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diff --git a/43272-0.txt b/43272-0.txt new file mode 100644 index 0000000..9dd7fc2 --- /dev/null +++ b/43272-0.txt @@ -0,0 +1,27245 @@ +*** START OF THE PROJECT GUTENBERG EBOOK 43272 *** + + Transcribers Notes: + + Italics words are denoted by _underscores_. Bold words are denoted by =equals=. + + Whole and fractional parts are displayed as 7-3/4. + + Greek text has been transliterated and are denoted by ~tildes~. + + Male and Female symbols are represented by [M] and [F] respectively. + + + + + 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 + + + + + UNIVERSITY OF KANSAS + LAWRENCE + 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 + + +[Illustration: PLATE 1. Coloration of head and foreparts in ten +subspecies of long-tailed weasel, _Mustela frenata_. All figures are of +males, approximately × 1/2. + +In regions of heavy rainfall (see figs. 2 and 3) there is an increase +in pigmentation and extent of blackish color backward over the neck and +a decrease in extent of the white facial markings. In regions +progressively more arid (see figs. 3 to 7) there is a decrease in +pigmentation and extent of blackish color and an increase in extent of +the white facial markings. + +As shown by rearing mammals from humid regions in arid regions, and +_vice versa_, the color is not visibly altered in one or a few +generations; the color is an hereditary character. Beginning with the +southernmost subspecies (fig. 1) and continuing northward to the +northern subspecies (fig. 10) there is a darkening, next a lightening, +and finally a darkening closely conforming to amounts of precipitation +in the geographic regions concerned. A fuller discussion of this +correlation is given on page 51.] + +[Illustration: FIG. 1. Map showing localities of capture of specimens +depicted in plate 1.] + + + + +American Weasels + + +BY + + +E. RAYMOND HALL + + + + +CONTENTS + + + PAGE + + INTRODUCTION 7 + + PALEONTOLOGICAL HISTORY 10 + + SKELETON AND DENTITION 12 + + DISPARITY IN NUMBERS OF MALES AND FEMALES 19 + + MATERIALS, ACKNOWLEDGMENTS AND METHODS 21 + + VARIATION 24 + Variation with Age 24 + Secondary Sexual Variation 26 + Individual Variation 28 + Seasonal Variation 30 + Variation in Coloration and Molt 30 + Variations of Taxonomic Worth 44 + + DISTRIBUTION AND SPECIATION 54 + + HISTORY OF CLASSIFICATION 69 + Chronological List (annotated) of Specific and Subspecific + Names Applied to American Weasels 71 + + CHECK-LIST OF AMERICAN SPECIES AND SUBSPECIES OF THE GENUS + MUSTELA 81 + + ARTIFICIAL KEY TO AMERICAN SPECIES OF THE GENUS MUSTELA 83 + + DIAGNOSIS OF THE GENUS 83 + + EXPLANATION OF SYSTEMATIC TREATMENT 84 + + SYSTEMATIC ACCOUNTS OF SPECIES AND SUBSPECIES 87 + _Mustela erminea_ 87 + _Mustela rixosa_ 168 + _Mustela frenata_ 193 + _Mustela africana_ 406 + + EXPLANATION OF CRANIAL MEASUREMENTS 417 + + TABLE OF CRANIAL MEASUREMENTS 418 + + LITERATURE CITED 442 + + INDEX 461 + + + + +American Weasels + + +By E. RAYMOND HALL + + + + +INTRODUCTION + + +The weasel's agility and speed take it in and out of retreats, over +obstacles and across open places in amazingly rapid fashion and are +responsible for the animal's actions being described as "quick as a +flash." The common long-tailed weasel of the United States measures +approximately a foot and a half in length, of which the tail comprises +a third; but the round, slender body is scarcely more than an inch and +a half in diameter. Brown above and whitish below in summer dress, the +animal is sleek as well as lithe and graceful. It is easy to +understand, therefore, why the Bavarian name _Schönthierlein_ (pretty +little creature) and the Italian name _donnola_ (little lady) were +bestowed upon it. The Spanish name is _comadreja_ (godmother). + +In the winter, in temperate and northern regions, the coat becomes pure +white except for the black tail-tip. In this dress the correct name for +the animal is ermine, a mammal whose fur is known to all and justly +esteemed, especially for its luster in artificial light, where it is +scarcely excelled in enhancing the beauty of gems and their feminine +wearers. + +In relation to its weight, the weasel is thought to be unsurpassed, and +perhaps it is unequalled among mammals, in the effectiveness with which +it exercises its carnivorous heritage; it kills with speed and strength +a wide variety of animals including many much larger than itself; and +it has been known to attack even man himself when he stood between the +weasel and its intended prey. In structure and temperament it is so +highly specialized for offense that, when opportunity affords, it +sometimes kills, for storage in its larder, far more than enough to +meet its immediate needs. After speaking of this tendency, Elliott +Coues (1877:129) has said: + +"A glance at the physiognomy of the weasels would suffice to betray +their character. The teeth are almost of the highest known raptorial +character; the jaws are worked by enormous masses of muscles covering +all the side of the skull. The forehead is low and the nose is sharp; +the eyes are small, penetrating, cunning, and glitter with an angry +green light. There is something peculiar, moreover, in the way that +this fierce face surmounts a body extraordinarily wiry, lithe, and +muscular. It ends in a remarkable long and slender neck in such a way +that it may be held at right angle with the axis of the latter. When +the creature is glancing around, with the neck stretched up, and flat +triangular head bent forward, swaying from one side to the other, we +catch the likeness in a moment--it is the image of a serpent." Although +Coues' colorful description more closely links the weasel with the +symbol of evil than pleases me, his description does emphasize the +raptorial character of the weasel. + +Even though most weasels are intractable as pets, they have a value to +man, as, for instance, when he is plagued by mice. In a field where +mice and other small rodents are so abundant as to damage cultivated +crops, the weasel is the farmer's best friend. A weasel may inhabit one +den until the rodents thereabouts are almost exterminated in an area +two or three hundred yards across; in this way the weasel acts as a +control, locally, as well as a check more widely, on the increase in +size of populations of kinds of rodents upon which it preys. The +smaller species are mousers of remarkable efficiency and can, if +necessary, follow a mouse to the end of the mouse's burrow. The slender +body allows the weasel to pass through any burrow or hole into which it +can thrust its head. This ability in an organism as highly specialized +for killing other animals as is the weasel, has earned for it a bad +name in connection with poultry yards. Authentic instances are recorded +in which a weasel, gaining entrance through a knot-hole to a coop of +young chickens, killed several dozen of the fowls. In other instances, +however, weasels have lived under buildings close by a poultry yard +without even molesting the birds in the slightest; in the latter +instances the weasels probably were present because there was an +abundant supply of rats and mice. At least three poultry raisers (see +page 214) have encouraged weasels to live in their poultry yards +feeling that the good they do by destroying rats outweighs the damage +caused by the occasional weasel which turns to the fowls; the idea is +that the individual weasel can be eliminated if he becomes destructive. + +Although tending to be nocturnal, weasels are almost as active by day +as by night. Their young, numbering 4 to 9, are born in a nest in a +burrow and as with other members of the Order Carnivora, are blind, and +incapable of looking after themselves at the time of birth. In _Mustela +frenata_ of Montana, breeding occurs in July and August, and the young +are born in the following April and May. Wright (1948A:342) showed that +the gestation period could not have been less than 337 days in one +individual and that it averaged 279 (205-337) days in 18 instances. +Findings of the same author (1942B:109) showed that the embryos are +implanted only 21 to 28 days before the young are born. In the +preceding part of the "long gestation period, the embryos lie dormant +in the uterus as un-implanted blastocysts. The young female weasel [of +_M. frenata_] mates when 3 or 4 months old." Consequently, in the +spring, all females of this species may produce young (Wright, +1942A:348). The circumboreal species _Mustela erminea_ likewise has +been shown to have a delayed implantation of the ova. Each of these two +species, _M. frenata_ and _M. erminea_, has only one litter per year; +but the weasel, _Mustela nivalis_, of the Old World seems to lack the +delayed implantation, in this respect resembling the ferret (subgenus +_Putorius_) as it does also in its ability to have more than one litter +per year (see Deanesly, 1944). The manner of reproduction in the South +American species _M. africana_ and the circumboreal species _M. rixosa_ +at this writing is unknown. + +The genus _Mustela_ includes the true weasels, the ferrets and minks. +The ferrets commonly are treated as a subgenus, _Putorius_, along with +the Old World polecat. The minks usually are accorded subgeneric +distinction under the name _Lutreola_, and the true weasels comprise +the subgenus _Mustela_, the three subgenera together, along with some +other subgenera which are mostly monotypic, comprising the genus +_Mustela_. Considered in this way, the group of true weasels, subgenus +_Mustela_, has a geographic range roughly coextensive with that of the +genus _Mustela_. This range includes Asia and Europe, Northern Africa, +North America and northern South America. Java has its weasel. +Australia and nearly all the oceanic islands lack weasels, and the +animals are absent from roughly the southern half of Africa and the +southern half of South America. Other small mustelids, weasellike in +shape and with corresponding habits and dentition, take the place of +true _Mustela_ in the southern half of Africa and in the corresponding +part of South America. + +In America the subgenus _Mustela_ occurs from the northernmost land in +Arctic America southward to Lake Titicaca in the Andes of South +America, a distance of approximately 6900 miles. _Felis_, I think, is +the only other genus of land mammals in the western hemisphere that has +a geographic range as extensive from north to south. _Felis_ does not +range so far north but does range farther south. The one species, +_Mustela frenata_, ranges from Lake Titicaca northward to about 57° N +in British Columbia or for approximately 5000 miles in a north to south +direction and from within the Alpine Arctic Life-zone through the +Tropical Life-zone. In North America, weasels occur in almost every +type of habitat, being absent only in the extremely desert terrain of +western Arizona and western Sonora and in adjoining parts of California +and Baja California. Even this area, along the Colorado River, may +support some weasels; evidence suggesting that it does so is given in +the account of _Mustela frenata neomexicana_. + + + + +PALEONTOLOGICAL HISTORY + + +The paleontological record fails to show the precise ancestry of +_Mustela_. The genus has been found in deposits of Pleistocene age, +but, so far as I can ascertain, not in deposits of earlier times. The +Pleistocene remains are not specifically distinct from Recent (living) +species, and in only a few instances (see _M. f. latirostra_ and _M. e. +angustidens_) are they even subspecifically distinct from the Recent +weasel living in the same area today. It is true that fossil remains +from deposits of several stages of the Tertiary beds have in the past +been identified in the literature as _Mustela_, but most of these +identifications were made many years ago when the generic name +_Mustela_ was used in a far broader and more inclusive sense than it is +today and much of the fossil material was so fragmentary that the +generic identity could not be ascertained, at least at that time. +Because the generic identity could not be ascertained, the fossil +material was tentatively assigned to the genus _Mustela_, the "typical" +genus of the family Mustelidae instead of to some other more +specialized or less well-known genus of the family. To satisfy my +curiosity about these species of "_Mustela_" of a geological age +earlier than the Pleistocene I have personally studied nearly all of +the original specimens from North America and have found each to be of +some genus other than _Mustela_. Also, such study as I have been able +to make of the Old World fossils themselves that have been referred to +the genus _Mustela_ up to 1938, and my study of the illustrations and +descriptions of the others from there lead to the same conclusion; that +is to say, none that is true _Mustela_ is known up to now from deposits +older than the Pleistocene. + +When, in 1930 (pp. 146-147), I wrote about the taxonomic position of +three American genera of fossils (known only from lower jaws), each of +which had been previously referred to the genus _Mustela_, I said that +they pertained "to that section of the weasel family (Mustelidae) +which comprises the polecats, true weasels, ferrets, minks and martens. +The fossil specimens . . . are smaller than any other later Tertiary +members of the group yet described, and are more primitive than any of +the above mentioned Recent relatives. Of the three extinct genera . . . +_Miomustela_ [Lower Pliocene or Upper Miocene of the Lower Madison +Valley, Montana] is the most primitive and _Martinogale_ [Pliocene, 18 +mi. SE Goodland, Sherman County, Kansas] is the most advanced. This +view rests largely on the character of M_{=1} which in _Miomustela_ has +a deeply basined, short, narrow talonid with a thick, high metaconid +situated partly posterior to the protoconid. In _Martinogale_ the +talonid is incipiently trenchant, long, broad, and it has a lesser +developed metaconid which is situated more anterior [ly]. _Pliogale_ +[Lower Pliocene, Humboldt County, Nevada] is intermediate in this +respect. + +"These three forms are of special interest as possible ancestors of the +subgenus _Mustela_, true weasels. No members of this subgenus, nor +related forms which can with any degree of certainty be regarded as +directly ancestral to them, have yet been described from Miocene or +Pliocene deposits. _Palaeogale_ of the Old World and _Bunaelurus_ of +North America, each of Oligocene age, have been placed by Schlosser +(1888, p. 116) and Matthew (1902, p. 137) as members of the primitive +group of mustelids ancestral to _Mustela_. This course seems logical; +and with no truly intermediate links between these forms of the +Oligocene on the one hand, and _Mustela_ which first appears in the +Pleistocene, on the other, more definite statements about ancestral +positions of the small Oligocene forms can hardly be made. The deciding +considerations for authors who placed _Palaeogale_ and _Bunaelurus_ as +ancestral to _Mustela_ were the absence of a metaconid on M_{1} and the +trenchant talonid of that tooth. These characters are found also in +_Mustela_. On the other hand certain structures in the basicranial +region of _Palaeogale_ and more especially of _Bunaelurus_ indicate +that these genera possibly are not close to the ancestral form of +Mustela . . . _Martinogale_ may stand near the ancestral form of +_Mustela_ and . . . _Pliogale_ may be ancestral to _Martinogale_. +_Pliogale_, in turn, may have had an ancestor similar to _Miomustela_. +If this should prove to be the case, _Palaeogale_ and _Bunaelurus_ +might be regarded as an independent branch which displays merely a +parallelism to _Mustela_ in the loss of the metaconid on M_{1} and the +development of a trenchant talonid on that tooth. The writer would make +it clear that he does not hold such to be the case. The ancestral +relation of _Martinogale_ to _Mustela_ is presented merely to show the +possibility, and not the special probability, of such an origin for +_Mustela_. Knowledge of the tympanic bullae and other structures of the +basicranial region would go far toward answering the question and until +these structures are known [in mustelids of the Later Tertiary,] some +uncertainty will remain." + +At the present writing I can add to the above statement only a few +facts. The discovery of better material of _Bunaelurus_ than was +available to previous workers led Simpson (1946), correctly I think, to +synonymize _Bunaelurus_ with _Palaeogale_. Simpson figures the cranial +foramina in _Palaeogale_. The differences, between _Palaeogale_ and +_Mustela_, in cranial foramina, possibly are only the result of the +elongation of the tympanic bullae. The bullae of the subgenus _Mustela_ +are seen to be much elongated posteriorly if comparison is made with +the bullae of earlier mustelids. Consequently, it might be concluded +that there is nothing in the arrangement of the cranial foramina which +would preclude the derivation of _Mustela_ from _Palaeogale_. However, +the anterior situation of the carotid foramen--well forward along the +medial margin of the tympanic bulla--is a character typical of other +mustelids and the posterior location of this foramen in _Palaeogale_ +might indicate that it was not ancestral to _Mustela_. + + + + +SKELETON AND DENTITION + + +The outstanding features of a weasel's skeleton are its length and +slenderness. Whereas the length of the vertebral column measured from +the atlas (the first cervical vertebra) to the last sacral vertebra is +175 per cent of the length of the hind leg (as measured from the head +of the femur to the tip of the longest claw), the corresponding +percentage is only 116 in the raccoon. Stated in another way, the +vertebral column and the hind leg are of approximately equal length in +a raccoon, but in a weasel the vertebral column is one and +three-fourths times as long as the hind leg. + + +VERTEBRAE + +The vertebral column consists of 7 cervicals, and ordinarily 14 +thoracics, 6 lumbars, 3 sacrals and, depending on the species, 11 to 23 +caudals. For the three species of which skeletons were examined, +variations from the normal number of vertebrae are noted in the +following table: + +TABLE I + +Data on vertebrae in three species of the subgenus Mustela (Numerals in +parentheses indicate number of specimens) + + ===================+=========+=========+========= + |_Mustela_|_Mustela_|_Mustela_ + |_erminea_| _rixosa_|_frenata_ + -------------------+---------+---------+--------- + Number of cervical | (75) | (12) | (65) + vertebrae | 7 | 7 | 7 + -------------------+---------+---------+--------- + Number of thoracic | (71) | (12) | (54) + vertebrae | 14 | 14 | 14 + +---------+---------+--------- + | (4) | | (13) + | 15 | | 15 + -------------------+---------+---------+--------- + The dorsal vertebra| (18) | (12) | (40) + constituting the | 11th | 11th | 11th + anticlinal +---------+---------+--------- + | (7) | | (27) + | 12th | | 12th + -------------------+---------+---------+--------- + Number of lumbar | (2) | | (11) + vertebrae | 5 | | 5 + +---------+---------+--------- + | (73) | (12) | (54) + | 6 | 6 | 6 + -------------------+---------+---------+--------- + Number of sacral | (9) | | (3) + vertebrae | 2 | | 2 + +---------+---------+--------- + | (65) | (10) | (67) + | 3 | 3 | 3 + +---------+---------+--------- + | (1) | (2) | + | 4 | 4 | + -------------------+---------+---------+--------- + Number of | (73) | (12) | (57) + pseudosacral | 0 | 0 | 0 + vertebrae +---------+---------+--------- + | (2) | | (6) + | 1 | | 1 + -------------------+---------+---------+--------- + | | (1) | + | | 11 | + +---------+---------+--------- + | | (3) | + | | 14 | + +---------+---------+--------- + | (2) | (7) | + | 15 | 15 | + +---------+---------+--------- + | (3) | (1) | + | 16 | 16 | + +---------+---------+--------- + | (9) | | + | 17 | | + +---------+---------+--------- + Number of caudal | (28) | | + vertebrae | 18 | | + +---------+---------+--------- + | (11) | | (6) + | 19 | | 19 + +---------+---------+--------- + | | | (14) + | | | 20 + +---------+---------+--------- + | | | (14) + | | | 21 + +---------+---------+--------- + | | | (7) + | | | 22 + +---------+---------+--------- + | | | (1) + | | | 23 + -------------------+---------+---------+--------- + +Variation according to the species is evident in the number of caudal +vertebrae, but in the other categories of vertebrae no consistent +difference in number according to species was found in the material +examined. Apparently there is also some geographic variation in the +number of caudal vertebrae within a species. For example, the one +skeleton seen of _Mustela rixosa eskimo_ (no. 219036, U. S. Nat. Mus., +from St. Michaels, Alaska) has only 11 caudal vertebrae, whereas in the +11 _Mustela rixosa rixosa_ from Roseau County, Minnesota, the usual +number is 15 with extremes of 14 and 16. Similarly specimens of +_Mustela frenata_ from Idaho and California almost always have 1 or 2 +more caudal vertebrae than do individuals of the shorter-tailed +subspecies of the same species from eastern Kansas. + +Of the vertebrae, only the cervicals, of which there are 7, were found +to be constant in number. In _M. erminea_, two of the seven individuals +in which the anticlinal vertebra was the 12th (instead of the 11th) had +15 instead of the customary 14 thoracic vertebrae. In _M. frenata_, +seven of the twenty-seven individuals in which the anticlinal vertebra +was the 12th (instead of the 11th) had 15 instead of 14 thoracic +vertebrae. The one _M. erminea_ with a pseudosacral vertebra had only +two instead of the customary 3 sacral vertebrae but the same individual +had 15 thoracic vertebrae. Of the six _M. frenata_ with a pseudosacral +vertebra, two animals had only two instead of three sacral vertebrae. +Conceivably, therefore, the pseudosacral vertebra in each of the three +instances mentioned may represent merely an unfused sacral vertebra, +instead of a true pseudosacral as occurs in four individuals of _M. +frenata_. + + +TEETH + +In American weasels, for example in _Mustela frenata_, the permanent +dentition normally is + + I 3 C 1 P 3 M 1 + -, -, -, -, -, -, -, - or 34 teeth in all. In most respects the + i 3 c 1 p 3 m 2 + +dentition is typical for post-Tertiary mustelids but in several parts +is highly specialized for a diet of flesh, the degree of this +specialization being second only to that of the cats, family Felidae. +The outstanding specialization is in the first lower molar, in which, +as in the cats, the internal cusp (metaconid) is completely suppressed +and the heel (talonid) forms an elevated blade for cutting food rather +than a basin for crushing it. In one sense the tooth is simplified +since it owes its distinctive form to a reduction in number of parts; +nevertheless, the distinctive form of the lower molar clearly is +correlated with a diet of flesh, and the tooth is correctly to be +thought of as the lower blade of a pair of shears; the upper blade is +the fourth upper premolar. The reduction in size of the second (last) +lower molar and small size of the inner lobe of the one remaining upper +molar probably are additional modifications for a diet of flesh. + +The absence of the last two upper molars and last molar in the lower +jaw would be expected in any mammal as highly specialized for a diet of +flesh as is the weasel, but these teeth are absent also in other +Quaternary members of the family Mustelidae, many of which are +substantially less specialized for a diet of flesh than is the weasel. +Therefore, in the weasel, it is reasonable to regard the absence of +these teeth more as a heritage than as an indication of a special +adaptation. The absence of a first premolar above and below, as in the +weasel, is to be expected in any carnivore that has the first lower +molar and fourth upper premolar highly specialized for shearing, but +the loss of these premolars and the small size of the second premolars +may be as much the result of a slight shortening of the face as it is a +result of a lengthening of the third and especially the fourth +premolars. The lengthening of these more posteriorly-situated teeth +would appear to be an adaptation to a diet of flesh. The cause of the +lengthening of the mentioned teeth and the reason for the absence of +the first premolars probably will be unknown until the fossil record is +more complete. + +The teeth of American species vary little except in size. The absence +of P2 in _Mustela africana_ is the only difference of a qualitative +(presence or absence) nature that was detected. Also, the Central +American subspecies of _Mustela frenata_ exhibit a tendency to early +loss of P2 and thus foreshadow the condition typical of _M. africana_. + +As a whole the dentition of the weasel exhibits a high degree of +specialization for a diet of flesh and this specialization is fully as +evident in the deciduous dentition as in the permanent dentition. + +The deciduous, or milk, dentition, of _Mustela frenata_, as known from +immature specimens of _Mustela frenata noveboracensis_ and _Mustela +frenata frenata_ available for this study, is comprised of canines, one +on each side above and below, and 3 cheek teeth on each side above and +below. See figures 2-9. The upper cheek teeth from anterior to +posterior are: a minute peglike tooth in general similar to the first +premolar of the permanent dentition; a shearing tooth in general +similar to P4 of the permanent dentition; and an anteroposteriorly +compressed tooth in general similar to M1 of the permanent dentition. +In the lower jaw, behind the canine, there is first a minute peglike +tooth, second a two-rooted tooth similar in general outline to a +permanent third premolar, and finally a shearing tooth corresponding in +function to m1 of the permanent dentition. + +No postnatal specimens which show deciduous incisors have been +examined. + +Selected, outstanding differences between the permanent teeth and the +deciduous teeth are as follows: In the deciduous teeth the canine above +has on the posterior face a well-defined ridge extending from the tip +to the cingulum. This ridge is absent or at most faintly indicated in +the permanent tooth. The lower deciduous canine, in cross section is +seen to have a marked indentation on the anteromedial border in the +region of the cingulum; this indentation is lacking in the permanent +tooth. The anterior one of the deciduous cheek teeth, both above and +below, is single rooted and its crown-surface is only about +one-fifteenth as much as that of the anterior premolar of the permanent +dentition. The second deciduous cheek tooth below has two roots, +usually fused, and differs from p4 of the permanent dentition in having +the tip of the principal cusp more recurved, in having the anterior +basal cusp better developed and the posterior heel less well developed. + +The second deciduous cheek tooth above corresponds in function and +general plan of construction to P4 of the permanent dentition but +differs from that tooth in the more pronounced protostyle, longer +tritocone, more posteriorly located deuterocone and as noted by Leche +(1915:322) separation of the protocone and tritocone by a notch. The +third upper deciduous tooth has a single cusp internally and two cusps +laterally. Thus it reverses the relation of parts seen in M1 where the +internal moiety is larger than the lateral or buccal moiety. The third +deciduous tooth below differs from m1 in very much shorter talonid and +separation of the paraconid from the protoconid by a deeper notch. + +All the features in which the last two deciduous teeth, both above and +below, are described as differing from their functional counterparts in +the permanent dentition, are features found in the permanent teeth of +primitive fossil mustelids and certain fossil and Recent viverrids. +Even so, taking into account Leche's (1915) work, which shows that the +milk teeth of some carnivores have structures lacking in the +corresponding permanent teeth of the same individual animal and also in +the teeth of genera that seem to be ancestral, a person suspects that +some of the structural features mentioned above are not inheritances +of ancestral conditions but rather specializations of the milk +dentition. + +[Illustration: FIGS. 2-9. Views of permanent and deciduous teeth of +_Mustela frenata nigriauris_. Incisors not shown. In each instance +teeth are of the left side. + +Permanent dentition × 3. No. 32421, Mus. Vert. Zoöl., [M], adult; +Berkeley, Alameda County, California; obtained October 4, 1921, by D. +D. McLean. + +Deciduous dentition × 5. No. 132158, U. S. Nat. Mus., [M], juvenile; +Stanford University, Santa Clara County, California; obtained May 7, +1898, by W. K. Fisher. + +Figs. 2-3. Lateral views of upper teeth, of adult and juvenile +respectively. + +Figs. 4-5. Occlusolingual views of upper teeth of adult and juvenile +respectively. + +Figs. 6-7. Lateral views of lower teeth of adult and juvenile +respectively. + +Figs. 8-9. Occlusolingual views of lower teeth of adult and juvenile +respectively.] + +In other deciduous teeth there is clearer evidence of more +specialization for a diet of flesh in the deciduous teeth than in the +permanent teeth. For example, the upper carnassial of the milk +dentition is even more highly sectorial than is the permanent tooth +and strikingly like that of some of the cats. The lower tooth that is +effective in the shearing action bears no more trace of the metaconid +than does the permanent first lower molar. These features of the +deciduous dentition suggest that it is more specialized for a diet of +flesh than is the permanent dentition. If this be the fact, it may seem +especially remarkable because the commonly employed term "milk teeth" +suggests that the animal makes but little or no use of these teeth in +the short time that they are in place. Accordingly, the student may +credit the form of these teeth more to some indirect effects of +inheritance than to natural selection acting directly upon the teeth. +But, after all, natural selection probably is responsible for the form +of these teeth as is indicated by the observations of Hamilton +(1933:318-325). He found that these milk teeth are used for eating +solid food as soon as the principal shearing teeth are in place. This +is three weeks after birth and before all of the deciduous teeth have +broken through the gums. These shearing teeth are used for almost two +months before being replaced by the permanent teeth and it is, +therefore, evident that natural selection could operate to fully as +great a degree in determining the form of the deciduous teeth as it may +with the permanent teeth. + +Hamilton (1933:325-326) found that the permanent dentition was complete +at 75 days after birth in captive specimens of _Mustela frenata +noveboracensis_. In the same subspecies, he noted 28 days after birth +that the canines and carnassial teeth [second deciduous cheek tooth +above and third below] had erupted through the gums. Animals 45 days +old, Hamilton found, were losing the milk dentition, and had the gums +broken through by several of the permanent cheek teeth. + +Study of the cleaned skulls available of juveniles indicates that the +deciduous teeth which persist longest are, on each side of the mouth, +the second cheek tooth above and the third cheek tooth below. These +teeth persist until after the permanent P4 and m1 have come into use. +These permanent teeth are situated immediately behind their functional +counterparts of the milk dentition. P3 and p4 are the teeth of the +permanent dentition which ultimately push out the last milk teeth to be +lost. Accordingly, in the permanent dentition, P4 and M1 appear before +P3 does, and m1 and m2 make their appearance before p4. + + + + +DISPARITY IN NUMBERS OF MALES AND FEMALES (IN ZOOLOGICAL COLLECTIONS) + + +The question has frequently been asked why twice as many male as female +weasels are captured. This is the proportion in research collections, +as may be seen from table no. 2, and I am convinced that the specimens +in these collections are saved in approximately the same proportion as +that in which they are caught. Although it might be assumed, upon first +consideration, that there are twice as many males as females in nature, +selective factors enter into the catch. For example, because a male +weasel is approximately twice as heavy as a female, it may be necessary +for him, in a given length of time, to travel twice as far as the +female to obtain the required amount of food with the result that a +given number of traps or snares will catch twice as many males as +females. Indeed, Glover (1943B:8) shows that, on the average, in +_Mustela frenata noveboracensis_ in Pennsylvania, the male actually +does travel slightly more than twice as far as the female (704 feet +versus 346 feet). From table no. 2, it may be seen that in most winter +months the ratio is 3 males to one female. This ratio is reasonable +enough, in view of what has been said, if it is considered also that +the lighter weight of the female permits her safely to step on the pans +of traps that would be sprung by heavier males. + +If in the breeding season, which is April through August in _M. +frenata_, the female is passive and if the male is restlessly searching +for her, he may thus increase still more his chances of being caught in +traps set for weasels. + +My own studies of live weasels in nature indicate that in the season +when females are attending young which are half grown, or larger, the +adult male weasels live singly in dens of their own, separate and apart +from the females and their young (Hamilton, 1933:328, records adult +males living with the female and her young, but possibly this was when +the young were less than half grown). Perhaps these males at that time +travel no farther than is necessary to obtain food for themselves. +Females, at this time, forage not only to meet their own needs, but for +food to supply their young as well. At this time, in May and June, as +may be seen from table no. 2, almost as many adult females as adult +males _are_ caught. The reason why only relatively more females than in +other months, instead of actually more females than males, are caught +at this time probably is that the adult males also are extraordinarily +active at this time because they are in breeding condition. Perhaps +the explanation 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 springing +it whereas the heavier male does spring the trap and as a consequence +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. + +TABLE 2 + +Specimens of _Mustela frenata_ (north of the range of _M. f. frenata_) +arranged by sex and under each sex by age + + KEY: + A: adult [M] + B: [M] ad., % of total adults + C: subadult [M] + D: young [M] + E: juvenal [M] + F: total number of [M] + G: [M] % of total + H: adult [F] + I: [F] ad., % of total adults + J: subadult [F] + K: young [F] + L: juvenal [F] + M: total number of [F] + N: [F], % of total + O: total number of [M] and [F] + P: total number of adults, [M] and [F] + + /-----------Male---------\/--------Female--------\ + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + | A | B | C | D | E| F | G| H | I| J| K | L| M | N| O | P + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + May | 29| 55| 4| 14| 7| 54|59| 24|45| 1| 9| 3| 37|41| 91| 53 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + June | 42| 53| 14| 40| 8| 97|59| 38|47| 4| 25| 2| 69|41| 166| 80 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + July | 59| 70| 18| 55| 2|130|59| 25|30| 5| 58| 2| 90|41| 220| 84 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + August | 40| 77| 23| 55|..|113|74| 12|23| 2| 25|..| 39|26| 152| 52 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + September| 15| 79| 25| 12| 1| 51|75| 4|21| 4| 9|..| 17|25| 68| 19 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + October | 11| 58| 46| 7|..| 43|66| 8|42|13| 1|..| 22|34| 65| 19 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + November | 41| 70| 48| 1|..| 88|73| 18|30|12| 2| 1| 33|27| 121| 59 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + December | 59| 69| 43| 1|..|108|73| 26|31|15|...|..| 41|27| 149| 85 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + January | 80| 69| 32| 2| 1|126|72| 36|31|14|...|..| 50|28| 176|116 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + February | 45| 66| 19| 5|..| 82|73| 23|34| 4| 3|..| 30|27| 112| 68 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + March | 38| 72| 2|...|..| 57|70| 15|28| 8| 1|..| 24|30| 81| 53 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + April | 30| 67| 2| 4| 3| 39|67| 15|33|..| 2| 2| 19|33| 58| 45 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + Totals |489| 67|281|196|22|988|68|244|33|82|135|10|471|32|1,459|733 + ---------+---+---+---+---+--+---+--+---+--+--+---+--+---+--+-----+--- + +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 than 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 stuffed skins with the +skulls separate and each was accompanied by the customary data as to +locality of capture, date of capture, name of collector, external +measurements and sex recorded on the labels by the collectors. Skulls +unaccompanied by skins, nevertheless, comprised a large share of the +total and a small proportion was made up of skins unaccompanied by +skulls, mounted specimens, skeletons, and entire animals preserved 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, approximately 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 publication). 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 publication 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 Grinnell and Mr. Charles + D. Bunker, among others, gave truly valuable encouragement and + assistance. + + Collections containing weasels which were examined in the study + here reported 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 Criddle +Collection of John Cushing +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 +Collection Rocky Mt. Spotted Fever Lab. +Collection of Victor B. Scheffer +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 Zoölogy +Mus. Polonais d'Hist. Nat., Warsaw +Mus. Vert. Zoöl., Univ. California +Museum of Zoölogy, Univ. Michigan +National Museum of Canada +Naturhistoriska Ricksmuseum, Sweden +Neuchatel University Museum +New York State Museum +Ohio State Museum +Oklahoma Agric. and Mech. College +Ottawa University, Kansas +Paris Museum +Provincial Museum of British Columbia +Royal Ontario Museum of Zoölogy +San Diego Society of Natural History +State Hist. and Nat. Hist. Soc. Colo. +State Normal School, Cheney, Wash. +Texas Cooperative Research Collection +United States National Museum +University of Arkansas +Univ. California Mus. Palaeo. +University of Idaho +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 Zoölogy +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 a large share of the holotypes. + Specimens in all of the North American collections including + Canada and México 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 assembled, special effort, with considerable + success, was made in each of several winters, to obtain specimens + from areas not previously represented in collections. + + 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 California + Chapter of Sigma Xi, the John Simon Guggenheim Memorial Foundation + and the Kansas University Endowment Association. I am mindful also + of an obligation to those who appropriated funds, by legislative + action, for research 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 Blos 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. Abernathy 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. _f_ 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 terminal + 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 +that 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 individuals 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 geographically intermediate +between the nominal species, I found intergradation and therefore +arranged the nominal species as subspecies of a single species. +Intergradation here is understood to be the result 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 according them subspecific rank. Absence +of intergradation in nature 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 _Mustela_, 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 variation 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 localities. 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 +zoölogical 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 approximately +the 37th day. When 2 to 4 months old, the tail is pointed 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 +subspecies, 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 increasing age +in the color-pattern or pelage that cause the systematist 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 +ultimate adult size, intermediate in stage of development. The +permanent 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 zoölogist 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 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 differences 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 animal 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 differentiating 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 subadults. +Ordinarily the few adults obtained in these seasons can easily be +segregated from the subadults because ontogenetic development 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 revealed 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 apparently 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 difference +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 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, _M. e. haidarum_ and _M. e. +olympica_, 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 +approximately 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 subspecies 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_, although 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. f. 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. f. +noveboracensis_ and _M. e. arctica_, in addition to the secondary +sexual variation already mentioned in the skull, females have the +braincase 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 +accounts. 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 Schädel vieler Säugethiere . . . so gross +sind, dass man diese wie Schädel verschiedener species behandeln muss, +während in anderen Ordnungen (Rosores, Edentaten) die Schädel 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 (_Mustela 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 parents, +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 represent +different genetic strains within certain populations and variations +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 probably result +largely from failure to recognize age-variation. When skulls of a large +series from any one locality are arranged first by 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. + +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 between 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 comparison 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 individuals become white whereas others become white on +only the underparts, the upper parts changing only to lighter brown. +Probably 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 magnitude but no evidence was found of +repetition of an abnormality in 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 excluded from +consideration, in the same way that variations ascribable 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 longitudinal stripe of +brown on the middle of the light-colored underparts; 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 markings occur in subspecies of _M. frenata_ +from the southwestern United Stated 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 markings 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 +individual 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 almost 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 +indirectly, 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 (melanin) 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 provide 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 temperatures 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 correlation between coat color and climate. On the +Asiatic continent, several species, including _M. erminea_, provide +parallel correlations and nowhere are there any exceptions for the +subgenus _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 subject +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 color. Bachman (1839:228-232), Macgillivary +(1843?:158), Audubon and Bachman (1851 (vol. 2):62), Schwalbe +(1893:538), Pearson _et al._ (1913:447), Miller (1930, 1931A), Hamilton +(1933:300) and Rothschild (1942), among others, have been inclined 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 dealing with this question. + +To my mind, it has not so far been demonstrated that 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. Metchnikoff'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 slight +importance in the whitening 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 the 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. f. noveboracensis_ and, each morning, found clumps of brown hair on +the floor of its cage; this 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 _only_ 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 Rapid City, South Dakota, if taken individually, do +not, in macroscopic examination, show definite molt lines or other +absolutely convincing evidence of molt. However, these same specimens, +insofar as examined microscopically, do show overhairs all white, or +overhairs pigmented 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 +_Mustela frenata_ and in the other species of more northern +distribution. Schwalbe's (1893) work, including sectioning of the skin +and study of the hair follicles, led him to conclude that the underfur +was molted twice each year in _Mustela erminea_. + +In _Mustela frenata noveboracensis_, _M. f. nevadensis_, and _M. f. +nigriauris_, measurements taken on adult males show the overhairs to be +longer in the winter pelage than in the summer pelage of specimens from +the same locality. For example, in _M. f. nigriauris_ from 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. Although 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), Hamilton (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_, subspecies +included, the spring molt begins on the mid-dorsal line and proceeds +laterally, producing, at almost any given time, a relatively 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 lines. 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 difference 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 commenting 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 autumn, 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 instances, +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 approximately 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. f. +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 (_loc. cit._) +thinks also that in _M. erminea_ studied by him, the black tip of the +tail in autumn is replaced approximately one month in advance of the +pelage on the rest of the body. As indicated above, my specimens of +_Mustela frenata_, subspecies _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 remainder 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 acquaintance who have +suggested that molt occurs in spring, whereas the individual hairs +change color in autumn. Reference to plate 39 will show how gross +comparisons might lead one to this erroneous explanation of the color +change. + +As to time of molt: In eight subspecies of _Mustela frenata_, namely, +_noveboracensis_, _occisor_, _primulina_, _spadix_, _longicauda_, +_arizonensis_, _nevadensis_ and _effera_, material is available to +indicate that the autumnal molt begins in October and is completed in +November, and that the spring molt occurs in March or April. A +condensed list of specimens providing basis for this statement is as +follows: + + _M. f. noveboracensis_: 26 specimens in transitional pelage taken + in autumn and 14 taken in spring; _M. f. occisor_: One topotype + has acquired one-fifth of the winter pelage on October 22, 1896; + _M. f. primulina_: 2 in November, one in March, and 2 in April are + in process of change; _M. f. spadix_: 6 autumnal specimens and one + in April show pelage change; _M. f. longicauda_: 7 autumnal + specimens and one in April show pelage change; _M. f. + arizonensis_: 12 specimens in autumn and 3 in spring are in + process of molt; _M. f. effera_: One November-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 autumnal 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 +brown or white. By plotting on a map the localities 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 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. + +[Illustration: 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_.] + +Hamilton (1933:302) has pointed out that "Where half of the weasels +remain brown, these brown winter specimens are always 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 localities 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 different 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 addition 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 individuals 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 America 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 wide, 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 dependent 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 November 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. + +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 +the 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 south 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 winter 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. + +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._:246) "Reducing the daily periods of light induced molting and +regrowth of new fur. . . . In the Bonaparte weasels [_Mustela +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 +pituitary 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) injected 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 follicles contain +no melanin-forming enzyme (dopa-oxidase), 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 the cells contained granules of pigment in spring when, +as we know, the granules are incorporated 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 precisely accurate, +other parts are the result of inferences which need 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 +this 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 underparts. 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 _Mustela +erminea_ and _Mustela nivalis_. Although in the long-tailed weasels +(_Mustela frenata_) the underparts 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 richardsonii_, 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 regards 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 have been an albino _Mustela vulgaris_ from Yorkshire +and an albino _M. erminea_ from an unstated locality. Jäckel (1873:459) +mentions specimens of _Mustela erminea_ 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 erythristic or only partly albinistic. Among the 1550 +skins of _M. frenata_ which were in summer pelage or brown 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 mid-dorsal line behind +the ears. Markings of this kind are not abnormal in _M. f. peninsulae_, +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. f. 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 proximal part is of the normal brown color. The +black terminal part commences proximally at the usual place, but the +distal 11 millimeters of the fleshy part of the tail bear only pure +white hairs producing a terminal white pencil 35 millimeters long. + +The three other specimens abnormally colored are erythristic +individuals. An adult male of _M. f. 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 occurs +above each ear. Incidentally, this and other subspecies of _Mustela +frenata_ 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. +oregonensis_, no. 47149, Mus. Vert. Zoöl., 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 individuals of normal coloration are the white +or light orange facial markings, have the reddish cast of the remainder +of the body, although 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 +population of polecats, _Mustela putorius_, in Cardiganshire, England, +in which this erythristic variation is maintained in a state of nature. +In ferrets, _Mustela furo_, Pitt (_op. cit._:114) notes that ". . . +erythrism is certainly dependent on a Mendelian factor, being dominant +to albinism and recessive to the black-brown coloration. Both in the +ferret and polecat, erythrism seems to be correlated with increased +size, and certainly in the ferret is usually accompanied by a quick +temper and general increase in vitality." + + +Variations of Taxonomic Worth + +Variations of taxonomic worth usually are referred to as characters. +For example, shortness of the tympanic bulla is a character, and the +opposite condition, long tympanic bulla, is another character. Specific +variations, that is to say specific characters, are provided by the +color-pattern, length of tail, number of premolar teeth, shape of the +tympanic bullae, and length of the braincase in relation to the length +of the tooth-bearing parts of the skull. Subspecific characters are +provided by color-pattern, color itself, size as measured by weight of +the animal, and its linear 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 the 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 geographic +variation. These variations, characters in zoölogical parlance, when +plotted on maps, reveal the geographic occurrence of, say, the convex +shape of the skull. In 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 adjacent to the westward. On this +line and for several miles to either side of it weasels show varying +combinations of these three characters 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 between that of the unicolored face of _arthuri_ +and that of the bicolored face of _frenata_, (2) the long tail of +_frenata_ and (3) the convex skull of _arthuri_. In the sum of its +characters this specimen is exactly intermediate between typical +_arthuri_ 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 _arthuri_ but the tail fully as long, and the skull as +lacking in dorsal convexity, as in _frenata_. Now the 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 locality 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 +the average is for the area as a whole. + +The three individual animals of an intermediate sort are ordinarily +termed _intergrades_. This implies that their characters are the result +of mixed parentage--perhaps a female of _M. f. arthuri_ and a male of +_M. f. 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 +intergradation 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 the adjacent mainland, 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 degree of difference is greater when the insular kind +is compared than when only the kinds of the mainland are compared, the +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 +criterion is degree of difference, with the limitation of geographic +adjacency, rather than intergradation. + +The geographic variation (subspecific characters) found could be spoken +of as two kinds: First, there is the variation which is expressed 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 follows: "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 fluctuations are somewhat wavelike in character +and rise to central points of extreme development and then 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 variation +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, zoölogical 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 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. erminea_. There is no cline of sustained +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 +subspecies _neomexicana_ having the long tail and the two subspecies +_arthuri_ and _primulina_ having short tails. The geographic ranges of +_primulina_ and _neomexicana_ are contiguous. In _M. erminea_ there is +likewise no variation of a clinal nature in length of tail and +furthermore 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 populations, 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. f. +arizonensis_) to 59 mm. (_M. f. 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. +Geographically 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 the 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 conditions at low altitudes. +The conclusion seems unavoidable that climate, directly or indirectly, +determines the degree of hairiness. Less careful observations were made +on the hairiness of the soles of the feet in other species but it is +clear that the northern species _M. erminea_ has the most hair on the +foot-soles and that _M. africana_, the tropical weasel, has the least. +In this regard, _M. frenata_ is intermediate as it is also in +geographic position. + +[Illustration: 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. All × 1. + +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.] + +[Illustration: FIG. 16. Map showing the localities where the skulls, +represented in figures 11-15, were obtained.] + +The maximum length of facial and carpal vibrissae is attained in _M. +erminea_ in the far north. In weasels from north of the Arctic Circle +the longest facial vibrissae extend posteriorly beyond the posterior +border of the ear. In the tropical weasel, _M. africana_, the facial +vibrissae do not extend posteriorly beyond the ear and the carpal +vibrissae are not so long as the distance between their bases and the +apical pad of the first digit. The correlation of long vibrissae with +low temperature, is mentioned here merely because length and density of +pelage were under consideration. + +The most obvious and most exact correlation between change in climate +and change in the animal is furnished by color. This is well shown in +the one species, _Mustela frenata_, to which the following remarks +apply unless indication is given to the contrary. The color of the +upper parts varies from bay (blackish brown) in _M. f. panamensis_ to +buckthorn brown (light brown) in _M. f. neomexicana_. The color of the +head varies from solid brown (white chin excepted) to contrasting black +and white markings. + +Dark color of the upper parts is associated with a large area of this +color; the enlargement of this area is at the expense of the area of +light color on the underparts. In the weasels of darkest color the +upper parts occupy four-fifths of the circumference of the body (as +measured in the anterior lumbar region) but in the lightest-colored +weasels the upper parts comprise only two-thirds of the total +circumference. In these light-colored animals the color of the +underparts extends onto the underside of the tail and down the insides +of the legs and over the feet whereas in the animals with the darkest +upper parts the entire tail, feet, and legs below the knees ordinarily +are of the same dark color as the upper parts. The length of the black +tip on the tail varies inversely with the length of the tail, probably +because the lightest-colored weasel has the longest tail. In some +subspecies the black brush is almost half as long as the tail-vertebrae +but in others is less than a fourth as long as the tail-vertebrae. + +The extent of the color of the head, as well as the intensity of the +color there, varies markedly and is correlated with climatic +conditions. The extent and intensity of this dark color is greater in +weasels inhabiting regions of heavy rainfall than in those inhabiting +regions of sparse rainfall. Considering the geographic range of each +subspecies of _Mustela frenata_, that of _M. f. panamensis_ has the +maximum of rainfall. Reference to the colored plate (1) will show that +in _M. f. panamensis_ (2) the black of the head is extended over all of +the upper parts. _M. f. macrura_ (1) of Perú, to the southward, is from +an area of lesser rainfall and is correspondingly lighter colored. +Returning to _panamensis_ (2) as a starting point and proceeding +northward to the range of _nicaraguae_ (3), which also has lesser +rainfall, thence another step northward to Guatamala, which has still +less rainfall, the weasel there, _M. f. goldmani_ (4) has the black +extending posteriorly only to the shoulders. _M. f. leucoparia_ (5) +from Michoacán, and _M. f. frenata_ (6) from Tamaulipas are from +progressively more northern and also progressively drier regions. In +_M. f. frenata_ (6) the dark color extends posteriorly only to the ears +and is blackish rather than black. In _M. f. neomexicana_ (7) of the +extremely arid parts of Durango, Arizona, and New Mexico the dark +marking of the head is confined to a brown spot on the nose. Its +geographic range is the most arid of those of all of the subspecies. +The contrast between _neomexicana_ (7) and _panamensis_ (2) illustrates +the great range of geographic variation in color which occurs in the +one species. Continuing from the geographic range of _neomexicana_ +(specimen from Safford, Arizona) northwesterly 480 miles to Riverside, +California (see 8, _latirostra_), 430 miles north to Point Reyes, +California (see 9, _munda_), and finally 570 miles north to Tillamook, +Oregon (see 10, _altifrontalis_), each place with more rainfall than +the one farther south, another correlation of increasingly dark +coloration with increasing amount of rainfall is illustrated. + +This geographic variation, it should be remembered, is all within one +species. It is the more significant still when we remember that the +same correlation, with never an exception, occurs at hundreds of places +within the geographic range of the species. A particular feature of +climate, namely rainfall, and possibly therefore humidity, is concerned +in this correlation. The same correlation, heavy rainfall and dark +color, is shown also in the other species of North American weasels. +The conclusion is unavoidable that climate, directly or indirectly, +determines or influences the color of weasels. + +The light facial markings appear in American weasels in two separate +geographic areas. One is the southwestern United States, México and +northern Central America. The second area is in the same latitude, in +Florida and adjoining parts of Georgia and Alabama. In the western +weasels the markings are white south of latitude 32° N. North of this +latitude, the facial markings, if at all extensive, usually are of the +same yellowish color as the underparts of the body. Weasels of southern +California and its interior valley usually have these yellowish instead +of white facial markings. The light facial markings, in this instance, +white markings, attain their maximum extent in _M. f. leucoparia_ of +the southwestern margin of the tableland of México, at latitude 19° N. +A gradual decrease in area of the light facial markings occurs both to +the north and south; they disappear at 10° N in _M. f. costaricensis_ +and at 35° N at approximately the southern limits of range of _M. f. +arizonensis_ and _M. f. nevadensis_. In the mild climate of California +the light (yellowish) facial markings are found at still higher +latitudes. These light facial markings crop up as vestiginal remnants, +consisting of a few white hairs, in some individuals of nearly all +races of weasels. + +In certain parts of the skull there are trends, in size and shape, +which continue for long distances geographically. In other words, +clines can be recognized. Changes in size and shape in some other parts +of the skull are wavelike; change toward narrower rostrum, for example, +is not progressive in a given geographic direction for any great +distance. Length of the upper tooth-rows and zygomatic breadth, when +expressed as percentages of the basilar length, and also the actual +length of individual teeth vary geographically in the same wavelike +fashion as does the width of the rostrum. + +Size of the skull, on the other hand, shows a sustained trend for a +long distance; it becomes progressively smaller from the southern +United States southward to Columbia, South America. This clinal +variation can be demonstrated by plotting on a graph, the basilar +length, the zygomatic breadth, or the weight of the skull. Beginning at +Mérida, Venezuela, and proceeding southward to increasing elevations in +the mountains of South America, there is a reversal of the direction of +the variation in this cline; weight of skull, for example, increases to +the southward from Mérida for a considerable distance. A cline of +decreasing width of the postorbital constriction of the skull is +evident from Panamá north into Texas. + +Variations in the tympanic bullae provide many characters useful in +distinguishing weasels from different localities. Most of these +characters have to do with degree of inflation of the bullae. +Indirectly correlated with degree of inflation is first the extent of +removal of the anterior margin of the bulla from the glenoid fossa and +foramen ovale, and second the form (convex, flat, or concave) of the +part of the squamosal bone between the foramen ovale and the anterior +margin of the tympanic bulla. As one proceeds southward from, say, +southwestern Kansas through the geographic range of the species +_Mustela frenata_, there is a progressive deflation of the bulla, an +increase in length of the space between its anterior margin and the +foramen ovale, and the floor of the braincase in front of the bulla +changes from ventrally concave to ventrally convex. (See figs. _e_ and +_h_ of pl. 24 and figs, _e_ and _f_ of pl. 27.) + +One extreme of this variation in bulla is shown in _Mustela frenata +neomexicana_ (fig. _e_ of pl. 24), in which the anterior margin of the +bulla (viewed from the ventral side) rises vertically from the floor of +the braincase to form a 90-degree angle. The other extreme, the +uninflated bulla, is in _Mustela frenata panamensis_ (fig. _e_ of pl. +27), in which the anterior margin of the bulla is not raised above the +floor of the braincase. This variation is remarkable because it occurs +within a single species. Otherwise, in the family Mustelidae, +differences in the tympanic bullae as great as that between the two +subspecies _M. f. neomexicana_ and _M. f. panamensis_, occur only +between genera. The need for caution in inferring the limits of +variation for a particular structure in one species or genus, on the +basis of variation in another group, is therefore obvious. + +Speaking now of full species, the most inflated tympanic bullae in +American weasels are in _Mustela frenata_, and more restrictedly in +those subspecies of it which occur in the temperate region. Subspecies +of _M. frenata_ in Central and South America, as already noted, have +less inflated bullae. The tropical weasel, _Mustela africana_, of the +Amazon drainage of South America has the bullae still less inflated +(see fig. _i_ of pl. 39 and fig. _f_ of pl. 40). The bullae are less +inflated even than in the mink, subgenus _Lutreola_. In _M. africana_ +the cleidomastoideus, omotrachelian, levator scapulae, and rhomboideus +profundus muscles take origin from a fossa on the mastoid bone, whereas +in the forms with greatly inflated bullae these muscles take origin +from a raised ridge or tubercle. Using _Mustela frenata_ of the +temperate region as a starting point and proceeding northward, a +reduction in inflation of the tympanic bulla is seen also in that +direction in that _Mustela erminea_ has less inflated bullae. The +bullae are less inflated in southern than in far northern (arctic) +populations of _Mustela erminea_. In _erminea_ the lesser inflation is +real enough but at the same time there appears to be less inflation +than actually exists, for the squamosal floor of the braincase is +"pushed down." This places the anterior end of the tympanic bulla +farther in the braincase than it otherwise would be. Although the +anterior end of the bulla is flattened to the extent that it resembles +the sharp edge of a splitting-wedge, inspection of the lateral and +medial edges shows that in its central part the bulla is more inflated +than it is in the weasels of Central and South America. + +For reasons set forth later, _M. erminea_ is judged to resemble the +ancestral stem form more closely than does any one of the other three +American species of weasels. If this judgment is correct, the shape of +the tympanic bullae of the American weasels may be explained as +follows: In the subspecies of _Mustela frenata_ of the temperate +regions of North America the bullae have most nearly been pushed out of +the braincase and at the same time have undergone some enlargement. The +subspecies of this same species in Central and South America represent +an earlier stage in the evolution of American weasels and retain less +inflated bullae--less inflated even than those of the southern +subspecies of _erminea_. _M. africana_ probably separated from the stem +form at a still earlier time if we may judge by the lesser inflation of +its tympanic bullae. There are other reasons for thinking that +_africana_ separated from the stem form earlier than _M. frenata_ did. +During the time that elapsed since the separation of _M. frenata_ from +the stem form, the tympanic bullae of _M. erminea_ probably increased +slightly in size, as probably also did the brain but without shoving +the auditory complex forward from its former position. + + + + +DISTRIBUTION AND SPECIATION + + +Weasels of the subgenus _Mustela_ are known from the Pleistocene but +not from deposits laid down at an earlier time (see page 10). The +Pleistocene weasels from Rancho La Brea of southern California and from +Potter Creek Cave and Samwel Cave, both of northern California, are +subspecifically indistinguishable from the weasels living in those same +localities today. The other notable occurrence of weasels in the +Pleistocene is in the Conard Fissure of Arkansas. Brown (1908:181, 182, +pl. 17) names two kinds from the Fissure. One is an extinct subspecies +(_Mustela frenata gracilis_) possibly of the species which occurs in +the same region today and the other, _Mustela erminea? angustidens_, +is an extinct subspecies of a species which occurs only farther north +today. _M. erminea_ came south, probably in front of one of the ice +sheets, as did several other species of American mammals, now of more +northern distribution, that left their remains in Conard Fissure. +_Mustela rixosa_ is not recorded as a fossil in America although it is +known from the "Diluvial" deposits of the Old World; see Woldrich +(1884:1000), who employs the name "_Foetorius minutus_ n. sp.," and see +also Zimmerman (1943:295-296). + +The ermine, _Mustela erminea_, is the most generalized of the full +species. For example, the number of teeth is as large as in any other +species and greater than in certain species. The teeth are +sharp-pointed, uncrowded, and individually less specialized than in any +other American weasel. M1 has the inner half, or lobe, of approximately +the same size as the outer lobe instead of much larger than the outer +lobe (the outer lobe is the larger in several other species). The +tympanic bullae are less inflated and less protruded from the +braincase. The skull is rounded, and has no marked crests and ridges +whereas the skulls of the other species are more pronouncedly modeled +and sculptured. Therefore, it is possible to think of these other +species as derived from _M. erminea_. A derivation in the reverse +direction would be more difficult. From the foot soles of an ermine, or +a weasel closely resembling an ermine, the more complex soles of +_Mustela africana_ could have been derived by a decrease in hairiness, +although it would be necessary to suppose that the thenar pad has been +retained in _africana_ and has been lost in the living _erminea_. The +alternate possibility, namely, that the thenar pad was a relatively +recent acquisition in the _africana_ line seems less probable. The tail +of _erminea_ is of "average" length and in size of entire animal +_erminea_ is intermediate between the other American weasels. +Structurally, _Mustela erminea_ appears to be nearest the stem form +from which all of the living weasels ascended. Its present holarctic +distribution is in harmony with the view that it is a direct descendant +from the stem form because the stem forms of most of the known kinds of +mustelids appear to have lived in the holarctic region. To be sure, +_Mustela erminea_ is regarded as having undergone some progressive +change in structure, but less than the other weasels, in the period of +time when the weasels were evolving from the stem form. + +The least weasel, _Mustela rixosa_, seems to be an ancient type and to +judge from the size and proportions of its parts, was differentiated +from the _erminea_ stem at a time earlier than were the other American +Recent species of weasels. In size, in reduction of the tail, and in +proportions of the skull, _M. rixosa_ is, in each instance, the most +aberrant of all the weasels, _Mustela nivalis_ of Europe and western +Asia included. This aberrancy results from the retention of certain +primitive features, in the teeth and basicranial region, and from +specialization in proportions of the skull. The skull is long, deep, +and narrow. These proportions probably are adaptations permitting the +animal to follow the smaller kinds of mice into their burrows. In most +of that part of North America where _erminea_ and _rixosa_ occur +together, _erminea_ is a much larger animal and takes as prey almost +all kinds of land vertebrates that it is powerful enough to kill. These +include varying hares and ptarmigans. The least weasel, _rixosa_, can +hardly manage such large prey and lives on the smaller rodents. +_Mustela rixosa_ may eat numbers of insects (see page 176 beyond),--a +kind of food which _Mustela erminea_ is not known to eat. Apparently +the two species are able to live in the same areas because each eats a +somewhat different kind of food than does the other and hence they do +not compete to the point where one is crowded out by the other. This is +the case in the latitudes where the two species of weasels are of +different bodily size, but in the southernmost latitudes where these +two species occur, _erminea_ becomes almost as small as _rixosa_ and +only one of the species, to the exclusion of the other, occurs in a +given area. All through the Rocky Mountains, south of Montana and in +the territory west of these mountains all the way to the Pacific Coast, +only the small subspecies of _erminea_ is to be found. In the +Alleghenies of the eastern United States only _rixosa_ occurs. In New +England where _erminea_ approaches the size of _rixosa_, the latter is +unknown. Probably this exclusiveness results from competition for food, +although competition for dens, safe breeding places and other +requirements of life may be involved. + +The species _erminea_ invaded the western United States and in the +process of invasion probably developed there the small size appropriate +to permit _erminea_ to live in that latitude before it could do the +same thing in the Appalachian region. Later than _erminea_, the least +weasel, _Mustela rixosa_, which was small to begin with, also spread +southward from the holarctic region, stopped short in the western +United States at the northern boundary of the area in which _erminea_ +was of small size, but in the Appalachian region of the eastern United +States continued on southward to the limits of temperature tolerant for +it because _erminea_ had not yet penetrated into that region and no +other small carnivore was there to offer competition. + +The long-tailed weasel, _Mustela frenata_, occurs mostly south of the +regions inhabited by the ermine, and mostly south of the region +inhabited by the least weasel which appears to live as well with +_frenata_ as with _erminea_. It is true that _erminea_ and _frenata_ +occur in the same region, but this is a relatively narrow belt across +the United States; and from within it a person cannot go far either +north or south without reaching a region in which only one of the two +species occurs. Exception has to be made for the Rocky Mountains and +the Sierra Nevada, where _erminea_ is of exceptionally small size. In +these mountains and in the boreal mountainous parts of the intervening +region of the United States, _erminea_ and the large-sized _frenata_ +occur together over a wide area. Presumably the two occupy different +ecologic niches, much as _rixosa_ and _frenata_ probably do where they +occur together. + +Most of the geographic range of the long-tailed weasel, _M. frenata_, +is in the temperate region. Structurally, this species is the most +advanced of the American weasels. Its dentition is the most highly +specialized for cutting. M1 is relatively small and the inner lobe is +slightly larger than the outer lobe. The skull, throughout, is more +modeled than in the other species; the rostrum, the lower jaws and the +teeth--all parts of the offensive equipment--are well developed +relative to the corresponding structures in other weasels; the +basicranial region exhibits an advanced stage of development in that +the tympanic bullae show the maximum degree of inflation. Also, they +are thrust far out of the braincase, thereby providing more room for +the relatively larger brain which is protected by a more solidly built +braincase than in _erminea_. + +Several subspecies of _Mustela frenata_ occur in the tropics, that is +to say, south of the Mexican tableland and on the coastal plain to the +east of it. Each is structurally more primitive than subspecies of the +temperate region. As compared with _Mustela frenata frenata_ of the +temperate Mexican tableland the size in these tropical subspecies is +smaller; the tail is shorter; the braincase and entire skull are less +modeled; the postorbital breadth is more; the teeth are smaller; the +deuterocone of P4 is not so far anterior to the protocone; the tympanic +bullae are less inflated, are farther removed from the foramen ovale, +and a larger proportion of each bulla is contained within the +braincase. These features serve to set off from northern races of +_frenata_ all those subspecies of _frenata_ which occur from southern +México southward to the northern and western limits of the Amazon +drainage of South America. The Amazon Basin is inhabited by another +species, _Mustela africana_, having more primitive characters. + +In the species _frenata_, the explanation for this abrupt change in +characters between the animals of the temperate highlands and those of +the tropical lowlands may be this: In the early Pleistocene, after the +emergence of much or all of Central America took place, weasels +distributed themselves over the Isthmus and into South America. These +weasels were more generalized in structure than those now inhabiting +the uplands of México. Failure of this stock of weasels often to cross +some still-persisting water barrier, or failure of this stock to cross +some water barrier that was widened or reformed because of a rise in +sea level in some one of the interglacial periods of the Pleistocene +cut the _frenata_ stock into two or more parts. After the land +connection was established or re-established and when the necessary +precedent plants and rodents again had established themselves, the two +groups of weasels, one from the northern tableland of México, and the +other from the southern area of tropical complexion, met. The weasels +of the _frenata_ stock that reinvaded the area from the north probably +did so by following along the chain of high volcanic cones and narrow +uplifts. If and when a subsequent inundation occurred in some +part of Central America, weasels were stranded on the adjacent +mountains--converted into islands--only the higher parts of which were +above water. _Mustela frenata costaricensis_ and _Mustela frenata +goldmani_ may be examples of a northern stock of weasel that pushed +southward in the highlands and became stranded for a short time. +Following the latest emergence of land to provide a continuous highway +between the two continents, weasels from the south and the insular +populations, as for example, _M. f. costaricensis_, were the first to +invade the low tropical areas most recently under water. When the +Pleistocene history of Central America is better known, the facts will +provide a useful means of testing the hypothesis that has been outlined +immediately above. + +As explained above, fossil specimens of _M. frenata_ from deposits of +the last half of Pleistocene time show that no appreciable change +occurred in some areas, for example, in the vicinity of Hawver Cave and +Samwel Cave of California, and that but slight change occurred in other +areas, for example, in southern California (fossils from Rancho La +Brea) and probably in the central United States (fossil from Conard +Fissure). It is possible to imagine, therefore, that the two groups of +weasels, one occurring southward only as far as the highlands of +Central America and the other occurring in northern South America, had +not differentiated sufficiently in the period of their isolation to +prevent crossbreeding when they last came into contact. If the +separation of the two groups had been maintained for a longer period, +the two groups, tropical weasels and austral weasels, probably would +have been so different when the two met as to prevent crossbreeding and +they would have constituted two full species instead of only one. + +_Mustela africana_ is the most primitive of the American weasels. Some +of the most important structural features that mark it as such are in +the basicranial region. The tympanic bullae are less inflated than in +other weasels, are pointed anteriorly and posteriorly, and do not have +the lateral margins carried outward to the outer margins of the +braincase. The mastoid sinus is not involved, by inflation or marked +modification in the production of the auditory complex. Between the +alisphenoid and the squamosal there is a clear demarcation posteriorly +from a point directly lateral to the foramen ovale. This demarcation +permits a transverse rounding of the alisphenoid to form a longitudinal +ridge between the anterior margin of each bulla and the base of the +pterygoid of the same side. Nevertheless, there is no such +specialization of this primitive, structural feature such as occurs in +some African and Asiatic mustelids in which the tympano-pterygoid part +of the alisphenoid fuses with the tip of the hamulus of the pterygoid. +However, the tympano-pterygoid eminence has not been obliterated in _M. +africana_ as it has in the other American weasels. Another primitive +feature in the basicranial region of _M. africana_ is the tendency +toward separation of the paroccipital processes from the tympanic +bullae. The thenar pad of the foot probably is an inheritance from a +primitive ancestor since the pad is present in the viverrids and in a +majority of mustelids judged to be more primitive than _Mustela_. + +Some specializations are obvious in _Mustela africana_. One is the +reduction in number of premolars; p2 is absent whereas it is normally +present in the other weasels; P2 has one instead of two roots; and, in +relation to the other teeth, m2 is smaller. The shortness of the +preorbital part of the skull in relation to the length of the skull as +a whole may reflect the mentioned reduction of the premolars or +retention of a primitive shape of skull, or both. Also, certain +features which denote immaturity in other weasels are retained in +adults of this species, as for example, sutures on the dorsal face of +the preorbital region of the skull. + +[Illustration: FIGS. 17-22. Views of the feet of American weasels +(subgenus _Mustela_) to show differences in number and arrangement of +the pads and variation in degree of hairiness of the soles. × 1-1/2 In +each figure, left-forefoot on left, and left hind foot on right. + +FIG. 17. _M. rixosa rixosa_, Halifax, N.S.; juv., [F], 7425 U.S.N.M. + +Fig. 18. _M. erminea richardsonii_, Ft. Chimo; ad. [F], 14866 U.S.N.M. + +Fig. 19. _M. frenata noveboracensis_, Mich., July 7, 1913; ad. [M], +44689 M.Z. + +Fig. 20. _M. f. frenata_, Brownsville, June 1, 1892; yg. [M], 34043 +U.S.N.M. + +Fig. 21. _M. frenata panamensis_, Panamá, February 17, 1911; sad. [F], +type. + +Fig. 22. _M. a. africana_, Pará, Brazil, Sept., 1908; yg. [M], 37475 +A.M.N.H. + +Figs. 17, 18 and 19. Drawn from specimens preserved in alcohol. + +Figs. 20, 21 and 22. Drawn from relaxed feet of dried skins.] + +_Mustela africana_, all characters considered, is the most aberrant of +the American weasels. That is to say, greater difference prevails +between _M. africana_ and any other American weasel than exists between +any other two American weasels. The distinctive cranial and dental +characters, excepting the reduction in number of premolars, are of a +primitive nature. For example, 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_, the species which is regarded as most closely resembling the +stem form. Also, the mentioned characters in adults of _M. africana_ +resemble ontogenic stages passed through by other weasels. +Consequently, it is thought that _M. africana_ crossed the +filter-barrier from North America to South America, remained isolated +from the original stock for a length of time sufficient to permit +_africana_ to differentiate from North American weasels and _vice +versa_ to such a degree that crossbreeding with the _frenata_ stock was +prevented when _frenata_, at a later time, pushed southward over the, +then zoölogically less-effective, water barrier, or continental bridge +if it was by this time in existence. + +[Illustration: FIG. 23. Diagram indicating probable relationships of +the species of American weasels.] + +The four full species of American weasels may well be thought of as +having the same stem form of which _erminea_ is the most nearly direct +descendant. Geographic and climatic changes may have operated to +isolate, and then to foster morphologic differentiation of, first +_rixosa_ in Eurasia, next _africana_, third the _tropicalis_ section of +_M. frenata_, and finally _M. frenata_ itself, leaving _M. erminea_ as +a modern version, somewhat altered to be sure, of the stem form. Some +of these ideas are expressed in figure 16. The climate is different in +the ranges of the several species and the climate has changed through +time in the ranges of at least many subspecies. Natural selection of +morphological features best adapted to a particular kind of climate +probably has altered some species more than others. _M. erminea_ in +almost every one of its characteristics is generalized and potentially +progressive whereas _africana_ retains more characters which are truly +primitive along with a few which are specializations. _M. africana_ is +potentially the least progressive of any of the American weasels. The +most specialized weasels are the North American races of _Mustela +frenata_. A progressive series of increasing specialization is +comprised in (1) _M. africana_, (2) the _M. tropicalis_ (Central +American, lowland) section of _M. frenata_, and (3) the races of _M. +frenata_ in North America. + +Considering now features of the environment which have obviously +influenced the distribution and speciation of weasels, water barriers +are important. Bering Strait, Carquinez Strait (along with San +Francisco Bay) which opens through the Golden Gate, and the channels +between the islands of southeastern Alaska, have contributed to the +formation of subspecies. The difference is really slight on the two +sides of Bering Strait and San Francisco Bay and is slightly more on +two sides of each of several of the channels between the islands of +southeastern Alaska. The differences between the weasels on the two +sides of one of these water barriers supposedly result from the +preservation in animals on one side, or on one island, of small +mutations, which would be swamped by crossbreeding if the water barrier +were not present. The effect of this isolation is easily seen if +ermines from the Queen Charlotte Islands are compared with those of the +opposite mainland. The degree of morphological difference is great. +Isolationwise, the Queen Charlotte Islands are the seaward end of a +chain, beginning with Admiralty Island in southeastern Alaska, and are +farther from the mainland, zoölogically, than the distance in actual +miles across the water channel would suggest. Between any two islands +that are geographically consecutive, however, and between the mainland +and the first island of the chain, the difference in the ermines is +small. In other places, water barriers of equal or greater width have +contributed little if anything to the differentiation from one another +of weasels on the two sides of the water barrier. The strait between +eastern Canada and Newfoundland is an example. + +The absence of water, or scarcity of it to a degree that closely +approaches absence, in any large area appears to prevent weasels from +living there. At any rate, the one sizeable region of North America +from which weasels are unknown is the desert of the southwestern United +States and adjoining part of northwestern México. More precisely, in +western Arizona, the Mohave Desert and the desert of northwestern +Sonora, collectors of mammals have repeatedly sought small carnivores +without ever finding any weasels. + +Degree of moisture is closely correlated with color in weasels. +Humidity and cloudiness as well as actual precipitation seem to be +involved. Even if we take into account average annual rainfall alone, +the darkest-colored weasels are found in the areas of heaviest rainfall +and the lightest-colored weasels in areas of lightest rainfall (extreme +type of desert where no weasels occur being excepted). In any large +region where there is a geographic gradient in rainfall, the transition +from light to dark color almost exactly parallels the increase in +amount of rainfall. Within a given species the same color reappears in +widely separated areas that have the same amount and seasonal +distribution of rainfall. This correlation is repeated so often that +one can almost certainly say that heavy rainfall, or the associated +phenomena of high humidity and cloudiness, acting separately or +together, causes an increase in intensity of color. Relative extent of +the color of the upper parts and underparts and presence and absence of +light facial markings seem also to be correlated, in a more general +way, with differences in rainfall. A fuller discussion of the nature +and amount of the variation in color is given on page 51. + +Temperature seems not to be an important factor in directly limiting +the distribution of weasels, since _M. frenata_ occurs from the hottest +to some of the coldest parts of the Americas. Do _M. erminea_ and _M. +rixosa_ range no farther south, than they do at present, because high +temperatures constitute a barrier? No evidence is known to me which +provides an answer, one way or the other, to this question. Granting +that temperature is unimportant in limiting the distribution of +weasels, it seems to cause geographic variation. Increase in mean +annual temperature is correlated with decreased size in _M. erminea_ +and with increased size in _M. rixosa_. Temperature, it seems, causes +the hair to vary; the pelage is harsher and sparser in weasels from +tropical regions than in those from boreal regions. Difference in +number of hairs is especially well shown on the soles of the feet. In +the weasels from the far north, the pads are concealed by hair and in +the weasels from the tropical regions the soles are mostly bare. Also, +the hair on the soles of the feet is longer in northern than in +southern weasels. Furthermore there is seasonal change in length of the +hair on the soles of the feet; at a given locality in southern Canada +the hair of the white winter coat is so long on the soles of the feet +as to obscure completely the palmar and plantar pads whereas the hair +of the brown summer coat is shorter and leaves these pads boldly +exposed to view. This seasonal change, as would be expected, is most +marked in animals of northern regions and is not perceptible in those +from the tropics; it is correlated with increase in seasonal change as +the distance from the equator increases. + +Temperature and moisture acting together may cause extensive white +facial markings, that neither alone would cause. In _Mustela frenata_ +these markings occur where there is heavy rainfall and high mean annual +temperature. Where there is heavy rainfall and a low mean annual +temperature they do not occur and where there is high mean annual +temperature and light rainfall the markings are not pure white but are +of the same color as the underparts. Plate I and the description of +color on page 51 may be consulted in this connection. Extremely high +mean annual temperature together with extremely heavy rainfall may +inhibit the development of light facial markings. _M. f. meridana_, +_panamensis_ and _costaricensis_ are cases in point. In either +direction, north or south, from the territory inhabited by these three +subspecies a similar combination of temperature and rainfall is found +and similar light facial markings appear there. + +Considering the delicate response of structure to climate, a person +naturally questions whether or not natural selection accounts for all +of the differences between subspecies. To show that natural selection +determines the color of _Mustela frenata_, it would be necessary to +assume that climate, color, and utility of color are positively +correlated. Although climate (rainfall) and color are correlated in +such a manner that three subspecies of weasel in places as far apart as +New England, Perú, and the state of Washington are colored alike, other +features of the three environments are unlike. Kinds of animals which +the weasel catches for food, and flora in which the weasel finds +concealment, are dissimiliar. If natural selection alone determined the +color, some difference in color would be expected between the weasel +which needed to be obliteratively colored, that is camouflaged, the +better to catch a _Phyllotis_ in Perú and the weasel in Washington +which needed nature's aid in catching _Microtus_. _Mustela frenata +goldmani_ of the highlands of southern México, which is known to attack +the huge pocket gophers, _Orthogeomys_ and _Cratogeomys_, has a weaker +dental armature than _Mustela frenata texensis_ which does not have to +overcome prey so formidable as does _goldmani_. Equally formidable +enemies endanger _M. f. goldmani_ and _texensis_. Examples of this +nature could be multiplied. Without actually proving anything +concerning selection, these examples give reason for us to suppose that +some characters are not determined by natural selection. + +Another question upon which data obtained from a study of _Mustela_ has +some bearing, is this: Where the geographic ranges of two subspecies +meet, why does not the swamping effect of crossbreeding cause one +subspecies to disappear? Although swamping may have occurred in some +instances, it does not occur in the majority of instances. Witness the +long-continued existence of the living subspecies _Mustela frenata +nevadensis_ of which skulls are available from Pleistocene deposits. +Therefore, its distinctive characters, cranially at least, have been +maintained for a long time. Furthermore, these characters are +maintained over a large geographic region more than a thousand miles +across. On the eastern margin of its range, at the eastern base of the +Rocky Mountains in Colorado, _M. f. nevadensis_ intergrades in a +relatively narrow belt with the lighter-colored, longer-tailed and +cranially different _Mustela frenata longicauda_, which has a +geographic range almost equally extensive. _M. f. longicauda_ also is +uniform in its characters over a large area but at approximately 400 +miles east of the base of the Rocky Mountains, it begins to intergrade +with the darker-colored, shorter-tailed and cranially different +_Mustela frenata primulina_ and does so over a belt of 100 miles or +more in width. At any given locality within this wide belt of +intergradation the range of individual variation ordinarily does not +exceed that in animals from a given locality well within the geographic +range of _M. f. longicauda_. In the narrow belt of intergradation along +the eastern base of the Rocky Mountains, the range of individual +variation at several places is greater than in animals from a given +locality well within the geographic range of _M. f. longicauda_ or for +that matter from well within the geographic range of _M. f. +nevadensis_. + +Considering the dominance and recessiveness of genes and the genetic +mechanism in general by which characteristics of offspring are +inherited from their parents, it would seem that _M. f. longicauda_ and +for that matter _M. f. nevadensis_ and _M. f. primulina_ would lose +their distinctive characteristics because of the crossbreeding that is +every year going on between _longicauda_ and _nevadensis_ on the one +hand and between _longicauda_ and _primulina_ on the other hand. + +Sumner (1932:84) suggests that homogeneity is prevented by population +pressure. Applying his suggestion to the species _Mustela frenata_ we +could say that the subspecies _longicauda_ pressing westward meets +strong pressure from the subspecies _nevadensis_ pressing eastward and +that the width of the zone of intergradation between the two subspecies +varies inversely with the strength of the population pressure from the +two sides. Sumner recognizes that according to his hypothesis the two +contiguous races would remain distinct only so long as there was a +preponderance of centrifugal movement from both of the centers of +dispersal. Sumner (_op. cit._:85) recognizes that an abrupt change of +environmental conditions could account in part for the boundaries of +the ranges of the two subspecies and finally that his hypothesis does +not certainly answer the question of why crossbreeding does not result +in homogeneity between two subspecies with contiguous geographic +ranges. + +The hypothesis of harmoniously stabilized complexes of genes was +offered by Timofeeff-Ressovsky (1940:124) to explain why the swamping +effect of crossbreeding does not obliterate subspecies. The hypothesis +takes into account that any one of several characters of a subspecies +may be caused by several genes. Some characters of this kind may be +favored by natural selection more than others. In the belt of +intergradation between two subspecies, where two of these favored +characters meet, a "biological tension" as Huxley (1939:415) terms it +"will result, which will produce _partial discontinuity_ between the +two groups. Each group will evolve a gene-complex which is not only +broadly adapted to the external environment of the central area of its +range, but is also harmoniously stabilized, in adaptation to the +internal genetic environment, by the selection of modifiers." Crosses, +that is to say intergrades, between the two subspecies will lack this +stabilization and will therefore be at a selective disadvantage. The +zone of intergradation will therefore remain narrow; intermediates are +constantly being brought into existence there by crossing but are as +constantly being extinguished by selection. + +These two hypotheses are the best that geneticists yet have offered. +Neither has been tested and both, as originally proposed, would hardly +apply everywhere because there are some contradictions. + +I can offer no better explanation--in fact no original one as good--but +would emphasize that under similar climate, weasels remain constant in +character, or at most do not vary beyond certain limits. Crossing at +the margins of ranges of two subspecies does not result in homogeneity +of weasels. There is, therefore, some stabilizing influence, or +influences, that maintain, and even develop, structural characteristics +of weasels in opposition to the contrary tendency of crossing. + +That this influence not only maintains uniform characters over areas of +large extent, but also permitted their development over large +geographic areas, must logically be supposed, for otherwise, +considering the swamping effect of crossing, such variations would not +have made their appearance in more than a few individuals. Also, if the +races had been formed in response to some kind of physiological +differentiation, or other non-climatic cause, the characters of the +population in the belt of intergradation probably would disappear in a +short time. In any event the close correlation between degree of change +in weasels and degree of change in climate, at once makes one suspect +that climate has been the deciding factor. Finally, when one recalls +that in certain parts of the animal, certain characters invariably +appear under similar climates and never under dissimilar climates, the +evidence is almost conclusive that, given long enough time, the animals +vary in response to climate. The variations (characters) may be induced +indirectly, but are no less exactly reproduced than if they can be +shown to be induced directly. + +In considering how the species and subspecies of American weasels were +formed and in attempting to account for some of the individual +characters, it is profitable to view the facts in the light of some of +the theories of species-formation--theories that are accessory to that +of organic descent and that are concerned with the _modus operandi_ of +organic descent. + +In any group of closely related species some of them, by the laws of +chance, are almost certain to be more primitive than others. _Mustela_ +is no exception and the more primitive species closely match, in +several characters, ontogenetic stages passed through by more advanced +species. Jaeckel's (1902) theory of metakinesis, therefore, is to be +considered since it postulates that many cases of epistasis occur; that +is to say, that many sexually adult animals are arrested in +development in early otogenetic stages and undergo no further +development. Although this theory is appealing upon initial +consideration, it is less so when we recall that in _Mustela_ there is +a direct correlation of increasingly primitive structure with +decreasing latitude as one proceeds from the steppe of North America +southward to the equator. It follows that the conditions seen in +_Mustela_ can be explained even better than by metakinesis, by assuming +that the several species have differentiated from a stem form at +different times, have developed at different rates, have developed in +different directions and that ontogeny recapitulates phylogeny. + +The theory of Age and Area (see Willis, 1922) holds that the species of +widest distribution are, on the average, the oldest, and that the +species which are distributed over small areas are, in general, of +recent origin. So far as the weasels are concerned, little support is +given to this theory. The same can be said of any one of the teological +theories, including the orthogenesis of post-Darwinian writers. All of +these imply a determinate line of variation controlled by the inherent +qualities of the organism. The idea that the several species of +_Mustela_ result from mutations of large degree and sudden appearance +is contrary to the evidence accumulated. In fact the evidence rather +clearly indicates that the mutations which may have occurred were of +small degree and in most instances owe their preservation to natural +selection. + +The data obtained by the study of weasels accords almost exactly with +the theory of species-forming embodied in Matthew's (1915) "Climate and +Evolution." Although the essential features of this theory were made +out from a study of families and orders and therefore would not be +expected to apply to members of only a genus or subgenus, the facts +known about the present distribution of American _Mustela_, +nevertheless, are strikingly in accord with the ideas advanced by +Matthew. In the first place, climate is an important factor in the +evolution of the weasels. In the second place, the line of migration +seems to have been outward from the holarctic region. In the third +place, the geographic changes necessary to explain the present +distribution of the species of _Mustela_ are not extensive and do not +affect the permanency of oceans as defined by the continental shelf. +These three statements are, almost verbatim, those made in the first +three of the five points of Matthew's (1915:172-173) thesis. The +remaining two points of Matthew's thesis have to do with +generalizations based on evidence obtained from sources outside the +scope of the present study. + +Furthermore, the relative degrees of specialization of the different +species and subspecies in relation to their geographic distribution are +in accord with the ideas elaborated by Matthew. For instance, the most +primitive species is farthest south from the probable center of +dispersal, the holarctic region. Also the full species become +progressively more primitive as one proceeds southward from the +holarctic, or at least from the northern half of the nearctic, region. +Although, in view of the known geological changes that have occurred in +the Caribbean region, we cannot say that the more primitive species owe +their positions entirely to having been pushed farther south from the +center of dispersal by actual and _continuous_ contact and competition +with the more advanced species, this seems to have been the case in a +general way. At any rate the more primitive kinds seem to have been +prevented from pushing northward by the more advanced kinds which +developed there and the latter have actually pushed southward. + +Additionally and in review: There is strong indication that the +American species of weasels were formed by gradual and slow change. +Much of this change probably is the result of natural selection +operating on fortuitous variations of a minor nature, but, also, +particular features of the environment, especially climate, and more +especially amount of rainfall, seem to compel variations that +differentiate subspecies and that characterize full species--compel +some of them without the direct operation of natural selection, or at +least compel them within limits so wide that natural selection exerts +no exact control. + + + + +HISTORY OF CLASSIFICATION + + +In the earlier accounts of American weasels, from the time of Linnaeus +and before, up until 1890, names then in use for European weasels +frequently were applied also to those in North America. For the next 50 +years, and almost without exception after 1896, the American weasels +were regarded as specifically distinct from those in the Old World. In +this 50-year period many new names were proposed, usually as full +species, although now that material from more localities has been +brought together and studied, geographic intergradation is evident +between many of the named kinds and most of these names now therefore +take only subspecific rank. In 1933 Glover M. Allen showed that +_Mustela rixosa_ occurred also in the Old World, and in 1943 I +emphasized that a second American species, _Mustela erminea_, was +circumpolar in distribution. In neither _rixosa_, nor _erminea_, +however, were the subspecies the same in the two continents. To this +general outline of the nomenclature, exception must be made for weasels +of the southwestern United States, México and Central America, and +South America, because as early as 1813 a distinctive name was given to +one of these and weasels from the three areas mentioned were, so far as +I know, never given names of Old World kinds. + +The first paper that could be regarded as revisionary in nature was +"Remarks on the species of the genus Mustela" by the zoölogist and +world-traveler, Charles L. Bonaparte, in Charlesworth's Magazine of +Natural History, for 1838. In that paper three new names, _Mustela +cicognanii_, _M. richardsonii_ and _M. longicauda_, all still valid, +were proposed for American weasels. + +Audubon and Bachman in their "Quadrupeds of North America," which +appeared in parts from 1845 to 1853, recognized 5 species. Actually +they were dealing with only 3 taxonomically valid kinds. For one of +these, _Mustela frenata noveboracensis_, they were misled by the +difference in size between males and females, and in the males by the +presence of a brown coat in some and a white coat in others. The male +that was white in winter they regarded as _Putorius ermineus_ of the +Old World; the male that was brown in winter they designated by their +earlier proposed name _P. fuscus_, and the female they named _P. +agilis_. The ermine, subspecies _M. erminea cicognanii_, they called +_P. pusillus_. Their fifth name, _P. frenatus_, included at least some +animals that today are assigned to the subspecies _M. frenata frenata_. +Each of three and perhaps four of the five names employed by Audubon +and Bachman embraced individuals of more than one species and in that +sense the names were composite. + +Only five years later, in 1858, Professor Spencer Fullerton Baird's +great work, "The Mammals of North America," made it clear that no +American weasel was identical (in the modern subspecific sense) with +any Old World weasel, and he applied most of his names in a correct +zoölogical sense. It is true that he thought that the female weasel of +the eastern United States was specifically different from the male, +misapplied to it the name _richardsonii_, and did not correctly +allocate every one of the few poor specimens available to him of the +little ermine (_M. e. streatori_) of the Pacific Coast; but he did +recognize that the least weasel was a distinct kind and his treatment +in general was excellent. + +After Baird came a period of great confusion in which most writers did +no better than had Audubon and Bachman, ordinarily confusing the two +sexes as different species, and, in 1877 in his "Fur-bearing Animals," +Elliot Coues went rather to the other extreme and allowed only 4 kinds +to all of the Americas, regarding two of these, for purposes of +zoölogical nomenclature, as identical with the European species. + +But, in 1896 Outram Bangs published "A Review of the Weasels of Eastern +North America" in which he correctly recognized eight kinds. Although +some of these were treated by him as full species, whereas the material +accumulated since 1896 has shown that subspecific status is in order, +his names, still in use, were correctly applied in every instance, save +probably one. This was his use of _Putorius richardsonii_ for the +animal now known as _M. e. arctica_. Unlike the earlier, excellent +treatment by Baird, this accurate one by Bangs was heeded and followed +by subsequent writers. For example, Dr. C. Hart Merriam in the same +year, 1896, accepted Bangs' conclusions except for correcting the +application of the name _richardsonii_. The principal contributions of +Merriam's paper "Synopsis of the Weasels of North America" were first, +the wider geographic scope and second, the naming as new of several +kinds outside the geographic area studied by Bangs. Otherwise the work +was not up to Dr. Merriam's usual standard and the internal evidence of +haste in its preparation and the superficial study of some of the +material at his disposal explain why the weasels of North America since +that time have been but little better understood than in 1896. Baird +and Bangs, then, unquestionably did the best systematic work on the +American weasels. + +In 1916 Dr. Joseph A. Allen published a valuable paper on the South +American weasels. The material available to him was inadequate and +prevented a thoroughly satisfactory treatment. There are too few +specimens even today to permit of a thorough treatment of the South +American weasels in the present paper; nevertheless the material today +is more nearly adequate than it was in 1916 and it is hoped that the +systematic arrangement is correspondingly improved. + + +Chronological List (annotated) of Specific and Subspecific Names +Applied to American Weasels + +At least eighty-seven specific and subspecific names have been proposed +for American weasels. Of these sixty-nine are now regarded as valid +designations of recognizable subspecies. The average is 1.2 names per +subspecies. Some names in the following chronological list were a +second time applied wholly or in part to some other kind of weasel. In +general, mention of the second or any other later application is +omitted from the following list but two usages of _agilis_ (1844 and +1853) and of _americana_ (1865) are recorded. + + 1734. =javonica= (_Mustela_) Seba, Locupletissimi Rerum naturalium + Thesauri ..., 1:77, 78, pl. 48, fig. 4. The weasel to which this + name was applied was said to have come from Java. Since no animal + answering to the description has again been found in Java, and + because specimens from Central America or possibly some from + northern India, may do so, it is conceivable that Seba was the + first to distinguish by name an American weasel from those in the + Old World. My attempts to locate the specimen concerned in places + where it might have been preserved along with some of the other + specimens thought to have belonged to Seba have been fruitless. + Since it is impossible positively to link Seba's description with + any known weasel, no further use is made of the name _javonica_ in + the present account. + + 1772. =erminea= (_Mustela_) Forster [= _Mustela erminea + richardsonii_], Philos. Trans., London, 1772:373. Forster's use of + the name is one of the earliest applications of it to American + animals. The name dates from Linnaeus, Syst. Naturae, (10th ed.) + 1:46, 1758, with type locality in Europe. In the subspecific sense + the name applies to the ermine which occurs over most of the + Scandinavian Peninsula, if Miller (1912:387) be followed in + regarding the type locality as Upsala, Sweden. If, instead, + Cabrera (1913A:394-396) be followed in regarding the type locality + as in Switzerland, the name, in the subspecific sense, will apply + to the ermine of continental Europe. As the earliest available + name applied to the circumpolar species concerned, it is used now + as the name of the species in the New World as well as in the Old + World. From the time of Forster until approximately 1890 the name + _erminea_ by many, but not by all, authors was applied to the + American weasels in the belief that they were zoölogically + indistinguishable from those in the Old World. From 1896 to 1943 + the name was not used by American authors at all because the + ermine of America was in 1896 treated nomenclaturally by Merriam + as specifically distinct from the animal in the Old World. Since + 1943 _erminea_ has been used in the specific sense for American + animals in recognition of the circumpolar distribution of the + species. Some of the early allocations of American specimens to + _erminea_ probably resulted in a composite use of the name in that + one or another subspecies of the American species _Mustela + frenata_ may also have been included with individuals truly of the + species _erminea_. + + 1772. =nivalis= (_Mustela_), Forster, Philos. Trans., London, + 1772:373. This is one of the early applications of this name to + American weasels of small size, made in the belief that they were + taxonomically the same in America and Europe. Linnaeus, Syst. Nat. + (12th ed.) 1:69, 1766 is the authority for the name [_Mustela_] + _nivalis_, and the Province of Vesterbotten, Sweden, is regarded + as the type locality. The name is in use today for the common + weasel of Europe and parts of Asia. Animals of the species + _nivalis_ are intermediate in size between _Mustela erminea_ and + _Mustela rixosa_. The name as used for American animals by some + authors who wrote later than Forster did, probably was composite + in that these authors may have applied the name to the small + weasels of North America and thus may have intended it to apply + not only to _Mustela erminea cicognanii_ but also to females of + _Mustela frenata noveboracensis_, and conceivably to both sexes of + _Mustela rixosa_ of any American subspecies. + + 1813. =Brasiliensis= (_Mustela_) Sevastianoff, Mem. Acad. Imp. + Sci. St. Petersburg, 4:356-363, table (= plate) 4. This name was + proposed for a weasel brought to St. Petersburg by Capt. + Krusenstern on his return from a voyage around the world. The + animal was said to have come from Brazil, but to judge from the + description, came instead from México, Central America, or west of + the Andes in South America, and was based on some one of the + subspecies of _Mustela frenata_. Although the name was in use for + more than 60 years it was shown by Merriam (1896:27) to be + unavailable because it was preoccupied by _Mustela brasiliensis_, + a name earlier used by Gmelin (Syst. Nat., ed. 13, p. 93, 1788) + for a South American otter. + + 1815. =vulgaris= (_Mustela_), Ord, Guthrie's Geography as + reprinted by Rhoads in 1894, vol. 2, p. 291. This use by Ord is + one of the earliest applications of this name to American weasels, + in the belief that the smaller weasels of North America and Europe + were zoölogically the same; [_Mustela_] _vulgaris_ seems + originally to have been proposed in 1777 by Erxleben on p. 471 of + vol. 1 of his Syst. Regni Anim., for the weasel of the temperate + part of Europe and to be a synonym of _Mustela nivalis_ Linnaeus + (1766). Probably the name as used by Ord was composite in the + sense that he may have intended it to apply to females of _Mustela + frenata noveboracensis_ as well as to one or both sexes of + _Mustela erminea cicognanii_ and, if he ever saw them, to the two + sexes of _Mustela rixosa_ (one or several subspecies). + + 1818. =africana= (_Mustela_) Desmarest [= _Mustela africana + africana_], Nouv. Diction, d. Hist. Nat., 19:376. In 1808 E. + Geoffroy St.-Hilaire visited Portugal and was given several + African primates and the specimen of _Mustela_ named by Desmarest + in 1818 who wrongly supposed that it, like most of the primates, + came originally from Africa. After the name had been misapplied + for 95 years Angel Cabrera showed that it pertained instead to the + tropical weasel of Brazil. Of distinctive names applied to + American weasels today, this is the one first proposed. + + 1832. =frenata= (_Mustela_) Lichtenstein [= _Mustela frenata + frenata_], Darstellung neuer oder wenig bekannter Säugethiere, pl. + 42 and corresponding text unpaged. This name is the first one + available for the long-tailed weasel and therefore applies to the + species as a whole. + + 1838. =Cicognanii= (_Mustela_) Bonaparte [= _Mustela erminea + cicognanii_], Charlesworth's Mag. Nat. Hist., 2:38. The name + erroneously spelled _Cigognanii_ was correctly spelled on page 39. + For a detailed consideration of this name see the account of the + subspecies _cicognanii_ on page 120. + + 1838. =Richardsonii= (_Mustela_) Bonaparte [= _Mustela erminea + richardsonii_], Charlesworth's Mag. Nat. Hist., 2:39. Until 1896 + the name sometimes was applied to the subspecies now known as _M. + e. arctica_ and sometimes to part of the subspecies now designated + as _M. e. cicognanii_ under the principal treatment of which see + (page 120) for a detailed account of the basis of the name + _=richardsonii=_, and the reasons for regarding Fort Franklin as + the type locality. + + 1838. =longicauda= (_Mustela_) Bonaparte [= _=Mustela frenata + longicauda=_], Charlesworth's Mag. Nat. Hist., 2:39. The type + locality appears to be Carlton House, Saskatchewan, and the name + always seems to have been applied to the long-tailed weasel of the + Great Plains, although in some earlier accounts the name was used + in a more inclusive sense to refer also to animals now of + subspecies closely allied to _longicauda_. As with the two + preceding names, a detailed consideration of the basis for, and + application of, this name is given on pages 120-123 in the account + of _Mustela erminea cicognanii_. + + 1840. =Noveboracensis= (_Putorius_) Emmons [= _Mustela frenata + noveboracensis_], Quadrupeds of Mass., p. 45. This name was + credited by Emmons to De Kay who in the same year published it in + his report on the "Zoology of New York" but without a description + and De Kay's name is a _nomen nudum_. Emmons' was the first use of + the name accompanied by a recognizable description and therefore + the name must date from Emmons although this obviously was not his + intent since he credited the name to De Kay. + + 1842. =fuscus= (_Putorius_) Audubon and Bachman [= _Mustela + frenata noveboracensis_], Jour. Acad. Nat. Sci., Philadelphia, 8: + (pt. 2) 288. + + 1842. =pusilla= (_Mustela_) De Kay [= _Mustela erminea + cicognanii_], Nat. Hist. of New York, Zool., Pt. 1, Mammalia, p. + 34. This name was proposed for small weasels of 12 to 13 inches in + length of which the tail amounted to a fourth of the same and + although obviously applying in considerable part to the earlier + named _M. e. cicognanii_ seems to have included some individuals + of the also earlier named _M. f. noveboracensis_. + + 1843. =xanthogenys= (_Mustela_) Gray [= _Mustela frenata + xanthogenys_], Ann. and Mag. Nat. Hist., 11:118, February, 1843, + was applied to all of the long-tailed weasels of California that + had light-colored facial markings. Merriam in 1896 suggested that + San Diego was the type locality and in 1899 Bangs proposed the + name _mundus_ for the California weasel north of San Francisco Bay + thus restricting the application of the name _xanthogenys_. In + 1936 Hall further restricted the application of the name and + applied it to the long-tailed weasel of the big interior valley of + California, pointing out that the name was correctly applied to + this weasel of the big interior valley or possibly instead to the + race named _munda_. + + 1844. =agilis= (_Mustela_) Tschudi [= _Mustela frenata agilis_], + Untersuch. ü. die Fauna Peruana, p. 110, is a name applied today + to the race of weasel of the Temperate Zone of the western Andes + and intermountain valleys of Perú. + + 1851. =nigripes= (_Putorius_) Audubon and Bachman [= _Mustela + nigripes_], Quadr. N. Amer., 2:297, 1851, applies to the + black-footed ferret of North America. + + 1853. =agilis= (_Putorius_) Audubon and Bachman [= _Mustela + frenata noveboracensis_], Viv. Quadrupeds N. Amer., 3:184, pl. + 140. This name was proposed for the female in the mistaken belief + that it was specifically distinct from the larger male for which + several names already were available. Also Tschudi in 1844 had + already used the name _Mustela agilis_ for a South American + weasel. + + 1864. =aureoventris= (_Mustela_) Gray [= _Mustela frenata + aureoventris_], Proc. Zoöl. Soc. London, 1864:55, pl. 8, February + 9, 1864, is the name applicable to the dark-colored weasel of the + Pacific coastal region of Ecuador and Columbia. + + 1865. =americana= (_Mustela erminea_ Var. 3) Gray, Proc. Zoöl. + Soc. London, 1865:111. The larger individuals of American weasels + of both _Mustela erminea_ and _Mustela frenata_ from the Atlantic + Coast to as far west as Carlton House, Saskatchewan, were lumped + under this name because Gray desired more information than he then + had before recognizing as different from one another several + species proposed for America up to the time concerned. The name is + unavailable because it is preoccupied by _Mustela americana_ + Turton (1806) the name for the American marten. + + 1865. =americana= (_Mustela vulgaris_ Var.) Gray, Proc. Zoöl. Soc. + London, 1865:113. Under this name the smaller weasels of the + northern and northeastern part of North America were lumped by + Gray but the name is preoccupied and can be ignored. + + 1874. =affinis= (_Mustela_) Gray [= _Mustela frenata affinis_], + Ann. and Mag. Nat. Hist., 14 (ser. 4):375, 1874, from New Granada + [= Colombia], had the type locality restricted to Bogotá, + Colombia, by Allen in 1916, and is applied to the long-tailed + weasel of the tropical and temperate zones of the eastern Andes of + Colombia. + + 1874. =macrura= (_Mustela_) Taczanowski [= _Mustela frenata + macrura_], Proc. Zoöl. Soc. London, for 1874, p. 311, pl. 48, May + 19, 1874, applies to the long-tailed weasel of central Perú and + northern Ecuador. + + 1877. =culbertsoni= (_Putorius_) Coues [= _Mustela frenata + longicauda_], Fur-bearing animals ..., p. 136, 1877, is based on + specimens from Fort Laramie, Wyoming. In the past the name has + been regarded as a _nomen nudum_ but there is some reason for + regarding it as having nomenclatural status. In either event it is + here arranged as pertaining to the long-tailed weasel of the Great + Plains which takes the prior name _longicauda_. See the account of + _longicauda_ for a more detailed account of the name + _culbertsoni_. + + 1877. =aequatorialis= (_Putorius_ (_Gale_) _brasiliensis_) Coues + [= _Mustela frenata aureoventris_], Fur-bearing animals ..., p. + 142. Proposed "merely as a substitute for Gray's [supposedly] + preoccupied name," _aureoventris_. + + 1881. =stolzmanni= (_Mustela_) Taczanowski [= _Mustela africana + stolzmanni_], Proc. Zoöl. Soc. London, for 1881, p. 835, November + 15, 1881, is applied to the tropical weasel of the Upper Amazon + Basin. + + 1881. =jelskii= (_Mustela_) Taczanowski [= _Mustela frenata + macrura_], Proc. Zoöl. Soc. London, for 1881, p. 647, May 17, + 1881, was proposed for the female in the mistaken opinion that it + was specifically distinct from the larger male which the same + author previously had named _macrura_. + + 1891. =arizonensis= (_Putorius_) Mearns [= _Mustela frenata + arizonensis_], Bull. Amer. Mus. Nat. Hist., 3:234, June 5, 1891, + until 1936 was applied to long-tailed weasels of most of the + western United States west of the Great Plains but by restriction + since 1936 has been applied only to the animals in parts of + Arizona and New Mexico. + + 1894. =peninsulae= (_Putorius_) Rhoads [= _Mustela frenata + peninsulae_], Proc. Acad. Nat. Sci. Philadelphia, 1894:152, June + 19, 1894, applies to the weasel of central and southern Florida. + + 1896. =alascensis= (_Putorius richardsonii_) Merriam [= _Mustela + erminea alascensis_], N. Amer. Fauna, 11:12, June 30, 1896, with + type locality at Juneau, Alaska, has been used for the ermine of + southeastern Alaska ever since it was proposed. In 1944 separate + subspecific rank was accorded ermines on several of the islands of + southeastern Alaska which proportionately restricted the range + assigned to _alascensis_. + + 1896. =streatori= (_Putorius_) Merriam [= _Mustela erminea + streatori_], N. Amer. Fauna, 11:13, June 30, 1896, applies to the + ermine of the Pacific Coast from Puget Sound, Washington, south + nearly to the Golden Gate of California. + + 1896. =arcticus= (_Putorius_) Merriam [= _Mustela erminea + arctica_], N. Amer. Fauna, 11:15, June 30, 1896. Ever since it was + proposed, this name has been applied to the subspecies of ermine + of Alaska and the northern parts of Canada. + + 1896. =kadiacensis= ([_Putorius arcticus_]) Merriam [= _Mustela + erminea kadiacensis_], N. Amer. Fauna, 11:16, June 30, 1896, is a + valid name applied to the ermine of Kodiak Island, Alaska. + + 1896. =washingtoni= (_Putorius_) Merriam [= _Mustela frenata + washingtoni_], N. Amer. Fauna 11:18, June 30, 1896, applies to the + long-tailed weasel of the southern Cascades of Washington and the + northern Cascades of Oregon. + + 1896. =saturatus= (_Putorius_) Merriam [= _Mustela frenata + saturata_], N. Amer. Fauna, 11:21, June 30, 1896, was little used + until 1936 but applies to long-tailed weasel of limited region in + northern California and southern Oregon. + + 1896. =alleni= (_Putorius_) Merriam [= _Mustela frenata alleni_], + N. Amer. Fauna, 11:24, June 30, 1896, applies to weasel of Black + Hills region. + + 1896. =oregonensis= (_Putorius xanthogenys_) Merriam [= _Mustela + frenata oregonensis_], N. Amer. Fauna, 11:25, June 30, 1896, + applies to long-tailed weasel of parts of western Oregon and + northern California. + + 1896. =goldmani= (_Putorius frenatus_) Merriam [= _Mustela frenata + goldmani_], N. Amer. Fauna, 11:28, June 30, 1896, applies to the + long-tailed weasel of Chiapas, and parts of Guatemala and + Salvador. + + 1896. =leucoparia= (_Putorius frenatus_) Merriam [= _Mustela + frenata leucoparia_], N. Amer. Fauna, 11:29, June 30, 1896, + applies to the long-tailed weasel of Michoacán and Nayarit. + + 1896. =tropicalis= (_Putorius_) Merriam [= _Mustela frenata + tropicalis_], N. Amer. Fauna, 11:30, June 30, 1896, applies to the + long-tailed weasel of the Tropical Life-zone of Veracruz. + + 1896. =spadix= (_Putorius longicaudus_) Bangs [= _Mustela frenata + spadix_], Proc. Biol. Soc. Washington, 10:8, February 25, 1896, + applies to the long-tailed weasel of Minnesota and adjoining + areas. + + 1896. =rixosus= (_Putorius_) Bangs [= _Mustela rixosa rixosa_], + Proc. Biol. Soc. Washington, 10:21, February 25, 1896, applies to + the least weasel of Saskatchewan and adjoining areas and as the + first available name for the species has been used as the specific + name for the species in America since 1896. + + 1897. =paraensis= (_Putorius (Mustela) braziliensis_) Goeldi [= + _Mustela africana africana_], Zool. Jahrb., abt. f. systematik, + geogr. u. biol., 10:560, pl. 21, September 15, 1897, a synonym 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, applies to the long-tailed + weasel of New Mexico, Arizona, Durango and adjoining areas. + + 1898. =haidarum= (_Putorius_) Preble [= _Mustela erminea + haidarum_], Proc. Biol. Soc. Washington, 12:169, August 10, 1898, + applies to the ermine of the Queen Charlotte Islands, British + Columbia. + + 1899. =notius= (_Putorius noveboracensis_) Bangs [= _Mustela + frenata noveboracensis_], Proc. New England Zoöl. Club, 1:53, June + 9, 1899, was applied to the long-tailed weasel of the Carolinas + until 1936 since which time it has been regarded as a synonym of + _noveboracensis_. + + 1899. =occisor= (_Putorius_) Bangs [= _Mustela frenata occisor_], + Proc. New England Zoöl. 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 xanthogenys_) Bangs [= _Mustela frenata + munda_], Proc. New England Zoöl. Club, 1:56, June 9, 1899, is now + applied, and generally has been since 1899, to the long-tailed + weasel of the coastal district of California north of San + Francisco Bay. + + 1899. =muricus= (_Putorius (Arctogale)_) Bangs [= _Mustela erminea + muricus_], Proc. New England Zoöl. Club, 1:71, July 31, 1899, + applies to the diminutive ermine, often erroneously designated + least weasel, of the western United States. + + 1899. =oribasus= (_Putorius (Arctogale) longicauda_) Bangs [= + _Mustela frenata oribasus_], Proc. New England Zoöl. Club, 1:81, + December 27, 1899, applies to the long-tailed weasel of the Rocky + Mountains northward from Yellowstone National Park. + + 1900. =eskimo= (_Putorius rixosus_) Stone [= _Mustela rixosa + eskimo_], Proc. Acad. Nat. Sci. Philadelphia, 1900:44, March 24, + 1900, is applied 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, applies 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, + applies 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 the type specimen was taken instead in northern + Ellesmere Island and because the weasel there is subspecifically + indistinguishable 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 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 [= _Mustela + erminea angustidens_], Mem. Amer. Mus. Nat. Hist., 9(pt. 4):181, + pl. 17, is applied to an extinct subspecies known from fossil + remains of Pleistocene age from northern Arkansas. + + 1908. =gracilis= (_Putorius_) Brown [= _Mustela frenata + gracilis_], Mem. Amer. Mus. Nat. Hist., 9(pt. 4):182, 1908, + applies 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, applies to the long-tailed weasel of Costa Rica. + + 1913. =primulina= (_Mustela_) Jackson [= _Mustela frenata + primulina_], Proc. Biol. Soc. Washington, 26:123, May 21, 1913, + applies 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_) Howell [= _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_) Hollister [= _Mustela frenata + meridana_], 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 nicaraguae_], Bull. Amer. Mus. Nat. Hist., 35:100, April + 28, 1916, applies to the long-tailed weasel of Nicaragua. + + 1927. =arthuri= (_Mustela noveboracensis_) Hall [= _Mustela + frenata arthuri_], Proc. Biol. Soc. Washington, 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 applied to the ermine of Southampton + Island but after 1945 was applied 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 Panamá. + + 1932. =anguinae= (_Mustela cicognanii_) Hall [= _Mustela erminea + anguinae_], Univ. California Publ. Zoöl., 38:417, November 8, + 1932, applies to the ermine of Vancouver Island, British Columbia. + + 1935. =labiata= (_Mustela arctica_) Degerbøl [= _Mustela erminea + semplei_], Rept. 5th Thule Exped., 1921-1924, vol. 2, no. 4, p. + 25, 1935. When Degerbøl 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, Degerbøl's + 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 Perú. + + 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 referred 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, applies 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 Concepcion. 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-México + 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 Perú + and Bolivia. + + 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, southeastern 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 islands, 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, applies 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, Jour. Mamm., 26:84, + February 27, 1945, applies to the diminutive ermine of the + Cascades in Washington. + + 1945. =bangsi= (_Mustela erminea_) Hall, Jour. Mamm., 26:176, July + 19, 1945, is the name applied today to the ermine of the western + Great Lakes region. + +In 1925 when this study was begun, the American weasels (subgenus +_Mustela_ 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 number 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. + + + + +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 bangsi_ 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 + _Mustela 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_ (Mearns) 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) 309 + _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 boliviensis_ Hall 402 + _Mustela frenata gracilis_ (Brown) 404 + + +Subgenus =Grammogale= 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_ Peale 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) + + + + +ARTIFICIAL KEY TO AMERICAN SPECIES OF THE GENUS MUSTELA + + + PAGE + + 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 + condylobasal length. + + C Tail without a black pencil and with at most a few + black hairs at extreme tip; in both sexes mastoid + breadth ordinarily exceeds breadth of braincase, + _Mustela rixosa_, least weasel, p. 168 + + C' Tail with a black pencil; in females mastoid breadth + ordinarily 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; m1 lacking + even a trace of a metaconid; distance between upper + canines more than width of basioccipital as measured + between foramina situated midway along medial sides of + tympanic bullae, + _Mustela nigripes_, black-footed ferret. + + E' Abdomen dark brown, like back; face uniformly brown + without blackish mask; m1 with incipient metaconid; + distance between upper canines less than width of + basioccipital as measured between foramina situated + midway along medial sides of tympanic 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 (moderately in +_Lutreola_), cancellous, and with paroccipital processes closely +appressed to bullae; palate behind upper molars; dental formula: + + I 3 C 1 P 2-3 M 1 + -, -; -, -; -, ---; -, -; inner moiety of M1 larger than outer; P4 + i 3 c 1 p 3-2 m 2 + +with simple deuterocone; in m1 inner moiety of M1 larger than outer; P4 +with simple deuterocone; in m1 trigonid longer than talonid, metaconid +absent (incipiently developed in _Lutreola_), and talonid trenchant. + +For many years prior to 1911, the name _Mustela_ was applied to +martens, and _Putorius_ 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 zoölogical differences between ferrets 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 zoölogists _Lutreola vison_ and by +American zoölogists _Mustela vison_. + + + + +EXPLANATION OF SYSTEMATIC TREATMENT + + +For each full species there will be found under the account of it the +following 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 zoölogical name, synonyms, type, geographic range, +zoölogical characters for ready recognition, description (mentioning +size, certain external features including color, the skull and teeth) +historical material when warranted, remarks which may elaborate on +points made in preceding paragraphs, and other information thought to +be useful, and finally a list of specimens examined. + +In explanation of certain of these categories it should be said that in +the synonymy no attempt is made to list 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 regarded as especially important and +citations to them ordinarily are omitted in the synonymy. No comma is +placed between the zoölogical name and the name of the author who +coined and first used the name in accordance with the rules of +zoölogical nomenclature. Otherwise a comma is interposed between the +zoölogical name and the name of the user (author). When the accepted +(earliest available) name of a subspecies at the head of any one of the +following accounts is combined with a generic name different from that +with which 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, denote 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 distinguishing 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 the 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 the 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 Oberthür 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: I2 denotes the second incisor tooth in the upper jaw and +i2 denotes the second incisor tooth in the lower jaw; C1 and c1 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; M1 and +m1 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 differences 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 subspecies is described. For a subspecies +geographically next adjacent to the one described, only the +differences between the two are enumerated. This 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 +description, 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 osseous 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 surprising 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 +specimens at the time they are prepared. + + + + +SYSTEMATIC ACCOUNTS OF SPECIES AND SUBSPECIES + + +=MUSTELA ERMINEA= Linnaeus + +Ermine + +(Synonymy under subspecies) + + _Type._--_Mustela erminea_ Linnaeus, Systema Naturae, 10th ed., p. + 46, 1758. + + _Range._--From the British Isles and Atlantic Coast of Europe + across Eurasia and North America including Greenland, from the + northernmost land, south, in North America, to the lower margin of + the Canadian Life-zone; geographically south to Connecticut, New + York, Pennsylvania, northeastern Ohio, southern Michigan, + Wisconsin, northern Iowa, Minnesota, North Dakota, in the Rocky + Mountains to northern New Mexico, in the Sierra Nevada to Mono + County, California, 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 +together, 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; underparts whitish, ordinarily +continuous from chin to inguinal region but in subspecies in the humid +region along the Pacific Coast interrupted in some individuals by brown +of upper parts encircling body in the abdominal region. The soles of +the feet in each of the subspecies 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 character 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: 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 underparts, 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 the 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 the 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 maximum 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 the one hand, and either of the largest +races, _texensis_ or _macrophonius_, on the other hand although actual +weights are not available 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 comments "so far as I +know at present, there is no evidence to show that any short-tailed +weasels inhabit a broad strip of treeless territory 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 Buffalo 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 +coniferous belts, and grassy, semi-wooded swamplands. + +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 (_Mustela +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 females. + +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 seldom 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. muricus_, 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 slightly +behind. "The distance normally is about 19 inches, representing 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, soft +snow. Its actions are much like those of an Otter pursuing salmon. +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. Concerning +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. McIlwraith shot a bald +eagle at Hamilton Bay which had the bleached skull of a weasel +(probably of this species) clinging to the throat (Seton, 1929 +(2):603). + +A. B. Howell (1943:98) likens mustelid 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." + + +Food + +The killing of prey is described by Hamilton (1933:332) as follows: "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 lies on its +side, while the diminishing struggles of the rodent continue, 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 buyers 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; _Tamias_, 3.6. Close correspondence 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 abundance. +Snowshoe rabbits and red squirrels were not eaten. The Minnesotan data +were from 60 stomachs and 53 intestinal tracts recovered 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 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 microtine rodents were brought to a den of +the diminutive _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 ermines. + +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 _Microtus_).--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 (_Rattus_).--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_, following 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. + +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 decimation 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 altogether lacking." + +Wild birds (Class Aves).--Aldous and Manweiler (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 _Gallus_).--Criddle and Criddle (1925:145), who +published relatively extensive data on the three 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 the 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 fragments of a +small fish in one of 60 stomachs of ermine from Minnesota. + +Earthworm (Phylum Annelida).--Osgood (1936:64), presumably 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. cit._:321-325) described animals one day old from +New York State as being flesh-colored, having the long neck of the +adult and a fine growth of white hair two millimeters 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 life. 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. + +[Illustration: FIG. 24. _Mustela erminea richardsonii_, adult female, +Catalogue Number 14866, U. S. Nat. Mus., Fort Chimo, Ungava. × 1/2. + +Ventral view of body of a pregnant female to show details of mastology. +Note the five pairs of mammae characteristic of weasels, and the uneven +arrangement of mammae of the two sides which is also common among +weasels.] + +[Illustration: 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. 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 + + _Putorius arcticus_ Merriam, N. Amer. Fauna, 11:15, pl. 2, figs. 1, + 1a, and pl. 5, figs. 6, 6a, June 30, 1896. + + _Putorius_ (_Gale_) _erminea_, Coues, Fur-bearing animals, p. 109, + 1877 (part). + + _Putorius richardsonii_, Bangs, Proc. Biol. Soc. Washington, 10:16, + pl. 1, figs. 3, 3a, pl. 2, figs. 3, 3a, and pl. 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 audax_ Barrett-Hamilton, Ann. and Mag. Nat. Hist., + 13(ser. 7):392, May, 1904, type from Discovery Bay, Ellesmere + Island. + + _Putorius alascensis_, Heller, Univ. California Publ. Zoöl., 5:345, + March 5, 1910. + + _Mustela arctica arctica_, Miller, U. S. Nat. Mus. Bull., 79:97, + December 31, 1912; Dice, Journ. Mamm., 2:22, February 10, 1921. + + _Mustela arctica_, Hall, Univ. California Publ. Zoöl., 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, Journ. 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 obvious signs of fading. + + _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 Colden + 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 measurements 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 + (16_n_) of Ridgway and about tones 2 to 3 of Chocolate of Oberthür + and Dauthenay, pl. 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 antipalmar 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 underparts 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 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 available adult female +of _kadiacensis_, the skull of female _arctica_ is larger in all parts +measured, a fourth heavier, has tympanic bullae of almost 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. Differences 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; measured 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. + +Comparison with _semplei_ is made in the account of that subspecies. + +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. McFarlane, +R. Kennicott, E. W. Nelson and R. M. Anderson are names which appear +commonly. Of Alaskan specimens prepared according 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 +differences. 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, instead of +from Europe, are used the differences mentioned by Merriam 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 _richardsonii_. . . . It is interesting 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 above that +he correctly understood the zoölogical 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 (crossbreeding) ever takes place +across the narrow Bering Strait I do not know. I doubt that +crossbreeding occurs but considering the Diomedes (islands), that might +serve as a half way stopping point, and remembering 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 occasional 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 ascertained was imported as a skin with other furs from Siberia. + +Ognev (_op. cit._) who used the name combination _Mustela erminea +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 Kamchatka) 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 proposed the new name +_Mustela erminea digna_ for the ermine of Kamchatka. In comparing +material of these two Asiatic races with topotypes 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 therein 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 +inseparable from _Putorius arcticus_ Merriam) Barrett-Hamilton +erroneously designated the type locality as "Discovery Bay, North +Greenland" whereas he should have written Grinnell Land [= Ellesmere +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 there, but this +individual was the type specimen of _Mustela 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 the boundary-line between _arctica_ and _richardsonii_ +should be drawn remains to be ascertained. The one specimen available +from Hinchenbrook Island, no. 912 Mus. Vert. Zoöl., 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 between 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 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 Districts 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 (1[1], 1[2], 1[75], 4[1], 7[60], 6[74]); + Flaxman Island, 3; Collinson Point, 1[77]; Salirochet River, + 1[77]; Hulahula River, 1[2]; 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[75]; Kruzamepa, 1[75]; Tanana, 6; Boulder Creek, Chena River, 3; + Fort Reliance, 4; Yukon River, 20 miles above Circle, 2; Mts. near + Eagle, 42 (1[60]); Snake River, Nome, 1[9]; 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[74]; 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[60]; Hope, Cook Inlet, 1; Iak Lake, 1[68]; head + of Behring Riv., 1; Bethel, 2; Kenai Lake, 8; Kenai Peninsula, 13 + (2[2]); He[i]nchenbrook Island, 1200 ft., 1[74]; Sunshine Point, + Kaliekh River, Yakataga Dist., 1[8]; Cape Yakataga, 3[8]; Yakutat + Bay, 3[74]; Seward, 7; Seldovia, 22 (4[2]); Homer, 1[2]; 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[100]; Mt. Pavlof, 1[75]; Unimak + Island, 2 (1[75]). + + =District of Franklin.= Cape Sheridan, 1[2]; Discovery Bay, + Ellesmere Island, 1[7] (type specimen of _Putorius audax_ + Barrett-Hamilton); Axel Heiberg Island, 1[95]; Bache Peninsula, + Ellesmere Island, 1[77]; Bedford Pims Island, 4[75]; Craig Harbor, + 2[77]; Cape Kellett, Banks Island, 1[77]; Franklin Isthmus, 1[95]; + King William Island, 2[95]. + + =District of Keewatin.= Ualiak, Ogden Bay, 2[95]. + + =District of Mackenzie.= Baillie Island, 1[75]; Franklin Bay, 1; + Langton Bay, arm of Franklin Bay, 15 mi. S of, 1[2]; Cockburn + Point, 69°N, 115°W, 2[77]; Dolphin and Union Strait, 1[77]; + Bernard Harbor, 2[77]; Kent Peninsula, 4[95]; Horton Riv., near + Fort Anderson, 1; Fort Anderson, 6; Anderson River, 3; Barry + Island, Bathurst Inlet, 1[77]; Fort McPherson, 1; Peels River, 2; + Arctic Red River, 8[75]; Fort Good Hope, 6; Clinton Colden, 1[2]. + + =Yukon.= Kamarkak, 1[77]; Herschel Island, 1[75]; Lapierres House, + 2; Forty Mile, L. T. Coal Cr., 4[74]; head of Coal Cr., 1; + Macmillan River, Forks, 1; 20 mi. W. Ft. Selkirk, 1; Slims River, + near Kluane, 1[75]; head of Lake Lebarge, 1. + + +=Mustela erminea polaris= (Barrett-Hamilton) + +Ermine + + _Putorius arcticus polaris_ Barrett-Hamilton, Ann. and Mag. Nat. + Hist., 13 (ser. 7):393, May, 1904. + + _Mustela erminea_, Manniche, Meddelelser on Grønland, 45:80-85, 1 + fig., 1910. + + _Mustela arctica polaris_, Miller, U. S. Nat. Mus. Bull., 79:97, + December 31, 1912. + + _Mustela erminea polaris_, Hall, Journ. Mamm., 26:179, July 19, + 1945. + + _Type._--Probably female, skin only; no. 78. 6. 19. 11, Brit. Mus. + Nat. Hist.; Gap Valley, 7-1/4 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 [_j_] 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 Scøresby 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 life. + + _Color._--As described in _Mustela erminea arctica_ except that + upper parts in summer near (_j_) Buckthorn Brown and tone 4 of + Dark Fawn of plate 307 to tone 1 of Raw Umber of plate 301 of + Oberthür 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 lighter-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 + measurements. 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 + measurements. As described in _Mustela erminea arctica_ except + that basilar length 36.8 (35.9, 37.8), and length of tooth-rows + not more than length of tympanic 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_. + +_Remarks._--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) relative 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 catalogue 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 attached to the specimen +does however give the locality as "Hall Land" "N Greenland" and another +label has on it "Ermine, procured 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 Island], +across Robeson Channel and along the north coast of Greenland to the +west base of Mount Farragut near 50° 30´ W he mentions the ermine only +this once. For several other kinds of animals, Beaumont mentions +individuals seen or shot, often with the notation 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 Expedition, 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 7-3/4 miles northeast of Cape +Brevoort, one camp on either side of the 82° line, and separated from +each other by a distance of only 2-1/4 air line miles or 4-1/2 miles +march according 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 +beginning of this account of the subspecies. + +The light-colored upper parts and more intensely yellow underparts well +differentiate this subspecies from _arctica_ or _semplei_. +Intergradation is suggested by a skin, no. 1462, Copenhagen Zoological +Museum, from Axel Heibergs Land, the color of the underparts 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 and 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 Baffin +Island 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 _Mustela erminea_. + +The best material of this subspecies is in the University Zoological +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 Universitetets Zoologisk Museum, Købnhavn, + Danmark. + + Gap Valley, 7-1/4 mi. NE Cape Brevoort, 82 N, 59 20´ W, 1 (British + Mus.); Dragon Point, 1; Danmarks Havn (Fjeldene ved Baadskjeret, + 1; lille 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, Jamesonsland, 1 (Berlin 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_ Degerbøl, Rept. 5th Thule Exped., 2 (no. + 4):25, 1935, type from Malugsitaq, Melville Peninsula, Canada. + + _Mustela erminea semplei_, Hall, Journ. Mamm., 26:179, July 19, + 1945. + + _Type._--Male, subadult, skull and skin; no. 6470, Carnegie Mus.; + Coral Inlet, 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 × 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 off. 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 _M. 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 Primrose 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 + extreme 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 Island): 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 breadth more or less than (approximately same as) + distance between last upper molar and jugular foramen. + +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 M1 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 averages 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 mainland west +of Southampton Island may owe their smaller (than in _arctica_) size to +intergradation with _richardsonii_ almost as much as to intergradation +with _semplei_. + +Degerbøl's name _Mustela arctica labiata_ was applied to specimens, +which to me are indistinguishable from topotypes of _Mustela arctica +semplei_, which latter name has three years priority. Degerbøl +(1935:34) states that Malugsitaq, Melville Peninsula, is the type +locality. He did not designate a type specimen. Reference to his +account (_op. cit._:26) shows that he lists five specimens from the +type locality, or more precisely as "Malugsitaq, Lyon Inlet. 5 summer +skins. [M] [M] June-July 1922. P. F., CN. 2262-2266." On labels +attached to these specimens, "Lyon Inlet" is replaced with "Melville +Peninsula." On July 28, 1937, Degerbøl and I together examined these +specimens in his laboratory. Because no. 2262 is first mentioned I +regard it as the type. It is a juvenal male, skull and skin, no. 2262 +(20.5 1931.8), Univ. Zool. Mus. Copenhagen, obtained in June or July of +1922 by Peter Freuchen whose original number was / s 2324. The specimen +is one of 5 males taken at the same locality by the same collector and +they bear identical data as to date. They look to be of the same +litter for all are roughly of the same size and each retains milk +teeth. + +Additional females, with external measurements carefully taken, are +much needed from Southampton Island, because the available females are +insufficient to show the degree of sexual dimorphism. If the meager +data available be accepted, the difference in size between the two +sexes is less than in other subspecies. My own feeling is that a better +sample of females would show the secondary sexual difference in size to +be more than available data indicate. + + _Specimens examined._--Total number, 183, arranged from north to + south by islands, or regions attached to the mainland, and from + north to south in each region or island. Unless otherwise + indicated, specimens are in the Zoological Museum, University of + Copenhagen, Denmark. + + =Baffin Island.= Pond[s] Inlet, 8; (5[77]); Tulukan (sometimes + spelled Tulukat), 6; Cape Eglinton, 1[7]; Gifford River, 2; Clyde, + 3[86]; head of Cumberland Sound, 1[91]; Pangnirtung, 2[77]; + Kingnait Fiord, 1[91]; Kikkulin Island, Cumberland Sound, 1[7]; + Blacklead Island, Cumberland Gulf, 1; merely Cumberland Gulf, + 1[7]; merely east Baffin Island, 34[7]; Cape Dorset, 2[2]; SW + coast of Baffin Island, 1[75]. + + =Melville Peninsula.= Iglulik, 3; Pingerqalik, 2; Kingadjuaq, + Amitsog, 3; Rae Isthmus, 3; Lyons Inlet, 13(9[2]); M[N?] + alugsitaq, Lyon Inlet, 5; Itibdjeriang, 2; Repulse Bay, 27 (22[2], + 2[19]); Drichetts Cove, Hurd Channel, 1[2]; Gore Bay, 1; Haviland + Bay, 1; Cleveland Harbor, Frozen Strait, 1. + + =Southampton Island and adjacent islands.= Danish Island, 11; + Vansittart Island, 4. Southampton Island: Coral Inlet, 19 (1[77], + 18[9]); Prairie Point, 1[9]; Munnimunnek Point, South Bay, 5[9]; + Native Point, 1[9]; Ranger Rim, 1[9]; Koodloatok (not found on + map), 1[77]; merely Southampton Island, 1[77]; Gore Bay, 1[2]; Fox + Channel, 2[2]. + + =Mainland to west of Southampton Island.= Cape Fullerton, 3 + (1[77], 2[2]); Chesterfield Inlet, 4 (1[77], 1[9]); Tavane, 1[77]; + N of Wagner Inlet, 1; Eskimo Point, 1[86]. + + +=Mustela erminea kadiacensis= (Merriam) + +Ermine + +Plates 2, 3, 4, 9, 10 and 11 + + [_Putorius arcticus_] subspecies _kadiacensis_ Merriam, N. Amer. + Fauna, 11:16, June 30, 1896. + + _Putorius kadiacensis_, Preble, Proc. Biol. Soc. Washington, + 12:169, August 10, 1898. + + _Mustela kadiacensis_, Miller, U. S. Nat. Mus. Bull., 79:97, + December 31, 1912. + + _Mustela erminea kadiacensis_, Hall, Journ. Mamm., 26:179, July 19, + 1945. + + _Type._--Male, subadult, skull and skin; no. 65290, U. S. Nat. + Mus., Biol. Surv. Coll.; Kodiak Island, Alaska; April 25, 1894; + obtained by B. J. Bretherton, original no. 304. + + The skull lacks the basioccipital, part of the basiphenoid, the + occipital region on the right side and the posterior part of the + right tympanic bulla. The third, upper, left incisor is missing. + Otherwise the teeth all are present and entire. + + The white, winter skin is only moderately well stuffed 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 averaging more than a third of + basilar length; interorbital breadth more than distance 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 length, 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 considerably +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 _Putorius arcticus_, Merriam (1896:16) wrote: "A small +form of _arcticus_ occurs on Kadiak Island. . . . It is probably worthy +of recognition as subspecies _kadiacensis_." The informality of this +description possibly was in part due to the describer's recognition of +the fact that the degree of difference between _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 pronounced distinction of +_kadiacensis_ and Merriam's male type specimen is smaller than any of +the five additional males saved from Kodiak Island since that time. +Even so the differences fully warrant 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 pronounced 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 Zoöl. 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. Zoöl., + 54:511, July, 1913. Type from Bay St. George, Newfoundland. + + _Mustela cicognanii_, Sheldon, Journ. Mamm., 13:201, August 9, + 1932. + + _Mustela cicognanii richardsonii_, Miller, U. S. Nat. Mus. Bull., + 79:95, December 31, 1912; Hall, Univ. California Publ. Zoöl., + 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 Franklin, Canada; presented to + British Museum on or before March 3, 1843; may be the type. + + 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_, _semplei_ 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. bangsi_, + 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 Islands is 69 + (64-78) grams. Weight of adults would be more. + + _Color._--Winter pelage all white except tip of tail. Summer + pelage with upper parts uniform in color and darker (16_n_) than + Raw Umber, and about tones 3 to 4 of Chocolate of Oberthür and + Dauthenay, pl. 343. Underparts Sulphur Yellow, Colonial Buff, or + Primrose Yellow, often nearly white on chin and insides of + forelegs; color of underparts extends narrowly over upper lips, + distally on posterior sides of forelegs onto antipalmar faces of + toes and sometimes 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 Athabasca 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; light-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 _bangsi_. In _invicta_ the + underparts 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 with _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 tympanic 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 tympanic + 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 lighter than that of + the male. + +Comparison of the skull with that of _arctica_, _polaris_, _semplei_, +_kadiacensis_, _haidarum_, _cicognanii_, _bangsi_, _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 +respect to the other races, is more abundantly represented by study +specimens in zoölogical 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 +accordingly is more fully described than any one of the other races +each of which by reference to _richardsonii_ is described in +comparative 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 companions, 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 previously well +represented. + +The nomenclatural history of this subspecies begins with references in +the literature that identify the animal as the Old World species, +_Mustela 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 +Boreali-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 _richardsonii_ +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 halfway 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 representative of _richardsonii_ the +far northern and hence large-sized animals (now separated as _M. e. +arctica_), his halfway 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 +approximately 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 that 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 between the +Musqwa and Prophet rivers, I for a second time examined 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 involved 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 intermediate 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 Simpson, 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_ +because _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 _richardsonii_ +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 +distinct 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. + +The name _Mustela cicognanii mortigena_ Bangs, 1913, proposed for the +ermine of Newfoundland, is placed as a synonym of _richardsonii_ 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 measurements and comparisons go, to be in nowise +different from those in Newfoundland. Females in Newfoundland may have, +on the average, 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 between 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 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 2-1/3:1; in _richardsonii_, 2-1/2:1; in +the south (_muricus_), 1-2/5:1. Indicated in still another way in +_richardsonii_ the skull of the female is 56 per cent lighter than that +of the male and the skull of the male is 127 per cent 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 _semplei_-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 intended, 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 difference between +the two subspecies just mentioned is approximately twice as much as +between _richardsonii_ and _cicognanii_. 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 differ appreciably from those taken at, +say, locality B, 50 miles farther south, although ermines from A and +those from a third locality, C, say, 130 miles south, clearly show +differences indicative of geographic variation. + + _Specimens examined._--Total number, 1035, as follows. Arranged + alphabetically by provinces and districts and from north to south + in each province or district. Unless otherwise indicated, + specimens are in the United States National Museum. + + =Alberta.= 15 mi. above Smith Landing, 2; Fort Smith, 2 (1[77]); + Smith Landing, 2; LaButte, Fitzgerald, 1[77]; Egg Lake, 15 mi. NW + Ft. Chippewyan, 4 (2[75]); 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[75]; 18 mi. below Peace Point, 1; Embarass River, 7 + (4[75]); Athabasca River, 1[2]; Ft. McMurray, 1; Athabasca River, + Middle Rapid, 2; 60 mi. above Grand Rapids, 1; Boiler Rapid, 1; + Entrance, 3[2]; St. Albert, 2. + + =British Columbia.= Fort Halket, 1; Shesley, 1[2]; Dorothy Lake, + Mts. W of Ft. Nelson, 4000 ft., 3[2]; valley between Musqwa and + Prophet rivers, 3800 ft., SW of Ft. Nelson, 2[2]; Sikanni Chief + Riv., 1; Telegraph Creek, 7 (6[2]); head of Bad River, 2350 ft., + on lake, 1; Six Mile, 5[74]; Tuchodi Lake, 2[2]; Iskoot River, + 2[14]; Level Mtn., 1[2]; head of Tatletuey Lake, 12 mi. W Thudade + Lake, 2; Robb Lake District, 5[2]; Ft. Grahame, 12 (2[77]); Sustut + Mts., on trib. Sustu Riv., 25 mi. SE Thudade Lake, 2; Laurier + Pass, 1; Omineca Mts., 1[85]; Point Creek and Clearwater River, 2; + Kispiox Valley, 23 mi. N Hazelton, 5[74]; Hazelton, 3[77]; NW arm + Tacla Lake, 7; N end Babine Lake, 1; Pt. Simpson, 1; Metlakatla, + 1; Stuart Lake, 27; S Fk. Salmon Riv., 1[77]; mouth Salmon Riv., + 1[77]; Vanderhoof, 4[77]; Wistaria P. O., near Burns Lake, 1[77]; + Kruger Lake, 9[74]; Indianpoint Lake, 23[74]; Quesnel, 1; Ahbau + Lake, 3[74]; Isaacs Lake, 6[74]; Beaver Pass, 56[74]; Lightning + Creek, 54[74]; LaFontaine, 16[74]; Barkerville, 1[74]; Barkerville + District, 34[74]; Swift River, 27[74]; Cunningham Creek, 34[74]; + Itcha Mts., 1[31]; Anahim Lake, 1[74]; Chezacut Lake, 8[31]; + Kleena Kleene, 18[74]; 158 mi. House (Cariboo on labels), 3[60]; + Rivers Inlet, 6 (5[94]; 1[77]); Horse Lake, 4[22]; Kingcome Inlet, + 8[77]; Loughborough Inlet, 7[77]; McGillivary Creek, 1; Camel + Back, Pemberton Meadows, 1[31]; Arrow Rapids, mainland opposite + Stuart Island, 1[77]; Butte Inlet, 9[77]; Green Lake, 1[31]; Mt. + Whistler, 1[86]; Alta Lake, 2 (1[31]; 1[21]); Mons, 1[31]. + + =Keewatin.= Foot of Baker Lake, 1. + + =Labrador.= Okak, 3[75]; Nain, 22 (11[75]; 11[60]); Hopedale, + 24[75]; Kippokak Bay, 7[75]; Ailik, 1; Makkovik, 26[75]; Labrador, + 55° N, 3; Hamilton Inlet, 2[75]; NW River Post, interior Labrador, + 5[1]; Cartwright, 5; Paradise, 12; Sandwich Bay (Muddy Bay, 6; + North River, 6), 12; Battle Harbor, 1[7]; St. Marys River, 3[7]; + Black Bay, 16 (15[75]; 1[76]); Lanceau Loup, 17 (1[75]). + + =Mackenzie.= Ft. Franklin, 1[2]; Ft. Rae, 12; Fairchild Point, + 6[9]; Fort Simpson, 10 (2[2]); Hot Springs (61°, 125°), 1[2]; + 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[77]; Govt. Hay Camp, Wood Buffalo Park, 2[77]. + + =Manitoba.= Egg Is., Rabbit Point, 1; Ft. Churchill, 1; Ft. York, + W Hudsons Bay 57° N, 1[7]; Oxford House, 11; Gypsumville, 1[86]; + Lake St. Martin. + + =New Brunswick.= _Restigouche County_: Bird Bait, north Camp, 6 + mi. NE Nictau Lake, 2[59]; Red Brook, Tobique River, 1[59]. + _Victoria County_: Trousers Lake, 3[2]. _Glouchester County_: + Youghall, 1[77]; Miramichi Road, 15 mi. from Bathurst, 13[77]. + _York County_: Scotch Lake, 2. + + =Newfoundland.= Nicholsville, 3[75]; Bay St. George, 48 (26[75]; + 2[7]; 1[9]); Codroy, 9 (7[75]; 2[60]). + + =Nova Scotia.= _Victoria County_: Cape North, 2[77]. _Inverness + County_: Fizzleton, 3[77]. _Richmond County_: St. Peters, 1[77]. + _Pictou County_: Glengary, 1[4]. _Guysborough County_: East Roman + Valley, 5[77]. _Kings County_: Wolfville, 5 (3[74], 2[77]); near + Wolfville, 1[77]. _Halifax County_: Hammond Plains, 1. _Annapolis + County_: Annapolis Royal, 1. _Digby County_: Digby, 3. No locality + more definite than Nova Scotia, 3. + + =Ontario.= Severn River, 1[77]; R. C. Mission, Yellow Creek, near + mouth of Albany, 2[86]; Ft. Albany, 4; Charlton Island, 1; Moose + Factory, 10 (7[9]; 3[77]); Abitibi, 1[4]. + + =Quebec.= Fort Chimo, 10[77]; Ungava Forks, 1; Belcher Islands, + Hudsons Bay (Tukarak Island, 29; Eskimo Harbor, 2; Innetalling + Island, 1; S tip Gibson Peninsula, 2; Flaherty Island, 1), 35[9]; + Cairn Island, Richmond Gulf, 2[9]; Manitounuk Sound, 4[9]; about + 15 mi. S Great Whale River, 1[9]; Ft. George, 1[9]; Charlton + Island, 1[9]; Waswonaby Post, 1[77]; Mistassinnay Post, 3[77]; + Godbout, 36; Mt. Albert, 7 (4[78]; 3[2]); St. Anne River, 1500 + ft., 1[77]; Ste. Anne des Monts, 3[2]; "Federal Mine," 1[77]; + Berry Mountain Camp, 1[77]; Berry Mountain Brook, 1[2]; Cascapedia + River (Middle Camp, 2; Tracadie, 2; Square Forks, 1), 5[2]. + + =Saskatchewan.= Poplar Point, Athabasca Lake, 1[75]; Fair Point, + Athabasca Lake, 1[75]; Emma Lake, 1[74]; Harper Lake, 2[77]; + Livelong, 3[55]; Fairholme, 2[74]; Touchwood Hills, 2[7]; Indian + Head, 1[86]. + + =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; Nisutlin River, 1; + Nisutlin Flats, 2; Wolf River, 1; Wolf Lake, 5), 21[77]. + + +=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. + + _Putorius vulgaris_, Emmons, Quadrupeds of Massachusetts, p. 44, + 1840. + + _Mustela pusilla_ DeKay, Zool. of New York, Pt. 1, Mammalia, p. 34, + pl. 14, fig. 1, 1842. Type from New York State. + + _Putorius pusillus_, Audubon and Bachman, Vivip. Quadrupeds of N. + Amer., 2:100, pl. 64, 1851 (pl. 1846) and erroneously labeled + _Mustela fusea_, as pointed out on p. 102 of text. + + _Putorius cicognanii_, Baird, Mamm. N. Amer., p. 161, 1858. + + _Putorius richardsoni cicognani_, Bangs, Proc. Biol. Soc. + Washington, 10; 18, figs. 4, 4a of pls. 1 and 2, and pl. 3, figs. + 2, 2a, February 25, 1896 (part). + + _Putorius cicognani_, Merriam, N. Amer. Fauna, 11:10, pl. 2, figs. + 3, 3a, 4, 4a and pl. 5, figs. 2, 2a, June 30, 1896. + + _Mustela cicognanii cicognanii_, Miller, U. S. Nat. Mus. Bull., + 79:95, December 31, 1912; Bishop, Journ. Mamm., 4:26, February 9, + 1923. + + _Mustela cicognanii_, Jackson, Journ. Mamm., 3:15, February 8, + 1922. + + _Mustela erminea cicognanii_, Hall, Journ. Mamm., 26:77, February + 27, 1945; Hall, Journ. Mamm., 26:180, July 19, 1945. + + _Type._--No type specimen designated; type locality, eastern + United States. + + The restriction of the type locality from the general region of + northeastern 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 by Bonaparte's remarks in + connection with the proposal of the name _cicognanii_. 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 constantly 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 different . . . as the + Roman and the United States. . . ." Clearly he had in mind + principally, if not exclusively, the animal of the 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._--Differs 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 lighter skull (0.92 versus 1.1 grams); + from _M. e. bangsi_, in males hind foot less instead of more than + 40, linear 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 underparts 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). + +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 slightly longer. In skulls of females of +_cicognanii_, linear measurements average 3 (0-6) per cent less, the +weight is 16 per cent less and the 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 measurements +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 Natural +History, C. L. Bonaparte proposed for three kinds of American weasels +the names _Mustela cicognanii_, _Mustela richardsonii_ and _Mustela +longicauda_. + +In this paper Bonaparte indicates that he previously had written (for +his Iconografia della Fauna Italica ...) an account of _Mustela +cicognanii_ using this same name. Fasciola XXII of the Iconogr. d. +Fauna Italica, presenting his account of _Mustela_, like the English +paper was published in the year 1838. In his article in Charlesworth'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 the 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) mentions that "all +the [American?] naturalists" used the name _M. vulgaris_ 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 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, +there appears: + + 43.3.3.3 Mustela longicauda _Bonap_ N amer presented + by Dr. J. Richardson + 4 Mustela 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, "killed 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 size of by giving measurements of +a specimen taken "in the neighborhood 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. vulgaris_ 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 conceivable 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 involved, 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 +application. The same is true of _Mustela richardsonii_. If the basis +of _Mustela cicognanii_ Bonaparte [the diagnosis in the Iconografia d. +Fauna Italica ... 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 Richardson's account +of _M. vulgaris_, the name will apply to a different 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 statement in Charlesworth's Magazine (1838:37) +that "I have _now_ [Italics 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 Charlesworth'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. +Conceivably 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 immediately become as +important as the one in London. Any American weasel or weasels (then +alive or preserved in a zoölogical collection) 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 provinces 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 County_: S. Woodstock, Woodstock Lake, + 1[2]. _Hartford County_: Windsor, 1[5]. _New London County_: + Liberty Hill, 3[75]. + + =Maine.= _Aroostook County_: Quimby, 1[75]; Ashland 2[75]. + _Piscataquis County_: tableland on top of Mt. Katahdin, 1; Chimney + Pond, 3; T. 5, R. 13, 3[5]; "vicinity of Chesnucook," 1[5]; T. 4, + R. 13, 1[5]; Moosehead Lake, 7[75]; Grenville, 10[75]; Barnard, 3 + (1[86]). _Penobscot County_: South Twin Lake, 1[2]; Lincoln, 11 + (7[1], 2[14], 2[50]). _Franklin County_: Seven Pond Township, + 7[75]. _Oxford County_: Umbago Lake, 1[75]; Upton, 4[86]; Bethel, + 1[75]. _Hancock County_: Bucksport, 17[75]; Naskeag, 1. _Lincoln + County_: Booth Bay, 1[5]. + + =Massachusetts.