5.2 5.5 4.4 2.2 12.7 21.1 13.0 6^6 '2^9 4.9 4.5 4.7 3.7 1.7 12.3 10.8 28.0 32.0 16.9 8.8 3.3 4.6 2.2 15.0 27.7 33.5 16.3 8.4 3.5 h'.i "5.9 'ga 4.7 2.7 15.3 li.'t leucoparia 28.3 32.9 16.0 7.7 3.5 5.9 5.7 5.7 4.3 2.2 15.5 14.0 6.8 6.5 6.8 5.0 2.6 14.3 22.4 26.3 15.0 7.0 3.2 5.9 5.5 6.0 4.5 2.1 14.0 11.9 428 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial Measurements — Continued o O « a a o s O ■V a eS M o ••a o o P3 o o «^ o J3 -*^ T3 0) o .a •o 03 pa J2 .2 5 o U. S. N. M. 68197 c? ad. U. S. N. M. 54278 9 ad. U. S. N. M. 133253 U. S. N. M. 133254 U. S. N. M. 77519 V. M. N. H. 15953 Dickey 12523 F. M. N. H. 14063 U. S. N. M. 132528 U. S. N. M. 54994 A. M. N. H. \nn M. C. Z. G514 U. S. N. M. 54993 U. S. N. M. 1060 M. C. Z. 2605 F. M N. H. 14050 U. S. N. M. 100041 U. S. N. M. 132997 U. S. N. M. 132996 U. S. N. M. 218036 U. S. N. M. 65422 A. M. N. H. 30754 A. M. N. H. 28331 A. M. N. H. 29280 U. S. N. M. B. M. 11408 3216 U. S. N. M. N. H. R. S. 1-138 B. Z. M. A 59. 13 M. C. Z. A. M. N. H. M. C. Z. 10112 18848 10113 Cerro San Felipe, Oaxaca. Type specimen 49.2 17.3 43.5 15.5 U. S. N. M. 170970 A. N. S. P. 18434 d' ad. 20 mi. SE Teopisca, Chiapas. . . 50.4 18.0 cf ad. 20 mi. SE Teopisca, Chiapas... 49.6 17.5 cf ad. Pinabete, Chiapas 50. 7 18.3 cf ad. near Tecpam, Guatamala 50.0 17.0 cf ad. Los Esesmiles, Salv 51.3 17.5 av. 5 50.4 17.7 cf ad. Achotal, Veracruz 64. 1 19.2 9 ad. P6rez, Veracruz 43.5 15.3 cf ad. Jico, Veracruz 47.8 17.2 (f sad. Jalapa, Veracruz 45.5 16.8 c?" sad. Orizaba, Veracruz +46.0 16.4 9 ad Jico, Veracruz 36.0 13.0 9 ad. Mexico 39.0 14.0 9 ye. Jalpa, Veracruz 38.7 13.8 9 yg. Xuchil, Veracruz 39. 0 14. 2 (f sad. Teapa, Tabasco 46. 1 17.0 cf sad. San Vicente, Chiapas 45.3 16.7 cf sad. San Cristobal, Chiapas 16.9 av. 3 45.7 16.9 9 sad. State of Chiapas 40.0 14.6 9 sad. Cateniaco, Veracruz 40.4 14.5 c? sad. Matagalpa, Nicaragua 44.8 17.2 cf ad. Matagalpa, Nicaragua 44.8 16.7 cf sad. San Rafel Del Norte 45.5 17.2 cf ad. Costa Rica *49.0 18.3 cf ad. Vic. San Jos6, Costa Rica 49.3 18.6 cf yg. San Jos^, Costa Rica 48.2 18.0 cf ad. Azahar Cartago, Costa Rica .. . 47.8 17.4 9 sad. I razu, 3000M., Costa Rica 38.8 14.4 cf ad. Boquete, Panama 48.3 17.1 cf ad. Boquete, Panama 44.5 16.3 cf ad. Bocjuete, Panama 42.8 15.8 av. 3 45.2 16.4 9 sad. near Gatun, Panama 41.3 15.3 9 ad. Siola, Panama 39.3 14.2 Mustela frenata 13.9 11.7 15.9 12.3 10.3 14.0 MuBtela frenata 15.0 12.5 15.9 13.8 10.9 16.2 13.5 i2;6 ieii 14.5 11.7 16.2 14.2 11.8 16.1 Mustela frenata 15.6 12.9 17.8 12.5 10.2 14.5 Mustela frenata 13.7 10.7 15.8 13.7 13.2 11.4 16.0 15.5 11.0 11.0 10.3 11.6 9.2 9.6 8.3 9.3 12.1 12;6 13.0 Mustela frenata 14.4 11.1 15.3 13.9 12.0 15.5 13.3 10.6 16.0 13.9 11.2 15.6 11.6 11.5 9.4 9.1 13.5 13.2 Mustela frenata 12.8 10.5 15.0 13.6 11.5 15.2 13.5 11.0 15.5 Mustela frenata 15.2 9.4 *18.0 15.0 12.7 16.9 14.0 14.9 11.7 11.7 16.0 15.6 12.5 10.1 14.2 Mustela frenata 15.0 12.5 15.7 14.0 12.0 15.0 14.0 12.0 15.6 14.3 12.2 15.4 12.5 10.4 14.3 11.3 9.4 12.5 Cranial Measurements 429 J3 ■3 1 a o u >, 6Q Tyi mpanic Bulla E a « £ i Depth of Skull at Ant. margin of basioccipital -J "2 'o J3 n "3 1 5 1 t-, a cog perotae 25.0 29.2 15.5 6.8 2.5 6.1 5.3 5.8 4.1 2.1 15.5 13.3 23.2 25.5 15.0 7.0 2.0 5.7 5.2 5.7 4.1 2.1 12.0 13.7 goldmani 26.5 26.4 26.5 26.2 26.7 26.5 32.3 31.8 31.8 31.5 30.6 31.6 15.6 15.8 15.3 15.5 15.8 15.6 7.1 7.4 7.5 7.0 7.7 7.3 2.9 3.0 2.9 's.i 3.0 6.4 6.1 6.6 5.9 6.3 6.3 5.9 5.5 5.8 5.6 5.5 5.7 6.2 5.8 6.1 6.0 5.8 6.0 5.0 4.5 5.0 4.5 4.5 4.7 2.5 2.5 2.4 2.2 2.3 2.4 16.5 15.0 16.0 16.5 16.0 13.7 13.7 ii.'i 13.8 macrophonius 28.5 33.6 16.8 7.6 2.9 7.1 6.4 6.8 5.2 2.9 16.8 15.3 23.1 26.5 15.0 6.5 2.6 5.5 5.2 5.7 4.1 1.9 14.2 12.2 tropicalis 24.5 28.2 15.5 6.3 2.9 6.2 5.6 5.9 4.5 2.2 15.5 13.7 24.0 30.0 15.4 15.0 6.7 8.0 3.0 6.4 6.3 5.7 5.7 5.9 6.0 4.7 5.0 2.2 2.5 15.2 16.5 13.6 14.5 19.8 21.0 19.4 20.5 22.6 22.5 23^5 12.4 1217 13.5 6.1 5.8 6.1 2.5 5.0 4.9 4.9 5.3 4.5 4.6 4.5 4.9 4.7 4.9 4.6 5.3 4.0 3.9 3.5 4.4 1.7 1.9 1.5 2.0 12.5 is. 7 15.3 11.6 11.5 10.9 11.5 pcrda 24.4 24.0 24.2 28.5 27.4 28;6 14.9 14.5 14^7 6.6 6.7 6;7 2.2 2.8 '2;5 6.4 5.5 6.4 6.1 5.5 5.1 5.5 5.4 6.0 5.4 6.0 5.8 4.6 4.2 4.5 4.4 2.5 2.2 2.5 2.4 16.4 15.8 ieii 13.8 13.0 13.0 13.3 20.2 21.2 23.2 23.0 13.0 13.7 5.6 6.5 2.1 2.1 5.2 4.8 4.8 4.7 5.0 5.2 3.7 3.8 1.7 1.9 13.2 14.2 11.4 10.8 nicaraguae 23.4 25.5 22.7 26.8 23.5 27.4 14.0 15.2 14.2 6.5 6.7 6.8 2.9 6.3 5.8 6.3 6.0 5.1 6.0 6.3 5.5 6.3 4.6 4.4 4.7 2.5 2.4 2.5 15.5 14.7 15.5 13.4 13.4 13.1 rostaricensis 26.0 30.5 25.9 31.3 15.0 15.3 7.5 7.3 '2^9 6.5 6.6 5.9 6.0 6.4 6.5 5.0 4.9 2.6 2.5 16.7 16.8 14.2 15.1 24.8 24.3 29.0 ♦30.1 15.0 15.1 7.2 7.5 2.7 7.0 6.6 6.0 5.9 6.5 6.2 4.8 4.8 2.9 2.5 16.5 14.0 20.0 23.6 12.8 6.3 2.9 4.9 4.7 5.0 3.7 1.8 13.7 11.5 panamensis 25.1 28.3 22.5 29.1 27.1 23.8 28.2 14.1 14.4 13.6 14.0 7.0 6.1 5.5 6.2 2.7 2.7 2;7 6.3 6.2 6.1 6.2 5.7 5.7 5.4 5.6 5.9 6.0 5.7 5.9 4.5 4.5 4.3 4.4 2.5 2.3 2.2 2.3 16.7 16.0 15.8 16.3 14.0 13.5 13.5 13.7 23.0 20.1 26.8 22.5 12.5 13.0 6.1 6.0 5.7 5.0 5.3 4.6 5.5 5.0 4.1 3.7 2.1 2.0 15.5 13.2 12.5 10.8 430 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial Measxirements — Continued I a a 3 a eg M V GQ O M a; c8^ n I B E o t3 03 £ a C9 a S m ^ A. M. N. H. U. S. N. M. U. S N. M. A. M. N. H. A. M. N. H. U. S. N. M. U. S. N. M. U. S. N. M. IT. S. N. M. U. S. N. M. A. M. N. H. A. M. N. H. A. M. N. H. A. M. N. H. 33154 137517 172959 24309 33155 123341 137516 143667 143666 143665 24308 24311 21343 24310 U. S. N. M. 241314 U. S. N. M. 239946 A. M. N. H. 35805 U. S. N. M. 241313 U. S. N. M. 241315 A. M. N. H. 34677 F. M N. H. F. M. N. H. 24133 24132 F. M. N. H. 24136 F. M. N. H. 24135 F. M. N. H. 24134 B. M. 8.1.10.1 F. M. N. H. 21147 M. P. H. N. 565 M. P. H. N. 561 M. P. H. N. 562 B. M. 20.2.1.2 U. S. N. M. 148528 M. P. H. N. 564 A. M. N. H. A. M. N. H. N. H. R. S. N. H. R. S. N. H. R. S. N. H. R. S. 60508 61406 2 5 7 14 N. H. R. S. 10 A. M. N. H. 72587 B. Z. M. 602 U. S. N. M. 137513 cf ad. M^rida, Venezuela 44.3 16.7 cf ad. M6rida, Venezuela 43.4 10.0 c?" ad. M6rida, Venezuela 44.0 16.5 d" ad. M6rida, Venezuela 44.2 16.1 cf ad. M^rida, Venezuela 42.3 16.7 av. 5 43.6 16.4 cf sad. M^rida, Venezuela 43.5 16.9 cT sad. M^rida, Venezuela 43.6 16.2 (^ sad. M6rlda, Venezuela 15. 4 9 ad. M^rida, Venezuela 38.2 14. 1 9 ad. M6rida, Venezuela 36.4 13.4 9 ad. M^rida, Venezuela 36.3 13.4 9 ad. M6rida, Venezuela 37.7 13.9 av. 4 37.2 13.7 9 sad. M6rida, Venezuela 37.3 13.6 9 sad. M^rida, Venezuela 35.0 12.6 d" ad. Choachi. Colombia *46.0 17.4 cf ad. Choachi, Colombia 16. 1 cf ad. Quetame, Colombia 45.5 16.6 av. 3 45.8 16.7 cf yg. Bogota, Columbia 44.7 10.7 cf yg. Choachi, Columbia 45.1 16.7 c? yg. Gualea, Ecuador 45.6 17.6 c? ad. Rio Chinchao 44.0 16.4 cf ad. Rio Chinchao 45.3 17.0 9 ad. Huanuco, Peru 35.3 13.0 9 sad. Ambo, Peru 36.1 13.6 9 sad. Ambo, Peru 38.1 14.0 d' ad. Lima, Peru 42.5 10.0 9 ad. Macate, Peru 35.2 12.7 9 ad. Peru *36.0 12.6 cf ad. Junin, Peru 45.5 16.6 cf ad. Junin, Peru 40.8 15.0 cf ad. Yana Mayo, Peru 42.6 15.9 cf ad. Marcapata, Peru 16. 0 9 ad. Cutervo, Peru *38.0 13.8 cf ad. El Chiral, Ecuador 45.5 17.1 cf sad. Guainche, Ecuador 48.2 17.3 cf ad. Panecillo, Ecuador 48.0 16.8 cf ad. San Antonio, Ecuador 44.0 16.3 cf ad. Carapungo, Ecuador 46.8 17.4 cf ad. Nara Papallacta, Ecuador 45.4 17.3 9 ad. Guapulo, Ecuador 37.9 13.5 cf ad. Nequejahuira, Bolivia 41.6 15.3 cf yg. Limbani, Peru 42.4 15.5 cf yg. Limbani, Peru 40.3 15.2 Mustela frenata 14.2 12.0 15.0 14.1 11.8 15.2 14.1 11.6 15.4 13.7 11.5 15.4 13.8 11.5 15.5 14.0 11.7 15.3 13.8 12.1 15.1 13.5 11.4 15.5 12.5 10.5 15.4 11.9 11.5 11.4 11.4 11.6 11.1 11.1 9.7 9.9 9.7 9.8 9.8 9.2 8.7 13.2 13.0 11.8 13.8 13.0 13.2 12.0 Mustela frenata 13.6 11.8 15.6 12.7 10.8 14.3 13.7 11.0 15.3 13.3 11.2 15.1 13.9 13.5 12.3 11.5 15.6 15.2 Mustela frenata 14.0 11.8 15.4 Mustela frenata 14.5 12.5 14.9 13.9 11.7 15.5 10.5 8.9 12.1 10.9 9.0 12.4 10.6 9.1 12.6 Mustela frenata 13.4 11.1 14.9 10.0 8.4 12.0 10.7 9.2 12.5 Mu stela frenata 14.0 11.6 15.1 13.6 11.3 15.1 13.0 10.2 15,0 14.5 11.2 9.5 13,0 13.8 11.5 14.5 14.5 12.3 15.7 13.3 14.8 14.0 11.9 14.8 13.5 11.1 14.9 ll.l 12.6 15.8 11.3 9.5 12.7 Mustela frenata 12.2 10,0 14.7 Cranial Measurements 431 5 Tympanic BuUa s s ■^<— E Xi •a o!"S / * > ^ — o 09 0) .a ^^ Xi ^O e of S mai ccip V- --t ^^ ■v jd ja o t; lo OQ i IS ^3 J Q. Q a (a 'i 73 •a OJ « J3 B Depth Ant. basic Depth posts of M in<>ridana 23.9 23.5 23.6 23.6 23.0 23.5 28.5 27.4 28.0 27.3 27.6 27.8 13.0 13.8 13.9 14.2 13.5 13.7 6.6 6.5 7.0 6.4 6.5 6.6 3.0 2.4 '2!3 2.6 6.2 6.0 6.2 5.9 5.8 6.0 5.4 5.5 5.6 5.2 5.6 4.4 4.1 4.7 4.5 4.3 4.4 2.5 2.3 2.4 2.3 2.1 2.3 15.0 15.6 15.7 15.3 15.5 16.0 13.4 13.3 12.9 13.0 13.7 13.3 23.0 23.4 27.5 27.9 25.0 13.2 13.4 13.2 6.4 2.2 6.0 5.6 5.5 5.0 5.7 5.7 5.3 4.5 4.2 4.2 2.5 2.3 2.3 16.0 15.0 13.0 13.2 20.3 23.0 12.4 5.9 1.8 20.1 23.4 12.1 5.9 2.0 18.8 22.0 11.7 5.5 2.0 4.9 4.5 4.6 3.5 1.6 19.5 12.1 5.9 1.8 4.9 4.7 4.9 3.7 1.8 19.7 22.8 12.1 5.8 1.9 13.0 10.5 11.2 19.2 19.3 22.5 21.8 11.9 11.5 5.6 5.8 2.1 1.8 5.1 4.7 4.6 4.4 4.8 4.7 3.5 3.5 1.7 1.6 13.5 13.8 11.1 11.4 