Part 13

Female: Three adults from Great Slave Lake (Willow River, Fairchild Point, and Fort Resolution) yield average and respective measurements as follows: Total length, 252 (237, 238, 282); length of tail, 69 (63, 60, 85); length of hind foot, 32 (31, 32, 34). Corresponding, average measurements for three adults from Glacier Lake are 240, 60, 32 and for 3 adults from the Athabasca Delta, 243, 65, 30. Weight of 8 subadults from the Belcher Islands is 69 (64-78) grams. Weight of adults would be more.

_Color._--Winter pelage all white except tip of tail. Summer pelage with upper parts uniform in color and darker (16_n_) than Raw Umber, and about tones 3 to 4 of Chocolate of Oberthür and Dauthenay, pl. 343. Underparts Sulphur Yellow, Colonial Buff, or Primrose Yellow, often nearly white on chin and insides of forelegs; color of underparts extends narrowly over upper lips, distally on posterior sides of forelegs onto antipalmar faces of toes and sometimes over most of antipalmar surfaces of forefeet, on medial sides of hind legs to a point between knee and ankle but reappears on antiplantar faces of toes and in some individuals is narrowly continuous onto toes. Least width of color of underparts averaging, in a series of 12 males from the Athabasca Lake Region, 40 (25-54) per cent of greatest width of color of upper parts. Black tip of tail averaging 56 (45-63) mm. in 5 adult males from same region and thus 60 (48-70) per cent of length of tail-vertebrae.

From _arctica_, _polaris_, _semplei_ and _kadiacensis_, _richardsonii_ differs in: Color darker; ventral side of tail same color as upper parts; light-colored underparts a fifth narrower; black tip of tail by actual measurement a fifth shorter and averaging less than two-thirds rather than more than four-fifths of length of tail-vertebrae. From _cicognanii_, _richardsonii_ differs in that the underparts are a fourth wider and in some specimens more brightly colored. The width of the underparts is likewise a fourth more than in _bangsi_. In _invicta_ the underparts are not so brightly colored as in some specimens of _richardsonii_. From _fallenda_, _richardsonii_ differs in that the upper parts often are lighter colored, upper lips white rather than colored like upper parts, and underparts as wide again. In comparison with _alascensis_, the black tip of the tail averages three-fifths rather than a half of length of tail-vertebrae.

_Skull._--Male (based on 6 adults from 3 miles south of Big Island, Great Slave Lake): See measurements and plates 2-4; weight, 2.5 (2.1-2.9) grams; basilar length, 40.9 (39.6-43.7); length of tooth-rows less than length of tympanic bulla; breadth of rostrum measured across lacrimal processes less than a third of basilar length; interorbital breadth less than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth less than distance between last upper molar and jugular foramen.

Female (based on 4 adults: from Willow River, 1; Fort Resolution, 1; Athabasca Delta, 2; and 2 subadults, one from 3 mi. S Big Island and one from 15 mi. above Smith Landing): See measurements and plates 9-11; weight, 1.1 (0.9-1.4) grams; basilar length, 33.1 (31.5-34.2); length of tooth-rows less than length of tympanic bulla; breadth of rostrum less than 30 per cent of basilar length; interorbital breadth less than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth less than distance between last upper molar and jugular foramen.

The skull of the female averages 56 per cent lighter than that of the male.

Comparison of the skull with that of _arctica_, _polaris_, _semplei_, _kadiacensis_, _haidarum_, _cicognanii_, _bangsi_, _invicta_, _fallenda_, and _alascensis_ is made in the accounts of those subspecies.

_Remarks._--_M. e. richardsonii_ has the most extensive geographic range of any American race of _erminea_, is centrally located with respect to the other races, is more abundantly represented by study specimens in zoölogical collections than any other race, and is a sort of average for the species as a whole in most structural features. Therefore _richardsonii_ is used as a standard of comparison and accordingly is more fully described than any one of the other races each of which by reference to _richardsonii_ is described in comparative fashion. This comparative description has the virtue of more clearly indicating differences between subspecies and also makes for brevity.