= _Middlesex County_: Wilmington, 2; Burlington, 6 + (1[75]); _Worcester County_: Cambridge, 5 (1[5], 3[75]); Sterling, + 1[5]. _Plymouth County_: Middleboro, 7 (1[75]). + + =New Hampshire.= _Carroll County_: Ossipee, 5. _Rockingham + County_: Greenland, 1[76]. _Cheshire County_: Dublin, 1. + + =New York.= _St. Lawrence County_: Ogdensburg, 1[74]. _Franklin + County_: Malone, 1[58]. _Lewis County_: Locust Grove, 1. _Warren + County_: Lake George, 1. _Montgomery County_: Amsterdam, 1. + _Albany County_: Albany, 1[80]. _Rensselaer County_: Berlin, 2[2]; + Schoharie, 1[2]. _Thompkins County_: Cascadilla Creek, Ithaca, + 1[58]. _Allegany County_: Ford Brook, Wellsville, 1[58]. _Ontario + County_: Phelps, 1[50]. _Cattaraugus County_: Cattaraugus, 1[5]. + + =Ontario= (localities locally north to south, then west to east). + _Thunder Bay Dist._: Grand Bay, Lake Nipigon, 5[86]; Macdiarmid, + 2[86]; Oscar, 2[14]; 20 mi. SW Fort Williams, 1[76]; Michipicoten + Island, 3[104]. _Algoma Dist._: Michipicoten, 1; Franz, 1[74]; + Pancake Bay, 2[77]. _Parry Sound Dist._: French River, Georgia + Bay, 1[2]; Seguin Falls, Twp. Montieth, 1[86]. _Sudbury Dist._: + Casselman, Rathbun Twp., 1[86]. _Nipissing Dist._: Smoky Falls, + near Kapuskasing, 4[86]; Franks Bay, Lake Nipissing, 1[86]. + _Haliburton County_: Gooderham, 1[60]. _Simcoe County_: Orillia, + 1[2]; no locality more definite than county, 1[60]. _Carleton + County_: Britannia, 5 mi. W Ottawa, 1[77]; Ottawa, 1[77]; Constant + Bay, NE? of Ottawa, 1[77]. _Wellington County_: Mt. Forest, 2[75]; + Guelph, 1[31]. _Addington County_: Buckshot Lake, Abinger Twp., + 1[86]. _Fontenac County_: Clear Lake, Arden, 1[77]. + + =Pennsylvania= (by counties from west to east). _Crawford County_: + North Shenango Township, Pymatuning Swamp, 2[9]; Linesville (3 mi. + NW, 1; 3-1/2 mi. W, 2; 3 mi. W, 1; 2 mi. SW, 1; 7-1/2 mi. SW, 1) + 6[9]. _Potter County_: Cherry Springs Farm, Abbott Township, 1; 3 + mi. S Inez, South Fork Sinnamahoning Creek, 1[9]. _Sullivan + County_: Lopez, 1[74]. _Lackawanna County_: Scranton, 1[1]. _Wayne + County_: Waymart, 1. + + =Quebec= (west to east). _Labelle County_: Kamika [= Kiamika] + Lake, 2[77]; Lacoste, 2[77]; Trout Lake, probably in this county, + 2[77]. _Megantic County_: Black Lake, 1[77]. + + =Rhode Island.= _Newport County_: Middletown, 2[5]. + + =Vermont.= _Lamoille County_: Mt. Mansfield, 1. _Windsor County_: + Barnard, 1[5]. + + +=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_, Mearns, 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, Mamm. Illinois and Wisconsin, p. 375, + 1912. + + _Mustela cicognanii_, Aldous and Manweiler, Journ. Mamm., 23:250, + August 13, 1942. + + _Mustela cicognanii cicognanii_, Bailey, N. Amer. Fauna, 49:169, + January 8, 1927; Leraas, Journ. 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 Minnesota, 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 width of color of + underparts averages about a third, instead 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 that 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 adults 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 width 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. + + _Skull._--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 +indistinguishable, 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. + +_Remarks._--Before the subspecific name _bangsi_ was proposed, +individuals of this subspecies ordinarily were recorded in the +literature as _Mustela cicognanii_. The best single lot of material is +in the zoölogical 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 braincase 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 +_richardsonii_, 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 +interpretation 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 +referred to bangsi are not geographically intermediate between +_richardsonii_ 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 to the +alleged occurrence of this ermine in northwestern Ohio at New Bremen. +Henninger (1921:239) published the original account 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 misled 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 Mountains 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 provinces 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 Zoölogy. + + =Iowa.= _Dickinson County_: W side Lake Okobogie, 1[48]. + _Winnebago County_: Lake Mills, 7[65]. _Worth County_: Northwood, + 1[65]. _Clay County_: "Dewey's Pasture, near Ruthven," 1[76]. + + =Manitoba.= Aweme, 4[47]; Red River Settlement, 1[91]. + + =Michigan.= _Isle Royal_: Tobin Harbor, 1[76]; Bell Isle, 1[76]; + Washington Harbor, 3[76]. _Ontonagon County_: Ontonagon, 2 (1[76], + 1[14]); T. 51N, R. 43W, S. 17, Porcupine Mts., 1[76]. _Gogebic + County_: Little Girls Point, 5[76]; Ironwood, 1[76]. _Iron + County_: no locality more definite than county, 1[76]. _Luce + County_: Tahquamenon River Falls, 1[91]. _Chippewa County_: Sault + Ste. Marie, 2[76]. _Emmet County_: Wilderness State Park, 2[76]. + _Cheboygan County_: Univ. Mich. Biol. Station, 1[76]. _Washtenaw + County_: Ann Arbor, 1[76]. + + =Minnesota.= _Kittson County_: no locality more definite than + county, 1[2]. _Roseau County_: Deer Township, 1[14]; Falun + Township, 2[14]. _Marshall County_?: Moose River, 5[93]; Warren, + definitely in Marshall County, 1[93]. _Cook County_: Grand Marais, + 3 (2[76], 1[14]). _St. Louis County_: 2 mi E Babbitt, 14[93]; + Burntside [= Burnside] Lake, 1[91]. _Itasca County_: T. 61N, R. + 26W, 23. _Clay County_: Moorhead, 3[9]. _Aitkin County_: Aitkin, + 13 (11[60], 1[7], 1[4]). _Otter Tail County_: Arthur, 3[60]; Ten + Mile Lake, 1[76]; Parkers Prairie, 2[75]. _Chisago County_: North + Branch, 1[60]. _Sherburne County_: Elk River, 42 (16[91], 5[14], + 20[59], 1[74]). _Hennepin County_: Lake Minnetonka, 1[75]; + Minneapolis, 1[91]; Fort Snelling, 5 (4[2], 1[60]). + + =North Dakota.= _Pembina County_: Walhalla, 1[91]. _Nelson + County_: Stump Lake, 1[91]. _Eddy County_: Brantford, 2[76]. + + =Wisconsin.= _Douglas County_: T. 44N, R. 13W, 1; Gordon, 1. + _Bayfield County_: Brinks Camp, Washburn, 1[2]; "near Cable," 1. + _Ashland County_: Bear Lake, 2. _Iron County_: Fisher Lake, 4; + Mercer, 5. _Vilas County_: Mamie Lake, 16[91]; Ox Bow Lake, 1[91]. + _Oneida County_: Tomahawk Lake, 1[60]. _Langlade County_: T. 34N, + R. 11E, 3. _Rush County_: Ladysmith, 1. _Dunn County_: Colfax, 2. + _Door County_: Mink River, Ellison Bay, 1[76]. _Dodge County_: Fox + Lake, 1[50]; Beaver Dam, 12[50]. + + +=Mustela erminea invicta= Hall + +Ermine + +Plates 2, 3, 4, 9, 10, 11 and 41 + + _Mustela erminea invicta_ Hall, Journ. Mamm., 26:75, February 27, + 1945; Hall, Journ. Mamm., 26:180, July 19, 1945. + + _Putorius cicognanii_, Preble, N. Amer. Fauna, 27:230, October 26, + 1908. + + _Type._--Male, subadult, skull and skin; no. 101122, Mus. Vert. + Zoöl.; Benewah, 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 Alberta, 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 lighter 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 per cent 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 underparts 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 specimens 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_ except + 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 + measurements, which see): See plates 9-11. As described in + _Mustela erminea richardsonii_ 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. Females +show the same differences but the degree of difference is about as +great again as in males. The teeth 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 M1 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 M1 is actually shorter. Females of _invicta_ are 12 per +cent heavier; relative to the 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 anteroposterior +diameter of M1 which is the same; 50 per cent heavier; relative to the +basilar length the length of the tooth-rows and interorbital breadth +are less. In females the inner lobe of M1 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 anteroposterior 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 than 97 per +cent of the length of the tooth-rows. + +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 indicate 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 overall size is about the +same in the two subspecies concerned, 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_. +Intergradation 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 intergradation in +color occurs is fairly wide, the area in which intergradation in +cranial characters in females occurs, appears, from the inadequate +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. + + _Specimens examined._--Total number, 177, as follows. Arranged + alphabetically 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[77]; Shovel Pass, 2[77]; Jasper Park, + 10[77]; head of Smoky River, 9; Henry House, 2 (1[77]); Blindman + River, 1[2]; forks of Blindman River and Red Deer River, 2 (1[60], + 1[75]); "near Red Deer, Red Deer River," 1[77]; Red Deer River, 2 + (1[2], 1[60]); Red Deer, 2[60]; Rosebud, 2[77]; Prairie, 3000 ft., + 1; Didsbury, Little Red Deer River, 1; Canadian Nat'l Park, 1[60]; + Canmore, 1; Banff, 1[60]; High River, 1[86]; "Waterton Lake Park" + in Alberta, 6[77]. + + =British Columbia.= Grand Forks of Fraser River, 1; Canoe River, + 1[77]; Field, 1; Glacier, 1[58]; E side Beaverfoot Range, + 4000-4500 ft., 6 mi. SE Fraser Creek, 8[74]; Wentworth Lake, + 1[31]; Revelstoke, 2 (1[77], 1[60]); Spillimacheen[e]en River, + 2[2]; Sicamous, 2; Albert River, 7000 ft., 1[2]; Lumby, Creighton + Valley, 1[31]; Okanagan, 4 (2[75], 1[94], 1[2]); Kettle River + Lake, Gold Range, 4000 ft., 1; Crows Nest Station, 1[74]; Yale + District, 3; Fort Hope, 1; Chilliwack Lake, 1[77]; Skagit, 2 + (1[77], 1[31]); Skagit Valley, 1[77]; Skagit Summit, 1[77]; + Lightning Lakes, 2 mi. N International Boundary, 3; + Osoyoos-Bridesville Summit, 2; Westbridge, 1[77]; Rossland, 5[77]; + Creston, mouth Goat Creek, 3[77]; Yahk, 4[77]. + + =Idaho.= _Bonner County_: Coolin, 4. _Benewah County_: Benewah, + 1[55]. _Idaho County_: "Pete Kings, Lochsa River," 1[97]; 2 mi. + SSE Selway Falls, 1900 ft., 1[8]; 4 mi. SW Selway Falls, 5800 ft., + 3[8]; Newsome Cr., 12 mi. above jct. with S Fk. Clearwater River, + 2[74]; Iron Mt., to 14 mi. W thereof, 24[74]; Pilot Cr., 3/4 to + 2-1/2 mi. above Newsome Cr., 4[74]; Sawmill Cr., 1-1/4 mi. W + Newsome, 1[74]; between Selway River and S. Fk. Clearwater R., + 4[74]. + + =Montana.= _Teton County_ (of old arrangement of counties): Many + Glacier, 4900 ft., 1[74]; Duck Lake, 6 mi. NE St. Marys Lake, 1; + St. Marys, Glacier Park, 1[76]; Lower St. Marys Lake, 1[2]. + _Flathead County_: Stanton Lake, 5. _County_ in question: Bitter + Root Mts., 1. _Ravalli County_: Tin Cup District, 1[74]; Bass + Creek, 6800 ft., NW of Stevensville, 1; Capitan Peak, 7000 ft., 1; + Darby, 2[74]; Girds Creek, 1[74]; Charlos Heights, 2[74]. + + =Washington.= _Whatcom County_: Twin Lakes, Winchester Mts., 3 + (1[10]); Chilliwack River, 2600 ft., 2; Cooper Creek, near head, + 4500 ft., Hannegan Pass, 1; Cooper Cr., 4300 ft., Hannegan Pass, + 1[10]; Beaver Creek (2500 ft., and at McMillan Ranch, 1700 ft.), + 2; Barron, Bornite Mine, 5000 ft., 1. _Okanogan County_: Tungsten + Mine, 6800 ft., Bauerman Ridge, 4; Hidden Lakes, 4100 ft., 1; West + Fork Pasayten River, 4700 ft., 1. _Stevens County_: Orin, 1[51]. + _Pend Oreille County_: Ione, 2[51]. _Chelan County_: Lake Chelan, + 1[46]. + + +=Mustela erminea alascensis= (Merriam) + +Ermine + +Plates 2, 3, 4, 9, 10 and 11 + + _Putorius richardsoni alascensis_ Merriam, N. Amer. Fauna, 11:12, + pl. 2, figs. 2, 2a, June 30, 1896. + + _Putorius cicognanii alascensis_, Miller, U. S. Nat. Mus. Bull., + 79:96, December 31, 1912; Swarth, Univ. California Publ. Zoöl., + 7:140, January 12, 1911. + + _Mustela erminea alascensis_, Hall, Proc. Biol. Soc. Washington, + 57:36, June 28, 1944; Hall, Journ. Mamm., 26:180, July 19, 1945. + + _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, instead of more, than + distance between glenoid fossa and posterior border of external + auditory meatus; from _M. e. salva_, in males, by overall depth of + braincase including tympanic bullae less than 89 per cent of + orbitonasal length, females 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 average 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 erminea 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 richardsonii_ + 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 localities listed in the table of + cranial measurements): See measurements and plates 9-11. As + described in _Mustela 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 +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. M1 is larger but m1 and P4 are +smaller. In females the anteroposterior extent of the inner moiety of +M1 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 skull with those of _arctica_, _salva_, _initis_, +_celenda_, 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 +_richardsonii_. Although actual intergrades are lacking between +_alascensis_ 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 characters. 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 intermediate 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_. + +[Illustration: FIG. 26. Map showing known occurrences and probable +geographic ranges of the subspecies of _Mustela erminea_ in +southeastern Alaska.] + + _Specimens examined._--Total number, 24, arranged by localities + from north to south. Unless otherwise indicated, specimens are in + the Museum of Vertebrate Zoölogy, University of California. + + =Alaska.= Juneau, 5[91]; Taku River, 1; Windham, 9; Mitkof Island, + 1; St. John Harbor, Zarembo Island, 1; Wrangel, 1[91]; Helm Bay, + Cleveland Peninsula, 1; Cleveland Peninsula, 2[91]; Revillagigedo + Island, Carroll Inlet, 2[91]; Loring, 1[91]. + + +=Mustela erminea salva= Hall + +Ermine + +Plates 2, 3, 4, 9, 10 and 11 + + _Mustela erminea salva_ Hall, Proc. Biol. Soc. Washington, 57:35, + June 28, 1944; Hall, Journ. Mamm., 26:180, July 19, 1945. + + _Type._--Male, adult, skull only; no. 74641, Mus. Vert. Zoöl.; + Mole Harbor, Admiralty Island, Alaska, December 27, 1936; obtained + by A. Hasselborg. + + The skull (plates 2-4) shows 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 rostrum 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 approximately 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; +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. Comparisons 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--_salva_ 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 + Zoölogy, 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, Journ. Mamm., 26:180, July 19, 1945. + + _Type._--Male, adult, skull and skin; no. 289, Mus. Vert. Zoöl.; + 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 the 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. + + _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 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 millimeters as against +39.6 and 40.5 in the two adult males. The difference 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 protruded 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 island and the three smaller specimens being +descendants of an individual ermine, or of ermines, that were rafted or +otherwise transported 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 ascendancy. Therefore, +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 specimens, +carefully labeled as to sex. Yet another reason why additional +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 features. This +specimen from Chichagof Island is labeled as a male also. + + _Specimens examined._--Total number, 6, arranged by localities + from north to south, and in the Museum of Vertebrate Zoölogy, + 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, Journ. 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 I1, and the + left I1 and I2 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 third 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 like upper parts rather 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 Oberthür + and Dauthenay, pl. 342; lower throat and chest covered by a large + patch of same color as upper parts; color of underparts 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 + winter 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 of + patch of dark color on lower throat and chest, narrower + light-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 like 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 between last upper molar and jugular foramen. + + Female.--Complete skull of adult unavailable. + +Differences from _richardsonii_ are indicated in the formal description +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 +M1 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 dimensions 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 _kadiacensis_, differences are: 26 per cent lighter, +skull shorter; in relation to the basilar length, braincase shallower +as measured at the anterior end of the basioccipital, tooth-rows +shorter but orbitonasal 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 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 zoölogist could +with reason accord full specific rank to _celenda_. Nevertheless 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 specimen 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 + localities from north to south. Unless otherwise indicated, in U. + S. National Museum. + + =Alaska.= Prince of Wales Island: Craig, 18 (10 in Mus. Vert. + Zoöl., and 8 in Los Angeles Mus. Hist. Art and Sci.); Kasaan Bay, + 2; no locality more definite 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, Journ. Mamm., 26:181, July 19, 1945. + + _Type._--Male, adult, skull alone; no. 31232, Mus. Vert. Zoöl.; + Port Santa Cruz, Suemez Island, Alaska; March 24, 1920; obtained + by George Willett. + + The skull (plates 5-7) is complete and unbroken. Of the upper + incisors only right I3 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).--Differs + 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 than distance between glenoid fossa and + posterior margin of external auditory meatus; zygomatic breadth + more than distance between last upper molar and jugular foramen. + + Female.--Skull not available. + +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 subspecies. +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 haidarum_ 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, Journ. 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._--Differs from _M. e. celenda_ + in chest white (not mostly covered by brown patch), proximal + two-thirds of under side of tail colored like underparts instead + of upper parts, in males basilar length less than 38.2; from _M. + e. seclusa_, in male, in zygomatic breadth 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 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 underparts 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 richardsonii_ 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 +female, which is 23 per cent heavier, the width of the tympanic bulla +and anteroposterior extent of the inner lobe of M1 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 M1 which are about the same in all +three races concerned. The pre-and interorbital 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 immediately to the north and south. In combination, the features +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 otherwise indicated, + specimens are in the U. S. National Museum. + + =British Columbia.= Queen Charlotte Islands. Masset, 7 (4[74], + 1[2], 1[59]); Skidegate, 1; Graham Island, 5 (2[94], 1[77], 1[2]); + Cumsheva Inlet, 3; no locality more definite than Queen Charlotte + Islands, 1[2]. + + +=Mustela erminea anguinae= Hall + +Ermine + +Plates 5, 6, 7, 11, 12 and 13 + + _Mustela cicognanii anguinae_ Hall, Univ. California Publ. Zoöl., + 38:417, November 8, 1932. + + _Putorius cicognanii_, Baird, Mamm. N. Amer., p. 161, 1858 (part). + + _Putorius streatori_, Swarth, Univ. California Publ. Zoöl., 10:102, + February 13, 1912. + + _Mustela erminea anguinae_ Hall, Journ. Mamm., 26:79, February 27, + 1945; Hall, Journ. Mamm., 26:181, July 19, 1945. + + _Type._--Male, adult, complete skeleton (no skin); no. 12482, Mus. + Vert. Zoöl., French Creek, Vancouver Island, British Columbia; + found as a desiccated 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._--Differs from _M. e. + haidarum_, in both sexes, in light-colored underparts less than + half the width of dark-colored 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. fallenda_, in both sexes, anterior margin of tympanic + bullae flush with squamosal 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 Oberthür 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 _fallenda_ 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 squamosal 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-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 braincase. In +relation to the skull as a whole the preorbital and interorbital 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 midline 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. M1, 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. + +Differences from _olympica_, in males, are: M1 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, braincase 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 relation 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 zoölogists. + +_M. c. anguinae_ is noteworthy for the slight secondary sexual +variation in size; the disparity between the two sexes is less than in +any other American subspecies of _erminea_. By linear measurement 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 approximation 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 skull 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-lips were brown. + + _Specimens examined._--Total number, 40, listed by localities from + north to south as follows. Unless otherwise indicated, specimens + are in the National Museum of Canada. + + =British Columbia.= Vancouver Island: Cape Scott, 4; Shushartie, + 1; Quatsino, 1[74]; Jeune Landing, 1[74]; Port Alice, 5[15]; + Marble Creek, Quatsino Sound, 1[22]; Port Hardy, 5; Sayward, 2; + Bear Lake, 4; Bear River, 1; Comox, 4(3[85]); Stamp River, + Alberni, 1[31]; Errington, 1[74]; French Creek, 1[74]; Hilliers, + 1[74]; Craigs Crossing, 1[74]; Nanaimo, 2[22]; Cowichan Lake, + 1[22]; Duncan, 2[85]; Salt Spring Island, 1[85]. + + +=Mustela erminea fallenda= Hall + +Ermine + +Plates 5, 6, 7, 11, 12 and 13 + + _Mustela erminea fallenda_ Hall, Journ. Mamm., 26:79, February 27, + 1945; Hall, Journ. Mamm., 26:181, July 19, 1945. + + _Putorius streatori_ Merriam, N. Amer. Fauna, 11:13, June 30, 1896 + (part-Sumas). + +[Illustration: FIG. 27. Map showing known occurrences and probable +geographic ranges of the subspecies of _Mustela erminea_ in Washington +and parts of British Columbia and Oregon.] + + _Type._--Male, adult, skull and skin; no. 7096, Nat. Mus. Canada; + Huntingdon, British Columbia; May 21, 1927; obtained by C. H. + Young, original no. 317. + + The brown summer skin 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 + opposite 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._--Differs from _M. e. + haidarum_, in both sexes, in light-colored underparts less than + half the width of dark-colored 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 lips 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 difference 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 1-1/4 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 1-1/4 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 extreme 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 unknown 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 summer pelage except for slightly more + smoky tinge in winter in specimens from some localities; otherwise + as described in _Mustela erminea streatori_ except that least + width of color of underparts averaging, in seven adult topotypes, + 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_ + differs in darker color of upper parts and their extension at the + expense of the light-colored underparts 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. + + _Skull._--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 + described 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 interorbitally. +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 interorbital parts of skull larger, and mastoid breadth +more. + +Differences from males of _olympica_ are: size larger with no overlap +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 +interorbital 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 M1 is shorter +anteroposteriorly; otherwise all measurements of _fallenda_ average +larger 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 with 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 +M1 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 _haidarum_, _invicta_ and _anguinae_ 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 with _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 combination 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 +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 three 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 the +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 type 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 +intermediate 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[74]; Point Grey, 1[31]; Port Moody, 5[91]; Chilliwack, 8 (2[75], + 4[91], 1[60]); Sumas, 19 (18[75], 1[60]); Thurstons Ranch, 2; + Cultus Lake, 2; Mt. Baker Range, 5[75]; Lihumption Park, 1; + Huntingdon, 14; Tami Hy Creek, 1. + + =Washington.= _Whatcom County_: Semiahmoo, 1[91]; New Whatcom, + 1[68]; Lake Whatcom, 2[91]; 5 mi. W Glacier, 1[51]; Glacier (3 at + 900 ft.), 4[91]; E Side Easton Glacier, Mt. Baker, 1[55]; Morovitz + Guard Station, 831 ft., 1[55]. + + +=Mustela erminea olympica= Hall + +Ermine + +Plates 5, 6, 7, 12, 13 and 14 + + _Mustela erminea olympica_ Hall, Journ. Mamm., 26:81, February 27, + 1945; Hall, Journ. Mamm., 26:181, July 19, 1945. + + _Mustela rixosa_, Svihla and Svihla, Murrelet, 13:24, January, + 1932. + + _Mustela rixosa rixosa_, Svihla and Svihla, Murrelet, 14:39, May, + 1933. + + _Type._--Male, adult, skull and skin; no. 90738, U. S. Nat. Mus., + Biol. Surv. Coll.; near head of Soleduc 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, Washington, south to Olympia. See + figures 25, 27 on pages 95, 149. + + _Characters for ready recognition._--Differs 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 grams, 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 M1 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 females they are shorter as well. +Comparison with _anguinae_ and _fallenda_ has been made in the accounts +of those subspecies. + +_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 region, but +widens out posteriorly in the inguinal region and anteriorly over the +pectoral region, throat, chin and lower lips. The upper lips are brown. +The brown of the upper parts extends around in front of each foreleg, +the two brown areas not quite meeting 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 County_: Clallam Bay, 2 (1[74], 1[94]); + Elwha, 2[10]; Johnsons Ranch, 1[60]; Happy Lake, 1[60]; Boulder + Lake, 2[60]; near head of Soleduc River, 4500 ft., 1; 12 mi. S + Port Angeles, 1[10]. _Jefferson County_: Hayes Cr., 2000 ft., + Elwha River, 2; head N Fork Quinault River, 4000 ft., 1; + Duckabush, 3; N Fork Skokomish River, 1. _Mason County_: Lake + Cushman, 2[76]; 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, pl. 2, figs. + 5, 5a, 6, 6a, June 30, 1896. + + _Putorius cicognanii_, Baird, Mamm. N. Amer., p. 161, 1858 (part + unless 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. California Publ. Zoöl., + 40:101, September 26, 1933. + + _Mustela cicognanii streatori_, Hall, Murrelet, 12:22, January, + 1931; Hall, Univ. California Publ. Zoöl., 38:417, November 8, + 1932. + + _Mustela erminea streatori_, Hall, Journ. Mamm., 26:77, February + 27, 1945; Hall, Journ. Mamm., 26:181, July 19, 1945. + + _Mustela rixosa_, Beer, Journ. Mamm., 29:296, August 31, 1948. + + _Type._--Male, adult, skull and skin; no. 76646, U. S. Nat. Mus., + Biol. Surv. Coll.; Mount Vernon, Skagit Valley, Skagit County, + Washington; February 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, instead of longer than, upper tooth-rows; + weight of skull less than 1-1/4 grams, in female weight of skull + less than 0.7 grams, length of lateral side of P4 less than 4 mm.; + from _M. e. olympica_, 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. gulosa_ and _muricus_, in both sexes, + by upper lips brown (not white), light color of underparts + extending 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 lips white (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 extreme 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 + uniform and ranging from Raw Umber to slightly darker (16_n_), and + about tones 1 to 3 of Dark Chocolate of Oberthür and Dauthenay, + pl. 342; underparts white, in summer rarely with a faint buffy + suffusion in pectoral region; color of underparts extends from + chin, and often lower lips, 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 (1.0-1.2) 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 material of +it was obtained by Alex Walker in Tillamook County, Oregon, where he +resides. The almost ideal series of 30 specimens showed the range of +secondary sexual, age, and individual variation 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 individuals are of comparable size. + +Intergradation with each of the geographically adjoining subspecies, +_olympica_, _fallenda_, _invicta_, _gulosa_ and _muricus_ is shown by +specimens examined. With the last mentioned subspecies, intergradation +is shown by two specimens from as far south as Siskiyou 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 clines 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 +another, 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 topotypes 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 arrangement 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 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 topotypes than to specimens from +any other locality. + +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 arrangement 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 light 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 abdomen 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[74]. + _Mendocino County_: Russian Gulch State Park, 1[74]. _Sonoma + County_: Mouth of Gualala River, 1[74]. + + =Oregon.= _Clatsop County_: Astoria, 1. _Tillamook County_: + Tillamook, 16 (14[14], 1[59]); Blaine, 12 (7[14], 2[59], 1[93], + 2[76]). _Washington County_: Beaverton, 1[60]; Forest Grove, + 1[36]. _Clackamas County_: Oregon City, 1[46]. _Lincoln County_: + Newport, 1. _Linn County_: Sico, 1[46]. _Lane County_: Vida Fish + Hatchery, 2[101]; McKenzie Bridge, 1[101]; Mercer, 1[75]. _Klamath + County_: Deschutes River, 6 mi. E Crescent Lake, 1[101]. _Douglas + County_: Gardiner, 1[60]. _Curry County_: Port Orford, 1; Gold + Beach, 2[60]. + + =Washington.= _Skagit County_: N end Whidby Island opposite + Deception Pass, 1; Hamilton, 4; Mt. Vernon, 3. _Snohomish County_: + Oso, 550 ft., 1; Darrington, 600 ft., 1. _Pacific County_: + Wallicut River, 2 mi. E Ilwaco, 1[74]. _Wahkiakum County_: 4 mi. + E. Skamokawa, 3[74]. _Cowlitz County_: 4 mi. E mouth Kalama River, + 2[74]; 6 mi. E mouth Kalama River, 1[74]. _Skamania County_: 15 + mi. N Govt. Springs, 1300 ft., 1. + + +=Mustela erminea gulosa= Hall + +Ermine + +Plates 5, 6, 7, 12, 13 and 14 + + _Mustela erminea gulosa_ Hall, Journ. Mamm., 26:84, February 27, + 1945; Hall, Journ. 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 midventral 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-thirds of the right one. The + right m2 is represented only by an abortive stump or the broken + root, and i1 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 slight in females), in males hind foot less + than 33.5, weight of skull less than 1-1/4 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 light-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 than 0.90 grams, basilar length more than + 30.8 mm.; from _M. e. streatori_, in both sexes, by upper lips + white (not brown), light color of underparts extending 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 (pl. 342) into tone 4 of Chocolate (pl. 343) + of Oberthür and Dauthenay; underparts (always white in winter) in + summer Sulphur Yellow or more whitish; least 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., which 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 rostrum ordinarily more than 30 per cent of basilar + length. + +In comparison with _streatori_, skulls of males and females average +smaller in every cranial measurement taken. Teeth of about same size +and males 9 per cent, and females 8 per cent, lighter. In relation to +basilar length, skull of female shallower, tympanic bullae slightly +shorter and, on the average, zygomata less expanded. + +In comparison with _muricus_, males average larger in every measurement +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 shallower 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. +Nevertheless, there is a geographic area, the southern Cascades of +Washington, throughout which individual characters are combined in +essentially 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 subspecies in +contrast to the strongly 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 of the forelegs, as in the coastal +forms, in only one of the 13 specimens in summer pelage and in it on +one side only. The black tip of the tail is short as in the coastal +forms. One specimen is in transitional pelage. It has acquired +approximately half of the white winter pelage and was taken on October +12, 1897, at Keechelus 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[51]. _Kittitas + County_: Keechelus Lake, 3 (1[1]); Martin, 1[1]; Easton, 3. + _Pierce County_: 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 County_: Mt. Rainier Nat'l Park, 5 (1 each + from: Paradise Park, 5400 ft.; Reflection Lakes, 4900 ft.; + Ohanapecosh [Hot] Springs, 2000 ft.; Tahoma Creek, 1[72]; Bear + Prairie); also in Mt. Rainier Nat'l Park, Longmire, 3 (1[72], + 1[94]). _Skamania County_: Mt. St. Helens, 6000 ft., 1. _Klickitat + County_: Trout Lake, 18. + + +=Mustela erminea muricus= (Bangs) + +Ermine + +Plates 7, 8, 12, 13, 14 and 41 + + _Putorius (Arctogale) muricus_ Bangs, Proc. New England Zoöl. 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, California Mammals, p. 248, 1906. + + _Putorius cicognani_, Taylor, Univ. California Publ. Zoöl., 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, Journ. Mamm., 12:72, February 12, 1931; Whitlow and + Hall, Univ. California Publ. Zoöl., 40:246, September 30, 1933. + + _Mustela muricus_, Miller, U. S. Nat. Mus. Bull., 79:96, December + 31, 1912; Kellogg, Univ. California Publ. Zoöl., 12:358, January + 27, 1916. + + _Mustela cicognanii lepta_, Dice, Journ. Mamm., 1:12, November 28, + 1919; Hall, Mamm. Nevada, p. 184, July 1, 1946. + + _Mustela rixosa_, Seton, Journ. Mamm., 14:70, February 14, 1933. + + _Mustela cicognanii leptus_, Miller, Journ. Mamm., 14:368, November + 13, 1933; Bailey, N. Amer. Fauna, 55:293, August 29, 1936. + + _Mustela erminea murica_, Hall, Journ. Mamm., 26:84, February 27, + 1945; Hall, Journ. Mamm., 26:181, July 19, 1945. + + _Type._--Male, young, skull and skin; no. 9146, collection of E. + A. and O. Bangs in Mus. Comp. Zoöl.; Echo, 7500 ft., El Dorado + County, California; 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, 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); + typically 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 County 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 description of + color), least width of light-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), light 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, + California, measures: Total length, 227; length of tail, 55; + length of hind foot, 27. Corresponding 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, Calif., 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 9-1/2 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 + Oberthür 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 + relatively longer black tip of tail. + + _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 +measurement 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 relation 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, preorbital 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 specimens 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 comprised 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 _Putorius (Arctogale) muricus_ Bangs with +type locality at Echo, El Dorado County, California. Furthermore, +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 subspecies, say, in +northwestern Wyoming; there is therefore no evidence of progressive +decrease in size to the southward as in advance of study I supposed +existed in _muricus_. This erroneous supposition 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 southwestern +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 indicated, specimens are in the Museum of + Vertebrate Zoölogy, University of California at Berkeley. + + =California.= _Siskiyou County_: head of Rush Creek, 6400 ft., 1; + Castle Lake, 5434 ft., 1. _Tehama 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 + County_: Fallen Leaf Lake, 6500 ft., 1[33]; Echo, 1[75]. _Tuolumne + County_: Ten Lakes, 9200 ft., Yosemite Park, 1. _Mariposa County_: + Vogelsang Lake, 10350 ft., Yosemite Park, 1. _Mono County_: + Mammoth, 1[59]. + + =Colorado.= _Rio Blanco County_: Marvine, 1. _Boulder County_: + Camp Albion, 10600 ft., 1[60]; Boulder, 1[91]. _Denver County_: + Denver, 1[57]. _Park County_: Jefferson, 1[57]. _Gunnison County_: + near Placita in Gunnison County, 1[26]; Crested Butte, 9000 ft., 3 + (1[91], 2[19]). _El Paso County_: Turkey Creek, SW Colorado + Springs, 6000 ft., 1[19]. _Chaffee County_: Arbourville, 1[91]; + Hancock, 1. _Montrose County_: Coventry, 6800 ft., 1[19]. _San + Juan County_: Silverton, 1[91]; in San Juan County above + timberline, 1[87]. + + =Idaho.= _Bannock County_: West Fork of Rapid Creek, 9-1/2 mi. E + Pocatello, 1. + + =Montana.= _Meagher County_: Camas Creek, Big Belt Mts., 4 mi. S + Ft. Logan, 1[91]. _Beaverhead County_: Donovan, 1[91]. _County_ in + question: Yellowstone Park, 1[75]. + + =Nevada.= _Humboldt County_: Alder Creek, 6000 ft., Pine Forest + Mts., 1. _Ormsby County_: 1/2 mi. S Marlette Lake, 8150 ft., 1. + _Nye County_: South Twin River, Toyabe Mts., 1[91]. _White Pine + County_: Baker Creek (8500 ft., 8675 ft., 11100 ft.), 3. + + =New Mexico.= _Taos County_: Twining, 10700 ft., 1[91]. _Sandoval + County_: 9 mi. E Cuba, 9000 ft., 1. _Santa Fe County_: Saddle S of + Santa Fe Baldy, 11000 ft., Santa Fe Range, 1[1]. + + =Oregon.= _Wasco County_: Mill Creek, 20 mi. W Warmsprings, 1[91]. + _Klamath County_: Fort Klamath, 1[91]. + + =South Dakota.= _Pennington County_: 4 mi. SE Hill City, 5300 ft, + 2[76]; Pfander's Ranch, 3 mi. SSE Hill City, 5300 ft., 1[76]; + Palmer Gulch, 3 mi. SE Hill City, 5300 ft., 1[76]; Spring Creek, 2 + mi. W Oreville, 5500 ft., 1[76]. _Custer County_: 1/2 mi. E Sylvan + Lake, 6250 ft., 1[76]. + + =Washington.= _Columbia County_: Butte Creek, 1; Stayawhile + Spring, 5150 ft., 1: + + =Wyoming.= _Crook County_: 5 mi. NW Sundance, 5900 ft., 1[93]. + _Teton County_: Whetstone Creek, 2[76]; 1/4 mi. E Moran, 6700 ft., + 1[93]. _Sublette County_: 1/2 mi. NE Pinedale, 7500 ft., 1[93]. + _Albany County_: 30 mi. N and 10 mi. E Laramie, 6560 ft, 1[93]; 26 + mi. N and 4-1/2 mi. E Laramie, 6960 ft., 1[93]. _Carbon County_: 8 + mi. N and 19-1/2 mi. E Savery, 8800 ft., 2[93]. + + +=Mustela erminea? angustidens= (Brown) + +Plates 7, 12, 13 and 14 + + _Putorius cicognanii angustidens_ Brown, Mem. Amer. Mus. Nat. + Hist., 9 (pt 4):181, pl. 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 than + length of tympanic 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 tympanic 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 + American races of _erminea_ from the far north will show that + many characters are held in common--more than with more southern + subspecies of _erminea_. + +_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 percentage +of the total (condylobasal) length of the skull, amounts to: + + in _frenata_ in _erminea_ + [M] ordinarily less than 46 [M] ordinarily more than 46 + [F] less than 47 [F] 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 occurred. This one +skull, seemingly of the _frenata_ stock, is the type of _Putorius +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 otherwise from a +deposit, like that in Conard Fissure, six females of frenata and four +males of _erminea_ without a male _frenata_ or a female 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 subspecifically distinct from any +American Recent weasel. Furthermore, by this latter interpretation each +sex of this weasel, _angustidens_, is intermediate between the +_frenata_ and _erminea_ stocks in the feature of postglenoidal length +which feature, at any place 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, _angustidens_ agrees better with _erminea_ +than with frenata and this circumstance has influenced me to place +_angustidens_ as a subspecies 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_ resembles the +northernmost American subspecies of _erminea_. Females 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 subspecies 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 erminea fallenda_ and _Mustela erminea invicta_. + +The assemblage of mammals from Conard Fissure includes several 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 considered +that the distinctive cranial features of _angustidens_ may be the +result of evolutionary change in time as well as of geographic +variation resulting from horizontal placement. + + +=MUSTELA RIXOSA= (Bangs) + +Least Weasel + +(Synonymy under subspecies) + + + _Type._--_Putorius rixosus_ Bangs, Proc. Biol. Soc. Washington, + 10:21, February 25, 1896. + + _Range._--From Norway and Switzerland eastward through Siberia 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 Columbia 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 +erminea_ 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 +extreme 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 braincase is more nearly flat +and is wider above and in front of the 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 geographic 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. Zimmermann (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_ nowadays is relegated to subgeneric rank under the +generic name _Mustela_, we have for consideration the name-combination +_Mustela minuta_ (Pomel). Unfortunately for Zimmermann's conclusion, +_Mustela minuta_ Pomel is not available because it is preoccupied by +_Mustela minuta_ Gervais [= _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, depending 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 seemingly available for weasels, and in use for them today, +which might replace _rixosa_ as the name of the species, are, first, +_Mustela boccamela_ Bechstein, 1801, of Sardinia [= _Mustela nivalis +boccamela_ 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 boccamela_, 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 _minuta_ POM. zu stellen ist, soll +hier nicht untersucht werden, ebensowenig die von CABRERA vermutete +Zugehörigkeit der grossen nordafrikanischen _M. numidica_ PUCH. zur +'_iberica_-Gruppe'." The answer to this problem requires a taxonomic, +rather than a nomenclatural, decision. Whether either _M. numidica_ or +_M. boccamela_ are conspecific with the dwarf weasel I cannot at this +time ascertain for want of adequate specimens. Because these two names, +_M. boccamela_, 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, +_mé-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 recorded 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 _allegheniensis_, the +winter pelage is either brown or white and the time of the molt into +winter pelage is irregular; each of eleven individuals from +Pennsylvania, Michigan and Ohio, taken in December, January, February +and March is mostly white but retains some considerable 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, according to existing information, more +litters are produced in spring and in winter than in summer and autumn. +Many juveniles and young of _allegheniensis_ examined in study +collections clearly were born in spring but about as many seem to have +been born in midwinter 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 young escaped--marked like parent. [F] 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 rump and one on the middle of the back. Otherwise 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 internal secretion the hormones of which +promote embryonic growth and that these glands, or others controlled +by them, were in some way responsible for an abnormal progress of molt, +or for a reversal 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 earlier +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 progressively longer periods of daylight may be the +causal stimulus. The suggestion made above that _M. rixosa_ does not +live 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 line 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, Manitoba, 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 weasel at Tyonek, +Alaska, writes that: "The natives regard the capture 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 'big 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 individuals; 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 according 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 specimen +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. + + +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 Michigan 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 seconds. 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 with its sharp teeth. The sound +of the teeth piercing the bone was distinctly audible at a distance of +several feet. During 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 +seconds 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 preferred; they were not eaten when +an abundance of food was present. 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 immediately 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 snakelike 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 lifted 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 slightly 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 account, given beyond, +of a nest, Criddle (1947:69) records the Pennsylvania meadow mouse +(_Microtus pennsylvanicus drummondi_) 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 dwelling 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 including _Microtus +drummondi_, these last remains being chiefly indicated 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 inhabitants 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 abundant 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_, revealing 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 Youngstown, 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. After 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 behind, 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, the Mouse had +not been disturbed." + +There are two suggestions, but no proof that I know of, in the +literature 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." Although this +suggests _Mustela rixosa_, Abbott mentions on the next 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 _Mustela 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 descended 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 December 25, 1927, +in Washington County, Pennsylvania, "five full-sized, though +young . . . animals were caught under the same pile of corn fodder" +(Sutton, 1929:253). The first week of January seems to have been the +time of birth of a juvenile, no. 88080, University 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 Winecoff (1930:313), who records +other young having been taken in the same month as well as in February. +On March 10, a female from North Portal, Saskatchewan, gave birth 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, Minnesota, according to Swanson and +Fryklund (1935:125) who remark that: "The Pennsylvania and Minnesota +records show that least weasels may be born 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 born 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 litter 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 differs 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 increased 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, Washington County, +Pennsylvania, an animal was seen to enter a small hole in a creek bank. +After the observer dug in a distance of approximately six inches an +adult, female least weasel was seen and obtained. Back of the animal, +the hole, which turned sharply 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, Criddle's (1926:199-200) account +of the havoc wrought by least weasels among the meadow mice (_Microtus +ochrogaster minor_) has been given. In this account he mentions the +fur-lined nests of the weasels that had appropriated the homes of the +_Microtus_. Criddle'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 +continues 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. + +[Illustration: FIG. 28. Map showing occurrences and probable geographic +ranges of the subspecies of _Mustela rixosa_ in North America.] + +"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 because 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 adjoining 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, Journ. Mamm., 7:327, November 23, 1926; + Hall, Univ. California Publ. Zoöl., 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. McIlhenny. 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 and body, and in study skins reaching only + to heel instead of to point between 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 + reëntrant angle from upper lip to behind eye, rather than + delimited 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 lists + 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 pl. 343 of Oberthür and + Dauthenay; underparts white, extending over upper lip, insides of + limbs and over all four feet. Line of demarcation between + underparts and upper parts extends from upper lip 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 + underparts 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; otherwise 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. McIlhenny. + +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 inhabits +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 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 evidence 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. Nevertheless, the +short tail, and color pattern, namely reëntrant angle of white behind +the eye, is to be seen in all Alaskan specimens examined in the brown +pelage, even in no. 107591, from Tyonek 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 +discernible 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 specimens 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 + alphabetically by Territory and District and unless otherwise + indicated in the United States National Museum. + + =Alaska.= Barrow and Point Barrow, 19 (8[2] 7[74], 2[1], 1[50]); + Wainwright, 1[57]; Mts. back of Icy Cape, 1[77]; west of Beechey + Point, 1[2]; west edge of Colville River Delta, 1[2]; Koyukuk + River, 16 mi. above Beetles, 1; upper Yukon, 1; Fort Yukon, 1; + Stephens Village, 1; Wales, 1[57]; McDonald Creek, tributary of + Salcha Slough, 1; near head of Toklat River, 1; head of Kantishna + River, 1; St. Michael, 4 (2[74]); Tyonek [= Tyonek], 1; Bethel, 3; + vic. Bristol Bay, 1. + + =Yukon.= La Pierre's House, 1; Klotassin River, tributary of White + River, 1. + + +=Mustela rixosa rixosa= (Bangs) + +Least Weasel + +Plates 14 and 15 + + _Putorius rixosus_ Bangs, Proc. Biol. Soc. Washington, 10:21, pl. + 1, fig. 6, pl. 2, fig. 6, pl. 3, fig. 4, February 25, 1896; + Merriam, N. Amer. Fauna, 11:14, pl. 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. + + _Mustela rixosa_, Thomas, Proc. Zoöl. Soc. London, p. 168, March, + 1911. + + _Mustela rixosa rixosa_, Miller, U. S. Nat. Mus. Bull., 79:96, + December 31, 1912; Swenk, Journ. Mamm., 7:327, November 23, 1926. + + _Type._--Female, adult, skin and skull; no. 642 Bangs Coll. in + Mus. Comp. Zoöl.; Osler, 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._--Differs 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 lip to side of body well below eye and + ear, rather than with reëntrant 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 line of demarcation on side of head + between upper parts and underparts passes almost straight back + without the dorsally directed reëntrant area 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.)--See 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 individual of +this race from Pembina, Minnesota, as pertaining to a distinct species. +Although he used for it the specific name _pusillus_ 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 preparing 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 the light of later knowledge, show it to be of the +species _Mustela erminea_. Only a few other naturalists followed Baird +in distinguishing the least weasel as a separate species until Bangs in +1896 (p. 21) clearly differentiated it and proposed for it the name +_Putorius rixosus_, 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 _allegheniensis_ from, say, Pennsylvania, +become available with accurate external measurements taken in the flesh +and especially with complete 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 indicated +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. 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 summer pelage has spots of dark +brown color on the breast. Only about one specimen in three of _rixosa_ +from Saskatchewan is similarly 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 Calgary, +Alberta; and 1 Island S Comb Hills, Queb.) display lesions resulting +from infestation of the frontal sinuses by nematode parasites. None of +the young skulls shows such infestation. + + _Specimens examined._--Total number, 87 as follows. Arranged + alphabetically 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[77]; 5 mi. NW Camrose, 1[77]; Camrose, + 2 (1[77], 1[31]); "near Camrose," 2[77]; Forks Blindman and Red + Deer rivers, 1[60]; Innisfail, 1[86]; Veteran, 1[93]; Diddsbury [= + Didsbury], 1; Calgary, 2 (1[93], 1[2]); Shepard, 1[86]. + + =British Columbia.= Clarks Ranch, Halfway River, Peace River Dist, + 1[85]; 15 mi. E Atlin, 1[8]; Wistaria, P. O., 3 (2[77], 1[85]); + Ootsa Lake, 1[85]. + + =Labrador.= Davis Inlet, 1[60]; 30 mi. upriver and 20 mi. toward + Groswater Mts., Eagle River, 1; St. Michael Bay, 1. + + =Mackenzie.= Old Fort Reliance, 1[2]; Fort Resolution, 2; Fort + Smith, 1. + + =Manitoba.= Gypsumville, 1[86]; Lake St. Martin Reserve, 1[86]. + + =Minnesota.= _Roseau County_: Cedarbend, 2[14]; Grimstad, 1[14]; + America, 2 (1[14], 1[74]); Malung, 1[74]; Norland, 1[41]; Falun, 3 + (1[14], 1[74], 1[41]); Palmville, 1[41]; Spruce, 1[74]; Stokes, + 1[74]. No locality more definite than Marshall County, 1[14]. + _Clay County_: Moorhead, 1[36]. _Winona County_: "near" Whitman, + 1[34]; Altura, 1[98]; Crystal Springs, 1[98]. + + =Montana.= Sun River Valley, 1; Wibaux in Wibaux County, 1. + + =North Dakota.= _Walsh County_: Grafton, 15 (3[60], 1[93], 5[2], + 2[14], 1[74], 1[1], 1[76]). _McHenry County_: 4 and 4-1/2 mi. N + Upham, 2. _Wells County_: 1[36]. _Morton County_: Mandan, 1[60]. + + =Ontario.= Algoma Dist: Tatnall, near Oba, 1[86]. Moose Factory, + 1[75]. + + =Quebec.= Island S of Comb Hills, James Bay, 1[9]. _Saguenay + County_: Natashkwan, 1. + + =Saskatchewan.= Osler, 1[75]; "near Regina," 1[77]; Dollard, + 2[31]; Shaunavon (and "near" and 1 mi. NE), 9[77]; Klintowel P. O. + (about 15 mi. N of Eastend), 1[77]; Eastend and "near" Eastend, + 2[77]. + + +=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, Journ. 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 northwestern 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. + + 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 pl. 342 + of Oberthür and Dauthenay. Underparts white at least on thoracic + region; approximately 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 underparts 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 + underparts 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. listed 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. richardsonii_. 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 earliest reference to it that I can +identify in the literature. 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 without accurate external +measurements taken in the flesh. The majority 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 Pennsylvania +have the brown color of the upper parts meeting on the underparts. +Also, more of the specimens from Ohio are lighter colored and this +suggests intergradation with the subspecies _campestris_ 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 pectoral 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 suitable 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 mentioned 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 cannot 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, Virginia and +North Carolina there is one specimen each and each specimen displays +lesions. From Ohio, 17 of 23 specimens display lesions. From Michigan 3 +of 8 specimens display lesions; 2 adults and one subadult have lesions +and 5 subadults do not have lesions. + + _Specimens examined._--Total number, 102 as follows: Arranged + alphabetically by states and within each state by counties from + north to south. Unless otherwise indicated, specimens are in the + United States National Museum. + + =Indiana.= _Huntington County_: Roanoke, 1. _Wells County_: + Harrison Township, 1[76]. + + =Illinois.= _Lake County_: Deerfield, 3[60]; no locality more + definite than county, 1[60] _Cook County_: Northfield, 1[60]; La + Grange, 1[18]. + + =Michigan.= _Tuscola County_: 8 mi. N Caro, 1[76]. _Santilac + County_: Deckerville, 1[76]. _Allegan County_: Swan Creek Exp. + Station, 1[76]; Swan Creek Farm, 1[76]; T. 2N, R. 14W, 1[76]; + Allegan, 1[76]. _Livingston County_: George Reserve, 1[76]; 1/2 + mi. N Unadilla, 1. _Oakland County_: Rochester, 1[76]. _Macomb + County_: Romeo, 1[76]. _Washtenaw County_: 5 mi. SW Ann Arbor, + 1[76]. _Branch County_: vic. Coldwater, 1[76]. + + =North Carolina.= "near Marshall," 1. + + =Ohio.= Northern part of state, 1[81]. _Williams County_: Stryker, + 1[60]. _Lucas County_: Monclova, 1[60]. _Erie County_: Sandusky, + 2[76]; marsh near Sandusky, 1[76]; Berlin Heights, 1[76]; no + locality more definite than county, 1[2]. _Wood County_: 10 mi. NE + Bowling Green, 1[76]; Bowling Green, 4[76]; 3 mi. E Bowling Green + 1[76]; Plain Township, 1[2]; Portage Township, 1[60]. _Loraine + County_: Wellington, 1[81]. _Huron County_: west of Monroeville, + 1[76]. _Summit County_: Ira, 3[81]. _Portage County_: Suffield, + 1[81]. _Hancock County_: Vanburen, 1[76]; Findlay, 1[81]; 9 mi. S + Findlay, 1[76]; no locality more definite than county, 7 (2[76], + 2[81], 3[2]). _Mahoning County_: Ellsworth, 1. _Crawford County_: + "near Crestline," 1[81]. _Delaware County_: Sunbury, 1[2]; Lewis + Center, 1[81]; no locality more definite than county, 1[81]. + _Licking County_: Johnstown, 1[2]. _Fairfield County_: Baltimore, + 1[81]; Violet Township, 1[81]. _Meigs [= Gallia?] County_: Vinton, + 1[81]. + + =Pennsylvania.= _Erie County_: McKeen Twp. 1. _Crawford County_: + Springboro, 1[1]; Pymatuning Swamp, between Hartstown and + Shermansville, Sadsbury Twp., 3[9]. _Mercer County_: Shenango + Twp., 1. _Lawrence County_: Little Beaver Twp., 1. _Butler + County_: Leasuresville, 1[9]; Clearfield Twp., 1; Valencia, 1[9]. + _Armstrong County_: Ford City, Burrell Twp., 1. _Indiana County_: + Smicksburg, 1; N. Mahoning Twp., 2; White Twp., 1. _Allegheny + County_: South Hills, Pittsburgh, 1[9]; "near Pittsburgh," 1[9]; + Fair Oaks, 1[9]. _Westmoreland County_: Bolivar, 1. _Dauphin + County_: Middle Paxton Twp., 1. _Washington County_: Finleyville, + 1; Rea, 5; Beallsville, 1[1]; Claysville, 1. _Green County_: Deep + Valley, 1; Waynesburg, 1; Jefferson, 1; Cumberland Twp., 1. + _Fayette County_: Acme, 1[9]; _Somerset County_, 1. _Lancaster + County_, 1. + + =West Virginia.= _Randolph County_: Huttonsville, 1. + + =Wisconsin.= _Sauk County_: Sumpter Twp., 1[60]. _Dodge County_: + Beaver Dam, 1[50]. _Dane County_: 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. + + _Putorius pusillus_, Aughey, Sketches of the physical geography and + geology of Nebraska, p. 119, 1880, Omaha. + + _Mustela rixosa campestris_, Swenk, Journ. 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. + + The skull is unbroken. On the left side, C1 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 averaging 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_ except 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 + reëntrant 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 examination 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. + +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 + alphabetically by states and by counties, from north to south in + each state. Unless otherwise indicated, specimens are in the + United States National Museum. + + =Iowa.= _Howard County_: Chester, 1[12]. _Palo Alto County_: + Emmetsburg, 1[65]. _Kassuth County_; Algona, 1[65]. _Clayton + County_: National, 1. _Storey County_: Nevada, 1[65]. _Wapello + County_: Ottumwa, 1[65]. _Henry County_: Mount Pleasant, 1[66]. + + =Nebraska.= _Holt County_: Page, 1[35]. _Madison County_: Norfolk + 1[35]. _Cuming County_: Beemer, 1. _Hamilton County_: Chapman, + 1[35]. _Clay County_: Inland to 1 mi. east thereof, 7[35]. + + =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, 1[102]. + + +=MUSTELA FRENATA= Lichtenstein + +Long-tailed Weasel + +(Synonymy under subspecies) + + + _Type._--_Mustela frenata_ Lichtenstein, Darstellung neuer oder + wenig bekannter Säugethiere, pl. 42 and corresponding text + unpaged. 1832. + + _Range._--From southern Canada southward over all of the United + States, México, Central America, Venezuela, and the republics of + western South America to southern Perú 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, México, 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 species); from +_Mustela rixosa_ by presence of black pencil on tail, caudal vertebrae +more than a fourth (2/5-3/4) 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 pls. 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 +percentage of the condylobasal length, less than 47 in females and +ordinarily less than 46 in males; upper parts brown; light-colored +underparts, in summer pelage, tinged with buffy or yellowish and +continuous from chin to inguinal region; some subspecies (southwestern +United States, México, Central America, and Florida) with white or +yellowish facial markings which do not occur in any other American +species of the genus _Mustela_. + +_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._--Habitat and Numbers.--As has already been remarked, +the long-tailed weasel is absent from the extreme desert of the +southwestern United States and northwestern México. Possibly the +absence of water to drink is the limiting factor. In southern 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 canyon at the head of +Dwight Way and in the autumn and winter of 1928 in Strawberry Canyon on +the campus of the 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 population 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 +the 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, 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 between the two species +is to be expected because _M. frenata_ does not range so far northward +toward the polar regions as does _M. erminea_ 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 increase; 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, Criddle and Criddle +(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 Centre County, 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. + + +Behavior + +An adult female (now the holotype of _Mustela frenata nevadensis_) seen +running across a field, and, I think, unaware of my presence, at every +bound bent her back up so far that she reminded me of a measuring worm. +For part of the time when running, the tail was held off the ground +straight out behind, and then, for a while, inclined upward at an angle +of about 45°. Another weasel that I saw in the daytime, and that I +think was unaware 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 Río +Ramos in México (Davis, 1944:381). Also, Green (1936), in May, in +Gratiot County, Michigan, saw a weasel, running with a _Peromyscus_ in +its mouth. The weasel dropped the mouse, entered the water and swam to +a hole among stones. + +More instances of climbing, than of swimming, have been reported in the +literature for the long-tailed weasel. Seton (1929 (2):625) quotes +William M. Graffius 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 +vertically, 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 appeared practically to wrap its sinuous body around the limb." + +A male long-tailed weasel, from Colorado, which I kept captive 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 thus 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 instance, 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 intended 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 Lafayette, +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 puff +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. f. noveboracensis_, in a week, averaged +10 evacuations every twenty-four hours, that urination 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 suggested by Soper's +(1919:45) account of _Mustela frenata noveboracensis_ wherein he +repeats a story told to him of a hawk observed in unsteady flight, and +obviously in distress, which when it plummeted 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, mistakenly thought to be a +chipmunk, was seen to dash into a hollow log whereupon pounding on the +log brought out the weasel which bit and clung to the 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 dislodge what was thought to +be a pack-rat. The animal was a long-tailed weasel which attacked him. +It ran up his legs a number 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 off 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, before 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 attacked him several +times. Finally the weasel went around Wight to reach the hen. In +Wight's words "There was no evidence of infuriation, but rather a well +directed offense at the one object, regardless of its size, which stood +between the weasel and an opportunity 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 digestive 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, +therefore, 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, another +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 produced young. + +A weasel in the white winter coat was used by Audubon and Bachman +(1856:177, Quarto edit.) to drive rabbits out of their burrows 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 animal'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, 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. f. +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. f. peninsulae_ from Hudsons, Florida, according to +Rhoads (1894:155) ". . . was caught in the woods by a cat." Barber and +Cockerell (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 +_Citellus 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 embedded in its +neck, head, and shoulders." The remainder of Osgood's account implies +that the weasel may have turned to porcupine because the normal food +for weasels was scarce at the time. Porcupine quills, then, are a +hazard for weasels although it is unlikely 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 suffered it, was recorded by Soper +(1921:37). The animal had a stick lodged crosswise between the fourth +upper premolar teeth. + +The recorded actions of several kinds of animals which are too small to +be dangerous to the weasel suggest that 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 by 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 sparrows and a male towhee that were noisily mobbing a +Weasel, twittering 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 curiosity of birds +to a degree which permitted the weasel to come within 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 Washington 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 [_Lepus 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 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 overcome, but weasels are very +persistent when they once get into contact 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 certain coverts with +noticeable regularity, but rarely cruised the same area on two +consecutive nights." + +The killing technique of fifteen captive _Mustela frenata +noveboracensis_ 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 light 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 induced 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. + +Polderboer, Kuhn and Hendrickson (1941) describe a cottontail 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 cottontails, 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. f. +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 +City, Utah; three from Contra Costa County, California; 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 (_loc. cit._). + +Allen (1938:225-229) experimented with the ability of four different +males of _M. f. 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 Criddle and +Stuart Criddle (1925:143, 144), in my opinion, correctly explain the +killing of more prey than weasels need. "The fact that 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." + +Criddle and Criddle (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." + +Considerable 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 City, California, 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 County, Michigan, +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 _Mustela 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 shrew (_Blarina brevicauda_).--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 attempted +capture of one of these squirrels in Saskatchewan is recorded 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 +tridecemlineatus_).--Errington (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 _Peromyscus_ and 1 +unidentified); 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 in mortal terror and +squeaking continuously. Eventually the squirrel was thrown on its back +"and would have been speedily killed but for an interruption" (Criddle +and Criddle, 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 County, California, +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 County, California, 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. + +Chipmunk (subgenus _Neotamias_).--Stanford (1931:363) on November 11, +1931, at Fish Lake, Utah, saw a weasel pursuing a chipmunk. On August +5, 1910, "near Independence Lake," Nevada County, California, 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 seconds 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 Cisco, California, 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 it to drop . . . +a pocket gopher [_Thomomys 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 _Thomomys talpoides_ in the vicinity of +Treesbank, Manitoba, Criddle and Criddle (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, "Came 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." Criddle (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 burrows 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 Chalchicomula, 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 Cockerell (1898:189) found +remains of this mouse in the stomach of a weasel at Mesilla Park, New +Mexico. + +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 +[_Neotoma 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. +Contents 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 + Mearns 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 + +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. Items 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 cottontails 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 that the abundance of several prey species does not cause the +weasels to ignore the shrews 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 percentages 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. + +Criddle and Criddle (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, California, found an adult and four young weasels which fed on +several _Microtus montanus montanus_. Green (1936) in May, in Gratiot +County, Michigan, in the vicinity of a nest in which there were four +young weasels, found "several" dead _Microtus_. 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 of Mammal Genera in Fall +and Winter Food of 163 _Mustela noveboracensis_": _Microtus_, 33.6 per +cent; _Sylvilagus_, 17.3; mammals undetermined to genus but principally +mice, 17.1; _Peromyscus_, 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 concerning 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 Criddle (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. Cooper, 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 accounted for during the afternoon, and Mr. Cooper +assures us that the weasel 'kept 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 +California, on Parker Creek, H. C. Bryant frightened a weasel that +dropped a freshly killed jumping mouse (Grinnell, Dixon and Linsdale, +1937:232). + +Snowshoe rabbit (_Lepus 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. + +"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 the +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 stratagem, +leisurely hopped about to leave its zigzag trail. Then it sat down +quietly to wait. . . . The weasel['s] . . . nose led him through the +network with little 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 that in December of 1886 in the +sandhills northeast 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) mentions a month-old varying +hare that was rescued from a weasel and of approximately the same size +as the weasel. + +Criddle and Criddle (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 further (_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 beneath 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 authors (_op. cit._:143) say that no bird remains +were found in the pile of 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 Criddle'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 the weasel's den. The cache contained only two +cottontails, one partly eaten. Leopold (1937) records seeing a +_Mustela_ (probably a long-tailed weasel but possibly 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 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 between 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 cottontail. + +Brush rabbit (_Sylvilagus bachmani_).--Vestal (1937:364) in Contra +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, incapacitated 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, Michigan, 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. f. effera_, was seen by Booth (1946:439) attempting to enter the +nesting hole of a pair of flickers, _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 responsible. + +Criddle and Criddle (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, the +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." Criddle and Criddle (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 sometime (The specimen was a large +male). . . . + +"In the fall of 1924, Mr. A. Cooper, 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 accounted for during the afternoon, +and Mr. Cooper assures us that the weasel 'kept up the good work for +same days'. + +"Being a farmer of many years' standing, Mr. Cooper has naturally lost +some poultry through the agency of weasels, but while he remarks 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 Criddle, 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, Criddle and Criddle (1925:145) write as follows: "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 sufficient 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, exceed 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 Criddle and Criddle (1925:147) who write: "Hawks are not +always the aggressors, as is shown by an incident 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 expect 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 _Mustela frenata_. + + Moles (family _Talpidae_), 5 + Shrews (family _Soricidae_), 26 + Pigmy weasel (_Mustela rixosa_), 1 + Ground squirrels (genus _Citellus_), 23 + Chipmunks (genus _Tamias_), 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 + chickens, 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 restricted +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 the seventh +week of age, they consumed approximately 22 per cent of their body +weight per day; from the eighth to the tenth week (just before reaching +mature size) they consumed approximately 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." + +Criddle and Criddle (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 occasionally 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, 1948A, and +1948B) reporting on his detailed studies of _Mustela frenata_ +(subspecies _oribasus_ 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 months, the embryos resulting from these matings +become active in early spring and develop to full term in less than 27 +days after they become 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 approximately 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) describes 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. f. noveboracensis_ +that it ". . . was pink and much wrinkled. The wetness . . . did not +entirely obscure a few sparse, rather long, white hairs . . . over its +back and head. It had the pronounced 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 1-1/2 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 "hurried 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 +_Mustela 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 three feet long. . . . The meager nest +material consisted entirely 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 distance 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 headquarters 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 dislodged 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 three 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. f. longicauda_ at Aweme, Manitoba, 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 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 living 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 large male, was in a field of sweet clover two feet high in +the former burrow of a Franklin's ground squirrel. The nest cell, seven +inches in diameter and nine inches below the surface of the ground, +"was lined with grasses mixed with much rabbit and mouse fur. Some +scats, and bones and fur of mice and shrews were matted together in +layers at the bottom of the nest." When this den was abandoned the male +weasel occupied, for a month, another burrow, 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. Observation 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 the male and female do not live +together except in the breeding season. Consequently, I wonder if the +sign was wrongly read. + +[Illustration: FIG. 29. Map showing the geographic ranges of the +subspecies of _Mustela frenata_ and _Mustela africana_. + + 1. _M. f. noveboracensis_ + 2. _M. f. occisor_ + 3. _M. f. primulina_ + 4. _M. f. arthuri_ + 5. _M. f. olivacea_ + 6. _M. f. peninsulae_ + 7. _M. f. spadix_ + 8. _M. f. longicauda_ + 9. _M. f. oribasus_ + 10. _M. f. alleni_ + 11. _M. f. arizonensis_ + 12. _M. f. nevadensis_ + 13. _M. f. effera_ + 14. _M. f. washingtoni_ + 15. _M. f. saturata_ + 16. _M. f. altifrontalis_ + 17. _M. f. oregonensis_ + 18. _M. f. munda_ + 19. _M. f. xanthogenys_ + 20. _M. f. nigriauris_ + 21. _M. f. latirostra_ + 22. _M. f. pulchra_ + 23. _M. f. inyoensis_ + 24. _M. f. neomexicana_ + 25. _M. f. texensis_ + 26. _M. f. frenata_ + 27. _M. f. leucoparia_ + 28. _M. f. perotae_ + 29. _M. f. macrophonius_ + 30. _M. f. goldmani_ + 31. _M. f. tropicalis_ + 32. _M. f. perda_ + 33. _M. f. nicaraguae_ + 34. _M. f. costaricensis_ + 35. _M. f. panamensis_ + 36. _M. f. meridana_ + 37. _M. f. affinis_ + 38. _M. f. aureoventris_ + 39. _M. f. helleri_ + 40. _M. f. macrura_ + 41. _M. f. agilis_ + 42. _M. f. boliviensis_ + 43. _M. a. africana_ + 44. _M. a. stolzmanni_ + +Hamilton (1933:328), however, writes that _M. f. noveboracensis_ 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 fuscus_ Audubon and Bachman, Journ. Acad. Nat. Sci. + Philadelphia, 8 (Pt. 2):288, 1842; Audubon and Bachman, Vivip. + quadrupeds of N. Amer., 3:234, pl. 148, 1853 (pl. 1848). + + _Putorius noveboracensis_, DeKay, Zool. of New York, Pt. 1, + Mammalia, p. 34, 1842; Baird, Mamm. N. Amer., p. 166, 1858; + Merriam, N. Amer. Fauna, 11:16, pl. 4, figs. 1, 1a, 2, 2a, pl. 5, + figs. 3, 3a, text figs. 4-6, 30, June 30, 1896; Bangs, Proc. + Biol. Soc. Washington, 10:13, pl. 1, figs. 2, 2a, pl. 2, figs. 2, + 2a, and pl. 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, pl. 59, 1851. + + _Putorius agilis_ Audubon and Bachman, Vivip. quadrupeds of N. + Amer., 3:184, pl. 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, Journ. Mamm., 4:251, + November 1, 1923. + + _Mustela cicognanii_, Henninger, Journ. 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, Journ. 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._--Williamstown, Berkshire County, Massachusetts. Type + specimen not known to be in existence. + + _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 Atlantic Coast to St. + Croix and Mississippi rivers. See figure 29 on page 221. + + _Characters for ready recognition._--Differs: 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 distance + between articular faces of exoccipital condyle and glenoid fossa; + from _M. f. occisor_ by a number of average differences including + smaller size, relatively 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 upper parts, absence of color of underparts 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. f. 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 + squamosal; from _M. f. 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. + +_Description.--Size._--Male and Female: + + ======================================================================= + | Number | | |Per cent| | + LOCALITY | of | Total | Length | of | Length of | + | specimens | length | of tail | body- | hind foot | + | averaged | | | length | | + ----------+------------+-----------+-----------+--------+-------------+ + | 8 ad. [M] | 415 | 146 | 54% | 46.0 | + Massa- | | (390-432) | (127-159) | | (41.0-48.0) | + chusetts +------------+-----------+-----------+--------+-------------+ + | 4 ad. [F] | 311 | 104 | 50% | 33.9 | + | | (298-321) | (95-114) | | (31.5-37.0) | + ----------+------------+-----------+-----------+--------+-------------+ + | 10 ad. [M] | 411 | 141 | 52% | 47.1 | + Liberty | | (379-438) | (124-155) | | (43.0-51.5) | + Hill, +------------+-----------+-----------+--------+-------------+ + Conn. | 6 ad. and | 318 | 105 | 49% | 33.0 | + | sad. [F] | (303-338) | (80-123) | | (31.7-36.0) | + ----------+------------+-----------+-----------+--------+-------------+ + | 10 ad. [M] | 407 | 130 | 47% | 46.0 | + Beaver | | (372-431) | (113-143) | | (42.0-50.0) | + Dam, +------------+-----------+-----------+--------+-------------+ + Wisc. | 4 ad. [F] | 326 | 99 | 43% | 35.6 | + | | (303-338) | (86-108) | | (34.6-38.0) | + ----------+------------+-----------+-----------+--------+-------------+ + | 10 ad. [M] | 371 | 130 | 54% | 45.0 | + Washtenaw | | (350-405) | (115-140) | | (40.0-50.0) | + Co., +------------+-----------+-----------+--------+-------------+ + Mich. | 10 ad. [F] | 306 | 97 | 46% | 34.0 | + | | (290-335) | (90-120) | | (30.0-40.0) | + ----------+------------+-----------+-----------+--------+-------------+ + + The length of the 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 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 differences 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: [M] 258; [F] 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._--Upper parts, in summer, Vandyke Brown or darker than + tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304. + Sometimes approaching tone 2 of Warm Sepia of Oberthür and + Dauthenay, pl. 305. Underparts, in summer, ranging from white + through Napthalene Yellow (Peterboro, N. Y.), Pale Orange Yellow + (eastern Mass.), near Primuline Yellow (unusual specimen 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 Oberthür and Dauthenay, pl. 301, with underparts white or + sometimes tinged with yellowish. 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 to foot and sometimes over upper sides + of toes and on medial sides of hind limbs 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 m1. + + 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 M1 or than between anterior palatine foramen and + anterior margin of tympanic bulla; postorbital 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 5-1/2 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 than rostrum. + + The skull of the female averages 53 per cent lighter than that of + the average male. + +Comparisons of the skull with those of _M. f. olivacea_, _M. f. +spadix_, _M. f. primulina_, and _M. f. arthuri_, are made in the +accounts of those subspecies. As compared with that of _M. f. 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-Linnaean references to this weasel +mostly were under the specific name _erminea_ in the belief 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 confused with some other species. Audubon and Bachman in +1853 even proposed the name _agilis_ for the female in the mistaken +belief 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 mammals 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 accomplishments 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 subadult +males from western Wisconsin, namely, one from Gordon, 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 _noveboracensis_ 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 slightly 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. +f. 