aflinis 29.5 27.7 28^6 6.4 6.1 6.0 5.8 6.0 6.1 5.7 5.7 5.8 4.4 4.5 4.4 4.4 2.3 2.1 2.0 2.1 13.9 13.0 13.3 13.4 23.9 23.9 29.0 28.7 aureoventris 23.8 28.3 14.0 15.1 6.9 6.8 5.9 5.9 5.7 5.6 5.7 5.8 4.4 4.5 2.5 2.1 2.7 6.7 6.0 6.4 5.0 3.0 17.0 13.5 14.3 helleri 24.5 24.5 29.1 29.0 14.4 14.4 6.5 6.5 2.4 2.4 5.8 5.5 5.9 5.8 4.6 4.5 2.5 2.5 16.0 16.0 14.0 13.7 17.9 18.8 19.8 20.8 22.4 22.2 11.8 11.8 12.0 4.7 5.0 5.4 2.0 1.9 2.0 4.7 4.7 4.7 4.4 4.4 4.6 4.6 4.6 4.6 3.5 3.6 3.5 1.6 1.7 1.5 11.8 12.6 13.9 10.0 10.5 11.0 agilis 23.1 28.2 13.8 6.9 2.6 5.8 5.5 5.8 4.5 2.2 14.8 12.9 18.0 18.1 20.6 21.5 12.4 13.0 5.0 5.8 1.7 2.2 4.4 4.8 4.1 4.1 4.4 4.5 3.2 3.5 1.5 1.9 13.0 12.0 10.3 11.7 macrura 23.8 29.0 22.9 26.5 21.8 26.2 14.5 14.5 13.5 6.2 7.5 7.0 2.7 2.5 5.5 5.8 5.8 5.7 5.4 5.0 5.5 5.0 5.5 5.1 5.6 5.2 4.3 4.0 4.4 4.3 2.1 2.0 2.3 2.1 15.7 14.6 14.6 13.2 12.9 12.3 18.1 *23.0 4.9 4.6 4.9 3.6 1.9 11.7 24.2 25.6 23^5 23.7 24.5 23.2 30.3 28^6 29.0 29.9 15.0 15.6 14.5 14.2 14.6 14.3 7.0 7.2 7.0 7.0 7.0 7.1 2.3 3.0 '2^9 2.9 2.6 5.9 6.1 6.0 5.8 6.2 6.1 5.6 5.2 5.4 5.1 5.8 5.4 6.0 5.4 5.7 5.4 6.2 5.6 4.8 4.5 4.3 4.3 4.5 4.3 2.6 2.3 2.2 2.1 2.3 2.5 16.4 15.7 16.0 17.5 13.8 13.9 i4;i 14.2 9.5 19.7 22.5 boliviensis 22.2 25.0 12.4 13.4 5.9 7.4 2.0 2.2 5.1 5.3 5.5 5.1 4.5 4.7 3.7 5.0 5.0 4.8 5.2 5.4 5.1 4.4 4.2 3.9 1.7 2.2 2.2 1.9 12.9 14.8 11.3 12.4 432 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial Measurements — Continued a o o 1 a o tt O o9 s M T3 O e •3 et ■» a S O o .a »M 0 a 0 •<^ ■£ 19 B O 33 -*^ o ^,= c 3i a> 3 V u. a >— < u n J CQ o Mustela afrirana A. M. N. H. 374.75 c? sad. Para, Brazil 47.8 17.2 14.3 12.9 15.4 B. M. 5.1.25.1. d'? sad. Para, Brazil 44.9 16.4 13.0 10.2 14.2 B. M. 26.1.8.10. cf? ad. Para, Brazil 44.6 16.4 13.3 10.9 14.4 Mustela afrirana B. M. 24.12.12.21. 9 ad. Moyobamba, Peru 45.8 16.9 13.5 12.0 14.6 M. P. H. N. 563 9 ad. Yuriniaguas. Peru 17. 5 ± A. M. N. H 61813 9 yg. Vai.d.Peren^, Peru 44.6 16.0 13.0 11.0 15.4 Mustela ermines M. C. Z. 10012 d' ad. Pt. Barrow 43.3 15.9 15.2 12.0 16.2 F. M. N. H. 35894 cf ad. Pt. Barrow 41.8 15.6 15.3 13.3 16.0 A. N. S. P. 6909 d' ad. Pt. Barrow 42.3 15.1 14.8 11.6 15.3 A. N. S. P. 6910 d' ad. Pt. Barrow 16.2 16.0 12.7 16.7 N. M. C. 2445 d' ad. Salirochit River 42.8 15.7 15.4 13.0 16.0 av. 5 42.5 15.7 15.3 12.5 16.0 F. M. N. H. 35895 9 ad. Pt. Barrow 35.4 13.1 12.7 10.3 13.3 M. V. Z. 43286 9 ad. Pt. Barrow 35.4 12.9 12.0 10.0 13.2 U. S. N. M. 243489 9 sad. Alatna River 37.0 13.4 11.9 9.4 13.5 U. S. N. M. 243493 9 sad. 16 mi. below Bettles 36.2 13.0 12.0 9.4 13.0 U S. N. M. 180459 9 ad. N. Fk. Ku.skokim 35.3 12.9 11.2 8.8 12.8 U. S. N. M. 242205 9 ad. Fairbanks 34.5 12.4 11.5 9.3 13.0 U. S. N. M. 157306 9 sad. Bear Creek 36.9 13.5 12.6 9.5 13.6 U. S. N. M. 157305 9 sad. Bear Creek 35.2 12.5 11.4 9.6 12.8 av. 8 35.8 12.9 11.8 9.3 13.1 Mustela erminea M. C. Z. 29831 d'ad. Ymer Is 41.6 15.6 15.3 12.0 13.0 C. Z. M. 1245 d' ad. Danmarks Havn 40.7 15.0 14.7 11.9 14.9 C. Z. M. 1246 d" ad. Danmarks Havn 39.0 14.1 13.8 10.9 14.2 C Z M. 1247 d' ad. Danmarks Havn 41.0 15.2 14.5 11.5 14.4 C. Z. M. 1248 d' ad. Danmark.s Havn 42.3 15.5 15.5 12.0 15.3 C. Z. M. 1249 d' ad. Danmarks Havn 42.2 15.8 15.0 12.2 15.1 C.Z. M. 1871 d-ad. ScoresbySd 42.4 15.3 15.3 12.3 15.6 av. 7 41.3 15.2 14.9 11.8 14.9 C. Z. M. 1060 9 ad. Turner Sd 35.9 13.0 12.5 10.1 12.9 B. Z. M. 43965 9 ad. Kap Hoegh 37.8 13.4 12.7 10.4 13.2 Mustela erminea CM. 6688 d'ad. Prairie Point *39.9 14.5 13.5 11.5 14.6 average 1 f 37.5 13.8 13.1 10.6 13.8 maximum \ d' 1 ad. and Southampton Isl \ 39.9 14.5 13.5 11.5 14.8 minimum J 10 sad. 1 35.7 13.1 12.6 9.7 12.7 CM. 8474 9 ad. Minnimunnek Pt 32.6 11.9 11.3 9.2 12.2 average 1 f 34.2 12.4 11.7 9.5 12.6 maximum 9 1 ad. and Southampton Isl 35.1 13.0 12.7 9.9 13.1 minimum J 4 sad. ( 32.6 11.9 11.3 9.2 12.2 Mustela erminea U. S. N. M. 107406 d'ad. Kadiak 43.2 15.4 14.9 11.1 14.9 F. M. N. II. 7290 d'ad. Kadiak Id 42.1 14.9 14.0 10.9 14.4 U. S. N. M. 98042 9 ad. Kadiak 33.0 12.0 10.2 8.0 11.3 Cranial Measurements 433 x: Tympanic Bulla 2 s J3 -c ^ ■5 "3 -■2 J3 f ■^ \ ^ 1- u 1 coS.S' x- '^ •^ a 0 •^•— ^ -a c) JS 0-0 0 fe ro ■5 3 a •0 C9 *^ c 0 Q J3 e 1 "2 C 1-4 Q africana 26.4 32.2 16.9 8.2 2.8 6.6 5.8 6.2 4.7 2.4 17.8 14.9 23.3 26.8 14.4 7.0 2.7 5.9 5.4 5.5 4.5 1.8 16.4 13.7 24.4 29.4 14.4 6.7 2.5 6.0 5.6 5.7 4.5 1.8 15.5 13 6 stolzmanni 23.. 5 28.8 14.5 6.2 2.6 5.9 6.0 5.4 5.8 4.9 5.6 4.3 4.4 1.8 2.3 16.5 13.9 24! 1 28!9 15^7 7^0 2^8 6.2 5.6 5.9 4.5 2.2 16.8 13.4 arctica 23.5 27.6 15.5 8.4 5.6 5.1 5.1 3.9 2.0 13.3 13.2 23.2 27.6 15.1 8.5 6.0 5.2 5.3 4.0 2.2 14.1 13.5 22.9 26.2 15.2 8.4 5.9 5.7 5.1 5.4 5.3 5.7 4.1 4.3 2.4 2.1 14.5 12.5 23! 5 28^5 i5!2 '8^0 5.6 5.0 5.2 4.0 2.0 14.3 13^3 23.3 27.5 15.3 8.3 5.8 5.2 5.3 4.1 2.1 14.1 13.1 19.3 23.0 12.7 7.2 5.0 4.5 4.8 3.5 1.7 12.0 10.5 18.8 21.8 12.6 6.7 4.6 4.2 4.3 3.2 1.8 11.4 10.3 19.4 22.0 13.2 7.2 4.7 4.3 4.5 3.3 1.7 11.9 10.6 19.1 21.8 12.9 7.2 5.0 4.2 4.5 3.3 1.8 11.9 10.3 18.9 20.7 12.7 7.0 4.7 4.3 4.5 3.3 1.8 10.5 9.7 18.0 12.7 6.8 4.8 4.3 4.5 3.3 1.7 11.8 10.2 18.7 20^9 13.0 6.9 4.8 4.4 4.5 3.3 1.8 11.4 10.2 17.8 20.5 12.6 6.3 4.8 4.2 4.4 3.3 1.6 11.2 10.1 18.7 21.2 12.9 6.9 4.8 4.3 4.5 3.3 1.7 11.5 10.2 Polaris 22.3 26.1 14.8 8.0 5.8 5.2 5.7 4.1 2.2 13.7 12.3 22.3 25.5 14.6 7.8 5.9 5.1 5.6 3.9 2.0 13.4 12.2 21.3 24.7 14 5 7.3 5.3 4.8 5.0 3.7 1.9 12.0 11.2 22.3 26.7 14.4 7.6 5.7 5.1 5.5 4.1 2.0 13.1 12.1 23.2 27.9 15.1 8.1 5.8 5.5 5.8 4.0 2.3 13.8 12.8 23.2 28.0 15.2 8.1 5.9 5.2 5.7 4.2 2.3 13.7 12.4 15.4 8.3 5.7 5.1 5.3 3.9 2.0 14.4 12.8 2214 26;5 14.9 7.9 5.7 5.2 5.5 4.0 2.1 13.4 12.3 19.6 22.6 13.6 7.1 5.0 4.5 4.7 3.6 1.7 12.3 10.3 19.3 22.4 13.9 7.0 4.8 4.2 4.5 3.7 1.8 12.1 10.9 semplei 21.5 24.8 14.1 7.9 5.0 4.6 4.7 3.6 2.0 14.0 11.6 20.2 23.6 13.6 7.2 5.1 4.7 4.9 3.6 2.0 13.4 11.0 21.5 24.8 15.0 7.9 5.3 4.9 5.0 3.7 2.3 14.0 11.6 19.2 22.2 12.9 6.4 4.8 4.4 4.5 3.4 1.8 12.3 10.5 18.5 21.2 12.2 6.6 4.4 3.9 4.2 3.0 1.5 11.9 9.8 18.8 20.9 12.8 6.7 4.5 4.1 4.3 3.1 1.6 12.1 10.1 19.4 21.5 13.4 7.0 4.3 4.3 4.5 3.3 1.7 12.4 10.4 18.1 20.1 12.2 6.6 4.6 3.9 4.2 3.0 1.5 11.9 9.6 kadiarensis 22.1 26.0 15.1 7.6 5.5 5.1 5.1 3.8 2,0 14.0 12.2 21.6 26.0 14.1 7.1 5.7 5.2 5.2 3.8 1.7 13.5 11.5 16.9 19.4 11.5 6.0 4.4 4.0 4.0 2.8 1.4 11.5 8.9 434 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial MEASxmEMENTs — Continued =3 o O s 3 a a 3 do ■3 d o o Ml 09 T3 a X 03 o 1 a 3 J3 0) j3 ■w c .2 tc'o 0 0 ^ c g «M 0 j3 0 fl Is y 0 ^ ■^ a) t-. a m hJ « 0 average maximum minimum average maximum minimum J U. S. N. M. M. C. Z. U. S. N. M. U. S. N. M. U. S. N. M. M. C. Z. M. V. Z. U. S. N. M. A. M. N. H A. M. N. H. A. M. N. H. Cornell C. M. C. M. M. V. Z. U. S. N. M. B. S. N. C. M. CM. F. M. F. M. F. M. F. M. F. M. F. M. F. M. F. M. F. M. F. M. F. M. F. M. N. H. N. H. N. H. N. H. N. H. N. H. N. H. N. H. N. H. N. H. N. H. N. H. average maximum minimum Walker Walker Wise. U. Wise. U. Walker Wise. U. Wise. U. Snyder Snyder Snyder & ad. 6 cf sad. 7 136112 242866 129703 110682 235959 18776 53789 32240 44nj"6? 67869 67868 15841 494 7461 10264 53788 135570 994 7460 10252 18134 18135 18130 18133 18131 7222 18127 18129 18132 18441 18440 7219 377 11 8681 8679 A 58 8691 8674 2637 993 2999 Mustela erminea f 40.9 14.3 12.5 10.2 13.5 3 mi. S Big Isl { 43.7 15.0 13.2 11.1 14.0 l 39.6 13.8 11.7 9.4 12.9 f 40.2 14.1 12.2 9.8 14.0 3 mi. S Big Isl \ 41.5 14.7 12.6 10.4 14.2 I 38.4 13.6 12.0 9.5 13.7 9 ad. Willow River 33.0 11.5 8.8 7.6 11.4 9 sad. 3 mi. S Big Isl 32.3 11.0 9.0 7.2 11.7 9 ad. Ft. Resolution 34.2 12.0 10.4 8.2 11.5 9 sad. 15 mi. above Smith Landing. . . 33.8 11.3 9.8 7.4 11.0 9 ad. Athabaska Delta 33.1 11.7 9.3 7.3 11.0 9 ad. Athabaska Delta 31.5 10.8 8.9 7.6 11.2 av. 6 33.0 11.4 9.4 7.6 11.3 Mustela erminea cf ad. Ogdensburg, N. Y 36.6 12.9 11.5 9.1 13.2 cf sad. Amsterdam. N. Y 36.7 12.9 11.5 9.4 13.3 cfad. Berlin, N.Y 36.2 12.5 10.2 8.9 12.7 cf ad. Berlin, N. Y 34.8 12.1 9.8 8.2 12.7 cf ad. Schoharie, N. Y 33.8 11.7 9.7 7.7 11.9 cf sad. Cascadilla Creek, N. Y 34.8 12.2 10.4 8.3 12.5 cf sad. Pymatuning Swamp 34.2 12.0 9.6 8.1 12.8 cf sad. 3)4 mi. W Linesville 37.6 13.2 11.0 8.9 12.7 cf sad. Lopez, Penn 36.5 12.6 10.7 8.6 12.6 av. 9 35.7 12.5 10.5 8.6 12.7 9 ad. Lake George 33.3 11.8 9.3 7.8 11.4 9 sad. Cattaraugus 31.4 10.8 9.2 7.2 10.0 9 sad. Pvmatuning Swamp 32.0 10.9 9.2 7.5 11.0 9 sad. 3 mi. NW Linesville 32.7 11.2 9.5 7.6 11.3 av. 4 32.4 11.2 9.4 7.5 10.9 Mustela erminea cf ad. Aitkin, Minn 40.3 14.1 12.7 10.6 14.6 cf ad. Aitkin, Minn 14.1 12.5 10.4 14.4 cf ad. Aitkin, Minn 40.7 15.0 12.2 9.7 14.6 cf ad. Aitkin, Minn 39.3 13.9 11.2 9.0 13.0 cf ad. Aitkin, Minn 38.5 13.4 11.2 9.0 12.6 cf ad. Aitkin, Minn 13.6 12.4 9.5 13.3 cf sad. Aitkin, Minn 38.4 13.1 11.1 9.0 13.0 cf sad. Aitkin, Minn 39.4 13.7 11.1 9.1 13.3 cf sad. Aitkin, Minn 39.3 14.3 11.5 9.5 13.2 cf -sad. Aitkin, Minn 40.2 14.0 12.0 9.9 14.6 cf sad. Aitkin, Minn 38.8 13.8 11.3 9.3 13.2 cf sad. Aitkin. Minn 13.9 11.9 10.0 13.5 av. 12 39.4 13.9 11.8 9.6 13.6 f 37.9 13.2 11.4 9.1 13.1 cf ad. Sand Elk River \ 39.5 13.9 12.6 9.8 14.2 sad. 5 ( 34.8 12.0 10.5 8.5 12.2 9 ad. Deer 31.8 11.0 9.0 7.5 