Part 11

_Putorius cicognanii alascensis_, Osgood, N. Amer. Fauna, 19:43, October 6, 1900.

_Putorius kadiacensis_, Osgood, N. Amer. Fauna, 21:69, September 26, 1901.

_Putorius audax_ Barrett-Hamilton, Ann. and Mag. Nat. Hist., 13(ser. 7):392, May, 1904, type from Discovery Bay, Ellesmere Island.

_Putorius alascensis_, Heller, Univ. California Publ. Zoöl., 5:345, March 5, 1910.

_Mustela arctica arctica_, Miller, U. S. Nat. Mus. Bull., 79:97, December 31, 1912; Dice, Journ. Mamm., 2:22, February 10, 1921.

_Mustela arctica_, Hall, Univ. California Publ. Zoöl., 30:420, March 19, 1929.

_Mustela erminea arctica_, Ognev, The mammals of U.S.S.R. and adjacent countries, 3:31, 1935; Hall, Proc. California Acad. Sci, 23:559, August 22, 1944; Hall, Journ. Mamm., 26:179, July 19, 1945.

_Type._--Male, adult, skull and skin; no. 14062/23010, U. S. Nat. Mus.; Point Barrow, Alaska; July 16, 1883; obtained by John Murdock, original no. 1672.

The skull has a fracture, on the dorsal surface, extending from the anterior nares to the interorbital constriction and another fracture on the left margin of the nasal bone. The middle of the left zygomatic arch is broken away. Otherwise the skull is complete. Right incisor one, above and below, are missing. Otherwise the teeth are present and entire. The skin is in the brown summer pelage, well made, in a good state of preservation, and shows no obvious signs of fading.

_Range._--Arctic regions of Alaska and western Canada from the Pacific Ocean to Smith Sound; from the northern limit of land south approximately to a line from Skagway through Ft. Goodhope, north shore of Great Bear Lake, south shore of Clinton Colden Lake, north shore of Baker Lake, west end of Wagner Bay to south end of Committee Bay. See figure 25 on page 95.

_Characters for ready recognition._--Differs from _M. e. polaris_ in darker upper parts (Raw Umber rather than Buckthorn Brown) and less intensely colored underparts that are Sulphur Yellow, Colonial Buff or Primrose Yellow rather than Buff Yellow; from _M. e. semplei_, in males, in that hind foot more than 44 and basilar length more than 41 and in that females average larger, the skulls of females being only about 11 per cent heavier; from _M. e. kadiacensis_ in hind foot more than 33 in females, zygomatic breadth amounting to more, rather than less, than distance between last upper molar and jugular foramen irrespective of sex; from _M. e. richardsonii_, _alascensis_, _salva_ and _initis_, both sexes so far as known, by proximal two-thirds of under side of tail colored same as underparts rather than same as upper parts, and by zygomatic breadth amounting to more, rather than less, than distance between last upper molar and jugular foramen.

_Description.--Size._--Male: Six adults from Tanana, Alaska, yield average and extreme measurements as follows: Total length, 336 (310-350); length of tail, 93 (84-105); length of hind foot, 49 (45-51).

Female: Five adults, one each from Alatna River, mountains near Eagle, Kamarkak in Alaska, Arctic Red River and Baillie Island in Canada, yield average and extreme measurements as follows: Total length, 285 (272-304); length of tail, 77 (68-95); length of hind foot, 39 (34-43).

Weight of 5 subadult males from Tanana is 206 (163-248) grams; adults would be heavier.

_Color._--Winter pelage all white except tip of tail. Summer pelage with upper parts uniform in color and Raw Umber or darker (16_n_) of Ridgway and about tones 2 to 3 of Chocolate of Oberthür and Dauthenay, pl. 343, but in autumn some specimens have more light red than tones 2 or 3. Underparts Sulphur Yellow, Colonial Buff, or Primrose Yellow, often white on chin and insides of forelegs; color of underparts extends narrowly over upper lips, distally on posterior sides of forelegs onto antipalmar surface of forefeet, onto proximal two-thirds or three-fourths of underside of tail as length of tail is measured along tail-vertebrae, 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; rim of ear in some specimens with short, white or pale hairs giving ears distinct whitish border; least width of color of underparts averaging, in adult males from Alaska, 65 (46-93) per cent of greatest width of color of upper parts. Black tip of tail in 5 males in winter pelage from Tanana averaging 84 (70-93) mm. which is 91 (75-107) per cent of length of tail-vertebrae.