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 underparts 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 geographic range of _olivacea_. + +Notwithstanding the state of affairs described above, intergradation +seems to take place. Three specimens referred to _noveboracensis_ 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 approaches 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 tendency +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., [M] 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 subadult male, no. 4193, Mus. Comp. Zoöl., from Upton, +Maine, has a longer (51 mm.) hind foot than _noveboracensis_ although +it otherwise 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 +_noveboracensis_ and _occisor_ intergrade. Therefore they are treated +as two subspecies of the single species, _Mustela frenata_. + +Intergradation with _M. f. primulina_ has been commented on in the +discussion of that subspecies. Female, no. 159980, U. S. Nat. Mus., +from Golconda, Illinois, has many characters of _primulina_ 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 Michigan 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 County_: Fort Payne, 1. + + =Connecticut.= _Litchfield County_: Riverton, 1[5]; Gaylordsville, + 1. _Hartford County_: East Hartford, 4 (3[5]); Glastonbury, 2[5]; + South Glastonbury, 4[5]. _Windham County_: Plainfield, 2 (1[14]). + _Fairfield County_: Greenwich, 2[2]. _New London County_: Liberty + Hill, 35 (33[75], 2[7]). + + =District of Columbia.= Washington, 3; near Washington, 1; Eastern + Branch, 1; Congress Heights, 1; Benning, 1; no definite locality, + 1. + + =Georgia.= _Towns County_: Young Harris, 1. _Cherokee County_: + Canton, 1. _Cobb County_: Roswell, 1[81]. + + =Indiana.= _St. Joseph County_: Notre Dame, 2[99]. _Porter + County_: Hebron, 1. _Miami County_: Denver, 5 (4[75], 1[4]). + _Wells County_: Bluffton, 1. _Howard County_: Russiaville, 1. _Jay + County_: Salamonia, 1[2]. _Boone County_, 1[2]. _Knox County_: + Bicknell, 3. + + =Illinois.= _Lake County_: Camp Logan, 3[60]; Fort Sheridan, + 1[60]. _Cook County_: W Northfield, 2; Flossmoor, 1[60]; no + locality more definite than county, 1. _Du Page County_: + Bloomingdale Spg., 1[60]. _Carroll County_: Savanna, 1[87]. + _McLean County_: Normal, 1[7]. _Champaign County_: Harwood + Township, 1[7]. _Pike County_?: Milton Spring, 1[60]. _Pope + County_: Golconda, 3. + + =Kentucky.= _Woodford County_: Midway, 1. _Hancock County_: + Hawesville, 1. + + =Maine.= _Oxford County_: Upton, 1[75]; Bethel, 1[74]. + + =Maryland.= _Howard County_: Long Corner, 1; Hanover, 1. + _Montgomery County_: Gaithersburg, 1; Garret Park, 1; Chevy Chase, + 1; Bethesda, 1. _Prince Georges County_: Laurel, 18; Plummer + Island, 3; Oxon Hill, 1. _Talbot County_: Easton, 1. _Dorchester + County_: Cambridge, 5[40]. + + =Massachusetts.= _Middlesex County_: Wilmington, 6; Burlington, 6; + Lexington, 1[75]; Wayland, 2[75]. _Berkshire County_: New + Marlboro, 1[5]. _Worcester County_: Lunenburg, 2; Lancaster, + 1[75]; Princeton, 2[75]. _Norfolk County_: So. Weymouth, 1[75]. + _Plymouth County_: Wareham, 5[75]. + + =Michigan.= _Marquette County_: Michigamme, 1. _Charlevoix + County_: Thumb Lake, 1[76]; 1/2 mi. N Thumb Lake, 1[76]. _Leelanau + County_: Leland, 3[76]; Duck Lake, 2 mi. S Leland, 1[76]; Lost + Pond, 8-1/2 mi. S Leland, 1[76]. _Osceola County_: Le Roy, 2[76]. + _Huron County_: Rush Lake, 1[76]. _Saginaw County_: East Saginaw, + 1. _Oakland County_: Royal Oak, 4[76]; South Lyon, 1[76]. + _Livingston County_: Portage Lake, 1[76]. _Washtenaw County_: + Portage Lake, 6[76]; Waterloo, 2[14]; Lima, 1[76]; Ann Arbor, + 11[76]; 3 mi. E Ann Arbor, 1[76]; 2 mi. SE Ann Arbor, 1[76]; 2 mi. + S Ann Arbor, 1[76]; 3 mi. S Ann Arbor, 1[76]; Dixboro, 1[76]; + Pittsfield, 3 (2[76]); Saline, 1[76]; near Saline, 2[76]; 1 mi. S + Saline, 2[76]; York, 2[76]; Manchester, 2[76]. _Lenawee County_: + Morenci, 1[76]. _Cass County_: Marcellus Township, 1[76]. _Berrien + County_: Harbert, 1[76]; Warren Wood Preserve, 1[76]; Warren + Woods, 1[76]. + + =New Hampshire.= _Grafton County_: Franconia, 1[2]. _Carroll + County_: South Chatham, 4 (3[5]); Ossipee, 2; Intervale, 1[5]. + _Merrimack County_: Webster, 2[75]. + + =New Jersey.= _Morris County_: Morristown, 1. _Essex County_: West + Orange, 1[2]. _Mercer County_: Princeton, 1[1]. _Ocean County_: + Point Pleasant, 1[2]. _Camden County_: Haddonfield, 1[1]. + _Cumberland County_: Millville, 2[74]. + + =New York.= _St. Lawrence County_: Ogdensburg, 1[74]. _Clinton + County_: Rouses Point, 1[80]. _County_?: Adirondacks, 12. _Essex + County_: Elizabethtown, 1; Schroon Lake, 1; no locality more + definite than county, 1. _Lewis? County_: Locust Grove, 4; Lyons + Falls, 1. _Warren County_: Lake George, 6; Caldwell, 1. _Hamilton + County_: Beaver Brook, 1/2 mi. above mouth Indian Lake, 1[80]. + _Oswego County_: Scriba, 2[74]; Palermo, 1[74]. _Monroe County_: + Penfield, 3. _Madison County_: Peterboro, 6 (2[75]). _Schoharie + County_: Schoharie, 1[2]. _Rensselaer County_: East Shodack, + 1[80]. _Tompkins County_: Taughannock Falls, 2[58]; Ithaca, 4 + (3[58]); Glenside, Ithaca, 1[58]; 6 mi. Creek, Ithaca, 1[58]. + _Green County_: Lanesville, 1[2]. _Orange County_: Poplopen's + Pond, 1[2]; Highland Falls, 1[2]. _Putnam County_, 1[19]. + _Westchester County_: Sing Sing, 4; Armonk, 1[2]; Hastings, 3 + (2[2], 1[19]). _Nassau County_: Flushing Meadows, 1[2]; Flushing, + 1[58]; near Flushing, 1[2]; Oyster Bay, 2. _Long Island_: Cold + Spring Harbor, 1; Bridgehampton, 1[2]. _County_ in question: + Severance, 3; Lake Grove (Long Island?), 1. + + =North Carolina= (east to west by counties). _Wake County_: + Raleigh, 4 (1[2], 1[75], 2[76]). _Mitchell County_: Magnetic City, + foot of Roan Mountain, 6; Roan Mt., 1; Roan Mt., 6000 ft., 3. + _Buncombe County_?: Valley of Black Mts., 1[2]. _Madison County_, + 2[11]. + + =Ohio.= _Trumbull County_: Warren, 1[93]. _Seneca County_: Tiffin, + 1[81]. _Summit County_: Ira, 2[81]. _Crawford County_: Galion, + 1[81]. _Ashland County_: Loudonville, 1[76]. _Auglaize County_: + New Bremen, 3[81]. _Franklin County_: 3 mi. N Columbus, 1[81]; + Minerva Park, Columbus, 5[81]. _Fairfield County_: Sec. 32, + Pleasant Twp., 1[81]; Lancaster, 1[81]. _Clinton County_: + Reesville, 1; 1/2 mi. S and 1/2 mi. W Wilmington, 2[74]. _Pike + County_: Waverly, 1[81]. + + =Ontario.= _Sudbury District_: Metagama, 2[86]. _Carleton County_: + Ottawa, 2[77]. _Muskoka County_: Lake of Bays, 1; Bracebridge, 1. + _Haliburton County_: Gooderham, 1[60]. _Simcoe County_: Orillia, 4 + (2[2], 2[60].) _Prince Edward County_: Bloomfield, 1[77]. _York + County_: Toronto, 1[2]. _Waterloo County_: Branchton, 3[60]; + Preston, 2[77]; no locality save county, 1[60]. _Welland County_: + Ridgway, 1[14]. _Elgin County_: St. Thomas, 1[77]. _Essex County_: + Kingsville, 1[77]; Point Pelee, 1[77]. + + =Pennsylvania= (east to west by counties). _Crawford County_: + Pymatuning Swamp, 3-1/2 mi. W Linesville, 1[9]; Meadville, 2[9]. + _Beaver County_: Beaver, 1[9]; Raccoon Creek, 1[9]. _Butler + County_: Mars, 1[9]; Leasuresville, 4[9]. _Allegheny County_: + Allegheny, 1. _Warren County_: Bear Lake, 2[9]. _Westmoreland + County_: Kingston, 1[9]; Laughlinstown, 2[9]. _Somerset County_: + Confluence, 1[9]; Tub Mill Run, 2 mi. N Springs, 1[9]. _Jefferson + County_: Siegel, 1[9]. _Clearfield? County_: Penfield, 1[9]. + _Cambria County_: Cresson, 1[9]. _Fulton County_: Well's Tannery, + 1[9]. _Clinton County_: near Round Island, 2[1]. _Cumberland + County_: Carlisle, 1. _Snyder County_: 5 mi. S Selinsgrove, 1. + _Northumberland County_: Pottsgrove, 1. _Union County_: + Mifflinburg, 1. _Sullivan County_: Lopez, 7 (4[1], 3[74]). + _Chester County_: Westtown, 1[1]; Valley Forge, 1[1]; W Bradford + Township, 1[1]; no locality more definite than county, 3. + _Philadelphia County_: Holmsburg, 2[1]. _Bucks County_: 1[1]. + _Pike County_: Milford, 1. + + =Rhode Island.= _Providence County_: Chepachet, 1. _Washington + County_: Lake Warden, 2. + + =Quebec.= _Megantic County_: Black Lake 1[77]. _County_ in + question: Meach Lake, 1[77]. + + =South Carolina.= _York County_: 5 mi. E York, 1[11]. _Laurens + County_: Laurens, 1[39]. + + =Tennessee.= _Campbell County_: Highcliff, 1. _Carter? County_: + Roan Mts., 1[2]. _Hamilton County_: Walden Ridge, near Soddy, 3. + + =Vermont.= _Windsor County_: Hartland, 1[2]. + + =Virginia.= _Shenandoah County_: Toms Brook, 1. _Arlington + County_: Arlington, 1; Ballston, 1; Alexandria, 1. _Fairfax + County_: Falls Church, 3; Mt. Vernon, 2; no locality more + definite than county, 1. _Prince William County_: Occoquan, 1. + _Essex County_: Montague, 1. _Prince George County_, 1. _Norfolk + County_: Wallaceton, 1. _Grayson County_: Mt Rogers, 3. _County_ + in question: Dismal Swamp, 1; Massanutten Mt., 1. + + =West Virginia.= _Hardy County_, 1. _Pendleton County_: radius of + 2 mi. Smoke Hole, 1[74]. _Greenbriar County_: White Sulphur, + 2[60]. + + =Wisconsin.= _Douglas County_: Gordon, 1[104]. _Vilas County_: + Mamie Lake, 4. _Dunn County_: Colfax, 4[104]; Meridean, 1[104]. + _Door County_: state game farm, 17[104]; no locality more definite + than county, 1[104]. _Dodge County_: Rolling Prairie, 1[50]; + Beaver Dam, 52[50]. _Dane County_: Wingra Lake, 1[104]. _Waukesha + County_: Pewaukee, 2[104]. _Racine County_: Waterford Township, + 2[104]. _Rock County_: Milton, 1[104]; Bowers Lake, 1[104]. + _Walworth County_: Lane's Mill, 8 mi. N Elkhorn, 7 (1[104], + 6[54]); 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. Zoöl.; Bucksport, Hancock County, Maine; + January 15, 1899; obtained 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 zygomatic arch broken away; otherwise the skull + is unbroken and complete. Left I3 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._--Differs from _M. f. + 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 measurements 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. + + 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 differences 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 _Mustela frenata noveboracensis_. + + _Color._--As described in _Mustela frenata noveboracensis_ except + that black tail-tip in series of 10 males in full winter 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 that: + 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 material 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, Quebec, 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 characters 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 of _occisor_. +However, this specimen is much older than any _occisor_ examined. In a +female, no. 4260, Mus. Comp. Zoöl., from Liberty Hill, Connecticut, the +skull is longer (but narrower) 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 +comparison with true _noveboracensis_ without much question are +geographic variations. Whether or not these variations are of a degree +sufficient to warrant nomenclatural recognition is debatable. With +equally scanty material from other regions I have not named variations +seemingly as great as those shown by _occisor_. The paucity of material +of _occisor_ is a handicap in making a decision in this instance. + +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 Zoölogy. + + =Maine.= _Oxford County_: South Andover, 2 (Boston Soc. Nat. + Hist.); Umbagog Lake, 1. _Franklin County_: Seven Pd. Township, 1. + _Piscataquis County_: Grenville, [= Greenville?], 1. _Hancock + County_: 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, Journ. Mamm., 4:108, May 9, + 1923. + + _Mustela longicauda primulina_, Linsdale, Journ. Mamm., 9:141, May + 9, 1928. + + _Mustela frenata primulina_, Hall, Carnegie Instit. Washington + Publ. 473:104, November 20, 1936. + + _Mustela frenata_, Leopold and Hall, Journ. 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. + + _Range._--Upper and Lower Austral life-zones west of the + Mississippi River in Missouri and Arkansas, the southeastern half + of Iowa, eastern half of Kansas and Oklahoma, northern Louisiana + and northeastern Texas. Southern and southwestern limits of range + undetermined. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. + noveboracensis_ in males by interorbital breadth averaging less + than 24 per cent of basilar length, orbitonasal length averaging + less than 34 per cent of basilar length or 64 per cent of mastoid + breadth, tympanic bullae as much inflated anteromedially as + posteromedially, 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 than 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 width of color of upper parts, absence of color of + underparts on hind leg below knee, zygomatic breadth less than + 28.8 in males and 24.1 in females; from _M. f. neomexicana_ 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. f. frenata_ and _M. f. texensis_ by absence of + white facial markings and smaller size (basilar length 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 pls. 16, 17 and 18), greater + inflation of the tympanic bullae anteromedially and more nearly + straight (less convex) dorsal outline of skull. + + _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 differences 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. + + _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; hairiness 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 Oberthür and + Dauthenay, pl. 301. Chin and rarely upper lips white. Remainder of + underparts Picric Yellow to Primuline 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 underparts 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 M1 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 bulla; + 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 width of 2-1/2 to 5 upper incisors; height of + tympanic bulla more (except in one specimen) 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 + (except in one specimen) than rostrum; anterior margin of + masseteric fossa behind or just below posterior eighth of m1. + + 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 foramen and M1 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; tympanic + 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 rostrum. + + The skull of the female averages 41 per cent lighter than that of + the male. + +Compared with the skull of _M. f. noveboracensis_ from Massachusetts, +that of the male of _primulina_, in dorsal view, is seen to be shorter +anteriorly to the postorbital processes and to have a more marked +postorbital constriction. In lateral view the dorsal outline of the +skull of _primulina_ is less concave in the postorbital region. In +ventral view the skull of _primulina_ 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 difference is in +some parts greater, for example, in the tympanic bullae. In +_primulina_, the bulla is in general like that of the male +_noveboracensis_, whereas in the female _noveboracensis_ it is less +inflated, especially anteromedially, shorter, relatively narrower, and +in ventral view projects little or none below the squamosal floor of +the braincase. The breadth of the bulla averages 51 per cent of its +length in _primulina_ but only 47 per cent in _noveboracensis_. The +bullae project 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 +developed 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 _noveboracensis_ 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. Expressed in +percentages of the basilar length, the two depth measurements of the +skulls are not significantly different, but, excluding the measurements +of the bullae and teeth, the other cranial measurements 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 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. longicauda_, 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 M1 +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 narrowness of +the skull of _primulina_. Upon direct comparison the narrowness is +especially noticeable in the interorbital region, mastoid 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 +corner of bulla rounded rather than "square," and in males a +transversely convex rather than flat interorbital region. + +Compared with _M. f. frenata_ and _M. f. 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 +zygomatic breadth and are less constricted interorbitally. + +Compared with the skull of _M. f. 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. f. 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 southwestern Missouri. Later, +through the efforts of Charles D. Bunker, 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, Lawrence V. Compton obtained a topotype for the California +Museum of Vertebrate Zoölogy, 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. +noveboracensis_, 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 7-1/2 miles +southeast of Carthage, a subadult male in brown, winter pelage, is +almost exactly intermediate between that of _noveboracensis_ from +Massachusetts and _primulina_ of Douglas County, Kansas, and Boone +County, Arkansas. _M. f. primulina_ 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 available +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. + +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 female is smaller than +_primulina_ and in this small size and in general configuration of the +skull, viewed dorsally, is more nearly like _noveboracensis_. As a +whole, the population averages almost exactly intermediate. The same is +true of 3 males and one female from Muscatine. 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 mastoidal region, but otherwise is more +like that of _primulina_. Two males and one female from Tipton, +although in each instance variously 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 Vinton, one adult is typical of +_primulina_ but the other, a subadult, is practically indistinguishable +from female _noveboracensis_, from Ann Arbor, Michigan. Three males +from Vinton agree well with _primulina_ 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 _primulina_. + +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 example, 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 frequently found in the +_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 underparts, 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 the account of _spadix_. + +The specimen from Swartz, Louisiana, suggests intergradation with +_arthuri_ in that the 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 pattern 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 +prevents judging of its relationships on the basis of relative +proportions of the skull. + +The two specimens, skins only, available from Oklahoma, are +provisionally referred to _primulina_. These are remarkable for the +restriction 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 distribution of the +color of the underparts cannot be made out. + +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. f. 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 _longicauda_ 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 indicated, specimens are in the University of + Kansas, Museum of Natural History. + + =Arkansas.= _Boone County_: 3 mi. E Bergman, 4[74]; 3 mi. SE + Bergman, 1[74]; 3 mi. S Bergman, 1[74]; 3 mi. SW Bergman, 1[74]; 4 + mi. SE Bergman, 2[74]; 5 mi. SE Bergman, 1[74]; 4-1/2 mi. SE + Bergman, 3[74]; 5 mi. SE Bergman, 1[74]; 5 mi. S Bergman, 2[74]; 5 + mi. SW Bergman, 2[74]. _Washington County_: Fayetteville, 1[96]. + _Crawford County_: 10 mi. S Winslow, 1. _Sebastian County_: Fort + Smith, 1[91]. + + =Iowa.= _Fayette County_: Fayette, 3[12]. _Dubuque County_: + Dubuque, 1[12]; Green's Island, 1[65]. _Butler County_: New + Hartford, 1[12]. _Hardin County_: Union, 1[65]. _Benton County_: + Vinton, 5[12]. _Linn County_: Cedar Rapids, 1[12]. _Boone County_: + Worth Township, Sec. 21, 1[65]; 2 mi. E Ledges St. Park, 1[65]. + _Story County_: Kelley, 1[65]. _Cedar County_: Tipton, 3[12]. + _Scott County_: Davenport, 2[12]. _Muscatine County_: Muscatine, + 4[12]. _Henry County_: Hillsboro, 5[91]. _Van Buren County_: + Keosaqua, 1[65]; no locality more definite than county, 1[50]. + _Taylor County_, 1. + + =Kansas.= _Riley County_: near Winkler, 1. _Pottawatomie County_: + Onaga, 1[83]. _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; 1-1/2 mi. W Lawrence, + 1; 6 mi. S Lawrence, 1; 7 to 7-1/2 mi. SW Lawrence, 14; 10 mi. W + Lawrence, 1; Clinton, 4; Baldwin, 1; no locality more than county, + 29 (2[74]). _Woodson County_: 1-1/2 mi. S Neosho Falls, 1[59]. + _Greenwood County_: 8 mi. SW Toronto, 2. _Pratt County_: Pratt, 1. + + =Louisiana.= _Quachita Parish_: Swartz, 1[71]. + + =Missouri.= _Carroll County_: Bridge Creek, 1[91]. _Lincoln + County_: Silex, 1[74]; 1 mi. E Silex, 1[74]. _Reynolds County_: 4 + mi. S Lesterville, 1[74]. _Jasper County_: 5 mi. NE Avilla, 1[91]; + 7-1/2 mi. SE Carthage, 1[74]. + + =Oklahoma.= _Payne County_: 8 mi. NW Stillwater, 1[82]. _Cleveland + County_: Norman, 1[100]. + + =Texas.= _Hopkins County_: 10 mi. S Sulphur Springs, 1[43]. + + +=Mustela frenata arthuri= Hall + +Long-tailed Weasel + +Plates 16, 17 and 18 + + _Mustela noveboracensis 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, skull and skin; no. 37515, Mus. Vert. + Zoöl.; 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 unbroken. 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 + outline of skull in longitudinal axis (see pls. 16-18); from _M. + f. noveboracensis_, 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 Oberthür + and Dauthenay, pl. 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 Oberthür + and Dauthenay, pl. 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 few scattered hairs. Color of + underparts extends distally on medial sides of hind limbs only to + knees. Least width of color of underparts 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. + + _Skull and teeth._--Male (based on type and 2 subadults): See + measurements and plates 16-18. As described in _M. f. + 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 opticum + and anterior margin of tympanic bulla; least width of palate more + or less than length of P4; tympanic 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 westward. + +Compared with the skull of _M. f. olivacea_ that of _arthuri_ differs +as follows: Averaging smaller in every part measured; basilar length 5 +mm. less; by weight a fourth lighter; 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 tympanic 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 anteriorly, 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. f. 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. f. 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 +anteromedially. 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 Vertebrate Zoölogy. 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 +obtained 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, Assumption 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. +Zoöl., 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 _M. f. olivacea_ 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 extend farther west in the +latitude of northern Louisiana than anywhere 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 County_: 12 mi. N Eagle Lake, 1[43]; 5 mi. W + Eagle Lake, 1[43]; 3 mi. S Garwood, 1[43]. + + =Louisiana.= _East Baton Rouge Parish_: Baton Rouge, 4[71]. + _Livingston Parish_: Springville, 1[74]. _Saint James Parish_: + Convent, 1[91]; Remy, 2 (1[74], 1[45]). _Assumption Parish_: near + Lake Verret, 1[45]. + + =Mississippi.= _Harrison County_: Saucier, 1[71]. + + +=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. + + _Type._--Male, adult, skull and skin; no. 180802, U. S. Nat. Mus., + Biol. Surveys 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 extreme 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. f. + peninsulae_ in finer, softer pelage and shorter (less than 15.8 in + ad. [F]) tympanic bullae; from _M. f. noveboracensis_, 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. f. 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 pls. 16-18). + + _Description._--_Size._--Male and female: External measurements + of adults are available as follows: + + Length Length + Catalogue Sex Locality Total of of hind + no. length tail foot + 47165 [M] Box Springs, 454 160 48 + Talbot Co., Georgia + + 47166 [M] Box Springs, 435 147 47 + Talbot Co., Georgia + + 47167 [M] Box Springs, 422 145 45 + Talbot Co., Georgia + + 41023 [M] Thomas Co., 443 140 47 + Georgia + + 41025 [M] Grady Co., 395 142 47 + Georgia + + 223880 [M] Okefinokee Swamp, 416 145 49 + Georgia + + 198 [M] Okefinokee Swamp, 425 140 48 + Georgia + + Average 7 [M] 427 146 47 + + 49385 [F] Gainesville, 396 124 45* + Alachua Co., Florida + + 41024 [F] Thomas Co., 380 125 41 + Georgia + + 51527 [F] Talbot Co., 376 128 43 + Georgia + * [not typical] + + 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 measurements 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 shown in + figure 19. + + _Color._--Upper parts, in summer, near tone 4 of Burnt Umber of + Oberthür and Dauthenay, pl. 304. In winter lighter, between tones + 3 and 4 of Raw Umber of Oberthür and Dauthenay, pl. 301. Dark spot + at each angle of mouth present or absent. Underparts ranging from + Massicot Yellow to Cream Buff except on chin and upper lips 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 mouth 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 M1 + 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 + length; 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; 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 M1 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 I3) 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. f. olivacea_ +averages smaller and has relatively and actually smaller and less +inflated bullae. As compared with the skull of _M. f. 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 +differences 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 (including I3); 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. f. arthuri_ and _M. f. primulina_ are made +in the accounts of those subspecies. + +_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 obtained three instructive +specimens from Okefinokee Swamp. Really adequate material, for the +localities represented, owes its preservation 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. peninsulae_ is not +satisfactorily established due to inadequate material of _peninsulae_. +Differences 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, therefore, 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 _peninsulae_ would apply to all. Examination of +more material from Florida, especially from the southern half of +Florida, will be necessary to answer this question. + +This large weasel of the southeastern United States is remarkably +different from _noveboracensis_. Indeed, were it not for actual +intergrades 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 referred 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 occurs +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 _noveboracensis_ to +specimens from southern localities adjoining the territory occupied by +_olivacea_. One of these features subject to gradual change is the +color of the underparts. Beginning at the Adirondacks of New York +where a large number of the specimens have white underparts, the +underparts become more intensely yellowish 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 _peninsulae_. 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 intensity 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 _noveboracensis_ in lesser +size of the skull and agree with _noveboracensis_ in the narrowness of +the color of the underparts. Also, these specimens, like 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 _peninsulae_, 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 distinctly +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 _noveboracensis_ 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 Creek is referable to +_olivacea_, 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 _olivacea_ although the skull shows +some approach to that of _noveboracensis_. Of the two males from near +Nauvoo, no. 51652 is to me indistinguishable from _noveboracensis_, but +no. 51653 does have some characters of _olivacea_, although on the +whole, the latter, too, seems to be a little nearer _noveboracensis_ +than _olivacea_. However, because the mean of these seven specimens +from northwestern Alabama is nearer _olivacea_ than _noveboracensis_ +the former name may be applied. + +Another specimen from "Souinlonie" Creek, Clark County, Mississippi, +has the coloration and rostral configuration of _primulina_, narrow +mastoidal breadth and smaller teeth of _noveboracensis_ and skull of +large size with "full" braincase as in _olivacea_. No. 235364, U. S. +Nat. Mus., from the Mobile River at the "L. and N. RR. Crossing," +Mobile County, Alabama, although definitely _olivacea_, shows approach +to _arthuri_ in that the dorsal outline of the skull is longitudinally +more convex and the tympanic bullae are less inflated than in +_olivacea_ 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 unknown the identification +as _olivacea_ 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 Charleston 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. f. noveboracensis_. + + _Specimens examined._--Total number, 52, arranged alphabetically + by states and from north to south by counties in each state. + Except as otherwise indicated specimens are in the University of + California Museum of Vertebrate Zoölogy. + + =Alabama.= _Lawrence County_: White Creek, 1; Little Sand Mt., + Shoal Creek, 2. _Winston County_: 7-1/2 mi. N Nauvoo, 1; 8 mi. N + Nauvoo, 1. _Lauderdale County_: near Leighton, 9 mi. N Tennessee + River, 1[91]. _Colbert County_: Leighton, 1[91]. _Autauga County_: + Autaugaville, 1[91]. _Dale County_: Midland City, 1[91]. _Mobile + County_: Mobile River, 12 mi. NE Mobile, 1[91]. + + =Florida.= _Alachua County_: Gainesville, 4[61]. _Marion County_: + "Silver Springs," 1. + + =Georgia.= _Spalding County_, 1. _Lamar County_, 1. _Talbot + County_: southwest 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[91]; locality no more + definite than county, 4. _Thomas County_: Sinkola Plantation, 2; + locality no more definite than county, 2. _Charlton County_: 1/2 + mi. E Chesser's Island, Okefinokee Swamp, 1[58]. _County_ in + question: Billy's Island, Okefinokee Swamp, 1[91]; Okefinokee + Swamp, 1[58]. + + =Mississippi.= _Clark County_: Souinlonie Creek, 1. + + =South Carolina.= _Darlington County_: Society Hill, 1[91]. + _Sumter County_: Mayesville, 1[11]. _Calhoun County_: St. + Matthews, 2[11]. _Georgetown County_: Sampit, 1[11]. Charleston + County: Rantowles, 1[11]; 8 mi. N Charleston, 1[11]. _Beaufort + County_: Yemassee, 1[2]. + + +=Mustela 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, 1912. + + _Mustela p. peninsulae_, Bailey, Bailey Mus. and Library Nat. + Hist., 1(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, the 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 + light facial markings are more extensive than in any of the + referred specimens. In the type these light 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. + + _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; length 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 + Oberthür and Dauthenay, pl. 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 uniform color. Color of underparts + extends distally on legs over both sides of feet and on front legs + over wrists. Proximal part of tail slightly lighter 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 both 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 + measurements. 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. f. olivacea_, the insufficient material of _M. +f. peninsulae_ 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, +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. f. +olivacea_ from Gainesville, Florida, has well-developed light (white) +facial markings. Also of the four specimens of _M. f. 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[1]. _Pinellas County_: + Tarpon Springs, 1[1]. _Hernando County_, 1[91]. _Polk County_: + Auburndale, 1[91]; no locality more definite than county, 1[91]. + _Orange County_: Apopka, 3[61]. _Volusia County_: Enterprise, + 1[60]. _Seminole County_: Osceola, 1[2]. + + +=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, February 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 Edgar A. Mearns; original no. 812. + + The skull is complete although there are fractures on the top of + the braincase, on the right side of the braincase and at the + middle of the right zygomatic arch. The teeth all are present and + entire. The skin, although overstuffed, 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. f. 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 three 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: Three adults from Elk River, Minnesota, yield average and + extreme 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 (approximately 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 lips 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 to ankle, over antiplantar faces of toes and + distomedial fourth of each tarsus, 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 the greater width of the light-colored + underparts and relatively short black tip of the tail, both + features of _M. f. longicauda_, _spadix_ is variously + intermediate, depending on locality, 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 + 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 4 to 5 upper + incisors; height of tympanic bulla more or less than distance from + its anterior margin to foramen ovale; length of tympanic bulla + less than length of rostrum; anterior margin of masseteric fossa + below talonid of m1. + + 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 width 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 the female averages 33 per cent lighter 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 the 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 tympanic 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 braincase, measured at the +anterior end of the basioccipital, amounts to only 56 per cent of the +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 appearance 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 shallower braincase and is +relatively deeper through the postorbital processes, but these +differences 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 _longicauda_. + +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; squamosal +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 discussion of that +subspecies. + +_Remarks._--Edgar A. Mearns in 1889 and the early nineties took several +specimens of this weasel and it was principally on these that Bangs in +1896 (p. 8) based his description. The best material, however, 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 geographically 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 the characters ascribed +to _spadix_, including large size itself, undergoes change from one +part of its geographic range to another; the characters 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 localities have been found from which the specimens can be said +really to agree in characters. + +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 lighter than in +_noveboracensis_. Farther westward the average coloration is a little +lighter and farther westward yet, toward the range of the extremely +light colored _longicauda_, the average coloration is lighter still. +Although all these animals are darker than _longicauda_ 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 geographically 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. +f. 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 +characteristic 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 below 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 diluted than is the _noveboracensis_ type of underparts. +Indeed, within the range of _noveboracensis_ itself, the broad extent +of the _longicauda_ type of underparts is manifest. This is, of course, +near the western margin of the range of _noveboracensis_. + +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 assigned 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 with size as a unique character distinguishing _spadix_ from +_longicauda_ and _noveboracensis_. 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 +_noveboracensis_. This is true, in _spadix_, of the relatively longer +(in comparison 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 +_noveboracensis_ 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 regarded as +specifically distinct from the _noveboracensis_ type of animal, comment +is offered below on selected specimens, referred to _spadix_, which are +regarded as intergrades with _noveboracensis_ or with other subspecies. + +No. 8722, Univ. Wisconsin, adult male, in the white winter coat, from +north central Itasca County, Minnesota, obviously has characters of _M. +f. spadix_ or _longicauda_ that occur to the west and _M. f. +noveboracensis_ of the east. Selected outstanding characters of +_longicauda_ are its long tail, anteriorly truncate tympanic bullae +and large teeth. Characters indicating its affinities with +_noveboracensis_ 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 +truncate bullae constitute basis for here referring the specimen to +_spadix_. However, the seemingly small size of the body and the narrow +skull clearly show relationship to _noveboracensis_. + +Specimens, referred to _spadix_, from northern Iowa, are instructive as +showing what happens where the ranges of _noveboracensis_, _primulina_, +_spadix_, and perhaps _longicauda_, meet. No. 47167, Univ. Mich. Mus. +Zoöl., 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 underparts onto the feet, agrees with +_spadix_ or _longicauda_ as does also the long tooth-row. The overall +length of the skull is intermediate between that of _spadix_ and +_primulina_. The proportions of the anterior part of the skull and of +the tympanic bullae resemble those found in _primulina_. A subadult +male skull only, no. 123846, American 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 _longicauda-spadix_ +type of light-colored underparts and color pattern, and is slightly +darker above than true _longicauda_. Another subadult 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 restriction of the +area of the light color of the underparts and in less expanded +zygomatic arches. The teeth are intermediate in size between 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_. + +An adult female, no. 426a, Iowa State College, from Barnum, in the +brown winter coat, agrees with _primulina_ except that the orbitonasal +length of the skull is more as in _spadix_ and the presence of some +light color on the lower part of the hind legs suggests _spadix_. The +skull only, no. 440a, Iowa State College, labeled merely Webster +County, Iowa, is almost a duplicate of no. 426a. A subadult male, no. +427a, Iowa State College, from Moorland, Iowa, only about six miles +southeast of Barnum, likewise is indistinguishable from _primulina_ +except for having a white winter coat and in being relatively broad in +the mastoidal region. Nevertheless, 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 Alberta. 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 practically 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 _noveboracensis_ 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 all +respects typical of _spadix_. This is the more remarkable because +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 _noveboracensis_. 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 restricted +somewhat although not so much as in _noveboracensis_ or _primulina_. +Of four males from the same place, also in the collection 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, uninterruptedly, 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 departures they show some approach to _primulina_ and to +_noveboracensis_. Another male, subadult, no. 2867, Coe College, from +Decorah, 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 _primulina_. + +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 narrower 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 together at the Coe +College Museum by the late B. H. Bailey. Most of them were obtained +from trappers who did not supply the conventional external measurements +taken in the flesh. Even though these are lacking, the specimens +clearly show that actual intergradation occurs where the ranges of _M. +f. 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 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 Sisseton, South Dakota, no. 188407, +United States National Museum, figured by Merriam (1896, p. 20, figs. +7-9), is almost exactly intermediate 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. f. longicauda_, _M. f. primulina_ +and _M. f. spadix_. One skin is in white winter pelage and the other, a +female, is in summer pelage which in coloration and color pattern +agrees with that of _spadix_. External measurements 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 _longicauda_ or _spadix_ although nearer the latter in this +respect. In dorsal aspect, the skulls especially posteriorly to the +orbital region, resemble _primulina_. All points considered, the +animals seem best referred to _spadix_. + +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; Armour, South Dakota and Clay County, South +Dakota. In summary, 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 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.= _Lyon County_: Granite, 1[65]. _Howard County_: Chester, + 6[12]. _Winneshiek County_: Decorah, 1[12]; 8 mi. NE Ossian, + 1[76]. _Allamakee County_: Lansing, 2[12]. _Clay County_: Island, + 1[76]; Webb, 1[2]. _Palo Alto County_: Ruthven, 1[76]; no locality + more definite than county, 1[76]. _Calhoun County_: Manson, 1[65]. + _Webster County_: Barnum, 1[65]; Moorland, 1[65]; no locality more + definite than county, 1[65]. _Boone County_: Pilot Mound, 3[12]; + Amaqua Township, Sec. 19, 1[65]. _Story County_: Ames, 1[65]. + + =Minnesota.= _Kittson County_, 1[2]. _Roseau County_: 2-1/2 mi. SW + Roseau, Jadis Township, 1[14]. _Itasca County_: T. 61N, R. 26W, + 1[104]. _Clay County_: Moorhead, 2[9]. _Atkin County_: Atkin, + 1[50]. _Otter Tail County_: Lake Lizzie, 1[9]; Parkers Prairie, + 1[57]. _Grant County_: 3 mi. NW Barrett, 1[76]. _Benton_ (now + Mille Lacs?) _County_: Princeton, 1[91]. _Sherburne County_: Elk + River, 14 (6[59], 4[14], 3[91], 1[74]). _Hennepin County_: Fort + Snelling, 6 (5[2], 1[91]). _Carver? County_: Chaska, 1[60]. _Lac + qui Parle County_: Madison, 5 (3[91], 2[1]); no locality more + definite than county, 2 (1[68], 1[75]). _Yellow Medicine County_: + Wood Lake, 1[2]. _Blue Earth County_: Rapidan, 1[64]. _County_ in + question: Moore Lake, 1[91]. + + =Nebraska.= _Cuming County_: Beemer, 5[91]. + + =North Dakota.= _Cass County_: Fargo, 1[91]; Casselton, 1[91]. + _Dickey County_: Oakes, 1[91]. + + =South Dakota.= _Roberts County_, 1[102]. _Marshall County_: Fort + Sisseton, 1[91]. _Douglas County_: Armour, 1[14]. _Clay County_, + 1[102]. + + +=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, 1838. + + _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, 1a of pls. 1, 2 and 3, February 25, + 1896; Merriam, N. Amer. Fauna, 11:19, pl. 3, figs. 3, 3a, 4, 4a, + pl. 5, figs. 1, 1a, 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, Journ. Mamm., 29:126, May 14, 1948. + + _Type._--Possibly not in existence. No. 43.3.3.3 [from Carlton + House, Saskatchewan] in the British Museum of Natural History has + been regarded by several zoölogists as the type. It is a subadult + female, skull and skin, 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, southward from central Alberta, Saskatchewan and southern + Manitoba through eastern Montana, the Dakotas and Nebraska into + southeastern Wyoming, northeastern Colorado and western Kansas. + See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. + primulina_ in near (_h_) Clay Color rather than Brussels Brown of + upper parts, least width 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 lighter 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 (14_n_) 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 (14_n_ to 1) Brussels Brown of upper parts, basilar + length more than 40 in females and averaging more than 45 in + males; from _M. f. 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 average 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. + + The average differences in external measurements of the two sexes + are: Total length, 37; length of tail, 13; length of hind foot, 7. + General comparisons 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 extending 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 Oberthür and Dauthenay, pl. 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. f. 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. oribasus_ and _M. f. 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 underparts, and light rather than + dark-colored hind feet. + + _Skull and teeth._--Male (based on 5 adults from Alberta): See + measurements 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 M1 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; 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 I3) 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 m1 + or anterior half of m2. + + Female (based on 5 adults: Alberta, 3; N. D., 1; Sask., 1.): See + measurements and plates 31-33; weight, 3.1 (2.8-3.5) grams; + basilar length, 42.3 (40.0-43.7); zygomatic breadth more or less + (approximately equal to) than distance between condylar foramen + and M1 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 I3) 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 lighter 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 lines +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 (= 121 + 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 cicognanii_. + +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 indicative 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, in Montana] itself, which, +however, is even darker than the two specimens taken on the Canadian +side of the line and hence is referred to _oribasus_. 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 respect may show approach to _oribasus_, 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 throughout the skull. The only other +specimens relating to the Turtle Mountains that have been seen are two +male, skins without measurements 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 specimens +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 intergradation 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 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 _longicauda_. 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 _primulina_ in narrowness but has +the large teeth of _longicauda_. + +Intergradation 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 culbertsoni_ 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 International Rules of Zoölogical 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 deciding on the +subspecific application of the name _Putorius culbertsoni_ Coues, the +skull of the type must be principally relied 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 measurements. + +The ranges as now known of three subspecies of _Mustela frenata_ +approach near to Fort Laramie. These are _M. f. longicauda_, _M. f. +alleni_, and _M. f. 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 diagnostic, +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 +_culbertsoni_. 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 example, at Lay, +Colorado, may represent a recognizable race characterized 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, 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 provinces and states and further by districts or counties from + north to south except as otherwise indicated. Unless otherwise + indicated specimens are in the collection of the United States + National Museum. + + =Alberta.= St. Albert, 1; S. Edmonton, 3; Islay, 4[77]; Battle + River, south of Camrose, 1[77]; Daysland, 1[77]; Moose Pass, 1; + Blindman River, 2 (1[75], 1[2]); Red Deer, 3 (2[2], 1[60]); + Bearberry Creek near Sundre, 1[77]; Canad. Nat. Park, N.W. + Territory, 1[60]; Red Deer River, Didsbury, 1; Canmore, 1; + Calgary, 11 (6[60], 2[1], 1[86]); Red Deer River, 3[2]; Little + Sandhill Creek, Red Deer River, 1[77]; Waterton Lake Park, 2[77]; + Sweetgrass Hills, 1[77]; Alberta, 1[14]. + + =Colorado.= _Yuma County_: Wray 4 (1[88], 3[74]). + + =Kansas.= _Rawlins County_: 7 mi. N, 3 mi. W Beardsley, 1[74]; 6 + mi. S and 2 mi. E Atwood, 1[74]; 15 mi. SE Atwood, 1[74]. _Thomas + County_: near Brewster, 2[93]; no locality more definite than + county, 2[93]. _Trego County_, 2 (1[2]). _Cloud County_: Glasco, + 1[67]. + + =Manitoba.= Portage la Prairie, 3[75]; Carberry, 2 (1[2], 1[1]); + Carman, 1[60]; Max Lake, Turtle Mt., 1[77]. + + =Montana.= _Glacier County_: St. Marys Lake, 1; Blackfoot, 1: + Blackfoot Agency, 1. _Blaine County_: 6 mi. east Chinook, 1[74]. + _Pondera County_: 1/2 mi. SE Conrad, 1[74]. _Toole County_: Shelby + Junction, 1. _Hill County_: Havre, 1. _Fergus County_: Moccasin + Mts., 5 mi. NW Hilger, 1; 7 mi. NE Hilger, 1. _Rosebud County_: + 3/4 mi. N Ingomar, 1. _County_ in question, Milk River, 2. + + =Nebraska.= _Dawes County_: Chadron, 2[35]. _Cherry County_: + Kennedy, 1; no locality more definite than county, 1. _Brown + County_: Long Pine, 1[68]. _Antelope County_: Neligh, 1[35]. + _Adams County_: Hastings, 2[2]. _Clay County_: Inland, 4[35]. + + =North Dakota= (arranged by counties from west to east). _Divide + County_: Crosby, 1. _Mountrail County_: Lostwood, 1. Little + Missouri River, 1. _Golden Valley County_: Sentinel Butte, 1. + _Billings County_: Medora, 1[60]. _McLean County_: 3 mi. W + Elbowoods, 1. _Oliver County_: Ft. Clark, 2. _Morton County_: + Mandan, 1. _Sioux County_: 3 mi. N Cannonball, 1. _Logan County_: + 6 mi. SW Napoleon, 1. _Rolette County_: Turtle Mts., 1[76]; Fish + Lake, 1. _Benson County_: Ft. Totten, 3[14]; Sully Hill Nat. Park, + 1. _Ramsey County_: Devils Lake, 2. Stump Lake or Turtle Mts., + 2[2]. _Nelson County_: Stump Lake, 1. _Grand County_: Larimore, 1. + _Walsh County_: Grafton, 11 (4[76], 3[74], 2[2]). _Stutsman + County_: Jamestown, 1. _Barnes County_: Valley City, 1. + + =Saskatchewan.= Wingard, 5; Osier, 2[75]; Simpson, 1[2]; Touchwood + Hills, 4[7]; South arm Last Mountain Lake, 1[77]; Rush Lake + (Assiniboia, N.W.T.), 2[75]. + + =South Dakota.= _Pennington County_: Rapid City, 1. + + =Wyoming.= _Goshen County_: Fort Laramie, 2. + + +=Mustela frenata oribasus= (Bangs) + +Long-tailed Weasel + +Plates 16, 17, 18, 31, 32, 33 and 40 + + _Putorius (Arctogale) longicauda oribasus_ Bangs, Proc. New England + Zoöl. 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, December 31, 1912. + + _Mustela longicauda oribasa_, Hall, Univ. California Publ. Zoöl., + 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. Zoöl.; 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 I^3 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 Chilcotin rivers south to Alta Lake, in the + Caribou and Monashee mountains, probably in the Selkirks and + Rockies, and through the Rockies of Montana into extreme northern + Wyoming. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. + 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, exclusive 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 hind 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 three males are: Total length, 49; length + 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) slightly less than shown in figure 19. + + _Color._--Upper parts, in summer, near (14 _n_) Brussels Brown, + more blackish and less reddish than tone 4 of Burnt Umber of + Oberthür and Dauthenay, pl. 304; in type near tone 4, pl. 301 of + Oberthür 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 forelegs over feet, on medial sides of hind + limbs 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 subspecies. + + _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 tympanic bulla; breadth of rostrum less than length of + tympanic bulla; anterior margin of tympanic bulla as far posterior + to foramen ovale as width of 2-1/2 to 5 upper incisors; length of + tympanic bulla not less than length of lower molar and premolar + tooth-row and shorter than rostrum. + + 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^{4}; tympanic bulla as far posterior to + foramen ovale as width of 4-1/2 upper incisors; height of tympanic + bulla less than distance from anterior margin of tympanic bulla to + foramen ovale; length of tympanic bulla less than length of + rostrum. If more than one skull were available of the female of + _oribasus_ 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 anterior +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 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 Barkerville +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 Montana 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 features, 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 Valley of Montana, has a slightly +longer hind foot than those from Florence, but a female from Hamilton, +agrees more nearly with _nevadensis_. 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, Montana, 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 +mountainous 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 collected +from along the upper reaches of Okanagan Lake, British Columbia, by +Major Allan Brooks and Mr. J. A. Munro and by a series of skulls from +Ione, 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 referable to +_oribasus_ although neither is quite typical. One has a saturated +coloration suggestive of that of _altifrontalis_ and the skull is +shorter and broader than in other specimens of _oribasus_. The female +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 measurement 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 +River, 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, Provincial Museum, British +Columbia, respectively. The characters distinguishing _longicauda_ and +_oribasus_ are not shown by white winter skins; the skull shows some +features of _longicauda_, and the reference of these 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 skulls of this +subspecies from a total of 238 weasel skulls gathered by local +trappers. The others were _Mustela 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[7]; 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[85]; Wistaria, 3[85]; Four Mile + Creek, 4 mi. NE Quesnel, 1[21]; Isaacs Lake, 3200 ft., 1[74]; + Barkerville region, 1[74]; Clear River, 4800 ft., 1[74]; Chezacut, + 1[31]; Lillooet 3 (2[77], 1[85]); Alta Lake, 1[31]; source of + Kettle River, 7500 ft., 1[75]; E side Beaverfoot Range, 4000 to + 4500 ft. between Fraser Creek and 6 mi. SE of Fraser Creek, 1[74]; + Cranbrook, 1[86]; head of Cross River, 10 mi. below Assiniboine + Pass, 1[7]; camp east of "Kootanie," 1[7]; camp east of Kootanie + River, 1[7]. + + =Alberta.= Thoral Creek, 7000 ft., 50 mi. NE Jasper, 1[2]. + + =Montana.= _Glacier? County_: Chief Mt. Lake (= Waterton Lake), 1. + _Flathead County_: Columbia Falls, 1. _Chouteau? County_: Highwood + Mts., 1. _Fergus? County_: Big Snowy Mts., 1. _Wheatland County_: + Harlowton, 1[74]. _Ravalli County_: Florence, 2; Hamilton, 1[56]; + Darby, 1[56]; Carlos [= Charlos] Heights, 2[74]; Tin Cup District, + 2[74]; no locality more definite than county, 2[74]. _Beaverhead + County_: Donovan, 1. _Madison County_: Sheridan, 1[74]. _Gallatin + County_: Ranch 7-11, Eldridge, 1[60]. _Stillwater County_: Tacy, + 2[76]. _County_ in question: Gallatin Valley, 1; Yellowstone Park, + 1[75]. + + =Wyoming.= Glen Creek, Mammoth Hot Springs, 1. _Park County_: Four + Bears, 1[2]. + + +=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._--Differs from _M. f. + longicauda_ in smaller size, adult males having a total length of + less than 400, hind foot less than 45, 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 _l_) 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. M1 #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. + + _Skull 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 described 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 postpalatal 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 incisors; 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 + lighter than the average for the two males. + + Comparison with _M. f. longicauda_ and _M. f. nevadensis_ reveals + that the tympanic bullae average more nearly flat and that 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 possibly +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. +longicauda_ 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. +Specimens 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 +available from Chadron, Nebraska, and here referred to as _longicauda_, +the female, M1 #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 County_: Sundance, 1[91]. + + =South Dakota.= _Pennington County_: Hill City, 1[2]; 20 mi. N Elk + Mt, 1[91]. _County_ in question: Black Hills, 1[2]. _Custer + County_: Custer, 2 (1[91], 1[2]). + + =Nebraska.= _Scottsbluff County_: Mitchell, 4[35]. + + +=Mustela frenata arizonensis= (Mearns) + +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 + puncture in the right squamosal. The incisors above and below and + M^2 and P^2 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 slightly 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 western New Mexico. See figure + 29 on page 221. + + _Characters for ready recognition._--Differs 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; from _M. f. + 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 specimens 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 Oberthür and + Dauthenay, Pl. 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 + antipalmar faces of feet and wrists, 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, + averaging 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. f. 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; 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 + more or less than medial length of P4; anterior margin of tympanic + bulla as far posterior to foramen ovale as width of 3-1/2 + (including I3) 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 longer or shorter than rostrum; anterior margin of masseteric + fossa below talonid of m1. + + 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 M1 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 I3) 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 lighter than that of + the male. + +Compared with the skull of _M. f. 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 that the animals from the plateau region of Arizona are +recognizably different from those farther north in the Sierra Nevada of +California 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 proposed. + +The smaller size, especially of the skull, and the greater inflation of +the tympanic bullae are the outstanding characters which distinguish +_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 available of +this subspecies are smaller in most parts measured than any of the +scores of _nevadensis_ of similar age that 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 measurements as large as are +found in the minimum-sized _nevadensis_ from farther north. + +Intergradation with the two subspecies whose geographic ranges adjoin +that of _arizonensis_ is indicated by specimens at hand. One of these +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 subadult 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 characters 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 +_arizonensis_, the specimen is here placed with the latter. The +subadult male from Springerville, Arizona, is of larger size than the +topotypical 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 specimen . . . +is an immature female and is of unusually small size. A male obtained +by him [Mearns] near the same place is of the normal 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 measurements, is not of normal (_i. e._ average) size, but +is smaller than the average for the other populations of similarly +colored weasels referred 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. + + _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 County_: VT Park, Kaibab Plateau, 1; Deadman + Flat, 6400 ft., 1[74]; Little Spring, 1[59]; Government Prairie, + near Parks, 1[74]; _Coconino? County_: San Francisco Forest + (Yavapai Co., in 1886), 1[2]; 25 mi. SE Flagstaff, 1; Quaking Asp + Settlement, 1[2]. _Apache County_: Springerville, 1; North Fork + White River, White Mts., 8200 ft., 4[87]; head San Francisco + River, Judd Ranch, Alpine, 1[74]; 2 mi. SE Big Lake Knoll, 8700 + ft., 24 mi. S Springerville, 1[74]. _Greenlee County_: S end Blue + Range, 9000 ft., Prieto Plateau, 1; Beaver Creek, 7000 ft., 1[74]. + + =New Mexico.= _Grant County_: Mogollon 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); Merriam, N. Amer. Fauna, 5:83, July 30, 1891. + + _Putorius (Gale) brasiliensis 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 California, p. + 247, 1906. + + _Mustela arizonensis_, Grinnell and Swarth, Univ. California Publ. + Zoöl., 10:376, October 31, 1913; Whitlow and Hall, Univ. + California Publ. Zoöl., 40:247, September 30, 1933. + + _Mustela arizonensis arizonensis_, Grinnell, Univ. California Publ. + Zoöl., 40:102, September 26, 1933. + + _Mustela frenata_, Boyer, Journ. Mamm., 24:99, February 20, 1943. + + _Type._--Female, adult, skull and skin; no. 41053, Mus. Vert. + Zoöl.; 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 Washington, Idaho, Utah, and Nevada to northeastern + Arizona and northern New Mexico; westward from the eastern base of + the Rocky Mountains 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 _l_) 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 to + less than 59 per cent of the mastoid breadth; from _M. f. alleni_ + by near (14 _n_ to _l_) Brussels Brown rather than near (_h_) Clay + Color of upper parts in summer; from _M. f. neomexicana_ by near + (14 _n_ to _l_) 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 _l_) + Brussels Brown rather than near (16 _j_) 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. + xanthogenys_ by absence of light facial markings and near (14 _n_ + to _l_) Brussels Brown rather than Buckthorn Brown color of upper + parts; from _M. f. 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. f. + 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. f. + 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. f. altifrontalis_ by + lighter colored upper parts which are tones 1 to 3 of Raw Umber, + pl. 301, rather than tone 4 of Brownish Drab, pl. 302, of Oberthür + 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. f. + effera_ by larger size, males averaging 12-1/2 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 Mountains of central + Colorado are as follows: 347 (325-375); 123 (111-141); 40 + (32-43). + + 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 + California 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 _l_) Brussels + Brown or tones 1 to 3 of Raw Umber of Oberthür and Dauthenay, pl. + 301, darker on top of head from nose to line 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 (_j_) Snuff Brown or lighter 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-fourths 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 California, 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 California): 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 between condylar foramen and M1 (save + in four instances) and more than distance between anterior + palatine foramen and anterior margin of tympanic bulla (save in + two specimens); mastoid breadth more (80 per cent of specimens) 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 lacerum + 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 rostrum; + anterior margin of masseteric fossa not carried farther forward + than point directly below hypoconid of m1. + + Female (based on ten adults from Sierra Nevada of California): 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 M1 and + more (save in two specimens) 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 width of palate more or less than 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 5-1/2 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. f. longicauda_, that of both sexes +averages smaller in every measurement taken. Males of _nevadensis_, on +the average, relative to the basilar length, are narrower in the +interorbital region and across the zygomata but have the orbitonasal +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 those which appear in _longicauda_ with +increasing age. The differences mentioned 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 relative proportions. Even so, there is but little +overlap in actual size. Comparisons with the skulls of _M. f. +oribasus_, _alleni_, _neomexicana_, _arizonensis_, _inyoensis_, +_pulchra_, _xanthogenys_, _munda_, _saturata_, _oregonensis_, +_washingtoni_, _altifrontalis_, and _effera_ are made in the accounts +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 _Putorius longicauda_. + +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. Comparison 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 mentioned differences are the only ones found +among the great number of points investigated, except that as remarked +by Merriam (1896:23) the Sierran animal has the yellow of the +underparts reaching farther up under the chin, the underside of the +tail on the average 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 remarked by Merriam +(_loc. cit._), not only this but the other color features are +inconstant in addition to being slight. When the occurrence of the dark +spots near the angles of the mouth are tabulated, 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 indicated 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 specimens of +_nevadensis_ from most other places in its range. This tendency 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 to the basilar +length, narrowness of the rostrum, interorbital region, zygomatic +expanse and the shallowness through the region of the postorbital +processes place it with _nevadensis_. Of two other specimens 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 +considered except size of the molar teeth which are as large as in +_longicauda_ 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: +[M], 408, 155, 44; [F], 361, 134, 40. The two other specimens examined +from this general locality, a young female, no. 2711, Mus. Vert. Zoöl., +from Fort Bridger, and a subadult 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 Wyoming, might provide grounds for +recognizing as a different subspecies the animals in this large area +comprising parts of Colorado and Wyoming from which so few specimens +now are available. Possibly the name _Putorius culbertsoni_ Coues would +apply. Decision on that point will require adequate material from the +type locality, Fort Laramie. See discussion of this name under _M. f. +longicauda_. + +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 _longicauda_ from the Great Plains, that this +population from southeastern Idaho is nearest to _longicauda_ in linear +measurements of the orbitonasal length, mastoid breadth, length of +tympanic bullae, and as expressed in percentage of the basilar length, +length of tooth-row, breadth of rostrum, 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 between +_nevadensis_ and _longicauda_. Similarly, the specimens from +Scottsbluff County, Nebraska, here referred to _M. f. 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 _oribasus_ and _nevadensis_, +however, there is no lack of material showing intergradation. As set +forth in the account of _oribasus_, 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 _nevadensis_ on the basis of +cranial characters. Indeed, some show no approach to _washingtoni_ in +cranial characters. As might be expected on geographic grounds, the +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 the tympanic bullae, and relatively greater length +of the preorbital part of the skull. However, the greater width of the +light color of the underparts and relatively great breadth across the +mastoid processes and zygomatic arches are points of agreement with +_nevadensis_. Similarly, 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. However, 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 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 intergradation with _effera_ is shown by specimens from +southern Oregon. The change from small _effera_ to the larger +_nevadensis_ 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 different from one another than are other kinds which +definitely are known to intergrade leave no doubt but that material +from the intervening localities would show complete intergradation. + +Intergradation between _nevadensis_ and _munda_ 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 +that specimens from suitable localities will show actual +intergradation. That intergradation occurs directly with the bridled +weasel of the interior valleys of California, _M. f. xanthogenys_, is +shown by specimens from along the west-facing flank of the southern +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 referred 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 Mineral King, 7850 feet. Of the specimens from 7850 feet, the adult +male has no light 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, 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 southwestward especially _latirostra_. One of the +five juveniles is lighter colored over all of the upper parts than +_nevadensis_ and is suggestive of _xanthogenys_ 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 specimens happen to be available from the region +in which intergradation 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 subspecies of _Mustela 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 malformed 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 National Museum. + + =Arizona.= _Apache County_: 15 mi. E Luka Chu Kai Navajo School, + 8000 ft., 2. + + =British Columbia.= Monte Cr., 20 mi. E Kamloops, 1[21]; Sicamous, + 2; Okanagan, 18 (7[2], 6[85], 1[75], 1[86]); Monashee Pass, 1[31]; + Swan Lake, near Okanagan Landing, 1[22]; Okanagan Landing, 11 + (2[74], 3[31], 3[86], 3[22]); Vernon, 1[74]; Hope-Princeton + Summit, 5600 ft., 1[77]; Hope, 1[20]; Similkameen, 1[77]; + Osoyoos-Bridesville Summit, 1[77]; Anarchist Mt., Osoyoos, 1[31]; + Myer's Creek, 1[77]; Rossland, Mt. Glory, 7000 ft, 1[77]; Cascade, + 1[77]; Nelson, 1. + + =California.= _Siskiyou County_: Hornbrook, 1; Tule Lake Refuge, + 5[74]; Upper Mud Creek, 6700 ft., Mt. Shasta, 3; Mt. Shasta, 1. + _Modoc County_: Goose Lake, 1[20]; Joseph Creek, 1[74]; 5280 ft., + Parker Creek, near Alturas, 1[74]; Warner Mts., near Alturas, + 1[8]; 5 mi. NW Eagle Peak, 7000 ft., 2[74]; Shields Creek, 5000 + ft., 1[74]; Jess Valley, 1[8]. _Shasta County_: Cassel, 1. _Lassen + County_: 3 mi. W Eagle Lake, 5800 ft., 1[74]; 4 mi. S Eagle Lake, + 6000 ft., 2[74]; Mill Creek, 5000 ft., S base Mt. Lassen, 1; 6 mi. + SW Calneva, 1. _Tehama County_: Dale's, 600 ft., on Paines Creek, + 1[74]. _Plumas County_: Kelly's, 2 mi. S Willow Lake, 5200 ft., + 3[74]; Quincy, 4[68]; Beckwith, Sierra Valley, 1. _Butte County_: + Jonesville, 1[74]. _Sierra_ _County_: Little Truckee River, 6500 + ft., 3 mi. N Independence Lake, 2[42]. _Nevada County_: + Independence Lake, 1[74]. _Placer County_: Donner, 3; 2 mi. W Soda + Springs Station, 6500 ft., 1[74]; Blue Canyon, 5000 ft., 2 + (1[74]); 4 mi. S Tahoe City, 1[74]. _Eldorado County_: 5 mi. S + Tallac, 6300 ft., 1; Gilmore Lake, Mt. Tallac, 2[74]; Mt. Tallac, + 1[68]; Phillips, 1[59]. _Alpine County_: 8000 ft., Hope Valley, 1; + 8000 ft., Silver Creek, 1. _Tuolumne County_: Strawberry, 5200 + ft., 1[74]; 9300 ft., Ten Lakes, Yosemite Park, 1[74]; Tuolumne + Meadows, 8600 ft., Yosemite Park, 1[74]; Tuolumne Meadows (Soda + Springs), 1; Tuolumne Meadows, 8500 ft., Yosemite Park, 1[74]; + Sequoia, 1. _Mariposa County_: Chinquapin, 6256 ft., 2[74]; Merced + Grove Big Trees, 5400 ft., 1[74]; Wawona, 1; no locality more + definite than county, 1. _Madera County_: Bass Lake, 1[74]. _Mono + County_: Tioga Crest, near Tioga Pass, 4[74]; Warren Creek, 1[74]; + Tioga Lake, 1[74]; Ellery Lake, 9600 ft., 1[74]; Mono Lake P. O., + Mono Lake, 1[74]; Walker Lake, 8000 ft., 2[74]; Pine City, 1; + Mammoth, 13 (12[59], 1[14]); 10300 ft., near Big Prospector + Meadow, White Mts., 2[74]. _Inyo County_: Little Onion Valley, + 7500 ft., 1[74]; N Fork Bishop Cr., 10500 ft., 1[74]; S fork + Bishop Cr., Andrews Camp, 8000 ft., 1[74]; South Lake, S Fk. + Bishop Cr., 9750 ft., 1[74]; Lamarck Cr., 9900 ft., 15 mi. SW + Bishop, 1[74]. _Fresno County_: Hume, 1. _Tulare County_: Mt. + Whitney, 2; Whitney Meadow, 9800 ft., 1[74]; Monache Meadow, 8000 + ft., 3[74]; E fork Kaweah River, 9800 ft., 1; 1/2 mi. S Mineral + King, 7850 ft., 7[52]; Quaking Aspen Meadow, 7500 ft., 1[52]. + + =Colorado.= _Moffat County_: Lay, 1[19]. _Routt County_: Steamboat + Springs, 2 (1[19]); no locality more definite than county, 1[57]. + _Jackson County_: Higho, North Park, 8400 ft., 1; Buffalo or + Illinois Creek, "near Rand," 6[74]. _Washington County_: 6 mi. NE + Hillrose, 1[74]. _Larimer County_: Estes Park, 2 (1[2], 1[7]); + Pinewood, 1; Loveland, 2 (1[57]); no locality more definite than + county, 1[7]. _Rio Blanco County_: Compass Creek, 9000 ft., 1[2]; + White River, 6200 ft., 1[21]; Piceance Creek, 6200 ft., 1[2]; Dry + Fork, 6200-6600 ft., 4[2]; Meeker, 1; Marvine, 1[74]. _Grand + County_: Crembling [= Kremmling?], 1[50]; Middle Park, 1[57]. + _Boulder County_: Foot Mt. Meeker, 8700 ft., 1[2]; Silver Lake + Mine, 1[60]; Boulder, 1[60]; Dixie Lake, 2 (1[2], 1[57]); Caribou, + 1[2]; no locality more definite than county, 1. _Clear Creek + County_?: Grays Peak, 1[93]. _Jefferson County_: 7000 ft., Mt. + Parks, 1[57]; 6 mi. W Denver, 1[57]. _Adams County_: Barr, 1[2]; + near East Lake, 2[57]. _Denver County_: Denver, 2 (1[2], 1[74]). + _Arapahoe County_: Littleton, 1[19]. _Summit County_: + Breckenridge, 1[57]. _Eagle County_: Eagle, 9500 ft., 1[104]. + _Park County_: Jefferson, 4 (1[2]); 12800 ft., Mt. Bross, 1[57]. + _Mesa County_: Tunnel, 1. _Montrose County_: near Crawford, Clear + Fork of Smiths Fork, 1[19]; Coventry, 3 (1[19]); Naturita, 1; + Paradox, 1[94]; West Paradox Valley, 1[57]. _Pitkin County_: + Placita, 2[26]. _Gunnison County_: Marble, 1[26], Crested Butte, + 2[19]; Deckers Ranch, Crested Butte, 2[19]; Sapinero, 7245 ft., + 1[19]. _Chaffee County_: Buena Vista, 1[76]; Hancock, 1[16]; + Salida, 5[19]. _Teller County_: Glencore, Pikes Peak, 1[76]. _El + Paso County_: Monument, 1[76]; Seven Lakes, 1[19]; Lake Moraine, + 10250 ft., 1[19]; Colorado Springs, 6000 ft., 1[19]; 5 mi. E Sand + Creek, Colorado Springs, 1[19]; no locality more definite than + county, 1[50]. _Saguache County_: Villa Grove, 1[19]; Pierce + Place, Cochetopa Nat. Forest, 1; Houselog Creek, Cochetopa Nat. + Forest, 1; P. Tevebaugh's Ranch, near Cochetopa Pass, 1; P. + Tevebaugh's Ranch, 9 mi. S Cochetopa Pass, 1. _Rio Grande County_: + between Monte Vista and Del Norte, 1[88]. _Archuleta County_: + Upper Navajo River, 2[57]; Navajo River, 5 (4[57], 1[2]); Chromo, + 2[57]. _Conejos County_: Osier, 3[57]. _Montezuma County_: Ure + Peak, 1[57]. _County_ in question: Del Norte Peak, 1[76]; no + locality more definite than state, 4[75]. + + =Idaho.= _Latah County_: Cedar Mt., 4000 ft., 12 mi. NE Moscow, + 1[55]; Moscow and 1/2 mi. W, 2[97]. _Idaho County_: Lochsa River + (= Locksaw Fork), 1; between Selway Riv., and S Fork Clearwater + Riv., 8[74]; Selway Divide, 8[74]; Pilot Creek, 2[74]; Newsome + Cr., 1[74]. _Lemhi County_: Salmon River Mts., (now Lemhi Mts.), + 8000 ft., 5; Leadore, 3. _Adams County_: summit Smith Mt., 7500 + ft., 1[41]. _Washington County_: Midvale, 2. _Custer County_: + Pahsimeroi Mts., 1; Double Springs, 16 mi. NE Dickey, 1[74]; + Mackay?, 1; Stanley Lake, 1. _Payette County_: 2 mi. S Payette, + 1[74]. _Fremont County_: 17 mi. E, 4 mi. N Ashton, 6275 ft., + 2[74]. _Teton County_: 3 mi. S Victor, 1[74]. _Jefferson County_: + 20 mi. W Camas, 1. _Blaine County_: Sawtooth City, 1; Ketchum, 5 + (3[50], 2[75]). _Canyon County_: Nampa 3. _Clark County_: Dry + Creek, Targhee Nat. Forest, 1[2]; Birch Creek, 2. _County_ in + question: North fork of Teton River, 1. _Bingham County_: Shelley, + 1; Alridge, 2; Springfield, 1. _Lincoln County_: Shoshone, 1. + _Minidoka County_: 1/4 mi. E Heyburn Bridge, 1[74]. _Power + County_: 4 mi. NW American Falls, 1[74]. _Bannock County_: 3 mi. N + Schutt's Mine, Ross Creek, 1[74]; 3 mi. N Pocatello, 1[74]; near + (within 10 miles of) Pocatello, 1[74]; 3 mi. S Pocatello, 1[74]; 1 + mi. E Portneuf, 1[74]; 2 mi. up Mink Creek, 2 (1[74], 1[41]); + Inkom, 2; Swan Lake, 1. _Owyhee County_: 5 mi. SE Riddle, 1; Three + Creek, 2. _Cassia County_: Elba, 1[52]. _Bear Lake County_: + Geneva, 6171 ft., 1[74]; Montpelier, 1; Paris, 6000 ft., 1[6]; + Pegram, 2. + + =Nevada.= _Humboldt County_: Alder Creek, 7000 ft., Pine Forest + Mts., 1[74]; head of Big Creek, 8000 ft., Pine Forest Mts., 1[74]; + Cottonwood Range, 1; Calico Mt., Little Owyhee R., 1; Mahogany, + Little Owyhee R., 2; Sulphur, 1. _Pershing County_: Lovelocks, 1. + _Elko County_: Mountain City, 3; Three Lakes, Ruby Mts., 1[41]. + _Washoe County_: Pyramid Lake, 1; 3 mi. E Reno, 1[74]; Incline + Creek, 7100 ft., 1[74]; 2-1/2 mi. S Incline, 6250 ft., 1[74]; E + side Marlette Lake, 8000 ft., 1[74]; Marlette Lake, 8000 ft., + 1[74]. _Ormsby County_: 1/2 mi. S Marlette Lake, 8150 ft., 1[74]. + _Churchill County_: 4 mi. W Fallon, 1[74]; 3 mi. W Fallon, 1[74]; + 2 mi. W Fallon, 1[74]; Fallon, 3970 ft., 1[74]. 5 mi. S Fallon, + 4000 ft., 1[74]; 8 mi. S and 3 mi. E Fallon, 1[74]. _Douglas + County_: Mt. Siegel, 1[60]. _Mineral County_: Lapon Cañon, 8900 + ft., Mt. Grant, 1. _Nye County_: Arc Dome, 1; 10700 ft., 1/2 mi. + SW Jefferson Peak, Toquima Range, 1[74]. _White Pine County_: 3 + mi. E Baker, 1[74]; Baker Creek, 6600 ft., 4[74]; Baker Creek, + 8400 to 8450 ft., 4[74]; Gleason Creek, 7500 ft., 1[74]. + _Esmeralda County_: Arlemont, 4850 ft., Fish Lake Valley, 1[74]. + _Lincoln County_: 3 mi. S Crystal Spring, 3900 ft., Pahranagat + Valley, 1[27]. + + =New Mexico.= _Taos County_: 2 mi. N Twining, 10500 ft., 1; Taos, + 2. _Santa Fe County_: 11600 ft., Pecos Baldy, 1. _San Miguel + County_: 8000 ft., above Willis, Pecos River, Forest Reserve, + 2[75]; Ribera, 1. + + =Oregon= (by counties from west to east). _Jackson County_: + Rustler Peak, Crater Nat. Forest, 1[46]; Siskiyou (probably south + of), 2. _Klamath County_: 20 mi. W Crescent, 1[101]; Anna Creek, + Mt. Mazama, 2; S Boundary Crater Lake Nat. Park, 1[74]; Fort + Klamath, 15; Upper Klamath Lake, 2[4]; Klamath Falls, 1[75]. _Lake + County_: Dog Lake Ranger Station, 30 mi. SW Lakeview, 1. _Harney + County_: Camp Harney, 2[75]; Burns, 2 (1[101]); 20 mi. S Burns, + 1[46]; Narrows, 1[59]; Voltage, 1; Shirk P. O., 2; Keiger Gorge, + Steen Mts., 4. _Malheur County_: Riverside, 1; 2 mi. NW Riverside, + 2; Barren Valley, Cord, 1; Cedar Mts., 2; Cow Creek Lake, 1; + Jordan Valley, 1. _County_ in question: Sageview, 1. + + =Utah.= _Cache County_: Logan, 1[74]. _Rich County_: 8000 ft., + near Laketown, 1. _Boxelder County_: Willard, 1[103]. _Salt Lake + County_: Salt Lake City, 1[74]; Barclay, 6500 ft., Wasatch Mts., + 1; Mill Creek, 1[103]. _Utah County_: Provo Bench, 2[6]; Aspen + Grove, Mt. Timpanogos, 1[6]; Payson, 1[6]. _Juab County_: between + Santaquin and Starr, 1[103]. _Uinta County_: Dry Fork Canyon, 20 + mi. NW Vernal, 1[9]. _Carbon County_: Sunnyside, 1[44]; Range + Creek, 1[44]. _Millard County_: Deseret, 1[74]. _Sevier? County_: + Fish Lake Plateau, 1. _Grand County_: Warner Ranger Station, La + Sal Mts., 1[6]. _Beaver County_: Britts Meadows, 11000 ft., Beaver + Range, 1[2]; Britts Meadows, Beaver Range, 1; Puffer Lake, 1[44]. + _Garfield County_: Boulder, 2[6]. _Washington County_: Pine + Valley, 1[44]; St. George, 1. _San Juan County_: Geyser Pass, La + Sal Mts., 2[6]. _County_ in question: Salt Lake, 2; Wasatch Mts., + 1; La Sal Mts., 11000 ft., 1. + + =Washington.= _Okanogan County_: Bald Mt., 6800 ft., 1; Bauerman + Ridge, 6800 ft., Tungsten Mine, 1; Hart Pass, Methow River Trail, + 1[46]; Conconully, 2 (1[51], 1[49]); 5 mi. NW Loomis, 1; Molson, + 3800 ft., 1; Tunk Mt., 3500 ft., 1. _Whatcom County_: Barron, 5000 + ft., 1. _Stevens County_: Colville, 1; Orin, 11[51]. _Pend Oreille + County_: Ione, 6[51]. _Chelan County_: Chelan Mts., 1[2]; Lake + Chelan, 1[46]; Manson, 1; Entiat River, 1680 ft., 20 mi. from + mouth, 7; Dryden, 2[49]; Wenatchee, 1. _Kittitas County_: Easton, + 2 (1[51]); Ellensburg, 1[51]; 4 mi. E Ellensburg, 1[51]. _Grant + County_: Neppel, 1[51]. _Lincoln County_: Sprague, 1. _Spokane + County_: Spokane, 1[94]; Cheney 2[89]. _Whitman County_: Pullman, + 11 (6[55], 1[68], 1[10]); 6 mi. S Pullman, 1. _Garfield County_: + Snake River, 1. _Yakima County_: Yakima, 1[74]; 1 mi. W Moxee, + 1[74]. + + =Wyoming.= NW Wyoming, 1[75]. _Yellowstone National Park_: Lamar + River, 1; Yellowstone Lake, 1. _Park County_: Greybull River, + 1[80]. _Teton County_: Crystal Creek, 2; Jackson, 1; Whetstone + Creek, 2[76]. _Johnson County_: Buffalo, 4 (2[93]). _Fremont + County_: Continental Divide, 20 mi. NW Dubois, 1[75]. _Sublette + County_: Bronx, 1[75]. _Carbon County_: Medicine Bow Mts., 1[75]; + 15 mi. SE Parco, 1[74]. _Albany County_: Garrett, 1; 12 mi. W + Laramie, 1[74]; 7 mi. W Laramie, 2[74]; 5 mi. W Laramie, 4[74]; + "near" Laramie, 1[74]; 3 mi. SW Laramie, 1[74]; 12 mi. S Laramie, + 1[74]. _Uinta County_: Fort Bridger, 6800 ft., 1[74]; Lonetree, 1; + Bridger Pass, 1. _County_ in question: Laramie River, 2. No + locality more definite than state, 1. + + +=Mustela frenata effera= Hall + +Long-tailed Weasel + +Plates 19, 20 and 21 + + _Mustela frenata effera_ Hall, Carnegie Instit. Washington Publ. + 473:93, November 20, 1936. + + _Mustela arizonensis_, Dice, Journ. Mamm., 1:12, November 28, 1919. + + _Type._--Male, adult, skull and skin; no. 33637, Amer. Mus. Nat. + Hist.; Ironside, 4000 ft., Malheur County, Oregon; September 8, + 1912; obtained by H. E. Anthony; original no. 267. + + The skull (plates 19-21) is complete and unbroken. The teeth all + are present and entire. The skin, in summer pelage, is well made. + + _Range._--Upper Sonoran to Arctic Alpine life-zones of northern + two-thirds of Oregon east of the Cascades, and southeastern + Washington, south of the Snake River. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. + nevadensis_ in small size, males averaging 12-1/2 per cent smaller + in external measurements, 8 per cent smaller in linear + measurements of skull, and 22 per cent in weight of skull, total + length averaging 360 rather than 400, condylobasal length + averaging 40.5 rather than 43.6; from _M. f. oregonensis_ in + absence of frontonasal white patch, presence of light color of + underparts on ventral face of tail and smaller skull with basilar + length averaging less than 41.7 in males; from _M. f. washingtoni_ + in presence of light color of underparts on ventral face of tail, + in male skull by linear measurements averaging 7 (5-12) per cent + shorter and relative to basilar length shorter in preorbital + region and broader across mastoid processes and zygomatic arches. + + _Description._--_Size._--Male: Eight (6 adult and 2 subadult) + males from northeastern Oregon yield average and extreme + measurements as follows: Total length, 360 (340-378); length of + tail, 129 (122-136); length of hind foot, 42 (40-44). Tail + averages 56 (52-59) per cent as long as head and body. Length of + hind foot more or less than (about same as) basal length. + + Female: No. 212423 from Vale, and no. 566 V. B. Scheffer, from 15 + mi. E Ukiah, measure, respectively: Total length, 312, 306; length + of tail, 113, 114; length of hind foot, 35, 35. Tail averages 57 + per cent as long as head and body. + + Differences in external measurements between the one adult female + and the average of the males are: Total length, 51; length of + tail, 16; length of hind foot, 7. + + _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 stage 4 of figure 19. + + _Color._--Upper parts, in summer, near (14 _n_ to _l_) Brussels + Brown or tones 1 to 3 of Raw Umber of Oberthür and Dauthenay, pl. + 301, darker on top of head from nose to, or slightly behind, line + connecting posterior margins of ears. Chin and usually all of + upper lips white. Remainder of underparts Buff-Yellow to Straw + Yellow. In winter all white except tip of tail or upper parts near + (_j_) Snuff Brown or lighter than Brussels Brown with a smoked + effect, with 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 toes and wrists, on + medial sides of hind legs to ankles over antiplantar faces of + toes, distomedial third of tarsus and usually over proximal fourth + to three-fourths of ventral side of tail. Least width of color of + underparts averaging, in 15 males, 53 (36-69) per cent of greatest + width of color of upper parts. Black tip of tail averaging 47 + (38-67) mm. long. Thus averaging longer than hind foot, and 36 per + cent of length of tail-vertebrae. + + _Skull and teeth._--Male (based on 6 adults from northeastern + Oregon): See measurements and plates 19-21. As described in + _Mustela frenata nevadensis_ except that: Weight, 2.9 (2.5-3.4) + grams; basilar length, 40.5 (39.3-41.8). + + Female (based on no. 212423, adult from Vale): In so far as parts + of the broken skull permit a person to judge, the skull is as + described in _M. f. nevadensis_ except that: Smaller; lighter; + postorbital breadth more than width of basioccipital measured from + medial margin of one foramen lacerum posterior to its opposite. + +As compared with the skull of _M. f. nevadensis_ that of _effera_ +seems, on the average, to have the preorbital part relatively smaller. +Otherwise, the skull is a miniature of the skull of _nevadensis_, +averaging about eight per cent smaller in linear measurements and +weighs twenty-two per cent less. Comparisons of the skull with those of +_M. f. washingtoni_ and _M. f. oregonensis_ are made in accounts of +those subspecies. + +_Remarks._--This geographic race has long borne the name of _Mustela +arizonensis_ (Mearns). Small size differentiates _effera_ from +_nevadensis_ and specimens have been allocated to one or the other +subspecies on the basis of size, or average size when several +individuals are available from one locality. Complete intergradation +with each adjoining subspecies is indicated by numerous specimens, more +of which are assigned to these adjoining subspecies than to _effera_ +itself. + +The minimum of size in _M. f. effera_ is found in the Blue Mountain +region of northeastern Oregon. Specimens from the area intervening +between these mountains and the Cascades average larger but are nearer +the mean of typical _effera_ than they are to the means of +_washingtoni_, _oregonensis_ or _nevadensis_. + +Two males, nos. 204883, adult, and 204884, young, from Sisters, Oregon, +near the eastern base of the Cascades, show approach structurally to +_M. f. washingtoni_ as it is represented at the nearby locality, +Permilia Lake, at the west base of Mount Jefferson. Everything +considered, however, the two specimens from Sisters are nearer to +_effera_. A male from Condon, Oregon, shows approach to the Cascade +race in slightly increased size. + +No perfect skulls of adult females are available from the part of +northwestern Oregon in which _effera_ reaches its typical state of +development as judged by the small size of the skull of the adult male. +Skulls of adult females are available, however, from more nearly +marginal localities. These, though smaller than in _nevadensis_, show +relatively less difference in size when compared with _nevadensis_ than +do skulls of males. Even so the females at these marginal localities +are smaller than those of _nevadensis_ of comparable age and adequate +material of adult female _effera_ from the region where the males +attain their extreme of small size probably will show about the same +relative difference in size between _nevadensis_ and _effera_ as is +known to exist between the adult males of these two subspecies. The +small size of a subadult female, no 74631, U. S. Nat. Mus., from +Asotin, Washington, constitutes partial basis for this opinion. + +Of 14 adults examined none showed malformation of the frontal sinuses +due to infestation by parasites. + + _Specimens examined._--Total number, 53, arranged within each + state by counties from north to south. Unless otherwise indicated + specimens are in the collection of the United States National + Museum. + + =Oregon.