10.9 9 ad. Grand Maris 32.7 10.8 11.0 9 ad. T. 61N, R. 26W 32.9 10.8 9.2 7.4 11.3 9 ad. T. 61N, R. 26W 33.6 11.6 9.8 7.6 12.2 9 ad. Elk River 32.8 11.5 9.9 8.1 11.3 av. 5 32.8 11.1 9.5 7.7 11.3 9 ad. Fi.sherLake 31.5 10.6 9.5 7.7 10.8 9 ad. Gordon 32.8 11.5 9.8 7.2 11.4 9 ad. Beaver Dam 32.9 11.0 9.8 7.8 11.7 9 ad. Beaver Dam 34.1 11.3 9.4 7.5 11.8 9 ad. Beaver Dam 31.9 10.8 9.2 7.4 11.2 av. 5 32.6 11.0 9.5 7.5 11.4 Cranial Measurements 435 :S 1 Tympanic Bulla s ^•3 lat rders 1 0 r 1—1 \ f \ f Skul! iiargin cipital f Skui: ior bo 1 1 t Breadth Depth a 1 1 m a Depth 0 Ant. n basioc Depth 0 poster of Ms richardsonii 21.2 24.2 14.9 7.9 5.3 4.7 4.8 3.8 2.1 13.0 11.9 22.2 25.5 15.5 8.3 6.0 5.0 5.2 4.2 2.3 14.2 11.4 20.0 22.9 14.2 7.2 4.9 4.5 4.6 3.7 2.0 12.5 12.3 20.6 23.3 14.6 7.7 5.2 4.7 4.9 3.7 2.1 13.6 11.8 21.5 24.4 15.1 8.2 5.5 4.9 5.0 3.8 2.5 14.5 12.2 20.0 22.0 13.7 7.0 5.0 4.5 4.7 3.5 2.0 12.8 11.4 15.9 18.0 12.5 5.7 4.0 3.8 3.9 2.7 1.5 11.6 8.9 16.6 17.7 12.3 6.5 4.0 3.7 3.9 3.0 1.5 10.0 8.8 17.2 19.7 12.3 6.5 4.2 4.0 4.2 3.2 1.7 12.0 9.6 16.5 18.3 12.1 6.0 4.2 3.8 3.9 2.9 1.5 11.2 9.3 15.7 18.1 11.6 5.6 4.0 3.8 3.8 3.0 1.6 10.5 8.6 16.3 18.3 11.6 6.3 3.8 3.7 3.8 3.1 1.6 11.0 8.7 16.4 18.3 12.1 6.3 4.0 3.8 3.9 3.0 1.6 11.1 9.0 cicngnanii 18.5 21.7 13.3 7.0 4.7 4.4 4.5 3.4 1.8 12.3 10.4 19.1 21.5 13.7 6.9 4.6 4.4 4.5 3.4 2.0 13.2 10.8 18.8 20.6 12.9 6.7 4.3 3.8 4.0 3.3 1.9 11.5 10.0 17.4 19.8 12.3 6.3 4.5 3.9 4.1 2.9 1.7 11.5 9.7 17.3 20.0 11.8 6.4 4.2 3.8 4.0 2.9 1.5 11.8 9.5 17.9 19.3 12.4 6.6 4.4 4.1 4.2 3.0 1.6 12.2 10.3 17.3 19.0 11.9 6.4 4.3 3.9 4.2 3.0 1.9 11.5 9.2 18.8 20.6 12.9 7.2 4.9 4.3 4.5 3.5 2.0 12.6 10.6 18.6 20.3 13.1 6.8 4.5 4.1 4.3 3.2 1.6 10.9 9.9 18.2 20.3 12.7 6.7 4.5 4.1 4.3 3.2 1.8 11.9 10.0 11.8 5.8 4.4 3.8 4.0 3.0 1.8 9.1 15.3 10.7 5.7 3.8 3.7 3.6 2.7 1.5 16:3 8.2 15.9 17 ^5 11.1 5.9 3.7 3.4 3.6 2.8 1.5 10.3 8.2 16.0 18.0 11.8 6.2 3.9 3.6 3.7 3.1 1.7 10.9 8.8 15.7 17.8 11.4 5.9 3.9 3.6 3.7 2.9 1.6 10.5 8.6 bangs! 20.6 23.5 14.3 7.7 4.9 4.6 4.9 3.6 1.9 13.3 11.8 24.3 5.3 4.5 4.6 3.7 2.1 12.2 26! 2 22.8 14^6 rs '.'.'.'. 5.2 5.0 5.1 3.8 2.6 13^5 12.0 19.5 22.0 14.3 7.2 5.1 4.5 4.6 3.5 2.0 13.0 11.2 18.8 21.0 13.7 7.3 4,8 4.4 4.5 3.4 1.8 12.0 10.5 14.4 7.9 4.8 4.4 4.4 3.5 1.8 11.2 19.5 22^0 14.3 6.9 4.6 4.5 4.5 3.5 1.9 is.i 11.0 19.0 21.6 13.6 7.2 4.7 4.3 4.5 3.4 2.0 12.8 11.1 20.2 22.8 14.3 7.2 4.9 4.7 4.7 3.5 2.1 12.5 11.2 19.6 22.6 15.0 7.1 5.2 4.7 4.8 3.7 2.0 13.3 11.5 19.7 22.0 13.8 7.6 5.1 4.5 4.7 3.7 2.0 13.0 11.1 24.7 14.6 7.3 5.3 4.6 4.8 4.0 2.1 11.8 19^7 22.7 14.3 7.4 5.0 4.6 4.7 3.6 2.0 13^0 11.4 19.3 21.6 14.4 7.2 4.7 4.3 4.4 3.4 1.9 12.7 10.8 20.5 22.8 15.3 8.0 5.1 4.4 4.6 3.6 2.0 14.0 11.7 18.0 20.8 12.4 6.3 4.3 4.1 4.1 3.1 1.7 12.0 10.0 16.6 18.4 11.7 5.9 4.2 3.7 3.8 2.8 1.6 11.2 9.3 15.9 17.3 12.2 5.6 3.9 3.8 3.8 2.9 1.6 10.7 8.5 15.8 16.8 12.3 6.2 3.8 3.6 3.6 2.8 1.4 10.0 8.3 15.7 17.1 11.9 5.7 4.0 3.6 3.7 3.0 1.6 9.0 8.9 16.8 18.3 11.8 6.4 4.0 3.8 4.0 2.9 1.7 10.5 9.2 16.2 17.6 12.0 6.0 4.0 3.7 3.9 2.9 1.6 10.4 8.8 15.8 17.4 11.2 5.6 3.8 3.5 3.7 2.6 1.4 10.4 8.5 15.7 16.7 11.6 5.7 4.2 3.8 3.8 2.8 1.6 9.6 8.9 16.9 18.9 12.2 6.4 4.1 3.8 3.8 3.2 1.5 11.3 9.7 16.7 18.0 12.8 6.1 4.4 3.8 3.8 3.0 1.7 10.8 9.0 15.9 17.5 12.4 5.8 4.0 3.6 3.8 3.0 1.6 11.4 9.2 16.2 17.7 12.0 5.9 4.1 3.7 3.8 2.9 1.6 10.7 9.1 436 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial Measurements — Continued a o O i a 3 fl o s « faA "O 0 a 1 o 00 ■■a o Is n ■s "S 0 ■s .a Xi .a -*3 ■n> -TJ o g 1^ 1— ( I-) n .a Ji ►ii O Mastela emiinea average 1 f 37.0 12.8 11.4 9.1 13.1 maximum \ cf ad. 5 Idaho Co \ 39.8 14.1 13.0 10.0 14.2 minimum J I 35.8 12.2 10.6 8.6 12.0 M. V. Z. 90763 9 ad. Pilot Creek, Idaho 31.6 10.8 9.2 7.0 10.6 average ) 1 32.2 ina.ximum \ 9 ad. 1 and Idaho Co \ 32.8 minimum j sad. 4 I 31.6 10.6 11.2 10.8 9.0 7.1 11.1 9.2 7.2 12.2 8.5 7.0 10.5 average maximum } minimum J U. S. N. M. M. V. Z. M. V. Z. U. S. N. M. M. V Z. cf ad. 8 74422 995 473 74773 78243 Windham 9 ad. .Juneau. . . . 9 ad. Juneau. . . . 9 ad. Helm Bay. 9 ad. Wrangel. . . 9 sad. Windham av. 5 37.5 38.9 36.5 33.2 33.1 32.9 32.2 31.9 32.7 average maximum minimum I cf ad. to sad. 12 Mole Harbor. average ] maximum \ 9 ad. 2 and minimum j sad. 4 Mole Harbor M. V. Z. M. V. Z. 289 d' ad. Saook Bay. 286 d' ad. Saook Bay . average i maximum [ cf ad. 5 minimum ) average ] maximum \ cf ad. 5 and minimum ) sad. 15 M. V. Z. M. V. Z. 31223 9 sad. 31232 cf ad. U. S. N. M. 94430 cf ad. average 1 maximum \ . N Breadt Depth Length ~5, S. o! -t3 n j3 Depth Ant. basic Depth pOSt£ 1 of M fallf^nda 18.3 20.8 13.1 6.8 .. **4.7 **4.2 **4.4 **3.4 **1.9 11.7 10.5 19.6 22.8 14.1 7.6 5.1 4.6 4.7 3.6 2.1 12.4 11.0 17.0 19.4 12.2 6.2 .. 4.3 3.9 3.9 3.2 1.7 11.0 10.0 15.4 17.4 11.1 5.5 3.7 3.5 3.6 2.8 1.5 10.5 8.6 16.0 18.5 11.5 5.6 4.1 3.7 3.8 2.9 1.5 10.0 9.0 15.8 17.2 11.0 5.3 3.8 3.5 3.7 2.7 1.5 11.0 8.7 14.7 15.4 10.5 5.2 3.8 3.5 3.8 2.7 1.5 9.5 8.3 15.7 17.4 11.5 5.5 3.8 3.4 3.7 2.6 1.2 9.3 8.2 oKmpica 15.3 17.9 11.6 5.3 .. 4.0 3.6 3.9 2.9 2.1 9.9 8.9 16.3 18.2 11.7 5.6 4.2 3.6 3.7 2.9 1.8 10.0 9.2 15.0 17.2 10.7 5.2 4.2 3.5 3.7 2.7 1.7 10.0 8.6 16.6 18.6 11.2 5.9 4.0 3.7 3.9 3.0 1.7 10.2 9.3 16.9 18.8 11.4 6.3 4.3 3.8 3.9 3.1 1.7 10.3 9.2 16.0 18.1 11.3 5.7 4.2 3.7 3.9 2.9 1.8 10.1 9.0 13.2 15.3 10.2 4.6 3.4 3.0 3.3 2.5 1.3 8.5 7.2 13.1 14.4 9.7 4.8 3.4 3.1 3.4 2.3 1.3 9.0 7.3 13.7 15.4 9.5 4.8 .. 3.3 3.0 3 1 2.1 1.2 8.3 7.2 13.3 15.0 9.8 4.7 3.4 3.1 3.3 2.3 1.2 8.6 7.2 strcntori 17.0 19.2 11.8 6.4 4.3 3.8 3.9 3.1 1.8 10.8 9.8 18.0 19.8 12.6 0.9 4.4 4.0 4.1 3.5 2.1 11.4 10.3 10. 1 18.5 11.1 6.0 4.1 3.6 3.8 2.9 1.6 10.3 9.5 14.3 15.9 10.6 5.2 .. 3.6 3.2 3.5 2.6 1.6 9.4 8.1 14.8 16.3 11.2 5.4 3.7 3.3 3.6 2.7 1.7 10.0 8.4 !4.1 15.5 10.0 5.0 .. 3.5 3.1 3.3 2.5 1.5 9.2 7.9 gulosa 16.3 18.3 11.3 5.9 4.2 3.8 4.1 2.9 1.6 10.5 9.4 16.9 18.5 11.8 5.9 4.2 3.7 3.7 2.8 1.8 10.6 9.4 16.4 18.4 11.5 5.9 4.2 3.7 3.9 2.9 1.7 10.3 9.2 17.2 19.3 12.1 6.3 4.5 3.8 4.1 3.2 2.0 11.2 9.7 15.6 17.8 10.8 5.5 3.9 3.5 3.6 2.7 1.6 9.6 8.5 14.1 15.8 10.9 5.4 3.7 3.2 3.5 2.7 1.6 8.4 7.4 13.6 15.6 10.0 4.8 3.6 3.3 3.5 2.5 1.6 9.0 7.6 13.8 15.4 10.1 4.9 3.6 3.2 3.3 2.3 1.4 8.1 7.3 14.5 15.6 10.0 5.0 3.5 3.2 3.4 2.5 1.4 8.6 7.5 13.6 15.2 9.7 4.7 3.8 3.4 3.5 2.7 1.5 8.1 7.2 13.9 15.5 10.1 5.0 .. 3.6 3.3 3.4 2.5 1.5 8.4 7.4 muricus *16.0 17.5 5.7 4.0 3.6 3.7 2.8 1.6 11.2 8.8 17.4 li'.s 5.6 4.3 3.7 4.0 3.0 1.6 8.7 15.9 17.5 10.8 5.6 4.2 3.5 3.6 2.8 1.8 '9.6 8.4 15.1 17.1 10.7 5.4 4.0 3.5 3.5 2.7 1.5 9.8 8.0 16.2 18.3 10.8 5.7 4.3 3.6 3.9 2.9 1.6 8.5 15.8 17.6 10.9 5.6 4.2 3.6 3.7 2.8 1.6 i6.'2 8.5 14.9 16.0 10.9 5.3 3.6 3.2 3.3 2.7 1.7 8.8 7.2 14.0 15.6 10.1 5.0 3.5 3.2 3.3 2.5 1.4 8.3 7.2 14.3 ?6.5 10.2 4.7 3.6 3.2 3.4 2.7 1.5 9.1 7.8 15.2 17.1 10.0 5.3 3.6 3.1 3.4 2.6 1.5 8.2 13.9 15.8 10.0 5.0 3.4 2.9 3.1 2.4 1.3 'i.'% 7.2 14.1 15.5 10.5 5.1 3.6 3.2 3.3 2.6 1.4 8.9 7.6 14.4 16.1 10.3 5.1 3.6 3.1 3.3 2.6 1.5 8.8 7.6 440 University of Kansas Publs., Mus. Nat. Hist. Table of Cranial Measurements — Concluded o O S 3 C -a a a K V CO =3 o o pa B 13 03 n j3 .0 £ A O A. M. N. H. A. M. N. H. A. M. N. IT. A. M. N. H. A. M. N. H. A. M. N. H. A. M. N. H. A. M. N. H. A. M. N. H. A. M. N. H. average maximum minimum averase maximum minimum 12432 9 ad. Conard Fissure, Ark 35. 1 12433 9 ad. Conard Fissure, Ark 12435 9 ad. Conard Fissure, Ark *32. 5 1 1766 9 ad. Conard Fissure, Ark 34. 5 av 34.0 12437 cf .«ad. Conard Fissure. -■Vrk 39. 2 12441 d" ad. Conard Fissure. Ark 38.5 12436 cf ad. Conard Fi.ssure, Ark 12444 cf ad. Conard Fissure, Ark 11769 d" sad. Conard Fissure. Ark 12438 (/• vs. Conard Fissure, Ark 36. 6 av 38.1 f 29. 5 cf ad. 6 Point Barrow \ 3o! 1 [ 27.6 f 27.8 9 ad. and sad. 4 Point Barrow \ 28. 5 [ 27.0 average 1 maximum | d" ad. 2, and minimum J sad. 4 average 1 maximum > 9 ad. 3, and minimum J sad. 1 Swenk, Mr. Swenk, IMr. 5 8 Swenk, Mr. 10 U. S. N. M. 171490 & ad. 'pes of the genera and the exact bases and localities of the species. Proc. Zool. Soc. London, 1911:120-158, March, 1911. 1920. Report on the Mammalia collected by Mr. Edmund Heller during the Peruvian expedition of 1915 under the auspices of Yale Uni- versity and the National Geographic Society. Proc. U. S. Nat. Mus., 58:217-249, pis., 14-15. Thubber, W. a. 1940. A weasel attacks a varying hare. Jour. Mamm., 21:356, August 14, 1940. TSCHUDI, J. J. 1844. Untersuchungen iiber die Fauna Peruana. Therologie. Druck und verlag von Scheitlin und Zollikofer, St. Gallen, xxx -\- 262 pp., 18 pis. Vestal, E. H. 1937. Activities of a weasel at a woodrat colony. Jour. Mamm., 18:364, August 14, 1937. 1938. Biotic relations of the woodrat (Neotoma fuscipes) in the Berkeley Hills. Jour. Mamm., 19:1-36, figs. 1-4, February 14, 1938. Warren, E. R. 1924. Ground squirrels and weasels. Jour. Mamm., 5:265-266, Novem- ber 15, 1924. 1932. When do weasels mate? Jour. Mamm., 13:71-72, February 9, 1932. Wight, H. M. 1932. A weasel attacks a man. Jour. Mamm., 13:163-164, May 11, 1932. Willis, J. C. 1922. Age and Area: A study in geographic distribution and origin of species. University Press, Cambridge, England, x -f- 259 pp., il- lustrated. 