John Richardson, Bernard R. Ross, and names of their companions, as written on the labels of the older specimens recall to the student's mind early explorations of the north country. Edward A. Preble obtained important specimens at several places and in recent years J. Kenneth Doutt and G. G. Goodwin have made the reviser's work easier by preparing specimens in series from areas not previously well represented.

The nomenclatural history of this subspecies begins with references in the literature that identify the animal as the Old World species, _Mustela erminea_--an identification which the study here reported upon shows to have been correct in the specific, although not in the subspecific, sense. Richardson, for example, in his "Fauna Boreali-Americana" published in 1829 so identified the animal. In 1838, Bonaparte, basing his description on Richardson's account of 1829, proposed the new name _richardsonii_. Richardson himself, the following year in the "Zoology of Beechey's Voyage," accepted Bonaparte's name and it has been applied to the animal in the central part of the northern timber-belt of North America ever since, except as authors used the name _Mustela erminea_ in the belief that _richardsonii_ was not distinct from _erminea_.

The north and south boundaries of the range assigned to _richardsonii_ varied according to the notions of the particular writer who was employing the name. Until Merriam in 1896 named _arctica_ as distinct, animals from the far north were generally included under the name _richardsonii_ along with populations to which the latter name now is applied. Because _richardsonii_ grades gradually into the smaller _cicognanii_ of more southern occurrence the boundary between the two has been set farther north by one writer and farther south by another, depending probably upon what the writer felt was the halfway point in size. This point of course depended upon the samples selected as typical of _richardsonii_ on the north and _cicognanii_ on the south. Because Bangs, in 1896, took as representative of _richardsonii_ the far northern and hence large-sized animals (now separated as _M. e. arctica_), his halfway point in size between them and the small _cicognanii_ of New England naturally fell farther north than it would have had he used as representative of _richardsonii_ specimens from places south of the range of _arctica_.

In 1903 J. A. Allen proposed the name _Putorius microtis_ for a specimen from Shesley, northwestern British Columbia, a place approximately 50 miles northwest of Telegraph Creek. Considering the great disparity in size between this one specimen and the other larger specimens of normal size, from the general region, available to Allen at that time, it is not surprising that he thought two full species were represented. In 1943 when G. G. Goodwin called to my attention two males, as small as the type of _microtis_ and taken by him approximately 300 miles east of Shesley, in the valley between the Musqwa and Prophet rivers, I for a second time examined all available specimens and data with the possibility in mind that _microtis_ was a species or subspecies distinct from _M. e. richardsonii_, but again concluded that only one subspecies was involved because no character except size was found to distinguish the large from the small individuals of a given sex and there are, preserved from northern British Columbia, individuals of intermediate size. _Putorius microtis_ Allen seems to have been based on an individual of _M. e. richardsonii_ near the lower limit of size for that subspecies and _microtis_ is regarded as a synonym.

Barrett-Hamilton in 1904 named the animal at "Fort Simpson, British Columbia" _Putorius arcticus imperii_. Preble (1908:232) pointed out that Fort Simpson on the Mackenzie undoubtedly was the place intended, and arranged _imperii_ as a synonym of _M. e. richardsonii_. The type specimen of _imperii_ was stated to have been received from B. M. Ross who is known to have collected specimens, including specimens of this species (now in U. S. Nat. Mus.), at Fort Simpson on the Mackenzie. I know of no Fort Simpson in British Columbia. If, as seems improbable, Port Simpson, British Columbia, was the place that Barrett-Hamilton intended to designate (where so far as I know Ross did not collect), the name _imperii_ still would seem to be a synonym of _richardsonii_ because _richardsonii_ seems to be the race of weasel at Port Simpson. In proposing the name _Putorius arcticus imperii_, Barrett-Hamilton stressed that the weasel, which he was naming, was a subspecies of _P. arcticus_, gave characters which applied perfectly to _richardsonii_ but made no reference to _richardsonii_. Barrett-Hamilton did not refer to _richardsonii_ possibly because he relied on Merriam's classification of 1896 wherein _richardsonii_ is treated as a species distinct from _arctica_. Merriam, it will be remembered, held that slight degree of morphological difference rather than intergradation was the criterion for subspecies. Although I have no record of having examined the type specimen of _imperii_ I have but little hesitancy in treating it as a synonym, and would have no hesitancy at all in so doing if the type was certainly known to have been obtained at Fort Simpson on the Mackenzie.

