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Chapter V: Introduction (4)

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1772. =nivalis= (_Mustela_), Forster, Philos. Trans., London,
1772:373. This is one of the early applications of this name to
American weasels of small size, made in the belief that they were
taxonomically the same in America and Europe. Linnaeus, Syst. Nat.
(12th ed.) 1:69, 1766 is the authority for the name [_Mustela_]
_nivalis_, and the Province of Vesterbotten, Sweden, is regarded
as the type locality. The name is in use today for the common
weasel of Europe and parts of Asia. Animals of the species
_nivalis_ are intermediate in size between _Mustela erminea_ and
_Mustela rixosa_. The name as used for American animals by some
authors who wrote later than Forster did, probably was composite
in that these authors may have applied the name to the small
weasels of North America and thus may have intended it to apply
not only to _Mustela erminea cicognanii_ but also to females of
_Mustela frenata noveboracensis_, and conceivably to both sexes of
_Mustela rixosa_ of any American subspecies.

1813. =Brasiliensis= (_Mustela_) Sevastianoff, Mem. Acad. Imp.
Sci. St. Petersburg, 4:356-363, table (= plate) 4. This name was
proposed for a weasel brought to St. Petersburg by Capt.
Krusenstern on his return from a voyage around the world. The
animal was said to have come from Brazil, but to judge from the
description, came instead from México, Central America, or west of
the Andes in South America, and was based on some one of the
subspecies of _Mustela frenata_. Although the name was in use for
more than 60 years it was shown by Merriam (1896:27) to be
unavailable because it was preoccupied by _Mustela brasiliensis_,
a name earlier used by Gmelin (Syst. Nat., ed. 13, p. 93, 1788)
for a South American otter.

1815. =vulgaris= (_Mustela_), Ord, Guthrie's Geography as
reprinted by Rhoads in 1894, vol. 2, p. 291. This use by Ord is
one of the earliest applications of this name to American weasels,
in the belief that the smaller weasels of North America and Europe
were zoölogically the same; [_Mustela_] _vulgaris_ seems
originally to have been proposed in 1777 by Erxleben on p. 471 of
vol. 1 of his Syst. Regni Anim., for the weasel of the temperate
part of Europe and to be a synonym of _Mustela nivalis_ Linnaeus
(1766). Probably the name as used by Ord was composite in the
sense that he may have intended it to apply to females of _Mustela
frenata noveboracensis_ as well as to one or both sexes of
_Mustela erminea cicognanii_ and, if he ever saw them, to the two
sexes of _Mustela rixosa_ (one or several subspecies).

1818. =africana= (_Mustela_) Desmarest [= _Mustela africana
africana_], Nouv. Diction, d. Hist. Nat., 19:376. In 1808 E.
Geoffroy St.-Hilaire visited Portugal and was given several
African primates and the specimen of _Mustela_ named by Desmarest
in 1818 who wrongly supposed that it, like most of the primates,
came originally from Africa. After the name had been misapplied
for 95 years Angel Cabrera showed that it pertained instead to the
tropical weasel of Brazil. Of distinctive names applied to
American weasels today, this is the one first proposed.

1832. =frenata= (_Mustela_) Lichtenstein [= _Mustela frenata
frenata_], Darstellung neuer oder wenig bekannter Säugethiere, pl.
42 and corresponding text unpaged. This name is the first one
available for the long-tailed weasel and therefore applies to the
species as a whole.

1838. =Cicognanii= (_Mustela_) Bonaparte [= _Mustela erminea
cicognanii_], Charlesworth's Mag. Nat. Hist., 2:38. The name
erroneously spelled _Cigognanii_ was correctly spelled on page 39.
For a detailed consideration of this name see the account of the
subspecies _cicognanii_ on page 120.

1838. =Richardsonii= (_Mustela_) Bonaparte [= _Mustela erminea
richardsonii_], Charlesworth's Mag. Nat. Hist., 2:39. Until 1896
the name sometimes was applied to the subspecies now known as _M.
e. arctica_ and sometimes to part of the subspecies now designated
as _M. e. cicognanii_ under the principal treatment of which see
(page 120) for a detailed account of the basis of the name
_=richardsonii=_, and the reasons for regarding Fort Franklin as
the type locality.