458 University of Kansas Publs., Mus. Nat. Hist, WiNECOFF, T. E. 1930. Least weasel in Pennsylvania. Jour. Mamm., 11:312-313, August, 1930. WOLDRICH, J. N. 1884. Diluviale Fauna von Zuzlavi^itz bei Winterberg im Bohmerwalde. Dritter Theil. . . . Sitzungsberich. d. K. Akad. d. Wisseu. Math.— Natur. Classe, 88 (heft 3, erste abth.): 978-1057, 3 pis., 2 figs, in text. Wright, P. L. 1942A. Delayed implantation in the long-tailed vi'easel (Mustek frenata), the short-tailed weasel (Mustela cicognani), and the marten (Martes americana). Anat. Rec, 83:341-353, 2 pis., July, 1942. 1942B. A correlation between the spring molt and spring changes in the sexual cycle in the weasel. Jour. Exp. Zool., 91:103-110, October, 1942. 1947. The sexual cycle of the male long-tailed weasel (Mustela frenata). Jour. Mamm., 28:343-352, 1 pi., 3 figs, in text, November 19, 1947. 1948A. Breeding habits of captive long-tailed weasels (Mustela frenata). Amer. Midland Nat., 39:338-344, March, 1948. 1948B. Preimplantation stages in the long-tailed weasel (Mustela fre- nata). Anat. Rec, 100:595-607, 2 pis., April, 1948. 1950. Effects of gonadotrophic hormone on the pelage of white winter long-tailed weasels; abstract. Anat. Rec, 106(no.2):130, Febru- ary, 1950. ZiMMERMANN, K. 1943. Zur Kenntnis deutscher Maus- und Zwerg-Wiesel. Zeitschr. fur Saugetierkunde, 15(3):289-298, 2 pis., 2 figs, in text, 3 tables, January 11, 1943. Transmitted May 22, 1950. Plate 2. ^ a. i ' 4 ^ K CO ' 00 > 9- h. L ; ;^v ^ '^ ^. ^ \ J I .r- k- 2 V Plate 2. Photographs, retouched, of skulls in dorsal view of nine subspecies of Mustela erminea. Natural size. a. Mustela erminea arctica, $ ad., 178405, U. S. Nat. Mus., Tanana, Alaska. h. Mustela er-minea semplei, $ sad., 6499, Carnegie Mus., Southhampton Island. c. Mustela erminea kadiacensis, $ ad., 107496, U. S. N. M., Kodiak Island, Alaska. i Mustela erminea richardsonii, $ ad., 133847, U. S. N. M., Ft. Franklin, MacK. «. Mustela erminea cicognanii, $ ad., 53788, Mus. Vert. Zool., Lopez, Pennsylvania. /. Mustela erminea bangsi, $ ad., 18130, Field Mus. Nat. Hist., Aitkin, Minn. ?. Mustela erminea invicta, S ad., 90759, Mus. Vert. Zool., Pilot Creek, Idaho. h. Mustela erminea alascensis, $ ad., 74665, Mus. Vert. Zool., Windham, Alaska. !■ Mustela erminea salva, $ ad., 74641, M. V. Z., Mole Harbor, Admiralty Id., Alaska. Plate 3, .1 > f '■ ) \ \ 1 -^^^ >^ a. ■J9- '^ V" -4 s . / /; \ ( ■Tk ' / 1 \ .• ^^i^ d. -"t ( Vv- 6. I y-^f H. ^•.■■w ^- h 1 1 , y ( ( c. / Plate 3. Photographs, retouched, of skulls in ventral view of nine subspecies of Mustela erminea. Natural size. a. Mustela erminea arctica, $, ad., 178405, U. S. Nat. Mus., Tanana, Alaska. h. Mustela erminea semplei, $ sad., 6499, Carnegie Mus., Southampton Island. c. Mustela erminea kadiacensis, S ad., 107496, U. S. N. M., Kodiak Island, Alaska. d. Mustela erminea richardsonii, $ ad., 133847, U. S. N. M., Ft. Franklin, MacK. e. Mustela erminea cicognanii, S ad., 53788, Mus. Vert. Zool., Lopez, Pennsylvania. /. Mustela erminea bangsi, S ad., 18130, Field Mus. Nat. Hist., Aitkin, Minn. g. Mustela erminea invicta, S ad., 90759, Mus. Vert. Zool., Pilot Creek, Idaho. h. Mustela erminea alascensis, £ ad., 74665, Mus. Vert. Zool., Windham, Alaska. i. Mustela erminea salva, $ ad., 74641, M. V. Z., Mole Harbor, Admiralty Id., Alaska. Plate 4. i/*> H-o^ a. #*' •^^TVy^ -"X c. . r 41 \v .^^'V'^ j^^ ^ o^^ f. m -?>-. 9- **^ f^'^. f (I Hi* i MM ' '^ : .If b. c. d. e. *.« ^''V /'i.i^ /» r ^ V Iju Plate 6. Photos, retouched, of skulls of 9 subspecies of msteJa ermxnea, x 1. a. Mustela erminea initis, $ ad., 289, Mus. Vert. Zool., Saook Bay, Alaska. b. Mustela erminea celenda, S ad., 1053, Los Angeles Mus., Craig, Alaska. c. Mustela erminea seclusa, S ad.. 31232, M. V. Z., Port Santa Cruz Alaska. d. Mustela erminea haidarum, $ ad., 230777, U. S. N. M., Graham Island B C e. Mustela erminea anguinae, $ ad., 13508, Nat. Mus. Canada, Capo Scott V 1. B. «.. /. Mustela erminea fallenda, S ad.. 7096. Nat. Mus. Canada, Huntingdon BC g. Mustela e. olympica, $ ad., 90738, U. S. N. M near head of Soleduc ^:'-^^^^- h. Mustela erminea streatori, S ad., 133, Coll. of Alex Walker, Blaine, Oiegon. i. Mustela erminea gulosa, i ad., 82177, U. S. Nat. Mus., Trout Lake, Wash. Plate 7. ^ ^ a. b ^ ^ i ^ / ' m. V ^ \J/\) Plate 7. Photos, retouched, of skulls, of subspecies of Mnstela erminea. Natural size. a. Mustcla erminea celeuda, i ad., 1053, Los Angeles Mus., Craig, Alaska. b. Musfela erminea seclnsa, S ad., 31232, M. V. Z., Port Santa Cruz, Alaska. c. Musfela erminea haidanim, S ad., 230777, U. S. N. M., Graham Island, B. C. d. Mnstela erminea angi(i)iae, i ad., 13508, Nat. Mus. Canada, Cape Scott, V. I., B. C. e. Mnstela erminea fallenda, i ad., 7096, Nat. Mus. Canada, Huntingdon, B. C. /. Mnstela erminea olympica, S ad., 90738, U. S. N. M., near head of Soleduc Riv., Wash. g. Mnstela erminea streatori, i ad., 133, Coll. of Alex Walker, Blaine, Oregon. h. Mnstela erminea gnlosa, S ad., 82177, U. S. Nat. Mus., Trout Lake, Wash. i, j, k. Mnstela erminea mnricus, i ad., 41501, M. V. Z., Baker Creek, 8675 ft., Nev. /, m. Mnstela erminea angnstidens, 6 ?, sad., 12437, A. M. N. H., Conard Fissure, Ark. n. Mnstela erminea angnstidens, S ?, ad., 12441, A. M. N. H., Conard Fissure, Ark. i^S^ -r- %^ Plate 8. ^ ^ %. a. r^,* << M 11^ • « 'H ^ »•' d. - H 4 S »^ y •» PLATE 8. Photos, retouched, of Mnsfete erminea angushdevs All in Amer. Mus. Nat. Hist., from Conard Fissure. Arkansas. Pleistocene in age x 1. a. Adult, probably male, 12441. e. Subadult, probably male 12437. b. Subadult, probably male, 12437. /. Young probably male 12438 c. Adult, probably male, 12444. g, h. Adult, type probably female, 12432. d. Adult, probably male, 12441. i. Adult, probably female, 12433. a. ti ;^ ''=*»v 38^ \ c I '^ k K \ J 9- h. Plate 9 ^ i / 1*M i r li -C\ -^ -^v /. ^^.i^w, ^^^-^N ,>!<.. .> r '• r. Plate 9. Photographs, retouched, of skulls in dorsal view of 9 subspecies of Musfela cnninca. Natural size. a. Mustela erminea arctica, 9 ad., 35895, Field Mus. Nat. Hist., Point Barrow, Alaska, b. Mustela erminea semplei, 9 ad., 6600, Carnegie Mus., Southhampton Island. c. Mustela erminea kadiacensis, 9 ad., 98042, U. S. Nat. Mus., Kadiak, Alaska. d. Mustela erminea richardsonii, 9 ad., 129703, U. S. N. M., Fort Resolution, MacK. e. M)istela erminea cicognanii, 9 ad., 7460, Carnegie Mus., Pymatuning Swamp, Pa. /. Mustela erminea bangsi, 9 ad., 8679, Univ. Wisconsin, T.61N, R.26W, Minn. g. Mustela erminea invicta, 9 ad., 90820, M. V. Z., IVz mi. W Iron Mtn., Idaho. h. Mustela erminea alascensis, 9 ad., 74422, U. S. Nat. Mus., Juneau, Alaska. i. Mustela erminea salva, 9 ad., 74655, Mus. Vert. Zool., Mole Harbor, Alaska. i v **^ ff- ;i. Plate 10. I i^^ ^ *TN \ Ml,. A X. Plate 10. Photographs, retouched, of skulls in dorsal view of 9 subspecies of Mustela erminea. Natural size. a. Mustela erminea arctica, $ ad., 35895, Field Mus. Nat. Hist., Point Barrow, Alaska. 6. Mustela erminea seniplei, 9 ad., 6600, Carnegie Mus., Southhampton Island. c. Mustela erminea kadiacensis, 9 ad., 98042, U. S. Nat. Mus., Kadiak, Alaska. d. Mustela erminea richardsonii, 9 ad., 129703, U. S. N. M., Fort Resolution, MacK. «. Mustela erminea cicognanii, 9 ad., 7460, Carnegie Mus., Pymatuning Swamp, Pa. /. Mustela erminea bangsi, 9 ad., 8679, Univ. Wisconsin, T.61N, R.26W, Minn. g. Mustela ermiyiea invicta, 9 ad., 90820, M. V. Z., W^ mi. W Iron Mtn., Idaho. ^. Mustela erminea alascensis, 9 ad., 74422, U. S. Nat. Mus., Juneau, Alaska, i. Mustela erminea salva, 9 ad., 74655, Mus. Vert. Zool., Mole Harbor, Alaska. Plate 11, 'f^-'^S^f' ^^^^--i ys*^* Iftf?^-"^ d. >»v # '*^o ^.i -'.^ ■ i .1. 5 Ti* "*«s& ,f^-^ 9. h. T^^ ^ fOoh'Jt'^ r/ [pBl! P^iNS^^ ^^ Plate 11. Photographs, retouched, of skulls in lateral view of twelve subspecies of Mustela erminea. Natural size. a. Mustela erminea arctica, 9 ad., 35895, Field Mus. Nat. Hist., Point Barrow, Alaska, h. Mustela erminea semplei, 9 ad., 6600, Carnegie Mus., Southhampton Island. c. Mustela erminea kadiacensis, 9 ad., 98042, U. S. Nat. Mus., Kadiak, Alaska. d. Mustela erminea richardsonii, 9 ad., 129703, U. S. Nat. Mus., Fort Resolution, MacK, e. Mustela erminea cicognanii, 9 ad., 7460, Carnegie Mus., Pymatuning Swamp, Pa; /. Mustela erminea bangsi, 9 ad., 8679, Univ. Wisconsin, T.61N, R.26W, Minn. g. Mustela erminea invicta, 9 ad., 90820, M. V. Z., IVe mi. W Iron Mtn., Idaho. h. Mustela erminea alascensis, 9 ad., 74422, U. S. Nat. Mus., Juneau, Alaska. i. Mustela erminea salva, 9 ad., 74655, Mus. Vert. Zool., Mole Harbor, Alaska. j. Mustela erminea haidarum, 9 ad., 100624, U. S. Nat. Mus., Moresby Island, B. C. A:. Mustela erminea anguinae, 9 ad., 13673, Nat. Mus. Canada, Cape Scott, V. I., B. C. /. Mustela erminea fallenda, 9 ad., 7284, Nat. Mus. Canada, Huntingdon, B. C. Plate 12. y tu o CD -P 9^ 4 W \ 1 -"4 ^. .