= _Wasco County_: 4 mi. S The Dalles, 1[74]; Wapinita, 1; + Antelope, 2; 7 mi. E Antelope, 5. _Gilliam County_: Condon, 1[46]. + _Morrow County_: 10 mi. S Hardman, 1. _Umatilla County_: Umatilla, + 2; 15 mi. E Ukiah, 4000 ft., 1[49]. _Union County_: Elgin, 1; 20 + mi. E Lehman, 1[46]. _Wallowa County_: Horse Creek, 15 mi. N + Paradise, 1; Enterprise, 1[46]; Wallowa Lake, 1[46]; Wallowa + Mts., 8300 ft., 1. _Baker County_: Haines, 1[49]; Anthony, 3[2]; + Bourne, 2. _Grant County_: Long Creek, 1[46]; Canyon Creek, 1[46]; + Strawberry Mts., 2; Silvies, 1[14]. _Crook County_: Prineville, 4. + _Deschutes County_: Sisters, 2; Bend, 1. _Lake County_: 3 mi. W + Stauffer, 1; Fort Rock, 1[46]. _Harney County_: 25 mi. NW Burns, + 1. _Malheur County_: 4000 ft., Ironside, 2[2]; 1-1/2 mi. S Vale, + 2. + + =Washington.= _Walla Walla County_: Prescott, 4 (2[76], 1[60], + 1[74]); Ft. Walla Walla, 2 (1[75]); Wallula, 1[76]. _Asotin + County_: Asotin, 1. + + +=Mustela frenata washingtoni= (Merriam) + +Long-tailed Weasel + +Plates 19, 20, 21, 34, 35 and 36 + + _Putorius washingtoni_ Merriam, N. Amer. Fauna, 11:18, pl. 4, figs. + 3, 3a, 4, 4a, June 30, 1896. + + _Mustela washingtoni_, Miller, U. S. Nat. Mus. Bull., 79:98, + December 31, 1912. + + _Mustela frenata washingtoni_, Hall, Carnegie Instit. Washington + Publ. 473:106, November 20, 1936. + + _Type._--Male, adult, skin and skull; no. 76322, U. S. Nat. Mus., + Biol. Surv. Coll.; Trout Lake, Mt. Adams, Klickitat (?) County, + Washington; December 15, 1895; obtained by D. N. Kaegi; original + no. 2. + + The skull is unbroken. The left incisors above are missing. + Otherwise the teeth are present and entire. The skin is well made, + in brown winter pelage, lacks collector's measurements, has no + bones in the feet, but by large size is judged to be a male. + + _Range._--Altitudinally from near 2000 feet at Trout Lake up to + the highest parts of the Cascade Range from Mount Jefferson, + Oregon, north to Mount Rainier, Washington; Upper Sonoran + Life-zone to Arctic Alpine Life-zone. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. + altifrontalis_ in lighter color of upper parts and underparts, + latter ranging from Buff-Yellow to Naples Yellow rather than near + (14 _a_ to 16 _c_) Ochraceous-Buff, in shallower skull in both + sexes (see measurements), in males, a longer preorbital region, + narrower skull with shorter bullae, and in females, a smaller + skull with interorbital breadth averaging less than 24 per cent of + basilar length; from _M. f. nevadensis_ in absence of light color + of underparts on ventral face of tail, in skulls of males, by + longer preorbital region and narrower skull across mastoid + processes and zygomatic arches, in skulls of females, by shorter + preorbital region, and smaller bullae (see measurements); from _M. + f. effera_ in absence of light color of underparts on ventral face + of tail, in skulls of males, by linear measurements averaging 7 + (5-12) per cent larger, and relative to basilar length, longer in + the preorbital region and narrower across mastoid processes and + zygomatic arches; from _M. f. oregonensis_ in absence of + frontonasal white patch, longer skull in males, which in + percentage of basilar length has, on the average, orbitonasal + length amounting to more than 35, mastoid breadth less than 55, + and zygomatic breadth less than 63, and in females, smaller skull + with least width of palate less than length of P4, upper + tooth-rows less than 38-1/2 per cent of basilar length, bullae + smaller, averaging less than 13.4 in length. + + _Description._--_Size._--Male: Fifteen subadult topotypes yield + average and extreme measurements as follows: Total length, 400 + (357-437); length of tail, 149 (122-171); length of hind foot, + 47.6 (42-59). Tail averages 59 per cent as long as head and body. + Length of hind foot averaging more than basal length. + Corresponding measurements of one adult and 3 young from Mount + Rainier are: 415 (405-423); 155 (145-164); 51 (50-53). + + Female: Five adult topotypes yield average and extreme + measurements as follows: Total length, 349 (330-393); length of + tail, 124 (114-133); length of hind foot, 38 (36-39). Tail + averages 55 per cent as long as head and body. Length of hind foot + averaging about same as basal length. Corresponding measurements + of two adults and 6 young from Mount Rainier are: 338 (320-360); + 121 (115-132); 36 (34-40). + + The average differences in external measurements of the two sexes, + from Mount Adams, are: Total length, 51; length of tail, 25; + length of hind foot, 9.6. Corresponding differences between the + specimens from Mount Rainier are: 77; 34; 15. + + _Externals._--Longest facial vibrissae black or brown (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 slightly less + than shown in figure 19. + + _Color._--Upper parts in summer near (14 _n_) Argus Brown or tone + 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304; one topotype + Buckthorn Brown or tone 3 to 4 of Snuff Brown of Oberthür and + Dauthenay, pl. 303. Dark spot at each angle of mouth present or + absent, and when present, often fused with color of upper parts, + which rarely covers lower lips. Chin, and usually lower lips, + white. Remainder of underparts Buff-Yellow to Naples Yellow. In + winter, all white except tip of tail which is at all times black, + or upper parts near (14) Brussels Brown to near (_j_) Snuff Brown + with smoked effect and underparts white, rarely with trace of + yellowish. Color of underparts extends distally on posterior sides + of forelegs over toes onto antipalmar faces of feet and usually + all of wrists, on medial sides of hind legs anywhere from knee to + tips of toes. Least width of color of underparts averaging in ten + topotypes, 24 (10-37) per cent of greatest width of color of upper + parts. Black tip of tail in same series averaging 55 (45-60) mm. + long, thus longer than hind foot and averaging 37 per cent of + length of tail-vertebrae. + + The color of the underparts is not so narrow in the specimens from + Mount Rainier and it is believed that the slender bodies used in + stuffing the topotypes has accentuated in them the appearance of + narrowness of the light-colored underparts. + + _Skull and teeth._--Male (based on 22 adult topotypes): See + measurements and plates 19-21; weight, 3.5 (2.8-4.7) grams; + basilar length, 43.7 (40.0-47.7); zygomatic breadth more or less + than distance between condylar foramen and M1 or than between + anterior palatine foramen and anterior margin of tympanic bulla; + mastoid breadth more or less than postpalatal length; postorbital + breadth 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 than distance between foramen opticum and anterior margin of + tympanic bullae; breadth of rostrum less (except in no. 82180) + than length of tympanic bulla; least width of palate more (except + in no. 81954) than length of P4; anterior margin of tympanic bulla + as far posterior to foramen ovale as width of 2 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 (except in two instances) than length of lower molar and + premolar tooth-row and shorter (except in two instances) than + rostrum; anterior margin of masseteric fossa below m2. + + Female (based on 11 ad. topotypes): See measurements and plates + 34-36; weight, 2.0 (1.8-2.2) grams; basilar length, 37.6 + (37.0-38.9); zygomatic breadth less (except in no. 70945) than + distance between condylar foramen and M1 or than between anterior + palatine foramen and anterior margin of tympanic bulla; + 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; least width of palate + less (except in one specimen) than greatest length of P4; tympanic + bulla as far posterior to foramen ovale as width of 3-1/2 to 5-1/2 + 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. + +Compared with _M. f. nevadensis_, the skull of the male of +_washingtoni_ averages more slender, as shown by the mastoid and +zygomatic breadths and has the preorbital part longer, on the average, +as shown by the greater ratio (to the basilar length) of the length of +the tooth-rows and orbitonasal length. Also, on the average, the +postorbital constriction is longer than in _nevadensis_ and the +tympanic bullae are smaller. In females, the skull is lighter, the +tooth-rows are shorter, the tympanic bullae are smaller, and the +preorbital part of the skull is shorter and narrower as shown by the +orbitonasal length and interorbital breadth. Except that the tympanic +bullae are actually, although not relatively, smaller in males of +_effera_, it differs from _washingtoni_ in the same way as does +_nevadensis_ as regards relative proportions, but, of course, the +actual difference in size is greater since _effera_ is smaller than +_nevadensis_. Comparison of the skull with that of _oregonensis_ is +made in the account of that subspecies. + +_Remarks._--_M. f. washingtoni_ was described and named in 1896 by +Merriam as a distinct species. Subsequently, specimens which here are +regarded as intergrades between _altifrontalis_ and _nevadensis_, were +classified as _washingtoni_. + +The external measurements given for the specimens from Mount Adams are +those recorded on the labels in inches and fractions thereof. Instead +of total length there sometimes is written "tip to tip." In the series +of 19 winter-taken topotypes the hairs project beyond the end of the +caudal vertebrae for an average distance of 28 (19-40) millimeters. If +the hairs on the end of the tail were included in the measurements, 28 +millimeters should be subtracted from the averages. Probably the +measurements should stand as given, since an adult male topotype, no. +226758, U. S. Nat. Mus., taken subsequently by Walter P. Taylor +measures 405; 152; 51. + +_Mustela frenata washingtoni_ is not a strongly marked geographic race. +In many features it is intermediate between _M. f. altifrontalis_ and +_M. f. nevadensis_. This is especially true of coloration. In the +series from Mount Adams and that from Mount Rainier, some individuals +have the light color of the underparts extended down the hind legs over +the feet and over the proximal face of the ventral third of the tail as +in _nevadensis_, whereas others from the same place have the light +color of the underparts absent from the tail and extending no farther +down the hind limbs than the knees. The light color of the underparts +in the series of topotypes is so restricted that the transverse extent +at the narrowest place amounts to only 24 (10-37) per cent of the +greatest width of the color of the upper parts. This narrowness of the +color of the underparts has been likened by Merriam (1896:18) to the +condition in _Mustela frenata noveboracensis_. So it is, but it is +similar to the condition found also in the geographically adjoining _M. +f. altifrontalis_. + +Of the 37 skulls of subadults and a few adults, 11 had the frontal +sinuses malformed as a result of infestation by parasites. + + _Specimens examined._--Total number, 56, arranged within each + state by localities from north to south. Unless otherwise + indicated specimens are in the collection of the United States + National Museum. + + =Oregon.= Mt. Jefferson, Permilia Lake, 1. + + =Washington.= _Pierce County_: 5500 ft., Spray Park, Mt. Rainier, + 1; Spray Park, 1[74]; 5935 ft., Glacier Basin, Mt. Rainier, 5 + (1[10]); 5051 to 5100 ft., Owyhigh Lakes, Mt. Rainier, 7 (1[10]), + Tahoma Creek, 1[72]; Nisqually entrance, 1[72]; Longmire, 1[72]; + Mt. Rainier Nat'l Park, 2[72]. _Klickitat County_: Trout Lake, S + Base Mt. Adams, 35; 3500 ft., Gotchen Creek, Mt. Adams, 1. + + +=Mustela frenata saturata= (Merriam) + +Long-tailed Weasel + +Plates 19, 20, 21 and 30 + + _Putorius saturatus_ Merriam, N. Amer. Fauna, 11:21, June 30, 1896. + + _Mustela saturata_, Miller, U. S. Nat. Mus. Bull., 79:98, December + 31, 1912. + + _Mustela arizonensis saturata_, Grinnell, Univ. California Publ. + Zoöl., 40:102, September 26, 1933. + + _Mustela frenata saturata_, Hall, Carnegie Instit. Washington Publ. + 473:106, November 20, 1936. + + _Type._--Male, adult, skull and skin; no. 65930, U. S. Nat. Mus., + Biol. Surv. Coll.; Siskiyou, Jackson County, Oregon; June 6, 1894; + obtained by C. P. Streator; original no. 3905. + + The skull (plates 19-21, 30) lacks the middle part of each + zygomatic arch. The teeth all are present although much worn, + probably from gnawing at the trap which captured the animal. The + skin, in fresh summer pelage, is fairly well made. + + _Range._--Transition and Boreal life-zones of Siskiyou and Trinity + mountains in southern Oregon and northwestern California. See + figures 29 and 30 on pages 221 and 314. + + _Characters for ready recognition._--Differs from _M. f. + nevadensis_ in lacking light color of underparts on tail and ankle + and in greater average breadth across mastoid processes of skull + (see measurements); from _M. f. oregonensis_ in lacking white + nasofrontal spot, in having color of underparts interrupted at + ankle; from _M. f. munda_ in lacking white nasofrontal spot, in + smaller and relatively deeper skull of males and smaller skull of + the female. + + _Description._--_Size._--Male: Four adult males (the type, 1 from + Mt. Ashland and 2 from Jackson Lake) yield average and extreme + measurements as follows: Total length, 414 (402-437); length of + tail, 150 (136-160); length of hind foot, 46 (43-50). Tail + averages 57 (49-62) per cent as long as head and body. Length of + hind foot more or less than basal length. + + Female: One young from the summit of the Trinity Mountains east of + Hoopa and one nontypical adult from 5500 feet elevation on South + Fork Mountain, Humboldt County, measure respectively as follows: + Total length, 330, 325; length of tail, 115, 123; length of hind + foot, 37, 37. Tail is 53 and 61 per cent as long as head and body. + Length of hind foot less than basal length. + + Average differences in external measurements between the two + sexes, indicated by the unsatisfactory material available, are: + Total length, 86; length of tail, 31; length of hind foot, 9. + + _Externals._--Longest facial vibrissae black or dark brown and + extending beyond ear; carpal vibrissae same color as underparts + and extending as far as apical pad of fifth digit; hairiness of + foot-soles, in summer pelage, as shown in figure 19. + + _Color._--Upper parts, in summer, Brussels Brown to near (_n_) + Brussels Brown or lighter than tone 3 of Raw Umber of Oberthür and + Dauthenay, pl. 301, usually darkest on nose and forehead. Chin + white. Remainder of underparts Buff-Yellow to Warm Buff. Tip of + tail black. Winter pelage unknown. Color of underparts extends + distally on posterior sides of forelegs over toes onto antipalmar + faces of feet and sometimes wrists, on medial sides of hind legs + only to ankles, but toes sometimes with isolated white markings. + Least width of color of underparts in the type and 2 adults from + Jackson Lake averaging 35 (30-40) per cent of greatest width of + color of upper parts. Black tip of tail averaging 54 (53-55) mm. + long; thus longer than hind foot and averaging 37 per cent of + length of tail-vertebrae. + + _Skull and teeth._--Male (based on 4 adults: Type, Mt. Ashland, 1; + Jackson Lake, 2): See measurements and plates 19-21, 30. As + described in _Mustela frenata nevadensis_ except that: Weight, 3.8 + (3.5-4.3) grams; basilar length, 44.4 (42.6-45.8); zygomatic + breadth more or less than distance between condylar foramen and M1 + or than distance between anterior palatine foramen and anterior + margin of tympanic bulla; mastoid breadth more than postpalatal + length; least width of palate less than medial length of P4 + (except in one specimen). + + Female (based on one adult possibly not typical, from 5500 ft., + South Fork Mt.): See measurements. As described in _Mustela + frenata nevadensis_ except that: Weight, 2.2 grams; basilar + length, 38.1; zygomatic breadth less than distance between + condylar foramen and M1 and less 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. + +The skull of the male of _saturata_, relative to the basilar length, is +broader across the mastoids and narrower across the rostrum and +interorbital region than that of _nevadensis_. Skull not known +certainly to differ from that of _oregonensis_. Compared with the skull +of _munda_, that of the male of _saturata_ is smaller in every part +measured except depth of tympanic bullae which averages 3.6 +millimeters, rather than 3.5 as in _munda_. Also, the skull of +_saturata_ has a less-marked postorbital constriction, is less heavily +ridged, less angular, does not have the impressions of the temporal +muscles carried so far forward on the frontal bones and is relatively +much narrower across the zygomatic arches. + +_Remarks._--In 1896, Merriam named _M. f. saturata_ as a distinct +species on the basis of one specimen, taken by Clark P. Streator at +Siskiyou, Oregon, and a second specimen taken the year previously by +Allan C. Brooks at Chilliwack, British Columbia. On the basis of these +two specimens, Merriam (1896:22) ascribed to the race a range ". . . on +the Cascade and Siskiyou mountains of Oregon and Washington, reaching a +short distance into British Columbia." Since that time, this name, +_saturata_, has been employed for the dark-colored weasels, of the +coastal region of Oregon, Washington, and extreme southwestern British +Columbia, which here are arranged under the name _M. f. altifrontalis_. +_M. f. saturata_ proves to be restricted to the humid mountainous +region inland from the coast in northern California and in the Siskiyou +Mountains of southern Oregon. Its range is separated by that of _M. f. +oregonensis_ from the range of the darker-colored, deeper-skulled, _M. +f. altifrontalis_ of the humid costal region proper. + +On May 5, 1933, Mr. Clark P. Streator, informed the writer that he +remembered taking the type specimen of _Mustela frenata saturata_ +(Merriam) in the town of Siskiyou, Oregon. The exact place, he said, +was reached, at the time of his work there, by going one or two blocks +east of the depot, then through a garden into the thick woods where +there were springs and numerous burrows of the rodent, _Aplodontia_. +Two other weasels labeled as taken at Siskiyou, on September 28 and +29, 1893, by Mr. Streator, are much lighter colored than the type of +_saturata_ and have the color of the underparts extended distally on +the hind legs to the tips of the toes and in other features of +coloration are more like _nevadensis_, the subspecies to which they are +referred, than _saturata_. Probably these did not come from exactly the +same place that the type specimen of _saturata_ did. Although Mr. +Streator does not remember the taking of these particular specimens in +1893, he does remember that on this visit to Siskiyou, he walked +southward through the railroad tunnel and collected on the opposite +side of the ridge from Siskiyou. Here on more southern exposures, the +country was markedly different than in the thick forest at Siskiyou. +Probably these two specimens taken in 1893, and referred to +_nevadensis_, came from a little way south of Siskiyou and from a +different habitat and life-zone than the type specimen of _M. f. +saturata_. + +Of the 6 specimens examined, only one, the type, shows malformation of +the frontal sinuses such as result from infestation by parasites. + + _Specimens examined._--Total number, 6, as follows: + + =California.= _Siskiyou County_: Jackson Lake, 5900 ft., 2, Mus. + Vert. Zoöl. _Humboldt County_: South Fork Mt., 5500 ft., 1, Mus. + Vert. Zoöl. _County_ in question, Trinity Mts., summit east of + Hoopa, 5800 ft., 1, U. S. Nat. Mus. + + =Oregon.= _Jackson County_: Mt. Ashland, 1, Univ. Oreg.; Siskiyou, + 1, U. S. Nat. Mus. + + +=Mustela frenata altifrontalis= Hall + +Long-tailed Weasel + +Plates 1, 19, 20, 21, 34, 35 and 36 + + _Mustela frenata altifrontalis_ Hall, Carnegie Instit. Washington + Publ. 473:94, November 20, 1936. + + _Putorius (Gale) brasiliensis frenatus_, Coues, Fur-bearing + animals, p. 142, 1877 (part). + + _Putorius saturatus_ Merriam, N. Amer. Fauna, 11:21, June 30, 1896 + (part). + + _Mustela saturata_, Miller, U. S. Nat. Mus. Bull., 79:98, December + 31, 1912. + + _Type._--Male, adult, skull and skin; no. 42093, Mus. Vert. Zoöl.; + Tillamook, Tillamook County, Oregon; July 10, 1928; obtained by + Alex Walker; original no. 717. + + The skull is complete and unbroken. P3 on the left side is + missing; otherwise the teeth all are present and entire. The skin + is well made and the enlarged scrotal pouch shows the collector's + sexing of the specimen to have been correct. + + _Range._--Altitudinally from sea level up to at least 4800 feet + (Mount Baker) in the Transition Life-zone of the humid, coastal + region of Oregon, Washington and extreme southwestern British + Columbia. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. + nevadensis_ in tone 4 of Brownish Drab, pl. 302, rather than tones + 1-3, of Raw Umber, pl. 301, of Oberthür and Dauthenay of upper + parts, in near (14 _a´_ to 16 _c´_) Ochraceous-Buff rather than + Buff-Yellow to Straw Yellow of underparts, in that least width of + color of underparts amounts to less than 37 per cent of greatest + width of color of upper parts, in absence of color of underparts + on ventral side of tail and on hind leg distal to knee, and in + greater depth of skull through frontal region; from _M. f. + washingtoni_ in darker color of upper parts and underparts, latter + near (14 _a_´ to 16 _c_´) Ochraceous-Buff rather than ranging from + Buff-Yellow to Naples Yellow, in deeper skull in both sexes (see + measurements), in males a shorter preorbital region, broader skull + with longer bullae and in females a larger skull with interorbital + breadth averaging more than 24 per cent of basilar length; from + _M. f. oregonensis_ in frontonasal white patch absent, color above + darker (tone 4 of Brownish Drab, pl. 302, rather than tone 2 to 3 + of Raw Umber, pl. 301 of Oberthür and Dauthenay), light-colored + underparts narrower and not extended distally beyond knee, in + females tooth-row shorter, amounting to less than 38 per cent of + basilar length. + + _Description._--_Size._--Male: Eight adult topotypes yield average + and extreme measurements as follows: Total length, 426 (392-445); + length of tail, 160 (148-170); length of hind foot, 47 (42-53). + Tail averages 60 per cent as long as head and body. Length of hind + foot averages more than basal length. + + Female: Five adults from Tillamook and Blaine, Oregon, yield + average and extreme measurements as follows: Total length, 347 + (320-370); length of tail, 125 (114-131); length of hind foot, 38 + (35-44). Tail averages 56 per cent as long as head and body. + Length of hind foot less than basal length. + + The average differences in the external measurements are: Total + length, 79; length of tail, 35; length of hind foot, 9. + + _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 or + beyond apical pad of fifth digit; hairiness of foot-soles (in + summer pledge) slightly less than shown in figure 19. + + _Color._--Upper parts, in summer, near (_n_) Argus Brown or tone 4 + of Brownish Drab of Oberthür and Dauthenay, pl. 302. Dark spot at + each angle of mouth well developed; often fused with color of + upper parts which sometimes covers lower lips. Chin white. + Remainder of underparts near (14 _a´_ to 16 _c´_) Ochraceous-Buff. + In winter, upper parts near (14) Argus Brown with smoked effect + and Warm Buff to Naples Yellow below. 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 usually all + of wrists, on medial side of hind legs typically only to knee but + sometimes to ankle. Tips of toes of hind feet almost always marked + with color of underparts. Least width of color of underparts + averaging in a series of 14 males from Blaine, Oregon, 23 (14-36) + per cent of greatest width of color of upper parts. Black tip of + tail in 8 adult males from Blaine, Oregon, averaging 59 (47-70) + mm. long; thus longer than hind foot and averaging 37 per cent of + length of tail-vertebrae. + + _Skull and teeth._--Male (based on 9 adults from Blaine, Tillamook + Co., Oregon): See measurements and plates 19-21; weight, 4.4 + (3.3-5.3) grams; basilar length, 45.6 (42.4-47.7); zygomatic + breadth more or less (usually more) than distance between condylar + foramen and M1 or than between anterior palatine foramen and + anterior margin of tympanic bulla; mastoid breadth more or less + (usually more) than postpalatal length; postorbital breadth less + (except in some instances of malformations of frontal sinuses + which result from infestation by parasites) 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; 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 + more or less than length of P4; anterior margin of tympanic bulla + as far posterior to foramen ovale as width of 3 to 4 (including + I3) 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 + more or less than orbitonasal length; anterior margin of + masseteric fossa directly below m2. + + Female (based on 4 adults): See measurements and plates 34-36; + weight, 2.2 (2.2-2.3) grams; basilar length, 38.1 (37.8-39.7); + zygomatic breadth more or less (less in three of four specimens) + than distance between condylar foramen and M1 or than between + anterior palatine foramen and anterior margin of tympanic bulla; + relation of postorbital breadth to other measurements in doubt + because of malformation of frontal sinuses by parasites; least + width of palate not less than greatest length of P4; tympanic + bulla as far posterior to foramen ovale as width of 3-1/2 to 5-1/2 + 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. + +Compared with the skull of _M. f. washingtoni_ that of each sex of +_altifrontalis_ averages slightly larger in every measurement taken, +except measurements of teeth which are approximately the same, and is +relatively deeper through the frontal region and through the braincase +as measured at the anterior margin of the basioccipital. Skulls of +females of _altifrontalis_ have a relatively broader interorbital +region. Skulls of males of _altifrontalis_ further differ in having +relatively, as well as actually, longer tympanic bullae, relatively +lesser orbitonasal length and a greater relative breadth across the +mastoids and across zygomata. Compared with _M. f. nevadensis_, the +skull of the male of _altifrontalis_ averages slightly larger and +heavier although the skulls of females are of approximately the same +size and weight. Relative to the basilar length, the skulls of both +sexes are deeper through the braincase and narrower across the +mastoids; the rostrum is broader, especially in males; the tooth-rows +are shorter and the interorbital breadth less, especially in females. +Comparison with the skull of _oregonensis_ is made in the account of +that subspecies. + +_Remarks._--Until the present study was begun, animals of this race +have gone under the name _Mustela saturata_ (Merriam). The United +States National Museum has a juvenile taken, in 1858, by Wayne at +Astoria, O. T.; the Samuel N. Rhoads collection contained one specimen +taken in 1891, at Tacoma, Washington; one in the Bangs' collection was +taken at Chilliwack, British Columbia, in 1895, and the Field Museum +has one taken on the Olympic Peninsula in 1898. The best material is +that collected by Alex Walker, at Tillamook, Oregon. + +Intergradation with _nevadensis_ is indicated by several specimens. The +coloration of the one adult female, no. 90, Chas. R. Conner Mus., from +Swamp Creek, Washington, has the color of the underparts extended down +the hind legs over the feet, and over the proximal third of the ventral +face of the tail as in _nevadensis_ although the other two specimens +from the same place have the color pattern of _altifrontalis_. Of the +four specimens from British Columbia referred to this subspecies, only +the specimen from Chilliwack is typical as regards color pattern. The +one from Cultus Lake has the color pattern of _nevadensis_ and might be +referred to that race almost as well as to _altifrontalis_. The two +specimens from Lihumption Park are intermediate between the two races +in tone of color. Neither has the color of the underparts extended onto +the tail or continuously over the hind feet as in _nevadensis_ but each +does have the color of the underparts less restricted and of lighter +hue than in _altifrontalis_. Only one of the specimens, no. 7848 Canad. +Nat. Mus., from Lihumption Park is adult and it has a skull which +agrees with that of _altifrontalis_ rather than _nevadensis_. + +After writing the above, a good representation of the weasel population +along the eastern side of Puget Sound was made available by friends in +that area. Study of the weasels from there shows that their color is +intermediate between that of _altifrontalis_ and _nevadensis_. On the +whole, they (specimens from Bellingham, for example) resemble one +subspecies about as much as the other. In cranial characters some +specimens, in certain features, approach _nevadensis_ but most +specimens agree with _altifrontalis_ and all are more nearly like +_altifrontalis_ to which race all are referred. + +The color of these animals is to me indistinguishable from that of +_washingtoni_. The color of _washingtoni_ is merely intermediate +between that of _nevadensis_ and _altifrontalis_. Nevertheless, the +race _washingtoni_ has cranial characters (long narrow skull) which set +it off from both _altifrontalis_ and _nevadensis_. This shape of skull +is not found in the specimens from along the eastern side of Puget +Sound; these animals have skulls like that of _altifrontalis_ and when +departures from this occur they are in the direction of _nevadensis_ +and not _washingtoni_. + +The above, then, explains why specimens which are colored like those +of _washingtoni_ are not referred to that race but instead to the race +_altifrontalis_. + +Of 23 adult skulls examined, 19 have the frontal sinuses malformed as +the result of infestation by parasites. + + _Specimens examined._--Total number, 80, arranged within states by + counties from north to south. Unless otherwise indicated specimens + are in the United States National Museum. + + =British Columbia.= _Chilliwack_, 1[74], Lihumption Park, 4750 + ft., 2[77]; Cultus Lake, 1[77]. + + =Oregon.= _Clatsop County_: Old Fort Clatsop, 1[74]; Astoria, 1. + _Tillamook County_: Tillamook, 12 (7[14], 2[74], 2[2], 1[46]); + Netarts, 1[46]; Blaine, 16 (13[14], 1[93], 1[76], 1[59]). _Lane + County_: Reed, 1; Mercer, 1[46]. _Curry County_: Langlois, 1[46]. + + =Washington.= _Whatcom County_: Nooksack River, 2000 ft., 14 mi. E + Glacier, 1; Swamp Creek, 2050 ft., Nooksack River, 3[10]; Lookout, + 4800 ft., Mt. Baker, 2[10]; Bellingham, 8[25]; 5 mi. S Bellingham, + 1[49]. _Skagit County_: Rockport, 300 ft., 1. _King County_: + Bothell, 2[94]; N Seattle 1[51]; Seattle, 1[49]; Tye, 1[51], 2 mi. + E Skykomish, 1[51]; 7 mi. E Kent, 1[76]; Auburn, 3[94]. _Pierce + County_: Tacoma, 1[1]. _Clallam County_: Sequim, 1[49]; Soleduc + Riv., near [_sic._] Sappho, 1[49]; Happy Lake, 1[60]; mouth of + Boulder Creek, Elwha River, 560 ft., Olympic Mts., 1; Hume's + Ranch, 1000 ft., Elwha River, 1; Bogachiel Riv., 1[49]. _Mason + County_: Lake Cushman, 2; 4 mi. N Shelton, 1[51]. _Thurston + County_: Olympia, 2[49]; Tenino, 1[51]. _Pacific County_: 2-1/2 + mi. SE Chinook, 3[74]. + + +=Mustela frenata oregonensis= (Merriam) + +Long-tailed Weasel + +Plates 19, 20, 21, 30, 34, 35 and 36 + + _Putorius xanthogenys oregonensis_ Merriam, N. Amer. Fauna, 11:25, + June 30, 1896; Bangs, Proc. New England Zoöl. Club, 1:57, June 9, + 1899. + + _Mustela xanthogenys oregonensis_, Miller, U. S. Nat. Mus. Bull., + 79:99, December 31, 1912. + + _Mustela xanthogenys munda_, Grinnell, Univ. California Publ. + Zoöl., 40:102, September 26, 1933 (part). + + _Mustela frenata oregonensis_, Hall, Carnegie Instit. Washington + Publ. 473:107, November 20, 1936. + + _Type._--Male, subadult, skull and skin; no. 32019/43828, U. S. + Nat. Mus., Biol. Surv. Coll.; Grants Pass, Rogue River Valley, + Josephine County, Oregon; December 19, 1891; obtained by C. P. + Streator; original no. 1404. + + The skull (plates 19-21, 30) is complete and unbroken. P3 on the + left side is missing. Otherwise the teeth all are present although + worn probably as a result of gnawing at the trap which captured + the specimen. The skin, in brown, winter pelage, is fairly well + made. + + Although the label on the skin and the label in the skull vial + each give the sex of the specimen as female, and although Merriam + (1896:25) regarded the specimen as a female, the present writer + regards the specimen as a male. + + It is as large as other undoubted males and larger than any known + female of this subspecies. The labels with the skull and skin give + the locality as "Rogue River Valley, Oregon." The listing here of + the more restricted locality, Grants Pass, is made on the basis of + Merriam's (1896:25) original description of the subspecies. + + _Range._--Transition and Canadian life-zones along coast of + northern California and southern Oregon from Humboldt County, + California, north through Curry County, Oregon, thence inland, + west of the Cascades, north to the Columbia River. See figures 29 + and 30 on pages 221 and 314. + + _Characters for ready recognition._--Differs from _M. f. + altifrontalis_ in presence of frontonasal white patch, lighter + color above (tone 2 to 3 of Raw Umber, pl. 301, rather than tone 4 + of Brownish Drab, pl. 302, Oberthür and Dauthenay), wider extent + of light color of underparts which is extended distally beyond + knee, and in females, longer tooth-row which amounts to more than + 38 per cent of basilar length; from _M. f. munda_ in shorter hind + foot of males which is less than 50, and in both sexes, smaller, + less rugose skull (see measurements and plates); from _M. f. + saturata_ in presence of frontonasal white patch, in having color + of underparts extended uninterruptedly over ankle onto foot; from + _M. f. nevadensis_ in presence of frontonasal white patch, lack of + light color of underparts on ventral face of tail, and longer + skull, which relative to its length in males, is shallower through + braincase; from _M. f. effera_ in presence of frontonasal white + patch, lack of light color of underparts on ventral face of tail, + and larger skull with basilar length averaging more than 41.7 in + males; from _M. f. washingtoni_ in presence of frontonasal white + patch, shorter skull in males, which in percentage of basilar + length has, on the average, orbitonasal length amounting to less + than 35, mastoid breadth more than 55, and zygomatic breadth more + than 63; and in females larger skull with least width of palate + more than length of P4, upper tooth-rows more than 38-1/2 per cent + of basilar length, bullae larger and averaging more than 13.4 + long. + + _Description._--_Size._--Male: Five males (3 adults and 2 + subadults from Eureka, Ferndale, and Carlotta, California) yield + average and extreme measurements as follows: Total length, 392 + (347-430); length of tail, 138 (110-160); length of hind foot, 46 + (43-50). Tail averages 54 (46-61) per cent as long as head and + body. Length of hind foot more or less than basal length. The type + specimen, and an adult from Goldbeach measure, respectively, as + follows: Total length, 412, 386; length of tail, 155, 137; length + of hind foot, 44, 46. + + Female: Three adults (2 from Fortuna and 1 from Carlotta, + California) yield average and extreme measurements as follows: + Total length, 367 (360-374); length of tail, 130 (123-134); length + of hind foot, 40 (39-40). Tail averages 55 (52-57) per cent as + long as head and body. Length of hind foot less than basal length. + A subadult from Goldbeach, an adult from 13 mi. SW Grants Pass, + and an adult from Medford, measure, respectively, as follows: + Total length, 316, 344, 294; length of tail, 114, 120, 122; length + of hind foot, 36, 40, 38. + + The average differences in external measurements of the two sexes + in the vicinity of Carlotta, are: Total length, 25; length of + tail, 8; length of hind foot, 6. Corresponding differences, at + Goldbeach, are: 70, 23, 10. Probably the females at Fortuna + reflect the large size of _munda_ more than do the males at + Carlotta and the differences between the measurements of the two + sexes probably, therefore, are actually more than are indicated by + the figures above. + + _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 20. + + _Color._--Upper parts, in summer, near (16 _l_) Brussels Brown or + tone 2 of Raw Umber of Oberthür and Dauthenay, pl. 301, to + slightly darker than tone 3 of same plate. Darker on nose and top + of head, usually with frontonasal white patch but lacking white + bar in front of each ear, except in the type and 2 specimens from + Salem. Chin, lower lips, angle of mouth, and usually posterior + seventh of upper lip white. Remainder of underparts Pale + Orange-Yellow. In winter usually lighter above with underparts + Warm Buff to Straw Yellow. 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 side of + hind leg, typically over ankle in extremely narrow line which + widens out over distal phalanges of antiplantar faces of toes but + sometimes interrupted at ankle. Least width of color of underparts + averaging, in twenty available specimens, 39 (27-54) per cent of + greatest width of color of upper parts. Black tip of tail in five + adults averaging 50 (43-60) mm. long; thus averaging longer than + hind foot and 33 per cent of length of tail-vertebrae. + + _Skull and teeth._--Male (based on 4 adults and subadults from + Eureka, Requa, Goldbeach, and Grant Pass): See measurements and + plates 19-21, 30. As described in _Mustela frenata nevadensis_ + except that: Weight, 3.5 (3.5-4.1) grams; basilar length, 42.9 + (41.8-44.0); least width of palate more or less than medial length + of P4. + + Female (based on 2 adults, one from Carlotta and one from 13 mi. + SW Grants Pass): See measurements and plates 34-36. As described + in _Mustela frenata nevadensis_ except that: Weight, 2.4 (2.2-2.6) + grams; basilar length, 37.7 and 39.5; zygomatic breadth less than + distance between condylar foramen and M1 and less than distance + between anterior palatine foramen and anterior margin of tympanic + bulla. See under "_Remarks_" for additional data on variation in + size of skulls of females. + + The skulls of the female averages 31 per cent lighter than that of + the average male. + +Because there is much geographic variation between specimens here +referred to _oregonensis_, the person who is guided by the present +account should keep in mind that results, here reported, of comparisons +of the skull with those of other races, were obtained by employing +specimens of _oregonensis_ from Carlotta and Eureka, California. These +specimens from California are judged to have more of the characters of +the subspecies _munda_ than do specimens of _oregonensis_ from more +northern localities. + +Compared with that of _M. f. washingtoni_ the skull of the male is +shorter, especially in the preorbital region and is relatively broader +across the mastoidal processes and zygomatic arches. The skull of the +female is longer in the preorbital region, has a less cylindrical +braincase and differs less from the male skull than is the case in _M. +f. washingtoni_. Compared with _M. f. effera_, the skull of the male is +smaller in every part measured and relative to the basilar length is +broader across the mastoids and has relatively shorter tympanic bullae. +From _M. f. nevadensis_ the skull of the male differs in the same way +except that size is about the same. The skull of the female +_oregonensis_ is more heavily ridged and is relatively broader across +the mastoids than that of _effera_. From _M. f. saturata_, +_oregonensis_ is not surely known to differ in cranial characters. From +_M. f. munda_, _oregonensis_ differs in having the skull of both sexes +smaller, and on the average, in all parts measured, has a less marked +postorbital constriction, relatively narrower interorbital region and +relatively more expanded zygomata. From _M. f. altifrontalis_, males of +_oregonensis_ differ on the average, in having larger teeth, and +relative to the basilar length, a greater mastoid breadth and a +shallower braincase as measured at the anterior margin of the +basioccipital. Females of _oregonensis_ differ in larger average size +of skull, except for breadth of rostrum and interorbital breadth which, +therefore, are relatively less in _oregonensis_, as also is the +relative depth of the skull measured at the posterior borders of the +upper molars and at the anterior margin of the basioccipital. However, +skulls of females of _oregonensis_ have relatively longer tooth-rows +and are relatively broader across the zygomata and mastoidal processes. + +_Remarks._--In 1896, Merriam named _oregonensis_ as a subspecies of the +California bridled weasel on the basis of a single specimen taken by +Clark P. Streator. Three additional specimens were acquired in later +years, by workers of Dr. Merriam's bureau, from near the type locality +and specimens from farther north in Oregon have been accumulated at the +University of Oregon. The most satisfactory material is that saved from +Humboldt County by the late H. E. Wilder, which, when brought together, +is adequate to give some idea of the range of variation that can be +expected in a given population. + +Of two specimens from Goldbeach, one shows approach to _altifrontalis_ +in that the color of the underparts stops at the ankle, and in one, the +angle of the mouth is dark colored. Specimens from Eugene and vicinity +lack the white facial markings, and in this feature approach the +adjoining _washingtoni-effera-nevadensis_ stock. A specimen from 6 +miles south of Medford shows approach to _saturata_ in the +interruption, on the ankle and lower tibial region, of the color of the +underparts. One adult female, no. 1413, Univ. Oregon, from the Rogue +River Valley, 13 miles southwest of Grants Pass, stands out +prominently, among the other specimens from extreme southern Oregon and +northwestern California, by reason of the near (18) Apricot Yellow +color of the underparts, but this same color occurs in specimens from +the more northerly localities of Buchanan, Eugene, Vida Fish Hatchery, +and McKenzie Bridge, as well as in no. 2178, Univ. Oregon, from Cresent +Lake. The last mentioned specimen is here referred to _nevadensis_. + +Two females referred to _oregonensis_ from southern Oregon differ so +greatly in size of skull that they challenge one's imagination in any +attempt to provide an explanation for so wide a range of variation in +one subspecies. One of these, no. 244520, U. S. Nat. Mus., is an adult +female from Medford. The other, no. 224034, U. S. Nat. Mus., is a +subadult female (though labeled male) from 43 miles northeast of Grants +Pass. The skull of the adult from Medford has a basilar length of 41.5, +upper tooth-rows, 16.1 in length, and a weight of 2.75 grams, whereas +corresponding figures for the subadult are only 33.8, 12.9, and 1.4. +Two other adult females are intermediate in size: No. 1413, Univ. +Oregon, from 13 miles southwest of Grants Pass, Oregon, approaches the +specimen from Medford in size, and the second specimen, no. 34325, Mus. +Vert. Zoöl., from Carlotta, California, is smaller. + +Not only is there a difference in length between the skulls of the two +extremes of the females but this difference extends to all other +dimensions of their skulls, and is most pronounced in the preorbital +region. The differences in breadth of the braincase and other parts of +the skull are relatively less than the differences in length. +Differences of the same nature, although of lesser degree than found in +the females, are to be seen in two males. The skull of an adult no. +51590, Mus. Vert. Zoöl., from 6 miles south of Medford, has a basilar +length of 46.4, upper tooth-rows, 17.6 mm. long, and a weight of 4.0 +grams, whereas corresponding figures for the subadult type specimen +from Grants Pass, are only 43.0, 16.2, and 3.3. + +The wide range of variation in size of skull of both sexes, together +with the considerable variation in color pattern of the specimens here +referred to _oregonensis_ raises the suspicion that we are using the +name in a composite sense; nevertheless, to recognize more than one +subspecies with the material now available would be unwise. + +A subadult female, of abnormal color, no. 47149, Mus. Vert. Zoöl., +taken by Mr. H. E. Wilder at Carlotta, California, on December 20, +1930, in a region where weasels do not turn white in winter, is white, +except for the black tip of the tail, but has a suffusion of orange. +This specimen, discussed at greater length on page 43, is instructive +in that it suggests that there are separate determiners for the brown +and red elements of the pelage. It is interesting also as suggesting +how natural selection may tend to eliminate from the population a +conspicuous color-variation of this kind. At any rate, Mr. Wilder (Ms.) +states: "This specimen was picked up in a field, where it evidently had +been dropped by a hawk or an owl." The braincase of the skull is +crushed in three places as though by a raptor's beak. None of the +several other weasels, all normally colored, saved by Mr. Wilder from +this general locality gives evidence of having fallen a victim to a +raptor. + +Only 2 skulls of the 12 adults and subadults examined show malformation +of the frontal sinuses such as results from the presence of parasites. + + _Specimens examined._--Total number, 29, arranged within states + from north to south by counties. Unless otherwise indicated + specimens are in the collection of the United States National + Museum. + + =California.= _Del Norte County_: Requa, 1[8]. _Humholdt County_: + Eureka, 2 (1[74], 1[75]); Ferndale, 1[74]; Fortuna, 2[63]; + Carlotta, 6 (3[74], 3[59]); 12 mi. E Bridgeville, 1[59]; 2 mi. W + Bridgeville, 1[59]. + + =Oregon.= _Washington County_: Forest Grove, 1. _Marion County_: + Salem, 2. _Benton County_: Buchanan, 1. _Lane County_: McKenzie + Bridge, 1[101]; Vida Fish Hatchery, 1[101]; Eugene, 1[101]. + _Douglas County_: Anchor, 1. _Curry County_: Gold Beach, 2[60]. + _Josephine County_: Rogue River Valley (Grants Pass), 1; 13 mi. SW + Grants Pass, 1[101]. _Jackson County_: Medford, 2; 6 mi. S + Medford, 1[74]. + + +=Mustela frenata munda= (Bangs) + +Long-tailed Weasel + +Plates 1, 19, 20, 21, 22, 23, 30, 34, 35, 36 and 40 + + _Putorius xanthogenys mundus_ Bangs, Proc. New England Zoöl. Club, + 1:56, June 9, 1899; Stephens, California mammals, p. 247, 1906. + + _Mustela frenata_, Audubon and Bachman, Journ. Acad. Nat. Sci. + Philadelphia, 8 (Pt. 2):291, 1842 (North California about 40° + latitude). + + _Mustela xanthogenys munda_, Miller, U. S. Nat. Mus. Bull., 79:99, + December 31, 1912. + + _Mustela frenata munda_, Hall, Carnegie Instit. Washington Publ. + 473:107, November 20, 1936. + + _Type._--Male, adult, skull, os penis and skin; no. 5459, + collection of E. A. and O. Bangs, but now in collection of Mus. + Comp. Zoöl.; Point Reyes, Marin County, California; June 19, + 1896; obtained by C. A. Allen; original no. 931. (See comments + under "Remarks," below, on places in California to which the name + Point Reyes has been applied.) + + The skull (pls. 19-21, 30) is complete and unbroken. I1 on each + side and right I2 are broken away; p2 and p3 on each side have + been aborted and the only alveoli remaining are two for the right + p3. Otherwise all teeth are present and entire. The skin is fairly + well made and in good condition. + + Cranially, the type is a "runt"; its small size and the + circumstance that the tympanic bulla is longer than the lower + molar and premolar tooth-row and longer than the rostrum are + features which differentiate the type from any other specimen seen + of this race. + + _Range._--Sea level to at least 6,000 feet (South Yolla Bolly + Mountain, Trinity County, California); Upper Sonoran and + Transition life-zones of the coast and Coast Range of northwestern + California from the Golden Gate northward into southern Humboldt + and Trinity counties. See figures 29 and 30 on pages 221 and 314. + + _Characters for ready recognition._--Differs from _M. f. + oregonensis_ in longer hind foot of males which is more than 50 + mm., and in both sexes, larger, more prominently ridged skull (see + measurements and plates); from _M. f. saturata_ by presence of + nasofrontal white spot, larger and relatively shallower skull of + males and larger skull of female; from _M. f. nevadensis_ by + presence of well-developed, white, facial markings; absence of + color of underparts on ventral face of proximal third of tail; and + hind foot of males more than 50; from _M. f. xanthogenys_ by near + (_l_) Sudan Brown to near (_l_) Antique Brown rather than + Buckthorn Brown colors of upper parts and greater size, and in + adult male basilar length more than 45 and hind foot more than 47; + from _M. f. nigriauris_ by having inside of ears same color as + back rather than much darker than back. + + _Description._--_Size._--Male: Three adults and two young from + Point Arena and Gualala, Mendocino County, yield average and + extreme measurements as follows: Total length, 447 (434-470); + length of tail, 167 (150-185); length of hind foot, 53 (50-60). + Corresponding measurements of three adults from 5 and 6 miles west + of Inverness, Marin County, are: 430 (420-440), 154 (141-160), 48 + (48-49). Corresponding measurements of four individuals (3 adults + and 1 young of large size) from South Yolla Bolly Mountain, + Trinity County, are: 383 (374-400), 134 (130-138); 44 (43-44). The + tail averages 60 per cent as long as the head and body in the + series from Point Arena, 56 per cent in the series from Point + Reyes, and 53 per cent in the series from South Yolla Bolly + Mountain. In every specimen except two, length of hind foot less + than basal length. The two exceptions are no. 19720, M.V.Z., male + adult from Point Arena in which the hind foot is recorded as 60 + (probably an error in measurement), and no. 19721, M.V.Z., from + the same place, in which the skull has not yet attained its full + growth. + + Female: One adult from Point Arena measures as follows: Total + length, 383; length of tail, 134; length of hind foot, 43. + Corresponding measurements of an adult from seven miles north of + Laytonville, Mendocino County, are: 336, 121, 33 (= 36 on dried + skin). Corresponding measurements of an adult from South Yolla + Bolly Mountain, Trinity County, are, 326, 113, 37. In these three + specimens, the tail is, in the order given, 54, 56, and 53 per + cent as long as the head and body. Length of hind foot more than + basal length. + + Differences in external measurements of the two sexes as indicated + by the five males and one female from Point Arena, are: Total + length, 64; length of tail, 33; length of hind foot, 10. Weights + of 2 adult males are 265 and 221 grams and of one adult female 155 + grams. + + _Externals._--As described in _Mustela frenata nigriauris_. + + _Color._--Spot between eyes, narrow band or spot confluent with + color of underparts on each side of head anterior to each ear, + chin, lower lips, and rarely posterior third or less of each upper + lip white; dark spot posterior to each angle of mouth uniformly + present and of large size; tip of tail black; remainder of upper + parts near (14 _l_) Sudan Brown and tone 4 of Raw Umber of + Oberthür and Dauthenay, pl. 301; occasionally, slightly darker + brown on forehead, nose, and about eyes. Underparts near (_a_ to + _c_) Ochraceous-Buff and sometimes Orange-Buff. 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 over antiplantar faces of toes. Least width of color of + underparts averaging, in a series of 5 males from Mendocino + County, 57 (46-67) per cent of greatest width of color of upper + parts; 38 (35-40) in 3 males from Point Reyes, Marin County. Black + tip of tail in Mendocino County series averaging 53 (46-60) mm., + which is same length as hind foot and 32 per cent of length of + tail. In Point Reyes males, black tip of tail averages 44 (34-52) + mm., which is less than length of hind foot and 45 per cent as + long as tail-vertebrae. + + Several specimens of the smaller, inland variant (see under + "_Remarks_") are near (_l_) Antique Brown rather than near (14 + _l_) Sudan Brown above and hence do not differ in this respect + from _nigriauris_. + + _Skull and teeth._--Male (based on 3 adults from Mendocino + County): See measurements and plates 19-23, 30. As described in + _Mustela frenata nigriauris_ except that: Weight, 6.0 (5.4-6.3) + grams; basilar length, 47.6 (46.5-48.2); length of tympanic bulla + more than length of lower molar and premolar tooth-row. + + Female (based on no. 19723, M.V.Z., from Point Arena): See + measurements and plates 34-36, 40. As described in _M. f. + nigriauris_ except that: Weight, 3.0 grams; basilar length, 42.3. + + The skull of the female is 50 per cent lighter than that of the + average male. + +Compared with the skull of the male of _nevadensis_ that of _munda_ +averages larger in every part measured and specimens from Point Arena +are nearly as heavy again, have relatively more expanded zygomata and +mastoid processes but are relatively narrower anteriorly as shown by +the breadth of the rostrum, interorbital breadth and postorbital +breadth. Also the braincase is less inflated anteriorly, the tympanic +bullae are lower and the skull is more angular. Females show the same +differences although in different degree. Compared with the skull of +the male of _M. f. nigriauris_, that of _munda_ from Point Arena +averages larger in every part measured except for the length of the +upper tooth-rows. Relative to the basilar length, the skull of _munda_ +averages broader across the mastoids and across the zygomata, is deeper +through the braincase at the anterior end of the basioccipital, and +has a greater development of the lambdoidal crest. + +_Remarks._--The skin and part skull, no. 536/1849, U. S. Nat. Mus., +taken by Lieutenant W. P. Trowbridge at San Pablo Bay, is the first +specimen known to have been saved of this subspecies. Since 1899 when +O. Bangs diagnosed _munda_ as of small size, the weasel of the humid +costal belt north of San Francisco Bay has been regarded as smaller +than bridled weasels from farther south in the State. Actually, +however, the weasel of the humid costal belt shares with _M. f. +pulchra_ the distinction of being one of the two largest weasels in +California. + +_M. f. munda_ may be a composite subspecies, for the variation in +facial markings, in coloration otherwise, in external measurements and +in size and shape of skull is great. At one time in the course of the +present study, manuscript accounts of two subspecies were prepared for +the animals now all called _munda_ and there is still much +justification for recognizing two subspecies, one, along the coast +proper, the larger, darker-colored animal with reduced white facial +markings and large, wide, heavily ridged skull from Point Arena, and 6 +miles south of Laytonville, Mendocino County, along with the specimens +from 5 and 6 miles west of Inverness, Marin County, and the other, an +inland race, which is a smaller, lighter-colored animal with more +extensive white facial markings and a smaller, narrower, skull, known +by specimens from Point Reyes [station?], Nicasio, 15 mi. north of San +Rafael, Freestone, Vallejo, and Mount Sanhedrin. The differences +between these two lots of specimens are of great degree. However, a +female from Fort Bragg proves to be no larger than three females +labeled as from Point Reyes. Also, a male from 2 miles south and one +mile east of Stewarts Point on the coast has a skull no larger than the +animal from Vallejo, whereas the skin alone of an adult female from 3 +miles south of Stewarts Point is large and agrees with the specimens +from Point Arena. Consequently, no logical ranges can be worked out for +the two variants with the material now available. + +Finally, the type specimen of _munda_ is a "runt," smaller than any +other male seen. This specimen, purchased by E. A. and O. Bangs from C. +A. Allen, who collected and sold specimens widely, was labeled as from +Point Reyes. So far as this place-name is concerned, it might refer to: +(1) The point of land by that name which projects out into the Pacific +Ocean, (2) an abandoned ranch house bearing that name at the head of +Drakes Bay, 6 miles north and 3-3/4 miles east of the actual point, or +(3) the railway station by the same name at the head of Tomales Bay, +12 miles east and 4-3/4 miles north of the actual point. Allen, +himself, lived near San Geronimo (then Nicasio) about nine miles +southeast of the Point Reyes railway station. All these places are in +Marin County, but differ markedly as regards climate and flora. The +first two are treeless, windswept and have much fog, whereas Point +Reyes Station is more often sunny, and is situated in a shallow valley, +inland, where the open grass-covered west-facing slopes meet the +east-facing wooded ones. From which one of these three places the type +specimen came, I do not know. The same may be said of the three female +specimens labeled Point Reyes; two of these are in the United States +National Museum and one in the Field Museum. + +The specimens in the Museum of Vertebrate Zoölogy from 5 and 6 miles +west of Inverness and those from near the same place in the collection +of John Cushing come from within a couple of miles or less of the Point +Reyes represented by the abandoned ranch house. These specimens, as +remarked above, agree with those from Point Arena in large size, +reduced facial markings and wide skull. These are points of difference +from the smaller variant suspected of being a recognizable subspecies. +It is the smaller variant which the type specimen approaches in size, +and with which it agrees in relatively well-developed white facial +markings. This suggests that the type specimen came from Point Reyes +Station rather than from either of the two other places bearing the +name "Point Reyes," from one of which, as just stated, the variant of +large size is known. The three females labeled "Point Reyes" also have +well-developed white facial markings and are of lesser size than the +female of similar age from Point Arena, Mendocino County. The +presumption is that these three females also came from Point Reyes +Station. + +The smaller, inland variant seems to agree in size, cranial characters, +and coloration with _M. f. nigriauris_ to the southward of San +Francisco Bay, but lacks the black on the head which characterizes +_nigriauris_. The larger variant, on which the description here used +for _munda_ is based, comprises animals which differ from _nigriauris_ +in larger size, darker color, reduced white facial markings, and +larger, relatively wider skull. Both of the variants mentioned above +are sharply distinct from _nigriauris_ on the basis of coloration of +the inside of the ear which is blackish in nigriauris like the dark +facial markings, and in _munda_ is colored like the back. _M. f. munda_ +lacks the dark facial markings; an occasional specimen has at most, a +trace of the markings but this does not extend back so far as the ears. +This difference, blackish versus non-blackish face, persists eastward +of San Francisco Bay to at least as far as the Carquinez Straits, where +a specimen of _munda_ is available from 4 miles north of Vallejo and +one of _nigriauris_ from Glen Frazer Station on the south shore +opposite Vallejo. + +[Illustration: FIG. 30. Map showing the geographic distribution of +subspecies of _Mustela frenata_ in California] + +Intergradation with _M. f. nevadensis_ and possibly with _M. f. +saturata_ is indicated by specimens from South Yolla Bolly Mountain, +Trinity County. In them the external measurements and measurements of +the skull are intermediate. Also the white frontal spot is much reduced +in size. The white bars in front of the ears are absent in three +specimens, and weakly developed in the other two. The relative +proportions of the skulls as a whole are nearer those of _nevadensis_ +or _saturata_ than _munda_. The skull of one of the three adult males +and the skull of the adult female suggests _M. f. oregonensis_ in +certain features; for example, the dorsal outline of the skull in +longitudinal axis is slightly convex as it is in _oregonensis_. + +None of the specimens shows malformation of the frontal sinuses such as +results from infestation by parasites. + + _Specimens examined._--Total number, 37, arranged by counties from + north to south. Unless otherwise indicated specimens are in the + Museum of Vertebrate Zoölogy. + + =California.= _Trinity County_: S. Yolla Bolly Mt., 3[91]; 1/2 mi. + S S. Yolla Bolly Mt., 1. _Tehama County_: 2 mi. S S. Yolla Bolly + Mt., 1. _Mendocino County_: 6 mi. N Laytonville, 1; Mt. Sanhedrin, + 1[87]; Ft. Bragg, 1; Gualala, 1; Point Arena, 5. _Sonoma County_: + 2 mi. S and 1 mi. E Stewarts Point, 1; 3 mi. S Stewarts Point P. + O., 1; Freestone, 1. _Napa County_: 6 mi. SSW, Napa, 1; 4 mi. N + Vallejo, 1. County in question: San Pablo Bay, 1[91]. _Marin + County_: 6 mi. W Inverness, 2; 5 mi. W Inverness, 2(1[28]); Point + Reyes, 4 (2[91] 1[60], 1[75]); Nicasio, 2 (1[60], 1[75]); Kehoes + Ranch, Pierce Point, 1[28]; Drakes Bay, 1[28]; Tomales Point, + about 1/2 mi. SW White Gulch, 1; Point Reyes School, 3-3/4 mi. W + Inverness, 1; 15 mi. (by road) N San Rafael, 1[52]; Hurley Ranch, + 2 mi. W Tomales, 1. No locality more definite than California, + 1[7]. + + +=Mustela frenata xanthogenys= Gray + +Long-tailed Weasel + +Plates 21, 22, 23, 28, 30, 34, 35 and 36 + + _Mustela xanthogenys_ Gray, Ann. and Mag. Nat. Hist., 11:118, 1843. + + _Putorius (Gale) brasiliensis frenatus_, Coues, Fur-bearing + animals, p. 142, 1877 (part). + + _Putorius xanthogenys_, Merriam, N. Amer. Fauna, 11:25, June 30, + 1896; Bangs, Proc. New England Zoöl. Club, 1:56, June 9, 1899. + + _Mustela xanthogenys xanthogenys_, Miller, U. S. Nat. Mus. Bull., + 79:99, December 31, 1912. + + _Mustela frenata xanthogenys_, Hall, Carnegie Instit. Washington + Publ. 473:107, November 20, 1936. + + _Type._--Male, adult, skull and skin; skull no. 197a-43.6.4.55, + skin no. 234a-42.11.21.4, British Museum (Nat. Hist.); from the + bank of Sacramento River below mouth of Feather River, or from + north shore of San Francisco Bay, California; taken in "1837 or + 1838"; presented by Captain Edward Belcher. + + The skull (plate 28) lacks the occiput, the right mandible + posterior to m1, and the right pterygoid; the right zygomatic arch + is fractured. The teeth are not greatly worn. The skin was + originally mounted for exhibition (R. I. Pocock in Litt.) but in + 1937 when I saw the skin, it was prepared as a conventional study + skin. The skin is in fairly good condition; some hair is missing + on the hind quarters and the skin of the tail is torn at one + place. + + _Range._--Altitudinally, less than 600 feet (Fair Oaks); Lower + Sonoran and Upper Sonoran life-zones of all but southern end of + the San Joaquin Valley, and probably Sacramento Valley, + California. See figures 29 and 30 on pages 221 and 314. + + _Characters for ready recognition._--Differs from _M. f. + nevadensis_ by presence of light facial markings and Buckthorn + Brown rather than near (14_n_ to _l_) Brussels Brown color of + upper parts; from _M. f. munda_ by Buckthorn Brown rather than + near (_l_) Sudan Brown, or near (_l_) Antique Brown color of upper + parts and lesser size, in adult males basilar length less than 45 + and hind foot less than 47; from _M. f. nigriauris_ by lighter + color in same way as from _munda_ and also by having inside of + ears same color as back rather than much darker than back; from + _M. f. pulchra_ in hind foot of males less than 46 and narrower + skull, in males having breadth of rostrum less than 13.9 and + mastoid breadth less than 26.0, see comparison of skulls in the + account of _pulchra_. + + _Description._--_Size._--Male: Three adults, from Fresno, Selma + and Los Banos, measure, respectively as follows: Total length, + 425, 417, 450; length of tail, 152, 154, 180; length of hind + foot,--, 43, 44. Tail averages 61 per cent as long as head and + body. Length of hind foot less than basal length. + + Female: Adults from Selma, Los Banos, and 4 mi. SW Turlock, + measure respectively as follows: Total length, 357, 365, 395; + length of tail, 133, 132, 145; length of hind foot, 40, 38, 41. + Tail averages 58 per cent as long as head and body. Length of hind + foot less than basal length. + + The average differences in external measurements between the two + sexes, as represented by these six specimens, are: Total length, + 65; length of tail, 25; length of hind foot, 3.5. One adult male + weighs 274 grams and 2 adult females 182 and 214 grams. + + _Externals._--As described in _Mustela frenata nigriauris_. + + _Color._--Spot between eyes, band confluent with color of + underparts on each side of head extending anterodorsally anterior + to each ear, and posterior half to third of each upper lip white, + or whitish tinged with some shade of yellowish; chin and lower lip + white; dark spot posterior to each angle of mouth of varying size + but uniformly present; tip of tail black; remainder of upper parts + Buckthorn Brown of Ridgway or a trifle browner than tone 4 of + Brown Pink of Oberthür and Dauthenay, pl. 297. Upper parts of + uniform color except for slight darkening of head-markings + anterior to ears. Underparts Ochraceous-Buff to Warm Buff. 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 over antiplantar faces of toes and sometimes tarsal + region. Least width of color of underparts averaging, in 9 + specimens from Fresno, Selma and Los Banos, 54 (32-74) per cent of + greatest width of color of upper parts. Black tip of tail in three + males (one subadult and 3 adults) averages 55 (50-60) mm. long. + Thus longer than hind foot and averaging 34 per cent of length of + tail-vertebrae. + + _Skull and teeth._--Male (based on 2 adults from Fresno and one + from Selma): See measurements and plates 21-23, 30. As described + in _M. f. nigriauris_ except that: Weight 3.8 grams; basilar + length, 43.7 (43.4-43.9); least width of palate more or less than + lateral length of P4; length of tympanic bulla more than length of + lower molar and premolar tooth-rows. + + Female (no. 2626 W. E. Snyder, from Selma): See measurements and + plates 34-36. As described in _M. f. nigriauris_ except that: + Weight, 2.5 grams; basilar length, 39.4. + + The skull of the female is 34 per cent lighter than the average + for the three males. + +Compared with skulls of _nevadensis_ from the Sierra Nevada, those of +the two adult males from Fresno differ as follows: M1 wider +(transversely); tympanic bullae narrower; preorbital part of skull +smaller. Comparison with _pulchra_ is made in the account of that +subspecies. Compared with skulls of adult males of _nigriauris_, from +Santa Clara County, the two skulls from Fresno are generally smaller +and in basilar length, length of tooth-rows and measurements of the +teeth fall below the minimum for _nigriauris_. Relative proportions of +the skulls are approximately the same. Comparison with _munda_ reveals +essentially the same differences as does comparison with _nigriauris_ +except that the difference in size is greater. + +_Remarks._--The name _Mustela xanthogenys_ Gray was long applied to all +the weasels of the interior valleys of California and of the coast of +that state south of San Francisco Bay. Gray, when he named the species +and when referring to it in later accounts, never defined the locality +whence the specimen came more definitely than "California." In 1896, +Merriam (1896:25) gave the type locality as "Southern California, +probably vicinity of San Diego" and later writers have not contradicted +him. The type specimen was obtained in the course of the voyage of the +British ship Sulphur, under command of Sir Edward Belcher. Examination +of Belcher's (1843, vol. 1, p. 129) narrative of the voyage indicates +the following places in California at which the specimen of weasel, +described by Gray, could have been obtained: Fort Ross, Bodega, +vicinity of San Francisco Bay and up Sacramento River to the mouth of +the Feather River, Monterey, Santa Barbara, Buenaventura, San Pedro, +San Juan, and San Diego. + +Reginald I. Pocock has kindly compared the type specimen in the British +Museum with several specimens sent for that purpose. In the first +place, comparison of skulls shows that the type specimen is a member of +one of the races north of San Diego. In the second place, comparison of +skins shows that the inside of the ears are not blackish but similar in +color to the back. In fact, Pocock writes under date of February 12, +1929, regarding the type specimen, that "It is practically uniformly +colored from the snout to the base of the tail, there being scarcely a +trace of the darkening of the head, or muzzle, observable in your +specimens [those sent for comparison]." This character of coloration of +the ear excludes all the weasels of the Coast region of California from +San Francisco Bay southward, namely, _M. f. latirostra_ and _M. f. +nigriauris_. My own examination of this type specimen at a date later +than that on which Pocock compared it satisfies me as to the accuracy +of his statement above. + +Accordingly, the name _xanthogenys_ would seem to apply to one of the +two subspecies here called _munda_ and _xanthogenys_. Perusal of +Belcher's narrative of the voyage (_loc. cit._) shows that little, if +any, opportunity was afforded to obtain vertebrate specimens at Fort +Ross or Bodega, both localities within the range of the subspecies here +called _munda_. Furthermore, the type specimen is smaller than +individuals of _munda_ from 5 to 6 miles west of Inverness and from +Point Arena with which the animals from Fort Ross and Bodega would be +expected to agree in size. Weasels from along the north shore of San +Francisco Bay are smaller than those on the coast north of the bay. +Possibly the type specimen of _xanthogenys_ came from the north side of +San Francisco Bay but probably it came from the bank of the Sacramento +River and almost certainly not farther up stream from San Francisco Bay +than the junction of the Sacramento and Feather rivers. The statement +of Belcher (1843, vol. 1, p. 129), regarding the trip up the Sacramento +River as far as Point Victoria, lat. 38°46´47" north, and return to +San Francisco Bay, that "Cuyote or jackal--fox, racoon, land otter, +weasel, and squirrel were obtained" lends strong probability to the +idea that this type specimen was taken along the Sacramento River, +possibly in the vicinity of the existing city of Sacramento. +Unfortunately no specimens are available from the Sacramento Valley. If +some were available, a comparison of them and specimens of _munda_ from +along the north side of San Francisco Bay and Carquinez Straits with +the type specimen of _xanthogenys_ should determine the correct +application of the name. For the present it seems best to retain the +name _munda_ and apply the name _xanthogenys_ to the weasels inhabiting +the northern part of the San Joaquin Valley and presumably the southern +part of the Sacramento Valley. + +Efforts to obtain specimens of weasels from the Sacramento Valley have +been in vain. A juvenal specimen taken five miles south of Fair Oaks, +Sacramento County, by Mr. John Fitzgerald, Jr., in December, 1927, was +examined at his home and found to agree in coloration with specimens +from farther south. Geographically, this specimen probably is more +nearly a topotype than any other examined. + +Most of the specimens examined are immature and adequate adult cranial +material has not been seen. Two adults, one of each sex, from Los Banos +have skulls of large size which agree with those of _nigriauris_. The +same is true of one adult and one young female from 4 miles southwest +of Turlock, which, unlike the animals from Los Banos, show a darkening +of the head extending in reduced degree even to the inside of the ears, +as in _nigriauris_. The slightly darker than average (for +_xanthogenys_) color on the back may indicate intergradation with +_nevadensis_. Intergradation with _M. f. nevadensis_ is shown by +specimens, from the southern part of the Sierra Nevada, mentioned in +the account of _nevadensis_. + +None of the skulls shows malformation of the frontal sinuses such as +result from infestation by parasites. + + _Specimens examined._--Total number 30, arranged by counties from + north to south. + + =California.= _Sacramento County_: Bank of Sacramento River, 1[7]; + 5 mi. S Fair Oaks, 1[29]. _San Joaquin County_: 4 mi. W Stockton, + 1[74]. _Merced County_: Tegner School, 4 mi. SW Turlock, 2; Los + Banos, 4 (2[74], 1[91] 1[87]). _Fresno County_: Mendota, 1[74]; + Biola, 1[30]; Clovis, 1[55]; Fresno, 5 (1[74], 1[91], 2[55], + 1[1]); 5 mi. W Fresno, 1[14]; Selma, 3 (2[50], 1[104]); 4 mi. NW + Sanger, 1[55]; 5 mi. S Selma, 1[62]. _Tulare County_: Monson, + 1[74]; 1-1/2 mi. N Goshen, 1[74]; Milo, 1[91]; 2 mi. N Tipton, + 1[74]; Poplar, 2[53]. No locality more definite than California, + 1[4]. + + +=Mustela frenata nigriauris= Hall + +Long-tailed Weasel + +Plates 22, 23, 24, 34, 35, 36 and 41 + + _Mustela frenata nigriauris_ Hall, Carnegie Instit. Washington + Publ. 473:95, November 20, 1936. + + _Putorius xanthogenys_, Baird, Mamm. N. Amer., 1858, p. 176 (part). + + _Mustela xanthogenys_ Gray, Ann. and Mag. Nat. Hist., 14(ser. + 4):375, 1874 (part?). + + _Putorius (Gale) brasiliensis frenatus_, Coues, Fur-bearing + animals, p. 142, 1877 (part). + + _Putorius xanthogenys xanthogenys_, Grinnel, Proc. California Acad. + Sciences, fourth series, 3:292, August 28, 1913. + + _Mustela xanthogenys xanthogenys_, Miller, U. S. Nat. Mus. Bull., + 79:99, December 31, 1912; Grinnell, Univ. California Publ. Zoöl., + 40:102, September 26, 1933. + + _Type._--Male, adult, skeleton and skin; no. 32820, Mus. Vert. + Zoöl.; Half Moon Bay, San Mateo County, California; received at + Museum of Vertebrate Zoölogy, May 4, 1922, through A. L. + Hagedoorn, after having been in captivity a few days where death + occurred owing to injuries received in trap; original no. 1590. + + The skull has each of the zygomatic arches and the anterior end of + the nasals broken through. The only part missing is the central + two millimeters of the left zygomatic arch. The teeth all are + present and entire. The skeleton appears to be complete except for + the bones of the feet, which are preserved within the skin. The + skin is well made and in good condition. + + _Range._--Altitudinally, sea level to more than 4000 feet; Sonoran + and Transition life-zones of Coast Range and coast of California + from San Francisco Bay south to Point Conception, Santa Barbara + County, California. See figures 29 and 30 on pages 221 and 314. + + _Characters for ready recognition._--Differs from _M. f. munda_, + _xanthogenys_, and _pulchra_ by having inside of ears darker than + back rather than same color as back, and from _xanthogenys_ and + _pulchra_ in near (_l_) Antique Brown color of upper parts rather + than Buckthorn Brown or near (16 _j_) Buckthorn Brown to near + (_h_) Yellow Ocher respectively; from _M. f. latirostra_ by + postorbital breadth, of adult males and females, less, rather than + more, than width of basioccipital measured from medial margin of + one foramen lacerum posterior to its opposite. + + _Description._--_Size._--Male: Five adults from Palo Alto, Santa + Clara County, yield average and extreme measurements as follows: + Total length, 447 (412-465); length of tail, 167 (147-175); length + of hind foot, 46 (45-47). Corresponding measurements of four + adults from San Francisco are: 412 (394-435); 153 (145-160); 43.5 + (41-46). Corresponding measurements of five adults and subadults + from Berkeley, Alameda County, are: 419 (390-448); 148 (135-160); + 44 (42-47). Tail averages 59 per cent as long as head and body in + series from Palo Alto and in one from San Francisco. The average + of 55 for the Berkeley series probably reflects a lesser average + age. Length of hind foot less than basal length. The type specimen + measures, 415, 150, 43. It is smaller than the mean. + + Female: A subadult from Palo Alto measures: Total length, 368; + length of tail, 126; length of hind foot, 39. An adult and two + subadults from Berkeley measure, respectively, as follows: Total + length, 347, 365, 340; length of tail, 134, 123, 125; length of + hind foot, 37, 38.4, 36.5. In these four females the tail averages + 55 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 represented by specimens from Berkeley, Alameda County, are: + Total length, 68; length of tail, 21; length of hind foot, 7. + Eight adult males weigh 249 (217-335) grams and one adult female + 123 grams. + + _Externals._--Longest facial vibrissae brownish like dark color of + head and extending beyond ear; carpal vibrissae mostly color of + underparts and extending to apical pad of fifth digit; hairiness + of foot-soles slightly more than shown in figure 20. + + _Color._--Spot between eyes, band, confluent with color of + underparts, on each side of head extending anterodorsally anterior + to ear, and posterior third of each upper lip tinged with color of + underparts or, less often, pure white; chin and lower lips white; + remainder of sides and top of head posteriorly to, or a little + behind, a line connecting posterior margins of ears, blackish; + inside of pinna of ear, and sometimes outside of pinna, blackish; + dark spot posterior to each angle of mouth present on each side in + three-fourths of specimens; tip of tail black; remainder of upper + parts near (_l_) Antique Brown, and with more yellow than tone 3 + of Raw Umber of Oberthür and Dauthenay, pl. 301. Often with more + blackish and red in winter. Underparts near (_a_ to _c_) + Ochraceous-Buff or Ochraceous-Salmon. Ochraceous-Salmon in some + juveniles. 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 limbs over antiplantar faces of toes. + Least width of color of underparts averaging, in 17 adult males + (Berkeley, 5; San Francisco, 5; Palo Alto, 7), 55 (40-73) per cent + of greatest width of color of upper parts. Black tip of tail in + same series of males averaging 51 (35-60) mm., thus averaging + longer than hind foot and 33 per cent of length of tail (Palo Alto + and San Francisco, 31 per cent; Berkeley, 35 per cent). In 8 adult + females, least color of underparts amounts to 55 (47-62) per cent + of greatest width of color of upper parts. Black tip of tail + averages 41.5 (28-50) mm., thus averaging longer than hind foot + and 32 per cent of length of tail-vertebrae. + + _Skull and teeth._--Male (based on six adults from Stanford Univ. + and vicinity): See measurements and plates 22-24; weight (four + adults), 5.4 (5.0-5.9) grams; basilar length, 47 (46.1-48.1); + zygomatic breadth more than distance between condylar foramen and + M1, 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 (less than + distance between posterior borders of P4 and P2) and 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 rostrum less than length of tympanic + bulla; least width of palate less than lateral length of P4; + anterior margin of tympanic bulla as far posterior to foramen + ovale as width of 3 or 4 (including I3) upper incisors; height of + tympanic bulla more than distance from its anterior margin to + foramen ovale; length of tympanic bulla more or less than (about + equal to) length of lower molar and premolar tooth-row and longer + or shorter (usually shorter) than rostrum; anterior margin of + masseteric fossa below anterior half of m2. + + Female (based on three adults, Hayward, Palo Alto, and Morro): See + measurements and plates 34-36; weight (no. 43574, from Morro) 2.7 + grams; basilar length, 41.2 (40.2-42.2); zygomatic breadth more or + less than distance between condylar foramen and M1 and more or + less than distance between anterior palatine foramen and anterior + margin of tympanic bulla; postorbital breadth less than length of + upper premolars and less than width of basioccipital measured from + medial margin of one foramen lacerum posterior to its opposite; + least width of palate less than lateral length of P4; tympanic + bulla as far posterior to foramen ovale as width of 3 (including + I3) 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 50 per cent lighter than that of + the average male. + + Comparisons of the skull of the male with those of _M. f. + latirostra_, _pulchra_, _xanthogenys_, and _munda_ are made in the + accounts of those subspecies. + +_Remarks._--Like _M. f. latirostra_, _nigriauris_ long bore the name +_xanthogenys_. The fairly adequate lot of specimens is divided between +the collections of several institutions. The most satisfactory material +in any one collection is in the Stanford University Natural History +Museum where local specimens have been accumulated over a period of +many years. + +No actual intergrade between _nigriauris_ and _xanthogenys_ has been +seen, although the specimens from Los Banos, referred to _xanthogenys_, +have large skulls as in _nigriauris_. Intergradation with _latirostra_ +is shown by specimens, referred to _latirostra_, from the Los Angeles +area. Also the one adult male from 5 miles southeast of Santa +Margarita, San Luis Obispo County, is of small size and in this respect +approaches _latirostra_. The range of _nigriauris_ is separated from +that of _munda_ by San Francisco Bay, Carquinez Straits, and I suppose +by the lower part of the San Joaquin River. On the basis of color of +the inside of the pinna of the ear, the two subspecies are uniformly +distinct. Intergradation is assumed to occur through the subspecies +_xanthogenys_. + +None of the 26 adult and subadult specimens examined for evidences of +infestation of the frontal sinuses by parasites shows malformation of +the sinuses. + + _Specimens examined._--Total number, 103, arranged by counties + from north to south. Unless otherwise indicated specimens are in + the Museum of Vertebrate Zoölogy. + + =California.