Part 9

1900. =eskimo= (_Putorius rixosus_) Stone [= _Mustela rixosa eskimo_], Proc. Acad. Nat. Sci. Philadelphia, 1900:44, March 24, 1900, is applied to the least weasel of Alaska and adjacent parts of boreal North America.

1901. =allegheniensis= (_Putorius_) Rhoads [= _Mustela rixosa allegheniensis_], Proc. Acad. Nat. Sci. Philadelphia, 1900:75, March 25, 1901, applies to the least weasel of the eastern United States.

1902. =perdus= (_Putorius tropicalis_) Merriam [= _Mustela frenata perda_], Proc. Biol. Soc. Washington, 15:67, March 22, 1902, applies to the long-tailed weasel of the Lower Tropical Life-zone from southern Veracruz into Guatemala.

1903. =microtis= (_Putorius_) Allen [= _Mustela erminea richardsonii_], Bull. Amer. Mus. Nat. Hist., 19:563, October 10, 1903, is a name applied to an individual ermine of small size from Shesley, British Columbia, which Allen thought was specifically distinct from the ermine of the Hudsonian Life-zone and adjacent territory. Now the name is arranged as a synonym of _richardsonii_.

1904. =audax= (_Putorius_) Barrett-Hamilton [= _Mustela erminea arctica_], Ann. and Mag. Nat. Hist., ser. 7, 13:392, May, 1904. In the original description the type locality, Discovery Bay, was erroneously stated to be in Greenland and the name _audax_ until 1945 was applied to the kind of weasel occurring in northern Greenland whereas the type specimen was taken instead in northern Ellesmere Island and because the weasel there is subspecifically indistinguishable from ermines from farther west, _audax_ is a synonym of _Putorius arcticus_.

1904. =imperii= (_Putorius arcticus_) Barrett-Hamilton [= _Mustela erminea richardsonii_], Ann. and Mag. Nat. Hist., ser. 7, 13:392, May, 1904, based on an animal from Fort Simpson, Mackenzie, Canada, proves to be inseparable from _richardsonii_ which has priority.

1904. =polaris= (_Putorius arcticus_) Barrett-Hamilton [= _Mustela erminea polaris_], Ann. and Mag. Nat. Hist., ser. 7, 13:393, May, 1904, is the name used for the ermine of eastern Greenland and since 1945 has been used for the weasel of Greenland as a whole.

1905. =macrophonius= (_Putorius_) Elliott [= _Mustela frenata macrophonius_], Proc. Biol. Soc. Washington, 18:235, December 9, 1905, applies to the long-tailed weasel of the mountains along the eastern border of Veracruz.

1906. =leptus= (_Putorius streatori_) Merriam [= _Mustela erminea murica_], Proc. Biol. Soc. Washington, 16:76, May 29, 1903, until 1945 was applied to the diminutive ermine of the Rocky Mountains from Wyoming south to northern New Mexico but proves to be a synonym of _muricus_ with type locality in the Sierra Nevada of California.

1908. =angustidens= (_Putorius cicognanii_) Brown [= _Mustela erminea angustidens_], Mem. Amer. Mus. Nat. Hist., 9(pt. 4):181, pl. 17, is applied to an extinct subspecies known from fossil remains of Pleistocene age from northern Arkansas.

1908. =gracilis= (_Putorius_) Brown [= _Mustela frenata gracilis_], Mem. Amer. Mus. Nat. Hist., 9(pt. 4):182, 1908, applies to a Pleistocene weasel known from a single skull from northern Arkansas.

1912. =costaricensis= (_Mustela_) Goldman [= _Mustela frenata costaricensis_], Proc. Biol. Soc. Washington, 25:9, January 23, 1912, applies to the long-tailed weasel of Costa Rica.

1913. =primulina= (_Mustela_) Jackson [= _Mustela frenata primulina_], Proc. Biol. Soc. Washington, 26:123, May 21, 1913, applies to the long-tailed weasel of the central part of the United States in eastern Kansas and adjoining areas.

1913. =campestris= (_Mustela_) Jackson [= _Mustela rixosa campestris_], Proc. Biol. Soc. Washington, 26:124, May 21, 1913, applies to the least weasel of the Great Plains region.