^ 4 e. 0 ^v^ ii^M ■v^ -'-^ fi'. t. Plate 12. Photographs, retouched, of skulls in dorsal view of eight subspecies of Miisfcla cDninea. Natural size. a. Mxsfela erminea haidarum, 9 ad., 100624, U. S. N. M., Moresby Island, B. C. 6. Mxstela erminea angniime 9 ad., 13673, N. M. Canada, Cape Scott, V. I., B. C. c. Mtistela erminea fallenda, 9 ad., 7284, Nat. Mus. Canada, Huntingdon, B. C. d. Mustela erminea olympica, 9 ad., 242133, U. S. Nat. Mus., Hayes Creek, Wash. e. Mustela erminea streatori, 9 ad., 9040, D. R. Dickey Coll., Blaine, Oregon. /. Mustela erminea gulosa, 9 ad., 77370, U. S. Nat. Mus., Trout Lake, Wash. g. Mustela erminea mnriciis, 9 ad., 41502, Mus. Vert. Zool., Baker Creek, Nevada. h. Mustela erminea angustidens, 9 ?, ad., 12435, A. M. N. H., Conard Fissure, Ark. t. Mustela erminea ang^istidens, 9 ?, ad., 11766, A. M. N. H., Conard Fissure, Ark. Plate 13. ,11 ^ J a. ^ r b. A \ ft \ if. /. i^^ ff «'• Plate 13. Photographs, retouched, of skulls in ventral view of eight subspecies of Mustcla erminca. Natural size. a. Mnstela erminea haidarum, 9 ad., 100624, U. S. N. M., Moresby Island, B. C. b. MHKfela erminea nnyidnae, 9 ad., 13fi73, Nat. Mus. Canada, Cape Scott, B. C. r. MKstcIa ermi»ra falloidn, 9 ad., 7284, Nat. Mus. Canada, Huntingdon, B. C. d. Mnstela erminea olympica 9 ad., 242133, U. S. Nat. Mus., Hayes Creek, Wash. e. Mnstela erminea streatori, 9 ad., 9040, D. R. Dickey Coll., Blaine, Oregon. /. Mnstela erminea gulosa, 9 ad., 77370, U. S. Nat. Mus., Trout Lake, Wa.sh. g. Mnstela ertninea mnriens, 9 ad., 41502, Mus. Vert. Zool., Baker Creek, Nevada. )). Mnstela erminca angnstidens, 9 ?, ad., 12435, A. M. N. H., Conard Fissure, Ark. /. Mnstela erminea angustidens, 9 ?, ad., 11766, A. M. N. H., Conard Fissure, Ark. Plate 14. a- b. # ^^. c. d. 0^ ■A ■ t :^«^ /c. I. Plate 14. Photographs, retouched, of M. ermirtea and M. minuta. Natural size. Mustela erminea oiympica, 9 ad., 242133, U. S. Nat. Mus., Hayes Creek, Wash. Mnstela erminea streatori, 9 ad., 9040, D. R. Dickey Coll., Blaine, Oregon. Mustela erminea gulosa, 9 ad., 77370, U. S. Nat. Mus., Trout Lake, Wash. Mustela erminea muricus, 9 ad., 41502, Mus. Vert. Zool., Baker Creek, Nevada. Mustela erminea angustidens, adult, probably female, type, 12432, Amer. Mus. Nat. Hist., with lower jaw, Conard Fissure, Ark. M. e. angustidens, 9 ?, ad., 12435, A. M. N. H., Conard Fissure, Ark. Mustela rixos.a eskimo, i sad., 43288, Mus. Vert. Zool., Barrow, Alaska. Mustela rixosa eskimo, 9 sad., 40059, Mus. Vert. Zool., Barrow, Alaska. Mustela rixosa rixosa, i ad., 11743, Nat. Mus. Canada, Shaunavon, Sask. Mustela rixosa rixosa, 9 ad., 12079, Nat. Mus. Canada, south of Shaunavon, Sask. Mustela r. allegheniensis, i ad., 35381, Field M. N. H., Portage Twp., Ohio. Mnstela rixosa allegheniensis, 9 ad., 33021, Field M. N. H., Stryker, Ohio. Plate 15. Photographs, retouched, of Mustela 7-ixosa. Natural size. a. Mustela rixosa eskimo, (J sad., 43288, Mus. Vert. Zool., Barrow, Alaska. b. Mustela r-ixosa rixosa, S ad., 11743, Nat. Mus. Canada, Shaunavon, Saskatchewan. c. Mustela rixosa allegheyiiensis, i ad., 33581, Field Mus. Nat. Hist., Portage Township, Wood County, Ohio. d. Mustela rixosa campestris, $ ad., 261830, U. S. Nat. Mus., shore of Sand Lake, South Dakota. e. Mustela rixosa eskimo, $ sad., same specimen shown in a. f. Mustela rixosa rixosa, $ same specimen shown in b. g. Mustela rixosa allegheniensis, S ad., same specimen shown in c. h. Mustela rixosa campestris, $ ad., same specimen shown in d. i. Mustela rixosa eskimo, 9 sad., 40059, Mus. Vert. Zool., Barrow, Alaska. j. Mustela rixosa rixosa, 9 ad., 12679, Nat. Mus. Canada, south of Shaunavon, Saskatchewan. k. Mustela 7-ixosa allegheniensis, 9 ad., 33021, Field Mus. Nat. Hist., Stryker, Ohio. /. Mustela rixosa allegheniensis, 9 ad., same specimen shown in k. m. Mustela rixosa eskimo, 9 sad., same specimen shown in i. n. Mustela rixosa rixosa, 9 ad., same specimen shown in ;'. 0. Mustela rixosa campestris, $ ad., same specimen shown in d and h. Plate 15. ■^ a. ^ 'Of \n c a '^ ^ V 6. ^ J ^^»^ *4 ^ ,^ I*- ? (• V>.^ fl'. .'^' •«'\ A .1 V V^^ • A zc>isSo U- m. n. Plate 16. Photographs, retouched, of skulls in dorsal view of nine subspecies of Mustela frenata. Natural size. a. Mtistela frenata noveboracensis, $ ad., 77112, U. S. Nat. Mus., Wil- mington, Massachusetts. b. Mustela frenata occisor, $ ad., 7267, Mus. Comp. Zool., Moosehead Lake, Maine. c. Mttstela frenata primulina, $ ad., 3325, Mus. Nat. Hist., Univ. Kansas, Clinton, Kansas. d. Mustela frenata arthuri, $ sad., 37515, Mus. Vert. Zool., type, Remy, Louisiana. e. Mustela frenata oUvacea, S ad., 180802, U. S. Nat. Mus., type. Bio- logical Surveys Collection, Autaugaville, Alabama. /. Mustela freyiata peninsulae, $ ad., 49387, Florida State Mus., Apopka, Florida. g. Mustela frenata spadix, $ ad., 53745, Mus. Vert. Zool., Elk River, Minnesota. h. Mustela frenata longicauda, S ad., 15875, Amer. Mus. Nat. Hist., Red Deer, Alberta. i. Mustela frenata oribasa, S ad., 43817, Mus. Vert. Zool., Isaacs Lake, British Columbia. Plate IG. i > \ I >* < a. d. ^ (I 4 ^ « * V 1 %/ V ^N^ to «S) l^ o « r- , OO f1 ' o e. ■^ r^ ^1^ ^' ¥ •5 y\ (a ^ rj h W > fif. f k I r-^\ I. Plate 17. Photos, retouched, of skulls of males, in ventral view, of 9 subspecies ol Mustela frenata, x 1. Data for a to i are given on Plate 18. a. M. f. novebovacensis b. M. f. occisor c. M. f. primnlina d. M. f. arthnri e. M. f. olivacea f. M. f. peninsiilae g. M. f. spadix h. M. f. longicauda i. M. f. oribasns Plate 18. v-^-» •»-?■ " ■* '♦■^■^ ■ V^ i ^ f. ,^ ■^.,7 ff- ^ ^' '^ Plate 18. Photographs, retouched, of skulls in lateral view of ten subspecies of Miistela frenata. Natural size. a. Miistela frenata novehoracensis, S ad., 77112, U. S. Nat. Mus., Wilmington, Mass. h. Miistela frenata occisor, $ ad., 7267, M. C. Z., Moosehead Lake, Maine. c. Miistela frenata primulina, 6 ad., 3325, Mus. Nat. Hist., Univ. Kansas, Clinton, Kans. d. Mustela frenata arthuri, S ad., 37515, Mus. Vert. Zool., type, Remy, Louisiana. e. Mnstela frenata olivacea, S ad., 180802, U. S. Nat. Mus., type. Biological Surveys Collection, Autaugaville, Alabama. /. Mustela frenata peninsulae, S ad., 49387, Florida State Mus., Apopka, Florida. g. Mustela frenata spadix, $ ad., 53795, Mus. Vert. Zool., Elk River, Minnesota. h. Mustela frenata longicauda, i ad., 15875, Amer. Mus. N. H., Red Deer, Alberta. i. Mustela frenata oribasus, i ad., 43817, Mus. Vert. Zool., Isaacs Lake, B. C. j. Mustela frenata alleni, 6 ad., 7440/9136, A. M. N. H., Hill City, S. D. Plate 19. Photographs, retouched, of skulls in dorsal view of nine subspecies of Mustela frenata. Natural size. a. Mustela frenata alleni, 6 ad., 7440/9136, Amer. Mus. Nat. Hist., Hill City, South Dakota. 6. Mustela frenata arizonensis, $ ad., 55211, Mus. Vert. Zool., Govern- ment Prairie, Arizona. c. Mustela frenata nevadeyisis, i ad., 22116, Mus. Vert. Zool., Chinqua- pin, California. d. Mustela frenata effera, i ad., 33637, Amer. Mus. Nat. Hist., Ironside, Oregon. e. Mustela frenata washiyigtoni, i ad., 226758, U. S. Nat. Mus., Gotchen Creek, Washington. /. Mustela freiiata saturata, $ ad., 65930, U. S. Nat. Mus., type, Siskiyou, Oi'egon. g. Mustela frenata altifrontalis, $ ad., 391, Coll. Alex Walker, Blaine, Oregon. h. Mustela frenata oregonensis, $ sad., 43828/32019, U. S. Nat. Mus., Grants Pass, Oregon. i. Mustela frenata munda, i ad., 5459, Mus. Comp. Zool., type. Point Reyes, California. Plate 19. > 1 ^ ^-^ / .■- %. V s. 4 \ \ c. a. ^ V ■J. ' 4 >< 4i %, r s i ^ V X or /^' ■ i ^ ^ \ b. No fif- h. Plate 20. f A V a. <, A c. r. ( \ n Plate 20. Photos, retouched, of skulls of males in ventral view of 9 subspecies of Mustela frenata, x 1. Data for a to i on Plates 18 and 21. a. M. f. alleni b. M. f. arizoyiensis c. M. f. nevadensis d. M. f. effera e. M. f. washingtoni f. M. f. saturata g. M. f. altifrontalis h. M. f. oregonensis i. M. f. munda, 5459. Plate 21, a. 'msr-^ (. ' ^ >t:S: *• >^-5-« !^^' iT^^ ^.y. t--^'^r'\^^ e. ^ ^ A. f..:\^' \ ^j^' **^^ ■s^.