_Skull._--Male (based on 5 adult topotypes): See measurements and plates 2-4. As described in _Mustela erminea richardsonii_ except that: Weight, 3.5 (3.1-3.9) grams; basilar length 42.5 (41.8-43.3); length of tooth-rows more than length of tympanic bulla; breadth of rostrum measured across lacrimal processes averaging more than a third of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more than distance between last upper molar and jugular foramen.

Female (based on 2 adult topotypes and 2 adults and 4 subadults from central Alaska): See measurements and plates 9-11. As described in _Mustela erminea richardsonii_ except that: Weight, 1.5 (1.2-2.0) grams; basilar length, 35.7 (34.5-37.0); length of tooth-rows more than length of tympanic bulla; breadth of rostrum more than 30 per cent of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more than distance between last upper molar and jugular foramen (except in specimens from Ellesmere Island where two distances are approximately equal).

Cranial differences from _Mustela erminea kaneii_ (which occurs on the Asiatic side of Bering Strait), in both males and females, are: larger size relatively as well as actually, broader except in mastoidal region where relatively (to basilar length) the width is less; preorbital part of skull broader as well as longer.

From _kadiacensis_ differences in the skull of the male are: size less; 13 per cent heavier, relatively (to basilar length) narrower across interorbital region and zygomatic arches; tympanic bullae relatively as well as actually narrower. Judging by the single available adult female of _kadiacensis_, the skull of female _arctica_ is larger in all parts measured, a fourth heavier, has tympanic bullae of almost twice the volume and the interorbital and preorbital regions, relative to the braincase, are much reduced in whatever plane measured.

Differences from _richardsonii_, additional to those noted above in the formal description of the skull, between the males, are: larger in all parts measured except length of tympanic bulla which is about the same; 42 per cent heavier; relative to basilar length, skull broader with preorbital part longer as well as broader; tympanic bullae more inflated posteriorly. The same differences prevail between females except that the skull is 36 per cent heavier and in _arctica_ the length of the bulla is actually more (although relative to the basilar length less) and its greater inflation posteriorly is hardly perceptible. Differences from _alascensis_, additional to those indicated in the formal descriptions of the skulls of the two, in males, are: larger in every part measured; 95 per cent heavier; relative to the basilar length, skull broader with preorbital part longer as well as broader; measured at a point opposite the foramen lacerum anterius, the width of the pterygoid space is more, rather than less, than 40 per cent of its length. Excepting this difference in width of interpterygoid space, the same differences prevail between females, those of _arctica_ being 56 per cent heavier.

Comparison with _semplei_ is made in the account of that subspecies.

Skull indistinguishable from that of _polaris_.

_Remarks._--The person who studies specimens of this subspecies finds labels inscribed with the names of naturalists well known to all readers of literature on the Arctic. Sir John Franklin, R. McFarlane, R. Kennicott, E. W. Nelson and R. M. Anderson are names which appear commonly. Of Alaskan specimens prepared according to modern methods, a large share was obtained by O. J. Murie and L. R. Dice.

The ermine was observed in the far north by early explorers and was mentioned in the literature, almost always under the name then used for the ermine of northern Europe and Asia. In 1896 Bangs misapplied to it the name _richardsonii_ but Merriam in the same year corrected the application of this name and proposed as new for this weasel the name _arctica_, the name in use today. For almost 50 years after Merriam and Bangs wrote about it, _arctica_ was treated, nominally at least, as a species distinct from its other relatives in both the Old-and New-World. The subspecific status of _arctica_ was emphasized in 1944 (555) by the present writer in reporting in detail upon the specimens, of _Mustela erminea_, from Eastern Asia which were made available on loan by Professor B. S. Vinogradov and the late Anatol I. Argyropulo of the Leningrad Academy of Sciences. Specimens of _Mustela erminea kaneii_ from the Asiatic side of Bering Strait and _Mustela erminea arctica_ from the American side are distinguishable by slight cranial characters but in coloration and external measurements I can detect no differences. Merriam's (1896:16) mention of more golden-colored upper parts and darker underparts in American specimens than in _erminea_ was the result of his comparison of Alaskan and northern European specimens. When Old World specimens from eastern Siberia, instead of from Europe, are used the differences mentioned by Merriam do not apply. Incidentally, many Siberian specimens have the white border, on the ear, which Merriam (_loc. cit._) noted as a distinguishing feature of _arctica_. When Merriam named _arctica_ he said (1896:15, 16) "_Putorius arcticus_ . . . has heretofore been confounded with _erminea_ or _richardsonii_. . . . It is interesting to find in this country an Arctic circumpolar weasel which, though specifically distinct, is strictly the American representative of the Old World _erminea_." Bearing in mind that Merriam's concept of species and subspecies (see Merriam, 1919:6) differed from that of nearly all modern systematists it is clear from his statement quoted above that he correctly understood the zoölogical relationship obtaining between the ermines of the Old and New Worlds.

