_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.

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