= _Contra Costa County_: Glen Frazer Station, 1; 2 mi. + W Pinole, 1[13]; 1 mi. E Pinole, 1; Richmond, 1[13]; Lafayette, 1; + 7 mi. E Clayton, 1; Moraga Valley, 1; Pinehurst, Redwood Canyon, + 1; Concord, 1. _Alameda County_: Berkeley, 11; Oakland, 1; + Piedmont, 1; Haywards, 2; near Haywards, 2; 10 mi. E Haywards, + 1[91]; Redwood Canyon, 1; Calaveras Dam, 1. _San Francisco + County_: San Francisco, 11 (5[8], 2[91], 1[60], 1[7]); Ocean View, + 1[68]; Visitation Valley, 1. _San Mateo County_: Moss Beach, 1; + Half Moon Bay, 1; Redwood City, 1[87]; Menlo Park, 9 (5[87], + 2[68]); no locality more definite than county, 1[8]. _Santa Clara + County_: 1/4 mi. N Milpitas, 1; 1/4 mi. S Milpitas, 1; Stanford + University, 6 (4[68], 2[91]); Palo Alto, 11 (6[41], 2[60], 1[75], + 1[87]). _Santa Cruz County_: 3 mi. E Santa Cruz, 1; 2-1/2 mi. E + Santa Cruz, 1; Santa Cruz, 6 (2[91], 1[68], 1[4]). _Monterey + County_: 1 mi. E mouth Salinas River, 10 ft., 1[37]; Pacific + Grove, 1[8]; Monterey, 2 (1[7]); Carmel, 1[8]; Carmel Valley, + 1[68]; Point Lobos, 1; Gonzales, 1. _San Luis Obispo County_: 5 + mi. SE Santa Margarita, 1; Morro, 1[91]; 3-1/2 mi. S Oceano, 6. + _Santa Barbara County_: Santa Maria, 1[87]; 5 mi. N Las Cruces, 1; + 7 mi. W Gaviota, 1; Gaviota, 1. + + +=Mustela frenata latirostra= Hall + +Long-tailed Weasel + +Plates 1, 22, 23, 24, 34, 35 and 36 + + _Mustela frenata latirostra_ Hall, Carnegie Instit. Washington + Publ. 473:96, November 20, 1936. + + _Putorius xanthogenys_, Baird, Mamm. N. Amer., p. 176, 1858 (part); + Stephens, California mammals, p. 246, 1906; Merriam, N. Amer. + Fauna, 11:25, June 30, 1896 (part). + + _Putorius (Gale) brasilianus frenatus_, Coues, Fur-bearing animals, + p. 142 (part). + + _Mustela xanthogenys xanthogenys_, Miller, U. S. Nat. Mus. Bull., + 79:99, December 31, 1912; Grinnell, Univ. California Publ. Zoöl., + 40:102, September 26, 1933. + + _Mustela arizonensis_, Grinnell and Swarth, Univ. California Publ. + Zoöl. 10, 376, October 31, 1913. + + _Type._--Male, adult, skull and skin; no. 3257, Mus. Vert. Zoöl.; + San Diego, San Diego County, California; May 20, 1907; obtained by + Frank X. Holzner. + + Right M1 is missing and the part of the jaw bearing this tooth is + broken away. With this exception the skull is complete and + unbroken and the teeth are all present and entire. The skin is + fairly well made and in good condition except that it is slightly + soiled. + + _Range._--Altitudinally sea level to 8000 feet (Tahquitz Valley, + San Jacinto Mountains); Sonoran and Transition life-zones of coast + and mountains west of Mohave and Imperial deserts of southern + California from Point Conception and Cuyama Valley southward at + least to Mexican boundary. See figures 29 and 30 on pages 221 and + 314. + + _Characters for ready recognition._--Differs from _M. f. + nigriauris_ by having postorbital breadth of adult males and + females, more, rather than less, than width of basioccipital + measured from medial margin of one foramen lacerum posterior to + its opposite; from _M. f. pulchra_ by having tympanic bulla longer + than rostrum (orbitonasal length) and by near (_l_) Antique Brown + rather than near (16 _j_) Buckthorn Brown to near (_h_) Yellow + Ocher color of upper parts. + + _Description._--_Size._--Male: Six adults and subadults from San + Diego yield average and extreme measurements as follows: Total + length, 439 (428-449); length of tail, 153 (142-160); length of + hind foot, 45 (40-47). Corresponding measurements for a series of + eight adult males from the vicinity of Los Angeles are: 416 + (394-428); 158 (151-166); 44 (40-47). In the series from San Diego + the tail averages 54 per cent as long as head and body. In the + series from Los Angeles the average is 61 per cent. Length of hind + foot in each series, less than basal length. The type specimen + measures, 435, 142, 42. + + Female: No. 5070, adult, from San Diego, measures 367, 141, 38. + Nos. 22 and 6748 from Santa Ysabel, measure: 359, 380; 130, 140; + 39, 35. No. 7194 from Jamacha measures, 358, 125, 35. Three adult + females from Los Angeles yield the following: Total length, 373, + 345, 368; length of tail, 150, 122, 134; length of hind foot,--, + 41, 41. In no. 5070 the tail is 62 per cent as long as the head + and body and in the three from Los Angeles it averages 60 (55-67) + per cent. Length of hind foot, in each case, less than basal + length. + + The average differences in external measurements of the two sexes + as shown by the six males from San Diego and the four females + from San Diego County are: Total length, 73; length of tail, 19; + length of hind foot, 8. Corresponding differences shown by the + eight males and three females from Los Angeles are: 54, 23, 3. + + _Externals._--Longest facial vibrissae brownish, like dark color + of head and extending beyond ear; carpal vibrissae mostly color of + underparts and extending to apical pad of fifth digit; hairiness + of foot-soles slightly more than shown in figure 20. + + _Color._--Spot between eyes, band confluent with color of + underparts on each side of head extending anterodorsally anterior + to ear, and posterior third of each upper lip tinged with color of + underparts or, less often, white; chin and lower lips white; + remainder of sides and top of head posteriorly to near line + connecting posterior margins of ears, blackish; inside of pinna of + ear, and sometimes outside of pinna, blackish; dark spot posterior + to each angle of mouth present on each side in three-fourths of + specimens; tip of tail black; remainder of upper parts near (_l_) + Antique Brown, and with more yellow than tone 3 of Raw Umber of + Oberthür and Dauthenay, pl. 301. Underparts Ochraceous-Buff to + Warm Buff and in some specimens from Los Angeles and Ventura + counties Ochraceous-Orange, especially in young and juveniles. + 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 limbs over antiplantar faces of toes. Least + width of color of underparts averaging, in 15 adult and subadult + males from San Diego County, 54 (35-75) per cent of greatest width + of color of upper parts. Black tip of tail in same series of males + averaging 54.5 (46-60) mm. long. Thus averaging longer than hind + foot and 35 per cent of length of tail-vertebrae. + + _Skull and teeth._--Male (based on 6 adults from San Diego + County). See measurements and plates 22-24. As described in _M. f. + nigriauris_ except that: Weight (4 specimens), 3.9 (3.8-4.0) + grams; basilar length 43.8 (41.9-47.0); postorbital breadth more + 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; anterior margin of tympanic bulla as far + posterior to foramen ovale as width of 2 to 2-1/2 (including I3) + upper incisors; length of tympanic bulla more than length of lower + molar and premolar tooth-row and longer than rostrum; anterior + margin of masseteric fossa below m2. + + Female (based on 4 adults from San Diego County): See + measurements. As described in _M. f. nigriauris_ except that: + Weight, 2.6 (2.2-2.8) grams; basilar length, 40.0 and 40.1; + postorbital breadth more than width of basioccipital measured from + medial margin of one foramen lacerum posterior to its opposite; + length of tympanic bulla more than length of rostrum. + + The skull of the female averages 34 per cent lighter than that of + the average male. + +The skull of the male of _latirostra_, compared with that of +_nigriauris_, is by weight, more than one-fourth lighter, has a lesser +basilar length, a lesser mastoid breadth, a lesser zygomatic breadth +and a narrower M1. In these features no overlap has been observed +between adults from the general vicinities of the type localities of +the two forms. In adult males of _latirostra_ the postorbital breadth, +with one exception, is more than the combined length of P4 and P3 +whereas the reverse is true in adult males of _nigriauris_. Both males +and females of _latirostra_ have a generally smaller skull with +relatively broader interorbital and postorbital parts and the tympanic +bullae are relatively larger, rounder and more inflated. + +Compared with the skull of the male of _pulchra_ that of _latirostra_ +is, by weight, more than one-fourth lighter, has a lesser basilar and +orbitonasal length, lesser zygomatic and mastoid breadth and a more +nearly flat braincase. In these features no overlap has been observed +between adults from the general vicinities of the type localities of +the two subspecies. Also, in _latirostra_ the tympanic bulla is longer +than the rostrum whereas the opposite is true in _pulchra_. The skull +of _latirostra_ is generally smaller and relatively, on the average, +has the preorbital part of the skull deeper and broader with longer +tooth-rows, although with shorter rostrum, while the zygomatic and +mastoid breadths are less. Study of skulls of subadult females of +_pulchra_ indicate that females of _latirostra_ and _pulchra_ differ in +the same fashion as do males. + +_Remarks._--This subspecies long has gone by the name _M. xanthogenys_ +and the type locality was generally supposed to be in the vicinity of +San Diego. This supposition seems to have originated with Merriam's +(1896:25) statement that the type locality was "Southern California, +probably vicinity of San Diego." Nevertheless, as set forth in the +account of _M. f. xanthogenys_ the type specimen concerned now is +thought to have come from much farther north. + +Although 76 Recent specimens are available from southern California, +additional adults are needed to understand the geographic variation +there. _M. f. latirostra_ may be a composite--made up of more than one +geographic race. Specimens from San Diego County differ so much in +relative length of the tail that at one stage in the present study it +was thought that a difference in this respect existed between the +coastal animals and those from farther inland. Material received later +did not wholly substantiate this view and because of the uniformly +small size of all of the skulls from that county, the animals were +later regarded as of the same subspecies. Eventually, even this +supposed common feature proved to be inconstant for an adult male from +Jamacha, no. 7098, of the San Diego Society of Natural History, and +another adult male from San Marcos, no. 8869, collection of Ralph +Ellis, were later examined and found to have skulls as large as those +of average-sized, adult males of _nigriauris_. + +Despite these puzzling local variations, it is established that the +long-tailed weasels of southern California are smaller than those from +farther north. Also, the southern animal averages smaller in weight and +size of skull, and the skull is differently proportioned. Specimens in +series from Los Angeles County definitely are intermediate in size and +shape of skull between _latirostra_ from San Diego County and +_nigriauris_ from, say, Santa Clara County, but definitely more closely +resemble _latirostra_ from San Diego County than they do _nigriauris_. +A skull of a young animal, not here identified to subspecies, from +Potholes, in the Colorado River Valley, 10 miles northeast of Bard, +Imperial County, California, may have closest relationship to _M. f. +latirostra_. Additional comment on this specimen is offered in the +account of _M. f. neomexicana_. + +From the asphalt pits of Rancho La Brea, in Los Angeles County, a total +of 57 skulls have been examined, more than half of which are reasonably +complete. I have been unable to learn whether these came from pits +regarded by students of the deposit as wholly Recent, from pits +regarded as of Pleistocene age, or from both. Suffice to say that only +two specimens were found which could be distinguished from skulls of +the subspecies of weasel living in that area today. + +These two specimens, lent to me by Professor Chester Stock, were with +other skulls received from the Los Angeles Museum of History, Art and +Science and bore identifying numbers as follows: 16/20-27, the anterior +part of the skull of an adult, and 16, the skull posterior to the +cribiform plate of a subadult or possibly young individual. The latter +has a mastoid breadth of 28.0 millimeters, a tympanic bulla 16.1 long +and other measurements in proportion. It is larger than any specimen of +weasel, of any subspecies, seen from California and in the subgenus +_Mustela_ seems to be exceeded in size only by certain individuals of +_M. f. texensis_. _M. f. neomexicana_ attains relatively large size and +comparisons were made with individuals of that subspecies. However, the +young specimen from Rancho La Brea differs from _neomexicana_ in that +the tympanic bullae rise less steeply on the medial sides and the +inferior lip of the external auditory meatus is less developed +laterally. Age considered, the sagittal crest is less developed and the +mastoid processes project more abruptly from the skull. The anterior +part of the skull of the adult, no. 16/20-27 is larger than any +specimen seen of _M. f. latirostra_ or adjoining subspecies, and among +California-taken specimens is equaled in size only by the largest males +of _M. f. munda_ from the northwest coastal district in Mendocino +County. This adult from Rancho La Brea differs from _neomexicana_, sex +and age taken into account, in greater postorbital breadth, lesser +rostral width in comparison with the interorbital breadth, and in +having the temporal ridges at the anterior end of the sagittal crest +spread out into a Y-shaped, rather than a T-shaped, pattern. All these +differences from _neomexicana_ are features of agreement with the +California bridled weasels of the subspecies _latirostra_, +_nigriauris_, and _munda_. The same is true of the characters which set +apart the young specimen from _neomexicana_. In summary: of 57 weasel +skulls examined from the asphalt pits at Rancho La Brea, Los Angeles +County, all but two are indistinguishable from the skulls of the Recent +weasel living in that region today. These two skulls agree in +qualitative characters with animals of the California coastal +subspecies now living from Los Angeles northward to Humboldt County, +but are larger. For the time being these two may be thought of as +giants of the same type of animal inhabiting the Los Angeles region +today. + +Only one of 41 adult and subadult skulls examined for malformation of +the frontal sinuses shows infestation by parasites. + + _Specimens examined._--Total number, 142, listed by counties from + north to south. Unless otherwise indicated specimens are in the + Museum of Vertebrate Zoölogy. + + =California.= _Santa Barbara County_: Rincon Point, 1. _Ventura + County_: Cuyama Valley, 2200 ft., 1[91]; Nordhoff, 3[59]; Santa + Paula, 1[59]; Ventura, 7. _Los Angeles County_: near Owensmouth, + 1[24]; Cahuenga, 1[91]; Llano, 10 mi. E Littlerock, 1; Flint + Ridge, Pasadena, 1[59]; Pasadena, 3; Lankershim, 1[24]; 1 mi. S + Lankershim, 1[24]; Duarte, 1[59]; Covina, 1[59]; Claremont, 1[91]; + El Monte, 4 (2[75], 1[24]); Montebello, 1; Alhambra, 6 (5[2], + 1[91]); El Nogal, 2[8]; Gardena, 1[26]; Palos Verdes Estate, 3; + Rancho La Brea asphalt deposits, 57[70]^{ and }[92]. _San + Bernardino County_: San Bernardino Valley, 1[75]; San Bernardino, + 4 (2[20], 1[91]); Redlands, 2 (1[38]); Bluff Lake, 2 (1[59], + 1[33]). _Riverside County_: West Riverside, 1; Arlington, 800 ft., + 1[17]; 3-1/2 mi. E and 1/2 mi. N Beaumont, 2600 ft., 1; Banning, + 1[91]; Cabazon, 1[91]; San Jacinto Plain, 1[20]; Tahquitz Valley, + 8000 ft., 1; Elsinore, 1[1]. _San Diego County_: Twin Oaks, 1[91]; + San Marcos, 2 (1[87], 1[41]); Escondido, 1; Witch Creek, 1[91]; + Ballena, 1[20]; Santa Ysabel, 3 (2[20], 1[87]); Julián, 1; La + Jolla, 1; Lakeside, 1[91]; El Cajon, 1[91]; El Vido (not found on + map), 1[91]; San Diego, 9 (1[91], 1[20], 1[87], 1[32]); Jamacha, + 2[87]; Chula Vista, 1[20]. + + +=Mustela frenata pulchra= Hall + +Long-tailed Weasel + +Plates 22, 23 and 24 + + _Mustela frenata pulchra_ Hall, Carnegie Instit. Washington Publ. + 473:98, November 20, 1936. + + _Type._--Male, adult, skeleton and skin; no. 16668, Mus. Vert. + Zoöl.; Buttonwillow, Kern County, California; April 30, 1912; + obtained by J. Grinnell; original no. 1953. + + The skull (plates 22-24) is complete and unbroken (a fracture in + the right jugal has healed). All teeth are present and entire. The + skeleton lacks the os penis, left fibula, shaft of left tibia and + the distal three or four caudal vertebrae. Some of the bones of + the feet distal to the radius and tibia are with the skeleton, and + the remainder probably are in the skin. The skin is fairly well + made and in good condition, except for the left hind leg which was + torn when the animal was captured. A well-developed scrotal pouch + shows the specimen to have been a male. + + _Range._--Altitudinally around 300 feet in San Joaquin Valley to + 2500 feet at Isabella; Upper Sonoran and Lower Sonoran life-zones + of southern end of San Joaquin Valley and in mountains at southern + end of Valley, California. See figures 29 and 30 on pages 221 and + 314. + + _Characters for ready recognition._--Differs from _M. f. + nevadensis_ in presence of light facial markings, and from _M. f. + nevadensis_ and _M. f. inyoensis_ in near (16 _j_) Buckthorn Brown + to near (_h_) Yellow Ocher rather than near (14 _n_ to _l_) + Brussels Brown color of upper parts, and greater size with hind + foot more than 40 in females and basilar length averaging more + than 46.0 in males; from _M. f. latirostra_ in having rostrum + (orbitonasal length) longer than tympanic bulla and from _M. f. + latirostra_ and _M. f. nigriauris_ by color of upper parts as + stated above rather than near (_l_) Antique Brown, and by having + inside of ears same color as back rather than much darker than + back; from _M. f. xanthogenys_ in hind foot of males more than 46 + and broader skull which in males has breadth of rostrum more than + 13.9 and mastoid breadth more than 26.0. + + _Description._--_Size._--Male: The type specimen and five other + adults yield average and extreme measurements as follows: Total + length, 454 (428-477); length of tail, 178 (153-184); length of + hind foot, 50 (47-55). Tail averages 65 per cent as long as head + and body. Length of hind foot approximately equal to basal length. + The type specimen measures, 460, 184, 49. + + Female: Three subadult topotypes yield average and extreme + measurements as follows: Total length, 399 (383-411); length of + tail, 154 (140-161); length of hind foot, 42 (41-42). Tail + averages 63 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 + are: Total length, 55; length of tail, 24; length of hind foot, 8. + + _Externals._--As described in _Mustela frenata nigriauris_. + + _Color._--Spot between eyes, band confluent with color of + underparts, on each side of head extending anterodorsally anterior + to each ear, posterior third of each upper lip, lower lips and + chin white or more often darker than Ochraceous-Buff and + therefore same color as belly; dark spot posterior to each angle + of mouth present but small; tip of tail black; remainder of upper + parts near (16 _j_) Buckthorn Brown to near (_h_) Yellow Ocher and + from tone 2 to 4 of Brown Pink of Oberthür and Dauthenay, pl. 297, + but with a trifle more reddish brown. Upper parts of uniform color + except for occasional slight darkening of nose, forehead, and + areas around eyes. Underparts darker (_a_) than Ochraceous-Buff. + 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 over antiplantar faces of toes, tarsal + region and sometimes in diluted fashion on proximal third of + underside of tail. Least width of color of underparts averaging, + in 6 male topotypes, 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., type 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 3-1/2 (including I3) 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 m1, M1, P4, and depth 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 distinguished from those of +_nigriauris_ by the combination of the following mentioned, average +differences: Tympanic bullae larger in each of three dimensions; +preorbital and interorbital parts of skull broader and notably heavier; +interorbital breadth greater; zygomatic 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 Co., Calif.), skulls of adult males of _pulchra_ +average larger in every measurement taken and no overlap exists in +basilar length, orbitonasal length, mastoid breadth, zygomatic breadth, +length of tympanic bulla, or depth of skull at either the anterior +margin of the basioccipital or at the posterior margins of the upper +molars. Relatively, 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 collections of the United States +Bureau of Biological Survey, another by J. Grinnell for the Museum of +Vertebrate Zoölogy in 1912, and in 1933, another by L. M. Huey, for the +San Diego Society of Natural 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 Coalinga, 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 unsexed +and without external measurements. Skulls of adults from Coalinga are +needed to permit of more positive identification of the subspecies +found there. The female from 4 miles east of Coalinga, taken on +February 21, 1936, is in process of molt on the underparts, and the +longer hair which is near (20´) Naples Yellow contrasts strongly with +the incoming shorter hair which is near (10 _c_) Salmon-Orange. The +skin alone, no. 16270, Mus. Vert. Zoöl., from Isabella, was made up +from a decayed animal and is of but little use. It is referred to +_pulchra_ purely because of geographic nearness 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 unlike _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 Vertebrate Zoölogy. + + =California.= _Fresno County_: Coalinga, 1[23]; 4 mi. E Coalinga, + 1; 3 mi. S Coalinga, 1[8]. _Tulare County_: Alila (= Earlimart), + 2[91]. _Kern County_: Delano, 1[91]; Buttonwillow, 9 (6[8], + 2[91]); Isabella, 1; Willow Spring, 1[91] _San Luis Obispo + County_: 2 mi. SW Simmler, 1[87]. + + +=Mustela frenata inyoensis= Hall + +Long-tailed Weasel + +Plates 22, 23 and 24 + + _Mustela frenata 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. Zoöl.; 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, California. 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 (_l_) Brussels Brown rather than near (16 _j_) + Buckthorn Brown to near (_h_) Yellow Ocher color of upper parts + and basilar length of less than 45 in males; from _M. f. + latirostra_ 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 + following 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 lips and in some specimens + part or all of upper lips 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 Oberthür + and Dauthenay, pl. 301; slightly darker brown on forehead, nose + and about eyes. In winter near (_j_) 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 +difference 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 the skull. Comparison of the skull with that of _M. f. +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. f. inyoensis_ as now known may be thought of as closely similar to +_M. f. 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. +nigriauris_. 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 southwestward +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 that the 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 County_: Alvord, 4000 ft., 1 (U. S. Nat. + Mus.); 2 mi. N Independence, 1 (Mus. Vert. Zoöl.); 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 neomexicanus_ 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, December 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. Zoöl.; + Armstrong's Lake, Mesilla Park, Dona Ana County, New Mexico; + February 1, 1898; obtained by A. C. Tryson; original no. 58 of C. + M. Barber. + + 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 locality) to 9000 feet (Cloudcroft, + N. Mex.); Upper Sonoran and Lower Sonoran life-zones of northern + México, 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 postpalatal length; from _M. f. + 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 than 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. f. + longicauda_ by Buckthorn 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. f. 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 than 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 length, 81; length + of tail, 40; length of hind foot, 7. + + Compared with _M. f. 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 the "long bristles of the upper lip" + are white as pointed out by Barber and Cockerell (_op. cit._: + 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 anterior to each ear, confluent with white spot + between eyes and with color of underparts; posterior half or all + of each upper lip 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 Oberthür and Dauthenay; 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, pl. 297, of Oberthür and Dauthenay); underparts + Antimony Yellow or near (_c_) Warm Buff of Ridgway, and Brown Pink + (tone 1, pl. 297, of Oberthür and Dauthenay); color of underparts + extends distally on legs over forefeet and hind feet. Least width + of color of underparts averaging 46 (41-55) 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. f. frenata_, _neomexicana_ differs in: upper parts and + underparts much lighter colored; white facial markings more + extensive; color of underparts more extended onto feet. From _M. + f. leucoparia_, _neomexicana_ differs as follows: above and below, + much lighter 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 Berino, 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 than distance between condylar foramen and M1 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 lighter than that of + the male. + +As compared with the skull of the male of _M. f. frenata_, that of +_neomexicana_ is decidedly more angular and ridged. The postorbital +constriction is narrower, the mastoid breadth greater (it is less than +the postpalatal length in some subadult males), the sagittal crest much +higher with impressions of the temporal and masseter 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 addition 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 concave as in _neomexicana_. Compared with _M. f. +longicauda_, _neomexicana_ is relatively narrower in the interorbital +region, has relatively shorter tooth-rows, a V-shaped rather than a +U-shaped interpterygoid 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 difference in relative length of tooth-row is less. The same +differences exist also in comparison with _M. f. arizonensis_ except +that its interorbital breadth, relative to the rest of the skull, is +about the same. Difference in size is especially marked here; even +females of _neomexicana_ 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 recently 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 Kansas. +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._:198) 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 +Langtry, 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. The +large size (basilar length of 46.5 mm.) may reflect some relationship +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 subspecies to the list of mammals +of Texas is made certain by the female (no. 1572, Texas Cooperative +Research Collection) taken on July 28, 1940, 1-1/2 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 showing the approach, if any, +in color, of _neomexicana_ to _M. f. leucoparia_. + +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. f. frenata_ is not shown by the material +at hand. The two females from Albuquerque, although typically +_neomexicana_ as regards color, have smaller, less prominently ridged +skulls than females of _neomexicana_ of the same age from farther south +and approach _M. f. nevadensis_. + +Probably the geographic ranges of _M. f. neomexicana_ and _M. f. +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 California, which intervenes between the ranges of the two +subspecies is the skull of a young individual, no. 68842, Mus. Vert. +Zoöl., from 10 miles northeast of Bard, Imperial County, California. +There, on December 29, 1932, Jack C. vonBloeker, Jr., retrieved the +weathered skull with some of the vertebrae attached, from the top of a +wood rat'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 specimen itself for +the tympanic bullae are larger than in _nevadensis_ and the skull is +larger than the largest males seen of _arizonensis_, the two upriver +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 conceals any other diagnostic +cranial features, and prevents referring it certainly to either +_neomexicana_ or _latirostra_. In any event the 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 member of an unnamed race resident in the +lower part of the valley of the Colorado River. + +Whereas _M. f. panamensis_ and _M. f. aureoventris_ are the +darkest-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 specimen was +shot in the grass on the shore of Armstrong's Lake. . . ." Bailey +(1928:97) found the tracks of one of these animals "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 County_: Safford, 1. _Cochise County_: 2 mi. W + Willcox, 1; Willcox, 1; 8000 ft., Chiricahua Mts., 1; 6000 ft., + Pinery Canyon, Chiricahua Mts., 1[33]; Tombstone, 1; Sulphur + Spring Valley, 1[74]. + + =Durango.= "Durango City," 1. + + =Kansas.= _Seward County_: Liberal, 1[93]. + + =New Mexico.= _Bernalillo County_: 3 mi. NW Albuquerque, 2. + _Lincoln County_: 7800 ft., South Fork Eagle Creek, White Mts., 1. + _Chaves County_: Pecos River, 10 mi. E Roswell, 8[74]; Dexter, + 1[74]. _Otero County_: Cloudcroft, 9000 ft., 1[90]. _Dona Ana + County_: Mesilla Park, 2 (1[75], 1[7]); Berino, 2. + + =Texas.= _Culberson County_: 1-1/2 mi. NW Kent, 1[90]. _Terrel + County_: 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). + + + _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 between posterior borders of P4 and P2; + from _M. f. 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. Zoöl., 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 masseteric 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 Collection, 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 preservation 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 central +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 correspondingly large. +The skin alone, no. 427, from near Waco, Texas, has, 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, +México. 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 +_macrophonius_ on the one hand and _perotae_, _frenata_ and _texensis_ +on the other, has relatively uniform color, color pattern and body +proportions over a large region but at its northern extremity develops +a "giant" population, _M. f. macrophonius_ and _M. f. texensis_, +respectively. The skull of the type specimen of _M. f. texensis_ is the +largest 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. f. +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 between 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 probably +does take place with _M. f. neomexicana_ and possibly with _M. f. +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 County_: 5 mi. N Waco, 1[3]; Erath, 1[3]. + _Gillespie County_: Fredericksburg, 1[90]. _Kerr County_: 4[75]; + 1[2]; and 2[90] trade skins. + + +=Mustela frenata frenata= Lichtenstein + +Long-tailed Weasel + +Plates 1, 22, 23, 24, 36, 37, 38 and 40 + + _Mustela frenata_ Lichtenstein, Darstellung neuer oder wenig + bekannter Säugethier, 1832, pl. 42, and corresponding text, + unpaged; Seton, Lives of game animals, 2:576, 1929. + + _Mustela brasiliensis_ Sevastianoff, Mem. de l'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. + Zoöl. Soc. London, 1865:114. + + _Putorius frenatus_, Baird Mamms. N. Amer., p. 173, 1858; Merriam, + N. Amer. Fauna, 11:26, pl. 3, figs. 1, 1a, 1b, 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] preoccupied 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 + Matamoros)"]. + + _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., México City, México; 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, México); from southern Texas as far south as México + City; Lower Sonoran to at least Transition life-zone. See figure + 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. perotae_ + in nonextension 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 wider (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. f. arthuri_ by white facial markings and + postorbital breadth less than distance between posterior borders + of P4 and P2; from _M. f. tropicalis_ by nonextension 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 + Brownsville, 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). 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, México, are: 505, 203, 53. Another adult + male, from Miquihuana, Tamaulipas, México, measures: 520, 215, 52. + + Female: Six adults, subadults and young 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 (with 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 hind 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 + underparts, on each side of head extending anterodorsally anterior + to each ear, and posterior two-thirds to one-half of each upper + lip, 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 + extends 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 M1 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 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 3 or 4 (including I3) 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 to young) 43.3 (41.3-47.3); zygomatic + breadth more or less than distance between condylar foramen and M1 + 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 bulla as far posterior to foramen + ovale as width of 2 to 3-1/2 (including I3) 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 lighter than that of + the average 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. f. 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 brasiliensis_' to a +weasel brought to St. Petersburg by Capt. A. J. Krusenstern 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 applied the name _Mustela frenata_ +to the weasels of the vicinity of México City and that name was used +for bridled weasels from México and the southwestern United States by +most subsequent German writers and by several Americans. In 1896 +Merriam (1896:27) showed that Sevastianoff's _Mustela brasiliensis_, +1813, although probably the same as _Mustela frenata_, was preoccupied +by Gmelin's _Mustela brasiliensis_, 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 México south to +México 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 specimen of +Sevastianoff's _Mustela brasiliensis_. + +The geographic range of this subspecies is relatively large and, as +might therefore be expected, specimens show geographic variation. 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 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 M1, 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 approach 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 Brownsville, 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 resembles +_leucoparia_. Better material from the western part of the state of +México may show the range of _leucoparia_ to extend eastward 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 this 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 +difference noted, therefore, seems to be of geographic significance. +However, there is from Alvarez, San Luis Potosí, approximately 115 +miles south of Miquihuana, a young (nearly subadult) female, no. 21968, +which is as light colored as specimens from Brownsville, Texas, or +Tlalpam, México. The only distinctive feature of this specimen is the +much greater extent of its white facial markings; they are more +extensive even than in the specimen from Miquihuana. + +Finally, the series from Brownsville, Texas, indicates that the animal +there is smaller than _frenata_ from the vicinity of México (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 +Brownsville. The similarities between specimens from the two +localities, Tlalpam and Brownsville, are striking; 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 localities 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. f. arthuri_ but it is supposed that frenata +intergrades with each of these subspecies. The difference 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 locality 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, Beeville, 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, permits +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 measurements 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 enables the student the better +to evaluate cranial characters of nearby 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 connection 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 specimens and on one +side only in three other specimens. + +In eleven specimens from México, 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 specimen. 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 nineteen 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 present 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, [F], three weeks +old, taken on February 15; no. 17318/24239, [M], four weeks old, taken +on March 16; no. 45899, [F], forty days old, taken on May 21; no. +21778/36481, [M], 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: 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 +within the chaparral and cactus lands, where Wood Rats, Rabbits 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 México by states, from north to south. Unless otherwise + indicated specimens are in the collection of the United States + National Museum. + + =Texas.= _Bexar County_: San Antonio, 2 (1[2]). _Goliad County_: + Charco, 1. _Nueces County_: Corpus Christi, 1[2]. _San Diego + County_ (not found), 1. _Hidalgo County_: La Hacienda, 1. _Duval + County_: San Diego, 2[7]. _County_ in question: Lower Rio Grande, + 1. _Cameron County_: Brownsville, 34 (3[2], 4[1], 3[93], 2[75], + 1[59], 1[60], 1[4]); no locality more definite than county, 2. + + =Nuevo León.= Río Ramis, 20 mi. NW Montemorelos, 1[90]. + + =Tamaulipas.= Matamoros, 6; Miquihuana (now in Nuevo León), 1[75]. + + =San Luis Potosí.= Alvarez, 1[75]. + + =México=: Region montagneuse des environs de Toluca, Nevada + Toluca, 3200 M., 1[84] + + =Distrito Federal.= City of México, 2 (1[4]); Tlalpam, 2. No + locality more definite than México, 4 (1[4], 3[7]). + + +=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.; Pátzcuaro, Michoacán, México; July 27, + 1892; obtained by E. W. Nelson; original no. 2960. + + The skull (plates 29 and 30) lacks most of the braincase; a + fragment, consisting 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 differs 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 + México (city) in Michoacán and Nayarit. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. 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 ventrally + 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 halfway to shoulders and by narrower (less + than 7.8) tympanic bullae; from _M. f. 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 Pátzcuaro, Michoacán, yield average and extreme + measurements as follows: 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 + Pátzcuaro, Michoacán, measure, respectively, as follows: Total + length, 412, 400; length of tail, 159, 159; length 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 differences in external measurements of the two sexes + are: Total length, 108; length of tail, 47; length of hind foot, + 13. + + _Mustela 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 anterior 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; 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 width of color of underparts averaging, in eight specimens, + 54 (extremes 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 averaging 25 per cent of length of + tail-vertebrae. + + As compared with _M. f. frenata_ 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 masseteric 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 M1, 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 subspecies. As +compared with that of _frenata_ the main difference is the less +inflated tympanic bulla, the height of which is approximately 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 Río 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 México 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). The five specimens from Michoacán were taken +by Nelson or Nelson 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." +Although the form is not strongly marked, the two additional specimens +from Michoacán and better comparative material than Merriam had confirm +several of the differential 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 Artenkiki (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. f. frenata_. Unlike _frenata_, +_leucoparia_ 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. f. frenata_ and _M. +f. goldmani_. The latter subspecies has the least inflated, narrowest +and least projecting tympanic bullae of the three. The black color of +the head extends, on the average, farther posteriorly than in _M. f. +frenata_ but not so far as in _M. f. goldmani_. The general color, too, +is intermediate between that of _M. f. frenata_ and that of the much +darker _M. f. goldmani_. The white facial markings are more extensive +than in either _M. f. frenata_ or _M. f. goldmani_. 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. f. neomexicana_, whose range possibly +meets that of _M. f. leucoparia_, also has more extensive white facial +markings than _M. f. frenata_ but less extensive markings than _M. f. +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 northern part of +the range of _leucoparia_ closely resemble _frenata_ in color, +constitute sufficient evidence for regarding the two as only +subspecifically distinct. The female, no. 26153 from Artenkiki, 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. f. 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 México. + Localities are listed by states from north to south. Specimens + from Michoacán are in the United States National Museum; one from + Río Santiago is in the British Museum of Natural History; all + others are in the American Museum of Natural History. + + =Nayarit.= Tepic, 1. + + =Jalisco.= Río Santiago, 1; Los Masos, 1; "Artenkiki" (J. A. + Allen, 1906, p. 238, writes "Artenkikil" and, on p. 259, + "Artenkiki."), 1. + + =Michoacán.= Zamora, 1; Los Reyes, 1; Pátzcuaro, 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, 11: 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, + México; 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 + (Popocatépetl), Upper Sonoran, Transition and Boreal life-zones of + mountains along Puebla-México boundary, eastward to western + central Veracruz and south into Oaxaca. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from _M. f. 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 width 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 grams); postorbital breadth + less than combined length of upper premolars; m1 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 antipalmar face of + forefoot, least width of color of underparts 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 extends 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 M1 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 M1 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 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 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. f. tropicalis_ is made in the +account of that subspecies. Compared with the skull of _M. f._ +_macrophonius_, that of the female of _perotae_ is more flattened, has +the longitudinal dorsal outline distinctly concave rather than flat +just behind the postorbital processes, and much wider tympanic bullae. +Accordingly, the basioccipital is slightly 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. f. 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. f. frenata_, the skull of the female of +_perotae_ differs mainly in the lesser inflation 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], pl. 3, fig. 2) as _Putorius frenatus_. The referred +nontypical specimen from Cerro San Felipe, 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 +published references to this subspecies, or specimens of it, have been +seen. In 1941 and 1942, W. B. Davis and associates took four specimens +along the boundary between the states of Puebla and México. + +Although the specimen from Cerro San Felipe, Oaxaca, is referred to +_Mustela frenata perotae_, to the description of which it answers best, +that specimen, on account of its structural characters and geographic +position relative to adjacent races, is in reality an intergrade +between several of the adjacent races. Some of its intermediate +characters are pointed out in the discussion of _M. f. goldmani_. In +the specimens from 45 and 55 kilometers ESE of México (city) the black +color of the top of the head does not extend so far behind the ears as +in the holotype of _M. f. perotae_ and in this feature the two +specimens show intergradation between the two subspecies, _perotae_ and +_frenata_. + +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 normal molt. + +As would be expected from its geographic position, _M. f. perotae_ +resembles _M. f. frenata_ of northern México and the high mountain +forms of southern México 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 supralacrymal 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 intermediate in degree between the condition in +_M. f. macrophonius_ and that in _M. f. frenata_. Thus _M. f. perotae_ +combines several characters of _M. f. 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, _Mustela +frenata tropicalis_. Between _M. f. perotae_ and _M. f. tropicalis_ +there is marked differentiation in the skulls with much less +differentiation in coloration. The differences in typical skulls of the +two subspecies 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._:31) are a topotype +of _M. f. tropicalis_ from Jico and the one which now is the type +specimen of _M. f. perotae_. They show the great difference in size and +proportions and are females of comparable ages, not of different ages +as I suspected before examining the skulls. However, despite 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 connection with _M. f. +tropicalis_ (see p. 366). + +None of the seven skulls shows marked deformity of the interorbital +region, but two of the three adults appear to have had these parts +infested with nematodes. + + _Specimens examined._--Total number, 7, all from México, listed by + localities from north to south. Specimens from Veracruz and Oaxaca + in the United States National Museum; remainder in Texas + Cooperative Research Collection. + + =México=: Monte Río Frío, 45 Km. ESE México City, 1; 55 Km. ESE + México City, 1; N slope Mt. Popocatépetl, 13,555 ft., 1. + + =Puebla.= Río 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, México; February 10, 1896; + obtained by E. A. Goldman (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 the 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 I2 and I3, right and left i3, and the medial parts + of the paraconid and protoconid of right m1 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 represented 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 Tecpám, Guatemala), Upper Tropical Life-zone of + mountains and western coasts of southern México, Guatemala and + Salvador. See figure 29 on page 221. + + _Characters for ready recognition_ (characters based on + males).--Differs from _M. f. nicaraguae_ and _M. f. 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. f. 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 Brown or darker rather than + Brussels Brown; nonextension of color of underparts onto hind + feet; from _M. f. 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 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 measurements 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 + slightly 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 underparts near (16' _c_) Ochraceous-Buff; 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 underparts, in five adult males, averaging + 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 M1 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 premolars + 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 tympanic 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 immediately behind + m2. + + Female: Typical skull unknown. + +Comparison of male skull with that of _M. f. perda_ made in discussion +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_. + +As compared with the skull of the male of _M. f. 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. f. 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 M1 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 _goldmani_ is shorter and relatively as well as actually +narrower and lighter 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 mastoid +breadths or length or height of tympanic bullae. In depth, the skull of +_goldmani_ averages actually and relatively greater. Its teeth are +smaller. The squamosal anterior to each tympanic bulla is convex +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 important diagnostic +characters, which additional specimens show pertain 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. f. macrophonius_. The altitude at which the two specimens 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 Dueñas, 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. f. +leucoparia_ or _macrophonius_. As compared with _M. f. frenata_, +_goldmani_ is longer, has an actually as well as relatively shorter +tail, and a much longer hind foot. + +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 southeast of +Teopisca, taken on May 12, the molt is well advanced. Another 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 underparts 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 specimen +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 characters. The +zygomatic breadth is less than, rather than more than, or equal to, the +distance between the condylar foramen and M1 or than that between the +anterior palatine foramen and the anterior margin of the tympanic +bulla. This difference appears to be correlated 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_, _leucoparia_ and +possibly _frenata_, there is the specimen from Cerro San Felipe, +Oaxaca. The degree of restriction of the color of the underparts is +intermediate 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 immediately +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 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 eastern 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 specimen 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. f. 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 +lowland races. + +These four specimens indicate that the lowland population on the +western side of the divide is smaller than the mountain population. 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 pattern of geographic variation in +weasels of this region that will require 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 end +of the tympanic bulla. Considering the far eastern location of Puebla +(just north of Río 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 elevation 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 remains 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 infested +with parasites. + + _Specimens examined._--Total number, 15, listed by localities from + north to south, and unless otherwise indicated in the American + Museum of Natural History. + + =México=: _Chiapas_: 20 mi. SE Teopisca, 2[91]; Pinabete, 1[91]. + + =Guatemala=: Puebla, 1; Finca Porvenir, 3500 ft., S slope Volcan + Tajumulco, 1[60]; Sierra [=? Cerro] Santa Elena, 9500 ft. (near + Tecpám), 1[60]; Carolina, 1; Volcano San Lucas, 1; "Finca El + Cipres," 1; "Finca Cipres," 2500 ft., 1; Finca San Isidro, San + Sebastión, Dept. Retalhuleu, 1[4]; Dueñas, 1[7]; no locality more + definite than Guatemala, 1[7]. + + =El Salvador=: Los Esesmiles, 8000 ft., Chalatenango, 1[59]; La + Cebia, 2150 ft., near San Salvador, 1[59]. + + +=Mustela frenata macrophonius= (Elliot) + +Long-tailed Weasel + +Plates 24, 25, 26, 30, 37, 38 and 39 + + _Putorius macrophonius_ Elliot, Proc. Biol. Soc. Washington, + 18:235, December 9, 1905. + + _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 skin; no. 14063, Field Mus. Nat. + Hist.; Achotal, Veracruz, México; 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 life-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. f. 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. f. + goldmani_ by larger size of skull (see above) and entire animal + and wider tympanic bullae; from _M. f. tropicalis_ and _M. f. + 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 female from Pérez, Veracruz, shows this sex to be + correspondingly large. Because the skin is understuffed and + because the hind feet are skinned out, reliable 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-soles 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 width 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 M1 or that between anterior palatine foramen and + anterior 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 incisors; 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 M1 and more or less than (in the + single specimen, equal to) that between 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 lighter than that of the + male. + +Comparison of the skull with that of _M. f. 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. tropicalis_ and _perda_, the skull of the male of +_macrophonius_ is larger in every measurement taken. The postorbital +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 Pérez for the collection of the United +States Bureau of Biological Survey. These specimens were tentatively +referred to _Mustela tropicalis_. In the following January, Edmund +Heller and Charles M. Barber obtained the adult male that was made the +type specimen by Elliot who did not see, or if he did, did not mention, +the specimens from Pérez. He did, however, refer a young female from +Xuchil, Veracruz, to his _Putorius macrophonius_. This young female is +here referred to _Mustela 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 the type specimen of _M. f. +texensis_. The other cranial measurements taken are greater in _M. f. +texensis_. The skull of the female from Pérez 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 understuffed 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 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 the 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. f. macrophonius_ most closely resembles _M. f. goldmani_ but in the +relatively flattened braincase, deep constriction of the postorbital +region and general angularity of the skull approaches _M. f. perotae_ +and _M. f. 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 México, listed by + localities from north to south. + + =Veracruz.= Achotal, 1 (Field Mus. Nat. Hist.); Pérez, 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, pl. 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, México; 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, México. 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. f. 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. f. macrophonius_ by lesser basilar length (not + more than 48) and in postorbital breadth exceeding length of upper + molar and premolar tooth-row. + + _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 vibrissae do not reach apical pad of fifth digit and + hairiness of foot soles is less. + + _Color._--As described in _M. f. frenata_ except that: Blackish of + head extends half way or more from ears to shoulders; upper parts + near (14) Brussels Brown or slightly faded tone 2 of Maroon of + Oberthür and Dauthenay, pl. 341; underparts 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; thus 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 slightly darker; ventral side of tail noticeably + darker; color of underparts more restricted, averaging + 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 slightly more extensive; blackish of head less extended + over neck; upper parts markedly lighter; 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 M1 + 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 I3) 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 tympanic bulla; least + width of palate more than greatest length of P4; height of + 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. f. +perda_, the nearest relative, has been made in the discussion of that +subspecies. Some of the features that readily distinguish skulls of _M. +f. tropicalis_ from those of _M. f. 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 _macrophonius_ +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. f. tropicalis_ is smaller, weighing less +than 3 grams; basilar length less than 41; postorbital breadth more +than 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 synonymy 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 México 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. f. 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 México City, are +assumed to be typical. The latter specimen certainly came from an +elevation lower than that of México City because _M. f. 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 underparts. There are three specimens labeled +as from Orizaba that indicate intergradation with _perotae_ as does +also the coloration of the juvenal female from 5 kilometers north of +Jalapa. The specimens 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. f. +perotae_. Merriam called attention to the great difference in size +between the skulls of the two sexes of _M. f. 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 juvenal female of _M. f. 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. f. perotae_ as +well as with _M. f. perda_. _M. f. tropicalis_ differs from _M. f. +perotae_ and _M. f. frenata_ in about the same way that _M. f. perda_ +differs from _M. f. goldmani_ and _M. f. macrophonius_. _M. f. +tropicalis_ and _perda_ each is smaller and more intensely colored than +_goldmani_ 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 México, listed + by localities from north to south. Unless otherwise indicated + specimens are in the collection of the United States National + Museum. + + =Tamaulipas.= 50 mi. S Victoria, 1[71]. Near? México City, 1. + + =Veracruz.= Jico, 3; 5 km. N Jalapa, 1[90]; Jalapa, 2 (1[2], + 1[75]); Xuchil, 1[60]; Orizaba, 4 (2[75], 1[4]). + + +=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. + + _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, México; 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 (Chichén Itzá) to 4000 feet (San Vicente) in + Lower Tropical Life-zone south from southern Veracruz through + southern México 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 zygomatic + breadth more than 27.4); from _M. f. 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. f. 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 Oberthür + and Dauthenay, pl. 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 (extremes 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 M1 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 opposite; 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 4-1/2 to 5-1/2 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 + measurements 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 M1 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. f. nicaraguae_ has +been made in the account of that subspecies. The skull of the male as +compared with that of _M. f. tropicalis_ has shorter tympanic bullae, +deeper braincase at anterior margin of basioccipital, lesser zygomatic +and palatal breadth and smaller P4 and m1. The skull of the female is +larger in every measurement taken except those reflecting width of the +preorbital portion. This part is actually narrower 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. f. goldmani_ and show no overlap are: +lesser basilar length, lesser weight, greater relative length of upper +tooth-rows, greater relative 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 +posterior margin of M1. Features in which _perda_ averages less are: +length of tooth-rows, interorbital breadth, orbitonasal length, +relative zygomatic breadth, length of m1, outside and inside lengths of +P4, width and length of M1, and depth of skull at posterior margin of +M1. Features in which _perda_ averages more than _goldmani_ are: +relative interorbital breadth, relative mastoid breadth and depth of +skull at anterior margin of basioccipital. The length of the inner +half of M1 averages the same. As compared with _goldmani_, 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 immediately 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. goldmani_ of the higher mountains just as _M. +f. tropicalis_ corresponds to _M. f. 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. f. +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. f. panamensis_ +among Central American weasels. Evidence of intergradation with +_goldmani_ is furnished by the specimens from Cobán, Guatemala, and the +nearby locality San Cristóbal in Verapaz, Guatemala. Reduced size as +compared with _goldmani_ suggests affinity with _perda_ but the greater +width of the light-colored underparts, which averages 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 Cristóbal 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 Cristóbal (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[7]). + + =Chiapas.= San Cristóbal, 1; San Vicente, 1; no locality more + definite than state, 1. + + =Yucatán.= Chichén-Itzá, 1[76]. + + =Guatemala=: Cobán, 2 (1[7], 1[4]); Finca la Providenci, S. + Cristóbal, Verapaz, 3[76]; central Guatemala, 1; no locality more + definite than Guatemala, 5 (2[7]). + + +=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 unbroken. 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. f. + 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 underparts (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 that shown in figures 20 and + 21) is less, slightly less even than in _M. f. perda_. + + _Color._--As described in _Mustela frenata goldmani_ except that: + Back near (_n_) Argus Brown, or Carbon Brown, tone 4 of Oberthür + and Dauthenay, pl. 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 adult + 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 width of four to five upper + incisors; length of tympanic 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. f. costaricensis_ +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 shallower 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. f. perda_ and relatively narrower. + +_Remarks._--When naming this form, Allen (1916:100) characterized it as +"Similar to _M. tropicalis tropicalis_ but general coloration much +darker and the white face markings somewhat reduced in area." In the +sentence preceding the one quoted, _Putorius tropicalis perdus_ was +placed as a synonym of _Putorius tropicalis tropicalis_. _M. f. +nicaraguae_ and _M. f. perda_ are nearly alike in color and color +pattern but differ in cranial characters. _M. f. perda_ and _M. f. +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. f. goldmani_, +_macrophonius_, _perotae_ and _frenata_. + +The weasels from Honduras definitely are not typical of _nicaraguae_ 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 outline of the skull is more +nearly flat (less convex) in some. In these and several other +differential features studied, the average of specimens from +Tegucigalpa is intermediate toward _goldmani_, but everything +considered the animals seem best placed with _nicaraguae_ rather than +with _goldmani_ or _perda_, to which latter also, they show some +resemblance. With better material from Nicaragua 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 _goldmani_ should be 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 upper 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. +costaricensis_ 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. f. 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. f. costaricensis_. + +_M. f. nicaraguae_ in typical form, then, is thought of as a small, +lowland, tropical subspecies only slightly differentiated from _M. f. +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. f. 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[75]; Comayagüela, 1[75]; vicinity of Tegucigalpa, 2; no + locality more definite than Honduras, 1[4]. + + =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; Lönnberg, Arkiv för Zool., 14(no. + 4):16, 1921. + + _Mustela frenata costaricensis_, Hall, Carnegie Instit. Washington + Publ. 473:109, November 20, 1936. + + + _Type._--Male, young, skull and skin; no. 13770/37149, U. S. Nat. + Mus.; San José, 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-thirds of the tail. The head is somewhat shrunken. + The color is possibly faded but if so only to a slight degree. + Otherwise, the skin is in good condition. 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 Oberthür and Dauthenay, pl. 344) and + longitudinally flat interorbital region of skull; from _M. f. + nicaraguae_ in darker color of upper parts (of Oberthür and + Dauthenay, tone 2 of pl. 344 rather than tone 4 of pl. 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 subadult from Irazú, 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. f. 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 Oberthür and + Dauthenay, pl. 344; chin, lips, 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 specimens, 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 José and no. 11408, U. S. Nat. Mus., from "Costa + Rica"): See measurements and plates 25-30; weight, 5.9 grams; + basilar length 49 +; zygomatic breadth more than distance between + condylar foramen and M1 or than between anterior palatine foramen + and anterior margin of tympanic bulla; mastoid breadth less than + postpalatal length; postorbital breadth in undiseased skulls less + than length of upper premolars (less than distance between + posterior borders of P2 and P4) and 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 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 anterior 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. f. 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 braincase. Indeed the skull +is more like that of _M. f. 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 affinis_. 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. f. goldmani_ which occur to the northward and from _M. f. +panamensis_ which occurs to the southward and is a large, +heavy-skulled, dark-colored animal with white facial markings +restricted or absent. 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 nasofrontal spot, 10 x 12 mm. The young female from Cachí +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 the upper parts is only a +little less dark than those of _M. f. panamensis_. Owing 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. f. costaricensis_ is a large animal and among its geographic +neighbors is approached in size only by a specimen of _panamensis_ from +Boquete, Panamá. Also the young male from Cervantes suggests +_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 José, is suggestive of +_M. f. 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. f. 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=: Irazú (Frasu or Irasu on label), 3000 M., 1[4]; + Cervantes, 1[2]; San José, 1[91]; vicinity of San José, 2[7]; + Azahar Cartago, 1[78]; Tucurrique, 1[7]; Cachí, 1[7]; El Muñco [= + Muñeco?] (Río Nivarro [= Navarro?]), 4000 ft., 10 mi. S Cartago, + Caribbean Slope, 1[76]; Navarro, 1[91]; Candelaria Mts., 1[75]; no + locality more definite than Costa Rica, 3[91]. + + +=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. Zoöl., 39:49, April, + 1902; Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, + 1914. + + _Mustela affinis_, Goldman, Proc. Biol. Soc. Washington, 25:10, + January 23, 1912; Hollister, 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.; Río Indio, Canal Zone, near Gatún, + Panamá; February 17, 1911; obtained by E. A. Goldman; original no. + 20897. + + The skull is complete and unbroken. The left lower incisor is + broken off 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 + Panamá. See figure 29 on page 221. + + _Characters for ready recognition._--Differs from both _M. f. + meridana_ and _M. f. costaricensis_ in darker tone (tone 4 of + Oberthür and Dauthenay, pl. 344) of color of upper parts and in + convex dorsal outline of skull (Compare figures of mentioned + subspecies on plates 25-27). + + _Description._--_Size._--Male: Two adults from Boquete in the + Museum of Comparative Zoölogy, 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 Chiriquí, 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 specimen 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 differences 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 lip 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, pl. 344 of Oberthür and Dauthenay; chin + and lips, whitish; remainder of underparts Warm Buff or near (16´ + _c_) Ochraceous-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 underparts, 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 M1 or than between anterior + palatine foramen and anterior margin of tympanic bulla; mastoid + breadth less than postpalatal length; postorbital breadth more + than length of upper premolars and more 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 approximately 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 4-1/2 to 5-1/2 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 masseteric + 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 M1 and more or less than (about equal + to) that between anterior palatine foramen and anterior margin of + tympanic bulla; postorbital breadth more than combined 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 Chiriquí is 58 per cent lighter +than the average of the two adult males. + +The skull of the male of _M. f. 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, orbitonasal +length, interorbital breadth and zygomatic breadth averaging narrower. +Mastoid breadth always narrower. Tympanic bullae longer, narrower, and +usually slightly less protruded. P4 and m1 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 developed. Differences between skulls of +females, in so far as known, similar to those described between males. + +As compared with _M. f. 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, M1, and m1 larger. Dorsal outline 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. + +_Remarks._--This subspecies had not been recognized by previous workers +because specimens from Panamá were supposed to be _Mustela affinis_ +Gray up until 1916, when Allen (1916:100) restricted the type locality +of _M. affinis_ to Bogotá, Colombia. At that time Allen referred +specimens from Panamá to _Mustela affinis costaricensis_, 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. f. +costaricensis_ in size more than do those from farther 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 variation 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 females 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 specimen 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 +Panamá weasel that is strikingly like that of a mink. _M. f. +panamensis_ is one of the two blackest weasels; _M. f. aureoventris_ is +the other. Each of these subspecies occurs in a region of heavy +rainfall and there clearly is a positive correlation 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 subspecies of the species. + +Adequate specimens from central and southern Panamá may reveal the +existence of one or more additional subspecies since animals from each +of the three localities now represented differ from those from the +other two and some of these differences are correlated with geographic +position. However, specimens from all three 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 differentiated from +any other American weasel. Nevertheless, _M. f. panamensis_ is clearly +a link between the North and South American subspecies and _panamensis_ +intergrades with the adjacent subspecies. 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. f. costaricensis_. +The smaller size of skull and teeth of no. 178970 from Mt. Pirre are +points of resemblance to _M. f. meridana_. + +The type specimen was selected from a region where _M. f. panamensis_ +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 +characters 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. + + =Panamá=: Boquete, 10 (3[75], 1[8], 1[2], 3[4], 1[7]); Río Gariche + [é], 5300 ft., 1[1]; Siola, 1[1]; Chiriquí, 1[7]; Río Indio, near + Gatún, 1; Mt. Pirre, 3 (2[1]); Calovébora, 1[7] (locality not + found, possibly misspelling of Calovébora); no locality more + definite than Panamá, 1[4]. + + +=Mustela frenata meridana= Hollister + +Long-tailed Weasel + +Plates 25, 26, 27, 37, 38 and 39 + + _Mustela meridana_ Hollister, Proc. Biol. Soc. Washington, 28:143, + July 10, 1914. + + _Putorius affinis_, Robinson and Lyon, Proc. U. S. Nat. Mus., + 24:147, October, 1901; Allen, Bull. Amer. Mus. Nat. Hist., + 30:256, December 2, 1911. + + _Mustela affinis_, Osgood, Field Mus. Nat. Hist. Publ. 155, zoöl. + 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. + + + _Type._--Male, subadult, skull and skin; no. 123341, U. S. Nat. + Mus., 1630 meters elevation, Montes de Mérida, near Mérida, + Venezuela; August 14, 1903; obtained by S. Briceno. + + The skull (plates 25 and 26) lacks the right exoccipital condyle + and posterior half of the right zygomatic arch. The teeth all are + present, unworn and entire. The skin is well made and complete. + + _Range._--Near sea level (San Julián) to 8500 feet (Montes de + Culata, Mérida, 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. frenata panamensis_ in + lighter color of upper parts (tone 3 rather than tone 4, pl. 344, + Reddish Black, Oberthür and Dauthenay), flat rather than convex + dorsal outline of skull immediately behind postorbital processes + (see pl. 27); from _M. f. 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, pl. 344, Reddish Black, Oberthür and Dauthenay) and + by smaller skull and teeth (basilar length less than 45, length of + m1 less than 6.3, width of M1 less than 4.8, outside length of P4 + less than 5.7). + + _Description._--_Size._--Male: 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, 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. Corresponding + measurements of no. 22191, a young male from Mérida, measured by + Osgood or Conover, are 439, 165, 54. The adult male no. 18703, + from Páramo de Tama (eastern boundary of Venezuela) has the + following measurements written 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 collectors' 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 Río Zapata, Colombia, measured + (by J. H. Batty), 315, 138, 39. No. 32182, adult, from Mira + Flores, Cauca, Colombia, measured (by W. B. Richardson), 375, 150, + 43. + + The average differences in external measurements of the two sexes, + at Mérida, 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 slightly greater than shown in + figure 21. + + _Color._--As described in _Mustela frenata panamensis_ except + that: Posterior fourth of each upper lip rarely, and small spot in + front of ear usually, white; black of head proper not extending + back of ears and grading insensibly 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 (pl. 344, Oberthür and Dauthenay) or tone 3 in + freshest, unfaded pelage. Least width of color of underparts (in + ten males from Mérida) 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 type 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 M1 or than between anterior palatine foramen and + anterior margin of tympanic bulla; mastoid breadth less than + postpalatal length; postorbital breadth 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 maximum) 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 M1 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 width 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 width 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 lighter than that of the + average male. + +Comparisons of the skull with those of _M. f. panamensis_ and _affinis_ +have been made in the accounts of those subspecies. As compared with +the skull of the male of _M. f. 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 margin of upper molars, or +measurements of teeth. Relative to the basilar length, most of the +measurements are greater in _meridana_. 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 compared it +with _M. f. affinis_ and most of the differential characters which he +ascribed to _meridana_ were merely "more than" or "less than" in +_affinis_. In _affinis_, Hollister included specimens from Chiriquí, +Panamá, 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 Bogotá, Colombia, and synonymized _Mustela +meridana_ with _M. a. affinis_. Hollister probably would not have named +_meridana_ had he had specimens from Bogotá for comparison and had he +regarded them as topotypes of _affinis_ for the difference is slight. +Nevertheless, within the large geographic range of _M. f. meridana_ +there is some geographic variation. There is more of such variation in +the color of the pelage than in shape and size of the skull. The +specimen from San Julián is darker than the average and in this respect +approaches true _panamensis_. San Julián is situated at a relatively +low elevation on the coast of Venezuela. + +_M. f. 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 unquestionably are, _on the +average_, different, and specimens from the southeastern part of the +range of _affinis_ probably are individually distinguishable from +topotypes of _meridana_. + +Variation in the skulls of the series from Mérida is relatively small. +This applies 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. All are of nearly the same tone except juveniles or +young which are, as in the case of _panamensis_, much brighter colored +on the underparts. 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 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 follicles. The white facial markings +vary relatively little in the 45 specimens carefully examined 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 specimen 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 confluent with the +color of the upper parts. + +A young male from San Julián, Robinson and Lyon (1901:147) state ". . . +was shot . . . as it ran over some bowlders in a ravine. Its eyes shone +with the same greenish light as do the eyes of our common weasel, and +it emitted the same strong odor." No. 14463, Am. Mus. Nat. Hist., from +Río Zapata, Colombia, according to data on the label, was "taken in +timber belt in valley in balk hills" and the native name is Cosonebi. +Two specimens taken on the Páramo de Tama, head of Tachira River, +Venezuela and Colombia 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 'house' 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 mentioned +in "Specimens examined" are described and located by Chapman +(1917:640-656, pl. 41) in his "Distribution of Bird-life 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 Río Barrotow, Río +Oscuro, Río Zapata, Río Japata, Guasca and El Baldro. Sonson may or +may not be the town of that name situated some eighty miles northwest +of Bogotá 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 Mérida are labeled 1630 meters, Montes de Mérida. +San Julián 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. Páramo de Tama is on the +Venezuelan-Colombian border near the source of the Tachira River (see +Osgood, 1912:35). Mt. Duida is shown as at 3° 30´ N and 65° 40´ W by +Chapman (1931:13) and Mt. Auyán-tepui 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 Julián, 1[91]; Carácas, 2; Galipare, Cerro del + Avila, 6500 feet, 1; San Esteban, 1[2]; Mérida, 45 (10[91], 14[2], + 10[4], 2[60], 2[14], 1[78]); Páramo de Tama, 1[60]. + + =Colombia=: Páramo de Tama, 1[60]; Cincinnati, 3[9]; Valdiva, 3800 + ft., 1; Medellín, 2; 7200 ft., Barro Blanco, 1[2]; Santa Elena, + 9000 ft., 1[2]; Santa Elena, 1[2]; Sonson, 2 (1[91], 1[2]); Mt. + Auyan-tepui, 1[2]; Pueblo Rico, 5200 ft., 1[91]; Mira Flores, + 1[2]; Jerico, near Cauca River, 1; Tornel, 20 mi. NE Quitichao, 1; + Mt. Duida, 1[2]; El Tambo, Cauca, 1[78]; El Baldro, 1[2]; Río + Japata, 2[2]; Río Zapata, 4500 ft., 1; Río Oscuro, 3300 ft., 1; + Río Barrotow, 3300 ft., 1; Guasca, 1[75]; 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 numbered 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 Bogotá, 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 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 Carmen), 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 _M. 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. f. aureoventris_ by lighter-colored upper + parts (tone 2 rather than tone 4, pl. 344, Reddish Black of + Oberthür and Dauthenay); from _M. f. macrura_ by darker color + (Reddish Black, tone 2, pl. 344, Ober. and Dauth., rather than + Chocolate, tone 3, pl. 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. Proportions 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 (pl. 344, Oberthür and Dauthenay). Least width of + color of underparts (in five males from vicinity of Bogotá) 24 + (15-29) 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 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±; interorbital breadth not + greater than distance between foramen opticum and anterior 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 relatively less. +Thus the skull of _affinis_ is longer and broader, but the facial +region is actually, as well as relatively, smaller. As compared 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 specimen of +_affinis_ are the measurements of length of tooth-rows or breadth of +rostrum, actually, or relatively, as great as in _aureoventris_. The +same is true of all measurements taken of M1, P4 and m1. 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 intergradation of _affinis_ with +_macrura_. + +_Remarks._--_Mustela affinis_ was named by John Edward Gray in 1874 (p. +375) on the basis of a specimen from New Granada. Although usually +synonymized with _Mustela brasiliensis_ by later authors until 1896 +when Merriam (1896:31) applied the name to weasels from Costa Rica, +nearly all the South American and several 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 mentioned the specimen, that later +became the holotype. In 1916 (p. 98) Allen restricted the type locality +to Bogotá, 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), Bogotá has long been the shipping point for +Colombian vertebrate specimens, many of which were obtained in the +mountains 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 Bogotá agree with Gray's description of the type +specimen. + +_Mustela frenata affinis_, as here defined, constitutes one of the +several slight geographic variants met with, on the sides of, and +between, 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 Bogotá 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 +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 Bogotá show well the +color and the color pattern but lack collectors' measurements. + +None of the ten skulls examined shows malformation of the frontal +region due to infestation of the frontal sinuses by parasites. Possibly +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 Carmen, 1[2]; W. Cundinamarca, 1[7]; Muzzo [= + Muzo?], 1[4]; Bogotá, 1; Castillo, near Bogotá, 1[7]; Fambrias, + near Bogotá, 1[75]; Bogotá district, 1[2]; Choachí, 9 (1[75], + 2[7], 1[84]); Páramo de Choachí, 2 (1[2], 1[84]); Laguna del + Verjón (= City of Bogotá), 1[75]; Quetame, 2[2]; Fusagasuga, 1; + Caqueta, 2[2]; no locality more definite than Colombia, 3 (1[7]). + + +=Mustela frenata aureoventris= Gray + +Long-tailed Weasel + +Plates 27, 28 and 29 + + _Mustela aureoventris_ Gray, Proc. Zoöl. Soc. London, 1864:55, pl. + 8, 1864; Gray, Proc. Zoöl. 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_, Lönnberg, Arkiv för 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 (locality 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 life-zones. 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 + rather than one root on P2; from _Mustela frenata macrura_ by + Reddish Black, tone 4, plate 344 rather than Chocolate, tone 3, + pl. 343 (of Oberthür and Dauthenay), or slightly darker color of + upper parts; from _M. f. affinis_ and _M. f. meridana_ by darker + color (tone 4 rather than tone 2, Reddish Black of Ober. and + Dauth.) of upper parts and larger size of teeth (M1 with length + more than 2.4 and breadth more than 4.7; P4 with outside length + more than 5.9; length of m1 more than 6.2). + + _Description._--Unless otherwise stated, information concerning + this subspecies 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 Oberthür and Dauthenay, pl. + 344; chin whitish; remainder of underparts Warm Buff, deep orange + in juvenile, type specimen, according to Gray (1864, pl. 8); color + of underparts extending distally on posterior sides of forelegs to + wrists but not reaching 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 length 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; zygomatic breadth + approximately equal to distance between condylar foramen and M1 + 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 premolars + 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 + width 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 I3) upper incisors; height of tympanic + bulla not greater than distance 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. f. aureoventris_, and _Mustela frenata + macrura_ from the 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. f. affinis_. Gray +(1864:55) states that the type specimen measures "Length of body and +head 6 inches, of tail 4-1/2 inches." The plate (pl. 8) accompanying +Gray's original description (_loc. cit._) is marked one-half natural +size and represents the animal as having 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 +northern section of _M. f. macrura_, but in color to _M. f. +panamensis_. _M. f. aureoventris_ and _M. f. panamensis_ are the two +darkest-colored subspecies and each occurs in a region of extremely +heavy rainfall. There is a skin only, no. 32620, Amer. Mus. Nat. Hist., +from Munchique, obtained on June 1, 1911, which is appreciably darker +than specimens of _M. f. affinis_ in corresponding pelage and is +intermediate between _M. f. affinis_ and _M. f. aureoventris_ in color +as it is geographically. 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 description which in +pagination precedes the colored plate of the animal that is labeled +_Mustela aureoventris_. _Putorius brasiliensis_ var. _aequatorialis_ +Coues (1877:142) is the only name known to the writer that has been +proposed as a substitute for _Mustela aureoventris_ 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. f. 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 _aureoventris_ as compared with an adult from +New Granada [= Colombia] 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 original +descriptions of _aureoventris_ and _macrura_ well illustrate the +difference stated in the written descriptions. My examination of the +type specimens of _M. macrura_ and _M. f. aureoventris_ shows them to +have been fairly accurately portrayed in the plates accompanying 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, pl. 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 specimens from the Temperate Zone of +Ecuador and northern Perú. Because Quito, Ecuador, is in the Temperate +Life-zone and because the available specimens from this zone in Ecuador +and northern Perú 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 +Subtropical 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 vicinity of the city, from +the 'Napo' region on the Amazonian slopes of the Andes, and from +Nanegal, Gualea, and other localities on the Pacific side rarely below +the Subtropical Zone." It is also pointed out that only some of the +specimens are labeled with their approximate 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 intergrade tending +toward the lighter-colored _Mustela f. macrura_ of the Temperate Zone +although sufficiently 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, Río Chinchao, Perú; 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 locality) to 6700 feet (Ambo), + Tropical and Subtropical life-zones of eastern Perú. 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, pl. 342 rather than Chocolate, + tone 3, pl. 343, Oberthür and Dauthenay) of upper parts. + + _Description._--_Size._--Male: Measurements of the type specimen + and topotype, 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, 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 + figure 20. + + _Color._--Rarely a few white hairs anterior to each ear; posterior + fifth of each upper lip white; top of head, posteriorly to + slightly 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 (pl. 342, Oberthür 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 M1 or than between anterior + palatine foramen and anterior margin of tympanic bulla; 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 rostrum. + + The skull of the female averages 62 per cent lighter 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. + +This subspecies is insufficiently known, especially as to geographic +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 measurements, 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 strikingly +like those of _Mustela stolzmanni_. Possibly the similar climatic +conditions under which the two live have left their impress in similar +fashion in this part of each of the two species. The teeth, tympanic +bullae, and certain other parts of the skull are, however, so +differently proportioned as to show that the skulls represent two +species. The referred male has a much longer skull than the type +specimen and the relative proportions of breadth and depth of the two +skulls differ widely. Judging from large series of weasels examined +from localities outside the range of _M. f. helleri_, the two skulls +probably represent almost the maximum of individual variation 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. + + =Perú=: 3500 ft., Hacienda Buena Vista, Río Chinchao, 1; 3000 ft., + Hacienda San Antonio, Río 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. + Zoöl. Soc. London, 1865:113, 1865; Taczanowski, Proc. Zoöl. Soc. + London, 1874:311, 1874; Taczanowski, Proc. Zoöl. 