Ognev (1935:31) seems to have been the first to use the name combination _Mustela erminea arctica_ for Alaskan specimens. Thereby he expresses the view adopted here, namely that the American ermine is subspecifically but not specifically distinct from the Old World animal. Whether actual intergradation (crossbreeding) ever takes place across the narrow Bering Strait I do not know. I doubt that crossbreeding occurs but considering the Diomedes (islands), that might serve as a half way stopping point, and remembering Mr. Charles Brower's oral statement to me that he had seen tracks of ermine as far as 10 miles from the northern shore of Alaska out on the ice, the possibility must be granted of an occasional individual crossing from one side to the other of Bering Strait on the ice in winter or of being carried across when the ice broke up and drifted. If transfers of this kind occurred often one would expect ermines to occur also on Saint Lawrence Island where apparently they do not. The one skin (U. S. Nat. Mus. no. 259046) seen as labeled from there, my friend, Otto William Geist ascertained was imported as a skin with other furs from Siberia.

Ognev (_op. cit._) who used the name combination _Mustela erminea arctica_ for Alaskan specimens, applied it also to animals from Kamchatka. At the same time he recognized the animal from the eastern mainland of Siberia (as opposed to the peninsula of Kamchatka) under the name _Mustela erminea orientalis_ Ognev 1928. Hall (1944:556) applied the earlier proposed name _Putorius kaneii_ Baird 1857, to the animal on the eastern mainland of Asia and proposed the new name _Mustela erminea digna_ for the ermine of Kamchatka. In comparing material of these two Asiatic races with topotypes and other specimens of _M. e. arctica_ from Alaska, it seemed to me that the degree of relationship, one with the other, was about the same. _M. e. digna_ has a slightly larger preorbital region than _M. e. kaneii_, and the skull is longer. In both of these particulars _digna_ approaches closer to _arctica_. _M. e. kaneii_ has longer tympanic bullae and a wider skull than _digna_ and therein approaches more towards _arctica_ than toward _digna_. As nearly as I can make out, _digna_ and _kaneii_ show a nearly equal degree of resemblance to _arctica_. Also the degree of difference between _digna_ and _kaneii_ is about the same as between either one of them and _arctica_. In view of the above considerations the ermines of the New and Old worlds are here regarded as only subspecifically distinct.

In the original description of _Putorius audax_ (here regarded as inseparable from _Putorius arcticus_ Merriam) Barrett-Hamilton erroneously designated the type locality as "Discovery Bay, North Greenland" whereas he should have written Grinnell Land [= Ellesmere Island of modern terminology] in place of Greenland. As reference to Nares (1877 and 1878) will readily reveal, Discovery Bay is near 65° W and 81° 40´ N, across Robeson Channel, to the west, from Greenland. The label on the type specimen and the specimen register in the British Museum of Natural History each designates the locality for this specimen, the type of _audax_, as Discovery Bay without mention of Greenland. The published accounts of Feilden (1878) and Nares (1877 and 1878) state that specimens of ermine were obtained at Discovery Bay. Probably H. C. Hart is the collector of the specimen; he was the naturalist attached to H. M. S. Discovery which wintered at Discovery Bay while H. W. Feilden was the naturalist attached to H. M. S. Alert which wintered a few miles southeast of Cape Sheridan, also on the eastern coast of Ellesmere Island.

It is true that from these ships a trip was made into Greenland and an ermine (only one individual it seems) was obtained there, but this individual was the type specimen of _Mustela erminea polaris_, in the account of which race something of the history of this specimen is given.

With the material available--and it is not entirely adequate--I can detect no features by which animals from the type locality of _audax_ can be distinguished from typical _arctica_ which latter name has priority.