1913. =olivacea= (_Mustela peninsulae_) Howell [= _Mustela frenata olivacea_], Proc. Biol. Soc. Washington, 26:139, May 21, 1913, applies to the long-tailed weasel of the southeastern United States excepting most of Florida.

1914. =meridana= (_Mustela_) Hollister [= _Mustela frenata meridana_], Proc. Biol. Soc. Washington, 27:143, July 10, 1914, applies to the long-tailed weasel of northern South America.

1916. =nicaraguae= (_Mustela tropicalis_) Allen [= _Mustela frenata nicaraguae_], Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916, applies to the long-tailed weasel of Nicaragua.

1927. =arthuri= (_Mustela noveboracensis_) Hall [= _Mustela frenata arthuri_], Proc. Biol. Soc. Washington, 40:193, December 2, 1927, applies to the long-tailed weasel of Louisiana and adjoining areas.

1932. =semplei= (_Mustela arctica_) Sutton and Hamilton [= _Mustela erminea semplei_], Ann. Carnegie Mus., 21(2):79, February 13, 1932, originally was applied to the ermine of Southampton Island but after 1945 was applied also to the ermine of Baffin Island, Melville Peninsula and the west side of Hudsons Bay as far south as Eskimo Point.

1932. =panamensis= (_Mustela frenata_) Hall, Proc. Biol. Soc. Washington, 45:139, September 9, 1932, applies to the long-tailed weasel of Panamá.

1932. =anguinae= (_Mustela cicognanii_) Hall [= _Mustela erminea anguinae_], Univ. California Publ. Zoöl., 38:417, November 8, 1932, applies to the ermine of Vancouver Island, British Columbia.

1935. =labiata= (_Mustela arctica_) Degerbøl [= _Mustela erminea semplei_], Rept. 5th Thule Exped., 1921-1924, vol. 2, no. 4, p. 25, 1935. When Degerbøl wrote his description and proposed this name he was unaware that Sutton and Hamilton had three years before based a new name on weasels from Southampton Island. Because the two names apply to the same subspecies, Degerbøl's name, _labiata_, must fall as a synonym of _semplei_ which has priority.

1935. =helleri= (_Mustela frenata_) Hall, Proc. Biol. Soc. Washington, 48:143, August 22, 1935, applies to the long-tailed weasel of eastern Perú.

1936. =nevadensis= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 91, November 20, 1945, applies to the long-tailed weasel of the western United States. For many years, animals of this subspecies were referred to _longicauda_ and from 1891 until 1936 to _arizonensis_.

1936. =effera= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 93, November 20, 1945, applies to the long-tailed weasel of the Blue Mountains region. From 1891 until 1936 this animal was referred to under the name _arizonensis_.

1936. =altifrontalis= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 94, November 20, 1936, applies to the long-tailed weasel of the humid coastal district from Puget Sound southward into Oregon.

1936. =nigriauris= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 95, November 20, 1936, applies to the long-tailed weasel of the coastal district of California from San Francisco Bay southward to Point Concepcion. Previous to 1936, _xanthogenys_ was the name applied to this race of weasel.

1936. =latirostra= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 96, November 20, 1936, applies to the long-tailed weasel of southern California which previously had borne the name _xanthogenys_.

1936. =pulchra= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 98, November 20, 1936, is applied to the long-tailed weasel of the southern end of the San Joaquin Valley of California.

1936. =inyoensis= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 99, November 20, 1936, is applied to the long-tailed weasel of Owens Valley, California.

1936. =texensis= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 99, November 20, 1936, applies to the long-tailed weasel of central Texas which previously had been assigned to the subspecies _frenata_.

1936. =perotae= (_Mustela frenata_) Hall, Carnegie Inst. Washington, publ. no. 473, p. 100, November 20, 1936, applies to long-tailed weasel of the mountains along the Puebla-México boundary.

1938. =boliviensis= (_Mustela frenata_) Hall, Proc. Biol. Soc. Washington, 51:67, May 18, 1938, applies to the southernmost known long-tailed weasel which is in the Lake Titicaca region in Perú and Bolivia.

1944. =salva= (_Mustela erminea_) Hall, Proc. Biol. Soc. Washington, 57:35, June 28, 1944, applies to the ermine of Admiralty Island, southeastern Alaska.

1944. =initis= (_Mustela erminea_) Hall, Proc. Biol. Soc. Washington, 57:37, June 28, 1944, applies to the ermine of Baranof and Chichagof islands, southeastern Alaska.