The name _Mustela cicognanii mortigena_ Bangs, 1913, proposed for the ermine of Newfoundland, is placed as a synonym of _richardsonii_ only after repeated, detailed comparisons. In advance of study I supposed that the isolation of the ermine, in Newfoundland, had contributed to its differentiation, which, however, the original describer, Bangs, indicated was slight. Bangs was a careful worker and I am confident that the differences he described really existed between his specimens. Material more nearly adequate than he had from the mainland, shows the males, so far as my measurements and comparisons go, to be in nowise different from those in Newfoundland. Females in Newfoundland may have, on the average, slightly longer hind feet than on the opposite mainland but I am not certain that they do and even if there is a slight difference in this regard as suggested by available data, I think it insufficient basis, alone, for according subspecific status to the insular animal.

The name _richardsonii_ was based by Bonaparte on Richardson's description which in turn was drawn from a specimen taken at Fort Franklin, that thus becomes the type locality. It is fortunate that Preble, in 1903, succeeded in taking specimens there because the place is near the belt of intergradation between _arctica_ and _richardsonii_. Of Preble's two adult males (see Preble, 1908:232) I have examined no. 133847, which is in transitional pelage and therefore gives no clue in so far as coloration is concerned, as to affinities with _arctica_ versus _richardsonii_. Specimens in the summer pelage are much to be desired from Fort Franklin. Regardless of what their coloration may be, specimen no. 133847, in external measurements and most certainly in cranial features is of the race to the south and not the race that Merriam named _arctica_. Because all specimens from localities to the south of Fort Franklin likewise differ from _arctica_ of the barren grounds, considerable additional confidence is felt in allocating the name _richardsonii_ to the animal which ranges from Fort Franklin southward rather than to the one, here designated _arctica_, that occurs to the northward of Fort Franklin.

Although in most structural features _richardsonii_ is a sort of average for the American races of the species, it is the extreme in high degree of sexual dimorphism. The difference in size between the males and females is greater than in any other race except possibly _M. e. kadiacensis_ in which so little is known of the female that the difference between the two sexes cannot be accurately judged. It will aid in understanding the high degree of secondary sexual difference in _richardsonii_ to visualize two kinds of weasels distributed over the northern half of the continent, thinking now of the geographic area in America occupied by the whole species _Mustela erminea_ of which the subspecies _richardsonii_ is only a part. One of the two kinds of weasel is the male ermine and the other the female. The decrease in size of the male, as measured by the weight of the skull, is in the ratio of 7 in the north to 2 in the south. This decrease is gradual whereas the corresponding decrease from 3 to 1 in the female is not gradual; half of the decrease in the female occurs in the short north to south distance comprised in the belt of intergradation, along the northern boundary of _richardsonii_, between it and _arctica_. As a result _richardsonii_ is composed of females with medium sized skulls and males with relatively large skulls, the ratio by weight being approximately 5 to 2. The disproportion in races of ermines both to the north and to the south is less. Actually in the north (_arctica_) the approximate ratio by weight is 2-1/3:1; in _richardsonii_, 2-1/2:1; in the south (_muricus_), 1-2/5:1. Indicated in still another way in _richardsonii_ the skull of the female is 56 per cent lighter than that of the male and the skull of the male is 127 per cent heavier than that of the female. Intergradation with races whose ranges border on that of _richardsonii_ is complete. On the northern boundary of the range of _richardsonii_ along the western shore of Hudsons Bay for perhaps a hundred miles north of Eskimo Point, there are intergrades with _arctica_. As judged by their lesser size, individuals of this population are influenced by the _semplei_-stock. Otherwise, intergradation on the northern boundary, with _arctica_, is abrupt whereas intergradation at the south, between _richardsonii_ and _cicognanii_, is gradual. Intergradation is similarly gradual between _richardsonii_ on the one hand and _bangsi_ and _invicta_ on the other. By speaking of the intergradation as abrupt, it is intended, in this instance, to indicate that in a relatively narrow belt, between the geographic ranges of _arctica_ and _richardsonii_, ermines intermediate in color-pattern, shape of skull, and size, bridge the gap between the ermine of the tundra (_arctica_) and that in the forest belt (_richardsonii_). It may be added that the degree of difference between the two subspecies just mentioned is approximately twice as much as between _richardsonii_ and _cicognanii_. The intergradation between _cicognanii_ and _richardsonii_ is gradual. By gradual it is meant that the change from one kind to the other is achieved in a wider area where ermines from locality A do not differ appreciably from those taken at, say, locality B, 50 miles farther south, although ermines from A and those from a third locality, C, say, 130 miles south, clearly show differences indicative of geographic variation.