1838. =longicauda= (_Mustela_) Bonaparte [= _=Mustela frenata
longicauda=_], Charlesworth's Mag. Nat. Hist., 2:39. The type
locality appears to be Carlton House, Saskatchewan, and the name
always seems to have been applied to the long-tailed weasel of the
Great Plains, although in some earlier accounts the name was used
in a more inclusive sense to refer also to animals now of
subspecies closely allied to _longicauda_. As with the two
preceding names, a detailed consideration of the basis for, and
application of, this name is given on pages 120-123 in the account
of _Mustela erminea cicognanii_.

1840. =Noveboracensis= (_Putorius_) Emmons [= _Mustela frenata
noveboracensis_], Quadrupeds of Mass., p. 45. This name was
credited by Emmons to De Kay who in the same year published it in
his report on the "Zoology of New York" but without a description
and De Kay's name is a _nomen nudum_. Emmons' was the first use of
the name accompanied by a recognizable description and therefore
the name must date from Emmons although this obviously was not his
intent since he credited the name to De Kay.

1842. =fuscus= (_Putorius_) Audubon and Bachman [= _Mustela
frenata noveboracensis_], Jour. Acad. Nat. Sci., Philadelphia, 8:
(pt. 2) 288.

1842. =pusilla= (_Mustela_) De Kay [= _Mustela erminea
cicognanii_], Nat. Hist. of New York, Zool., Pt. 1, Mammalia, p.
34. This name was proposed for small weasels of 12 to 13 inches in
length of which the tail amounted to a fourth of the same and
although obviously applying in considerable part to the earlier
named _M. e. cicognanii_ seems to have included some individuals
of the also earlier named _M. f. noveboracensis_.

1843. =xanthogenys= (_Mustela_) Gray [= _Mustela frenata
xanthogenys_], Ann. and Mag. Nat. Hist., 11:118, February, 1843,
was applied to all of the long-tailed weasels of California that
had light-colored facial markings. Merriam in 1896 suggested that
San Diego was the type locality and in 1899 Bangs proposed the
name _mundus_ for the California weasel north of San Francisco Bay
thus restricting the application of the name _xanthogenys_. In
1936 Hall further restricted the application of the name and
applied it to the long-tailed weasel of the big interior valley of
California, pointing out that the name was correctly applied to
this weasel of the big interior valley or possibly instead to the
race named _munda_.

1844. =agilis= (_Mustela_) Tschudi [= _Mustela frenata agilis_],
Untersuch. ü. die Fauna Peruana, p. 110, is a name applied today
to the race of weasel of the Temperate Zone of the western Andes
and intermountain valleys of Perú.

1851. =nigripes= (_Putorius_) Audubon and Bachman [= _Mustela
nigripes_], Quadr. N. Amer., 2:297, 1851, applies to the
black-footed ferret of North America.

1853. =agilis= (_Putorius_) Audubon and Bachman [= _Mustela
frenata noveboracensis_], Viv. Quadrupeds N. Amer., 3:184, pl.
140. This name was proposed for the female in the mistaken belief
that it was specifically distinct from the larger male for which
several names already were available. Also Tschudi in 1844 had
already used the name _Mustela agilis_ for a South American
weasel.

1864. =aureoventris= (_Mustela_) Gray [= _Mustela frenata
aureoventris_], Proc. Zoöl. Soc. London, 1864:55, pl. 8, February
9, 1864, is the name applicable to the dark-colored weasel of the
Pacific coastal region of Ecuador and Columbia.

1865. =americana= (_Mustela erminea_ Var. 3) Gray, Proc. Zoöl.
Soc. London, 1865:111. The larger individuals of American weasels
of both _Mustela erminea_ and _Mustela frenata_ from the Atlantic
Coast to as far west as Carlton House, Saskatchewan, were lumped
under this name because Gray desired more information than he then
had before recognizing as different from one another several
species proposed for America up to the time concerned. The name is
unavailable because it is preoccupied by _Mustela americana_
Turton (1806) the name for the American marten.

1865. =americana= (_Mustela vulgaris_ Var.) Gray, Proc. Zoöl. Soc.
London, 1865:113. Under this name the smaller weasels of the
northern and northeastern part of North America were lumped by
Gray but the name is preoccupied and can be ignored.

1874. =affinis= (_Mustela_) Gray [= _Mustela frenata affinis_],
Ann. and Mag. Nat. Hist., 14 (ser. 4):375, 1874, from New Granada
[= Colombia], had the type locality restricted to Bogotá,
Colombia, by Allen in 1916, and is applied to the long-tailed
weasel of the tropical and temperate zones of the eastern Andes of
Colombia.