^ / '^ ^ V, ; '/;? ' \ M i •^' Plate 23. Ventral views of same skulls shown in Plate 22. a. munda; b. xanthogenys; c. nigriauris; d. latirostra; e. pulchra; f. inyoensis; g. neomexicanus; h. texensis; i. frenata. Plate 24. **^':^V|g^ «»? i^^ /0>\^y^ 4 f. I ,* i 1*^*^ »Jfc. «*.J Plate 24. Photographs, retouched, of skulls, in lateral view, of ten subspecies of Mustela frenata. Natural size. a. Mustela frenata nigriauris, $ ad., 487, Stanford Univ., Palo Alto, California. b. Mustela frenata latirostra, $ ad., 52702, U. S. Nat. Mus., El Cajon, California. c. Mustela frenata pulchra, $ ad., 16668, Mus. Vert. Zool., type, Buttonwillovr, Calif. d. Mustela frenata inyoensis, $ ad., 25907, Mus. Vert. Zool., type, 2 mi. N Independ- ence, California. e. Mustela frenata neomexicana, $ ad., 1485, M. N. H., Univ. Kansas, Liberal, Kansas. /. Mustela frenata texensis, $ ad., 14821, A. M. N. H., Kerr County, Texas. g. Mxistela frenata frenata, $ ad., 50826, U. S. Nat. Mus., Tlalpam, Mexico, D. F. h. Mustela frenata leucoparia, 6 ad., 125972, U. S. N. M., Los Reyes, Michoacan. i. Mustela frenata macrophonius, $ ad.. Field Mus. N. H., type, Achotal, Veracruz. J. Mustela frenata goldmani, S ad., 77519, U. S. Nat. Mus., Pinabete, Veracruz. Plate 25. Photographs, retouched, of skulls, in dorsal view of nine subspecies of Mustela frenata. Natural Size. a. Mustela frenata leucoparia, $ ad., 125972, U. S. Nat. Mus., Los Reyes, Michoacan. b. Mustela frenata macrophonius, S ad., 14063, Field Mus. Nat. Hist., type, Achotal, Veracruz. c. Mustela frenata goldmani, $ ad., 133253, U. S. Nat. Mus., 20 mi. SE Teopisca, Chiapas. d. Mustela frenata tropicalis, 6 ad., 54994, U. S. Nat. Mus., type, Jico, Veracruz. e. Mustela frenata perda, $ sad., 100041, U. S. Nat. Mus., type, Teapa, Tabasco. /. Mustela frenata nicaraguae, $ sad., 30754, Amer. Mus. Nat. Hist., type, Matagalpa, Nicaragua. g. Mustela frenata costaricensis, S ad., 3.2.1.6., British Mus. Nat. Hist., San Jose, Costa Rica. h. Mustela frenata panamensis, S ad., 18848, Amer. Mus. Nat Hist., Boquete, Panama. i. Mustela frenata meridana, 6 ad., 123341, U. S. Nat. Mus., type, Merida, Venezuela. Plate 25. im i'f 4 IK J"^ ^!V^ 4 #\ V. • v "I a. b. is 4 ^ f'- d. 0h 4 ^ t •^ ' t e. Mir fe. , -c r '^^0£ /. (/' / Plate 26. ^- I*: t ft' V ^ ^ iSj 'i ■far.. f Plate 26. Ventral views of same skulls shown in Plate 25. a. leucoparia; b. macrophonius ; c. goldmani; d. tropicalis; e. perda; f. nicaraguae; g. costaricensis ; h. panarnensis; i. meridana. Plate 27. a. c. ( \-^ /• * ,m%. J^ % .-^,» J Plate 27. Photographs, retouched, of skulls, in lateral view of ten subspecies of Mustela frenata. Natural size. a. Mustela frenata tropicalis, S ad., 54994, U. S. Nat. Mus., type, Jico, Veracruz. b. Mustela frenata perda, S sad., 100041; U. S. Nat. Mus., type, Teapa, Tabasco. c. Mustela frenata nicaraguae, $ sad., 30754, Amer. Mus. Nat. Hist., type, Matagalpa, Nicaragua. d. Mustela frenata costaricensis, $ ad., 3.2.1.6., British Mus. Nat. Hist., San Jose, Costa Rica. e. Mustela frenata panamensis, S ad., 18848, Amer. Mus. Nat. Hist., Boquete, Panama. /. Mustela frenata meridana, $ ad., 24309, Amer. Mus. Nat. Hist., Merida, Venezuela. g. Mustela frenata aureoventris, 6 yg-, 34677, Amer. Mus. Nat. Hist., Gualea, Ecuador. h. Mustela frenata helleri, S ad., 24133, Field Mus. Nat. Hist., type, Rio Chinchao, Peru. i. Mustela frenata macrura, S ad., 561, Mus. Polonais d' Hist., Nat., type, Junin, Perii. j. Mustela frenata agilis, $ ad., 8.1.10.1., British Mus. Nat. Hist., Lima, Peru. Plate 28. Photographs, retouched (except /) , of skulls, in dorsal view of nine kinds (species and subspecies) of Mustela. Natural size. a. Mustela frenata aureoventris, S yg-, 34677, Amer. Mus. Nat. Hist., Gualea, Ecuador. b. Mustela frenata helleri, $ ad.^ 24133, Field Mus. Nat. Hist., type, Rio Chinchao, Peru. c. Mustela frenata macrura, $ ad., 561, Mus. Polonais d' Hist. Nat., type, Junin, Peru. d. Mustela frenata agilis, S ad., 8.1.10.1., British Mus. Nat. Hist., Lima, Peru. e. Mustela frenata boliviensis, S ad., 72587, Amer. Mus. Nat. Hist., type, Nequejahuira, Bolivia. /. Mustela frenata xanthogenys, S ad., 43.6.4.55., British Mus., Nat. Hist., type, California. g. Mustela frenata costaricensis, £ yg., 37149, U. S. Nat. Mus., type, San Jose, Costa Rica. h. Mustela frenata panamensis, $ yg., 178970, U. S. Nat. Mus., Mt. Pirre, Panama. i. Mustela africana africana, $ yg., 37475, Amer. Mus. Nat. Hist., Para, Brazil. Plate 28. 41^ /^ \ X %. V^ ^ a. 4 . -fV « ^, nr %ttl^ % *4 r e. ># ^ V % ^^«s o> li \ "^ \ M^ Plate 29. Photographs, retouched, of skulls, in ventral view, of nine kinds (species and subspecies) of Mustela. Natural size. a. Mustela frenata aureoventris, 6 yg-, 34677, Amer. Mus. Nat. Hist., Gualea, Ecuador. b. Mustela frenata helleri, $ ad., 24133, Field Mus. Nat. Hist., type., Rio Chinchao, Peru. c. Mustela frenata macrura, $ ad., 561, Mus. Polonais d' Hist. Nat., type, Junin, Peru. d. Mustela frenata agilis, $ ad., 8.1.]'^.!., British Mus. Nat. Hist., Lima, Peru. e. Mustela frenata boliviensis, S ad., 72587, Amer. Mus. Nat. Hist., type, Nequejahuira, Bolivia. /. Mustela frenata leucoparia, 6 ad., 47179/34914, U. S. Nat. Mus., type, Patzcuaro, Michoacan. g. Mustela frenata costaricensis, $ yg., 37149, U. S. Nat. Mus., type, San Jose, Costa Rica. h. Mustela frenata panamensis, S yg., 17897-0, U. S. Nat. Mus., Mt. Pirre, Panama. i. Mustela africana africana, 6 yg., 37475, Amer. Mus. Nat. Hist., Para, Brazil. Plate 29. V*^^^V " > f 4 ^5.^ M >' ^^ / I w^ ( /! V ' \ 9- Plate 30. Photographs, retouched (except e and q) of skulls and lower jaws of Mnstela. a. Mustela frenata bolivensis, $ ad., 72587, Amer. Mus. Nat. Hist., type, Nequejahuira, Bolivia. b. Mustela frenata xanthogenys, S ad., 43.4.6.55, British Mus. Nat. Hist., type, California. c. Mustela frenata costaricensis, S yg., 37149, U. S. Nat. Mus., type, San Jose, Costa Rica. d. Mustela frenata panamensis, $ yg., 178970, U. S. Nat. Mus., Mt. Pirre, Panama. e. Mustela frenata affinis, S ad., 54.6.3.4, British Mus. Nat. Hist., type, New Granada [ = Colombia]. /. Mustela africana africana, $ yg., 37475, Amer. Mus. Nat. Hist., Para, Brazil. g. Mustela frenata saturata, <5 ad., 65930, U. S. Nat. Mus., type, Siski- you, Oregon. h. Mustela frenata oregonensis, S ad., 43828/32019, U. S. Nat. Mus., type, Grants Pass, Oregon. i. Mustela frenata miaida, S ad., 5459, Mus. Comp. Zool., type, Point Reyes, California. j. Mustela frenata leucoparia, $ ad., 47179/34914, U. S. Nat. Mus., type, Patzcuaro, Michoacan. k. Mustela frenata macrophonius, i ad., 14963, Field Mus. Nat. Hist., type, Achotal, Veracruz. I. Mustela frenata goldmani, $ ad., 77519, U. S. Nat. Mus., type, Pina- bete, Chiapas. m. Mustela frenata tropicalis, S ad., 54994, U. S. Nat. Mus., type, Jico, Veracruz. n. Mustela frenata perda, $ sad., 100041, U. S. Nat. Mus., type, Teapa, Tabasco. o. Mustela frenata nicaraguae, S sad., 30754, Amer. Mus. Nat. Hist., type, Matagalpa, Nicaragua. p. Mustela frenata costaricensis, S yg., 37149, U. S. Nat. Mus., type, San Jose, Costa Rica. q. Mustela frenata affinis, S ad., 54.6.3.4, British Mus. Nat. Hist., type. New Granada [ = Colombia]. r. Mustela frenata ynacrura, 6 ad., 561., Mus. Polonais d' Hist. Nat., type, Junin, Peru. Plate 30. .1^ a. 4 c. d. ^ r 9- h. i. l. 4: m. 0. r. Plate 31. '^ >% m\ ^ i vc96^» ^f* « » V Oj 9- ^,#.-^ '<', 1.1 Plate 31. Photos, retouched, of skulls of 9 subspecies of M. freiiata, x 1. a. Mustela frenata noveboracensis, 9 ad., 64087, U. S. N. M., Wilmington, Mass. b. Mustela frenata occisor, 9 ad., 9101, Mus. Comp. Zool., Bucksport, Maine. c. Mustela frenata prhnnlina, 9 ad., 3G38, U. K. M. N. H., 7 mi. SW Lawrence, Kans. d. Mustela frenata olivacea, 9 ad., 41024, Mus. Vert. Zool., Sinkola Plantation, Ga. e. Mustela frenata spadix, 9 ad., 188410, U. S. Nat. Mus., Elk River, Minn. /. Mustela frenata longicauda, 9 ad., 75483, U. S. Nat. Mus., Wingard, Sask. g. Mustela frenata oribasus, 9 ad., 9058, M. C. Z., type, source of Kettle River, B. C. h. Mustela frenata alleyii, 9 ?, ad., 7441, A. M. N. H., Black Hills, S. D. I. Mustela frenata arizonensis, 9 ad., 188G, A. M. N. H., type, S. F. Forest, Ariz. Plate 32. a. J-^% 9- t. Plate a. Mustela b. Mustela c. Mustela d. M^istela e. Mustela f. Mustela g. Mustela h. Mustela i. Mustela 32. Photos, retouched, of skulls of 9 subspecies of M. frenata, x 1. frenata noveboracensis, 9 ad., 64687, U. S. N. M., Wilmington, Mass. frenata occisor, 9 ad., 9101, Mus. Comp. Zool., Bucksport, Maine. frenata prinmlma, 9 ad., 3638, U. K. M. N. H., 7 mi. SW Lawrence, Kans. frenata olivacea, 9 ad., 41024, Mus. Vert. Zool., Sinkola Plantation, Ga. frenata spadix, 9 ad., 188410, U. S. Nat. Mus., Elk River, Minn. /. longicauda, 9 ad., 75483, U. S. Nat. Mus., Wingard, Sask. frenata oribasus, 9 ad., 9058, M. C. Z., type, source of Kettle Riv., B. C. frenata alleni, 9 ?, ad., 7441, A. M. N. H., Black Hills, S. D. frenata arizonensis, 9 ad., 1886, A. M. N. H., type, S. F. Forest, Ariz. Plate 33. eviisy ^f^;*^^^'^ 1: c. ^^ ^f^^S^^"^ e. 1^ ■,^'*Ka ..40*v flf. ,^^^,v . ^. .aX. ■* Plate 33. Photographs, retouched, of skulls in lateral view of ten subspecies of Mustela frenata. Natural size. a. Mustela frenata ywveboracensis, 9 ad., 64687, U. S. N. M., Wilmington, Mass. b. Mustela frenata occisor, 9 ad, 9101, Mus. Comp. Zool., Bucksport, Maine. c. Mustela frenata primulina, 9 ad., 3638, Univ. Kansas Mus. Nat. Hist., 7 mi. SW Lawrence, Kansas. d. Mustela frenata olivacea, 9 ad., 41024, Mus. Vert. Zool., Sinkola Plantation, Ga. e. Mustela frenata spadix, 9 ad., 188410 (2196), U. S. Nat. Mus., Elk River, Minn. /. Mustela frenata lungicauda, 9 ad., 75483, U. S. Nat. Mus., Wingard, Sask. g. Mustela frenata oribas7is, 9 ad., 9058, Mus. Comp. Zool., type, source of Kettle River, British Columbia. /(. Mustela frenata alleni, 9 ?, ad., 7441, Amer. Mus. N. H., Black Hills, S. D. /. Mustela frenata arizonensis, 9 ad., 1886, Amer. Mus. Nat. Hist., type, San Francisco, Forest, Arizona. ;. Mustela frenata nevadensis, 9 ad., 41503, M. V. Z., type, 3 mi. E Baker, Nev. Plate 34. 1 4 y i. >. ( ^ ^ On K ^ J A > 4 I •3 NT /. M ^ Plate 34. Photographs, retouched, of skulls in dorsal view, of eight subspecies of Mustela frenata. Natural size. a. Mustela frenata nevadensis, $ ad., 41503, M. V. Z., type, 3 mi. E Baker, Nevada. 