_Specimens examined._--Total number, 1035, as follows. Arranged alphabetically by provinces and districts and from north to south in each province or district. Unless otherwise indicated, specimens are in the United States National Museum.

=Alberta.= 15 mi. above Smith Landing, 2; Fort Smith, 2 (1[77]); Smith Landing, 2; LaButte, Fitzgerald, 1[77]; Egg Lake, 15 mi. NW Ft. Chippewyan, 4 (2[75]); Lobstick Island, near Ft. Chippewyan, 1; Athabasca Delta, 9 mi. above mouth of main branch, 1; Athabasca Delta, Long Creek, 1 mi. W of main branch, 2; Ft. Chippewyan, 1; Peace Point, 1[75]; 18 mi. below Peace Point, 1; Embarass River, 7 (4[75]); Athabasca River, 1[2]; Ft. McMurray, 1; Athabasca River, Middle Rapid, 2; 60 mi. above Grand Rapids, 1; Boiler Rapid, 1; Entrance, 3[2]; St. Albert, 2.

=British Columbia.= Fort Halket, 1; Shesley, 1[2]; Dorothy Lake, Mts. W of Ft. Nelson, 4000 ft., 3[2]; valley between Musqwa and Prophet rivers, 3800 ft., SW of Ft. Nelson, 2[2]; Sikanni Chief Riv., 1; Telegraph Creek, 7 (6[2]); head of Bad River, 2350 ft., on lake, 1; Six Mile, 5[74]; Tuchodi Lake, 2[2]; Iskoot River, 2[14]; Level Mtn., 1[2]; head of Tatletuey Lake, 12 mi. W Thudade Lake, 2; Robb Lake District, 5[2]; Ft. Grahame, 12 (2[77]); Sustut Mts., on trib. Sustu Riv., 25 mi. SE Thudade Lake, 2; Laurier Pass, 1; Omineca Mts., 1[85]; Point Creek and Clearwater River, 2; Kispiox Valley, 23 mi. N Hazelton, 5[74]; Hazelton, 3[77]; NW arm Tacla Lake, 7; N end Babine Lake, 1; Pt. Simpson, 1; Metlakatla, 1; Stuart Lake, 27; S Fk. Salmon Riv., 1[77]; mouth Salmon Riv., 1[77]; Vanderhoof, 4[77]; Wistaria P. O., near Burns Lake, 1[77]; Kruger Lake, 9[74]; Indianpoint Lake, 23[74]; Quesnel, 1; Ahbau Lake, 3[74]; Isaacs Lake, 6[74]; Beaver Pass, 56[74]; Lightning Creek, 54[74]; LaFontaine, 16[74]; Barkerville, 1[74]; Barkerville District, 34[74]; Swift River, 27[74]; Cunningham Creek, 34[74]; Itcha Mts., 1[31]; Anahim Lake, 1[74]; Chezacut Lake, 8[31]; Kleena Kleene, 18[74]; 158 mi. House (Cariboo on labels), 3[60]; Rivers Inlet, 6 (5[94]; 1[77]); Horse Lake, 4[22]; Kingcome Inlet, 8[77]; Loughborough Inlet, 7[77]; McGillivary Creek, 1; Camel Back, Pemberton Meadows, 1[31]; Arrow Rapids, mainland opposite Stuart Island, 1[77]; Butte Inlet, 9[77]; Green Lake, 1[31]; Mt. Whistler, 1[86]; Alta Lake, 2 (1[31]; 1[21]); Mons, 1[31].