1874. =macrura= (_Mustela_) Taczanowski [= _Mustela frenata
macrura_], Proc. Zoöl. Soc. London, for 1874, p. 311, pl. 48, May
19, 1874, applies to the long-tailed weasel of central Perú and
northern Ecuador.

1877. =culbertsoni= (_Putorius_) Coues [= _Mustela frenata
longicauda_], Fur-bearing animals ..., p. 136, 1877, is based on
specimens from Fort Laramie, Wyoming. In the past the name has
been regarded as a _nomen nudum_ but there is some reason for
regarding it as having nomenclatural status. In either event it is
here arranged as pertaining to the long-tailed weasel of the Great
Plains which takes the prior name _longicauda_. See the account of
_longicauda_ for a more detailed account of the name
_culbertsoni_.

1877. =aequatorialis= (_Putorius_ (_Gale_) _brasiliensis_) Coues
[= _Mustela frenata aureoventris_], Fur-bearing animals ..., p.
142. Proposed "merely as a substitute for Gray's [supposedly]
preoccupied name," _aureoventris_.

1881. =stolzmanni= (_Mustela_) Taczanowski [= _Mustela africana
stolzmanni_], Proc. Zoöl. Soc. London, for 1881, p. 835, November
15, 1881, is applied to the tropical weasel of the Upper Amazon
Basin.

1881. =jelskii= (_Mustela_) Taczanowski [= _Mustela frenata
macrura_], Proc. Zoöl. Soc. London, for 1881, p. 647, May 17,
1881, was proposed for the female in the mistaken opinion that it
was specifically distinct from the larger male which the same
author previously had named _macrura_.

1891. =arizonensis= (_Putorius_) Mearns [= _Mustela frenata
arizonensis_], Bull. Amer. Mus. Nat. Hist., 3:234, June 5, 1891,
until 1936 was applied to long-tailed weasels of most of the
western United States west of the Great Plains but by restriction
since 1936 has been applied only to the animals in parts of
Arizona and New Mexico.

1894. =peninsulae= (_Putorius_) Rhoads [= _Mustela frenata
peninsulae_], Proc. Acad. Nat. Sci. Philadelphia, 1894:152, June
19, 1894, applies to the weasel of central and southern Florida.

1896. =alascensis= (_Putorius richardsonii_) Merriam [= _Mustela
erminea alascensis_], N. Amer. Fauna, 11:12, June 30, 1896, with
type locality at Juneau, Alaska, has been used for the ermine of
southeastern Alaska ever since it was proposed. In 1944 separate
subspecific rank was accorded ermines on several of the islands of
southeastern Alaska which proportionately restricted the range
assigned to _alascensis_.

1896. =streatori= (_Putorius_) Merriam [= _Mustela erminea
streatori_], N. Amer. Fauna, 11:13, June 30, 1896, applies to the
ermine of the Pacific Coast from Puget Sound, Washington, south
nearly to the Golden Gate of California.

1896. =arcticus= (_Putorius_) Merriam [= _Mustela erminea
arctica_], N. Amer. Fauna, 11:15, June 30, 1896. Ever since it was
proposed, this name has been applied to the subspecies of ermine
of Alaska and the northern parts of Canada.

1896. =kadiacensis= ([_Putorius arcticus_]) Merriam [= _Mustela
erminea kadiacensis_], N. Amer. Fauna, 11:16, June 30, 1896, is a
valid name applied to the ermine of Kodiak Island, Alaska.

1896. =washingtoni= (_Putorius_) Merriam [= _Mustela frenata
washingtoni_], N. Amer. Fauna 11:18, June 30, 1896, applies to the
long-tailed weasel of the southern Cascades of Washington and the
northern Cascades of Oregon.

1896. =saturatus= (_Putorius_) Merriam [= _Mustela frenata
saturata_], N. Amer. Fauna, 11:21, June 30, 1896, was little used
until 1936 but applies to long-tailed weasel of limited region in
northern California and southern Oregon.

1896. =alleni= (_Putorius_) Merriam [= _Mustela frenata alleni_],
N. Amer. Fauna, 11:24, June 30, 1896, applies to weasel of Black
Hills region.

1896. =oregonensis= (_Putorius xanthogenys_) Merriam [= _Mustela
frenata oregonensis_], N. Amer. Fauna, 11:25, June 30, 1896,
applies to long-tailed weasel of parts of western Oregon and
northern California.