Intergradation with _richardsonii_ probably occurs completely across the continent. Intergrades here referred to _arctica_ include those from Fort Goodhope. The one defective specimen from Lake Lebarge, Yukon, is not certainly identified as _arctica_ and how far west of Teslin Lake the boundary-line between _arctica_ and _richardsonii_ should be drawn remains to be ascertained. The one specimen available from Hinchenbrook Island, no. 912 Mus. Vert. Zoöl., an adult female, is doubtfully referred to _arctica_ because the damaged tympanic bullae appear to be no larger than in _alascensis_, and the size of the skull is more as in _alascensis_ although intermediate between that race and _arctica_. Shape of the skull is more as in _arctica_. Possibly more nearly adequate material would show the existence on Hinchenbrook Island of an insular race differing in about the same degree from _arctica_ of the mainland as does the insular _kadiacensis_. Nevertheless, the males from farther south at Cape Yakataga are in all respects _arctica_ and this argues against near relationship to _alascensis_ of the animal on Hinchenbrook Island. The three animals seen from Yakutat Bay are so young as not to display clearly the cranial characters of the subspecies but the extension of the color of the underparts onto the underside of the tail in them and also in the skin without corresponding skull from Glacier Bay, Alaska, is as in _arctica_, the race to which they are referred, and gives substantial basis for showing the geographic range of _arctica_ as extending this far south along the Pacific Coast.

_Specimens examined._--Total number, 281, arranged alphabetically by Districts and from north to south in each District. Unless otherwise indicated, specimens are in the collection of the United States National Museum.

=Alaska.= Point Barrow, 22 (1[1], 1[2], 1[75], 4[1], 7[60], 6[74]); Flaxman Island, 3; Collinson Point, 1[77]; Salirochet River, 1[77]; Hulahula River, 1[2]; 69°20´ & 141°, 1; Rampart House, 1; Yukon River, mouth of Porcupine River, 18; Alatna River, 30 mi. from mouth, 1; Koyakuk Riv., 16 mi. below Bettles, 4; Shelton, 1[75]; Kruzamepa, 1[75]; Tanana, 6; Boulder Creek, Chena River, 3; Fort Reliance, 4; Yukon River, 20 miles above Circle, 2; Mts. near Eagle, 42 (1[60]); Snake River, Nome, 1[9]; Nulato, 3; No[e]wikakat Riv., 1; Kantishna, 3; Fairbanks, 5 (1 20 mi. E and 1 33 mi. E); Richardson, 1; N. Fk. Kuskokwim R. at base of Mt. Sischo, 1; N. Fk. Kuskokwim R. at Junction with McKinley Fk., 1; Nenana Riv., mouth of Maurice Cr., 1; Ober Cr., trib. of Jarvis Cr., Delta Riv. region, 1; head of Savage Riv., near Jennie Cr., 1; Wonder Lake, 1[74]; Bear Cr., 3; Unlakleet, 3; St. Michaels, 11; 125 mi. E and a little N of Knik, Cook Inlet, on S side Matanuska Range, 1[60]; Hope, Cook Inlet, 1; Iak Lake, 1[68]; head of Behring Riv., 1; Bethel, 2; Kenai Lake, 8; Kenai Peninsula, 13 (2[2]); He[i]nchenbrook Island, 1200 ft., 1[74]; Sunshine Point, Kaliekh River, Yakataga Dist., 1[8]; Cape Yakataga, 3[8]; Yakutat Bay, 3[74]; Seward, 7; Seldovia, 22 (4[2]); Homer, 1[2]; Cape Elizabeth, 18; Akchookuk Lake, 1; Lake Weelooluk, 1; Kokwok Riv., 80 mi. up, 4; Nushagak, 1; Nushagak Riv., 1; Kolukuk, 1; Egooshik River at mouth, 1; Glacier Bay, 1; Becharof Lake, between Portage Bay and Becharof Lake, 1; Ugashik Riv., 4; Chignik, 7; East base Frosty Peak, 1; Pavlov Bay, 1[100]; Mt. Pavlof, 1[75]; Unimak Island, 2 (1[75]).

=District of Franklin.= Cape Sheridan, 1[2]; Discovery Bay, Ellesmere Island, 1[7] (type specimen of _Putorius audax_ Barrett-Hamilton); Axel Heiberg Island, 1[95]; Bache Peninsula, Ellesmere Island, 1[77]; Bedford Pims Island, 4[75]; Craig Harbor, 2[77]; Cape Kellett, Banks Island, 1[77]; Franklin Isthmus, 1[95]; King William Island, 2[95].

=District of Keewatin.= Ualiak, Ogden Bay, 2[95].

