American Weasels

CHECK-LIST OF AMERICAN SPECIES AND SUBSPECIES OF THE GENUS MUSTELA

ARTIFICIAL KEY TO AMERICAN SPECIES OF THE GENUS MUSTELA

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:

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.