1896. =goldmani= (_Putorius frenatus_) Merriam [= _Mustela frenata
goldmani_], N. Amer. Fauna, 11:28, June 30, 1896, applies to the
long-tailed weasel of Chiapas, and parts of Guatemala and
Salvador.

1896. =leucoparia= (_Putorius frenatus_) Merriam [= _Mustela
frenata leucoparia_], N. Amer. Fauna, 11:29, June 30, 1896,
applies to the long-tailed weasel of Michoacán and Nayarit.

1896. =tropicalis= (_Putorius_) Merriam [= _Mustela frenata
tropicalis_], N. Amer. Fauna, 11:30, June 30, 1896, applies to the
long-tailed weasel of the Tropical Life-zone of Veracruz.

1896. =spadix= (_Putorius longicaudus_) Bangs [= _Mustela frenata
spadix_], Proc. Biol. Soc. Washington, 10:8, February 25, 1896,
applies to the long-tailed weasel of Minnesota and adjoining
areas.

1896. =rixosus= (_Putorius_) Bangs [= _Mustela rixosa rixosa_],
Proc. Biol. Soc. Washington, 10:21, February 25, 1896, applies to
the least weasel of Saskatchewan and adjoining areas and as the
first available name for the species has been used as the specific
name for the species in America since 1896.

1897. =paraensis= (_Putorius (Mustela) braziliensis_) Goeldi [=
_Mustela africana africana_], Zool. Jahrb., abt. f. systematik,
geogr. u. biol., 10:560, pl. 21, September 15, 1897, a synonym for
the weasel of the lower Amazon area.

1898. =neomexicanus= (_Putorius frenatus_) Barber and Cockerell [=
_Mustela frenata neomexicana_], Proc. Acad. Nat. Sci.
Philadelphia, p. 188, May 3, 1898, applies to the long-tailed
weasel of New Mexico, Arizona, Durango and adjoining areas.

1898. =haidarum= (_Putorius_) Preble [= _Mustela erminea
haidarum_], Proc. Biol. Soc. Washington, 12:169, August 10, 1898,
applies to the ermine of the Queen Charlotte Islands, British
Columbia.

1899. =notius= (_Putorius noveboracensis_) Bangs [= _Mustela
frenata noveboracensis_], Proc. New England Zoöl. Club, 1:53, June
9, 1899, was applied to the long-tailed weasel of the Carolinas
until 1936 since which time it has been regarded as a synonym of
_noveboracensis_.

1899. =occisor= (_Putorius_) Bangs [= _Mustela frenata occisor_],
Proc. New England Zoöl. Club, 1:54, June 9, 1899, applies to the
long-tailed weasel of central and northern Maine. Until 1936,
occisor was ordinarily used as the name of a full species but
since then has been arranged as a subspecific name under _Mustela
frenata_.

1899. =mundus= (_Putorius xanthogenys_) Bangs [= _Mustela frenata
munda_], Proc. New England Zoöl. Club, 1:56, June 9, 1899, is now
applied, and generally has been since 1899, to the long-tailed
weasel of the coastal district of California north of San
Francisco Bay.

1899. =muricus= (_Putorius (Arctogale)_) Bangs [= _Mustela erminea
muricus_], Proc. New England Zoöl. Club, 1:71, July 31, 1899,
applies to the diminutive ermine, often erroneously designated
least weasel, of the western United States.

1899. =oribasus= (_Putorius (Arctogale) longicauda_) Bangs [=
_Mustela frenata oribasus_], Proc. New England Zoöl. Club, 1:81,
December 27, 1899, applies to the long-tailed weasel of the Rocky
Mountains northward from Yellowstone National Park.

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.

Capitalized color terms, unless otherwise indicated, refer to Ridgway's (1912) _Color Standards and Color Nomenclature_. Some use is made of color terms taken from Oberthür and Dauthenay (1905) because those authors show a much larger number of shades between dark brown and black than does Ridgway (1912). The colors of the upper parts of most weasels are some shade or other of dark brown. Color terms that do not have the initial letter capitalized do not refer to any one standard and consequently are used in a general sense.

Relative extents of the color of the upper parts and underparts are computed from measurements of the circumference of the body at the place where the color of the underparts is narrowest. Ordinarily this place is in the lumbar region rather than in the thoracic region.