1944. =celenda= (_Mustela erminea_) Hall, Proc. Biol. Soc. Washington, 57:38, June 28, 1944, applies to the ermine of Prince of Wales, Dall and Long islands, Alaska.

1944. =seclusa= (_Mustela erminea_) Hall, Proc. Biol. Soc. Washington, 57:39, June 28, 1944, applies to the ermine of Suemez Island, southeastern Alaska.

1945. =invicta= (_Mustela erminea_) Hall, Jour. Mamm., 26:75, February 27, 1945, applies to the ermine of the Rocky Mountains for several hundred miles both north and south of the United States-Canadian boundary.

1945. =fallenda= (_Mustela erminea_) Hall, Jour. Mamm., 26:79, February 27, 1945, applies to the ermine of the coastal mainland in southern British Columbia and northern Washington.

1945. =olympica= (_Mustela erminea_) Hall, Jour. Mamm., 26:81, February 27, 1945, applies to the diminutive ermine of the Olympic Peninsula, state of Washington.

1945. =gulosa= (_Mustela erminea_) Hall, Jour. Mamm., 26:84, February 27, 1945, applies to the diminutive ermine of the Cascades in Washington.

1945. =bangsi= (_Mustela erminea_) Hall, Jour. Mamm., 26:176, July 19, 1945, is the name applied today to the ermine of the western Great Lakes region.

In 1925 when this study was begun, the American weasels (subgenus _Mustela_ proper) were arranged as belonging to 47 kinds (including subspecies) of 29 full species. In the present account a total of 68 kinds, belonging to 4 full species are recognized in the subgenus _Mustela_. The increase in number of subspecies and the decrease in number of species are the nomenclatural results ordinarily obtained in this decade from a systematic study of a genus of American mammals.

CHECK-LIST OF AMERICAN SPECIES AND SUBSPECIES OF THE GENUS MUSTELA

Subgenus =MUSTELA= Linnaeus

PAGE _Mustela erminea_ 87 _Mustela erminea arctica_ (Merriam) 96 _Mustela erminea polaris_ (Barrett-Hamilton) 103 _Mustela erminea semplei_ Sutton and Hamilton 105 _Mustela erminea kadiacensis_ (Merriam) 108 _Mustela erminea richardsonii_ Bonaparte 110 _Mustela erminea cicognanii_ Bonaparte 118 _Mustela erminea bangsi_ Hall 124 _Mustela erminea invicta_ Hall 128 _Mustela erminea alascensis_ (Merriam) 131 _Mustela erminea salva_ Hall 135 _Mustela erminea initis_ Hall 136 _Mustela erminea celenda_ Hall 139 _Mustela erminea seclusa_ Hall 141 _Mustela erminea haidarum_ (Preble) 142 _Mustela erminea anguinae_ Hall 145 _Mustela erminea fallenda_ Hall 148 _Mustela erminea olympica_ Hall 153 _Mustela erminea streatori_ (Merriam) 155 _Mustela erminea gulosa_ Hall 159 _Mustela erminea muricus_ (Bangs) 161 _Mustela erminea angustidens_ (Brown) 165

_Mustela rixosa_ 168 _Mustela rixosa eskimo_ Stone 181 _Mustela rixosa rixosa_ Bangs 184 _Mustela rixosa allegheniensis_ Rhoads 187 _Mustela rixosa campestris_ Jackson 190

