Part 31

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_.