An explanation of how cranial measurements were taken is given on page 417. In designating teeth, capital letters are used for teeth in the upper jaw and lower case letters are used for teeth in the lower jaw. For example: I2 denotes the second incisor tooth in the upper jaw and i2 denotes the second incisor tooth in the lower jaw; C1 and c1 refer to the canine tooth of the upper jaw and lower jaw, respectively; P3 and p3 refer to the third premolar of the upper jaw and lower jaw, respectively, bearing in mind that the first (anterior) premolar is absent in the lower jaw and upper jaw of weasels (see fig. 31 on page 416), as also, in some kinds of weasels, is the second premolar; M1 and m1 refer to the first molar of the upper jaw and lower jaw respectively.

In describing the skull and teeth the two sexes are treated separately because differences in shape as well as size are the rule. Unless otherwise indicated, the skulls on which descriptions are based are of adults. Weights of skulls include the weight of the lower jaws. In general, every second subspecies is described. For a subspecies geographically next adjacent to the one described, only the differences between the two are enumerated. This method of description indicates also likenesses and is more economical of words than some other methods of description. Also, by use of this method, cross reference is reduced to one other subspecies. Following this formal description, there is a comparison of the cranial and dental characters with those of geographically adjacent subspecies.

In the paragraph headed "Remarks" the two words "character" and "structure" frequently appear. The word structure here is used to mean some part of an animal, as for example, a hair, a muscle, a bone, or an internal organ. A structure is not a system, as for example, the digestive system or osseous system. A character is some weight, linear dimension, volume, shape, color, or other perceptible attribute of a structure, of a system, or of an entire organism.

In recording the localities of capture of specimens examined, effort has been made to be exactly as precise as the locality data on the labels of the specimens permit. The word "County" is written out in full when the name of the county is written on the label of each specimen listed from that county. When one specimen, or more, here assigned to a given county lacks the name of the county on the label, then the abbreviation "_Co_." is used. The surprising frequency with which the same place name is repeated in a given state or province makes it desirable for the collector to write the name of the county, or corresponding minor political subdivision, on labels of study specimens at the time they are prepared.

SYSTEMATIC ACCOUNTS OF SPECIES AND SUBSPECIES

=MUSTELA ERMINEA= Linnaeus

Ermine

(Synonymy under subspecies)

_Type._--_Mustela erminea_ Linnaeus, Systema Naturae, 10th ed., p.
46, 1758.

_Range._--From the British Isles and Atlantic Coast of Europe
across Eurasia and North America including Greenland, from the
northernmost land, south, in North America, to the lower margin of
the Canadian Life-zone; geographically south to Connecticut, New
York, Pennsylvania, northeastern Ohio, southern Michigan,
Wisconsin, northern Iowa, Minnesota, North Dakota, in the Rocky
Mountains to northern New Mexico, in the Sierra Nevada to Mono
County, California, and on the Pacific Coast to the Golden Gate.

_Characters for ready recognition._--Differs from _Mustela rixosa_ in presence of black pencil on tail, tail-vertebrae more than a fourth of length of head and body, and in regions where the two species occur together, basilar length of skull more than 32.5 in males and more than 31.0 in females; from _Mustela frenata_, in regions where the two species occur together, by tail less than 44 per cent of length of head and body and by postglenoidal length of skull more than 46 per cent of condylobasal length in males and more than 48 per cent in females.

_Characters of the species._--Size medium to small (total length 225 to 340 mm. in males and 190 to 290 mm. in females); tail 30 to 45 per cent of length of head and body, with distinct black pencil; caudal vertebrae 16 to 19; skull with long braincase and short precranial portion; postglenoidal length, when expressed as a percentage of the condylobasal length, more than 48 in females and ordinarily more than 46 in males; upper parts brown; underparts whitish, ordinarily continuous from chin to inguinal region but in subspecies in the humid region along the Pacific Coast interrupted in some individuals by brown of upper parts encircling body in the abdominal region. The soles of the feet in each of the subspecies are densely haired in winter and have only a relatively small area of the foot-pads exposed in summer, the intervening areas being well haired even at that season. The uniformity throughout the species as regards hairiness of the foot-soles and also the character of the vibrissae makes it unnecessary to describe these features in the accounts of the subspecies of _erminea_.

