American Weasels :: HISTORY OF CLASSIFICATION

HISTORY OF CLASSIFICATION

In the earlier accounts of American weasels, from the time of Linnaeus and before, up until 1890, names then in use for European weasels frequently were applied also to those in North America. For the next 50 years, and almost without exception after 1896, the American weasels were regarded as specifically distinct from those in the Old World. In this 50-year period many new names were proposed, usually as full species, although now that material from more localities has been brought together and studied, geographic intergradation is evident between many of the named kinds and most of these names now therefore take only subspecific rank. In 1933 Glover M. Allen showed that Mustela rixosa occurred also in the Old World, and in 1943 I emphasized that a second American species, Mustela erminea, was circumpolar in distribution. In neither rixosa, nor erminea, however, were the subspecies the same in the two continents. To this general outline of the nomenclature, exception must be made for weasels of the southwestern United States, MÉxico and Central America, and South America, because as early as 1813 a distinctive name was given to one of these and weasels from the three areas mentioned were, so far as I know, never given names of Old World kinds.

The first paper that could be regarded as revisionary in nature was "Remarks on the species of the genus Mustela" by the zoÖlogist and world-traveler, Charles L. Bonaparte, in Charlesworth's Magazine of Natural History, for 1838. In that paper three new names, Mustela cicognanii, M. richardsonii and M. longicauda, all still valid, were proposed for American weasels.

Audubon and Bachman in their "Quadrupeds of North America," which appeared in parts from 1845 to 1853, recognized 5 species. Actually they were dealing with only 3 taxonomically valid kinds. For one of these, Mustela frenata noveboracensis, they were misled by the difference in size between males and females, and in the males by the presence of a brown coat in some and a white coat in others. The male that was white in winter they regarded as Putorius ermineus of the Old World; the male that was brown in winter they designated by their earlier proposed name P. fuscus, and the female they named P. agilis. The ermine, subspecies M. erminea cicognanii, they called P. pusillus. Their fifth name, P. frenatus, included at least some animals that today are assigned to the subspecies M. frenata frenata. Each of three and perhaps four of the five names employed by Audubon and Bachman embraced individuals of more than one species and in that sense the names were composite.

Only five years later, in 1858, Professor Spencer Fullerton Baird's great work, "The Mammals of North America," made it clear that no American weasel was identical (in the modern subspecific sense) with any Old World weasel, and he applied most of his names in a correct zoÖlogical sense. It is true that he thought that the female weasel of the eastern United States was specifically different from the male, misapplied to it the name richardsonii, and did not correctly allocate every one of the few poor specimens available to him of the little ermine (M. e. streatori) of the Pacific Coast; but he did recognize that the least weasel was a distinct kind and his treatment in general was excellent.

After Baird came a period of great confusion in which most writers did no better than had Audubon and Bachman, ordinarily confusing the two sexes as different species, and, in 1877 in his "Fur-bearing Animals," Elliot Coues went rather to the other extreme and allowed only 4 kinds to all of the Americas, regarding two of these, for purposes of zoÖlogical nomenclature, as identical with the European species.

But, in 1896 Outram Bangs published "A Review of the Weasels of Eastern North America" in which he correctly recognized eight kinds. Although some of these were treated by him as full species, whereas the material accumulated since 1896 has shown that subspecific status is in order, his names, still in use, were correctly applied in every instance, save probably one. This was his use of Putorius richardsonii for the animal now known as M. e. arctica. Unlike the earlier, excellent treatment by Baird, this accurate one by Bangs was heeded and followed by subsequent writers. For example, Dr. C. Hart Merriam in the same year, 1896, accepted Bangs' conclusions except for correcting the application of the name richardsonii. The principal contributions of Merriam's paper "Synopsis of the Weasels of North America" were first, the wider geographic scope and second, the naming as new of several kinds outside the geographic area studied by Bangs. Otherwise the work was not up to Dr. Merriam's usual standard and the internal evidence of haste in its preparation and the superficial study of some of the material at his disposal explain why the weasels of North America since that time have been but little better understood than in 1896. Baird and Bangs, then, unquestionably did the best systematic work on the American weasels.

In 1916 Dr. Joseph A. Allen published a valuable paper on the South American weasels. The material available to him was inadequate and prevented a thoroughly satisfactory treatment. There are too few specimens even today to permit of a thorough treatment of the South American weasels in the present paper; nevertheless the material today is more nearly adequate than it was in 1916 and it is hoped that the systematic arrangement is correspondingly improved.

Chronological List (annotated) of Specific and Subspecific Names Applied to American Weasels

At least eighty-seven specific and subspecific names have been proposed for American weasels. Of these sixty-nine are now regarded as valid designations of recognizable subspecies. The average is 1.2 names per subspecies. Some names in the following chronological list were a second time applied wholly or in part to some other kind of weasel. In general, mention of the second or any other later application is omitted from the following list but two usages of agilis (1844 and 1853) and of americana (1865) are recorded.

1734. javonica (Mustela) Seba, Locupletissimi Rerum naturalium Thesauri ..., 1:77, 78, pl. 48, fig. 4. The weasel to which this name was applied was said to have come from Java. Since no animal answering to the description has again been found in Java, and because specimens from Central America or possibly some from northern India, may do so, it is conceivable that Seba was the first to distinguish by name an American weasel from those in the Old World. My attempts to locate the specimen concerned in places where it might have been preserved along with some of the other specimens thought to have belonged to Seba have been fruitless. Since it is impossible positively to link Seba's description with any known weasel, no further use is made of the name javonica in the present account.
1772. erminea (Mustela) Forster [= Mustela erminea richardsonii], Philos. Trans., London, 1772:373. Forster's use of the name is one of the earliest applications of it to American animals. The name dates from Linnaeus, Syst. Naturae, (10th ed.) 1:46, 1758, with type locality in Europe. In the subspecific sense the name applies to the ermine which occurs over most of the Scandinavian Peninsula, if Miller (1912:387) be followed in regarding the type locality as Upsala, Sweden. If, instead, Cabrera (1913A:394-396) be followed in regarding the type locality as in Switzerland, the name, in the subspecific sense, will apply to the ermine of continental Europe. As the earliest available name applied to the circumpolar species concerned, it is used now as the name of the species in the New World as well as in the Old World. From the time of Forster until approximately 1890 the name erminea by many, but not by all, authors was applied to the American weasels in the belief that they were zoÖlogically indistinguishable from those in the Old World. From 1896 to 1943 the name was not used by American authors at all because the ermine of America was in 1896 treated nomenclaturally by Merriam as specifically distinct from the animal in the Old World. Since 1943 erminea has been used in the specific sense for American animals in recognition of the circumpolar distribution of the species. Some of the early allocations of American specimens to erminea probably resulted in a composite use of the name in that one or another subspecies of the American species Mustela frenata may also have been included with individuals truly of the species erminea.

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.

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