=Keewatin.= Foot of Baker Lake, 1.

=Labrador.= Okak, 3[75]; Nain, 22 (11[75]; 11[60]); Hopedale, 24[75]; Kippokak Bay, 7[75]; Ailik, 1; Makkovik, 26[75]; Labrador, 55° N, 3; Hamilton Inlet, 2[75]; NW River Post, interior Labrador, 5[1]; Cartwright, 5; Paradise, 12; Sandwich Bay (Muddy Bay, 6; North River, 6), 12; Battle Harbor, 1[7]; St. Marys River, 3[7]; Black Bay, 16 (15[75]; 1[76]); Lanceau Loup, 17 (1[75]).

=Mackenzie.= Ft. Franklin, 1[2]; Ft. Rae, 12; Fairchild Point, 6[9]; Fort Simpson, 10 (2[2]); Hot Springs (61°, 125°), 1[2]; Willow River, near Ft. Providence, 1; 35 mi. N Big Island, 7; Big Island, 9; 3 mi. S Big Island, 7; Ft. Resolution, 9; 100 mi. N Ft. Smith, 2; 75 mi. NW Ft. Smith, 1; Ft. Liard, 2; Sucker Creek, 4[77]; Govt. Hay Camp, Wood Buffalo Park, 2[77].

=Manitoba.= Egg Is., Rabbit Point, 1; Ft. Churchill, 1; Ft. York, W Hudsons Bay 57° N, 1[7]; Oxford House, 11; Gypsumville, 1[86]; Lake St. Martin.

=New Brunswick.= _Restigouche County_: Bird Bait, north Camp, 6 mi. NE Nictau Lake, 2[59]; Red Brook, Tobique River, 1[59]. _Victoria County_: Trousers Lake, 3[2]. _Glouchester County_: Youghall, 1[77]; Miramichi Road, 15 mi. from Bathurst, 13[77]. _York County_: Scotch Lake, 2.

=Newfoundland.= Nicholsville, 3[75]; Bay St. George, 48 (26[75]; 2[7]; 1[9]); Codroy, 9 (7[75]; 2[60]).

=Nova Scotia.= _Victoria County_: Cape North, 2[77]. _Inverness County_: Fizzleton, 3[77]. _Richmond County_: St. Peters, 1[77]. _Pictou County_: Glengary, 1[4]. _Guysborough County_: East Roman Valley, 5[77]. _Kings County_: Wolfville, 5 (3[74], 2[77]); near Wolfville, 1[77]. _Halifax County_: Hammond Plains, 1. _Annapolis County_: Annapolis Royal, 1. _Digby County_: Digby, 3. No locality more definite than Nova Scotia, 3.

=Ontario.= Severn River, 1[77]; R. C. Mission, Yellow Creek, near mouth of Albany, 2[86]; Ft. Albany, 4; Charlton Island, 1; Moose Factory, 10 (7[9]; 3[77]); Abitibi, 1[4].