_Geographic variation._--In the Old World 16 or more subspecies are currently recognized and there are 20 in North America. The features in which geographic variation is especially prominent are: First, size, as expressed by external measurements and weight, second, color pattern, depending on the extent, in relation to one another, of the dark-colored upper parts and light-colored underparts, and third, breadth and depth of the rostral region of the skull. Except in size, the variation in the skull is less than in _M. frenata_. Likewise in tone and shade of upper parts and hue or tint of underparts, _erminea_ is less variable than _frenata_ and has the face all of one color without the contrasting color-pattern of the face and head seen in many subspecies of _frenata_. _M. erminea_ exceeds _frenata_ as regards variation of the size of the area occupied by the light-colored underparts. At one extreme is the subspecies _arctica_ in which the area of the light color extends well up on the sides of the body, down the insides of the legs, over the feet and far out on the lower side of the tail whereas at the other extreme are the races _streatori_ and _olympica_ in which the light-colored underparts are restricted to two areas, one on the chin, throat and chest, and the other on the inguinal region. These areas may or may not be connected by a thin line of white color along the midline of the underparts. In size of animal, _erminea_ probably exhibits the maximum variation among American species of weasels; an average-sized male of the race _arctica_ weighs 4 times as much as one of the race _muricus_, and in the species _frenata_ I doubt that the difference is quite as great between individuals of the smallest race, _effera_, on the one hand, and either of the largest races, _texensis_ or _macrophonius_, on the other hand although actual weights are not available for these races of _frenata_. As elsewhere indicated, the small-sized individuals of _M. erminea_ are of the southern races and the large-sized individuals are of the northern races. This decrease in size southward occurs both in Asia and in America.

_Natural history._--habitat and numbers.--Along the International Boundary east of the Turtle Mountains, Soper (1946:136) found this species present only in timbered areas and absent from many untimbered areas. Of the same species to the westward he comments "so far as I know at present, there is no evidence to show that any short-tailed weasels inhabit a broad strip of treeless territory immediately north of the International Boundary in Canada from southwestern Alberta to southeastern Saskatchewan." The same author (1942) reports that in the general area of Wood Buffalo Park, Northwest Territory, south of Great Slave Lake, the ermine is uncommon on pine-grown sand ridge and rolling upland and common in lower spruce-aspen parklands, stream-side coniferous belts, and grassy, semi-wooded swamplands.

Nine ermines per square mile is the number that Soper (1919:46-47) estimated at Edmonton on the basis of the numbers that he trapped there in the winters of 1912-13 and 1913-14 and on the basis of the tracks of remaining ermines. From corresponding data he estimated the population in the winter of 1913 on the Hay River, north of Jasper Park, to be nine per square mile. In each of these instances he estimated ten weasels per square mile but he inclined to the view that one-tenth of the animals involved in his counts were long-tailed weasels (_Mustela frenata_). Osgood (1909B:30) and his field companion in the period July 31 to September 3, 1903, took a series of 42 specimens within a radius of 500 yards of their camp at the head of Seward Creek, Alaska, all caught in four traps, in one month. Of the 42 specimens, 28 are males and 14 are females.

Fluctuations of a multiannual nature are marked in this species. Bailey (1929:156) observes that in Sherburne County, Minnesota, when meadow mice are abundant for two or three years these weasels become abundant but that when the mice are scarce the weasels also become scarce. Manning (1943:56), on Southampton Island, noted "that the maximum and minimum points of the weasel cycle are much more sharply marked than those of the fox cycle and the increase and decrease are more rapid."

How far an ermine will travel in a given length of time has seldom been recorded but Hamilton (1933:293), on March 20, 1932, "followed the track of a small weasel, presumably a male _cicognanii_, for four miles in the fresh snow," and Ingles (1942) observed a diminutive ermine of the subspecies _M. e. muricus_, at Woods Lake, California, 286 yards from its den.

Behavior

As regards locomotion, Soper (1919:46), in reference to _Mustela cicognanii_, presumably in Ontario, Canada, writes that in the bounding gait the hind feet register almost, if not exactly, in the front-foot impressions, with the right front and hind feet lagging slightly behind. "The distance normally is about 19 inches, representing a regular rate of travel. . . . In traversing open spaces they resort to long, graceful leaps upwards of six feet in length. . . . I measured a record . . . of 8 feet, 2 inches."

Of _M. e. arctica_, Dice (1921:22) writes that when it runs "the tail is carried off the ground usually at an angle of about 45 degrees." Seton (1929 (2):598) states that "At Carberry [Manitoba] I have often seen this energetic little creature seeking for Mice in the deep, soft snow. Its actions are much like those of an Otter pursuing salmon. Sometimes it gallops along a log, or over an icy part of the drift; then plunges out of sight in a soft place, to reappear many yards away. . . ."

