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. ♀ 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.
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
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
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.