6. Mustela frenata washingtoni, 9 sad., 81953, U. S. N. M., Trout Lake, Wash. c. Mustela frenata altifrontalis, 9 ad., 392, coll. of Alex Walker, Blaine, Oregon. d. Mustela frenata oregonensis, 9 ad., 244520, U. S. Nat. Mus., Medford, Oregon. e. Mustela frenata munda, 9 ad., 91764, U. S. Nat. Mus., Point Reyes, California. /. Mustela frenata munda, 9 ad., 19723, Mus. Vert. Zool., Point Arena, California. g. Mustela frenata xanthogenys, 9 ad., 2626, coll. of W. E. Snyder, Selma, California. h. Mustela frenata nigriauris, 9 ad., 3761, M. V. Z., San Francisco, California. i. Mustela frenata neomexicana, 9 ad., 36482, U. S. N. M., Tombstone, Arizona. / A JUh, Plate 35. a. h. I. Plate 35. Photoe:raphs, retouched, of skulls in ventral view, of eight subspecies of Mustela frenata. Natural size. a. Mtistela frenata nevadensis, 9 ad., 41503, M. V. Z., type, 3 mi. E Baker, Nevada. h. Mustela frenata ivashingtoni, 9 sad., 81953, U. S. N. M., Trout Lake, Wash. c. Mustela frenata altifrontalis, 9 ad., 392, coll. of Alex Walker, Blaine, Oregon. d. Miistela frenata oregonensis, 9 ad., 244520, U. S. Nat. Mus., Medford, Oregon. e. Mustela frenata munda, 9 ad., 91764, U. S. N. M., Point Reyes, California. /. Mustela frenata munda, 9 ad., 19723, M. V. Z., Point Arena, California. g. Mustela frenata xanthogenys, 9 ad, 2626, coll. of W. E. Snyder, Selma, Calif. h. Mustela frenata nigriauris, 9 ad., 3761, M. V. Z., San Francisco, California. i. Mustela frenata neomexicana, 9 ad., 36482, U. S. N. M., Tombstone, Arizona. Plate 36. ^^^iJ* ^v*"-**** ^^'^^^^i^^ ^/ d. ^^tw' i^^^ h. ^:-i^ » — A A Plate 36. Photographs, retouched, of skulls in lateral view, of eight subspecies of Mustela frenata. Natural size. a. Mustela frenata washingtoni, 9 sad., 81953, U. S. N. M., Trout Lake, Wash. h. Miistela frenata altifrontalis, 9 ad., 392, coll. of Alex Walker, Blaine, Oregon. c. Mustela frenata oregonensis, 9 ad., 244520, U. S. N. M., Medford, Oregon. d. Mustela frenata munda, 9 ad., 91764, U. S. N. M., Point Reyes, California. e. Mustela frenata munda, 9 ad., 19723, Mus. Vert. Zool., Point Arena, Calif. 9 ad., 2626, coll. of W. E. Snyder, Selma, Calif, ad., 3761, Mus. Vert. Zool., San Francisco, Calif. 9 ad., 36482, U. S. N. M., Tombstone, Ariz. Brownsville, Texas, type, Mexico City, D. F. Mustela frenata munda, 9 ad, /. Mustela frenata xanthogenys, g. Mustela frenata nigriauris, 9 Mustela frenata neomexicana, Mtistela frenata frenata, 9 ad., 58685, U. S. Nat. Mus Mustela frenata frenata, 9 ad., 991, Berlin Zool.. Mus. h. i. 3- k. I Mustela frenata leucoparia, 9 ad., 26153, Amer. Mus. N. H., Artenkiki, Jalisco. Mustela frenata perotae, 9 ad., 54278, U. S. Nat. Mus., type, 12500 ft., Cofre de Perota, Veracruz. Plate 37. 4 ^ 4 ^ 1 f\ *" 'v \^ ^' 1 r,m_^ Mi \i^^' a. b. c. ^^ I 'r^^^ .-?N. * V V. V 1 d. e. M\ f. f^ ^. * ^ ^ ^ M >^ ',#«*■' 9. 9^ h. Plate 37. Photos, retouched, of skulls of 8 subspecies of Mustela frenata, x 1. a. Mustela frenata fretmta, 9 ad., 58685, U. S. Nat. Mus., Brownsville, Texas. b. Mustela frenata frenata, 9 ad., 991, Berlin Zool. Mus., type, Mexico City, D. F. c. Mustela frenata leucoparia, 9 ad., 26153, Amer. Mus. Nat. Hist., Artenkiki, Jalisco. d. Mustela f. perotae, 9 ad., 54278, U. S. N. M., type, Cofre de Perota, Veracruz. e. Mustela frenata macrophonius, 9 ad., 132528, U. S. Nat. Mus., Perez, Veracruz. /. Mustela frenata tropicalis, 9 ad., 54993, U. S. Nat. Mus., Jico, Veracruz. g. Mustela frenata perda, 9 sad., 65422, U. S. Nat. Mus., Catemaco, Veracruz. h. Mustela frenata meridana, 9 ad., 143665, U. S. N. M., Merida, Venezuela. i. Mustela f. macrura, $ ad., 564, M. P. H. N., type of Mustela jelskii, Cutervo, Peru. V t A- a. t ♦. t "I d. %r ^ * A J % \ \x 1 'W*;. /* f ■ ■ Plate 38. . ^ "*!' ^ >, ? /. Plate 38. Photos, retouched, skulls in ventral view, 8 subspecies of M. fvenata, x 1. a. Mustela frenata frenata, 9 ad., 58685, U. S. Nat. Mus., Brownsville, Texas. b. Mustela frenata frenata, 9 ad., 991, Berlin Zool. Mus., type, Mexico City, D. F. c. Mustela frenata teucoparia, 9 ad., 26153, Amer. Mus. Nat. Hist., Artenkiki, Jalisco. d. Mustela f. perotae, 9 ad., 54278, U. S. N. M., type, Cofre de Perota, Veracruz. e. Mustela frenata macrophonius, 9 ad., 132528, U. S. Nat. Mus., Perez, Veracruz. /. Mustela frenata tropicalis, 9 ad., 54993, U. S. Nat. Mus., Jico, Veracruz. g. Mustela frenata perda, 9 sad., 65422, U. S. Nat. Mus., Catemaco, Veracruz. h. Mustela frenata meridana, 9 ad., 143665, U. S. N. M., Merida, Venezuela. i. Mustela f. macrura, 9 ad., 564, M. P. H. N., type of Mustela jelskii, Cutervo, Peru. 0L a. 'niP^ Plate 39. b. -r X- ^f^**" X.^ 0k "^^^ii^ V d. .€^ Li* ',m , r. ^ 5-*t. ^ »•«*" ^^^«*v.^ ""%fc^^ Plate 39. Figs. a-h. Photos, retouched, of 10 kinds Mustela, x 1. a. Mustela frenata macrophonius, 9 ad., 132528, U. S. Nat. Mus., Perez, Veracruz. b. Mustela frenata tropicalis, 9 ad., 54993, U. S. Nat. Mus., Jico, Veracruz. c. Mustela frenata perda, 9 sad., 65422, U. S. Nat. Mus., Catemaco, Veracruz. d. Mustela frenata meridana, 9 ad., 143665, U. S. Nat. Mus., Merida, Venezuela. e. Mustela f. macrura, 9 ad., 564, M. P. H. N., type of Mustela jelskii, Cutervo, Peru. /. Mustela frenata agilis, 9 sad., 21147, Field Mus. Nat. Hist., Macate, Peru. g. Mustela frenata gracilis, 9 ?, ad., 12431, Amer. Mus. Nat. Hist., type, Conard Fissure, Arkansas, Pleistocene. h. Mustela a. stolzmanni, 9 sad., 24.12.12.24, Brit. M. N. H., Myobamba, Peru. Figs. i-l. Mustela frenata nevadensis, all males, from Colorado, showing seasonal change in color, x approximately 1/9. Note the sharply marked molt line in the pelage of spring and the absence of any definite molt line in autumn. i. No. 151415 U. S. Nat. Mus., Coventry, December 27, 1907. ;. No. 202741 U. S. Nat. Mus., Pierce Place, April 18, 1913. k. No. 201681 U. S. Nat. Mus., Jefferson, June 23, 1913. /. No. 41997 Amer. Mus. Nat. Hist., Navajo River, October 29, 1913. Plate 40. |k t a. % IS % I ,='%.■ .:*% ^^^'^riiss/ /• •>/Y<»^ /i. /,-. /. Plate 40. Photos, retouched, of skulls and lower jaws of Mnstcla, x 1. Mustela frenata agilis, 9 sad., 21147, Field Mus. Nat. Hist., Macate, Peru. Mustela frenata gracilis, 9 ?, ad., type, 12431, A. M. N. H., Conard Fiss., Ark. Mustela africana stolzmanni, 9 sad., 24.12.12.24, Brit. M. N. H., Myobamba, Peru Mustela frenata agilis, 9 sad., 21147, Field Mus. Nat. Hist., Macate, Peru. Mustela frenata gracilis, 9 ?, ad., type, 12431, A. M. N. H., Conard Fiss., Ark. Mustela africana stolzmanni, 9 sad., 24.12.12.24, Brit. M. N. H., Myobamba, Peru. Mustela africana stolzmanni, 9 ad., 503, Mus. Polonais d' Hist. Nat., type, Yuri- maguas, Peru. The palate is broken longitudinally and the two maxillae are slightly out of normal position. Mustela frenata oribasus, 9 ad., 9058, M. C. Z., type, source of Kettle Riv., B. C. Mustela frenata munda, 9 ad., 917(54, U. S. Nat. Mus., Point Reyes, Calif. Mustela frenata frenata, 9 ad., 991, Berlin Z. M., type, Mexico City, I). F. Mustela f. macrura, 9 ad., 504, Mus. Polonais d' Hist. Nat., type of Mustela jehkii. Mustela africana stolzmanni, 9 ad., 503, Mus. Polonais d' Hist. Nat., type, Yuri- maguas, Peru. Right half of lower jaw reversed. Plate 41 A 1^ i I MUSTELA ERMINEA ARCTICA ♦ f I I MUSTELA FRENATA MUSTELA ERMINEA INVICTA MUSTELA RIXOSA MUSTELA ERMINEA MURICUS MUSTELA AFRICANA Plate 41. Photographs, approximately }i, of stuffed study-skins of the four species of American weasels. For each pair the male is at the left and the female at the right. Photo, by W. C. Matthews. Mustela erminea arctica, both in U.S.B.S., from Mts. near Eagle, Alaska, $ 131256 and 5 131245. Mustela erminea invicta, both in U.S.B.S., from Tungsten Mine, Wasliing- ton, S 235236 and $ 235235. Mustela errninea muricus, both in M.V.Z., from Baker Creek, 8675 ft., Ne- vada, S 41501 and ? 41502. Mustela frenata nigriauris, both in M.V.Z., from Cahfomia, $ 51666 from Concord and $ 73109 from Berkeley. Mustela africana africana, $ 37475 A.M.N.H., from Para, Brazil. Mustela rixosa allcgheniensis, both in M.Z.U.M., from Michigan, 5 83260 Swan Creek Farm and $ 88079 from Unadilla. (459) 16—3758 INDEX TO VOLUME 4 Principal references are in boldface type aequatoriiilis, Mustela, 75 Putorius, 75, 341, 387 aestuarina, Mustela, 82 affinis, Mustela, 75, 375, 379, 384, 398, 409 Putorius, 372, 375, 379, 384 africana, Mustela, 73, 406, 409 agilis, Mustela, 74, 393 Putorius, 74, 222 alascensis, Mustela, 75, 131 Putorius, 75, 131 albigula, Neotoma, 208 allegheniensis, Mustela, 77, 187 Putorius, 77, 187 3.1lGli Nlustela, 76, 274 Putorius, 76, 274 altifrontalis, Mustela, 79, 300 americana, 74, 75 americanus, Lepus, 93, 201,210, 212,216 anguinae, Mustela, 79, 145 angustidens, Mustela, 78, 165 Putorius, 78, 165 Annelida, 93 arctica, Mustela, 76, 96 arcticus, Putorius, 76 arizonensis, Mustela, 75, 276, 280, 291. 