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 Perú] + designated. + + _Range._--High, barren Cordillera of Perú (see Tschudi, orig. + descr.); as here restricted, Temperate Life-zone and higher in + western Andes and intermountain valleys of Perú. See figure 29 on + page 221. + + _Characters for ready recognition._--Differs from _Mustela frenata + macrura_ by lighter color (Chocolate, tone 2 rather than 3, pl. + 343, Oberthür and Dauthenay) of upper parts; length of upper + tooth-rows, in females, less than 13; inside length of P4 more + than 4.6; from _M. f. aureoventris_ by smaller teeth (maximum size + just given for _agilis_); from _M. f. boliviensis_ by lighter + color, upper parts being Chocolate, tone 2, pl. 343, rather than + tone 4 or darker of Carbon Brown, pl. 342 (Oberthür and + Dauthenay). + + _Description._--_Size._--Male: The stuffed skin of an adult, from + Lima, measures: Total length, 460; length of tail, 125; length of + hind foot, 45.7. A skin alone from Huarochirí 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 + light-colored and extending 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 lips edged with white; + throat, breast, belly and higher parts of inner sides of + extremities 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 light, has all + the upper parts near (14´ _j_) Ochraceous-Tawny or Cinnamon, and + tone 4 of Oberthür 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 confluent with color of + upper parts); tip of tail probably black (tip missing); underparts + white, belly probably originally with slight tinge of yellow or + allied 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 underparts 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 lip white. Top of head near + (_n_) Mars Brown, and Carbon Brown, tone 3 (pl. 342, Ober. and + Dauth.); tip of tail black; remainder of upper parts near (16" + _j_) Tawny-Olive, and Chocolate (tone 2, of pl. 343 of Ober. and + Dauth.) or Raw Umber (tone 3 of pl. 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 lip + white; white frontonasal spot present, 11 x 11 mm.; antipalmar + faces of forefeet spotted with brown color of upper parts; 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 Huarochirí in color and color pattern closely + resembles no. 21147. It differs from no. 21147 in slightly lighter + color of upper parts, entirely 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, subadult), 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 differs 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. f. +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 possibly 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 Perú. Allen (1916:104) and Thomas +(1920:224) were not convinced that _Mustela agilis_ and _Mustela +macrura_ were distinct species or subspecies. + +Search on August 28, 1937, in the Musée d'Histoire Naturelle, at +Neuchatel, Switzerland, by Mr. Théodore Delachaux, assistant there, and +the writer, revealed no trace of weasels from Tschudi's collection, +although some other specimens of mammals that he figured in the "Fauna +Peruana" are preserved in that Museum. Not only were the collections of +specimens examined but the new catalogue and old catalogue of mammals +were vainly searched for mention of weasels deposited by Tschudi. +Later, at the British Museum 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 +description most nearly applies to the light-colored animals from +western Perú--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 Perú +is that by one line of reasoning, Taczanowski in naming _macrura_ +(_jelskii_ is a synonym of it) from farther eastward in Perú, and that +Hall in naming _helleri_ from still farther eastward, and _boliviensis_ +to the southeastward, geographically restricted the application of the +name _agilis_. Hall's action did this because he recognized geographic +variation and employed the subspecies concept. Taczanowski, 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_ geographically, even though later authors +may choose to rule that his naming of _macrura_ in effect did so +restrict the application of _Mustela 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 lines 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 measurements 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. + +Specimen no. 565, in the Polish Museum of Natural History, without +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, öden Hochebenen der Cordillera an + sonnigen Steinhaufen und Felsen gewöhnlich in Gesellschaft von + 8-12 Stücken. Diese Thierchen sind so ausserordentlich behende und + scheu, dass bei dem leisesten Geräusche die ganze Schaar mit + Blitzesschnelle verschwindet. Es ist uns auch nie gelungen, eines + derselben zu erlegen. Die Indianer aber verstehen es, dieselben + lebendig einzufangen und zu zähmen. Ein sehr zahmes sahen wir bei + einer uns befreundeten Dame in Tarma; gegen alle Fremden biss es + mit Wuth und liess sich nicht anfassen, während es sich von seiner + Herrin Alles gefallen liess; sie öffnete ihm den Mund und steckte + ihm den Finger hinein, ohne dass es eine böse Miene dazu machte, + während es bei der geringsten Bewegung, die wir machten, es zu + ergreifen, grimmig auf uns lossprang. Wenn es eingeschüchtert + wurde, versteckte es sich in den Busen seiner Gebieterin und kroch + ihr bald nachher zum Aermel heraus. An den Wänden und Meublen + kletterte es mit grosser Behendigkeit und schlüpfte durch so + kleine Ritzen und Löcher, dass wir fast an der Möglichkeit dieses + Hindurchdringens gezweifelt haben würden, wenn wir es nicht selbst + mit angesehen hätten. Wenn es unartig war, wurde es mit einer + Schnur an seinem kleinen Halsbande festgebunden; dadurch vermehrte + sich sein Zorn, so dass es zuweilen gegen die Dame auffuhr. + Mehrmals verschwand es während 8-10 Tagen und kam dann plötzlich + wieder zum Vorschein. Seine Nahrung bestand in Gemüse und Fleisch, + besonders aber liebte es Zuckerbrod in Milch aufgeweicht; einmal + machte es sich an einen Kanarienvogel, den es auch tödtete. Es + erhielt seine Strafe und verschwand dann für immer. Die Indianer + sollen dieses Wiesel zum Fange der Viscacha abrichten (davon + weiter unten). Sie nennen es Comadreja, auch Ardilla. ([footnote] + Ardilla ist spanisch und heisst Eichhörnchen. Mit diesem Namen + werden sehr verschiedene Thiere bezeichnet; ausser dem Sc. + variabilis und der Galictis agilis auch noch mehrere Nager und + einige Didelphysarten.) Die Indianerinnen verfertigen sich aus dem + kleinen Felle Geldbeutel. Des Sonntags trifft man unter den vielen + tausend Punaindianerinnen die nach den grossen Dörfern der Sierra + kommen, um ihre Einkäufe zu machen, kaum ein halbes Dutzend, die + nicht solche Börsen mit sich führten, und dann kann man auch die + verschiedensten Farbennuancen, die bei dieser Species vorkommen, + beobachten." + +None of the three skulls referred to this subspecies shows infestation +of the frontal sinuses by parasites. + + _Specimens examined._--Total number, 4. + + =Perú=: Macate, 1 (Field Mus. Nat. Hist.); Huarochirí, 1 (Mus. + Comp. Zool.); Lima, 1 (British Mus. Nat. Hist.); no locality more + definite than Perú, 1 (Mus. Polonais d'Hist. Nat.). + + +=Mustela frenata macrura= Taczanowski + +Long-tailed Weasel + +Plates 1, 27, 28, 29, 30, 37, 38, 39 and 40 + + _Mustela macrura_ Taczanowski, Proc. Zoöl. Soc. London, 1874:311, + pl. 48, May 19, 1874; _ibid_., 1881:647, May 17, 1881; _ibid_., + 835, November 15, 1881; Lönnberg, Arkiv för 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; Lönnberg, Arkiv för Zool., 14 (no. 4):11, 1921. + + _Putorius (Gale) braziliensis frenatus_, Coues, Fur-bearing + animals, p. 142, 1877. + + _Mustela jelskii_ Taczanowski, Proc. Zoöl. Soc. London, 1881:647, + May 17, 1881. + + _Mustela affinis_, Lönnberg, Arkiv för 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, with skull separate; no. 561, + Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Lake Junín, central + Perú; 1873; obtained by M. Jelski. + + The skull (plates 27-29, 30), mounted with 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 + Perú and Ecuador north from the states of Apurimac and Cuzco, + Perú, to San Antonio, northern Ecuador. 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 helleri_, _M. f. + affinis_ and _M. f. aureoventris_ by lighter color of upper parts + which are Chocolate tone 3, pl. 343, Oberthür and Dauthenay, + whereas, with reference to the same color standard, the colors + are: in _helleri_, Carbon Brown, tone 3, pl. 342; in _affinis_, + Reddish Black, tone 2, pl. 344; in _aureoventris_, Reddish Black, + tone 4, pl. 344; from _M. f. agilis_ by darker color (Chocolate, + tone 3 rather than 2, pl. 343, Oberthür 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, pl. 342 of Oberthür and Dauthenay, and larger + size (in males, hind foot more than 45 and m1 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 Yana Mayo, Río + Tarma, was measured by Hendee as 394, 134. Hind foot relaxed + measures, 47. Tail 55 per cent as long as head and body. Length + of 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; length 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 slightly behind eyes, near (_n_) + Chestnut-Brown (Ridgway) and Carbon Brown, tone 2 or darker (pl. + 342, Oberthür 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 (_l_) Russet (Ridgway) and + Chocolate, tone 3 (pl. 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 Yana 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 foot 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 foramen and M1 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 foramen opticum and anterior margin of tympanic + bulla; breadth of rostrum less than length of tympanic bulla; + 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 I3) 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, Perú, 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 5-1/2 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. + +As compared with that of _helleri_, the skull of the male of _macrura_ +from Junín 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 Junín. As one +proceeds northward the color of the weasels becomes progressively +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, Perú, to near Quito, Ecuador. West of the range of +_macrura_ there exists the still lighter-colored subspecies, _M. f. +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 Junín 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 distinct 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 differences +between the two skulls are not greater than those ordinarily found in +specimens from the same locality. No. 562 has the tympanic bullae +greatly, relative to the other South American weasels, inflated +posteriorly. Otherwise, the bullae agree with those of the type +specimen. + +Specimens from southwestern Ecuador, average large, and include the +largest specimens of the species _Mustela frenata_ seen from South +America. A subadult male, no. 61406, in the American Museum of Natural +History, is the largest. Its external measurements are 482, 191, 56. +The basilar length of the skull is 48.2 and the zygomatic 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 +Junín. + +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 Junín and the +one from Ollantaytambo. + +In size of teeth the female, type specimen of _M. jelskii_, from +Cutervo, shows an approach to the larger-toothed weasels of the +northern part of the range of _macrura_. + +The specimens in the Riksmuseum from the vicinity of Quito, Ecuador, +have been rather fully described by Lönnberg (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. f. 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 geographic +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 purchased 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, +therefore, it was taken not at Baeza, Ecuador, which I find to the +eastward of Quito at 77° 55' W and O° 25' S, but at some place of the +same name on the Pacific Slope, unless the locality has been altogether +wrongly 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 therefore +offers insufficient basis for the judging of its subspecific +relationships. + +Other specimens, in the British Museum of Natural History, recorded 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 Junín +proved to belong to the superfamily Oxyuriodea according to Professor +W. B. Herms and Mr. O. L. Williams, who have independently 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[7]; San Antonio, + 8000-8500 ft., 5 mi. N Quito, 4 (2[7], 2[78]); Nono, 10000 ft., 1; + Mindo, 1[78]; Zambiza, 8000-8100 ft., NE Quito, 4 (2[78], 2[95]); + Carapungo, 8500 ft, NE Quito, 1[78]; Panecillo, 10000 ft., near + Quito, 2[78]; Guapulo, 8800 ft., 3 mi. E of Quito, 1[78]; + Pichincha, 10500 ft., and 12000 ft., 2 (1[78], 1[95]); San + Ignacio, 11500 ft., Pichincha, 1; Santa Rosa, 9600 ft., Río Pita, + 2; near Santa Rosa, 9000 ft., 1; Río San Rafel, 9000 ft., 1; N + side Quito, 9000 ft., 1[78]; Quito, 1[4]; near Quito, 5[7]; Nára + Papallacta, 11000 ft., 1[78]; below Papallacta, 9000 ft., 1[78]; + Chillo Valley, 1[78]; "Hacienda Hda," 10000 ft., Pintag, Valencia, + 1; Baeza arriba, 1; Ambato, 2; San Francisco, 8000 ft., E of + Ambato, 1; Chunchí, Pagma Forest, 6400 ft., 1[1]; Canar, 2600 M., + 1[7]; Malletura, 7600 ft., 1; Contrayerbas, 11000 ft., 1; Sisig, + 8500 ft., 3[7]; El Chiral, 1; Almor, 1; Guainche, 3200 ft., 1; no + locality more definite than Ecuador, 4[95]; "Received from Quito," + 1[7]; Quisaya, 6000 ft. (locality not found), 1[7]; La Carolina + (locality not found), 1[78]. + + =Perú=: La Lejía, 1; Huancabamba, 4 (2[75]); Cutervo, 9000 ft., + 1[73]; Condechacha, 7000 ft., Río Utcubamba, 1[7]; San Pedro, + 8600-9400 ft., S of Chachapoyas, 1; Celendín, 1[7]; Junín, 2[73]; + Yana Mayo, Río Tarma, 1[7]; Ollantaytambo, 9000 ft., 3 (1[7], + 2[91]); Ocabamba, 1[7]; Anta Cuzco, 3400 and 3500 M., 2[4]; + Marcapata, 1[91]. + + +=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). + + + _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 + Limbaní, Perú, south to Nequejahuira, Bolivia; upper Subtropical + and Temperate life-zones. 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 of upper parts (tone 4 or darker of Carbon Brown, pl. + 342 rather than tone 3 of Chocolate, pl. 343, Oberthür and + Dauthenay) and lesser size (in males hind foot less than 45 and m1 + less than 5.6); from _Mustela frenata agilis_ by darker color of + upper parts (as given above rather than tone 2 of Chocolate, pl. + 343, of Oberthür and Dauthenay). + + _Description._--_Size._--Male: The type and two young specimens + from Limbaní, Perú, 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 (pl. 342, Oberthür + 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 underparts 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 their +way into a zoölogical collection were the two young males taken on +February 17, 1904, at Limbaní, 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 Limbaní, in the United States National Museum, an +account of this race was drawn up, but the account was discarded 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 previously recognized kinds. + +The collector has noted on the labels of the two young from Limbaní +that they were shot in the afternoon when running together beneath +bushes. The frontal sinuses of the type are malformed as a result of +infestation by parasites. + + _Specimens examined._--Total number, 3, as follows: + + =Perú=: Carabaya, Limbaní, 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, pl. 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 + measurements and plates 39 and 40; weight unknown; 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 individual +referred by Brown (1908) to this species. The remaining material of +weasels from this deposit was referred by Brown to his _Putorius +cicognanii angustidens_. Examination of the original materials +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 occasional skull +alone of a subadult male _erminea_ is extremely difficult certainly to +distinguish from a skull alone of an adult female _frenata_. This is +true among Recent specimens in the northern Mississippi Valley today; +more exactly in Iowa and southern Minnesota the females of _frenata_, +oftentimes intergrades between the subspecies _Mustela frenata +longicauda_, _M. f. noveboracensis_ and _M. f. primulina_, 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. f. +primulina_ which occurs in the same region today and to _M. f. +noveboracensis_, 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 inflated +anteromedially. _M. gracilis_ differs from both _noveboracensis_ (97 +[M] and 56 [F] with skulls of comparable age) and _primulina_ (64 [M] +and 24 [F] with skulls of comparable age) in that the zygomatic 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 M1, 3.4; length of inner + moiety of M1, 1.8; depth of skull at anterior margin of + 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. + + +=MUSTELA AFRICANA= Desmarest + +Tropical Weasel + +(Synonymy under subspecies) + + + _Type._--_Mustela africana_ Desmarest, Nouv. Dict. d'Hist. Nat., + vol. 19, p. 376. 1818. + + _Range._--Known from the headwaters of the Amazon in eastern Perú + 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._--Differs from _Mustela frenata_, +the only geographically adjacent species of the genus, by: presence of +thenar pad on forefoot; presence of a longitudinal, median, abdominal +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 pl. 29, fig. i and pl. 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; braincase +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 + + 3 1 2-3 1 + -, -, ---, -; teeth heavy; medial lobe of M1 but slightly larger than + 3 1 2 2 + +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 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 Perú. +However, the resemblance is not close. The tympanic bullae, although +unique among American weasels, are more like those of _M. f. 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 reduction in +number of premolars, are of a primitive nature. For example, 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 barrier in the Central American +region, for a length of time sufficient 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_ intergrades 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 include +the one species _africana_. I am inclined to accord _Grammogale_ only +subgeneric rank. + +It is possibly significant that _Mustela africana_ is intermediate in +several respects between _Lyncodon_ and typical _Mustela_. The median, +longitudinal, abdominal band of the same color as the upper parts in +_M. africana_ and the relative uniformity of the coloration 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, unicolored upper parts and dark-colored underparts +of the Patagonian weasel (_Lyncodon_) on the other hand. The number of +premolars, + + 2-3 + ---, is also intermediate between the numbers + 2 + + 3 2 + - and - of the North American _Mustela_ and the Patagonian + 3 2 + +_Lyncodon_, respectively. The 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 examined), 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 Córdoba, at +each place in the Pampean [= Pleistocene] formation of Argentine (see +also Cabrera, 1928:263) is the first and only fossil form of this group +recorded from the whole of South America. 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 Pleistocene. + +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 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 different sexes from +Pará, described and figured by Goeldi (1904:61-62, pls. 1, 2). The +undoubted female, type specimen of _Mustela africana stolzmanni_, is as +large as the undoubted male, no. 37475, of the same species, but of a +different subspecies, from Pará, 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 Pará +had been captured "swimming in the salt water of the estuary about half +a mile away from the shore." On the label of the specimen 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. Españ. de Hist. Nat., 13:429, + November, 1913; Cabrera, Bol. Real Soc. Españ. de Hist. Nat., + 14:175, pl. 1, March, 1914. + + _Putorius (Mustela) brasiliensis paraensis_, Goeldi, Zool. Jahrb. + abt. f. systematik, geogr. u., Biol., 10:556, pl. 21, September + 15, 1897, type from Pará, Brazil, near Pará, 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, pls. 1, + 2, 1904. + + _Mustela affinis paraensis_, Hollister, Proc. Biol. Soc. + Washington, 28:143, July 10, 1914. + + _Mustela paraensis_, Allen, Bull. Amer. Mus. Nat. Hist., 35:105, + April 28, 1916; Tate, Journ. Mamm., 12:253, August 24, 1931. + + _Mustela stolzmanni paraensis_, Hall, Carnegie Instit. Washington + Publ. 473:111, November 20, 1936; Hall, Physis, 16:167, pl. 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 determined from the characters of the animal; probably came + from Brazil, and for the present assumed to be from Pará. + + 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, pl. 1). Except for the loss of the distal part of + the tail, and fading because of exposure to light, the specimen + was in good condition. See also under remarks. + + _Range._--Known from the south side of the Amazon River, near its + mouth at Pará and Cametá, Río 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 pls. 29 and 30) and + from _Mustela africana stolzmanni_ by lighter color of upper parts + which although near Chestnut-Brown are in adults 10' _l_ (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 measurements 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 measurements written on the attached 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 Cametá, and a subadult labeled as male, from Pará + 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 hypothenar pad except + in one young specimen); longest facial vibrissae not extending to + posterior margin of ear; superior genal tuft not found, hairiness + of foot-soles as shown in figure 22. + + _Color._--Upper parts near (10 l) 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 (lips 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; distance from postorbital + process to anterior, nasal notch approximately equal to breadth + across exoccipital condyles; palatal region wide; tympanic bullae + less 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 between C1 and P4 less + than length of P4; teeth heavy; medial lobe of M1 only slightly + larger than lateral lobe; deuterocone of P4 heavy and often + inclined mesially; p2 absent (P2 present above on both sides in + only one of seven specimens seen or described); lower jaw heavy; + masseteric fossa not extending anteriorly to posterior fourth of + talonid of m1; paraconid of m1 low and base of cleft between it + and protoconid relatively low on tooth. + + Female: No skull examined but from figures published by Goeldi + (1904, pl. 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 _Putorius (Mustela) brasiliensis paraensis_ by +Goeldi (1897:556, pl. 21) from Pará, 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. 111) 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 specimen 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 before 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 +Pará and not of the darker chocolate color of specimens of _M. +stolzmanni_ from Perú. The specimen is indistinguishable from topotypes +of _P. paraensis_ 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, pls. 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. + +Pará, and Cametá, Brazil, places from which _Mustela africana africana_ +is known, are nearly 2000 miles from the localities in eastern Perú and +eastern Ecuador from which _M. a. stolzmanni_ is known, and no +specimens, from intermediate localities, are available 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 considerations, 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 Rodríguez 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 +Ethnographia, Pará, Brazil. + +The one young specimen seen, that from Cametá, is darker colored than +any of the four older specimens examined. It is almost exactly the +Chestnut Brown of Ridgway (1912) and therefore approaches 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: Pará, 2 (1[2], 1[7]); Pará Murutucu, 1[7]; Río + Tocantins, Cametá, 1[75]; type specimen, 1[84]. + + +=Mustela africana stolzmanni= Taczanowski + +Tropical Weasel + +Plates 39 and 40 + + _Mustela stolzmanni_ Taczanowski, Proc. Zoöl. 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, pl. 1, + figs. 5, 6, 1939. + + _Type._--Female, adult, mounted skin, with skull separate; no. + 563, Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Yurimaguas, + Perú; 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 M1 and adjacent part of maxilla are lost. + The skin is well mounted, in a good state of preservation and + shows no fading due to exposure to light. Inguinal mammae are + distinctly 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 + longitudinal, abdominal stripe, unmarked by color of the upper + parts. + + _Range._--Known from the Tropical Life-zone of eastern Ecuador and + Perú from Jatun Yacu south to Valle del Perené. 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 pls. 28, 29, 30, 39 + and 40) and from _Mustela africana africana_ by darker color of + upper parts which, although near Chestnut Brown, are 11' _n_ + instead of 10' _l_ as in _M. a. africana_. + + _Description._--_Size._--Male: unknown. + + Female: Taczanowski (1881:836) gives, among others, the following + measurements 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 posture of the + now mounted specimen, I get from it essentially the same + measurements. Collectors' measurements of a subadult from + Moyobamba and a young female from Valle del Perené, are + respectively, as follows: 469, 415; 184, 160; 57, 52. My own + measurements of the dry hind feet on the skins are respectively, + 48 and 49. + + _Externals._--As described in _M. a. africana_ except that the + length of the 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 lighter 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 Perú, 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 Yurimaguas (see Osgood, +1914:147), but secured no topotypes of this little-known weasel. C. O. +Schunke took the second specimen in the Valle del Perené in April, +1921; L. Rutter on January 25, 1924, took the third specimen, and W. +Clark-MacIntyre 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. f. 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. Zoöl. + + =Perú=: Yurimaguas, 1 in Mus. Polonais d'Hist. Naturelle, Warsaw; + Moyobamba, 2700 ft. [6° S, 77° W], 1 in Brit. Mus. Nat. Hist.; + Valle del Perené, 1200 meters, 1 in Amer. Mus. Nat. Hist. + +EXPLANATION OF CRANIAL MEASUREMENTS APPEARS ON PAGE 417 + +[Illustration: 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. f. primulina_, from 3 mi. E Bergman, +Boone County, Arkansas, obtained December 12, 1933, by B. G. Roberts; +ad. [F]. 62854 Mus. Vert. Zoöl. × 1-2/5.] + + + + +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 + alveoli of the first upper incisors. F to F' on fig. 31. + + Condylobasal length.--Least distance from a line 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 G' on fig. 31. + + 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. 31. + + Interorbital breadth.--Least distance across top of skull between + orbits (eye sockets). O to O' on fig. 31. + + 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 + perpendicular 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. + +m1, 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 (anterointernal cusp). + +M1.-- + + 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 Ms1.--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. + + + + + TABLE OF CRANIAL MEASUREMENTS + + + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela frenata + average} {45.1 16.6 13.7 + maximum} [M] ad. 10 Adirondacks, New York {47.0 17.4 14.6 + minimum} {43.2 15.9 13.0 + + average} {44.7 16.3 13.7 + maximum} [M] ad. 10 Massachusetts {47.0 16.8 14.9 + minimum} {43.3 15.9 13.1 + + average} {44.0 16.3 13.2 + maximum} [M] ad. 10 Liberty Hill, Connecticut {46.0 17.4 14.4 + minimum} {41.5 15.2 12.1 + + average} {44.4 16.5 13.3 + maximum} [M] ad. 10 Beaver Dam, Wisconsin {46.1 17.7 14.1 + minimum} {40.6 15.6 12.3 + + average} {43.0 15.6 13.0 + maximum} [M] ad. 10 Washtenaw Co., Michigan {45.4 16.5 13.4 + minimum} {39.7 14.7 11.6 + + average} {37.5 13.4 10.7 + maximum} [F] ad. 8 Adirondacks, New York {39.8 14.5 11.2 + minimum} {36.4 12.8 9.8 + + average} {36.5 13.4 10.5 + maximum} [F] ad. 5 Massachusetts {38.1 14.0 11.0 + minimum} {35.2 13.0 10.0 + + average} {37.3 13.5 10.8 + maximum} [F] ad. 6 Maryland {37.9 14.1 11.2 + minimum} {36.8 13.0 10.4 + + average} {37.2 13.4 10.6 + maximum} [F] ad. 5 Beaver Dam, Wisconsin {37.6 13.7 11.3 + minimum} {37.0 12.9 10.0 + + average} {36.5 13.1 10.5 + maximum} [F] ad. 9 Washtenaw Co., Michigan {37.5 13.7 11.0 + minimum} {35.9 12.8 9.8 + + Mustela frenata + A. N. S. P. 4137 [M] ad. Washington Co., Maine 45.8 16.5 13.3 + M. C. Z. 7267 [M] ad. Moose Head Lake, Maine 44.9 16.6 14.5 + M. C. Z. 5501 [M] ad. Bucksport, Maine 46.9 17.1 14.4 + M. C. Z. 9142 [M] ad. Bucksport, Maine 45.0 16.1 14.0 + av 45.7 16.6 14.1 + + M. C. Z. 9101 [F] sad. Bucksport, Maine 37.7 14.0 11.6 + M. C. Z. 9122 [F] sad. Bucksport, Maine 38.2 13.8 11.5 + + Mustela frenata + average} {44.8 16.8 13.5 + maximum} [M] ad. 10 Douglas Co., Kansas {46.0 17.8 14.1 + minimum} {43.8 16.2 12.9 + + average} {44.7 16.5 13.2 + maximum} [M] ad. and sad. 8 Boone Co., Arkansas {46.5 17.1 14.0 + minimum} {42.5 15.6 12.6 + + average} {38.9 14.4 11.3 + maximum} [F] ad. and sad. 11 Douglas Co., Kansas {40.7 15.3 12.0 + minimum} {37.6 13.8 10.8 + + average} {39.3 14.1 11.4 + maximum} [F] ad. and sad. 6 Arkansas {40.1 14.6 11.9 + minimum} {38.8 13.7 11.0 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + noveboracensis + 10.9 15.3 23.3 26.7 15.4 7.8 3.5 5.9 5.3 5.5 4.3 2.4 14.1 12.7 + 11.9 15.9 24.7 28.0 16.2 8.4 4.0 6.2 5.7 5.9 4.5 2.7 14.5 13.3 + 10.4 14.8 22.0 25.3 14.6 7.0 3.1 5.5 4.9 4.9 4.0 1.9 13.5 12.2 + + 11.3 15.7 23.2 26.5 15.4 7.7 3.4 5.9 5.2 5.5 4.4 2.4 14.1 12.7 + 12.0 16.7 25.6 28.2 16.4 8.2 3.7 6.3 5.5 5.9 4.7 3.0 14.7 13.0 + 10.5 14.8 21.8 25.1 14.3 7.3 3.1 5.6 5.0 5.2 4.0 2.0 13.5 12.0 + + 10.9 15.3 22.4 26.3 15.0 7.5 3.3 5.8 5.1 5.4 4.2 2.2 14.3 12.6 + 12.0 16.4 23.5 28.0 15.9 8.0 3.6 6.3 5.6 6.1 5.0 2.5 15.2 13.2 + 9.7 14.2 20.8 24.0 14.4 6.9 3.0 5.3 4.6 5.0 3.7 2.0 13.4 11.7 + + 10.6 15.6 23.0 26.3 15.2 8.0 5.8 5.3 5.5 4.3 2.4 13.9 12.6 + 11.3 16.4 23.8 27.4 16.2 8.2 6.1 5.7 5.8 4.6 2.7 14.5 13.2 + 9.8 13.7 22.1 23.9 14.2 7.7 5.6 4.7 5.1 3.9 2.1 13.0 11.8 + + 10.2 14.9 22.0 24.9 14.7 7.6 5.6 5.0 5.3 4.2 2.2 13.4 11.9 + 11.0 15.7 22.8 26.1 15.2 7.9 6.0 5.3 5.6 4.5 2.5 14.2 12.3 + 9.6 13.8 21.1 23.5 14.1 7.3 5.0 4.7 4.9 3.8 1.9 12.6 11.5 + + 8.7 12.9 18.7 20.9 13.5 6.5 2.8 4.8 4.3 4.6 3.5 1.7 11.4 10.2 + 9.3 13.4 19.3 22.1 14.3 7.0 2.9 5.2 4.7 5.0 3.7 1.9 12.1 10.8 + 7.9 12.1 17.5 20.0 12.6 5.9 2.5 4.5 4.0 4.2 3.2 1.7 10.3 9.4 + + 8.5 12.9 18.5 20.1 13.3 6.3 2.3 5.0 4.3 4.5 3.5 1.8 11.4 10.0 + 8.6 13.3 19.6 20.8 13.9 6.9 2.7 5.1 4.5 4.7 3.7 2.0 11.6 10.4 + 8.3 12.1 17.2 19.3 12.8 5.6 2.0 4.9 4.2 4.4 3.3 1.4 11.0 9.6 + + 8.6 13.0 18.8 21.0 13.1 6.6 4.9 4.4 4.7 3.6 1.9 11.9 10.9 + 8.9 13.5 19.6 21.7 13.5 6.8 5.2 4.6 4.9 4.1 2.3 12.6 11.4 + 8.5 12.8 18.2 20.1 13.0 6.4 4.7 4.2 4.5 3.1 1.4 11.6 10.3 + + 8.8 13.0 18.7 21.0 13.5 6.5 4.9 4.5 4.6 3.5 1.8 11.6 10.2 + 9.7 13.5 19.5 21.7 13.8 6.8 5.0 4.5 4.8 3.7 2.1 12.3 10.3 + 8.0 12.0 17.6 20.2 13.1 6.1 4.9 4.4 4.3 3.3 1.5 10.8 9.8 + + 8.4 12.6 18.5 20.5 13.2 6.3 4.8 4.3 4.5 3.5 1.8 11.8 10.3 + 8.8 13.1 19.1 21.3 13.8 6.7 5.0 4.5 4.7 3.7 2.0 12.2 10.7 + 7.8 12.0 18.1 19.5 12.9 6.0 4.6 4.1 4.3 3.3 1.5 11.3 10.0 + + occisor + 11.0 16.2 24.5 27.5 15.7 8.5 3.5 6.2 5.6 5.9 4.6 2.8 13.3 13.0 + 11.8 15.6 24.2 27.7 15.2 7.7 3.1 6.0 5.2 5.6 4.2 2.4 14.2 13.0 + 11.0 15.8 24.0 28.0 16.4 8.0 4.0 6.0 5.8 6.1 4.5 2.6 14.7 13.2 + 11.2 15.3 24.2 27.4 15.8 7.3 2.8 6.1 5.3 5.6 4.3 2.4 14.5 12.6 + 11.3 15.7 24.2 27.7 15.8 7.9 3.4 6.1 5.5 5.8 4.4 2.6 14.2 13.0 + + 9.1 13.0 19.0 21.7 14.1 6.3 2.6 5.2 4.6 4.6 3.8 2.1 12.5 10.5 + 9.2 13.6 19.5 21.8 13.6 6.5 2.5 4.6 4.3 4.5 3.5 1.7 12.3 10.3 + + primulina + 10.6 14.9 24.0 27.2 15.5 8.3 3.4 6.1 5.5 5.8 4.6 2.4 14.2 12.8 + 11.5 15.4 24.9 28.2 16.3 8.8 3.9 6.6 5.9 6.1 4.7 2.6 14.8 13.2 + 10.0 14.3 23.2 26.2 14.4 8.0 3.0 5.8 5.2 5.5 4.3 2.2 13.6 12.0 + + 10.5 15.0 24.1 26.9 15.5 8.1 3.6 6.0 5.5 5.6 4.3 2.2 14.2 12.3 + 11.8 16.3 24.8 27.7 16.4 8.8 4.2 6.5 6.0 6.2 4.9 2.5 14.6 13.0 + 9.7 14.1 22.8 26.1 14.9 7.5 3.1 5.8 5.2 5.2 4.0 2.0 13.9 11.7 + + 8.6 12.9 20.3 22.6 13.5 6.9 2.9 5.2 4.8 5.0 3.5 1.8 12.6 11.2 + 9.2 13.4 21.4 23.8 15.1 7.5 3.3 5.7 5.2 5.4 4.2 2.1 13.5 11.7 + 7.9 12.4 18.8 21.1 13.0 6.3 2.5 4.8 4.5 4.6 3.4 1.5 11.7 10.0 + + 8.7 13.1 20.3 23.1 14.0 6.9 2.9 5.2 4.8 5.0 3.8 1.9 12.5 10.7 + 8.8 13.7 21.0 23.8 14.5 7.2 3.0 5.5 5.0 5.3 4.2 2.0 13.0 11.8 + 8.5 12.6 19.8 22.5 13.5 6.5 2.8 5.0 4.5 4.7 3.5 1.7 12.3 10.3 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================ + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela frenata + U. S. N. M. 246345 [M] ad. Convent, Louisiana 43.4 15.4 13.7 + M. V. Z. 37515 [M] sad. Remy, Louisiana 43.8 16.4 12.7 + M. V. Z. 38901 [M] sad. Springville, Louisiana *45.0 *16.5 + + Mustela frenata + M. V. Z. 47165 [M] ad. Talbot Co., Georgia 50.1 18.3 15.0 + M. V. Z. 47144 [M] ad. Talbot Co., Georgia 49.2 18.3 13.8 + M. V. Z. 47166 [M] ad. Talbot Co., Georgia 48.5 17.6 14.6 + M. V. Z. 47167 [M] ad. Talbot Co., Georgia 45.8 17.3 14.2 + M. V. Z. 47147 [M] ad. Talbot Co., Georgia 48.1 18.0 14.0 + av. 5 48.3 17.9 14.3 + + M. V. Z. 41023 [M] ad. Thomas Co., Georgia 48.8 17.5 14.9 + M. V. Z. 41025 [M] ad. Grady Co., Georgia 44.8 16.9 13.2 + M. V. Z. 40934 [M] ad. Grady Co., Georgia 47.8 17.4 14.1 + M. V. Z. 40935 [M] ad. Grady Co., Georgia 47.4 18.0 13.6 + av. 4 47.2 17.5 14.0 + + U. S. N. M. 223880 [M] ad. Okefinokee Swamp 49.0 18.8 14.2 + Cornell 198 [M] ad. Okefinokee Swamp 47.3 18.0 14.0 + Cornell 652 [M] ad. Okefinokee Swamp 47.0 17.0 13.8 + U. S. N. M. 180802 [M] ad. Autaugaville, Alabama 46.5 17.3 13.4 + + M. V. Z. 51527 [F] ad. Talbot Co., Georgia 43.5 16.4 12.5 + M. V. Z. 41024 [F] ad. Thomas Co., Georgia 42.7 16.0 12.0 + M. V. Z. 41022 [F] ad. Thomas Co., Georgia 44.0 16.1 12.8 + + Mustela frenata + F. S. M. 49387 [M] ad. Apopka, Florida 49.8 18.6 15.1 + + A. N. S. P. 9379 [F] ad. Tarpon Springs, Florida 44.2 16.6 13.8 + A. N. S. P. 8515 [F] yg. Pasco Co., Florida 15.9 11.8 + + Mustela frenata + M. V. Z. 53795 [M] ad. Elk River, Minnesota 49.2 19.1 15.9 + Walker A23 [M] ad. Elk River, Minnesota 49.0 18.7 15.9 + Walker A37 [M] sad. Elk River, Minnesota 48.7 18.9 15.2 + Dickey A865 [M] sad. Elk River, Minnesota 46.8 17.6 15.4 + Dickey A846 [M] sad. Elk River, Minnesota 46.1 18.0 14.2 + + Dickey 11548 [F] ad. Elk River, Minnesota 42.3 16.4 13.1 + Walker A174 [F] ad. Elk River, Minnesota 43.2 16.2 13.8 + Dickey 9688 [F] ad. Elk River, Minnesota 43.2 16.8 12.7 + U. S. N. M. 188410 [F] ad. Elk River, Minnesota 43.0 16.2 13.2 + av. 4 42.9 16.4 13.2 + + Mustela frenata + A. M. N. H. 15875 [M] ad. Red Deer, Alberta 46.5 18.6 15.1 + N. M. C. 8060 [M] ad. Sweet Grass Hills, Alberta 45.5 18.1 15.5 + F. M. N. H. 7021 [M] ad. Canadian Nat. Park, Alberta 46.8 17.8 13.8 + F. M. N. H. 8567 [M] ad. Calgary, Alberta 44.7 17.2 13.9 + U. S. N. M. 75725 [M] ad. St. Albert, Alberta 46.5 18.3 15.0 + av. 5 46.0 17.9 14.7 + + N. M. C. 6968 [F] ad. Daysland, Alberta 43.7 16.8 13.1 + U. S. N. M. 68731 [F] ad. S. Edmonton, Alberta 42.5 16.0 13.4 + A. M. N. H. 16044 [F] ad. Blindman River, Alberta 40.0 15.1 12.2 + M. V. Z. 53792 [F] ad. Grafton, North Dakota 42.8 16.5 13.6 + U. S. N. M. 75483 [F] ad. Wingard, Sask. 42.3 16.9 12.9 + av. 5 42.3 16.3 13.0 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + arthuri + 11.5 14.5 22.3 26.6 15.4 7.7 3.3 5.4 4.9 5.1 3.7 2.0 14.8 12.4 + 10.9 15.0 22.3 25.5 15.5 7.5 3.7 4.9 5.7 5.8 4.2 2.3 14.2 12.4 + 15.5 16.0 8.3 5.8 5.5 5.7 4.6 2.5 + + olivacea + 12.5 16.5 26.7 30.8 17.2 9.2 4.0 6.8 6.2 6.2 4.7 2.5 15.0 13.8 + 11.7 16.9 26.0 29.0 17.9 9.4 4.2 6.8 6.0 6.3 4.8 2.8 15.5 13.4 + 11.5 16.5 27.0 30.9 17.5 9.8 4.3 6.8 5.7 6.0 4.5 2.5 15.6 13.2 + 11.5 16.2 25.7 29.4 15.9 8.7 4.1 6.1 5.6 6.0 4.9 2.3 15.4 13.7 + 10.9 16.9 26.2 28.9 17.2 8.8 4.2 6.1 5.7 6.2 4.6 2.3 14.5 13.2 + 11.6 16.6 26.3 29.8 17.1 9.2 4.2 6.5 5.8 6.1 4.7 2.5 15.2 13.5 + + 11.9 16.0 27.3 31.2 17.9 8.7 4.2 6.4 5.9 6.3 4.9 2.5 17.0 13.4 + 10.9 15.5 24.7 28.3 16.8 8.8 3.9 6.2 5.7 5.8 4.3 2.4 15.0 13.3 + 12.1 15.5 25.1 29.1 17.9 8.8 4.5 6.6 6.0 6.0 4.7 2.6 15.0 14.2 + 11.7 16.2 24.1 29.1 15.8 8.5 4.5 6.4 5.7 6.0 4.6 2.2 15.0 13.5 + 11.7 15.8 25.3 29.4 17.1 8.7 4.3 6.4 5.8 6.0 4.6 2.4 15.5 13.6 + + 11.6 14.6 26.1 30.4 16.5 8.6 4.4 6.1 5.7 6.0 4.6 2.4 15.6 13.2 + 11.4 16.3 26.8 29.8 17.2 8.8 4.3 6.5 5.8 6.3 4.6 2.5 14.8 13.2 + 11.8 16.0 25.8 30.5 17.3 8.6 4.5 6.2 5.8 6.0 4.5 2.5 15.5 13.6 + 10.7 16.3 25.5 29.2 15.6 8.3 3.7 6.8 5.8 6.1 4.9 2.6 14.5 13.2 + + 9.8 14.5 22.4 25.7 15.5 7.8 3.8 5.8 5.6 5.8 4.2 2.2 13.5 12.3 + 9.6 14.7 22.8 26.0 15.8 8.0 3.9 5.8 5.0 5.4 4.1 2.1 14.4 12.2 + 10.6 15.3 23.4 25.8 15.3 8.2 3.6 6.1 5.7 5.8 4.4 2.1 13.2 12.0 + + peninsulae + 12.2 15.8 27.8 31.4 17.6 10.2 4.8 6.3 5.9 6.2 4.7 2.4 15.3 14.6 + + 11.0 15.8 23.7 27.1 16.4 8.4 4.3 6.3 5.7 5.7 4.1 1.9 13.3 12.9 + 9.6 14.2 6.4 5.7 5.9 4.6 2.0 12.3 + + spadix + 11.9 17.1 28.0 31.7 15.9 9.5 3.6 6.9 6.5 6.7 5.4 2.7 15.5 14.0 + 13.5 16.5 28.1 32.1 16.4 9.7 4.0 6.8 6.2 6.7 4.9 2.0 15.9 15.0 + 11.9 16.5 27.0 29.9 15.5 9.2 3.8 6.6 6.1 6.5 5.0 3.0 15.0 13.8 + 16.7 26.1 15.2 8.4 3.3 6.2 5.5 5.9 4.3 2.3 + 10.6 15.5 24.9 28.2 14.8 8.5 3.7 6.6 5.9 6.6 4.8 2.9 15.1 12.8 + + 10.7 15.0 22.9 27.0 13.8 7.7 3.1 6.6 5.6 5.6 4.4 2.1 13.0 12.3 + 10.8 15.3 24.5 26.8 14.5 8.6 3.7 5.8 5.3 5.5 4.5 2.1 13.9 13.2 + 10.5 15.0 23.3 25.6 15.2 7.6 3.2 5.9 5.4 5.7 4.1 2.0 14.0 12.2 + 10.7 14.9 23.3 26.0 14.7 8.1 3.3 5.7 5.3 5.6 4.3 2.4 13.8 12.8 + 10.7 15.1 23.4 26.4 14.6 8.0 3.3 6.0 5.4 5.6 4.3 2.2 13.7 12.6 + + longicauda + 12.1 15.3 25.3 30.8 15.2 8.9 3.4 6.9 6.3 6.5 4.9 2.6 15.0 14.6 + 12.2 15.5 31.0 6.9 6.3 6.6 5.0 2.5 13.8 + 11.4 16.0 25.7 30.4 15.7 8.7 3.9 6.3 5.7 5.9 4.4 1.9 15.5 13.4 + 11.3 14.8 24.8 29.7 15.5 8.7 3.6 6.0 5.4 5.6 4.3 2.4 15.0 13.4 + 12.4 15.6 25.0 29.4 15.2 8.3 3.4 6.6 6.2 6.4 4.6 2.0 16.0 13.6 + 11.9 15.4 25.2 30.3 15.4 8.6 3.6 6.5 6.0 6.2 4.6 2.3 15.4 13.8 + + 14.9 24.0 26.1 14.7 8.3 3.1 5.7 5.2 5.4 4.3 2.0 13.3 12.2 + 10.3 14.3 24.1 26.7 14.5 8.8 3.3 5.6 5.4 5.5 4.3 2.3 14.0 12.5 + 8.8 13.2 22.5 24.5 13.4 8.2 2.8 5.6 5.0 5.0 4.0 2.0 12.8 11.5 + 10.5 14.9 23.2 26.1 15.3 8.0 2.8 6.1 5.4 5.6 4.3 2.4 14.0 12.3 + 9.8 15.5 23.3 25.9 14.8 8.3 3.2 6.4 5.9 6.1 4.8 2.4 14.3 12.1 + 9.9 14.6 23.4 25.9 14.6 8.3 3.0 5.9 5.4 5.5 4.3 2.2 13.7 12.1 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela frenata + M. V. Z. 44568 [M] sad. Barkerville reg., B. C. 48.8 19.1 17.2 + M. V. Z. 43817 [M] sad. Isaacs Lake, British Columbia 48.8 19.5 15.8 + Cowan 443 [M] ad. Quesnel, British Columbia 46.6 17.7 14.5 + N. M. C. 2676 [M] ad. Lillooet, British Columbia 47.5 18.5 14.3 + N. M. C. 2695 [M] ad. Lillooet, British Columbia 45.0 17.5 15.9 + + M. C. Z. 9058 [F] ad. Source Kettle River, B. C. 41.7 16.4 12.8 + M. V. Z. 62791 [F] ad. Beaverfoot Range, B. C. 42.0 16.4 13.5 + + Mustela frenata + U. S. N. M. 186451 [M] ad. Custer, South Dakota 40.9 15.0 13.8 + + A. M. N. H. 7440/9136 [M] ad. Hill City, South Dakota 41.0 15.6 13.7 + + A. M. N. H. 7441 [F] ad. Black Hills, South Dakota 37.6 14.1 12.2 + + Mustela frenata + M. V. Z. 55211 [M] ad. near Parks, Arizona 40.4 15.5 12.5 + M. V. Z. 65231 [M] ad. Alpine, Arizona 39.6 15.1 12.2 + U. S. N. M. 248993 [M] ad. Kaibab Plat., Arizona 40.4 15.6 12.9 + + A. M. N. H. 2490/1886 [F] ad. S. F. Forest, Arizona 35.5 13.8 10.8 + + Mustela frenata + average} {43.6 16.6 13.7 + maximum} [M] ad. 25 Sierra Nevada, California {46.1 17.6 14.9 + minimum} {40.6 15.2 12.5 + + average} {43.7 16.5 13.9 + maximum} [M] ad. 10 S and SW Colorado {44.6 17.3 14.8 + minimum} {41.6 16.0 12.8 + + average} {38.2 14.7 11.8 + maximum} [F] ad. 10 California {39.5 15.1 12.4 + minimum} {36.7 13.9 11.0 + + average} {38.5 14.8 12.2 + maximum} [F] ad. 14 Colorado {39.7 15.4 13.1 + minimum} {36.1 14.0 11.1 + + Mustela frenata + average} {40.5 15.2 12.3 + maximum} [M] ad. 6 NE Oregon {41.8 16.4 12.7 + minimum} {39.3 14.4 11.9 + U. S. N. M. 212423 [F] ad. Vale, Oregon 37.4 + + Mustela frenata + average} {43.7 16.7 13.7 + maximum} [M] ad. 22 Mt. Adams, Washington {47.7 18.0 15.4 + minimum} {40.0 15.6 12.5 + + average} {37.7 14.3 11.5 + maximum} [F] ad. 11 Mt. Adams, Washington {39.0 14.9 12.0 + minimum} {37.1 13.3 10.8 + + Mustela frenata + U. O. 3709 [M] ad. Mt. Ashland, Oregon 45.8 17.2 14.1 + U. S. N. M. 65930 [M] ad. Siskiyou, Oregon 42.6 15.9 14.0 + M. V. Z. 13778 [M] ad. Jackson Lake, California 45.5 18.0 13.2 + M. V. Z. 13779 [M] ad. Jackson Lake, California 43.8 16.9 12.7 + av. 4 44.4 17.0 13.5 + + M. V. Z. 52144 [F] ad. S. Fork Mt., California 38.2 14.7 10.8 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + oribasus + 13.0 17.7 28.5 32.2 17.4 9.8 3.2 6.8 6.2 6.4 5.3 2.8 15.5 14.6 + 12.0 17.3 28.4 31.7 15.9 9.7 4.0 6.9 6.3 6.4 5.3 2.9 15.5 13.7 + 10.1 15.9 26.4 30.7 15.8 9.2 3.9 6.2 5.8 6.1 4.6 2.5 15.0 13.1 + 10.3 15.9 25.5 30.1 15.2 9.1 3.8 6.4 5.9 6.3 4.6 2.5 15.0 13.1 + 11.8 15.9 25.6 31.0 15.2 8.9 3.0 6.2 5.6 6.1 4.7 2.9 15.0 13.4 + + 10.5 14.8 24.9 26.7 14.3 8.2 2.8 6.1 5.5 5.7 4.5 2.5 14.0 12.6 + 10.2 15.1 24.4 27.0 14.3 8.1 3.1 5.9 5.5 5.6 4.3 2.2 14.0 12.3 + + alleni + 11.0 14.1 22.0 27.7 13.7 7.9 3.0 5.7 4.9 5.3 3.9 2.5 14.3 12.3 + + 11.0 13.9 23.3 25.7 13.6 8.3 2.3 5.1 4.8 5.2 4.2 1.9 13.7 12.0 + + 9.1 13.2 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 + 14.2 23.3 26.0 14.3 8.9 3.1 5.7 5.2 5.5 4.0 2.2 13.5 12.1 + 9.7 13.7 22.4 25.6 13.7 8.3 3.2 5.4 4.9 5.1 4.0 2.1 14.0 11.4 + 9.8 14.5 22.9 26.3 14.1 8.4 3.2 5.8 5.5 5.5 4.0 1.7 14.8 12.0 + + 8.6 12.9 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 + 10.7 15.1 23.9 28.0 15.0 8.4 3.4 5.9 5.4 5.6 4.3 2.2 14.4 12.5 + 12.0 16.2 26.1 31.4 15.9 9.0 4.0 6.4 5.8 6.1 4.8 2.7 15.2 14.8 + 9.9 14.0 22.1 25.0 14.4 7.8 2.9 5.5 4.9 5.1 3.9 1.8 13.7 11.6 + + 10.6 15.0 24.2 27.9 15.1 8.6 3.5 6.1 5.5 5.7 4.5 2.1 14.5 12.6 + 11.1 16.4 25.3 29.3 15.8 9.2 4.0 6.8 5.9 6.0 4.8 2.6 15.6 13.0 + 9.9 13.8 23.1 26.5 14.4 8.2 3.0 5.7 5.2 5.3 4.0 1.9 13.9 12.1 + + 9.2 13.4 20.9 23.1 13.4 7.3 2.9 5.2 4.8 5.0 3.8 1.7 12.3 11.0 + 9.9 14.2 21.8 23.4 14.1 7.9 3.2 5.6 5.2 5.4 4.1 2.0 13.0 11.8 + 8.6 12.6 20.1 22.4 12.7 6.8 2.6 4.9 4.5 4.7 3.6 1.5 11.4 10.0 + + 9.3 13.4 20.6 23.1 13.4 7.6 2.8 5.4 4.9 5.1 3.9 1.9 12.9 11.1 + 10.2 14.4 22.1 24.6 13.9 8.1 3.2 5.6 5.3 5.4 4.1 2.3 13.8 11.8 + 8.5 12.5 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 + 9.6 13.7 22.1 25.6 14.1 7.8 3.3 5.5 5.0 5.3 3.9 2.0 13.5 11.8 + 10.0 14.4 23.3 27.3 15.0 8.4 3.5 5.9 5.3 5.8 4.1 2.6 14.3 12.2 + 9.2 13.1 20.5 25.0 12.3 7.2 3.2 5.0 4.7 5.0 3.6 1.6 12.6 11.4 + 9.2 19.5 22.0 13.1 6.9 3.0 5.4 4.9 5.0 3.7 1.8 12.3 + + washingtoni + 10.5 15.5 23.4 27.0 14.6 8.0 3.1 5.9 5.4 5.6 4.3 2.2 14.4 12.6 + 12.0 16.5 26.4 29.6 15.8 8.7 3.4 6.5 6.0 6.1 4.8 2.6 15.8 13.7 + 9.0 14.5 22.1 24.6 13.5 7.6 2.7 5.4 4.9 5.0 4.0 1.7 13.1 11.5 + + 8.8 12.8 20.2 22.5 12.9 7.2 2.8 5.1 4.7 4.8 3.9 1.9 12.2 10.4 + 9.3 13.2 21.1 24.5 13.6 7.7 3.0 5.6 4.9 5.0 4.2 2.1 13.0 11.0 + 8.2 12.1 19.4 21.3 12.3 6.8 2.3 4.7 4.4 4.2 3.4 1.7 11.2 9.8 + + saturata + 11.1 15.8 26.0 27.9 15.1 8.9 3.9 6.1 5.5 5.9 4.3 2.3 14.2 12.6 + 10.9 14.4 24.5 27.7 14.7 8.5 4.0 5.7 5.0 5.5 4.1 1.9 14.8 12.4 + 10.0 16.2 24.7 27.4 14.6 8.7 3.2 6.3 5.9 6.2 4.6 2.5 14.6 12.8 + 9.9 15.3 24.2 26.9 15.1 8.6 3.3 5.6 5.2 5.5 4.0 2.2 14.2 12.3 + 10.2 15.4 24.9 27.5 14.9 8.7 3.6 5.9 5.4 5.8 4.3 2.2 14.5 12.5 + + 8.1 13.5 19.8 21.8 12.4 7.2 2.7 5.0 4.6 4.9 3.7 1.8 12.2 10.4 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela frenata + average} {45.8 17.5 14.2 + maximum} [M] ad. 10 Tillamook Co., Oregon {48.0 18.9 15.0 + minimum} {42.4 16.1 13.2 + + Walker 392 [F] ad. Blaine, Oregon 39.7 15.1 12.8 + Walker 185 [F] ad. Blaine, Oregon 37.8 13.9 11.9 + Walker 89 [F] ad. Blaine, Oregon 38.3 14.3 11.3 + Walker 45 [F] ad. Tillamook, Oregon 37.8 14.1 11.0 + + Mustela frenata + M. V. Z. 11747 [M] ad. Eureka, California 44.0 16.8 13.2 + C. A. C. 3907 [M] ad. Requa, California 41.8 16.0 13.5 + F. M. N. H. 9595 [M] ad. Gold Beach, Oregon 43.0 16.9 13.1 + U. S. N. M. 32019 [M] sad. Grants Pass, Oregon 42.9 16.3 13.7 + + M. V. Z. 34325 [F] ad. Carlotta, California 37.8 14.7 10.8 + U. O. 1413 [F] ad. 13 mi. S Grants Pass, Oregon 39.4 14.7 12.0 + + Mustela frenata + M. V. Z. 19720 [M] ad. Point Arena, California 49.0 18.5 15.2 + M. V. Z. 19722 [M] ad. Point Arena, California 48.5 18.9 14.4 + M. V. Z. 19718 [M] ad. Gualala, California 45.7 17.2 13.7 + av. 3 47.7 18.2 14.4 + M. V. Z. 19714 [M] ad. 6 mi. W Inverness, California 48.2 18.4 13.9 + M. V. Z. 19715 [M] ad. 6 mi. W Inverness, California 48.2 18.9 14.5 + M. V. Z. 19716 [M] ad. 5 mi. W Inverness, California 46.5 17.5 13.8 + av. 3 47.6 18.3 14.1 + F. M. N. H. 9598 [M] ad. Nicasio, California 46.5 18.5 14.0 + M. C. Z. 8632 [M] ad. Nicasio, California 44.2 17.8 14.0 + M. C. Z. 5459 [M] ad. Point Reyes, California 39.7 15.5 13.3 + M. V. Z. 40302 [M] ad. 4 mi. N Vallejo, California 45.8 17.7 13.0 + av. 4 44.1 17.4 13.6 + + M. V. Z. 19723 [F] ad. Point Arena, California 42.3 16.0 12.3 + U. S. N. M. 135010 [F] ad. Point Reyes, California 38.7 15.3 11.0 + F. M. N. H. 9597 [F] ad. Point Reyes, California 39.5 15.2 11.1 + U. S. N. M. 91764 [F] ad. Point Reyes, California 38.7 15.4 12.0 + + Mustela frenata + Walker 1440 [M] ad. 5 mi. W Fresno, California 43.9 16.5 13.8 + A. N. S. P. 11863 [M] ad. Fresno, California 43.4 16.8 12.9 + Wisconsin U. 4232 [M] ad. Selma, California 43.7 16.2 13.3 + + Snyder 2626 [F] ad. Selma, California 39.4 15.0 12.0 + M. V. Z. 79640 [F] ad. Tegner School, California 43.4 16.5 12.6 + + Mustela frenata + Stanford U. 863 [M] ad. Palo Alto, California 48.1 18.7 15.0 + F. M. N. H. 6559 [M] ad. Palo Alto, California 48.0 18.7 13.9 + Stanford U. 1651 [M] ad. Menlo Park, California 47.1 17.8 13.3 + Stanford U. 487 [M] ad. Palo Alto, California 46.5 18.3 13.4 + F. M. N. H. 7031 [M] ad. Palo Alto, California 46.1 18.1 14.5 + Stanford U. 236 [M] ad. Menlo Park, California 46.1 17.8 13.5 + av. 6 47.0 18.2 13.9 + + M. V. Z. 5851 [F] ad. Hayward, California 40.7 15.3 11.8 + M. V. Z. 30327 [F] ad. Palo Alto, California 41.2 16.2 11.1 + U. S. N. M. 43574 [F] ad. Morro, California 42.2 16.1 12.2 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + altifrontalis + 11.2 15.9 25.1 29.2 15.6 8.5 3.5 6.1 5.5 5.7 4.5 2.5 15.4 13.6 + 12.0 16.8 26.0 31.6 16.5 9.0 3.6 6.5 6.0 6.0 4.9 2.8 16.2 14.0 + 10.0 14.8 23.9 26.0 15.0 7.7 3.2 5.6 5.0 5.2 4.0 2.1 14.5 12.4 + + 10.1 13.7 21.8 24.0 13.8 7.7 5.2 5.0 5.0 3.7 1.9 13.5 11.8 + 9.5 13.4 20.8 22.7 13.4 7.3 2.8 4.9 4.5 4.7 3.2 1.5 12.6 10.7 + 9.5 12.7 19.8 22.7 13.2 7.0 3.1 4.4 4.7 3.8 1.9 13.4 11.3 + 8.6 12.5 20.1 23.2 13.0 7.3 4.9 4.7 4.8 3.8 2.0 13.5 11.0 + + oregonensis + 10.7 15.3 24.3 28.0 14.6 8.3 3.4 6.0 5.4 5.8 4.3 2.2 13.6 12.7 + 10.9 14.0 22.9 26.6 14.2 7.6 3.1 5.8 5.3 5.4 4.0 2.2 13.4 12.3 + 10.5 15.5 23.4 26.2 15.1 8.2 3.3 5.8 5.2 5.5 4.4 2.0 14.7 12.5 + 10.5 15.7 23.2 26.5 15.2 8.8 4.0 5.5 4.8 5.2 4.3 2.1 14.0 12.5 + + 8.8 13.0 20.7 23.2 13.4 7.5 2.2 5.3 5.0 5.5 4.0 1.8 13.7 11.0 + 9.7 13.9 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 + 11.7 16.8 27.5 33.2 16.2 9.3 3.6 6.5 5.6 6.0 4.8 2.6 14.2 14.2 + 10.5 17.2 26.8 32.+ 16.0 8.5 3.8 6.5 5.6 6.2 5.0 2.9 14.7 13.8 + 10.5 15.5 26.4 31.7 15.5 8.5 3.1 6.5 5.7 5.8 4.9 2.3 13.4 14.2 + 10.9 16.5 26.9 32.7 15.9 8.8 3.5 6.5 5.7 6.0 4.9 2.6 14.1 14.1 + 11.3 17.2 26.2 30.5 15.7 8.5 3.5 6.5 5.6 6.0 4.8 2.6 14.2 14.2 + 11.4 16.6 26.5 30.0 16.3 8.7 3.5 6.3 5.6 6.2 5.0 2.9 14.7 13.8 + 11.0 15.9 25.2 30.3 15.4 7.9 3.5 6.1 5.7 5.8 4.9 2.3 13.4 14.2 + 11.2 16.6 26.0 30.3 15.8 8.4 3.5 6.3 5.7 6.0 4.9 2.6 14.1 14.1 + 10.5 15.4 25.2 30.5 15.0 8.8 3.5 6.9 5.9 6.3 5.2 2.7 15.2 13.0 + 11.4 16.9 24.4 27.6 15.7 8.2 3.7 6.4 5.7 5.9 5.0 2.6 12.6 13.7 + 9.0 13.4 23.1 26.5 13.7 7.7 3.2 6.1 5.1 5.5 4.4 2.5 13.6 12.1 + 10.7 15.9 24.4 26.9 15.1 8.0 2.8 6.3 5.7 6.2 4.8 2.3 14.5 13.0 + 10.4 15.4 24.3 27.9 14.9 8.2 3.3 6.4 5.6 6.0 4.9 2.5 14.0 13.0 + + 12.2 14.0 23.6 25.5 14.5 8.3 2.9 5.0 5.2 4.0 1.9 13.4 11.3 + 8.7 14.5 21.0 23.7 13.0 7.5 3.0 5.1 4.9 5.4 3.9 2.3 12.9 10.8 + 8.2 13.1 20.2 22.8 12.7 6.8 2.7 5.3 4.9 5.0 3.9 1.9 14.7 10.8 + 9.7 12.7 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 + 10.4 14.1 23.8 28.5 14.7 7.9 3.2 5.7 5.2 5.3 4.2 2.1 14.8 12.6 + 10.0 14.8 24.0 27.5 14.5 7.5 5.8 5.3 5.5 4.4 2.0 13.7 12.7 + 9.9 14.5 23.7 27.1 15.2 8.5 3.7 5.8 5.3 5.4 4.2 2.2 13.8 12.3 + + 8.9 13.2 21.3 24.3 13.7 7.2 5.3 4.0 13.0 11.4 + 9.3 15.1 22.8 24.9 15.0 7.5 3.0 5.6 5.5 5.9 4.5 2.0 13.3 12.2 + + nigriauris + 11.2 17.5 32.9 15.5 6.5 6.4 6.7 5.2 2.4 14.3 + 11.0 16.2 27.0 29.6 15.7 8.7 3.4 6.2 5.8 6.0 4.8 2.7 14.0 14.1 + 9.8 14.9 25.6 30.0 15.0 8.0 3.3 6.5 6.0 6.5 4.6 2.4 14.6 13.8 + 11.0 16.1 25.2 31.1 15.1 8.0 2.9 6.3 5.7 6.0 4.9 2.5 14.5 14.3 + 10.7 15.1 26.0 29.5 14.9 8.3 2.4 6.3 6.0 6.1 4.6 2.6 15.0 13.2 + 10.4 15.5 29.6 15.6 8.5 6.0 5.6 5.7 4.8 2.5 14.0 + 10.7 15.9 26.0 30.5 15.3 8.3 3.0 6.3 5.9 6.2 4.8 2.5 14.5 14.0 + + 8.8 13.9 21.8 24.9 14.2 7.9 3.0 5.2 4.9 4.9 4.1 1.8 12.4 11.0 + 8.6 14.3 21.7 24.8 13.4 7.8 3.0 5.3 5.0 5.4 4.1 2.1 12.6 11.0 + 8.9 14.2 22.7 24.3 15.1 8.1 2.9 5.5 5.4 5.6 4.2 2.0 13.2 11.4 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela frenata + M. V. Z. 3257 [M] ad. San Diego, California 44.4 17.8 13.4 + U. S. N. M. 52701 [M] ad. El Vido, California 43.7 16.9 12.7 + U. S. N. M. 52702 [M] ad. El Cajon, California 43.2 16.7 13.9 + M. V. Z. 3258 [M] ad. San Diego, California 42.5 16.6 14.1 + Stephens 20 [M] ad. San Jacinto Plain, California 41.9 16.9 13.1 + S. D. M. 7098 [M] ad. Jamacha, California 47.0 18.5 15.6 + av. 6 43.8 17.2 13.8 + + Stephens 22 [F] ad. Santa Ysabel, California 40.1 15.5 11.7 + Stephens 19 [F] ad. Ballena, California 40.0 15.0 12.3 + S. D. M. 6748 [F] ad. Santa Ysabel, California 42.0 16.1 12.9 + S. D. M. 7194 [F] ad. Jamacha, California 39.8 15.0 11.8 + av. 4 40.5 15.4 12.2 + + Mustela frenata + C. A. S. 335 [M] ad. Buttonwillow, California 48.6 18.8 15.2 + C. A. S. 337 [M] ad. Buttonwillow, California 48.6 18.8 15.0 + M. V. Z. 16668 [M] ad. Buttonwillow, California 48.1 18.8 14.8 + U. S. N. M. 137935 [M] ad. Buttonwillow, California 47.2 18.2 14.0 + C. A. S. 336 [M] ad. Buttonwillow, California 47.0 18.3 14.5 + C. A. S. 338 [M] ad. Buttonwillow, California 46.0 17.4 14.7 + av. 6 47.6 18.4 14.7 + + Mustela frenata + M. V. Z. 25907 [M] ad. 2 mi. N Independence, Calif. 44.7 17.3 13.3 + + Mustela frenata + A. M. N. H. 14821 [M] ad. Kerr Co., Texas 54.0 19.2 16.0 + M. C. Z. 15476 [M] yg. Kerr Co., Texas 53.3 18.9 16.3 + Baylor U. 2017 [M] sad. 5 mi. N Waco, Texas 52.0 18.3 16.5 + + Mustela frenata + Kansas U. 1485 [M] ad. Liberal, Kansas 50.5 17.9 15.4 + U. S. N. M. 131582 [M] ad. Berino, New Mexico 47.7 17.5 15.0 + + U. S. N. M. 36482 [F] ad. Tombstone, Arizona 45.5 16.5 12.8 + U. S. N. M. 230973 [F] ad. Willcox, Arizona 42.5 15.1 12.9 + U. S. N. M. 225629 [F] ad. Albuquerque, New Mexico 40.8 15.0 12.4 + + Mustela frenata + M. C. Z. 240 [M] ad. Brownsville, Texas 49.4 17.4 14.9 + A. N. S. P. 724 [M] ad. Brownsville, Texas 50.1 17.9 15.5 + U. S. N. M. 58684 [M] ad. Brownsville, Texas 48.2 17.3 14.2 + U. S. N. M. 63857 [M] ad. Brownsville, Texas 48.6 18.0 13.7 + U. S. N. M. 44976 [M] ad. Brownsville, Texas 50.9 18.0 + av. 5 49.4 17.7 14.6 + + A. M. N. H. 24405 [F] ad. Brownsville, Texas 47.3 16.2 12.5 + U. S. N. M. 58685 [F] sad. Brownsville, Texas 41.3 15.0 11.8 + + U. S. N. M. 36362/48732 [F] sad. Brownsville, Texas 42.5 15.4 12.5 + av. 3 43.7 15.5 12.3 + B. Z. M. 991 [F] ad. México 15.5 + B. Z. M. 992 [F] ad. México 13.9 10.7 + + M. C. Z. 20841 [M] ad. Miquihana, Nuevo León 50.2 18.0 15.3 + U. S. N. M. 50826 [M] ad. Tlalpam, D. F. 51.3 18.3 15.1 + + Mustela frenata + U. S. N. M. 125972 [M] ad. Los Reyes, Michoacán 51.2 17.5 15.0 + + U. S. N. M. 34914/47179 [M] ad. Pátzcuaro, Michoacán 18.7 14.3 + + A. M. N. H. 26153 [F] ad. Artenkiki, Jalisco 44.5 16.0 12.7 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + latirostra + 10.2 15.0 23.5 27.6 15.6 8.4 3.9 6.0 5.5 5.5 4.3 2.2 14.0 12.8 + 9.9 14.3 24.0 14.7 8.7 4.0 5.7 5.6 5.6 4.2 2.0 14.2 13.4 + 10.0 15.0 24.0 27.2 15.1 8.2 4.2 5.8 5.3 5.5 4.2 2.5 14.0 12.5 + 12.1 14.7 24.0 28.7 15.1 8.1 3.9 5.9 5.6 5.5 4.4 2.5 14.3 13.5 + 10.6 14.0 24.0 28.2 14.7 8.0 3.2 6.0 5.5 5.7 4.2 2.1 14.0 12.9 + 11.3 16.8 24.8 29.2 16.7 8.9 3.8 6.0 5.8 6.2 4.8 2.2 14.0 13.2 + 10.7 15.0 24.1 28.2 15.3 8.4 3.8 5.9 5.6 5.7 4.4 2.3 14.1 13.1 + + 9.3 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 + 9.3 13.5 21.6 23.5 13.7 8.1 3.2 5.1 4.9 5.0 4.0 2.1 12.0 11.2 + 9.4 14.2 22.0 24.5 14.0 7.8 3.0 5.8 5.3 5.7 4.5 2.0 13.4 11.5 + 9.1 13.7 20.2 23.7 13.7 7.0 2.9 5.2 4.8 4.9 3.9 1.9 12.6 11.5 + 9.3 13.6 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 + 11.6 17.2 27.4 31.4 16.7 9.1 3.6 6.6 5.9 6.4 5.1 2.7 16.2 13.7 + 11.8 17.2 27.1 32.7 16.6 9.0 3.5 6.6 6.3 6.3 5.0 2.4 16.5 14.3 + 12.0 17.1 27.7 31.2 16.4 9.2 3.6 6.4 5.7 5.9 5.1 2.8 15.2 14.0 + 10.3 16.0 26.1 29.5 15.6 8.5 3.3 6.0 5.6 5.9 4.1 2.0 15.0 12.9 + 11.5 16.5 27.0 29.5 15.5 9.1 3.2 6.3 5.6 5.9 5.0 2.3 15.0 13.0 + 11.3 16.1 26.3 32.6 15.9 8.9 3.7 6.1 5.7 5.8 4.6 2.2 15.0 13.3 + 11.4 16.7 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 + 10.8 15.9 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 + 12.6 17.2 28.6 35.1 17.5 8.0 3.5 6.9 6.5 6.7 5.0 2.5 16.7 15.1 + 12.8 18.3 28.2 34.8 18.0 8.3 4.0 6.4 6.3 6.5 4.8 2.4 16.0 15.0 + 13.5 16.7 29.2 18.0 9.1 6.6 6.3 6.4 4.9 2.6 16.3 15.6 + + neomexicana + 12.4 16.4 28.3 35.0 17.0 9.0 4.0 6.3 5.9 6.0 4.9 2.4 15.5 14.3 + 11.1 15.9 26.3 31.2 16.5 7.9 3.4 6.4 5.7 6.3 4.7 2.5 15.8 13.5 + + 9.7 15.5 22.3 26.6 15.2 7.5 3.3 5.9 5.6 5.9 4.5 2.2 13.8 12.0 + 9.9 14.0 22.6 26.5 14.5 7.0 3.1 5.5 5.3 5.6 4.1 1.8 14.0 12.0 + 9.4 13.3 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 + 11.6 15.4 25.9 33.0 16.7 7.9 4.3 5.9 5.9 6.1 4.5 2.1 15.0 13.8 + 12.3 15.5 27.0 32.2 16.5 8.5 4.1 6.5 6.2 6.3 4.8 2.7 16.0 14.0 + 11.0 15.3 27.2 31.0 16.6 8.3 4.2 6.3 5.9 6.1 4.8 2.7 15.5 13.5 + 11.1 16.5 26.0 31.0 16.0 8.2 4.8 6.5 5.7 6.1 4.8 2.6 15.0 13.6 + 16.9 26.9 16.6 7.9 3.4 6.5 5.6 6.1 4.7 2.3 16.0 13.4 + 11.5 15.9 26.6 31.8 16.3 8.2 4.2 6.4 5.9 6.2 4.7 2.5 15.5 13.7 + + 10.0 14.6 5.9 5.4 5.4 4.1 2.1 12.2 + 9.5 12.8 22.7 27.0 14.0 6.9 3.3 5.5 5.2 5.4 4.1 2.0 14.0 11.7 + + 10.0 14.3 23.8 26.7 14.3 7.5 3.2 5.9 5.5 5.7 4.2 2.0 13.5 11.8 + 9.8 13.9 23.3 26.9 14.2 7.2 3.3 5.8 5.4 5.5 4.1 2.0 13.8 11.9 + 12.2 13.8 22.8 27.0 13.7 5.7 5.2 5.5 4.4 2.2 12.7 + 8.9 12.9 21.1 13.0 6.6 2.9 4.9 4.5 4.7 3.7 1.7 12.3 10.8 + + 12.1 16.3 28.0 32.0 16.9 8.8 3.3 4.6 2.2 15.0 + 12.1 17.5 27.7 33.5 16.3 8.4 3.5 6.7 5.9 6.4 4.7 2.7 15.3 14.6 + + leucoparia + 12.0 16.0 28.3 32.9 16.0 7.7 3.5 5.9 5.7 5.7 4.3 2.2 15.5 14.0 + + 16.8 6.8 6.5 6.8 5.0 2.6 14.3 + + 10.0 14.4 22.4 26.3 15.0 7.0 3.2 5.9 5.5 6.0 4.5 2.1 14.0 11.9 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela frenata + U. S. N. M. 68197 [M] ad. Cerro San Felipe, Oaxaca 49.2 17.3 13.9 + + U. S. N. M. 54278 [F] ad. Type specimen 43.5 15.5 12.3 + + Mustela frenata + U. S. N. M. 133253 [M] ad. 20 mi. SE Teopisca, Chiapas 50.4 18.0 15.0 + U. S. N. M. 133254 [M] ad. 20 mi. SE Teopisca, Chiapas 49.6 17.5 13.8 + U. S. N. M. 77519 [M] ad. Pinabete, Chiapas 50.7 18.3 + F. M. N. H. 15953 [M] ad. near Tecpám, Guatemala 50.0 17.0 13.5 + Dickey 12523 [M] ad. Los Esesmiles, Salv. 51.3 17.5 14.5 + av. 5 50.4 17.7 14.2 + + Mustela frenata + F. M. N. H. 14063 [M] ad. Achotal, Veracruz 54.1 19.2 15.6 + + U. S. N. M. 132528 [F] ad. Pérez, Veracruz 43.5 15.3 12.5 + + Mustela frenata + U. S. N. M. 54994 [M] ad. Jico, Veracruz 47.8 17.2 13.7 + + A. M. N. H. 12764/11058 [M] sad. Jalapa, Veracruz 45.5 16.8 13.7 + M. C. Z. 6514 [M] sad. Orizaba, Veracruz *46.0 16.4 13.2 + + U. S. N. M. 54993 [F] ad. Jico, Veracruz 36.0 13.0 11.0 + U. S. N. M. 1060 [F] ad. México 39.0 14.0 11.0 + M. C. Z. 2605 [F] yg. Jalpa, Veracruz 38.7 13.8 10.3 + F. M. N. H. 14050 [F] yg. Xuchil, Veracruz 39.0 14.2 11.6 + + Mustela frenata + U. S. N. M. 100041 [M] sad. Teapa, Tabasco 46.1 17.0 14.4 + U. S. N. M. 132997 [M] sad. San Vicente, Chiapas 45.3 16.7 13.9 + U. S. N. M. 132996 [M] sad. San Cristóbal, Chiapas 16.9 13.3 + av. 3 45.7 16.9 13.9 + + U. S. N. M. 218036 [F] sad. State of Chiapas 40.0 14.6 11.6 + U. S. N. M. 65422 [F] sad. Catemaco, Veracruz 40.4 14.5 11.5 + + Mustela frenata + A. M. N. H. 30754 [M] sad. Matagalpa, Nicaragua 44.8 17.2 12.8 + A. M. N. H. 28331 [M] ad. Matagalpa, Nicaragua 44.8 16.7 13.6 + A. M. N. H. 29280 [M] sad. San Rafel Del Norte 45.5 17.2 13.5 + + Mustela frenata + U. S. N. M. 11408 [M] ad. Costa Rica *49.0 18.3 15.2 + B. M. 3216 [M] ad. Vic. San José, Costa Rica 49.3 18.6 15.0 + + U. S. N. M. 13770/37149 [M] yg. San José, Costa Rica 48.2 18.0 14.0 + N. H. R. S. 1-138 [M] ad. Azahar Cartago, Costa Rica 47.8 17.4 14.9 + + B. Z. M. A 59.13 [F] sad. Irazú, 3000M., Costa Rica 38.8 14.4 12.5 + + Mustela frenata + M. C. Z. 10112 [M] ad. Boquete, Panamá 48.3 17.1 15.0 + A. M. N. H. 18848 [M] ad. Boquete, Panamá 44.5 16.3 14.0 + M. C. Z. 10113 [M] ad. Boquete, Panamá 42.8 15.8 14.0 + av. 3 45.2 16.4 14.3 + + U. S. N. M. 170970 [F] sad. near Gatún, Panamá 41.3 15.3 12.5 + A. N. S. P. 18434 [F] ad. Siola, Panamá 39.3 14.2 11.3 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + perotae + 11.7 15.9 25.0 29.2 15.5 6.8 2.5 6.1 5.3 5.8 4.1 2.1 15.5 13.3 + + 10.3 14.0 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 + 12.5 15.9 26.5 32.3 15.6 7.1 2.9 6.4 5.9 6.2 5.0 2.5 16.5 13.7 + 10.9 16.2 26.4 31.8 15.8 7.4 3.0 6.1 5.5 5.8 4.5 2.5 15.0 13.7 + 26.5 31.8 15.3 7.5 2.9 6.6 5.8 6.1 5.0 2.4 16.0 + 12.0 16.1 26.2 31.5 15.5 7.0 5.9 5.6 6.0 4.5 2.2 + 11.7 16.2 26.7 30.6 15.8 7.7 3.1 6.3 5.5 5.8 4.5 2.3 16.5 14.1 + 11.8 16.1 26.5 31.6 15.6 7.3 3.0 6.3 5.7 6.0 4.7 2.4 16.0 13.8 + + macrophonius + 12.9 17.8 28.5 33.6 16.8 7.6 2.9 7.1 6.4 6.8 5.2 2.9 16.8 15.3 + + 10.2 14.5 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 + 10.7 15.8 24.5 28.2 15.5 6.3 2.9 6.2 5.6 5.9 4.5 2.2 15.5 13.7 + + 11.4 16.0 24.0 30.0 15.4 6.7 3.0 6.4 5.7 5.9 4.7 2.2 15.2 13.6 + 15.5 15.0 8.0 6.3 5.7 6.0 5.0 2.5 16.5 14.5 + + 9.2 12.1 19.8 22.6 12.4 6.1 2.5 5.0 4.5 4.7 4.0 1.7 12.5 11.6 + 9.6 21.0 22.5 4.9 4.6 4.9 3.9 1.9 11.5 + 8.3 12.6 19.4 12.7 5.8 4.9 4.5 4.6 3.5 1.5 13.7 10.9 + 9.3 13.0 20.5 23.5 13.5 6.1 5.3 4.9 5.3 4.4 2.0 15.3 11.5 + + perda + 11.1 15.3 24.4 28.5 14.9 6.6 2.2 6.4 5.5 6.0 4.6 2.5 16.4 13.8 + 12.0 15.5 24.0 27.4 14.5 6.7 2.8 5.5 5.1 5.4 4.2 2.2 15.8 13.0 + 10.6 16.0 6.4 5.5 6.0 4.5 2.5 13.0 + 11.2 15.6 24.2 28.0 14.7 6.7 2.5 6.1 5.4 5.8 4.4 2.4 16.1 13.3 + + 9.4 13.5 20.2 23.2 13.0 5.6 2.1 5.2 4.8 5.0 3.7 1.7 13.2 11.4 + 9.1 13.2 21.2 23.0 13.7 6.5 2.1 4.8 4.7 5.2 3.8 1.9 14.2 10.8 + + nicaraguae + 10.5 15.0 23.4 25.5 14.0 6.5 2.9 6.3 6.0 6.3 4.6 2.5 15.5 13.4 + 11.5 15.2 22.7 26.8 15.2 6.7 5.8 5.1 5.5 4.4 2.4 14.7 13.4 + 11.0 15.5 23.5 27.4 14.2 6.8 6.3 6.0 6.3 4.7 2.5 15.5 13.1 + + costaricensis + 9.4 *18.0 26.0 30.5 15.0 7.5 6.5 5.9 6.4 5.0 2.6 16.7 14.2 + 12.7 16.9 25.9 31.3 15.3 7.3 2.9 6.6 6.0 6.5 4.9 2.5 16.8 15.1 + + 11.7 16.0 24.8 29.0 15.0 7.2 2.7 7.0 6.0 6.5 4.8 2.9 16.5 14.0 + 11.7 15.5 24.3 *30.1 15.1 7.5 6.6 5.9 6.2 4.8 2.5 + + 10.1 14.2 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 + 12.5 15.7 25.1 28.3 14.1 7.0 2.7 6.3 5.7 5.9 4.5 2.5 16.7 14.0 + 12.0 15.0 22.5 29.1 14.4 6.1 2.7 6.2 5.7 6.0 4.5 2.3 16.0 13.5 + 12.0 15.5 27.1 13.6 5.5 6.1 5.4 5.7 4.3 2.2 15.8 13.5 + 12.2 15.4 23.8 28.2 14.0 6.2 2.7 6.2 5.6 5.9 4.4 2.3 16.3 13.7 + + 10.4 14.3 23.0 26.8 12.5 6.1 5.7 5.3 5.5 4.1 2.1 15.5 12.5 + 9.4 12.5 20.1 22.5 13.0 6.0 5.0 4.6 5.0 3.7 2.0 13.2 10.8 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela frenata + A. M. N. H. 33154 [M] ad. Mérida, Venezuela 44.3 16.7 14.2 + U. S. N. M. 137517 [M] ad. Mérida, Venezuela 43.4 16.0 14.1 + U. S. N. M. 172959 [M] ad. Mérida, Venezuela 44.0 16.5 14.1 + A. M. N. H. 24309 [M] ad. Mérida, Venezuela 44.2 16.1 13.7 + A. M. N. H. 33155 [M] ad. Mérida, Venezuela 42.3 16.7 13.8 + av. 5 43.6 16.4 14.0 + + U. S. N. M. 123341 [M] sad. Mérida, Venezuela 43.5 16.9 13.8 + U. S. N. M. 137516 [M] sad. Mérida, Venezuela 43.6 16.2 13.5 + U. S. N. M. 143667 [M] sad. Mérida, Venezuela 15.4 12.5 + + U. S. N. M. 143666 [F] ad. Mérida, Venezuela 38.2 14.1 11.9 + U. S. N. M. 143665 [F] ad. Mérida, Venezuela 36.4 13.4 11.5 + A. M. N. H. 24308 [F] ad. Mérida, Venezuela 36.3 13.4 11.4 + A. M. N. H. 24311 [F] ad. Mérida, Venezuela 37.7 13.9 11.4 + av. 4 37.2 13.7 11.6 + + A. M. N. H. 21343 [F] sad. Mérida, Venezuela 37.3 13.6 11.1 + A. M. N. H. 24310 [F] sad. Mérida, Venezuela 35.0 12.6 11.1 + + Mustela frenata + U. S. N. M. 241314 [M] ad. Choachí, Colombia *46.0 17.4 13.6 + U. S. N. M. 239946 [M] ad. Choachí, Colombia 16.1 12.7 + A. M. N. H. 35805 [M] ad. Quetame, Colombia 45.5 16.6 13.7 + av. 3 45.8 16.7 13.3 + + U. S. N. M. 241313 [M] yg. Bogotá, Columbia 44.7 16.7 13.9 + U. S. N. M. 241315 [M] yg. Choachí, Columbia 45.1 16.7 13.5 + + Mustela frenata + A. M. N. H. 34677 [M] yg. Gualea, Ecuador 45.6 17.6 14.0 + + Mustela frenata + F. M. N. H. 24133 [M] ad. Rio Chinchao 44.0 16.4 14.5 + F. M. N. H. 24132 [M] ad. Rio Chinchao 45.3 17.0 13.9 + + F. M. N. H. 24136 [F] ad. Huanuco, Perú 35.3 13.0 10.5 + F. M. N. H. 24135 [F] sad. Ambo, Perú 36.1 13.6 10.9 + F. M. N. H. 24134 [F] sad. Ambo, Perú 38.1 14.0 10.6 + + Mustela frenata + B. M. 8.1.10.1 [M] ad. Lima, Perú 42.5 16.0 13.4 + + F. M. N. H. 21147 [F] ad. Macate, Perú 35.2 12.7 10.0 + M. P. H. N. 565 [F] ad. Perú *36.0 12.6 10.7 + + Mustela frenata + M. P. H. N. 561 [M] ad. Junín, Perú 45.5 16.6 14.0 + M. P. H. N. 562 [M] ad. Junín, Perú 40.8 15.0 13.6 + B. M. 26.2.1.2 [M] ad. Yana Mayo, Perú 42.6 15.9 13.0 + U. S. N. M. 148528 [M] ad. Marcapata, Perú 16.0 + + M. P. H. N. 564 [F] ad. Cutervo, Perú *38.0 13.8 11.2 + + A. M. N. H. 60508 [M] ad. El Chiral, Ecuador 45.5 17.1 13.8 + A. M. N. H. 61406 [M] sad. Guainche, Ecuador 48.2 17.3 14.5 + N. H. R. S. 2 [M] ad. Panecillo, Ecuador 48.0 16.8 13.3 + N. H. R. S. 5 [M] ad. San Antonio, Ecuador 44.0 16.3 14.0 + N. H. R. S. 7 [M] ad. Carapungo, Ecuador 46.8 17.4 13.5 + N. H. R. S. 14 [M] ad. Nára Papallacta, Ecuador 45.4 17.3 14.1 + + N. H. R. S. 10 [F] ad. Guapulo, Ecuador 37.9 13.5 11.3 + + Mustela frenata + A. M. N. H. 72587 [M] ad. Nequejahuira, Bolivia 41.6 15.3 12.2 + B. Z. M. 602 [M] yg. Limbaní, Perú 42.4 15.5 + U. S. N. M. 137513 [M] yg. Limbaní, Perú 40.3 15.2 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + meridana + 12.0 15.0 23.9 28.5 13.0 6.6 3.0 6.2 5.4 5.6 4.4 2.5 15.0 13.4 + 11.8 15.2 23.5 27.4 13.8 6.5 2.4 6.0 5.5 5.7 4.1 2.3 15.6 13.3 + 11.6 15.4 23.6 28.0 13.9 7.0 6.2 5.6 5.7 4.7 2.4 15.7 12.9 + 11.5 15.4 23.6 27.3 14.2 6.4 2.3 5.9 5.2 5.6 4.5 2.3 15.3 13.0 + 11.5 15.5 23.0 27.6 13.5 6.5 2.6 5.8 5.4 5.6 4.3 2.1 15.5 13.7 + 11.7 15.3 23.5 27.8 13.7 6.6 6.0 5.4 5.6 4.4 2.3 16.0 13.3 + + 12.1 15.1 23.0 27.5 13.2 6.0 5.5 5.7 4.5 2.5 + 11.4 15.5 23.4 27.9 13.4 6.4 2.2 5.6 5.2 5.7 4.2 2.3 16.0 13.0 + 10.5 15.4 25.0 13.2 5.5 5.0 5.3 4.2 2.3 15.0 13.2 + + 9.7 13.2 20.3 23.0 12.4 5.9 1.8 + 9.9 13.0 20.1 23.4 12.1 5.9 2.0 + 9.7 11.8 18.8 22.0 11.7 5.5 2.0 4.9 4.5 4.6 3.5 1.6 10.5 + 9.8 13.8 19.5 12.1 5.9 1.8 4.9 4.7 4.9 3.7 1.8 13.0 11.2 + 9.8 13.0 19.7 22.8 12.1 5.8 1.9 + + 9.2 13.2 19.2 22.5 11.9 5.6 2.1 5.1 4.6 4.8 3.5 1.7 13.5 11.1 + 8.7 12.0 19.3 21.8 11.5 5.8 1.8 4.7 4.4 4.7 3.5 1.6 13.8 11.4 + + affinis + 11.8 15.6 29.5 6.1 5.9 6.1 4.4 2.3 13.9 + 10.8 14.3 27.7 6.0 5.3 5.7 4.5 2.1 13.0 + 11.0 15.3 6.4 5.8 5.5 5.7 4.4 2.0 13.3 + 11.2 15.1 28.6 6.0 5.6 5.8 4.4 2.1 13.4 + + 12.3 15.6 23.9 29.0 5.9 5.7 5.7 4.4 2.5 + 11.5 15.2 23.9 28.7 14.0 6.9 5.9 5.6 5.8 4.5 2.1 13.5 + + aureoventris + 11.8 15.4 23.8 28.3 15.1 6.8 2.7 6.7 6.0 6.4 5.0 3.0 17.0 14.3 + + helleri + 12.5 14.9 24.5 29.1 14.4 6.5 2.4 6.2 5.8 5.9 4.6 2.5 16.0 14.0 + 11.7 15.5 24.5 29.0 14.4 6.5 2.4 5.8 5.5 5.8 4.5 2.5 16.0 13.7 + + 8.9 12.1 17.9 20.8 11.8 4.7 2.0 4.7 4.4 4.6 3.5 1.6 11.8 10.0 + 9.0 12.4 18.8 22.4 11.8 5.0 1.9 4.7 4.4 4.6 3.6 1.7 12.6 10.5 + 9.1 12.6 19.8 22.2 12.0 5.4 2.0 4.7 4.6 4.6 3.5 1.5 13.9 11.0 + + agilis + 11.1 14.9 23.1 28.2 13.8 6.9 2.6 5.8 5.5 5.8 4.5 2.2 14.8 12.9 + + 8.4 12.0 18.0 20.6 12.4 5.0 1.7 4.4 4.1 4.4 3.2 1.5 13.0 10.3 + 9.2 12.5 18.1 21.5 13.0 5.8 2.2 4.8 4.1 4.5 3.5 1.9 12.0 11.7 + + macrura + 11.6 15.1 23.8 29.0 14.5 6.2 5.5 5.4 5.5 4.3 2.1 15.7 13.2 + 11.3 15.1 22.9 26.5 14.5 7.5 2.7 5.8 5.0 5.1 4.0 2.0 14.6 12.9 + 10.2 15.0 21.8 26.2 13.5 7.0 2.5 5.8 5.5 5.6 4.4 2.3 14.6 12.3 + 14.5 5.7 5.0 5.2 4.3 2.1 + + 9.5 13.0 18.1 *23.0 4.9 4.6 4.9 3.6 1.9 11.7 + + 11.5 14.5 24.2 23.2 15.0 7.0 2.3 5.9 5.6 6.0 4.8 2.6 16.4 13.8 + 12.3 15.7 25.6 30.3 15.6 7.2 3.0 6.1 5.2 5.4 4.5 2.3 15.7 13.9 + 14.8 14.5 7.0 6.0 5.4 5.7 4.3 2.2 + 11.9 14.8 23.5 28.0 14.2 7.0 2.9 5.8 5.1 5.4 4.3 2.1 16.8 14.1 + 11.1 14.9 23.7 29.0 14.6 7.0 2.9 6.2 5.8 6.2 4.5 2.3 16.0 14.2 + 12.6 15.8 24.5 29.9 14.3 7.1 2.6 6.1 5.4 5.6 4.3 2.5 17.5 9.5 + + 9.5 12.7 19.7 22.5 12.4 5.9 2.0 5.1 4.5 4.7 3.7 1.7 12.9 11.3 + + boliviensis + 10.0 14.7 22.2 25.0 13.4 7.4 2.2 5.3 5.0 5.2 4.4 2.2 14.8 12.4 + 5.5 5.0 5.4 4.2 2.2 + 5.1 4.8 5.1 3.9 1.9 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela africana + A. M. N. H. 374.75 [M] sad. Pará, Brazil 47.8 17.2 14.3 + B. M. 5.1.25.1. [M]? sad. Pará, Brazil 44.9 16.4 13.0 + B. M. 26.1.8.10. [M]? ad. Pará, Brazil 44.6 16.4 13.3 + + Mustela africana + B. M. 24.12.12.24. [F] ad. Moyobamba, Perú 45.8 16.9 13.5 + M. P. H. N. 563 [F] ad. Yurimaguas, Perú 17.5± + A. M. N. H. 61813 [F] yg. Val. d. Perené, Perú 44.6 16.0 13.0 + + Mustela erminea + M. C. Z. 10012 [M] ad. Pt. Barrow 43.3 15.9 15.2 + F. M. N. H. 35894 [M] ad. Pt. Barrow 41.8 15.6 15.3 + A. N. S. P. 6909 [M] ad. Pt. Barrow 42.3 15.1 14.8 + A. N. S. P. 6910 [M] ad. Pt. Barrow 16.2 16.0 + N. M. C. 2445 [M] ad. Salirochet River 42.8 15.7 15.4 + av. 5 42.5 15.7 15.3 + + F. M. N. H. 35895 [F] ad. Pt. Barrow 35.4 13.1 12.7 + M. V. Z. 43286 [F] ad. Pt. Barrow 35.4 12.9 12.0 + U. S. N. M. 243489 [F] sad. Alatna River 37.0 13.4 11.9 + U. S. N. M. 243493 [F] sad. 16 mi. below Bettles 36.2 13.0 12.0 + U. S. N. M. 180459 [F] ad. N. Fk. Kuskokim 35.3 12.9 11.2 + U. S. N. M. 242205 [F] ad. Fairbanks 34.5 12.4 11.5 + U. S. N. M. 157306 [F] sad. Bear Creek 36.9 13.5 12.6 + U. S. N. M. 157305 [F] sad. Bear Creek 35.2 12.5 11.4 + av. 8 35.8 12.9 11.8 + + Mustela erminea + M. C. Z. 29831 [M] ad. Ymer Is. 41.6 15.6 15.3 + C. Z. M. 1245 [M] ad. Danmarks Havn. 40.7 15.0 14.7 + C. Z. M. 1246 [M] ad. Danmarks Havn. 39.0 14.1 13.8 + C. Z. M. 1247 [M] ad. Danmarks Havn. 41.0 15.2 14.5 + C. Z. M. 1248 [M] ad. Danmarks Havn. 42.3 15.5 15.5 + C. Z. M. 1249 [M] ad. Danmarks Havn. 42.2 15.8 15.0 + C. Z. M. 1871 [M] ad. Scoresby Sd. 42.4 15.3 15.3 + av. 7 41.3 15.2 14.9 + + C. Z. M. 1060 [F] ad. Turner Sd. 35.9 13.0 12.5 + B. Z. M. 43965 [F] ad. Kap Hoegh 37.8 13.4 12.7 + + Mustela erminea + C. M. 6688 [M] ad. Prairie Point *39.9 14.5 13.5 + + average} {37.5 13.8 13.1 + maximum} [M] 1 ad. and Southampton Isl. {39.9 14.5 13.5 + minimum} 10 sad. {35.7 13.1 12.6 + + C. M. 8474 [F] ad. Minnimunnek Pt. 32.6 11.9 11.3 + + average} {34.2 12.4 11.7 + maximum} [F] 1 ad. and Southampton Isl. {35.1 13.0 12.7 + minimum} 4 sad. {32.6 11.9 11.3 + + Mustela erminea + U. S. N. M. 107496 [M] ad. Kadiak 43.2 15.4 14.9 + F. M. N. H. 7290 [M] ad. Kadiak Id. 42.1 14.9 14.0 + + U. S. N. M. 98042 [F] ad. Kadiak 33.0 12.0 10.2 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + africana + 12.9 15.4 26.4 32.2 16.9 8.2 2.8 6.6 5.8 6.2 4.7 2.4 17.8 14.9 + 10.2 14.2 23.3 26.8 14.4 7.0 2.7 5.9 5.4 5.5 4.5 1.8 16.4 13.7 + 10.9 14.4 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 + 12.0 14.6 23.5 28.8 14.5 6.2 2.6 5.9 5.4 4.9 4.3 1.8 16.5 13.9 + 6.0 5.8 5.6 4.4 2.3 + 11.0 15.4 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 + 12.0 16.2 23.5 27.6 15.5 8.4 5.6 5.1 5.1 3.9 2.0 13.3 13.2 + 13.3 16.0 23.2 27.6 15.1 8.5 6.0 5.2 5.3 4.0 2.2 14.1 13.5 + 11.6 15.3 22.9 26.2 15.2 8.4 5.9 5.1 5.3 4.1 2.4 14.5 12.5 + 12.7 16.7 5.7 5.4 5.7 4.3 2.1 + 13.0 16.0 23.5 28.5 15.2 8.0 5.6 5.0 5.2 4.0 2.0 14.3 13.3 + 12.5 16.0 23.3 27.5 15.3 8.3 5.8 5.2 5.3 4.1 2.1 14.1 13.1 + + 10.3 13.3 19.3 23.0 12.7 7.2 5.0 4.5 4.8 3.5 1.7 12.0 10.5 + 10.0 13.2 18.8 21.8 12.6 6.7 4.6 4.2 4.3 3.2 1.8 11.4 10.3 + 9.4 13.5 19.4 22.0 13.2 7.2 4.7 4.3 4.5 3.3 1.7 11.9 10.6 + 9.4 13.0 19.1 21.8 12.9 7.2 5.0 4.2 4.5 3.3 1.8 11.9 10.3 + 8.8 12.8 18.9 20.7 12.7 7.0 4.7 4.3 4.5 3.3 1.8 10.5 9.7 + 9.3 13.0 18.0 12.7 6.8 4.8 4.3 4.5 3.3 1.7 11.8 10.2 + 9.5 13.6 18.7 20.9 13.0 6.9 4.8 4.4 4.5 3.3 1.8 11.4 10.2 + 9.6 12.8 17.8 20.5 12.6 6.3 4.8 4.2 4.4 3.3 1.6 11.2 10.1 + 9.3 13.1 18.7 21.2 12.9 6.9 4.8 4.3 4.5 3.3 1.7 11.5 10.2 + + polaris + 12.0 13.0 22.3 26.1 14.8 8.0 5.8 5.2 5.7 4.1 2.2 13.7 12.3 + 11.9 14.9 22.3 25.5 14.6 7.8 5.9 5.1 5.6 3.9 2.0 13.4 12.2 + 10.9 14.2 21.3 24.7 14.5 7.3 5.3 4.8 5.0 3.7 1.9 12.0 11.2 + 11.5 14.4 22.3 26.7 14.4 7.6 5.7 5.1 5.5 4.1 2.0 13.1 12.1 + 12.0 15.3 23.2 27.9 15.1 8.1 5.8 5.5 5.8 4.0 2.3 13.8 12.8 + 12.2 15.1 23.2 28.0 15.2 8.1 5.9 5.2 5.7 4.2 2.3 13.7 12.4 + 12.3 15.6 15.4 8.3 5.7 5.1 5.3 3.9 2.0 14.4 12.8 + 11.8 14.9 22.4 26.5 14.9 7.9 5.7 5.2 5.5 4.0 2.1 13.4 12.3 + + 10.1 12.9 19.6 22.6 13.6 7.1 5.0 4.5 4.7 3.6 1.7 12.3 10.3 + 10.4 13.2 19.3 22.4 13.9 7.0 4.8 4.2 4.5 3.7 1.8 12.1 10.9 + + semplei + 11.5 14.6 21.5 24.8 14.1 7.9 5.0 4.6 4.7 3.6 2.0 14.0 11.6 + + 10.6 13.8 20.2 23.6 13.6 7.2 5.1 4.7 4.9 3.6 2.0 13.4 11.0 + 11.5 14.8 21.5 24.8 15.0 7.9 5.3 4.9 5.0 3.7 2.3 14.0 11.6 + 9.7 12.7 19.2 22.2 12.9 6.4 4.8 4.4 4.5 3.4 1.8 12.3 10.5 + + 9.2 12.2 18.5 21.2 12.2 6.6 4.4 3.9 4.2 3.0 1.5 11.9 9.8 + + 9.5 12.6 18.8 20.9 12.8 6.7 4.5 4.1 4.3 3.1 1.6 12.1 10.1 + 9.9 13.1 19.4 21.5 13.4 7.0 4.3 4.3 4.5 3.3 1.7 12.4 10.4 + 9.2 12.2 18.1 20.1 12.2 6.6 4.6 3.9 4.2 3.0 1.5 11.9 9.6 + + kadiacensis + 11.1 14.9 22.1 26.0 15.1 7.6 5.5 5.1 5.1 3.8 2.0 14.0 12.2 + 10.9 14.4 21.6 26.0 14.1 7.1 5.7 5.2 5.2 3.8 1.7 13.5 11.5 + + 8.0 11.3 16.9 19.4 11.5 6.0 4.4 4.0 4.0 2.8 1.4 11.5 8.9 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela erminea + average} {40.9 14.3 12.5 + maximum} [M] ad. 6 3 mi. S Big Isl. {43.7 15.0 13.2 + minimum} {39.6 13.8 11.7 + + average} {40.2 14.1 12.2 + maximum} [M] sad. 7 3 mi. S Big Isl. {41.5 14.7 12.6 + minimum} {38.4 13.6 12.0 + + U. S. N. M. 136112 [F] ad. Willow River 33.0 11.5 8.8 + M. C. Z. 242866 [F] sad. 3 mi. S Big Isl. 32.3 11.0 9.0 + U. S. N. M. 129703 [F] ad. Ft. Resolution 34.2 12.0 10.4 + U. S. N. M. 110682 [F] sad. 15 mi. above Smith Landing 33.8 11.3 9.8 + U. S. N. M. 235959 [F] ad. Athabasca Delta 33.1 11.7 9.3 + M. C. Z. 18776 [F] ad. Athabasca Delta 31.5 10.8 8.9 + av. 6 33.0 11.4 9.4 + + Mustela erminea + M. V. Z. 53789 [M] ad. Ogdensburg, N. Y. 36.6 12.9 11.5 + U. S. N. M. 32240/44066 [M] sad. Amsterdam, N. Y. 36.7 12.9 11.5 + + A. M. N. H. 67869 [M] ad. Berlin, N. Y. 36.2 12.5 10.2 + A. M. N. H. 67868 [M] ad. Berlin, N. Y. 34.8 12.1 9.8 + A. M. N. H. 15841 [M] ad. Schoharie, N. Y. 33.8 11.7 9.7 + Cornell 494 [M] sad. Cascadilla Creek, N. Y. 34.8 12.2 10.4 + C. M. 7461 [M] sad. Pymatuning Swamp 34.2 12.0 9.6 + C. M. 10264 [M] sad. 3-1/2 mi. W Linesville 37.6 13.2 11.0 + M. V. Z. 53788 [M] sad. Lopez, Penn. 36.5 12.6 10.7 + av. 9 35.7 12.5 10.5 + + U. S. N. M. 135570 [F] ad. Lake George 33.3 11.8 9.3 + B. S. N. 994 [F] sad. Cattaraugus 31.4 10.8 9.2 + C. M. 7460 [F] sad. Pymatuning Swamp 32.0 10.9 9.2 + C. M. 10252 [F] sad. 3 mi. NW Linesville 32.7 11.2 9.5 + av. 4 32.4 11.2 9.4 + + Mustela erminea + F. M. N. H. 18134 [M] ad. Aitkin, Minn. 40.3 14.1 12.7 + F. M. N. H. 18135 [M] ad. Aitkin, Minn. 14.1 12.5 + F. M. N. H. 18130 [M] ad. Aitkin, Minn. 40.7 15.0 12.2 + F. M. N. H. 18133 [M] ad. Aitkin, Minn. 39.3 13.9 11.2 + F. M. N. H. 18131 [M] ad. Aitkin, Minn. 38.5 13.4 11.2 + F. M. N. H. 7222 [M] ad. Aitkin, Minn. 13.6 12.4 + F. M. N. H. 18127 [M] sad. Aitkin, Minn. 38.4 13.1 11.1 + F. M. N. H. 18129 [M] sad. Aitkin, Minn. 39.4 13.7 11.1 + F. M. N. H. 18132 [M] sad. Aitkin, Minn. 39.3 14.3 11.5 + F. M. N. H. 18441 [M] sad. Aitkin, Minn. 40.2 14.0 12.0 + F. M. N. H. 18440 [M] sad. Aitkin, Minn. 38.8 13.8 11.3 + F. M. N. H. 7219 [M] sad. Aitkin, Minn. 13.9 11.9 + av. 12 39.4 13.9 11.8 + + average} {37.9 13.2 11.4 + maximum} [M] ad. 5 and Elk River {39.5 13.9 12.6 + minimum} sad. 5 {34.8 12.0 10.5 + + Walker 377 [F] ad. Deer 31.8 11.0 9.0 + Walker 11 [F] ad. Grand Maris 32.7 10.8 + Wisc. U. 8681 [F] ad. T. 61N, R. 26W 32.9 10.8 9.2 + Wisc. U. 8679 [F] ad. T. 61N, R. 26W 33.6 11.6 9.8 + Walker A 58 [F] ad. Elk River 32.8 11.5 9.9 + av. 5 32.8 11.1 9.5 + + Wisc. U. 8691 [F] ad. Fisher Lake 31.5 10.6 9.5 + Wisc. U. 8674 [F] ad. Gordon 32.8 11.5 9.8 + Snyder 2637 [F] ad. Beaver Dam 32.9 11.0 9.8 + Snyder 993 [F] ad. Beaver Dam 34.1 11.3 9.4 + Snyder 2999 [F] ad. Beaver Dam 31.9 10.8 9.2 + av. 5 32.6 11.0 9.5 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + richardsonii + 10.2 13.5 21.2 24.2 14.9 7.9 5.3 4.7 4.8 3.8 2.1 13.0 11.9 + 11.1 14.0 22.2 25.5 15.5 8.3 6.0 5.0 5.2 4.2 2.3 14.2 11.4 + 9.4 12.9 20.0 22.9 14.2 7.2 4.9 4.5 4.6 3.7 2.0 12.5 12.3 + + 9.8 14.0 20.6 23.3 14.6 7.7 5.2 4.7 4.9 3.7 2.1 13.6 11.8 + 10.4 14.2 21.5 24.4 15.1 8.2 5.5 4.9 5.0 3.8 2.5 14.5 12.2 + 9.5 13.7 20.0 22.0 13.7 7.0 5.0 4.5 4.7 3.5 2.0 12.8 11.4 + + 7.6 11.4 15.9 18.0 12.5 5.7 4.0 3.8 3.9 2.7 1.5 11.6 8.9 + 7.2 11.7 16.6 17.7 12.3 6.5 4.0 3.7 3.9 3.0 1.5 10.0 8.8 + 8.2 11.5 17.2 19.7 12.3 6.5 4.2 4.0 4.2 3.2 1.7 12.0 9.6 + 7.4 11.0 16.5 18.3 12.1 6.0 4.2 3.8 3.9 2.9 1.5 11.2 9.3 + 7.3 11.0 15.7 18.1 11.6 5.6 4.0 3.8 3.8 3.0 1.6 10.5 8.6 + 7.6 11.2 16.3 18.3 11.6 6.3 3.8 3.7 3.8 3.1 1.6 11.0 8.7 + 7.6 11.3 16.4 18.3 12.1 6.3 4.0 3.8 3.9 3.0 1.6 11.1 9.0 + + cicognanii + 9.1 13.2 18.5 21.7 13.3 7.0 4.7 4.4 4.5 3.4 1.8 12.3 10.4 + 9.4 13.3 19.1 21.5 13.7 6.9 4.6 4.4 4.5 3.4 2.0 13.2 10.8 + + 8.9 12.7 18.8 20.6 12.9 6.7 4.3 3.8 4.0 3.3 1.9 11.5 10.0 + 8.2 12.7 17.4 19.8 12.3 6.3 4.5 3.9 4.1 2.9 1.7 11.5 9.7 + 7.7 11.9 17.3 20.0 11.8 6.4 4.2 3.8 4.0 2.9 1.5 11.8 9.5 + 8.3 12.5 17.9 19.3 12.4 6.6 4.4 4.1 4.2 3.0 1.6 12.2 10.3 + 8.1 12.8 17.3 19.0 11.9 6.4 4.3 3.9 4.2 3.0 1.9 11.5 9.2 + 8.9 12.7 18.8 20.6 12.9 7.2 4.9 4.3 4.5 3.5 2.0 12.6 10.6 + 8.6 12.6 18.6 20.3 13.1 6.8 4.5 4.1 4.3 3.2 1.6 10.9 9.9 + 8.6 12.7 18.2 20.3 12.7 6.7 4.5 4.1 4.3 3.2 1.8 11.9 10.0 + + 7.8 11.4 11.8 5.8 4.4 3.8 4.0 3.0 1.8 9.1 + 7.2 10.0 15.3 10.7 5.7 3.8 3.7 3.6 2.7 1.5 10.3 8.2 + 7.5 11.0 15.9 17.5 11.1 5.9 3.7 3.4 3.6 2.8 1.5 10.3 8.2 + 7.6 11.3 16.0 18.0 11.8 6.2 3.9 3.6 3.7 3.1 1.7 10.9 8.8 + 7.5 10.9 15.7 17.8 11.4 5.9 3.9 3.6 3.7 2.9 1.6 10.5 8.6 + + bangsi + 10.6 14.6 20.6 23.5 14.3 7.7 4.9 4.6 4.9 3.6 1.9 13.3 11.8 + 10.4 14.4 24.3 5.3 4.5 4.6 3.7 2.1 12.2 + 9.7 14.6 20.2 22.8 14.6 7.5 5.2 5.0 5.1 3.8 2.6 13.5 12.0 + 9.0 13.0 19.5 22.0 14.3 7.2 5.1 4.5 4.6 3.5 2.0 13.0 11.2 + 9.0 12.6 18.8 21.0 13.7 7.3 4.8 4.4 4.5 3.4 1.8 12.0 10.5 + 9.5 13.3 14.4 7.9 4.8 4.4 4.4 3.5 1.8 11.2 + 9.0 13.0 19.5 22.0 14.3 6.9 4.6 4.5 4.5 3.5 1.9 13.4 11.0 + 9.1 13.3 19.0 21.6 13.6 7.2 4.7 4.3 4.5 3.4 2.0 12.8 11.1 + 9.5 13.2 20.2 22.8 14.3 7.2 4.9 4.7 4.7 3.5 2.1 12.5 11.2 + 9.9 14.6 19.6 22.6 15.0 7.1 5.2 4.7 4.8 3.7 2.0 13.3 11.5 + 9.3 13.2 19.7 22.0 13.8 7.6 5.1 4.5 4.7 3.7 2.0 13.0 11.1 + 10.0 13.5 24.7 14.6 7.3 5.3 4.6 4.8 4.0 2.1 11.8 + 9.6 13.6 19.7 22.7 14.3 7.4 5.0 4.6 4.7 3.6 2.0 13.0 11.4 + + 9.1 13.1 19.3 21.6 14.4 7.2 4.7 4.3 4.4 3.4 1.9 12.7 10.8 + 9.8 14.2 20.5 22.8 15.3 8.0 5.1 4.4 4.6 3.6 2.0 14.0 11.7 + 8.5 12.2 18.0 20.8 12.4 6.3 4.3 4.1 4.1 3.1 1.7 12.0 10.0 + + 7.5 10.9 16.6 18.4 11.7 5.9 4.2 3.7 3.8 2.8 1.6 11.2 9.3 + 11.0 15.9 17.3 12.2 5.6 3.9 3.8 3.8 2.9 1.6 10.7 8.5 + 7.4 11.3 15.8 16.8 12.3 6.2 3.8 3.6 3.6 2.8 1.4 10.0 8.3 + 7.6 12.2 15.7 17.1 11.9 5.7 4.0 3.6 3.7 3.0 1.6 9.6 8.9 + 8.1 11.3 16.8 18.3 11.8 6.4 4.0 3.8 4.0 2.9 1.7 10.5 9.2 + 7.7 11.3 16.2 17.6 12.0 6.0 4.0 3.7 3.9 2.9 1.6 10.4 8.8 + + 7.7 10.8 15.8 17.4 11.2 5.6 3.8 3.5 3.7 2.6 1.4 10.4 8.5 + 7.2 11.4 15.7 16.7 11.6 5.7 4.2 3.8 3.8 2.8 1.6 9.6 8.9 + 7.8 11.7 16.9 18.9 12.2 6.4 4.1 3.8 3.8 3.2 1.5 11.3 9.7 + 7.5 11.8 16.7 18.0 12.8 6.1 4.4 3.8 3.8 3.0 1.7 10.8 9.0 + 7.4 11.2 15.9 17.5 12.4 5.8 4.0 3.6 3.8 3.0 1.6 11.4 9.2 + 7.5 11.4 16.2 17.7 12.0 5.9 4.1 3.7 3.8 2.9 1.6 10.7 9.1 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela erminea + average} {37.0 12.8 11.4 + maximum} [M] ad. 5 Idaho Co {39.8 14.1 13.0 + minimum} {35.8 12.2 10.6 + + M. V. Z. 90763 [F] ad. Pilot Creek, Idaho 31.6 10.8 9.2 + + average} {32.2 10.6 9.0 + maximum} [F] ad. 1 and Idaho Co {32.8 11.2 9.2 + minimum} sad. 4 {31.6 10.8 8.5 + + Mustela erminea + average} {37.5 13.1 11.5 + maximum} [M] ad. 8 Windham {38.9 13.7 12.3 + minimum} {36.5 12.3 11.0 + + U. S. N. M. 74422 [F] ad. Juneau 33.2 11.4 10.5 + M. V. Z. 995 [F] ad. Juneau 33.1 11.3 9.4 + M. V. Z. 473 [F] ad. Helm Bay 32.9 11.2 9.5 + U. S. N. M. 74773 [F] ad. Wrangel 32.2 11.3 9.2 + M. V. Z. 78243 [F] sad. Windham 31.9 11.1 10.1 + av. 5 32.7 11.3 9.7 + + Mustela erminea + average} {37.8 13.0 11.9 + maximum} [M] ad. to sad. 12 Mole Harbor {39.5 13.7 13.0 + minimum} {36.4 12.5 10.7 + + average} {33.0 11.3 9.9 + maximum} [F] ad. 2 and Mole Harbor {33.6 11.9 10.2 + minimum} sad. 4 {32.0 10.9 9.5 + + Mustela erminea + M. V. Z. 289 [M] ad. Saook Bay 40.5 13.9 12.8 + M. V. Z. 286 [M] ad. Saook Bay 39.6 13.5 13.1 + + Mustela erminea + average} {39.5 14.0 13.6 + maximum} [M] ad. 5 Prince of Wales Id {40.7 14.4 14.5 + minimum} {38.9 13.9 13.1 + + average} {38.7 13.6 13.2 + maximum} [M] ad. 5 and Prince of Wales Id {40.9 14.4 14.5 + minimum} sad. 15 {36.7 13.0 11.8 + + M. V. Z. 31223 [F] sad. Prince of Wales Id 12.2 11.5 + + Mustela erminea + M. V. Z. 31232 [M] ad. Suemez Id 34.3 12.6 12.6 + + Mustela erminea + U. S. N. M. 94430 [M] ad. Massett 36.7 13.4 12.7 + + average} {36.7 13.4 12.7 + maximum} [M] ad. 7 Graham Id {37.5 13.6 12.9 + minimum} {35.6 13.0 12.2 + + M. V. Z. 31209 [F] ad. Massett 34.2 12.5 11.3 + U. S. N. M. 100624 [F] ad. Cumsheva Inlet 34.2 12.3 11.5 + + Mustela erminea + average} {34.0 11.7 10.8 + maximum} [M] 13 ad. Vancouver Id {35.6 12.2 11.3 + minimum} {32.5 11.0 10.1 + + average} {31.5 10.9 9.8 + maximum} [F] 5 ad. Vancouver Id {31.8 11.1 10.0 + minimum} {30.9 10.5 9.6 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + invicta + 9.1 13.1 19.0 21.3 13.6 7.0 4.5 4.1 4.2 3.2 1.7 11.7 10.5 + 10.0 14.2 19.7 22.6 14.2 7.1 4.9 4.3 4.5 3.5 1.9 12.3 11.3 + 8.6 12.0 18.2 20.5 13.3 6.8 4.2 4.0 4.0 3.1 1.4 10.9 9.7 + + 7.0 10.5 15.6 16.5 10.9 5.6 4.0 3.6 3.7 2.9 1.4 9.0 8.2 + + 7.1 11.1 16.3 17.2 12.0 5.9 3.9 3.7 3.7 2.8 1.4 9.5 8.3 + 7.2 12.2 17.0 17.8 12.7 6.5 4.3 3.9 3.9 2.9 1.5 10.0 8.7 + 7.0 10.5 15.6 16.5 10.9 5.6 3.5 3.6 3.6 2.7 1.3 9.0 8.0 + + alascensis + 9.4 13.2 19.4 21.9 13.2 6.9 4.8 4.2 4.4 3.5 1.9 12.5 10.7 + 10.1 14.3 20.5 23.7 13.7 7.4 5.0 4.4 4.7 3.9 2.2 13.8 11.6 + 8.6 12.0 18.5 20.4 12.9 6.6 4.5 4.1 4.2 3.3 1.8 11.8 10.4 + + 8.3 10.5 16.2 18.3 11.5 5.7 3.9 3.5 3.7 2.9 1.6 10.0 8.7 + 7.8 11.3 16.3 17.8 12.0 5.8 3.9 3.7 3.8 2.9 1.7 8.7 + 8.5 11.6 16.3 17.6 11.9 6.0 4.0 3.7 3.9 2.6 1.4 10.5 9.3 + 7.5 16.0 17.6 11.9 5.8 4.0 3.8 4.0 3.2 1.7 11.5 8.7 + 7.7 11.5 16.8 18.0 11.5 5.7 4.0 3.6 3.8 3.1 1.7 10.6 8.8 + 8.0 11.2 16.3 17.9 11.8 5.8 4.0 3.7 3.8 2.9 1.6 10.6 8.8 + + salva + 9.6 13.3 19.2 22.0 12.8 6.8 4.6 4.3 4.4 3.5 1.8 11.7 10.7 + 10.8 14.2 20.0 23.2 13.8 7.2 5.0 4.6 4.8 3.9 2.0 13.1 11.4 + 8.4 12.4 18.0 20.4 12.0 6.2 4.4 4.0 4.0 3.1 1.7 11.2 10.0 + + 8.1 11.6 16.5 18.2 11.5 5.8 4.0 3.8 3.8 3.0 1.5 10.1 9.1 + 8.7 12.3 17.1 18.7 12.0 6.2 4.2 3.9 3.9 3.2 1.6 10.9 9.4 + 7.5 11.1 15.4 17.1 11.0 5.3 3.8 3.6 3.6 2.9 1.5 8.9 8.4 + + initis + 10.6 14.4 22.1 24.5 14.8 7.6 5.2 4.7 5.0 4.1 1.9 12.8 11.0 + 11.4 15.0 21.0 24.3 13.6 7.6 5.0 4.7 4.9 3.6 2.1 12.3 12.0 + + celenda + 11.5 14.7 20.9 24.2 13.6 7.5 5.1 4.7 4.8 3.7 1.9 12.9 11.6 + 12.1 15.6 21.7 25.8 14.2 7.9 5.1 4.9 4.9 3.9 2.2 13.6 12.5 + 10.9 13.8 19.9 23.2 13.2 7.0 5.0 4.6 4.6 3.6 1.7 12.3 10.8 + + 11.2 14.4 20.3 23.3 13.2 7.3 5.0 4.6 4.7 3.6 1.8 12.9 11.4 + 12.1 15.6 21.7 25.8 14.2 7.9 5.3 5.0 4.9 3.9 2.2 13.6 12.5 + 10.2 13.4 19.0 21.3 12.3 6.8 4.6 4.3 4.2 3.3 1.6 12.1 10.8 + + 9.8 12.6 4.5 4.2 4.2 3.2 1.5 10.0 + + seclusa + 10.6 13.9 20.2 22.7 12.7 6.9 5.1 4.7 5.0 3.8 1.8 12.3 11.5 + + haidarum + 10.5 13.9 19.3 22.6 12.4 6.4 5.0 4.3 4.5 3.3 1.9 12.4 11.4 + + 10.9 14.3 18.9 21.8 12.6 6.8 5.0 4.3 4.6 3.4 1.7 12.6 11.3 + 11.2 14.8 19.6 22.4 13.0 7.1 4.8 4.4 4.7 3.4 1.9 13.1 11.7 + 10.5 14.0 18.0 21.1 12.3 6.4 5.1 4.2 4.4 3.3 1.6 12.1 10.8 + + 9.8 13.3 17.3 19.8 11.5 6.1 4.7 4.1 4.2 3.0 1.5 11.5 10.2 + 9.8 13.1 17.0 19.8 11.8 6.1 4.6 4.0 4.3 3.2 1.7 11.2 10.0 + + anguinae + 9.0 12.0 17.1 19.3 11.9 6.1 4.3 3.8 4.0 3.1 1.7 11.6 10.0 + 9.6 12.5 17.9 20.6 12.5 6.7 4.6 4.0 4.1 3.3 1.9 12.5 10.5 + 8.5 11.3 16.5 18.8 11.2 5.7 4.0 3.6 3.7 2.9 1.6 10.7 9.8 + + 8.2 11.5 15.8 17.5 10.8 5.5 4.0 3.6 3.8 2.9 1.6 10.5 9.3 + 8.8 12.4 16.1 17.8 11.1 5.7 4.2 3.8 4.0 3.0 1.6 11.5 10.0 + 7.9 10.6 15.6 17.3 10.4 5.4 3.8 3.5 3.6 2.8 1.5 10.0 8.9 + + + TABLE OF CRANIAL MEASUREMENTS--_Continued_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela erminea + average} {35.7 12.6 11.1 + maximum} [M] ad. 7 Topotypes {38.2 13.1 11.6 + minimum} {34.3 12.0 10.5 + + N. M. C. 7284 [F] ad. Topotype 29.4 10.1 9.1 + N. M. C. 7516 [F] ad. Topotype 31.1 10.1 9.6 + M. C. Z. 6852 [F] ad. Sumas 31.3 10.3 9.2 + M. C. Z. 3645 [F] ad. Sumas 29.4 10.2 8.6 + M. C. Z. 10728 [F] ad. Sumas 31.7 10.2 9.1 + + Mustela erminea + U. S. N. M. 90738 [M] ad. Type 31.9 11.6 10.0 + U. S. N. M. 241941 [M] ad. N. Fk. Quinault River 32.5 11.7 10.2 + U. S. N. M. 231829 [M] ad. Duckabush 30.6 10.9 9.2 + U. S. N. M. 231830 [M] ad. Duckabush 32.1 11.1 10.0 + M. Z. 53700 [M] ad. Lake Cushman 32.0 11.4 9.8 + av. 5 31.8 11.3 9.8 + + C. R. C. M. 96 [F] ad. Elwha River 27.5 9.4 8.3 + C. R. C. M. 1164 [F] ad. 12 mi. S Port Angeles 26.7 9.0 8.1 + U. S. N. M. 242133 [F] ad. Hayes Creek 27.2 9.2 8.4 + av. 3 27.1 9.2 8.3 + + Mustela erminea + average} {33.2 11.7 10.5 + maximum} [M] ad. 12 Tillamook Co {33.8 12.1 11.1 + minimum} {32.5 11.3 10.0 + + average} {28.5 9.9 8.9 + maximum} [F] ad. 7 Tillamook Co {29.5 10.2 9.2 + minimum} {27.6 9.6 8.7 + + Mustela erminea + U. S. N. M. 82177 [M] ad. Trout Lake 32.0 11.3 10.1 + U. S. N. M. 64768 [M] ad. Trout Lake 33.3 12.0 9.9 + + average} {32.3 11.5 10.0 + maximum} [M] ad. 2, sad. 13 Trout Lake {33.4 12.0 10.7 + minimum} {30.9 10.8 9.0 + + U. S. N. M. 232741 [F] ad. Reflection Lakes 28.4 9.6 8.7 + U. S. N. M. 90727 [F] ad. Mt. St. Helens 28.0 9.7 8.1 + U. S. N. M. 81919 [F] ad. Trout Lake 28.1 9.7 8.8 + U. S. N. M. 87039 [F] ad. Trout Lake 28.4 9.8 8.7 + U. S. N. M. 77370 [F] ad. Trout Lake 27.8 9.6 8.6 + av. 5 28.1 9.7 8.6 + + Mustela erminea + U. S. N. M. 231397 [M] ad. Donovan, Mont *31.2 10.5 9.5 + U. S. N. M. 206991 [M] ad. Mill Creek, Oreg *30.9 10.8 9.0 + M. V. Z. 34746 [M] ad. Black Butte, Calif 30.8 11.1 9.4 + M. V. Z. 41501 [M] ad. Wheeler Peak, Nev 29.8 10.4 9.3 + E. R. W. 3050 [M] ad. Crested Butte, Colo. *30.4 11.1 9.8 + av. 5 30.6 10.8 9.4 + + M. Z. 62111 [F] ad. Teton Co., Wyoming 28.0 9.7 8.3 + M. Z. 62112 [F] ad. Teton Co., Wyoming 27.3 9.7 8.1 + M. V. Z. 13776 [F] ad. Rush Creek, California 28.1 9.5 8.8 + M. V. Z. 13777 [F] ad. Castle Lake, California 29.4 9.9 8.8 + M. V. Z. 41502 [F] ad. Wheeler Peak, Nev 27.3 9.3 8.0 + F. M. N. H. 11440 [F] ad. Camp Albion, Colo 27.8 9.4 8.4 + av. 6 28.0 9.6 8.4 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + fallenda + 9.2 12.5 18.3 20.8 13.1 6.8 **4.7 **4.2 **4.4 **3.4 **1.9 11.7 10.5 + 9.9 13.0 19.6 22.8 14.1 7.6 5.1 4.6 4.7 3.6 2.1 12.4 11.0 + 8.3 12.0 17.0 19.4 12.2 6.2 4.3 3.9 3.9 3.2 1.7 11.0 10.0 + + 7.1 10.5 15.4 17.4 11.1 5.5 3.7 3.5 3.6 2.8 1.5 10.5 8.6 + 8.0 11.0 16.0 18.5 11.5 5.6 4.1 3.7 3.8 2.9 1.5 10.0 9.0 + 7.3 10.1 15.8 17.2 11.0 5.3 3.8 3.5 3.7 2.7 1.5 11.0 8.7 + 7.3 10.7 14.7 15.4 10.5 5.2 3.8 3.5 3.8 2.7 1.5 9.5 8.3 + 7.1 10.7 15.7 17.4 11.5 5.5 3.8 3.4 3.7 2.6 1.2 9.3 8.2 + + olympica + 7.9 11.9 15.3 17.9 11.6 5.3 4.0 3.6 3.9 2.9 2.1 9.9 8.9 + 8.2 12.0 16.3 18.2 11.7 5.6 4.2 3.6 3.7 2.9 1.8 10.0 9.2 + 7.4 10.3 15.0 17.2 10.7 5.2 4.2 3.5 3.7 2.7 1.7 10.0 8.6 + 8.2 12.3 16.6 18.6 11.2 5.9 4.0 3.7 3.9 3.0 1.7 10.2 9.3 + 8.0 11.0 16.9 18.8 11.4 6.3 4.3 3.8 3.9 3.1 1.7 10.3 9.2 + 7.9 11.5 16.0 18.1 11.3 5.7 4.2 3.7 3.9 2.9 1.8 10.1 9.0 + + 6.9 9.3 13.2 15.3 10.2 4.6 3.4 3.0 3.3 2.5 1.3 8.5 7.2 + 6.7 9.0 13.1 14.4 9.7 4.8 3.4 3.1 3.4 2.3 1.3 9.0 7.3 + 7.2 9.2 13.7 15.4 9.5 4.8 3.3 3.0 3.1 2.1 1.2 8.3 7.2 + 6.9 9.2 13.3 15.0 9.8 4.7 3.4 3.1 3.3 2.3 1.2 8.6 7.2 + + streatori + 8.5 11.7 17.0 19.2 11.8 6.4 4.3 3.8 3.9 3.1 1.8 10.8 9.8 + 9.1 12.5 18.0 19.8 12.6 6.9 4.4 4.0 4.1 3.5 2.1 11.4 10.3 + 8.2 11.1 16.1 18.5 11.1 6.0 4.1 3.6 3.8 2.9 1.6 10.3 9.5 + + 7.3 10.1 14.3 15.9 10.6 5.2 3.6 3.2 3.5 2.6 1.6 9.4 8.1 + 7.6 10.2 14.8 16.3 11.2 5.4 3.7 3.3 3.6 2.7 1.7 10.0 8.4 + 7.0 9.8 14.1 15.5 10.0 5.0 3.5 3.1 3.3 2.5 1.5 9.2 7.9 + + gulosa + 8.5 11.2 16.3 18.3 11.3 5.9 4.2 3.8 4.1 2.9 1.6 10.5 9.4 + 8.3 12.2 16.9 18.5 11.8 5.9 4.2 3.7 3.7 2.8 1.8 10.6 9.4 + + 8.3 11.5 16.4 18.4 11.5 5.9 4.2 3.7 3.9 2.9 1.7 10.3 9.2 + 8.8 12.4 17.2 19.3 12.1 6.3 4.5 3.8 4.1 3.2 2.0 11.2 9.7 + 7.4 10.8 15.6 17.8 10.8 5.5 3.9 3.5 3.6 2.7 1.6 9.6 8.5 + + 7.4 9.3 14.1 15.8 10.9 5.4 3.7 3.2 3.5 2.7 1.6 8.4 7.4 + 6.9 9.5 13.6 15.6 10.0 4.8 3.6 3.3 3.5 2.5 1.6 9.0 7.6 + 7.0 10.0 13.8 15.4 10.1 4.9 3.6 3.2 3.3 2.3 1.4 8.1 7.3 + 7.1 *10.8 14.5 15.6 10.0 5.0 3.5 3.2 3.4 2.5 1.4 8.6 7.5 + 6.6 *9.3 13.6 15.2 9.7 4.7 3.8 3.4 3.5 2.7 1.5 8.1 7.2 + 7.0 9.8 13.9 15.5 10.1 5.0 3.6 3.3 3.4 2.5 1.5 8.4 7.4 + + muricus + 8.0 11.3 *16.0 17.5 5.7 4.0 3.6 3.7 2.8 1.6 11.2 8.8 + 7.2 11.0 17.4 11.3 5.6 4.3 3.7 4.0 3.0 1.6 8.7 + 7.7 11.1 15.9 17.5 10.8 5.6 4.2 3.5 3.6 2.8 1.8 9.6 8.4 + 7.2 10.6 15.1 17.1 10.7 5.4 4.0 3.5 3.5 2.7 1.5 9.8 8.0 + 7.7 11.2 16.2 18.3 10.8 5.7 4.3 3.6 3.9 2.9 1.6 8.5 + 7.6 11.0 15.8 17.6 10.9 5.6 4.2 3.6 3.7 2.8 1.6 10.2 8.5 + + 6.7 10.0 14.9 16.0 10.9 5.3 3.6 3.2 3.3 2.7 1.7 8.8 7.2 + 6.5 9.2 14.0 15.6 10.1 5.0 3.5 3.2 3.3 2.5 1.4 8.3 7.2 + 7.4 9.7 14.3 16.5 10.2 4.7 3.6 3.2 3.4 2.7 1.5 9.1 7.8 + 7.3 10.8 15.2 17.1 10.0 5.3 3.6 3.1 3.4 2.6 1.5 8.2 + 6.3 9.5 13.9 15.8 10.0 5.0 3.4 2.9 3.1 2.4 1.3 8.8 7.2 + 6.9 9.5 14.1 15.5 10.5 5.1 3.6 3.2 3.3 2.6 1.4 8.9 7.5 + 6.9 9.8 14.4 16.1 10.3 5.1 3.6 3.1 3.3 2.6 1.5 8.8 7.5 + + + TABLE OF CRANIAL MEASUREMENTS--_Concluded_ + ================================================================================= + + Key: + + A Basilar length (of Hensel) + B Length of tooth rows + C Breadth of rostrum + + Catalog Sex and + Collection Number age Locality A B C + --------------------------------------------------------------------------------- + + Mustela erminea + A. M. N. H. 12432 [F] ad. Conard Fissure, Ark 35.1 12.7 10.4 + A. M. N. H. 12433 [F] ad. Conard Fissure, Ark 12.6 9.9 + A. M. N. H. 12435 [F] ad. Conard Fissure, Ark *32.5 12.4 10.0 + A. M. N. H. 11766 [F] ad. Conard Fissure, Ark 34.5 12.3 10.7 + av 34.0 12.5 10.3 + + A. M. N. H. 12437 [M] sad. Conard Fissure, Ark 39.2 14.5 11.7 + A. M. N. H. 12441 [M] ad. Conard Fissure, Ark 38.5 13.9 11.3 + A. M. N. H. 12436 [M] ad. Conard Fissure, Ark 13.5 11.7 + A. M. N. H. 12444 [M] ad. Conard Fissure, Ark 14.3 11.6 + A. M. N. H. 11769 [M] sad. Conard Fissure, Ark + A. M. N. H. 12438 [M] yg. Conard Fissure, Ark 36.6 13.5 12.2 + av 38.1 13.9 11.7 + + Mustela rixosa + average} {29.5 10.1 9.1 + maximum} [M] ad. 6 Point Barrow {30.1 10.6 9.9 + minimum} {27.6 9.3 8.6 + + average} {27.8 9.3 8.3 + maximum} [F] ad. and sad. 4 Point Barrow {28.5 9.5 8.5 + minimum} {27.0 9.0 7.9 + + Mustela rixosa + average} {29.5 10.1 8.2 + maximum} [M] ad. 2, and Shaunavon {30.4 10.5 9.0 + minimum} sad. 4 {28.4 9.6 7.4 + + average} {26.1 8.9 7.2 + maximum} [F] ad. 3, and Regina and Shaunavon {27.0 9.2 7.5 + minimum} sad. 1 {24.7 8.5 6.9 + + Mustela rixosa + Swenk, Mr. 5 [M] ad. 1 mi. E Inland 11.6 8.8 + Swenk, Mr. 8 [M] ad. Inland 30.7 10.5 8.2 + + Swenk, Mr. 10 [F] ad. Inland 28.0 9.8 7.6 + U. S. N. M. 171490 [F] ad. Type 28.8 7.7 + + Mustela rixosa + U. S. N. M. 249285 [M] sad. Finleyville, Pa 29.7 10.2 + U. S. N. M. 203173 [M] sad. Waynesburg, Pa 28.6 9.5 7.7 + U. S. N. M. 206340 [M] ad. Huttonsville, W. Va 28.5 9.9 8.5 + + C. M. 7543 [F] ad. Pymatuning Swamp 28.0 9.2 8.1 + A. N. S. P. 11279 [F] ad. Beallville, Pa 28.0 9.5 7.5 + U. S. N. M. 245843 [F] ad. near Marshall, N. C 27.5 9.4 7.3 + + ================================================================================= + + D Interorbital breadth + E Orbitonasal length + F Mastoid breadth + G Zygomatic breadth + H Length } + J Breadth }Tympanic Bulla + K Depth } + L Length m1 + M Lateral } + N Medial }P4 + P Breadth }M1 + Q Length } + R Depth of Skull at Ant. margin of basioccipital + S Depth of Skull at posterior borders of Msl + + |-Tympanic Bulla-| |----P4---| |----M1---| + D E F G H J K L M N P Q R S + -------------------------------------------------------------------------------- + + angustidens + 8.1 12.1 18.7 12.1 6.2 4.5 4.4 3.3 1.65 11.0 9.6 + 8.3 11.4 4.2 4.5 3.4 1.5 9.9 + *7.5 12.2 17.1 *19.0 11.4 5.8 3.8 4.0 2.9 1.4 8.6 + 8.4 11.8 18.2 12.5 6.6 4.1 4.3 3.2 1.5 10.0 9.5 + 8.1 11.9 18.0 12.0 6.2 4.2 4.3 3.2 1.5 10.5 9.4 + + 9.6 13.6 20.4 13.0 6.7 4.6 4.9 3.8 2.1 13.2 11.0 + 9.1 13.0 20.0 13.5 6.9 4.7 4.9 3.6 1.5 12.1 10.7 + 8.9 13.5 4.0 4.3 3.2 1.6 10.4 + 9.2 13.8 4.7 4.6 3.2 1.8 10.8 + 4.5 4.9 3.9 1.9 + 9.3 12.8 13.0 6.6 4.4 4.6 3.5 1.7 13.7 10.9 + 9.2 13.3 20.2 13.2 6.7 4.5 4.7 3.5 1.8 13.0 10.8 + + eskimo + 7.4 10.1 15.6 17.8 11.3 5.4 3.8 3.6 3.6 2.7 1.4 9.5 8.4 + 7.8 10.6 16.3 18.0 11.9 5.8 4.2 3.9 3.9 2.9 1.6 10.0 8.7 + 7.1 9.3 14.5 17.0 10.1 5.0 3.5 3.2 3.3 2.5 1.1 8.5 7.7 + + 6.9 9.6 14.1 15.7 10.5 5.1 3.4 3.2 3.2 2.5 1.2 9.5 7.7 + 7.2 9.7 15.0 16.5 11.1 5.6 3.7 3.3 3.4 2.6 1.3 10.0 8.0 + 7.0 9.5 13.6 15.0 10.2 4.8 3.2 3.0 3.0 2.3 1.1 9.2 7.4 + + rixosa + 6.6 9.9 15.1 16.4 11.0 5.2 3.7 3.3 3.6 2.6 1.4 10.0 8.4 + 6.9 10.5 16.1 17.1 11.5 5.5 3.9 3.5 3.8 2.7 1.5 10.4 8.8 + 6.3 9.2 14.0 15.2 10.7 5.0 3.5 3.1 3.3 2.4 1.3 9.4 8.0 + + 5.5 8.9 13.1 14.1 9.7 4.9 3.3 3.0 3.2 2.3 1.2 8.6 7.1 + 5.9 9.5 13.6 14.6 10.0 5.0 3.5 3.1 3.3 2.4 1.3 9.0 7.2 + 5.2 8.3 12.3 13.7 9.5 4.7 3.1 2.8 2.9 2.3 1.1 8.2 7.0 + + campestris + 7.6 11.1 16.1 18.0 4.1 3.6 3.8 2.7 1.6 11.5 9.7 + 7.0 10.5 15.9 17.9 10.9 5.7 3.8 3.4 3.5 2.6 1.5 8.6 + + 5.8 9.4 14.2 10.3 5.4 1.7 3.5 3.1 3.3 2.5 1.5 9.3 7.8 + 6.1 9.1 14.1 15.0 10.2 5.1 1.6 3.8 3.2 3.4 2.5 1.5 9.3 7.7 + + allegheniensis + 10.1 15.0 16.5 10.5 5.2 4.1 3.4 3.7 2.7 1.5 9.6 8.1 + 6.7 9.5 14.7 16.1 10.5 5.4 3.4 3.5 3.2 2.5 1.3 10.2 8.0 + 7.1 10.3 15.1 16.7 10.2 5.1 3.3 3.0 3.2 2.4 1.3 10.5 8.4 + + 9.5 13.6 10.0 5.2 3.4 3.0 3.1 2.4 1.3 + 6.2 9.7 13.5 14.6 10.0 5.1 3.7 3.3 3.5 2.6 1.4 8.7 7.8 + 6.4 9.4 15.0 9.3 5.0 3.6 3.2 3.4 2.5 1.3 7.7 + + ================================================================================= + + (Abbreviations used for names of collections in the table of + measurements of Mustela) + + A. M. N. H. American Museum of Natural History + A. N. S. P. Academy of Natural Sciences of Philadelphia + Baylor U. Baylor University + B. M. British Museum of Natural History + B. S. N. Boston Society of Natural History + B. Z. M. Berlin Zoological Museum + C. A. C. California Academy of Sciences + C. M. Carnegie Museum + C. R. C. M. Charles R. Conner Museum, Washington State College + C. Z. M. University Zoological Museum, Copenhagen, Denmark + Cornell Cornell University + Cowan Ian McTaggart-Cowan, private collection + Dickey Donald R. Dickey (deceased), private collection + E. R. W. Edward R. Warren, private collection + F. M. N. H. Field Museum of Natural History + F. S. M. Florida State Museum + Kans. U. University of Kansas, Museum of Natural History + M. C. Z. Museum of Comparative Zoölogy + M. P. H. N. Musée Polonais d'Histoire Naturelle (Warsaw, Poland) + M. V. Z. Museum of Vertebrate Zoölogy, University of California + M. Z. Museum of Zoölogy, University of Michigan + N. H. R. S. Naturhistoriska Riksmuseum + N. M. C. National Museum of Canada + S. D. M. San Diego Society of Natural History + Snyder W. E. Snyder, Beaver Dam, Wisconsin + Stan. U. Leland Stanford Junior University + Stephens Frank Stephens, private collection + Swenk, Mr. Myron H. Swenk, private collection + U. O. University of Oregon, Eugene, Oregon + U. S. N. M. United States National Museum + Walker Alex Walker, private collection + Wisc. U. University of Wisconsin + * Approximate + ** Average of 14 + + + + +LITERATURE CITED + + +ABBOT, C. C. + + 1884. A naturalist's rambles about home. D. Appleton and Co., New + York, 485 pp. + +ADDY, E. + + 1939. A weasel trails a rabbit. Jour. Mamm., 20:372-373, August 14, + 1939. + +ALDOUS, S. E., and MANWEILER, J. + + 1942. The winter food habits of the short-tailed weasel in northern + Minnesota. Jour. Mamm., 23:250-255, August 13, 1942. + +ALLEN, D. L. + + 1938. Notes on the killing technique of the New York weasel. Jour. + Mamm., 19:225-229, May 14, 1938. + + 1940. Two recent mammal records from Allegan County, Michigan. + Jour. Mamm., 21:459-460, November 14, 1940. + +ALLEN, G. M. + + 1933. The least weasel a circumboreal species. Jour. Mamm., + 14:316-319, November 13, 1933. + +ALLEN, J. A. + + 1889. Notes on a collection of mammals from southern México, with + descriptions of new species of the Genera Sciurus, Tamias, + and Sigmodon. Bull. Amer. Mus. Nat. Hist., 2:165-181, October + 21, 1889. + + 1891. On a collection of mammals from southern Texas and + northeastern México. Bull. Amer. Mus. Nat. Hist., 3:219-228, + April 17, 1891. + + 1894. On the mammals of Aransas County, Texas, with descriptions of + new forms of Lepus and Oryzomys. Bull. Amer. Mus. Nat. Hist., + 6:165-198, 1 map, May 31, 1894. + + 1896. On mammals collected in Bexar County and vicinity, Texas, by + Mr. H. P. Attwater, with field notes by the collector. Bull. + Amer. Mus. Nat. Hist., 8:47-80, April 22, 1896. + + 1904. Mammals from southern México and Central and South America. + Bull. Amer. Mus. Nat. Hist., 20:29-80, text figs. 1-18, + February 29, 1904. + + 1906. Mammals from the states of Sinaloa and Jalisco, México, + collected by J. H. Batty during 1904 and 1905. Bull. Amer. + Mus. Nat. Hist., 22:191-262, pls. 22-33, 3 figs. in text, + July 25, 1906. + + 1908. Mammals from Nicaragua. Bull. Amer. Mus. Nat. Hist., + 24:647-670, 12 figs. in text, October 13, 1908. + + 1911. Mammals from Venezuela collected by Mr. M. A. Carriker, Jr., + 1909-1911. Bull. Amer. Mus. Nat. Hist., 30:239-273, December + 2, 1911. + + 1912. Mammals from western Colombia. Bull. Amer. Mus. Nat. Hist., + 31:71-95, April 19, 1912. + + 1916. The neotropical weasels. Bull. Amer. Mus. Nat. Hist., + 35:89-111, April 28, 1916. + + 1916A. List of mammals collected in Colombia by the American Museum + of Natural History Expeditions, 1910-1915. Bull. Amer. Mus. + Nat. Hist., 35:191-238, 1 map, May 31, 1916. + +ALSTON, A. R. + + 1879-1882. Biologia Centrali-Americana. Mammalia, xx + 220 pp., + pls. 1-22. + +AMEGHINO, F. + + 1889. Contribución al conocimiento de los mamiferos fosiles de la + República Argentina. Imprenta de Pablo E. Coni É Hijos, + Especial para obras, xxxii + 1027 pp., pls. 1-94, numerous + figures in text. + +ANDERSON, R. M. + + 1945. Three mammals of the weasel family (Mustelidae) added to the + Quebec list with descriptions of two new forms. Ann. Rept. + Provancher Soc., 25th Anniversary, pp. 56-61, November 2, + 1945. + +ARTHUR, S. C. + + 1928. The fur animals of Louisiana. State of Louisiana, Dept. + Conservation Bull., 18:1-433, illustrated, November, 1928. + +AUDUBON, J. J., and BACHMAN, J. + + 1845-1853. The viviparous quadrupeds of North America: pls. in 3 + vols., elephant folios, each of 50 pls., vol. 1, 1845; vol. + 2, 1846; vol. 3, 1848. Text in 3 vols.: vol. 1, xiv + 389; + vol. 2, 1-334; vol. 3, iv + 257. + + 1851. The quadrupeds of North America, vol. 2, pp. 1-334, pls. + 51-100. Publ. by V. G. Audubon, New York. + + 1856. The quadrupeds of North America, vol. 2, pp. xiii-xiv + + i-viii + 2-383, pls. 1-50. Publ. by V. G. Audubon, New York. + +BACHMAN, J. + + 1839. Observations on the changes of colour in birds and + quadrupeds. Trans. Amer. Philos. Soc., 6:197-239. + +BAILEY, B. + + 1929. The mammals of Sherburne County, Minnesota. Jour. Mamm., + 10:153-164, May 9, 1929. + +BAILEY, H. H. + + 1930. Correcting inaccurate ranges of certain Florida mammals and + others of Virginia and the Carolinas. The Bailey Mus. and + Library of Nat. Hist., Bull. no. 3; 4 pages (Miami, Florida), + December 1, 1930. + +BAILEY, V. + + 1905. Biological survey of Texas. N. Amer. Fauna, 25:1-222, 16 + pls., 24 figs. in text, October 24, 1905. + + 1928. Animal life of the Carlsbad Cavern. Monograph, Amer. Soc. + Mammalogists, no. 3, pp. xiii + 195 pp., 67 figs. Williams + and Wilkins Co., Baltimore [Md.]