Little is recorded concerning swimming but on this score Seton (1929 (2):602) does quote J. W. Curran, who in July, 1899, at Lake Couchiching, Ontario, watched an ermine pursue a chipmunk into the water and for 100 yards before giving up the chase and wheeling around and making for shore. In swimming "The Weasel, I think, showed more of his body, and seemed to exert himself more" than the chipmunk.

As to voice, Dice (1921:22), at Tanana, Alaska, heard the ermine, when excited, bark somewhat like a mink but not so loud and Seton (1929 (2):606) quotes Manley Hardy to the effect that the species has a purring note.

Sense of smell was used by an _M. e. muricus_ that Dixon (1931:72) watched as the ermine followed a three-fourths-grown pika. Concerning the ermine at Carberry, Manitoba, Seton (1929 (2):598-599) writes that "The smell of blood must be as far-reaching as it is attractive to these sanguinary little creatures. I have frequently hung new-killed Rabbits and partridges temporarily in trees, and, after an absence, in some cases of a few minutes only, have found an Ermine mauling the game, though there was no sign of such a visitor when the cache was made."

Enemies

George Measham, of Winnipeg, found sign in the snow indicating that a great snowy owl had killed an ermine and T. McIlwraith shot a bald eagle at Hamilton Bay which had the bleached skull of a weasel (probably of this species) clinging to the throat (Seton, 1929 (2):603).

A. B. Howell (1943:98) likens mustelid mammals to domestic cats in their manner of crossing roads and thinks that mustelids loiter at the side of the road until the stimulus of the approaching car causes them to make a dash whereupon they are caught by the wheels and killed. Three of four weasels seen to cross the road were killed, one even having apparently crossed the road before turning back and being killed under the car. One weasel killed was _Mustela erminea cicognanii_. Dalquest (1948:190) in writing of this species in the state of Washington, says "I have seen only one abroad in the daytime. It dashed from a roadside thicket . . . and was crushed beneath the wheels of a car."

Food

The killing of prey is described by Hamilton (1933:332) as follows: "A rapid dash, and the bird or mouse is grabbed over the back of the skull, the fore legs encircle the animal as though hugging it, and the hind legs are brought up to scratch wildly at the captive. . . . If [the prey is] a large animal, as a rat, the weasel usually lies on its side, while the diminishing struggles of the rodent continue, but if a mouse or a small bird [is the object of attack], the weasel is apt to crouch over its prey. Little time is lost over the first [mouse] . . . if two mice are present [;] a strong bite through the brain case . . . [is] sufficient. If only one animal is present, the weasel dawdles over its kill some time after life has departed."

Hamilton's (1933:333) study of the contents of the digestive tracts of bodies of ermines obtained from fur trappers and fur buyers in New York enabled him to publish the following "Frequency Indices of Mammal Genera in Fall and Winter Food of 191 Mustela cicognanii": _Microtus_, 35.7 per cent; mammals undetermined to genus but principally mice, 16.3; _Blarina_, 15.1; _Peromyscus_, 11.4; _Sylvilagus_, 9.0; _Sorex_, 4.9; _Rattus_, 4.4; _Tamias_, 3.6. Close correspondence is shown by the following data of Aldous and Manweiler (1942) for the ermine from Lake of the Woods, Minnesota: mice, 58.7 per cent by number and 54.5 by volume; shrews 22.5 and 21.8 per cent; birds, 2.7 and 5.0 per cent. Of the mice in stomachs, 40 per cent were microtines, 15 per cent were _Peromyscus_ and 45 per cent were unidentified as to kind. Fragments of a small fish were found in one stomach. Summed up, the dominant winter foods were mice and shrews. Trapping of the mammal populations was done to see what the available food was and it was found that the small mammals were eaten in direct ratio to their relative abundance. Snowshoe rabbits and red squirrels were not eaten. The Minnesotan data were from 60 stomachs and 53 intestinal tracts recovered from 129 weasels trapped by use of scent (not bait) mostly from January 1 to February 7, 1939, although a few were trapped in 1938. Analyses of contents from stomachs gave approximately the same results as those from intestines. In 1939 at Lake of the Woods, weasels were concentrated where food was abundant but no such concentration was noted in the following winter.