_Mustela frenata_ 193 _Mustela frenata noveboracensis_ (Emmons) 222 _Mustela frenata occisor_ (Bangs) 230 _Mustela frenata primulina_ (Jackson) 232 _Mustela frenata arthuri_ Hall 241 _Mustela frenata olivacea_ Howell 244 _Mustela frenata peninsulae_ Rhoads 250 _Mustela frenata spadix_ (Bangs) 252 _Mustela frenata longicauda_ Bonaparte 262 _Mustela frenata oribasus_ (Bangs) 270 _Mustela frenata alleni_ (Merriam) 274 _Mustela frenata arizonensis_ (Mearns) 276 _Mustela frenata nevadensis_ Hall 280 _Mustela frenata effera_ Hall 291 _Mustela frenata washingtoni_ (Merriam) 294 _Mustela frenata saturata_ (Merriam) 297 _Mustela frenata altifrontalis_ Hall 300 _Mustela frenata oregonensis_ (Merriam) 304 _Mustela frenata munda_ (Bangs) 309 _Mustela frenata xanthogenys_ Gray 315 _Mustela frenata nigriauris_ Hall 319 _Mustela frenata latirostra_ Hall 323 _Mustela frenata pulchra_ Hall 328 _Mustela frenata inyoensis_ Hall 331 _Mustela frenata neomexicana_ (Barber and Cockerell) 333 _Mustela frenata texensis_ Hall 338 _Mustela frenata frenata_ Lichtenstein 341 _Mustela frenata leucoparia_ (Merriam) 347 _Mustela frenata perotae_ Hall 351 _Mustela frenata goldmani_ (Merriam) 355 _Mustela frenata macrophonius_ (Elliot) 360 _Mustela frenata tropicalis_ (Merriam) 363 _Mustela frenata perda_ (Merriam) 366 _Mustela frenata nicaraguae_ Allen 370 _Mustela frenata costaricensis_ Goldman 372 _Mustela frenata panamensis_ Hall 375 _Mustela frenata meridana_ Hollister 379 _Mustela frenata affinis_ Gray 384 _Mustela frenata aureoventris_ Gray 387 _Mustela frenata helleri_ Hall 391 _Mustela frenata agilis_ Tschudi 393 _Mustela frenata macrura_ Taczanowski 398 _Mustela frenata boliviensis_ Hall 402 _Mustela frenata gracilis_ (Brown) 404

Subgenus =Grammogale= Cabrera

_Mustela africana_ 406 _Mustela africana africana_ Desmarest 409 _Mustela africana stolzmanni_ Taczanowski 413

Subgenus =Putorius= Cuvier

(Black-footed Ferret--not treated in present work)

_Mustela nigripes_ (Audubon and Bachman)

Subgenus =Lutreola= Wagner

(Minks--not treated in present work)

_Mustela vison_ _Mustela vison vison_ Schreber _Mustela vison mink_ Peale and Beauvois _Mustela vison lutensis_ (Bangs) _Mustela vison evergladensis_ Hamilton _Mustela vison vulgivaga_ (Bangs) _Mustela vison letifera_ Hollister _Mustela vison lacustris_ (Preble) _Mustela vison energumenos_ (Bangs) _Mustela vison evagor_ Hall _Mustela vison aestuarina_ Grinnell _Mustela vison nesolestes_ (Heller) _Mustela vison melampelus_ (Elliot) _Mustela vison ingens_ (Osgood) _Mustela macrodon_ (Prentiss)

ARTIFICIAL KEY TO AMERICAN SPECIES OF THE GENUS MUSTELA

PAGE

A Length of upper tooth-rows less than 20 mm. in males and 17.8 mm. in females.

B Postglenoid length of skull more than 47 per cent of condylobasal length.

C Tail without a black pencil and with at most a few black hairs at extreme tip; in both sexes mastoid breadth ordinarily exceeds breadth of braincase, _Mustela rixosa_, least weasel, p. 168

C' Tail with a black pencil; in females mastoid breadth ordinarily exceeded by breadth of braincase, _Mustela erminea_, ermine, p. 87

B' Postglenoid length of skull less than 47 per cent of condylobasal length.

D Tail with distinct black tip; midventral line white, yellowish, orange, not same color as upper parts; p2 present; thenar pad on forefoot absent, _Mustela frenata_, long-tailed weasel, p. 193

D' Tail without black tip; midventral line same color as upper parts; p2 absent; thenar pad on forefoot present, _Mustela africana_, tropical weasel, p. 406

A' Length of upper tooth-rows more than 20 mm. in males and 17.8 mm. in females.

E Abdomen all white; face with blackish mask; m1 lacking even a trace of a metaconid; distance between upper canines more than width of basioccipital as measured between foramina situated midway along medial sides of tympanic bullae, _Mustela nigripes_, black-footed ferret.

E' Abdomen dark brown, like back; face uniformly brown without blackish mask; m1 with incipient metaconid; distance between upper canines less than width of basioccipital as measured between foramina situated midway along medial sides of tympanic bullae, _Mustela vison_, mink, American mink.