. + + 1932. Mammals of New Mexico. N. Amer. Fauna, 53:1-412, 22 pls., 58 + figs. in text, March 1, 1932. + +BAIRD, S. F. + + 1858. General report upon the zoölogy of the several Pacific + Railroad Routes. Part I, Mammals, xlvii + 757 pp., 60 pls., + July 14, 1858. + +BANGS, O. + + 1896. A review of the weasels of eastern North America. Proc. Biol. + Soc. Washington, 10:1-24, 3 pls., February 25, 1896. + + 1899. Three new weasels from North America. Proc. New England Zoöl. + Club, 1:53-57, June 9, 1899. + + 1899B. Description of a new weasel from the Rocky Mountains of + British Columbia. Proc. New England Zoöl. Club, 1:81-82, + December 27, 1899. + + 1902. Chiriquí Mammalia. Bull. Mus. Comp. Zoöl., 39:17-51, 27 figs. + in text, April, 1902. + +BARBER, C. M., and COCKERELL, T. D. A. + + 1898. A new weasel from New Mexico. Proc. Acad. Nat. Sci. + Philadelphia, 1898:188-189. + +BARRETT-HAMILTON, G. E. H. + + 1903. Abstract of a physiological hypothesis to explain the winter + whitening of mammals and birds inhabiting snowy countries, + and the more striking points in the distribution of white in + vertebrates generally. Proc. Royal Irish Acad., 24:303-314. + + 1904. Notes and descriptions of some new species and subspecies of + Mustelidae. Ann. and Mag. Nat. Hist., 13(ser. 7):388-395, + May, 1904. + +BELL, T. + + 1874. A history of British quadrupeds. John Van Voorst, London, xi + + 474 pp., illustrated. + +BELCHER, E. + + 1843. Narrative of a voyage round the world, ... in her majesty's + ship Sulphur, during the years 1836-1842.... London, H. + Colburn, 2 vols.: v. 1, xxxviii + 387; v. 2, vi + 474, + illustrated. + +BISHOP, S. C. + + 1923. Note on the nest and young of the small brown weasel. Jour. + Mamm., 4:26-27, 1 pl., February 9, 1923. + +BISSONNETTE, T. H., and BAILEY, E. E. + + 1940. Den and runway system for weasels and other small mammals in + the laboratory. Amer. Midland Nat., 24:761-763, November, + 1940. + + 1944. Experimental modification and control of molts and changes of + coat-color in weasels by controlled lighting. Ann. New York + Acad. Sci., 45:221-260, 7 pls., 1 fig., April 7, 1944. + +BONAPARTE, C. L. + + 1838. Remarks on the species of the genus Mustela. Charlesworth's + Mag. Nat. Hist., 2:37-38. + +BOOTH, E. S. + + 1946. Account of a weasel in a tree. Jour. Mamm., 26:439, February + 12, 1946. + +BORELL, A. E., and ELLIS, R. + + 1934. Mammals of the Ruby Mountains region of northeastern Nevada. + Jour. Mamm., 15:12-44, 6 pls., 1 fig. in text, February 15, + 1934. + +BOYER, R. H. + + 1943. Weasel versus squirrel in Sequoia National Park. Jour. Mamm., + 24:99-100, February 20, 1943. + +BROWN, B. + + 1908. The Conard Fissure, a Pleistocene bone deposit in northern + Arkansas: with descriptions of two new genera and twenty new + species of mammals. Mem. Amer. Mus. Nat. Hist., 9(pt. + 4):155-208, pls. 14-25, 3 figs. in text. + +BROWN, M. + + 1941. A gazetteer of entomological stations in Ecuador. Ann. Ento. + Soc. Amer., 34:809-851, 10 figs. in text, December 19, 1941. + +BURROUGHS, J. + + 1900. Squirrels and other fur-bearers. Houghton, Mifflin and + Company, 149 pp., illustrated. + +BURROUGHS, R. D. + + 1939. New York weasel preying on the cottontail rabbit. Jour. + Mamm., 20:253, May 14, 1939. + +CABRERA, A. + + 1913. Sobre algunas formas del género "Mustela." Bol. d. l. Real + Soc. Español d. Hist. Nat., 13:429-435, November, 1913. + + 1914. Sobre algunas formas del género "Mustela." Bol. d. l. Real + Soc. Español d. Hist. Nat., 14:175-176, 1 pl., March, 1914. + + 1928. Sobre Lyncodon patagonicus con descripción de una nueva + subespecie. Revista Chilean d. Hist. Nat., 32:259-263. + + 1940. Sobre carnívoros Sudamericanos. Inst. d. Mus. d. l. Univ. + Nac. d. La Plata, 5(no. 29):1-22. + +CHAPMAN, F. M. + + 1917. The distribution of bird-life in Colombia; a contribution to + a biological survey of South America. Bull. Amer. Mus. Nat. + Hist., 36: x + 729 pp., 41 pls., 21 figs. in text. + + 1926. The distribution of bird-life in Ecuador. Bull. Amer. Mus. + Nat. Hist., 55: xiii + 784 pp., 30 pls., 20 figs. in text. + + 1931. The upper zonal bird-life of mts. Roraima and Duida. Bull. + Amer. Mus. Nat. Hist., 63:1-135, 42 figs. + + 1937. The Phelps Venezuela Expedition. Nat. Hist., 40(no. + 5):760-761, December, 1937. + +COUES, E. + + 1877. Fur-bearing animals: A monograph of North American + Mustelidae. Dept. Int., U. S. Geol. Surv. Terr., Miscl. + Publ., No. 8, xiv + 348 pp., 20 pls., 1 fig. in text. + +CRIDDLE, N., and CRIDDLE, S. + + 1925. The weasels of southern Manitoba. Canadian Field-nat., + 39:142-148, September, 1925. + +CRIDDLE, S. + + 1926. The habits of Microtus minor in Manitoba. Jour. Mamm., + 7:193-200, August 9, 1926. + + 1930. The prairie pocket gopher, Thomomys talpoides rufescens. + Jour. Mamm., 11:265-280, 1 pl., August 9, 1930. + + 1947. A nest of the least weasel. Canadian Field-nat., 61:69, + April, 1947. + +DALQUEST, W. W. + + 1948. Mammals of Washington. Univ. Kansas Publ., Mus. Nat. Hist., + 2:1-444, 140 figs. in text, April 9, 1948. + +DANFORTH, C. N. + + 1925. Hair with special reference to hypertrichosis. Amer. Med. + Assoc., N Dearborn St., Chicago, Illinois, 152 pp., 64 figs. + +DAVENPORT, C. B. + + 1905. Evolution without mutation. Jour. Exp. Zoöl., 2:139-143, + April, 1905. + + 1906. The mutation theory in animal evolution. Science, N. S., + 24:556-558, November 2, 1906. + +DAVIS, W. B. + + 1944. Notes on Mexican mammals. Jour. Mamm., 25:370-403, 1 fig. in + text, December 12, 1944. + +DAVIS, W. B., and ROBERTSON, J. L., JR. + + 1944. The mammals of Culberson County, Texas. Jour. Mamm., + 25:254-273, September 8, 1944. + +DEANESLY, R. + + 1944. The reproductive cycle of the female weasel (_Mustela + nivalis_). Proc. Zoöl. Soc. London, 114(pt. 3):339-349. + +DEARBORN, N. + + 1932. Foods of some predatory fur-bearing animals in Michigan. + Univ. Michigan School Forestry and Conservation, Bull. No. + 1:1-52, 8 figs. in text, 22 charts, 10 maps. + + 1932B. Occurrence of the least weasel in Michigan. Jour. Mamm., + 13:277, August 9, 1932. + +DEGERBØL, M. + + 1935. Mammals. Part 1. Systematic notes. Rept. 5th Thule Exped...., + 2(no. 4):1-67, 12 figs. in text. + +DESMAREST, A. G. + + 1818. Nouveau Dictionnaire d' Histoire Naturelle.... Tome XIX, 619 + pp., Paris. + +DICE, L. R. + + 1921. Notes on the mammals of interior Alaska. Jour. Mamm., + 2:20-28, February 21, 1921. + +DIXON, J. + + 1931. Pikas versus weasel. Jour. Mamm., 12:72, February 12, 1931. + +DUNK, F. A. + + 1946. Least weasel in Saskatchewan. Jour. Mamm., 27:392, November + 25, 1946. + +ELLIOT, D. G. + + 1905. Descriptions of apparently new species and subspecies of + mammals from México and San Domingo. Proc. Biol. Soc. + Washington, 18:233-236, December 9, 1905. + +ERRINGTON, P. L. + + 1935. Food habits of mid-west foxes. Jour. Mamm., 16:192-200, + August 12, 1935. + + 1936. Food habits of a weasel family. Jour. Mamm., 17:406-407, + November 14, 1936. + +FARURICK, B. + + 1873. Ein Albino des kleiner Wiesels (_Mustela vulgaris_). Der + Zool. Garten, 14:17-18. + +FEILDEN, H. W. + + 1878. See as included under Nares, G. S., 1878. + +FLINTOFF, R. J. + + 1935. Stoats and weasels, brown and white. North Western Naturalist + [England], 10:214-229, September 30, 1935. + +FOLLETT, W. I. + + 1937. Prey of weasel and mink. Jour. Mamm., 18:365, August 14, + 1937. + +FREMONT, J. C. + + 1856. A narrative of the exploring expedition to Oregon and North + California, pp. 190-493, in The Life of Col. John Charles + Fremont, and his narrative of Explorations and Adventures, in + Kansas, Nebraska, Oregon and California. By Samuel M. + Smucker, New York and Auburn. + +GLOVER, F. A. + + 1942. Spring color change of the New York weasel. Pennsylvania Game + News, 13(no. 7):18, 32, October, 1942. + + 1943A. Killing techniques of the New York weasel. Pennsylvania Game + News, 13(no. 10):11, 23, January, 1943, issue. + + 1943B. A study of the winter activities of the New York weasel. + Pennsylvania Game News, 14(no. 6):8, 9, September, 1943. + +GOELDI, E. A. + + 1897 (for 1898). Ein erstes authentisches Exemplar eines echten + Wiesels aus Brasilien. Zoöl. Jahrb., Abt. f. systematik, + geogr. u. Biol., 10:556-562, pl. 21, September 15, 1897. + + 1901. O primeiro exemplar authentico deuna genuina dominha do + Brazil. Bol. do Mus. Paraense, 3:195-203, August, 1901. + + 1904. Catalogo de mammiferos. Bol. do Mus. Goeldi, 4:41-122, 6 + pls., February, 1904. + +GOBLE, F. C. + + 1942. The Guinea-worm in a Bonaparte weasel. Jour. Mamm., 23:221, + June 3, 1942. + +GOLDMAN, E. A. + + 1912. A new weasel from Costa Rica. Proc. Biol. Soc. Washington, + 25:9-10, January 23, 1912. + + 1920. Mammals of Panamá. Smithsonian Miscl. Coll., 69(no. 5):1-309, + pls. 1-39, 1 map. + +GRAY, J. E. + + 1844A. The zoölogy of the voyage of H. M. S. Sulphur, under the + command of Captain Sir Edward Belcher, R. N., C. B., F. R. G. + S., etc., during the years 1836-42. Mammalia, pp. 9-36, pls. + 1-18. + + 1864. Description of a new Mustela from Quito. Proc. Zoöl. Soc. + London, 1864:55, pl. 8. + + 1865. Revision of the genera and species of Mustelidae contained in + the British Museum. Proc. Zoöl. Soc. London, 1865:100-154, 1 + pl., 2 figs. in text. + + 1874. Notes on the varieties of the western-American weasels. Ann. + and Mag. Nat. Hist., 14(ser. 4):374-375. + +GREEN, C. V. + + 1936. Observations on the New York weasel. Jour. Mamm., 17:247-249, + August 14, 1936. + +GRINNELL, J. + + 1913. A distributional list of the mammals of California. Proc. + California Acad. Sci., 3(ser. 4):265-390, pls. 15, 16, August + 28, 1913. + + 1917. Field tests or theories concerning distributional control. + Amer. Nat., 51:115-128, February, 1917. + + 1917A. The niche-relationships of the California Thrasher. Auk, + 34:427-433, October, 1917. + + 1922. A geographical study of the kangaroo rats of California. + Univ. California Publ. Zoöl., 24:1-124, pls. 1-7, text figs. + A-X, June 17, 1922. + + 1926. Geography and evolution in the pocket gopher. Univ. + California Chronicle. July, 1926, pp. 247-262, 1 pl., 3 figs. + in text. + +GRINNELL, J., and DIXON, J. + + 1919. Natural history of the ground squirrels of California. + California State Comm. Hortic., Monthly Bull., 7:4 + 597-708, + 5 col. plates, 30 figs. in text, January 27, 1919. + +GRINNELL, J., DIXON, J., and LINSDALE, J. M. + + 1937. Fur-bearing mammals of California.... Univ. California Press, + 2 vols., xii + 375 pp., pls. 1-7, figs. 1-138, xiv + 377-777 + pp., pls. 8-13, figs. 139-345, July 22, 1937. + +GRINNELL, J., and SWARTH, H. S. + + 1913. An account of the birds and mammals of the San Jacinto Area + of southern California with remarks upon the behavior of + geographic races on the margins of their habitats. Univ. + California Publ. Zoöl., 10:197-406, pls. 6-10, 3 text figs., + October 31, 1913. + +GROSJEAN, M. S. + + 1942. A persistent weasel. Jour. Mamm., 23:443, December 30, 1942. + +GUNN, C. K. + + 1932. Color and primeness in variable mammals. Amer. Nat, + 66:546-559, 4 figs. in text, December, 1932. + +HADWEN, S. + + 1929. Color changes in Lepus americanus and other animals. Pp. + 1-12, pls. 1-5. Reprint, Canad. Jour. Research, 1:189-200, + July, 1929. + +HALL, E. R. + + 1926. Systematic notes on the subspecies of _Bassariscus astutus_ + with description of one new form from California. Univ. + California Publ. Zoöl., 30:39-50, pls. 2 and 3, September 8, + 1926. + + 1926A. Changes during growth in the skull of the rodent + Otospermophilus grammurus beecheyi. Univ. California Publ. + Zoöl., 21:355-404, 43 figs. in text, March 9, 1926. + + 1927. A new weasel from Louisiana. Proc. Biol. Soc. Washington, + 40:193-194, December 2, 1927. + + 1930. Three new genera of Mustelidae from the Later Tertiary of + North America. Jour. Mamm., 11:146-155, 2 pls., May 9, 1930. + + 1936. Mustelid mammals from the Pleistocene of North America with + systematic notes on some Recent members of the genera + Mustela, Taxidea and Mephitis. Carnegie Institution Publ., + no. 473:41-119, 5 pls., 6 figs. in text, November 20, 1936. + + 1937. Mustela cicognanii, the short tailed weasel, incorrectly + ascribed to Ohio. Amer. Midland Nat., 18:304, March, 1937. + + 1944. Classification of the ermines of eastern Siberia. Proc. + California Acad. Sci., 23(ser. 4):555-560, 1 fig. in text, + August 22, 1944. + + 1946. Mammals of Nevada. Univ. California Press, Berkeley, xi + + 710, frontispiece, colored, 11 pls., 485 figs. in text, + unnumbered silhouettes, July 1, 1946. + +HAMILTON, W. J., JR. + + 1928. Weasels eat shrews. Jour. Mamm., 9:249-250, August 9, 1928. + + 1933. The weasels of New York. Amer. Midland Nat., 14:289-344, 2 + maps, 3 figs., 4 pls., July, 1933. + +HANDLEY, C. O., JR. + + 1949. Least weasel, prey of barn owl. Jour. Mamm., 30:431, November + 14, 1949. + +HARPER, J. + + 1927. The mammals of the Okefinokee Swamp region of Georgia. Proc. + Boston Soc. Nat. Hist., 38(7):191-396, pls. 4-7, March, 1927. + +HELLER, H. H. + + 1924. Peruvian pets. Natural History, 24:479-493, 14 figs. in text, + July-August, 1924. + +HENNINGER, W. F. + + 1921. Two mammals new for Ohio. Jour. Mamm., 2:239, November 29, + 1921. + +HENSEL, R. + + 1881. Craniologische Studien. Nova Acta d. Ksl. Leop.-Carol. + Deutschen Akad. d. Nat., 42:127-195, pls. 6-13, tables A-T. + +HODGSON, B. H. + + 1841. Classified catalogue of mammals of Nepal (corrected to end of + 1841, first printed in 1832). Jour. Asiatic Soc. Bengal, + 10:907-916. + +HOLLISTER, N. + + 1913. A synopsis of the American minks. Proc. U. S. Nat. Mus., + 44:471-480, April 18, 1913. + + 1914. Descriptions of four new mammals from tropical America. Proc. + Biol. Soc. Washington, 28:141-144, July 10, 1914. + +HOLMES, S. J. + + 1925. "Age and Area" in relation to extinction. Science, N. S., + 61:77-79, January 23, 1925. + +HOWARD, W. J. + + 1935. Apparently neutral relations of weasel and squirrel. Jour. + Mamm., 16:322-323, November 15, 1935. + +HOWELL, A. B. + + 1943. An apparent mustelid trait. Jour. Mamm., 24:98-99, February + 20, 1943. + +HUXLEY, J. S. + + 1939. Ecology and taxonomic differentiation. Jour. Ecology, + 27:408-420, August, 1939. + +INGLES, L. G. + + 1939. Observations on a nest of the long-tailed weasel. Jour. + Mamm., 20:253-254, May 14, 1939. + + 1942. Observations on the short-tailed weasel in California. Jour. + Mamm., 23:446-448, 1 pl., December 30, 1942. + +JACKSON, H. H. T. + + 1913. Two new weasels from the United States. Proc. Biol. Soc. + Washington, 26:123-124, May 21, 1913. + +JÄCKEL, A. J. + + 1873. Ueber Leucismen unter den Mustelen. Der Zool. Garten, + 14:456-459. + +JAECKEL, O. + + 1902. Uber verschiedene Wege phylogenetischer Entwicklung [original + not seen--only pp. 289-290 of V. L. Kellogg's Darwinism + today]. + +KELLOGG, V. L. + + 1907. Darwinism today. Henry Holt and Co., New York, xii + 403 pp. + +KENNICOTT, R. + + 1859. The quadrupeds of Illinois, injurious and beneficial to the + farmer. Agric. Rept. for 1858, U. S. Patent Office Rept., pp. + 241-256. + +KIRK, G. L. + + 1921. Shrews and weasels. Jour. Mamm., 2:111, May 2, 1921. + +LANTZ, D. E. + + 1905. A list of Kansas mammals. Trans. Kansas Acad. Sci., + 19:171-178. + +LECHE, W. + + 1915. Zur Frage nach der stammesgeschichtlichen Bedeutung des + Milchgebisses bei den Säugetieren. Zool. Jahrb., Abt. f. + Syst., 38:275-370, 126 figs. in text. + +LEOPOLD, A. + + 1937. Killing technique of the weasel. Jour. Mamm., 18:98-99, + February 14, 1937. + +LICHTENSTEIN, M. H. C. + + 1832. Darstellung neuer oder wenig bekannter Säugethiere, pl. 42 + and corresponding text (unpaged), 1832 [not seen]. + +LLEWELLYN, L. M. + + 1942. Notes on the Alleghenian least weasel in Virginia. Jour. + Mamm., 23:439-441, December 30, 1942. + +LONG, W. S. + + 1938. The weasel an enemy of the pika. Jour. Mamm., 19:250, May 14, + 1938. + +LÖNNBERG, E. + + 1913. Mammals from Ecuador and related forms. Arkiv för Zool., + 8(no. 1):1-36, 1 pl., 1 text fig., July 12, 1913. + + 1921. A second contribution to the mammalogy of Ecuador with some + remarks on Caenolestes. Arkiv för Zool., 14(no. 4):1-104, pl. + 1, 8 figs. in text. + +LYMAN, C. P. + + 1943. Control of coat color in the varying hare _Lepus americanus_ + Erxleben. Bull. Mus. Comp. Zoöl., 93:393-461, 11 pls., + December, 1943. + +LYON, M. W., JR., and OSGOOD, W. H. + + 1909. Catalogue of the type-specimens of mammals in the United + States National Museum, including the Biological Survey + Collection. U. S. Nat. Mus. Bull., 62: ix + 325. + +MACGILLIVARY, W. + + 1848? The naturalists library, Vol. XVII. Mammalia, pp. 1-309, J. + Ogden and Co., London. + +MANNICHE, A. L. V. + + 1910. The terrestrial mammals and birds of northeast Greenland. + Meddelelser om Grønland, 45:1-200, 6 pls., 20 figs. in text, + 1910. + +MANNING, T. H. + + 1943. Notes on the mammals of south and central west Baffin Island. + Jour. Mamm., 24:47-59, 1 fig. in text, February 20, 1943. + +MATTHEW, W. D. + + 1902. On the skull of Bunaelurus, a musteline from the White River + Oligocene. Bull. Amer. Mus. Nat. Hist., 16:137-140, April 7, + 1902. + +MEARNS, E. A. + + 1891. Description of a new species of weasel, and a new subspecies + of the gray fox, from Arizona. Bull. Amer. Mus. Nat. Hist, + 3:234-238, June 5, 1891. + +MERRIAM, C. H. + + 1896. Synopsis of the weasels of North America. N. Amer. Fauna, + 11:1-33, 5 pls., 16 figs. in text, June 30, 1896. + + 1902. Five new mammals from México. Proc. Biol. Soc. Washington, + 15:67-69, March 22, 1902. + + 1906. Is mutation a factor in the evolution of higher vertebrates. + Science, N. S., 23:241-257, February, 1906. + + 1919. Criteria for the recognition of species and genera. Jour. + Mamm., 1:6-9, November 28, 1919. + +METCHNIKOFF, E. + + 1901. On the process of hair turning white. Proc. Royal Soc. + London, 69:156. + +MILLER, F. W. + + 1930. The spring molt of Mustela longicauda. Jour. Mamm., + 11:471-473, November 11, 1930. + + 1931A. The fall moult of Mustela longicauda. Jour. Mamm., + 12:150-156, 1 fig. in text, May 14, 1931. + + 1931B. A feeding habit of the long-tailed weasel. Jour. Mamm., + 12:164, May 14, 1931. + +MILLER, G. S., JR. + + 1912. Catalogue of the mammals of western Europe.... British Museum + of Natural History, London, xv + 1019 pp., 213 figs. in text. + + 1924. List of North American Recent mammals, 1923. Bull. U. S. Nat. + Mus., 128: xvi + 673 pp., April 29, 1924. + +MOORE, J. C. + + 1945. Life history notes on the Florida weasel. Proc. Florida + Academy of Science, 7(no. 4):247-263, for 1944. + +MORSE, M. + + 1939. A local study of predation upon hares and grouse during the + cyclic decimation. Jour. Wildlife Management, 3:203-211, + July, 1939. + +MULAIK, S. + + 1938. Notes on Mustela frenata frenata. Jour. Mamm., 19:104, + February 14, 1938. + +MURIE, A. + + 1935. A weasel goes hungry. Jour. Mamm., 16:321-322, November 15, + 1935. + +NARES, G. S. + + 1877. Arctic Expedition/1875-6/Journals and Proceedings/of + the/Arctic Expedition, 1875-6,/under the command of/Captain + Sir George S. Nares, R. N., K. C. B./[in continuation of + Parliamentary Papers C 1153 of 1875, and C 1560 of + 1876.]/presented to both Houses of Parliament by Command of + her Majesty, 1877./London:/printed for her Majesty's + stationary office, by/Harrison & Sons, printers in ordinary + to her majesty,/St. Martin's Lane,/[c.-1636.] Price £1 1s. + + 1878. Narrative of a voyage to the Polar Sea during 1875-6 in H. M. + ships "Alert" and "Discovery" by Captn. Sir G. S. Nares, R. + N.... commander of the expedition with notes on the Natural + History edited by H. W. Feilden, F. G. S., ... naturalist to + the expedition. In 2 vols., 2 ed., London, Sampson Low, + Marston, Searle, & Rivington, 1878. Vol. 1, xl + 395 pp.; + vol. 2, vii + 378 pp., both vols. illustrated. + +NELSON, A. W. + + 1934. Notes on Wisconsin mammals. Jour. Mamm., 15:252-253, August + 10, 1934. + +NELSON, E. W. + + 1909. The rabbits of North America. N. Amer. Fauna, 29:1-314, 13 + pls., 19 figs. in text, August 31, 1909. + + 1918. Smaller mammals of North America. Nat'l Geographic, + 33(5):371-493, with abundant illustrations, many in color, + May, 1918. + +NICHOLS, D. G., and NICHOLS, J. T. + + 1935. Notes on the New York weasel (Mustela noveboracensis). Jour. + Mamm., 16:297-299, November 15, 1935. + +NOBACK, C. V. + + 1935. Observations on the seasonal hair moult in a New York State + weasel (_Mustela noveboracensis_). Bull. New York Zoöl. Soc., + 38(no. 1):25-27, February, 1935. + +OBERTHÜR, R., and DAUTHENAY, H. + + 1905. Répertoire de Couleurs. Rennes, Impr. Oberthür; Paris, + Librairie horticole. 5p. 1., [9]-82p. illus., 3 pl. (2 col.) + _and_ 365 col. pl. in 2 portfolios. + +OEHLER, C. + + 1944. Notes on the temperament of the New York weasel. Jour. Mamm., + 25:198, May 26, 1944. + +OGNEV, S. I. + + 1935. The mammals of USSR and adjacent countries, vol. 3: viii + + 752 pp., 8 pls., 299 figs. in text, 19 maps, 52 tables. + Moskow. + +OSGOOD, F. L. + + 1935. Fluctuations in small mammal populations. Jour. Mamm., + 16:156, May 15, 1935. + + 1936. Earthworms as a supplementary food for weasels. Jour. Mamm., + 17:64, February 14, 1936. + +OSGOOD, W. H. + + 1901. Natural history of the Queen Charlotte Islands, British + Columbia; natural history of the Cook Inlet Region, Alaska. + N. Amer. Fauna, 21:1-87, 8 pls., 1 fig. in text, September + 26, 1901. + + 1909. Revision of the mice of the American genus Peromyscus. N. + Amer. Fauna, 28:1-267, 8 pls., 12 figs. in text, April 17, + 1909. + + 1909B. Biological Investigations in Alaska and Yukon Territory. N. + Amer. Fauna, 30:1-96, 5 pls., 2 figs. in text, October 7, + 1909. + + 1912. Mammals from western Venezuela and eastern Colombia. Field + Mus. Nat. Hist., Publ. no. 155, zoöl. ser., 10:33-66, pls. 4, + 5, January 10, 1912. + + 1914. Mammals of an expedition across northern Perú. Field Mus. + Nat. Hist., Publ. 176, zoöl. ser., 10(no. 12):143-185, April + 20, 1914. + +PEARCE, J. + + 1937. A captive New York weasel. Jour. Mamm., 18:483-488, November + 14, 1937. + +PEARSON, K., NETTLESHIP, E., and USHER, C. H. + + 1913. A monograph on albinism in man. Drapers Co., Research + Memoirs, Biometric series, VIII, Dept. Applied Statistics, + Univ. College, Univ. London. Text, P II, vii + 264-524 pp., + London. + +PEARSON, O. P., and PEARSON, A. K. + + 1947. Owl predation in Pennsylvania, with notes on the small + mammals of Delaware County. Jour. Mamm., 28:137-147, June 1, + 1947. + +PITT, F. + + 1921. Notes on the genetic behavior of certain characters in the + polecat, ferret, and in polecat-ferret hybrids. Jour. + Genetics, 11:99-115, 2 pls., 5 figs. in text, September, + 1921. + +POLDERBOER, E. B. + + 1942. Habits of the least weasel (_Mustela rixosa_) in northeastern + Iowa. Jour. Mamm., 23:145-147, June 3, 1942. + + 1948. Late fall sexual activity in an Iowa least weasel. Jour. + Mamm., 29:296, August 31, 1948. + +POLDERBOER, E. B., KUHN, L. W., and HENDRICKSON, G. O. + + 1941. Winter and spring habits of weasels in central Iowa. Jour. + Wildlife Management, 5:115-119, 1 pl., January, 1941. + +PREBLE, E. A. + + 1908. A biological investigation of the Athabasca-Mackenzie region. + N. Amer. Fauna, 27:1-574, 25 pls., 13 figs. in text, October + 26, 1908. + +QUICK, H. F. + + 1944. Habits and economics of the New York weasel in Michigan. + Jour. Wildlife Management, 8:71-78, January, 1944. + +RHOADS, S. N. + + 1894. Contributions to the mammalogy of Florida. Proc. Acad. Nat. + Sci. Philadelphia, pp. 152-161, June 19, 1894. + +RIDGWAY, R. + + 1912. Color standards and color nomenclature. Published by the + author, Washington, D. C., iii + 43 pp., 53 pls. + +ROBINSON, W., and LYON, M. W. + + 1901. An annotated list of mammals collected in the vicinity of La + Guaira, Venezuela. Proc. U. S. Nat. Mus., 24:135-162, + October, 1901. + +ROTHSCHILD, M. + + 1942. Change of pelage in the stoat Mustela erminea L. Nature, + 149:78, January 17, 1942. + +RUSSELL, W. C. + + 1930. Weasel badly injured by king snake. Jour. Mamm., 11:504-505, + November 11, 1930. + +SANDERSON, G. C. + + 1949. Growth and behavior of a litter of captive long-tailed + weasels. Jour. Mamm., 30:412-415, 1 fig. in text, November + 14, 1949. + +SCHLOSSER, M. + + 1888. Die Affen, Lemuren, Chiropteren, Insectivoren, Marsupialier, + Creodonten und Carnivoren des Europäischen Tertiärs. Alfred + Hölder, K. K. Hof.-und Univ.-Buchhändler, Wien. Part 2, pp. + 225-386, pls. 6-9. + +SCHUMACHER, S. + + 1928. Wie kommt die stellenweise Gelbfärbung des winterweissen + Wiesels (Mustela erminea L.) zustande? Zeitschr. f. Morph, u. + Ökologie der Tiere, 11:229-234, 1 fig., July 3, 1928. + +SCHWALBE, G. + + 1893. Ueber den Farbenwechsel winterweisser Thiere. Morphologische + Arbeiten, 2:483-600, Jena. + +SETON, E. T. + + 1929. Lives of game animals. Vol. 2, 1929, xvii + 746 pp., + illustrated; vol. 4, xxii + 949 pp., Doubleday, Doran & Co., + New York. + +SHAW, W. T. + + 1921. The nest of the Washington weasel (Mustela washingtoni). + Jour. Mamm., 2:167-168, August 19, 1921. + +SHELDON, W. G. + + 1932. Mammals collected or observed in the vicinity of Laurier + Pass, B. C. Jour. Mamm., 13:196-203, August 9, 1932. + +SIMPSON, G. G. + + 1946. Palaeogale and allied early mustelids. Amer. Mus. Novitates, + 1320:1-14, 4 figs. in text, May 28, 1946. + +SNYDER, L. L., and LOGIER, E. B. S. + + 1930. No. 3. A faunal investigation of King Township, York County, + Ontario. Trans. Royal Canadian Institute, 17(pt. 2):167-208. + +SOPER, J. D. + + 1919. Notes on Canadian weasels. Canadian Field-nat., 33:43-47, + September, 1919. + + 1921. Curious palatal obstruction in Mustela longicauda. Jour. + Mamm., 2:37-38, 1 fig. in text, February 10, 1921. + + 1942. Mammals of Wood Buffalo Park, northern Alberta and District + of Mackenzie. Jour. Mamm., 23:119-145, 2 pls., 1 map in text, + June 3, 1942. + + 1946. Mammals of the northern Great Plains along the International + Boundary in Canada. Jour. Mamm., 27:127-153, 1 fig. in text, + May 14, 1946. + + 1948. Mammal notes from the Grand Prairie-Peace River region, + Alberta. Jour. Mamm., 29:49-64, 1 pl., 1 fig. in text, + February 13, 1948. + +SOWLS, L. K. + + 1948. The Franklin ground squirrel, _Citellus franklinii_ (Sabine), + and its relationship to nesting ducks. Jour. Mamm., + 29:113-137, 3 pls., 3 figs. in text, May 14, 1948. + +STANFORD, J. S. + + 1931. Notes on small mammals of Utah. Journ. Mamm., 12:356-363, + November 11, 1931. + +STEPHENS, F. + + 1906. California mammals. West Coast Publishing Co., San Diego, + California, pp. 1-351, numerous figs. and maps. + +STRECKER, J. K. + + 1924. The mammals of McLennan County, Texas. The Baylor Bull., + Baylor Univ., Waco, Texas, 27(no. 1):1-20, September, 1924. + + 1926. A check-list of the mammals of Texas exclusive of the Sirenia + and Cetacea. The Baylor Bull., Baylor Univ., Waco, Texas, + 29(no. 3):1-48, August, 1926. + +STRONG, W. D. + + 1930. Notes on mammals of the Labrador Interior. Jour. Mamm., + 11:1-10, February 11, 1930. + +SUMNER, F. B. + + 1932. Genetic, distributional, and evolutionary studies of the + subspecies of deer mice (Peromyscus). Bibliographia Genetica, + 9:1-106, 24 figs. in text. + +SURBER, T. + + 1932. The mammals of Minnesota. Minnesota Dept. Conservation, Div. + Game and Fish, Saint Paul, Minnesota, pp. 1-84, illustrated. + +SUTTON, G. K. + + 1929. The Alleghenian least weasel in Pennsylvania. Jour. Mamm., + 252-254, 1 fig. in text, August 10, 1929. + +SVIHLA, A. + + 1931. Habits of the New York weasel in captivity. Jour. Mamm., + 12:67-68, February 12, 1931. + +SWANSON, G., and FRYKLUND, P. O. + + 1935. The least weasel in Minnesota and its fluctuation in numbers. + Amer. Midland Nat., 16:120-126, figs. 1-6, 1935. + +SWENK, M. H. + + 1926. Notes on Mustela campestris Jackson, and on the American + forms of least weasels. Jour. Mamm., 7:313-330, 1 fig. in + text, November 23, 1926. + +TACZANOWSKI, L. + + 1874. Description d' une nouvelle espéce de _Mustela_ du Pérou + central. Proc. Zool. Soc. London, 1874:311-312, pl. 48, May + 19, 1874. + + 1881. Description d' une nouvelle belette du Pérou septentrional. + Proc. Zool. Soc. London, 1881:647-649, May 17, 1881. + + 1881. Description d' une nouvelle espéce du genre _Mustela_ du + Pérou nord-oriental. Proc. Zool. Soc. London, 1881:835-836, + November 15, 1881. + +TATE, G. H. H. + + 1931. Random observations on habits of South American mammals. + Jour. Mamm., 12:248-256, August 24, 1931. + +THOMAS, O. + + 1911. The mammals of the tenth edition of Linnaeus; an attempt to + fix the types of the genera and the exact bases and + localities of the species. Proc. Zoöl. 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 + University and the National Geographic Society. Proc. U. S. + Nat. Mus., 58:217-249, pls. 14-15. + +THURBER, W. A. + + 1940. A weasel attacks a varying hare. Jour. Mamm., 21:356, August + 14, 1940. + +TSCHUDI, J. J. + + 1844. Untersuchungen über die Fauna Peruana. Therologie. Druck und + verlag von Scheitlin und Zollikofer, St. Gallen, xxx + 262 + pp., 18 pls. + +VESTAL, E. H. + + 1937. Activities of a weasel at a wood rat colony. Jour. Mamm., + 18:364, August 14, 1937. + + 1938. Biotic relations of the wood rat (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, + November 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 + 259 + pp., illustrated. + +WINECOFF, T. E. + + 1930. Least weasel in Pennsylvania. Jour. Mamm., 11:312-313, + August, 1930. + +WOLDRICH, J. N. + + 1884. Diluviale Fauna von Zuzlawitz bei Winterberg im Bohmerwalde. + Dritter Theil.... Sitzungsberich. d. K. Akad. d. Wissen. + Math.--Natur.--Classe, 88(heft 3, erste abth.):978-1057, 3 + pls., 2 figs. in text. + +WRIGHT, P. L. + + 1942A. Delayed implantation in the long-tailed weasel (Mustela + frenata), the short-tailed weasel (Mustela cicognani), and + the marten (Martes americana). Anat. Rec., 83:341-353, 2 + pls., July, 1942. + + 1942B. A correlation between the spring molt and spring changes in + the sexual cycle in the weasel. Jour. Exp. Zoöl., 91:103-110, + October, 1942. + + 1947. The sexual cycle of the male long-tailed weasel (Mustela + frenata). Jour. Mamm., 28:343-352, 1 pl., 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 + frenata). Anat. Rec., 100:595-607, 2 pls., April, 1948. + + 1950. Effects of gonadotropic hormone on the pelage of white + winter long-tailed weasels; abstract. Anat. Rec., 106(no. + 2):130, February, 1950. + +ZIMMERMANN, K. + + 1943. Zur Kenntnis deutscher Maus-und Zwerg-Wiesel. Zeitschr. fur + Säugetierkunde, 15(3):289-298, 2 pls., 2 figs. in text, 3 + tables, January 11, 1943. + +_Transmitted May 22, 1950_. + + +[Illustration: PLATE 2. Photographs, retouched, of skulls in dorsal +view of nine subspecies of _Mustela erminea_. Natural size. + + _a._ _Mustela erminea arctica_, [M] ad., 178405, U. S. Nat. Mus., + Tanana, Alaska. + + _b._ _Mustela erminea semplei_, [M] sad., 6499, Carnegie Mus., + Southhampton Island. + + _c._ _Mustela erminea kadiacensis_, [M] ad., 107496, U. S. N. M., + Kodiak Island, Alaska. + + _d._ _Mustela erminea richardsonii_, [M] ad., 133847, U. S. N. M., Ft. + Franklin, MacK. + + _e._ _Mustela erminea cicognanii_, [M] ad., 53788, Mus. Vert. Zoöl., + Lopez, Pennsylvania. + + _f._ _Mustela erminea bangsi_, [M] ad., 18130, Field Mus. Nat. Hist., + Aitkin, Minn. + + _g._ _Mustela erminea invicta_, [M] ad., 90759, Mus. Vert. Zoöl., Pilot + Creek, Idaho. + + _h._ _Mustela erminea alascensis_, [M] ad., 74665, Mus. Vert. Zoöl., + Windham, Alaska. + + _i._ _Mustela erminea salva_, [M] ad., 74641, M. V. Z., Mole Harbor, + Admiralty Id., Alaska.] + + +[Illustration: PLATE 3. Photographs, retouched, of skulls in ventral +view of nine subspecies of _Mustela erminea_. Natural size. + + _a._ _Mustela erminea arctica_, [M] ad., 178405, U. S. Nat. Mus., + Tanana, Alaska. + + _b._ _Mustela erminea semplei_, [M] sad., 6499, Carnegie Mus., + Southampton Island. + + _c._ _Mustela erminea kadiacensis_, [M] ad., 107496, U. S. N. M., + Kodiak Island, Alaska. + + _d._ _Mustela erminea richardsonii_, [M] ad., 133847, U. S. N. M., Ft. + Franklin, MacK. + + _e._ _Mustela erminea cicognanii_, [M] ad., 53788, Mus. Vert. Zoöl., + Lopez, Pennsylvania. + + _f._ _Mustela erminea bangsi_, [M] ad., 18130, Field Mus. Nat. Hist., + Aitkin, Minn. + + _g._ _Mustela erminea invicta_, [M] ad., 90759, Mus. Vert. Zoöl., Pilot + Creek, Idaho. + + _h._ _Mustela erminea alascensis_, [M] ad., 74665, Mus. Vert. Zoöl., + Windham, Alaska. + + _i._ _Mustela erminea salva_, [M] ad., 74641, M. V. Z., Mole Harbor, + Admiralty Id., Alaska.] + + +[Illustration: PLATE 4. Photographs, retouched, of skulls in lateral +view of ten subspecies _Mustela erminea_. Natural size. + + _a._ _Mustela erminea arctica_, [M] ad., 178405, U. S. Nat. Mus., + Tanana, Alaska. + + _b._ _Mustela erminea semplei_, [M] sad., 6499, Carnegie Mus., + Southampton Island. + + _c._ _Mustela erminea kadiacensis_, [M] ad., 107496, U. S. N. M., + Kodiak Island, Alaska. + + _d._ _Mustela erminea richardsonii_, [M] ad., 133847, U. S. N. M., Ft. + Franklin, MacK. + + _e._ _Mustela erminea cicognanii_, [M] ad., 53788, Mus. Vert. Zoöl., + Lopez, Pennsylvania. + + _f._ _Mustela erminea bangsi_, [M] ad., 18130, Field Mus. Nat. Hist., + Aitkin, Minn. + + _g._ _Mustela erminea invicta_, [M] ad., 90759, Mus. Vert. Zoöl., Pilot + Creek, Idaho. + + _h._ _Mustela erminea alascensis_, [M] ad., 74665, Mus. Vert. Zoöl., + Windham, Alaska. + + _i._ _Mustela erminea salva_, [M] ad., 74641, M. V. Z., Mole Harbor, + Admiralty Id., Alaska. + + _j._ _Mustela erminea initis_, [M] ad., 289, Mus. Vert. Zoöl., Saook + Bay, Alaska.] + + +[Illustration: PLATE 5. Photos, retouched, of skulls of 9 subspecies of +_Mustela erminea_, × 1. + + _a._ _Mustela erminea initis_, [M] ad., 289, Mus. Vert. Zoöl., Saook + Bay, Alaska. + + _b._ _Mustela erminea celenda_, [M] ad., 1053, Los Angeles Mus., Craig, + Alaska. + + _c._ _Mustela erminea seclusa_, [M] ad., 31232, M. V. Z., Port Santa + Cruz, Alaska. + + _d._ _Mustela erminea haidarum_, [M] ad., 230777, U. S. N. M., Graham + Island, B. C. + + _e._ _Mustela erminea anguinae_, [M] ad., 13508, Nat. Mus. Canada, Cape + Scott, V. I., B. C. + + _f._ _Mustela erminea fallenda_, [M] ad., 7096, Nat. Mus. Canada, + Huntingdon, B. C. + + _g._ _Mustela e. olympica_, [M] ad., 90738, U. S. N. M., near head of + Soleduc Riv., Wash. + + _h._ _Mustela erminea streatori_, [M] ad., 133, Coll. of Alex Walker, + Blaine, Oregon. + + _i._ _Mustela erminea gulosa_, [M] ad., 82177, U. S. Nat. Mus., Trout + Lake, Wash.] + + +[Illustration: PLATE 6. Photos, retouched, of skulls of 9 subspecies of +_Mustela erminea_, × 1. + + _a._ _Mustela erminea initis_, [M] ad., 289, Mus. Vert. Zoöl., Saook + Bay, Alaska. + + _b._ _Mustela erminea celenda_, [M] ad., 1053, Los Angeles Mus., Craig, + Alaska. + + _c._ _Mustela erminea seclusa_, [M] ad., 31232, M. V. Z., Port Santa + Cruz, Alaska. + + _d._ _Mustela erminea haidarum_, [M] ad., 230777, U. S. N. M., Graham + Island, B. C. + + _e._ _Mustela erminea anguinae_, [M] ad., 13508, Nat. Mus. Canada, Cape + Scott, V. I., B. C. + + _f._ _Mustela erminea fallenda_, [M] ad., 7096, Nat. Mus. Canada, + Huntingdon, B. C. + + _g._ _Mustela e. olympica_, [M] ad., 90738, U. S. N. M., near head of + Soleduc Riv., Wash. + + _h._ _Mustela erminea streatori_, [M] ad., 133, Coll. of Alex Walker, + Blaine, Oregon. + + _i._ _Mustela erminea gulosa_, [M] ad., 82177, U. S. Nat. Mus., Trout + Lake, Wash.] + + +[Illustration: PLATE 7. Photos, retouched, of skulls, of subspecies of +_Mustela erminea_. Natural size. + + _a._ _Mustela erminea celenda_, [M] ad., 1053, Los Angeles Mus., Craig, + Alaska. + + _b._ _Mustela erminea seclusa_, [M] ad., 31232, M. V. Z., Port Santa + Cruz, Alaska. + + _c._ _Mustela erminea haidarum_, [M] ad., 230777, U. S. N. M., Graham + Island, B. C. + + _d._ _Mustela erminea anguinae_, [M] ad., 13508, Nat. Mus. Canada, Cape + Scott, V. I., B. C. + + _e._ _Mustela erminea fallenda_, [M] ad., 7096, Nat. Mus. Canada, + Huntingdon, B. C. + + _f._ _Mustela erminea olympica_, [M] ad., 90738, U. S. N. M., near head + of Soleduc Riv., Wash. + + _g._ _Mustela erminea streatori_, [M] ad., 133, Coll. of Alex Walker, + Blaine, Oregon. + + _h._ _Mustela erminea gulosa_, [M] ad., 82177, U. S. Nat. Mus., Trout + Lake, Wash. + + _i_, _j_, _k_. _Mustela erminea muricus_, [M] ad., 41501, M. V. Z., + Baker Creek, 8675 ft., Nev. + + _l_, _m_. _Mustela erminea angustidens_, [M]?, sad., 12437, A. M. N. + H., Conard Fissure, Ark. + + _n._ _Mustela erminea angustidens_, [M]?, ad., 12441, A. M. N. H., + Conard Fissure, Ark.] + + +[Illustration: PLATE 8. Photos, retouched, of _Mustela erminea +angustidens_. All in Amer. Mus. Nat. Hist., from Conard Fissure, +Arkansas. Pleistocene in age, × 1. + + _a._ Adult, probably male, 12441. + + _b._ Subadult, probably male, 12437. + + _c._ Adult, probably male, 12444. + + _d._ Adult, probably male, 12441. + + _e._ Subadult, probably male, 12437. + + _f._ Young, probably male, 12438. + + _g_, _h_. Adult, type, probably female, 12432. + + _i._ Adult, probably female, 12433.] + + +[Illustration: PLATE 9. Photographs, retouched, of skulls in dorsal +view of 9 subspecies of _Mustela erminea_. Natural size. + + _a._ _Mustela erminea arctica_, [F] ad., 35895, Field Mus. Nat. Hist., + Point Barrow, Alaska. + + _b._ _Mustela erminea semplei_, [F] ad., 6600, Carnegie Mus., + Southhampton Island. + + _c._ _Mustela erminea kadiacensis_, [F] ad., 98042, U. S. Nat. Mus., + Kadiak, Alaska. + + _d._ _Mustela erminea richardsonii_, [F] ad., 129703, U. S. N. M., Fort + Resolution, MacK. + + _e._ _Mustela erminea cicognanii_, [F] ad., 7460, Carnegie Mus., + Pymatuning Swamp, Pa. + + _f._ _Mustela erminea bangsi_, [F] ad., 8679, Univ. Wisconsin, T. 61N, + R. 26W, Minn. + + _g._ _Mustela erminea invicta_, [F] ad., 90820, M. V. Z., 1-1/2 mi. W + Iron Mtn., Idaho. + + _h._ _Mustela erminea alascensis_, [F] ad., 74422, U. S. Nat. Mus., + Juneau, Alaska. + + _i._ _Mustela erminea salva_, [F] ad., 74655, Mus. Vert. Zoöl., Mole + Harbor, Alaska.] + + +[Illustration: PLATE 10. Photographs, retouched, of skulls in dorsal +view of 9 subspecies of _Mustela erminea_. Natural size. + + _a._ _Mustela erminea arctica_, [F] ad., 35895, Field Mus. Nat. Hist., + Point Barrow, Alaska. + + _b._ _Mustela erminea semplei_, [F] ad., 6600, Carnegie Mus., + Southhampton Island. + + _c._ _Mustela erminea kadiacensis_, [F] ad., 98042, U. S. Nat. Mus., + Kadiak, Alaska. + + _d._ _Mustela erminea richardsonii_, [F] ad., 129703, U. S. N. M., Fort + Resolution, MacK. + + _e._ _Mustela erminea cicognanii_, [F] ad., 7460, Carnegie Mus., + Pymatuning Swamp, Pa. + + _f._ _Mustela erminea bangsi_, [F] ad., 8679, Univ. Wisconsin, T. 61N, + R. 26W, Minn. + + _g._ _Mustela erminea invicta_, [F] ad., 90820, M. V. Z., 1-1/2 mi. W + Iron Mtn., Idaho. + + _h._ _Mustela erminea alascensis_, [F] ad., 74422, U. S. Nat. Mus., + Juneau, Alaska. + + _i._ _Mustela erminea salva_, [F] ad., 74655, Mus. Vert. Zoöl., Mole + Harbor, Alaska.] + + +[Illustration: PLATE 11. Photographs, retouched, of skulls in lateral +view of twelve subspecies of _Mustela erminea_. Natural size. + + _a._ _Mustela erminea arctica_, [F] ad., 35895, Field Mus. Nat. Hist., + Point Barrow, Alaska. + + _b._ _Mustela erminea semplei_, [F] ad., 6600, Carnegie Mus., + Southhampton Island. + + _c._ _Mustela erminea kadiacensis_, [F] ad., 98042, U. S. Nat. Mus., + Kadiak, Alaska. + + _d._ _Mustela erminea richardsonii_, [F] ad., 129703, U. S. Nat. Mus., + Fort Resolution, MacK. + + _e._ _Mustela erminea cicognanii_, [F] ad., 7460, Carnegie Mus., + Pymatuning Swamp, Pa. + + _f._ _Mustela erminea bangsi_, [F] ad., 8679, Univ. Wisconsin, T. 61N, + R. 26W, Minn. + + _g._ _Mustela erminea invicta_, [F] ad., 90820, M. V. Z., 1-1/2 mi. W + Iron Mtn., Idaho. + + _h._ _Mustela erminea alascensis_, [F] ad., 74422, U. S. Nat. Mus., + Juneau, Alaska. + + _i._ _Mustela erminea salva_, [F] ad., 74655, Mus. Vert. Zoöl., Mole + Harbor, Alaska. + + _j._ _Mustela erminea haidarum_, [F] ad., 100624, U. S. Nat. Mus., + Moresby Island, B. C. + + _k._ _Mustela erminea anguinae_, [F] ad., 13673, Nat. Mus. Canada, Cape + Scott, V. I., B. C. + + _l._ _Mustela erminea fallenda_, [F] ad., 7284, Nat. Mus. Canada, + Huntingdon, B. C.] + + +[Illustration: PLATE 12. Photographs, retouched, of skulls in dorsal +view of eight subspecies of _Mustela erminea_. Natural size. + + _a._ _Mustela erminea haidarum_, [F] ad., 100624, U. S. N. M., Moresby + Island, B. C. + + _b._ _Mustela erminea anguinae_, [F] ad., 13673, N. M. Canada, Cape + Scott, V. I., B. C. + + _c._ _Mustela erminea fallenda_, [F] ad., 7284, Nat. Mus. Canada, + Huntingdon, B. C. + + _d._ _Mustela erminea olympica_, [F] ad., 242133, U. S. Nat. Mus., + Hayes Creek, Wash. + + _e._ _Mustela erminea streatori_, [F] ad., 9040, D. R. Dickey Coll., + Blaine, Oregon. + + _f._ _Mustela erminea gulosa_, [F] ad., 77370, U. S. Nat. Mus., Trout + Lake, Wash. + + _g._ _Mustela erminea muricus_, [F] ad., 41502, Mus. Vert. Zoöl., Baker + Creek, Nevada. + + _h._ _Mustela erminea angustidens_, [F]?, ad., 12435, A. M. N. H., + Conard Fissure, Ark. + + _i._ _Mustela erminea angustidens_, [F]?, ad., 11766, A. M. N. H., + Conard Fissure, Ark.] + + +[Illustration: PLATE 13. Photographs, retouched, of skulls in ventral +view of eight subspecies of _Mustela erminea_. Natural size. + + _a._ _Mustela erminea haidarum_, [F] ad., 100624, U. S. N. M., Moresby + Island, B. C. + + _b._ _Mustela erminea anguinae_, [F] ad., 13673, Nat. Mus. Canada, Cape + Scott, B. C. + + _c._ _Mustela erminea fallenda_, [F] ad., 7284, Nat. Mus. Canada, + Huntingdon, B. C. + + _d._ _Mustela erminea olympica_, [F] ad., 242133, U. S. Nat. Mus., + Hayes Creek, Wash. + + _e._ _Mustela erminea streatori_, [F] ad., 9040, D. R. Dickey Coll., + Blaine, Oregon. + + _f._ _Mustela erminea gulosa_, [F] ad., 77370, U. S. Nat. Mus., Trout + Lake, Wash. + + _g._ _Mustela erminea muricus_, [F] ad., 41502, Mus. Vert. Zoöl., Baker + Creek, Nevada. + + _h._ _Mustela erminea angustidens_, [F]?, ad., 12435, A. M. N. H., + Conard Fissure, Ark. + + _i._ _Mustela erminea angustidens_, [F]?, ad., 11766, A. M. N. H., + Conard Fissure, Ark.] + + +[Illustration: PLATE 14. Photographs, retouched, of _M. erminea_ and +_M. minuta_. Natural size. + + _a._ _Mustela erminea olympica_, [F] ad., 242133, U. S. Nat. Mus., + Hayes Creek, Wash. + + _b._ _Mustela erminea streatori_, [F] ad., 9040, D. R. Dickey Coll., + Blaine, Oregon. + + _c._ _Mustela erminea gulosa_, [F] ad., 77370, U. S. Nat. Mus., Trout + Lake, Wash. + + _d._ _Mustela erminea muricus_, [F] ad., 41502, Mus. Vert. Zoöl., Baker + Creek, Nevada. + + _e._ _Mustela erminea angustidens_, adult, probably female, type, + 12432, Amer. Mus. Nat. Hist., with lower jaw, Conard Fissure, Ark. + + _f._ _M. e. angustidens_, [F]?, ad., 12435, A. M. N. H., Conard + Fissure, Ark. + + _g._ _Mustela rixosa eskimo_, [M] sad., 43288, Mus. Vert. Zoöl., + Barrow, Alaska. + + _h._ _Mustela rixosa eskimo_, [F] sad., 40059, Mus. Vert. Zoöl., + Barrow, Alaska. + + _i._ _Mustela rixosa rixosa_, [M] ad., 11743, Nat. Mus. Canada, + Shaunavon, Sask. + + _j._ _Mustela rixosa rixosa_, [F] ad., 12679, Nat. Mus. Canada, south + of Shaunavon, Sask. + + _k._ _Mustela r. allegheniensis_, [M] ad., 35381, Field M. N. H., + Portage Twp., Ohio. + + _l._ _Mustela rixosa allegheniensis_, [F] ad., 33021, Field M. N. H., + Stryker, Ohio.] + + +[Illustration: PLATE 15. Photographs, retouched, of _Mustela rixosa_. +Natural size. + + _a._ _Mustela rixosa eskimo_, [M] sad., 43288, Mus. Vert. Zoöl., + Barrow, Alaska. + + _b._ _Mustela rixosa rixosa_, [M] ad., 11743, Nat. Mus. Canada, + Shaunavon, Saskatchewan. + + _c._ _Mustela rixosa allegheniensis_, [M] ad., 33581, Field Mus. Nat. + Hist., Portage Township, Wood County, Ohio. + + _d._ _Mustela rixosa campestris_, [M] ad., 261830, U. S. Nat. Mus., + shore of Sand Lake, South Dakota. + + _e._ _Mustela rixosa eskimo_, [M] sad., same specimen shown in _a_. + + _f._ _Mustela rixosa rixosa_, [M] same specimen shown in _b_. + + _g._ _Mustela rixosa allegheniensis_, [M] ad., same specimen shown in + _c_. _c_. + + _h._ _Mustela rixosa campestris_, [M] ad., same specimen shown in _d_. + + _i._ _Mustela rixosa eskimo_, [F] sad., 40059, Mus. Vert. Zoöl., + Barrow, Alaska. + + _j._ _Mustela rixosa rixosa_, [F] ad., 12679, Nat. Mus. Canada, south + of Shaunavon, Saskatchewan. + + _k._ _Mustela rixosa allegheniensis_, [F] ad., 33021, Field Mus. Nat. + Hist., Stryker, Ohio. + + _l._ _Mustela rixosa allegheniensis_, [F] ad., same specimen shown in + _k_. + + _m._ _Mustela rixosa eskimo_, [F] sad., same specimen shown in _i_. + + _n._ _Mustela rixosa rixosa_, [F] ad., same specimen shown in _j_. + + _o._ _Mustela rixosa campestris_, [M] ad., same specimen shown in _d_ + and _h_.] + + +[Illustration: PLATE 16. Photographs, retouched, of skulls in dorsal +view of nine subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata noveboracensis_, [M] ad., 77112, U. S. Nat. Mus., + Wilmington, Massachusetts. + + _b._ _Mustela frenata occisor_, [M] ad., 7267, Mus. Comp. Zool., + Moosehead Lake, Maine. + + _c._ _Mustela frenata primulina_, [M] ad., 3325, Mus. Nat. Hist., Univ. + Kansas, Clinton, Kansas. + + _d._ _Mustela frenata arthuri_, [M] sad., 37515, Mus. Vert. Zoöl., + type, Remy, Louisiana. + + _e._ _Mustela frenata olivacea_, [M] ad., 180802, U. S. Nat. Mus., + type, Biological Surveys Collection, Autaugaville, Alabama. + + _f._ _Mustela frenata peninsulae_, [M] ad., 49387, Florida State Mus., + Apopka, Florida. + + _g._ _Mustela frenata spadix_, [M] ad., 53745, Mus. Vert. Zoöl., Elk + River, Minnesota. + + _h._ _Mustela frenata longicauda_, [M] ad., 15875, Amer. Mus. Nat. + Hist., Red Deer, Alberta. + + _i._ _Mustela frenata oribasa_, [M] ad., 43817, Mus. Vert. Zoöl., + Isaacs Lake, British Columbia.] + + +[Illustration: PLATE 17. Photos, retouched, of skulls of males, in +ventral view, of 9 subspecies of _Mustela frenata_, × 1. Data for _a_ +to _i_ are given on Plate 18. + + _a._ _M. f noveboracensis_ + + _b._ _M. f. occisor_ + + _c._ _M. f. primulina_ + + _d._ _M. f. arthuri_ + + _e._ _M. f. olivacea_ + + _f._ _M. f. peninsulae_ + + _g._ _M. f. spadix_ + + _h._ _M. f. longicauda_ + + _i._ _M. f. oribasus_] + + +[Illustration: PLATE 18. Photographs, retouched, of skulls in lateral +view of ten subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata noveboracensis_, [M] ad., 77112, U. S. Nat. Mus., + Wilmington, Mass. + + _b._ _Mustela frenata occisor_, [M] ad., 7267, M. C. Z., Moosehead + Lake, Maine. + + _c._ _Mustela frenata primulina_, [M] ad., 3325, Mus. Nat. Hist., Univ. + Kansas, Clinton, Kans. + + _d._ _Mustela frenata arthuri_, [M] ad., 37515, Mus. Vert. Zoöl., type, + Remy, Louisiana. + + _e._ _Mustela frenata olivacea_, [M] ad., 180802, U. S. Nat. Mus., + type, Biological Surveys Collection, Autaugaville, Alabama. + + _f._ _Mustela frenata peninsulae_, [M] ad., 49387, Florida State Mus., + Apopka, Florida. + + _g._ _Mustela frenata spadix_, [M] ad., 53795, Mus. Vert. Zoöl., Elk + River, Minnesota. + + _h._ _Mustela frenata longicauda_, [M] ad., 15875, Amer. Mus. N. H., + Red Deer, Alberta. + + _i._ _Mustela frenata oribasus_, [M] ad., 43817, Mus. Vert. Zoöl., + Isaacs Lake, B. C. + + _j._ _Mustela frenata alleni_, [M] ad., 7440/9136, A. M. N. H., Hill + City, S. D.] + + +[Illustration: PLATE 19. Photographs, retouched, of skulls in dorsal +view of nine subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata alleni_, [M] ad., 7440/9136, Amer. Mus. Nat. + Hist., Hill City, South Dakota. + + _b._ _Mustela frenata arizonensis_, [M] ad., 55211, Mus. Vert. Zoöl., + Government Prairie, Arizona. + + _c._ _Mustela frenata nevadensis_, [M] ad., 22116, Mus. Vert. Zoöl., + Chinquapin, California. + + _d._ _Mustela frenata effera_, [M] ad., 33637, Amer. Mus. Nat. Hist., + Ironside, Oregon. + + _e._ _Mustela frenata washingtoni_, [M] ad., 226758, U. S. Nat. Mus., + Gotchen Creek, Washington. + + _f._ _Mustela frenata saturata_, [M] ad., 65930, U. S. Nat. Mus., type, + Siskiyou, Oregon. + + _g._ _Mustela frenata altifrontalis_, [M] ad., 391, Coll. Alex Walker, + Blaine, Oregon. + + _h._ _Mustela frenata oregonensis_, [M] sad., 43828/32019, U. S. Nat. + Mus., Grants Pass, Oregon. + + _i._ _Mustela frenata munda_, [M] ad., 5459, Mus. Comp. Zool., type, + Point Reyes, California.] + + +[Illustration: PLATE 20. Photos, retouched, of skulls of males in +ventral view of 9 subspecies of _Mustela frenata_, × 1. Data for a to i +on Plates 18 and 21. + + _a._ _M. f. alleni_ + + _b._ _M. f. arizonensis_ + + _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.] + + +[Illustration: PLATE 21. Photographs, retouched, of skulls, in lateral +view, of nine subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata arizonensis_, [M] ad., 55211, Mus. Vert. Zoöl., + Government Prairie, Ariz. + + _b._ _Mustela frenata nevadensis_, [M] ad., 22116, Mus. Vert. Zoöl., + Chinquapin, California. + + _c._ _Mustela frenata effera_, [M] ad., 33637, Amer. Mus. Nat. Hist., + Ironside, Oregon. + + _d._ _Mustela frenata washingtoni_, [M] ad., 226758, U. S. Nat. Mus., + Gotchen Creek, Wash. + + _e._ _Mustela frenata saturata_, [M] ad., 65930, U. S. Nat. Mus., type, + Siskiyou, Oregon. + + _f._ _Mustela frenata altifrontalis_, [M] ad., 391, Coll. Alex Walker, + Blaine, Oregon. + + _g._ _Mustela frenata oregonensis_, [M] sad., 43828/32019, U. S. N. M., + Grants Pass, Ore. + + _h._ _Mustela frenata munda_, [M] ad., 5459, M. C. Z., type, Point + Reyes, Calif. + + _i._ _Mustela frenata munda_, [M] ad., 19722, Mus. Vert. Zoöl., Point + Arena, Calif. + + _j._ _Mustela frenata xanthogenys_, [M] ad., 1440, Coll. Alex Walker, 5 + mi. W Fresno, Calif.] + + +[Illustration: PLATE 22. Photographs, retouched, of skulls, in dorsal +view, of nine subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata munda_, [M] ad., 19722, Mus. Vert. Zoöl., Point + Arena, California. + + _b._ _Mustela frenata xanthogenys_, [M] ad., 1440, col. Alex Walker, 5 + mi. W Fresno, California. + + _c._ _Mustela frenata nigriauris_, [M] ad., 487, Stanford Univ., Palo + Alto, California. + + _d._ _Mustela frenata latirostra_, [M] ad., 52702, U. S. Nat. Mus., El + Cajon, California. + + _e._ _Mustela frenata pulchra_, [M] ad., 16668, Mus. Vert. Zoöl., type, + Buttonwillow, California. + + _f._ _Mustela frenata inyoensis_, [M] ad., 25907, Mus. Vert. Zoöl., 2 + mi. N Independence, California. + + _g._ _Mustela frenata neomexicana_, [M] ad., 1485, Mus. Nat. Hist., + Univ., Kansas, Liberal, Kansas. + + _h._ _Mustela frenata texensis_, [M] ad., 14821, Amer. Mus. Nat. Hist., + Kerr County, Texas. + + _i._ _Mustela frenata frenata_, [M] ad., 50826, U. S. Nat. Mus., + Tlalpam, México, D. F.] + + +[Illustration: 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_.] + + +[Illustration: PLATE 24. Photographs, retouched, of skulls, in lateral +view, of ten subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata nigriauris_, [M] ad., 487, Stanford Univ., Palo + Alto, California. + + _b._ _Mustela frenata latirostra_, [M] ad., 52702, U. S. Nat. Mus., El + Cajon, California. + + _c._ _Mustela frenata pulchra_, [M] ad., 16668, Mus. Vert. Zoöl., type, + Buttonwillow, Calif. + + _d._ _Mustela frenata inyoensis_, [M] ad., 25907, Mus. Vert. Zoöl., + type, 2 mi. N Independence, California. + + _e._ _Mustela frenata neomexicana_, [M] ad., 1485, M. N. H., Univ. + Kansas, Liberal, Kansas. + + _f._ _Mustela frenata texensis_, [M] ad., 14821, A. M. N. H., Kerr + County, Texas. + + _g._ _Mustela frenata frenata_, [M] ad., 50826, U. S. Nat. Mus., + Tlalpam, México, D. F. + + _h._ _Mustela frenata leucoparia_, [M] ad., 125972, U. S. N. M., Los + Reyes, Michoacán. + + _i._ _Mustela frenata macrophonius_, [M] ad., [14063,] Field Mus. N. + H., type, Achotal, Veracruz. + + _j._ _Mustela frenata goldmani_, [M] ad., 77519, U. S. Nat. Mus., + Pinabete, Veracruz.] + + +[Illustration: PLATE 25. Photographs, retouched, of skulls, in dorsal +view of nine subspecies of _Mustela frenata_. Natural Size. + + _a._ _Mustela frenata leucoparia_, [M] ad., 125972, U. S. Nat. Mus., + Los Reyes, Michoacán. + + _b._ _Mustela frenata macrophonius_, [M] ad., 14063, Field Mus. Nat. + Hist., type, Achotal, Veracruz. + + _c._ _Mustela frenata goldmani_, [M] ad., 133253, U. S. Nat. Mus., 20 + mi. SE Teopisca, Chiapas. + + _d._ _Mustela frenata tropicalis_, [M] ad., 54994, U. S. Nat. Mus., + type, Jico, Veracruz. + + _e._ _Mustela frenata perda_, [M] sad., 100041, U. S. Nat. Mus., type, + Teapa, Tabasco. + + _f._ _Mustela frenata nicaraguae_, [M] sad., 30754, Amer. Mus. Nat. + Hist., type, Matagalpa, Nicaragua. + + _g._ _Mustela frenata costaricensis_, [M] ad., 3.2.1.6., British Mus. + Nat. Hist., San José, Costa Rica. + + _h._ _Mustela frenata panamensis_, [M] ad., 18848, Amer. Mus. Nat. + Hist., Boquete, Panamá. + + _i._ _Mustela frenata meridana_, [M] ad., 123341, U. S. Nat. Mus., + type, Mérida, Venezuela.] + + +[Illustration: 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._ _panamensis_; + + _i._ _meridana_.] + + +[Illustration: PLATE 27. Photographs, retouched, of skulls, in lateral +view of ten subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata tropicalis_, [M] ad., 54994, U. S. Nat. Mus., + type, Jico, Veracruz. + + _b._ _Mustela frenata perda_, [M] sad., 100041; U. S. Nat. Mus., type, + Teapa, Tabasco. + + _c._ _Mustela frenata nicaraguae_, [M] sad., 30754, Amer. Mus. Nat. + Hist., type, Matagalpa, Nicaragua. + + _d._ _Mustela frenata costaricensis_, [M] ad., 3.2.1.6., British Mus. + Nat. Hist., San José, Costa Rica. + + _e._ _Mustela frenata panamensis_, [M] ad., 18848, Amer. Mus. Nat. + Hist., Boquete, Panamá. + + _f._ _Mustela frenata meridana_, [M] ad., 24309, Amer. Mus. Nat. Hist., + Mérida, Venezuela. + + _g._ _Mustela frenata aureoventris_, [M] yg., 34677, Amer. Mus. Nat. + Hist., Gualea, Ecuador. + + _h._ _Mustela frenata helleri_, [M] ad., 24133, Field Mus. Nat. Hist., + type, Rio Chinchao, Perú. + + _i._ _Mustela frenata macrura_, [M] ad., 561, Mus. Polonais d' Hist., + Nat., type, Junín, Perú. + + _j._ _Mustela frenata agilis_, [M] ad., 8.1.10.1., British Mus. Nat. + Hist., Lima, Perú.] + + +[Illustration: PLATE 28. Photographs, retouched (except _f_), of +skulls, in dorsal view of nine kinds (species and subspecies) of +_Mustela_. Natural size. + + _a._ _Mustela frenata aureoventris_, [M] yg., 34677, Amer. Mus. Nat. + Hist., Gualea, Ecuador. + + _b._ _Mustela frenata helleri_, [M] ad., 24133, Field Mus. Nat. Hist., + type, Rio Chinchao, Perú. + + _c._ _Mustela frenata macrura_, [M] ad., 561, Mus. Polonais d' Hist. + Nat., type, Junín, Perú. + + _d._ _Mustela frenata agilis_, [M] ad., 8.1.10.1., British Mus. Nat. + Hist., Lima, Perú. + + _e._ _Mustela frenata boliviensis_, [M] ad., 72587, Amer. Mus. Nat. + Hist., type, Nequejahuira, Bolivia. + + _f._ _Mustela frenata xanthogenys_, [M] ad., 43.6.4.55., British Mus., + Nat. Hist., type, California. + + _g._ _Mustela frenata costaricensis_, [M] yg., 37149, U. S. Nat. Mus., + type, San José, Costa Rica. + + _h._ _Mustela frenata panamensis_, [M] yg., 178970, U. S. Nat. Mus., + Mt. Pirre, Panamá. + + _i._ _Mustela africana africana_, [M] yg., 37475, Amer. Mus. Nat. + Hist., Pará, Brazil.] + + +[Illustration: PLATE 29. Photographs, retouched, of skulls, in ventral +view, of nine kinds (species and subspecies) of _Mustela_. Natural +size. + + _a._ _Mustela frenata aureoventris_, [M] yg., 34677, Amer. Mus. Nat. + Hist., Gualea, Ecuador. + + _b._ _Mustela frenata helleri_, [M] ad., 24133, Field Mus. Nat. Hist., + type, Rio Chinchao, Perú. + + _c._ _Mustela frenata macrura_, [M] ad., 561, Mus. Polonais d' Hist. + Nat., type, Junín, Perú. + + _d._ _Mustela frenata agilis_, [M] ad., 8.1.10.1., British Mus. Nat. + Hist., Lima, Perú. + + _e._ _Mustela frenata boliviensis_, [M] ad., 72587, Amer. Mus. Nat. + Hist., type, Nequejahuira, Bolivia. + + _f._ _Mustela frenata leucoparia_, [M] ad., 47179/34914, U. S. Nat. + Mus., type, Pátzcuaro, Michoacán. + + _g._ _Mustela frenata costaricensis_, [M] yg., 37149, U. S. Nat. Mus., + type, San José, Costa Rica. + + _h._ _Mustela frenata panamensis_, [M] yg., 178970, U. S. Nat. Mus., + Mt. Pirre, Panamá. + + _i._ _Mustela africana africana_, [M] yg., 37475, Amer. Mus. Nat. + Hist., Pará, Brazil.] + + +[Illustration: PLATE 30. Photographs, retouched (except _e_ and _q_) +of skulls and lower jaws of _Mustela_. + + _a._ _Mustela frenata boliviensis_, [M] ad., 72587, Amer. Mus. Nat. + Hist., type, Nequejahuira, Bolivia. + + _b._ _Mustela frenata xanthogenys_, [M] ad., 43.4.6.55, British Mus. + Nat. Hist., type, California. + + _c._ _Mustela frenata costaricensis_, [M] yg., 37149, U. S. Nat. Mus., + type, San José, Costa Rica. + + _d._ _Mustela frenata panamensis_, [M] yg., 178970, U. S. Nat. Mus., + Mt. Pirre, Panamá. + + _e._ _Mustela frenata affinis_, [M] ad., 54.6.3.4, British Mus. Nat. + Hist., type, New Granada [=Colombia]. + + _f._ _Mustela africana africana_, [M] yg., 37475, Amer. Mus. Nat. + Hist., Pará, Brazil. + + _g._ _Mustela frenata saturata_, [M] ad., 65930, U. S. Nat. Mus., type, + Siskiyou, Oregon. + + _h._ _Mustela frenata oregonensis_, [M] ad., 43828/32019, U. S. Nat. + Mus., type, Grants Pass, Oregon. + + _i._ _Mustela frenata munda_, [M] ad., 5459, Mus. Comp. Zool., type, + Point Reyes, California. + + _j._ _Mustela frenata leucoparia_, [M] ad., 47179/34914, U. S. Nat. + Mus., type, Pátzcuaro, Michoacán. + + _k._ _Mustela frenata macrophonius_, [M] ad., 14963, Field Mus. Nat. + Hist., type, Achotal, Veracruz. + + _l._ _Mustela frenata goldmani_, [M] ad., 77519, U. S. Nat. Mus., type, + Pinabete, Chiapas. + + _m._ _Mustela frenata tropicalis_, [M] ad., 54994, U. S. Nat. Mus., + type, Jico, Veracruz. + + _n._ _Mustela frenata perda_, [M] sad., 100041, U. S. Nat. Mus., type, + Teapa, Tabasco. + + _o._ _Mustela frenata nicaraguae_, [M] sad., 30754, Amer. Mus. Nat. + Hist., type, Matagalpa, Nicaragua. + + _p._ _Mustela frenata costaricensis_, [M] yg., 37149, U. S. Nat. Mus., + type, San José, Costa Rica. + + _q._ _Mustela frenata affinis_, [M] ad., 54.6.3.4, British Mus. Nat. + Hist., type, New Granada [=Colombia]. + + _r._ _Mustela frenata macrura_, [M] ad., 561., Mus. Polonais d' Hist. + Nat., type, Junín, Perú.] + + +[Illustration: PLATE 31. Photos, retouched, of skulls of 9 subspecies +of _M. frenata_, x 1. + + _a._ _Mustela frenata noveboracensis_, [F] ad., 64687, U. S. N. M., + Wilmington, Mass. + + _b._ _Mustela frenata occisor_, [F] ad., 9101, Mus. Comp. Zool., + Bucksport, Maine. + + _c._ _Mustela frenata primulina_, [F] ad., 3638, U. K. M. N. H., 7 mi. + SW Lawrence, Kans. + + _d._ _Mustela frenata olivacea_, [F] ad., 41024, Mus. Vert. Zoöl., + Sinkola Plantation, Ga. + + _e._ _Mustela frenata spadix_, [F] ad., 188410, U. S. Nat. Mus., Elk + River, Minn. + + _f._ _Mustela frenata longicauda_, [F] ad., 75483, U. S. Nat. Mus., + Wingard, Sask. + + _g._ _Mustela frenata oribasus_, [F] ad., 9058, M. C. Z., type, source + of Kettle River, B. C. + + _h._ _Mustela frenata alleni_, [F]?, ad., 7441, A. M. N. H., Black + Hills, S. D. + + _i._ _Mustela frenata arizonensis_, [F] ad., 1886, A. M. N. H., type, + S. F. Forest, Ariz.] + + +[Illustration: PLATE 32. Photos, retouched, of skulls of 9 subspecies +of _M. frenata_, × 1. + + _a._ _Mustela frenata noveboracensis_, [F] ad., 64687, U. S. N. M., + Wilmington, Mass. + + _b._ _Mustela frenata occisor_, [F] ad., 9101, Mus. Comp. Zool., + Bucksport, Maine. + + _c._ _Mustela frenata primulina_, [F] ad., 3638, U. K. M. N. H., 7 mi. + SW Lawrence, Kans. + + _d._ _Mustela frenata olivacea_, [F] ad., 41024, Mus. Vert. Zoöl., + Sinkola Plantation, Ga. + + _e._ _Mustela frenata spadix_, [F] ad., 188410, U. S. Nat. Mus., Elk + River, Minn. + + _f._ _Mustela f. longicauda_, [F] ad., 75483, U. S. Nat. Mus., Wingard, + Sask. + + _g._ _Mustela frenata oribasus_, [F] ad., 9058, M. C. Z., type, source + of Kettle Riv., B. C. + + _h._ _Mustela frenata alleni_, [F]?, ad., 7441, A. M. N. H., Black + Hills, S. D. + + _i._ _Mustela frenata arizonensis_, [F] ad., 1886, A. M. N. H., type, + S. F. Forest, Ariz.] + + +[Illustration: PLATE 33. Photographs, retouched, of skulls in lateral +view of ten subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata noveboracensis_, [F] ad., 64687, U. S. N. M., + Wilmington, Mass. + + _b._ _Mustela frenata occisor_, [F] ad., 9101, Mus. Comp. Zool., + Bucksport, Maine. + + _c._ _Mustela frenata primulina_, [F] ad., 3638, Univ. Kansas Mus. Nat. + Hist., 7 mi. SW Lawrence, Kansas. + + _d._ _Mustela frenata olivacea_, [F] ad., 41024, Mus. Vert. Zoöl., + Sinkola Plantation, Ga. + + _e._ _Mustela frenata spadix_, [F] ad., 188410 (2196), U. S. Nat. Mus., + Elk River, Minn. + + _f._ _Mustela frenata longicauda_, [F] ad., 75483, U. S. Nat. Mus., + Wingard, Sask. + + _g._ _Mustela frenata oribasus_, [F] ad., 9058, Mus. Comp. Zool., type, + source of Kettle River, British Columbia. + + _h._ _Mustela frenata alleni_, [F]?, ad., 7441, Amer. Mus. N. H., Black + Hills, S. D. + + _i._ _Mustela frenata arizonensis_, [F] ad., 1886, Amer. Mus. Nat. + Hist., type, San Francisco, Forest, Arizona. + + _j._ _Mustela frenata nevadensis_, [F] ad., 41503, M. V. Z., type, 3 + mi. E Baker, Nev.] + + +[Illustration: PLATE 34. Photographs, retouched, of skulls in dorsal +view, of eight subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata nevadensis_, [F] ad., 41503, M. V. Z., type, 3 + mi. E Baker, Nevada. + + _b._ _Mustela frenata washingtoni_, [F] sad., 81953, U. S. N. M., Trout + Lake, Wash. + + _c._ _Mustela frenata altifrontalis_, [F] ad., 392, coll. of Alex + Walker, Blaine, Oregon. + + _d._ _Mustela frenata oregonensis_, [F] ad., 244520, U. S. Nat. Mus., + Medford, Oregon. + + _e._ _Mustela frenata munda_, [F] ad., 91764, U. S. Nat. Mus., Point + Reyes, California. + + _f._ _Mustela frenata munda_, [F] ad., 19723, Mus. Vert. Zoöl., Point + Arena, California. + + _g._ _Mustela frenata xanthogenys_, [F] ad., 2626, coll. of W. E. + Snyder, Selma, California. + + _h._ _Mustela frenata nigriauris_, [F] ad., 3761, M. V. Z., San + Francisco, California. + + _i._ _Mustela frenata neomexicana_, [F] ad., 36482, U. S. N. M., + Tombstone, Arizona.] + + +[Illustration: PLATE 35. Photographs, retouched, of skulls in ventral +view, of eight subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata nevadensis_, [F] ad., 41503, M. V. Z., type, 3 + mi. E Baker, Nevada. + + _b._ _Mustela frenata washingtoni_, [F] sad., 81953, U. S. N. M., Trout + Lake, Wash. + + _c._ _Mustela frenata altifrontalis_, [F] ad., 392, coll. of Alex + Walker, Blaine, Oregon. + + _d._ _Mustela frenata oregonensis_, [F] ad., 244520, U. S. Nat. Mus., + Medford, Oregon. + + _e._ _Mustela frenata munda_, [F] ad., 91764, U. S. N. M., Point Reyes, + California. + + _f._ _Mustela frenata munda_, [F] ad., 19723, M. V. Z., Point Arena, + California. + + _g._ _Mustela frenata xanthogenys_, [F] ad., 2626, coll. of W. E. + Snyder, Selma, Calif. + + _h._ _Mustela frenata nigriauris_, [F] ad., 3761, M. V. Z., San + Francisco, California. + + _i._ _Mustela frenata neomexicana_, [F] ad., 36482, U. S. N. M., + Tombstone, Arizona.] + + +[Illustration: PLATE 36. Photographs, retouched, of skulls in lateral +view, of eight subspecies of _Mustela frenata_. Natural size. + + _a._ _Mustela frenata washingtoni_, [F] sad., 81953, U. S. N. M., Trout + Lake, Wash. + + _b._ _Mustela frenata altifrontalis_, [F] ad., 392, coll. of Alex + Walker, Blaine, Oregon. + + _c._ _Mustela frenata oregonensis_, [F] ad., 244520, U. S. N. M., + Medford, Oregon. + + _d._ _Mustela frenata munda_, [F] ad., 91764, U. S. N. M., Point Reyes, + California. + + _e._ _Mustela frenata munda_, [F] ad., 19723, Mus. Vert. Zoöl., Point + Arena, Calif. + + _f._ _Mustela frenata xanthogenys_, [F] ad., 2626, coll. of W. E. + Snyder, Selma, Calif. + + _g._ _Mustela frenata nigriauris_, [F] ad., 3761, Mus. Vert. Zoöl., San + Francisco, Calif. + + _h._ _Mustela frenata neomexicana_, [F] ad., 36482, U. S. N. M., + Tombstone, Ariz. + + _i._ _Mustela frenata frenata_, [F] ad., 58685, U. S. Nat. Mus., + Brownsville, Texas. + + _j._ _Mustela frenata frenata_, [F] ad., 991, Berlin Zool. Mus., type, + México City, D. F. + + _k._ _Mustela frenata leucoparia_, [F] ad., 26153, Amer. Mus. N. H., + Artenkiki, Jalisco. + + _l._ _Mustela frenata perotae_, [F] ad., 54278, U. S. Nat. Mus., type, + 12500 ft., Cofre de Perote, Veracruz.] + + +[Illustration: PLATE 37. Photos, retouched, of skulls of 8 subspecies +of _Mustela frenata_, × 1. + + _a._ _Mustela frenata frenata_, [F] ad., 58685, U. S. Nat. Mus., + Brownsville, Texas. + + _b._ _Mustela frenata frenata_, [F] ad., 991, Berlin Zool. Mus., type, + México City, D. F. + + _c._ _Mustela frenata leucoparia_, [F] ad., 26153, Amer. Mus. Nat. + Hist., Artenkiki, Jalisco. + + _d._ _Mustela f. perotae_, [F] ad., 54278, U. S. N. M., type, Cofre de + Perote, Veracruz. + + _e._ _Mustela frenata macrophonius_, [F] ad., 132528, U. S. Nat. Mus., + Pérez, Veracruz. + + _f._ _Mustela frenata tropicalis_, [F] ad., 54993, U. S. Nat. Mus., + Jico, Veracruz. + + _g._ _Mustela frenata perda_, [F] sad., 65422, U. S. Nat. Mus., + Catemaco, Veracruz. + + _h._ _Mustela frenata meridana_, [F] ad., 143665, U. S. N. M., Mérida, + Venezuela. + + _i._ _Mustela f. macrura_, [F] ad., 564, M. P. H. N., type of Mustela + jelskii, Cutervo, Perú.] + + +[Illustration: PLATE 38. Photos, retouched, skulls in ventral view, 8 +subspecies of _M. frenata_, × 1. + + _a._ _Mustela frenata frenata_, [F] ad., 58685, U. S. Nat. Mus., + Brownsville, Texas. + + _b._ _Mustela frenata frenata_, [F] ad., 991, Berlin Zool. Mus., type, + México City, D. F. + + _c._ _Mustela frenata leucoparia_, [F] ad., 26153, Amer. Mus. Nat. + Hist., Artenkiki, Jalisco. + + _d._ _Mustela f. perotae_, [F] ad., 54278, U. S. N. M., type, Cofre de + Perote, Veracruz. + + _e._ _Mustela frenata macrophonius_, [F] ad., 132528, U. S. Nat. Mus., + Pérez, Veracruz. + + _f._ _Mustela frenata tropicalis_, [F] ad., 54993, U. S. Nat. Mus., + Jico, Veracruz. + + _g._ _Mustela frenata perda_, [F] sad., 65422, U. S. Nat. Mus., + Catemaco, Veracruz. + + _h._ _Mustela frenata meridana_, [F] ad., 143665, U. S. N. M., Mérida, + Venezuela. + + _i._ _Mustela f. macrura_, [F] ad., 564, M. P. H. N., type of _Mustela + jelskii_, Cutervo, Perú.] + + +[Illustration: PLATE 39. FIGS. _a_-_h_. Photos, retouched, of 10 kinds +_Mustela_, × 1. + + _a._ _Mustela frenata macrophonius_, [F] ad., 132528, U. S. Nat. Mus., + Pérez, Veracruz. + + _b._ _Mustela frenata tropicalis_, [F] ad., 54993, U. S. Nat. Mus., + Jico, Veracruz. + + _c._ _Mustela frenata perda_, [F] sad., 65422, U. S. Nat. Mus., + Catemaco, Veracruz. + + _d._ _Mustela frenata meridana_, [F] ad., 143665, U. S. Nat. Mus., + Mérida, Venezuela. + + _e._ _Mustela f. macrura_, [F] ad., 564, M. P. H. N., type of _Mustela + jelskii_, Cutervo, Perú. + + _f._ _Mustela frenata agilis_, [F] sad., 21147, Field Mus. Nat. Hist., + Macate, Perú. + + _g._ _Mustela frenata gracilis_, [F]?, ad., 12431, Amer. Mus. Nat. + Hist., type, Conard Fissure, Arkansas, Pleistocene. + + _h._ _Mustela a. stolzmanni_, [F] sad., 24.12.12.24, Brit. M. N. H., + Myobamba, Perú. + + FIGS. _i_-_l_. _Mustela frenata nevadensis_, all males, from Colorado, + showing seasonal change in color, × 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. + + _j._ No. 202741 U. S. Nat. Mus., Pierce Place, April 18, 1913. + + _k._ No. 201681 U. S. Nat. Mus., Jefferson, June 23, 1913. + + _l._ No. 41997 Amer. Mus. Nat. Hist., Navajo River, October 29, 1913.] + + +[Illustration: PLATE 40. Photos, retouched, of skulls and lower jaws of +_Mustela_, × 1. + + _a._ _Mustela frenata agilis_, [F] sad., 21147, Field Mus. Nat. Hist., + Macate, Perú. + + _b._ _Mustela frenata gracilis_, [F]?, ad., type, 12431, A. M. N. H., + Conard Fiss., Ark. + + _c._ _Mustela africana stolzmanni_, [F] sad., 24.12.12.24, Brit. M. N. + H., Myobamba, Perú. + + _d._ _Mustela frenata agilis_, [F] sad., 21147, Field Mus. Nat. Hist., + Macate, Perú. + + _e._ _Mustela frenata gracilis_, [F]?, ad., type, 12431, A. M. N. H., + Conard Fiss., Ark. + + _f._ _Mustela africana stolzmanni_, [F] sad., 24.12.12.24, Brit. M. N. + H., Myobamba, Perú. + + _g._ _Mustela africana stolzmanni_, [F] ad., 563, Mus. Polonais d' + Hist. Nat., type, Yurimaguas, Perú. The palate is broken longitudinally + and the two maxillae are slightly out of normal position. + + _h._ _Mustela frenata oribasus_, [F] ad., 9058, M. C. Z., type, source + of Kettle Riv., B. C. + + _i._ _Mustela frenata munda_, [F] ad., 91764, U. S. Nat. Mus., Point + Reyes, Calif. + + _j._ _Mustela frenata frenata_, [F] ad., 991, Berlin Z. M., type, + México City, D. F. + + _k._ _Mustela f. macrura_, [F] ad., 564, Mus. Polonais d' Hist. Nat., + type of _Mustela jelskii_. + + _l._ _Mustela africana stolzmanni_, [F] ad., 563, Mus. Polonais d' + Hist. Nat., type, Yurimaguas, Perú. Right half of lower jaw reversed.] + + +[Illustration: PLATE 41. Photographs, approximately 1/2, 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, [M] 131256 and [F] 131245. + + _Mustela erminea invicta_, both in U. S. B. S., from Tungsten Mine, + Washington, [M] 235236 and [F] 235235. + + _Mustela erminea muricus_, both in M. V. Z., from Baker Creek, 8675 + ft., Nevada, [M] 41501 and [F] 41502. + + _Mustela frenata nigriauris_, both in M. V. Z., from California, [M] + 51666 from Concord and [F] 73109 from Berkeley. + + _Mustela africana africana_, [M] 37475 A. M. N. H., from Pará, Brazil. + + _Mustela rixosa allegheniensis_, both in M. Z. U. M., from Michigan, + [M] 83260 Swan Creek Farm and [F] 88079 from Unadilla.] + + + + +INDEX TO VOLUME 4 + +Principal references are in boldface type + + + aequatorialis, + 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 + + alleni, + Mustela, 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 + + armatus, Citellus, 200 + + arthuri, 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, 205 + lateralis, 206 + richardsonii, 205 + townsendii, 205 + tridecemlineatus, 205 + + Clethrionomys, 92 + gapperi, 175, 181 + + Colaptes, 213 + + comadreja, 7 + + Condylura, 210 + + costaricensis, Mustela, 78, =372=, 379, 387 + + cotton rat, 208 + + cottontail, 202, 203, 205, 212 + Mearns, 208 + + coyote, 200 + + Cratogeomys, 65, 207 + + culbertsoni, + Mustela, 75 + Putorius, 75 + + + deer mouse, 208, 209, 216 + + domestic cat, 174 + + donnola, 7 + + drummondi, Microtus, 175 + + + earthworm, 93 + + effera, Mustela, 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 + + Grammogale, 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 + + letifera, 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 + + + macrodon, Mustela, 82 + + macrophonius, + Mustela, 78, 360 + Putorius, 78, 360 + + macrura, Mustela, 75, 387, 393, 398, 402 + + macrurus, Putorius, 379 + + maniculatus, Peromyscus, 175 + + Martinogale, 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 + anguinae, 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, 398 + 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 + lutensis, 82 + lutreola, 170 + macrodon, 82 + macrophonius, 78, 360 + 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= + stolzmanni, 75, 409, =413= + streatori, 76, =155= + texensis, 79, 338 + tropicalis, 76, =363=, 367 + vison, =82= + vulgivaga, 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= + + nigriauris, Mustela, 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, + barn, 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 + + Peromyscus, 91, 92, 173, 196, 208, 209, 210, 216 + maniculatus, 175 + + perotae, Mustela, 79, =351= + + pheasant, 209 + ring-necked, 202 + + pigmy weasel, 216 + + pika, 92 + + pikas, 201 + + pipiens, Rana, 93 + + Pisces, 93 + + Pituophis sayi sayi, 213 + + Pliogale, 11 + + pocket gopher, 209, 216 + Mexican, 207 + northern, 206 + Shaw, 208 + + polaris, + Mustela, 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, 161 + 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 + neomexicanus, 76, 333 + nigripes, 74 + notius, 77, 222 + noveboracensis, 74, 222, 232 + numidicus, 170 + occisor, 77, 230 + oregonensis, 304 + oribasus, 77, 270 + paraensis, 76, 409 + peninsulae, 75, 250 + perdus, 77, 366 + polaris, 77, 103 + pusillus, 118, 155, 184, 190 + richardsonii, 96, 110 + rixosus, 76, 181, 184 + saturatus, 76, 294, 300 + spadix, 76, 252 + stolzmanni, 145, 148, 155, 159 + streatori, 76 + tropicalis, 76, 363, 370 + vulgaris, 118, 155, 181, 184 + washingtoni, 76, 294 + xanthogenys, 315, 319, 323, 331 + + putorius, Mustela, 43 + + + quail, 209, 213 + + + rabbit, + brush, 213 + snowshoe, 93, 201 + + racer, red, 213 + + Rana pipiens, 93 + + rat, + brown, 202 + cotton, 208, 216 + wood, 208 + + Rattus, 91, 92, 210, 216 + + rattlesnake, 200 + + red-backed mouse, 92 + + red + racer, 213 + squirrel, 206 + + Reithrodontomys, 175, 216 + + reptiles, 213 + + Richardson ground squirrel, 205 + + richardsonii, + Citellus, 205 + Mustela, 73, 110 + Putorius, 222 + + ring-necked pheasant, 202 + + rixosa, Mustela, 76, 153, 155, 161, 168, 184 + + rixosus, Putorius, 76, 181, 184 + + Rodentia, 216 + + russet-backed thrush, 204 + + + salva, Mustela, 80, 135 + + saturata, Mustela, 76, 297 + + sayi, Pituophis, 213 + + Scalopus, 209 + + Scapanus latimanus, 205 + + Schönthierlein, 7 + + Sciurus, 210 + + seclusa, Mustela, 80, 141 + + semplei, Mustela, 78, 105 + + short-tailed shrew, 208, 209 + + short-tailed weasel, 87 + + shrews, 216 + + Sigmodon, 216 + + Sigmodon hispidus, 208 + + slate-colored junco, 213 + + snake, + bull, 213 + king, 213 + + snakes, 216 + + snowshoe rabbit, 93, 210 + + snowy owl, 90 + + song sparrow, 201, 213 + + Sorex, 91, 209 + cinereus, 205 + + Soricidae, 216 + + spadix, + Mustela, 76, 252 + Putorius, 76, 252 + + sparrow, + song, 201, 213 + tree, 208, 209 + + spermophile, Uinta, 200 + + squirrel, + flying, 206 + red, 206 + tree, 216 + + stolzmanni, Mustela, 75, 409, 413 + + streatori, + Mustela, 76, 155 + Putorius, 76, 145, 148, 155, 159 + + striatus, Tamias, 206 + + Sylvilagus, 91, 93, 209, 210, 216 + bachmani, 213 + + + Talpidae, 216 + + talpoides, Thomomys, 207 + + Tamias, 91, 92, 206, 210, 216 + minimus, 206 + striatus, 206 + + Tamiasciurus, 206, 216 + + texensis, Mustela, 79, 338 + + thirteen-lined ground squirrel, 205 + + Thomomys, 206, 219 + bottae, 207 + talpoides, 207 + + thrush, russet-backed, 204 + + towhee, 201 + + Townsend ground squirrel, 205 + + townsendii, Citellus, 205 + + tree sparrow, 208, 209 + + tridecemlineatus, Citellus, 205 + + tropical weasel, 406 + + tropicalis, + Mustela, 76, 363, 367 + Putorius, 76, 363, 370 + + + Uinta spermophile, 200 + + + varying hare, 210, 216 + + vison, + Mustela, 82 + Lutreola, 84 + + vulgaris, + Mustela, 73 + Putorius, 118, 155, 181, 184 + + vulgivaga, Mustela, 82 + + Vulpes fulvus, 200 + + + washingtoni, + Mustela, 76, 294 + Putorius, 76, 294 + + weasel, + least, 168, 209 + long-tailed, 193 + Pigmy, 216 + short-tailed, 87 + tropical, 406 + + wild birds, 93, 213 + + wood rat, 208, 210, 216, 219 + + + xanthogenys, + Mustela, 74, 315, 331 + Putorius, 315, 319, 323, 331 + + + Zapus, 216 + minor, 210 + + +FOOTNOTES: + +[1] Acad. Nat. Sciences of Philadelphia + +[2] American Mus. Nat. History + +[3] Baylor University + +[4] Berlin Zoological Museum + +[5] Boston Society of Natural History + +[6] Brigham Young University + +[7] British Museum of Natural History + +[8] California Academy of Sciences + +[9] Carnegie Museum + +[10] Charles R. Conner Museum + +[11] Charleston Museum + +[12] Coe College + +[13] Collection of A. H. Miller + +[14] Collection of Alex Walker + +[15] Collection of Arthur Peake + +[16] Collection of D. D. Stone + +[17] Collection of Donald V. Hemphill + +[18] Collection of E. J. Koestner + +[19] Collection of Edward R. Warren + +[20] Collection of Frank Stephens + +[21] Collection of Ian McTaggart-Cowan + +[22] Collection of J. A. Munro + +[23] Collection of J. Arnold + +[24] Collection of J. E. Law + +[25] Collection of J. M. Edson + +[26] Collection of Jack C vonBloeker + +[27] Collection of Joe and Dean Thiriot + +[28] Collection of John Cushing + +[29] Collection of John Fitzgerald, Jr. + +[30] Collection of John Tyler + +[31] Collection of Kenneth Racey + +[32] Collection of L. M. Huey + +[33] Collection of Lloye H. Miller + +[34] Collection of Mr. Green + +[35] Collection of Myron H. Swenk + +[36] Collection of O. J. Murie + +[37] Collection of O. P. Silliman + +[38] Collection of R. D. Moore + +[39] Collection of R. H. Coleman + +[40] Collection of R. W. Jackson + +[41] Collection of Ralph Ellis + +[42] Collection of Robert T. Orr + +[43] Collection of Rollin H. Baker + +[44] Collection of Ross Hardy + +[45] Collection of Stanley C. Arthur + +[46] Collection of Stanley G. Jewett + +[47] Collection of Stuart Criddle + +[48] Collection of T. C. Stephens + +[49] Collection of Victor B. Scheffer + +[50] Collection of W. E. Snyder + +[51] Collection of Walter W. Dalquest + +[52] Collection of William B. Davis + +[53] Collection of William B. Richardson + +[54] Collection of William Bebb + +[55] Collection of William T. Shaw + +[56] Collection Rocky Mt. Spotted Fever Lab. + +[57] Colorado Museum of Natural History + +[58] Cornell University + +[59] Donald R. Dickey Collection + +[60] Field Museum of Natural History + +[61] Florida State Museum + +[62] Fresno State Junior College + +[63] Humboldt State Teachers College + +[64] Illinois Natural History Survey + +[65] Iowa State College + +[66] Iowa Wesleyan College + +[67] Kansas State Agric. College + +[68] Leland Stanford Junior University + +[70] Los Angeles Mus. Hist. Art and Sci. + +[71] Louisiana State University + +[72] Mt. Rainier Nat'l Park Collection + +[73] Mus. Polonais d'Hist. Nat., Warsaw + +[74] Mus. Vert. Zoöl., Univ. California + +[75] Museum of Comparative Zoölogy + +[76] Museum of Zoölogy, Univ. Michigan + +[77] National Museum of Canada + +[78] Naturhistoriska Ricksmuseum, Sweden + +[80] New York State Museum + +[81] Ohio State Museum + +[82] Oklahoma Agric. and Mech. College + +[83] Ottawa University, Kansas + +[84] Paris Museum + +[85] Provincial Museum of British Columbia + +[86] Royal Ontario Museum of Zoölogy + +[87] San Diego Society of Natural History + +[88] State Hist. and Nat. Hist. Soc. Colo. + +[89] State Normal School, Cheney, Wash. + +[90] Texas Cooperative Research Collection + +[91] United States National Museum + +[92] Univ. California Mus. Palaeo. + +[93] Univ. Kansas Mus. Nat. History + +[94] Univ. Washington Museum of Zoölogy + +[95] Univ. Zool. Mus., Copenhagen + +[96] University of Arkansas + +[97] University of Idaho + +[98] University of Minnesota + +[99] University of Notre Dame + +[100] University of Oklahoma + +[101] University of Oregon + +[102] University of South Dakota + +[103] University of Utah + +[104] University of Wisconsin + + * * * * * + + Transcribers Notes: + + Punctuation and spelling were made consistent when a predominant + preference was found in this book; otherwise they were not changed. + + Simple typographical and spelling errors were corrected. + + P. 162 changed Tahoma Creek, [72] to Tahoma Creek, 1[72]. + + P. 418 moved last two columns of "TABLE OF CRANIAL MEASUREMENTS" + part a to the begining of part b. + + Plate 24 added [14063,] for missing specimen number. + + + + + +End of the Project Gutenberg EBook of American Weasels, by E. Raymond Hall + +*** END OF THE PROJECT GUTENBERG EBOOK 43272 *** |