Big short-tailed shrew (_Blarina brevicauda_).--In New York State, the ermine preys on _Blarina_ as shown by Hamilton's (1933:330) seeing one being carried by a male ermine on May 6, 1931, and another being carried by a female on May 13, 1932. The same author (1928:249) found the remains of a _Blarina_ in a small female from Malone, New York. Kirk (1921) observed, however, that the ermine (_M. e. cicognanii_) avoided the shrew, _Blarina_, caught in a trap and that _Blarina_ avoided the weasel caught in a trap.

Chipmunk (genus _Tamias_).--Remains were found in a male ermine in New York on May 14, 1932 (Hamilton, 1933:330), and Seton (1929 (2):602) records a chipmunk at Lake Couchiching, Ontario, that was pursued into the water by an ermine.

Deer mice (genus _Peromyscus_).--As shown by Hamilton (1933:33) and Aldous and Manweiler (1942), _Peromyscus_ was second only to microtines in numerical abundance among the food items of ermines in New York and Minnesota. _Peromyscus_ and microtine rodents were brought to a den of the diminutive _M. e. muricus_ in early August, in Fresno County, California, according to Ingles (1942). He observed that an Alpine chipmunk was active under and around the tree and that juncos reared young 40 feet from the den but that the chipmunk and juncos were unmolested by the ermines.

Lemming (genus _Lemmus_).--One was recovered from a female ermine (with milk in her glands) at Laurier Pass, British Columbia (Sheldon, 1932:201).

Red-backed mouse (genus _Clethrionomys_).--Criddle and Criddle (1925:146) record that on "May 31, 1921.--Saw a Bonaparte's weasel capture a Red-backed Vole after a long hunt during which the pursuer never once lost track of its victim."

Meadow mice (genus _Microtus_).--As shown by the data of Hamilton (1933:333) and Aldous and Manweiler (1942) recorded above, _Microtus_ is the item of first importance in the diet of the ermine in New York and Minnesota. Criddle and Criddle (1925:146) write concerning the vicinity of Treesbank, Manitoba, that "October, 1918.--Following a severe outbreak of mice in 1916-17, Bonaparte's weasel increased enormously and very soon reduced the rodents to comparative rarity. This resulted in a scarcity of food for the weasels, which in their turn became greatly reduced in numbers."

Old World rat (_Rattus_).--Bishop (1923) found two headless rats near a nest of this species in Albany, New York.

Pika (_Ochotona_).--Dixon (1931:72) at Milner Pass, Colorado, on July 20, 1931, saw an ermine, of the subspecies _muricus_, following a three-fourths grown pika by scent and outrunning the pika. The pikas worked a relay system and the weasel abandoned the trail when the fourth pika became the object of the chase.

Cottontail (genus _Sylvilagus_).--Hamilton (1933:33), as noted above, found remains of cottontail in the digestive tracts of ermine that had been trapped for fur in winter. Possibly these remains were bait that had been placed at traps.

Snowshoe rabbit (_Lepus americanus_).--Morse (1939:210) in a study of predation on hares and grouse in the period of notable decimation of these two game species in 1935-1936 in the Cloquet Valley State Forest, in St. Louis County, Minnesota, found that "weasel predation on hares appeared to be of very low incidence or altogether lacking."

Wild birds (Class Aves).--Aldous and Manweiler (1942), as noted above, found that the remains of birds constituted five per cent by volume of the food of the ermine in winter in Minnesota.

Chicken (genus _Gallus_).--Criddle and Criddle (1925:145), who published relatively extensive data on the three species of weasels of Manitoba, write that: "We have no record of Bonaparte's weasel killing poultry, and we doubt whether it ever does so." However, Soper (1919:46) investigated the excited cackling of a hen brooding chicks at night and found a solitary ermine that had killed three chicks and that had the remainder under very active scrutiny.

Leopard frog (_Rana pipiens_).--One frog was found in a male ermine on November 20, 1931, in New York by Hamilton (1933:300).

Fish (Class Pisces).--Aldous and Manweiler (1942) found fragments of a small fish in one of 60 stomachs of ermine from Minnesota.

Earthworm (Phylum Annelida).--Osgood (1936:64), presumably at Rutland, Vermont, observed a pair of weasels from 2:15 P.M. to 5:00 P.M., in a barn and saw the female in that time make many trips for food for her young. Only earthworms were brought. Fifty traps in an adjacent, swampy field caught only one bull frog and no mice indicating that mice had been eliminated from the foraging territory of the ermine.

In handling food, Dice (1921:22) noted that the Alaskan ermine did not use the feet but only the mouth.

Reproduction

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American WeaselsChapter V: Introduction (4)

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