DIAGNOSIS OF THE GENUS

Genus =Mustela= Linnaeus

Weasels, Ferrets, Polecats, Minks

_Genotype._--_Mustela erminea_ Linnaeus.

_Diagnosis._--Legs short; body relatively long; adults 190 mm. to 700 mm. in total length; skull ranging in basilar length from 16 to 70 mm.; facial angle slight; tympanic bullae greatly inflated (moderately in _Lutreola_), cancellous, and with paroccipital processes closely appressed to bullae; palate behind upper molars; dental formula:

I 3 C 1 P 2-3 M 1 -, -; -, -; -, ---; -, -; inner moiety of M1 larger than outer; P4 i 3 c 1 p 3-2 m 2

with simple deuterocone; in m1 inner moiety of M1 larger than outer; P4 with simple deuterocone; in m1 trigonid longer than talonid, metaconid absent (incipiently developed in _Lutreola_), and talonid trenchant.

For many years prior to 1911, the name _Mustela_ was applied to martens, and _Putorius_ was regarded as the first available generic name for the weasels. In 1911 Thomas (1911:139) showed that _M. erminea_ (_Mustela_ of Gesner) by tautonymy was the type of _Mustela_ and subsequently the generic name _Mustela_ has been used for the true weasels which include the American weasels to which we now apply the specific names _erminea_, _rixosa_ and _frenata_. The mink, _Mustela (Lutreola) vison_, and the black-footed ferret, _Mustela (Putorius) nigripes_, since 1911 also have been referred by most American authors to the genus _Mustela_, the names _Lutreola_ and _Putorius_ being regarded by these authors as having no more than subgeneric status. European writers, on the other hand, accord greater taxonomic weight to the zoölogical differences between ferrets and weasels and, therefore, accord full generic rank to _Putorius_. Consequently, for the black-footed ferret, Europeans today write _Putorius nigripes_ and Americans write _Mustela nigripes_. For the same reasons, the name of the mink is written by some European zoölogists _Lutreola vison_ and by American zoölogists _Mustela vison_.

EXPLANATION OF SYSTEMATIC TREATMENT

For each full species there will be found under the account of it the following information: Type, statement of geographic range, selected characters for ready recognition, other characters of the species, a summary of geographic variation, and information on habits, in the order mentioned.

For each subspecies, information is presented in the following order: earliest available zoölogical name, synonyms, type, geographic range, zoölogical characters for ready recognition, description (mentioning size, certain external features including color, the skull and teeth) historical material when warranted, remarks which may elaborate on points made in preceding paragraphs, and other information thought to be useful, and finally a list of specimens examined.

In explanation of certain of these categories it should be said that in the synonymy no attempt is made to list every published reference to the subspecies concerned. It is aimed, however, to include at least one citation to each name-combination that has been applied, to the subspecies concerned, along with other especially important references. Mere records of occurrence are not regarded as especially important and citations to them ordinarily are omitted in the synonymy. No comma is placed between the zoölogical name and the name of the author who coined and first used the name in accordance with the rules of zoölogical nomenclature. Otherwise a comma is interposed between the zoölogical name and the name of the user (author). When the accepted (earliest available) name of a subspecies at the head of any one of the following accounts is combined with a generic name different from that with which it originally was placed, the authority for the name is set in parentheses. The same rule is followed with the name of a full species when it is written without any subspecific name following. Parentheses in such situations, therefore, denote that for the terminal part of the scientific name there has been a change in generic name with which the terminal part of the scientific name is here associated.

In the paragraph headed "characters for ready recognition," only a few characters, namely, those regarded as most useful for identification when the student has limited time, are mentioned. Other features useful for distinguishing the kind of animal in question from its near relatives are to be found in the description and comparisons.

In the description, external measurements, unless otherwise indicated, are those recorded by the collector on the label attached to the skin. Total length is the distance from the tip of the pad on the nose to the tip of the fleshy part of the tail when the relaxed animal is laid out straight, not stretched. This measurement does not include the hairs that project beyond the end of the fleshy part of the tail. Length of tail is the distance from the base of the tail, when it is bent at right angles to the long axis of the body, to the tip of the fleshy part of the tail excluding the hairs that project beyond the fleshy part of the tail. Length of tail and length of tail-vertebrae are synonymous. Length of hind foot is measured from the proximal end of the calcaneum to the tip of the longest claw.