323 Putorius, 75, 276, 280 annatus, Citellus, 200 artliuri, Mustela, 78, 241 audax, Mustela, 77 Putorius, 77 aureoventris, Mustela, 74, 387, 398 Aves, 93 bachmani, Sylvilagus, 213 bangsi, Mustela, 80, 124 barn owl, 173 Belding ground squirrel, 205 beldingi, Citellus, 205 big jumping mouse, 210 birds, small, 216 wild, 93, 213 Blarina, 91, 205, 209, 210 brevicauda, 205 bobwhite, 213 boccamela, Mustela, 170 boliviensis, Mustela, 79, 402 bottae, Thomomys, 207 boylii, Lampropeltis, 213 brasiliensis, Mustela, 73, 341, 372, 375 Putorius, 280, 300, 315, 319, 323, 384, 387, 398, 409 brevicauda, Blarina, 91, 205 brush rabbit, 213 bull snake, 213 Bunaelurus, 11 campestris, Mustela, 78, 190 cat, domestic, 174 celenda, Mustela, 80, 139 chicken, 93, 213, 216 chipmunk, 92, 196, 201, 206, 210, 216 cicognanii, Mustela, 73, 110, 118, 124, 222 Putorius, 118, 124, 128, 145, 155, 161 cinereus, Sorex, 205 Citellus, 216 armatus, 200 beldingi, 205 franklini, 203 lateralis, 206 richardsonii, 205 townsendii, 205 tridecemlineatus, 205 Clethrionomys, 92 gapped, 175, 181 Colaptes, 213 comadreja, 7 Condylura, 210 costaricensis, Mustela, 78, 372, 379, 387 cotton rat, 208 cottontail, 202, 203, 205, 212 Meams, 208 coyote, 200 Cratogcomys, 65, 207 culbertsoni, Mustela, 75 Putorius, 75 deer mouse, 208, 209, 216 domestic cat, 174 donnola, 7 druinmondi, Microtus, 175 — Univ. Kansas Publ., Mus. Nat. Hist., Voi.. 4, 1951. (461) 462 I>fDEX earthworm, 93 effera, Mustek, 79, 291 energumenos, Mustela, 82 ermine, 87 erminea, Mustela, 71 Putorius, 222 eskimo, Mustela, 77, 181 Putorius, 77, 181 evagor, Mustela, 82 evergladensis, Mustela, 82 Evotomys, 175 fallenda, Mustela, 80, 148 Felis, 9 ferrets, 43 fish, 91, 93 flickers, 213 floridana, Neotoma, 208 flying squirrel, 206, 216 Franklin ground squirrel, 205 franklini, Citellus, 205 frenata, Mustela, 73, 232, 252, 280, 309. 338, 341 frenatus, Putorius, 280, 300, 315, 319, 323, 341, 351, 363, 366, 372, 384, 398 frog, leopard, 93 fulvus, Vulpes, 200 furo, Mustela, 43 fusca, Mustela, 222 fuscipes, Neotoma, 208 fuscus, Putorius, 74, 222 Gallus, 93, 213 gapperi, Clethrionomys, 175, 181 Geomyidae, 216 getulus, Lampropeltis, 213 Glaucomys, 206, 216 golden-mantled ground squirrel, 206 goldmani, Mustela, 76, 355 Putorius, 76, 355 gracilis, Mustela, 78, 404 Putorius, 78,404 Granmogale, 407 grasshopper, 208 grasshopper mouse, 207, 216 great-horned owl, 173 ground squirrel, 210, 216 Belding, 205 Franklin, 205 golden-mantled, 206 Richardson, 205 thirteen-lined, 205 Townsend, 205 gulosa, Mustela, 80, 159 haidarum, Mustela, 76, 142 Putorius, 76, 142 hare, varying, 210, 216 harvest mouse, 208, 216 helleri, Mustela, 79, 391 hispidus, Sigmodon, 208 horned lark, 209 house mouse, 216 hyemalis, Junco, 213 imperii, Putorius, 77, 110 ingens, Mustela, 82 initis, Mustela, 80, 136 insects, 176, 209, 216 intergrades, 45 invicta, Mustela, 80, 128 inyoensis, Mustela, 79, 331 javonica, Mustela, 72 jelskii, Mustela, 75, 398 jumping mouse, 216 big, 210 Junco hyemalis, 213 kadiacensis, Mustela, 76, 108 Putorius, 76, 108 king-snake, 213 labiata, Mustela, 79 lacustris, Mustela, 82 Lampropeltis, boylii, 213 getulus, 213 lark, horned, 209 lateralis, Citellus, 205 latimanus, Scapanus, 205 latirostra, Mustela, 79, 323 least weasel, 168, 209 lemming, 92 Lemmus, 92 Leopard frog, 93 lepida, Neotoma, 208 lepta, Mustela, 161 leptus, Mustela, 161 Putorius, 78, 161 Lepus americanus, 93, 201, 210, 211, 212,216 letifcra, Mustela, 82 leucoparia, Mustela, 76, 347 Putorius, 76, 347 lizards, 216 longicauda, Mustela, 73, 232, 252, 262 Putorius, 280 long-tailed weasel, 193 lutensis, Mustela, 82 Lutra minor, 170 Lutreola, 84 lutreola. Mustela, 170 Lyncodon, 407 Index 463 macrodon, Mustek, 82 macrophonius, Mustela, 78, 360 Putorius, 78, 360 macrura, Mustela, 75, 387, 393, 398, 402 macrurus, Putorius, 379 maniculatus, Peromyscus, 175 Martinogaie, 11 meadow mice, 92, 208, 209, 216 Mearns cottontail, 208 melampelus, Mustela, 82 melodia, Melospiza, 213 Melospiza melodia, 213 meridana, Mustela, 78, 379 Mexican pocket gopher, 207 mexicanus, Putorius, 341 mice, meadow, 92, 208, 209, 216 microtis, Mustela, 110 Putorius. 77, 110 Microtus, 65, 91, 92, 174, 179, 181, 208,209,210,216,220 drummondi, 175 minor, 175 montanus, 209 ochrogaster, 179 pennsylvanicus, 175 minimus, Tamias, 206 mink, Mustela, 82 minor, Lutra, 170 Mustela, 169 minor, Zapus, 210 minuta, Mustela, 169 Palaeogale, 169 minutus, Putorius, 169 Miomustela, 11 mole, 210 moles, 216 montanus, Microtus, 209 mortigena, Mustela, 110 mouse, deer. 208, 209, 216 grasshopper, 207, 216 harvest. 208. 216 house, 208, 216 jumping, 210 meadow, 208, 209, 216 red-backed, 92 munda, Mustela, 77. 304, 309 mundus. Putorius, 77, 309 murica, Mustela, 161 muricus, Mustela, 77, 161 Putorius, 77, 161 Mus, 216 muskrat, 216 Mustela, 83 aequatorialis, 75 aestuarina, 82 affinis, 75, 375, 379, 384, 398, 409 africana, 73, 406, 409 agilis, 74, 393 alascensis, 75, 131 allegheniensis, 77, 187 alleni, 76, 274 altifrontalis, 79, 300 americana, 74, 75 anguinac, 79, 145 angustidens, 78, 165 arctica, 76, 96 arizonensis, 75, 276, 280, 291, 323 arthuri, 78, 241 audax, 77 aureoventris, 74, 387, 398 bangsi, 80, 124 boccamela, 170 boliviensis, 79, 402 brasiliensis, 73, 341, 372, 375 campestris, 78, 190 celenda, 80, 139 cicognanii, 73, 118,222 cigognanii, 118 costaricensis, 78, 372, 379, 387 culbertsoni, 75 digna, 100 effera, 79, 291 energumenos, 82 erminea, 72, 87, 103 eskimo, 77, 181 evagor, 82 evergladensis, 82 fallenda, 80, 148 furo, 43 frenata, 73, 193, 232, 252, 280, 309, 338, 341 fusca, 224 goldmani, 76, 355 gracilis, 78, 404 gulosa, 80, 159 haidarum, 76, 142 helleri, 79, 391 ingens, 82 initis, 80. 136 invicta, 80, 128 inyoensis, 79, 331 javonica, 72 jelskii, 75, 39^8 kadiacensis, 76, 108 kaneii, 99 labiata, 79, 105 lacustris, 82 latirostra, 79, 323 lepta, 161 leptus, 161 letifera, 82 leucoparia, 76, 347 longicauda, 73, 232, 252, 262 lutcnsis, 82 lutreola, 170 macrodon, 82 macrophonius, 78, 360 464 Index macrura, 75, 387, 393, 398, 402 . melampelus, 82 meridana, 78, 379 microtus, 77, 110 mink, 82 minor, 169 minuta, 169 munda, 77, 304, 309 murica, 161 muricus, 77, 161 neomexicana, 76, 333 neomexicanus, 333 nesolestes, 82 nevadensis, 79, 280 nicaraguae, 78, 370 nigriauris, 79, 319 nigripes, 74 nivalis, 72 notius, 77 noveboracensis, 74, 222, 252 numidica, 170 occisor, 77, 230 olivacea, 78, 244 olympica, 80, 153 oregonensis, 76, 304 oribasa, 270 oribasus, 77, 270 orientalis, 100 panamensis, 78, 375 paraensis, 76, 409 peninsulae, 75, 250 perda, 77, 366^ perotae, 79, 351 polaris, 77, 103 primulina, 78, 232 pulchra, 79, 328 pusilla, 74, 118 putorius, 43 richardsonii, 73, 110 rixosa, 76, 153, 155, 161, 168, 184 salva, 80, 135 saturata, 76, 297 seclusa, 80, 141 semplei, 78. 105 spadix, 76, 252 stohmanni, 75, 409, 413 streatori 76, 155 texensis, 79. 338 tropicalis, 76, 363, 367 vison, 82 NTilgivaga, 82 washingtoni, 76, 294 xanthogenys, 74, 315, 331 neomexicana, Mustela, 76, 333 neomexicanus, Mustela, 333 Putorius, 76, 333 Neotamias, 206 Neotoma, 208, 216 albigula, 208 floridana, 208 fuscipes, 208 lepida, 208 nesolestes, Mustela, 82 nevadensis, Mustela, 79, 280 nicaraguae, Mustela, 78, 370 nigriaiuris, Klustela, 79, 319 nigripes, Mustela, 74 Putorius, 74 nivalis, Mustela, 72, 169 northern pocket gopher, 206 notius, Putorius, 77, 222 noveboracensis, Mustela, 74, 222, 252 Putorius, 74, 222 numidica, Mustela, 170 numidicus, Putorius, 170 occisor, Mustela, 77, 230 Putorius, 77 Ochotona, 92 ochrogaster, Microtus, 179 olivacea, Mustela, 78, 244 olympica, Mustela, 80, 153 Ondatra, 210, 216 Onychomys, 207, 216 oregonensis, Mustela, 304 Putorius, 304 oribasa, Mustela, 270 oribasus, Mustela, 77, 270 Putorius, 77 Orthogeomys, 65 owl, bam, 173 great-horned, 173 snowy, 90 Palaeogale, 11 minuta, 169 panamensis, Mustela, 78, 375 paraensis, Mustela, 76, 409 Putorius, 76, 409 peninsulae, Mustela, 75, 250 Putorius, 75, 250 pennsylvanicus, Microtus, 175 perda, Mustela, 77, 366 perdus, Putorius. 77, 366 Peromvscus, 91, 92, 173, 196, 208, 209,210,216 maniculatus, 175 perotae, Mustela, 79, 351 pheasant, 209 ring-necked, 202 pin:my weasel, 216 pika, 92 pikas, 201 pipiens, Rana, 93 Index 465 Pisces, 93 Pituophis sayi sayi, 213 Pliogale, 11 pocket gopher, 209, 216 Mexican, 207 northern, 206 Shaw, 208 polaris, Mustek, 77, 103 Putorius, 77 polecats, 43 porcupine, quills of, 200 primulina, Mustela, 78, 233 pulchra, Mustela. 79, 323 pusilla, Mustela, 74, 118 pusillus, Putorius, 118, 155, 184, 190 Putorius, 76, 294, 300 aequatorialis, 75, 341, 387 affinis, 372, 375, 379, 384 agilis, 74, 222 alascensis, 75, 96, 131 allegheniensis, 77, 187 alleni, 76, 274 angustidens, 76, 165 arcticus, 76, 96 arizonensis, 75, 276, 280 audax, 77, 96 brasiliensis, 280, 300, 315, 319, 323, 384, 387, 398, 409 cicognanii, 96, 110, 118, 124, 128, 145, 155,J.61 culbertsoni, 75 erminea, 96, 110,222 eskimo, 77, 181 frenatus, 280, 300, 315, 319, 323, 341, 347, 351, 363, 366, 372, 384, 398 fuscus, 74, 222 goldmani, 76, 355 gracilis, 78, 404 haidarum, 76, 142 imperii, 110 kadiacensis, 76, 96, 108 kaneii, 100 leptus, 78, 161 leucoparia, 76, 347 longicauda, 262, 270, 280 macrophonius, 78, 360 macrotis, 77 macrurus, 370 mexicanus, 341 microtus, 100 minutus, 169 mundus, 77, 309 muricus, 77, 161 neoniexicanus, 76, 333 ni