=Quebec.= Fort Chimo, 10[77]; Ungava Forks, 1; Belcher Islands, Hudsons Bay (Tukarak Island, 29; Eskimo Harbor, 2; Innetalling Island, 1; S tip Gibson Peninsula, 2; Flaherty Island, 1), 35[9]; Cairn Island, Richmond Gulf, 2[9]; Manitounuk Sound, 4[9]; about 15 mi. S Great Whale River, 1[9]; Ft. George, 1[9]; Charlton Island, 1[9]; Waswonaby Post, 1[77]; Mistassinnay Post, 3[77]; Godbout, 36; Mt. Albert, 7 (4[78]; 3[2]); St. Anne River, 1500 ft., 1[77]; Ste. Anne des Monts, 3[2]; "Federal Mine," 1[77]; Berry Mountain Camp, 1[77]; Berry Mountain Brook, 1[2]; Cascapedia River (Middle Camp, 2; Tracadie, 2; Square Forks, 1), 5[2].

=Saskatchewan.= Poplar Point, Athabasca Lake, 1[75]; Fair Point, Athabasca Lake, 1[75]; Emma Lake, 1[74]; Harper Lake, 2[77]; Livelong, 3[55]; Fairholme, 2[74]; Touchwood Hills, 2[7]; Indian Head, 1[86].

=Yukon.= Hoole Canyon, 1; Teslin Lake (30 mi. N of, 1; Lake itself, 1; "near" the lake, 1; Mts. "near," 2; Snowden Mts., 2; Teslin Post, 2; Eagle Bay, 1; Morley Bay, 2; Nisutlin River, 1; Nisutlin Flats, 2; Wolf River, 1; Wolf Lake, 5), 21[77].

=Mustela erminea cicognanii= Bonaparte

Ermine

Plates 2, 3, 4, 9, 10 and 11

_Mustela cigognanii_ [_sic._] Bonaparte, Charlesworth's Mag. Nat. Hist., 2:37, 1838.

_Putorius vulgaris_, Emmons, Quadrupeds of Massachusetts, p. 44, 1840.

_Mustela pusilla_ DeKay, Zool. of New York, Pt. 1, Mammalia, p. 34, pl. 14, fig. 1, 1842. Type from New York State.

_Putorius pusillus_, Audubon and Bachman, Vivip. Quadrupeds of N. Amer., 2:100, pl. 64, 1851 (pl. 1846) and erroneously labeled _Mustela fusea_, as pointed out on p. 102 of text.

_Putorius cicognanii_, Baird, Mamm. N. Amer., p. 161, 1858.

_Putorius richardsoni cicognani_, Bangs, Proc. Biol. Soc. Washington, 10; 18, figs. 4, 4a of pls. 1 and 2, and pl. 3, figs. 2, 2a, February 25, 1896 (part).

_Putorius cicognani_, Merriam, N. Amer. Fauna, 11:10, pl. 2, figs. 3, 3a, 4, 4a and pl. 5, figs. 2, 2a, June 30, 1896.

_Mustela cicognanii cicognanii_, Miller, U. S. Nat. Mus. Bull., 79:95, December 31, 1912; Bishop, Journ. Mamm., 4:26, February 9, 1923.

_Mustela cicognanii_, Jackson, Journ. Mamm., 3:15, February 8, 1922.

_Mustela erminea cicognanii_, Hall, Journ. Mamm., 26:77, February 27, 1945; Hall, Journ. Mamm., 26:180, July 19, 1945.

_Type._--No type specimen designated; type locality, eastern United States.

The restriction of the type locality from the general region of northeastern North America, as given by Merriam (1896:10) to the less inclusive area of the eastern United States as earlier given by Bangs (1896:18) is supported by Bonaparte's remarks in connection with the proposal of the name _cicognanii_. He says (1838:37-38) "During my stay in the United States, I only saw a small species of _Mustela_, very common throughout the Union . . . ." This animal constituted basis for the name _cicognanii_ which name, he points out, is bestowed in order that the Americans ". . . should have constantly under their eye, this very common little animal, as the perpetual memorial . . ." to the Italian Governmental representative ". . . who, for upwards of fourteen years had served, in diplomatic and commercial concerns, . . . two countries, . . . so different . . . as the Roman and the United States. . . ." Clearly he had in mind principally, if not exclusively, the animal of the United States.