=District of Mackenzie.= Baillie Island, 1[75]; Franklin Bay, 1; Langton Bay, arm of Franklin Bay, 15 mi. S of, 1[2]; Cockburn Point, 69°N, 115°W, 2[77]; Dolphin and Union Strait, 1[77]; Bernard Harbor, 2[77]; Kent Peninsula, 4[95]; Horton Riv., near Fort Anderson, 1; Fort Anderson, 6; Anderson River, 3; Barry Island, Bathurst Inlet, 1[77]; Fort McPherson, 1; Peels River, 2; Arctic Red River, 8[75]; Fort Good Hope, 6; Clinton Colden, 1[2].

=Yukon.= Kamarkak, 1[77]; Herschel Island, 1[75]; Lapierres House, 2; Forty Mile, L. T. Coal Cr., 4[74]; head of Coal Cr., 1; Macmillan River, Forks, 1; 20 mi. W. Ft. Selkirk, 1; Slims River, near Kluane, 1[75]; head of Lake Lebarge, 1.

=Mustela erminea polaris= (Barrett-Hamilton)

Ermine

_Putorius arcticus polaris_ Barrett-Hamilton, Ann. and Mag. Nat. Hist., 13 (ser. 7):393, May, 1904.

_Mustela erminea_, Manniche, Meddelelser on Grønland, 45:80-85, 1 fig., 1910.

_Mustela arctica polaris_, Miller, U. S. Nat. Mus. Bull., 79:97, December 31, 1912.

_Mustela erminea polaris_, Hall, Journ. Mamm., 26:179, July 19, 1945.

_Type._--Probably female, skin only; no. 78. 6. 19. 11, Brit. Mus. Nat. Hist.; Gap Valley, 7-1/4 miles northeast Cape Brevoort, 82° N, 59° 20´ W, Northwestern Greenland; June 15 or 16, 1876; obtained by Lewis A. Beaumont.

The skin is in full, fresh summer pelage, fairly well stuffed except for the tail which is unstuffed; the whole is in a good state of preservation.

_Range._--North coast, and east coast as far south as Turner Sound (between 69 and 70 degrees) of Greenland. See figure 25 on page 95.

_Characters for ready recognition._--Differs from _M. e. arctica_ in lighter upper parts (near [_j_] Buckthorn Brown rather than Raw Umber or darker) and more intensely-colored underparts that are Buff Yellow rather than Sulphur Yellow, Colonial Buff, or Primrose Yellow; from _M. e. semplei_ in color in same fashion as from _arctica_ and in larger size of skull.

_Description._--_Size._--Male: One subadult and two adults (one ad. from Scøresby Sound and other two from Ymer Island) measure as follows, the average being given first: Total length, 318 (301, 320, 315); length of tail, 72 (69, 70, 73); length of hind foot, 46.5 (44, 46, 47).

Female: No measurements taken in the flesh available but hind foot, measuring 33.5 in the dried state and therefore approximately 35 in life.

_Color._--As described in _Mustela erminea arctica_ except that upper parts in summer near (_j_) Buckthorn Brown and tone 4 of Dark Fawn of plate 307 to tone 1 of Raw Umber of plate 301 of Oberthür and Dauthenay. Underparts Buff-Yellow. Least width of color of underparts averaging, in 3 males, 66 (57-72) per cent of greatest width of color of upper parts. Black tip of tail in same males averaging 71 (70-72) mm. which is 99 (99-104) per cent of length of tail-vertebrae.

The lighter-colored upper parts and more intensely yellow underparts are the distinguishing features of the subspecies _polaris_ in comparison with other races of American _M. erminea_.

_Skull._--Male (based on 5 adults from eastern Greenland): See measurements. As described in _Mustela erminea richardsonii_ except that: Weight more (not recorded); basilar length, 41.3 (39.0-42.4); length of tooth-rows more than length of tympanic bulla; breadth of rostrum measured across lacrimal processes averaging more than a third of basilar length; interorbital breadth more than distance between glenoid fossa and posterior border of external auditory meatus; zygomatic breadth more than distance between last upper molar and jugular foramen.

Female (based on 2 adults, Turner Sund and Kap Hoeg): See measurements. As described in _Mustela erminea arctica_ except that basilar length 36.8 (35.9, 37.8), and length of tooth-rows not more than length of tympanic bulla. Skulls of females not in hand when this comparison is written; only the recorded measurements are available.

To me the skull of _polaris_ is indistinguishable from that of _arctica_. Therefore the comparisons made of the skull of _arctica_ with those of other subspecies will apply also for _polaris_.