American Weasels (part 3 of 4)
Characters for ready recognition.--Differs: From M. f. olivacea, in males, by width of tympanic bulla which is less than rather than more than 8.5 mm., and in adult females by total length which is less than rather than more than 345 mm. and by mastoid breadth which is less than rather than more than distance between articular faces of exoccipital condyle and glenoid fossa; from M. f. occisor by a number of average differences including smaller size, relatively shorter tail and relatively narrower skull (see measurements); from M. f. spadix by least width of color of underparts amounting to less than 41 per cent of greatest width of color of upper parts, absence of color of underparts on ankles and feet, adults with hind foot less than 50 in males and 40 in females, orbitonasal length less than 15.5 in males and 13.5 in females, length of tooth-rows less than 18.0 in males and 15.7 in females, mastoid breadth less than 25.5 in males and 22.0 in females; from M. f. primulina in males by interorbital breadth averaging more than 24 per cent of basilar length, orbitonasal length averaging more than 34 per cent of basilar length or 64 per cent of mastoid breadth, tympanic bullae less inflated anteromedially, than posteromedially, and in females by orbitonasal length amounting to more than two-thirds of mastoid breadth, by zygomatic breadth averaging less than 21, and by anterolateral margin of tympanic bullae not projecting below squamosal; from M. f. arthuri in males, by zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla and by convex dorsal outline of skull in longitudinal axis.
Description.--Size.--Male and Female:
======================================================================= | Number | | |Per cent| | LOCALITY | of | Total | Length | of | Length of | | specimens | length | of tail | body- | hind foot | | averaged | | | length | | ----------+------------+-----------+-----------+--------+-------------+ | 8 ad. [M] | 415 | 146 | 54% | 46.0 | Massa- | | (390-432) | (127-159) | | (41.0-48.0) | chusetts +------------+-----------+-----------+--------+-------------+ | 4 ad. [F] | 311 | 104 | 50% | 33.9 | | | (298-321) | (95-114) | | (31.5-37.0) | ----------+------------+-----------+-----------+--------+-------------+ | 10 ad. [M] | 411 | 141 | 52% | 47.1 | Liberty | | (379-438) | (124-155) | | (43.0-51.5) | Hill, +------------+-----------+-----------+--------+-------------+ Conn. | 6 ad. and | 318 | 105 | 49% | 33.0 | | sad. [F] | (303-338) | (80-123) | | (31.7-36.0) | ----------+------------+-----------+-----------+--------+-------------+ | 10 ad. [M] | 407 | 130 | 47% | 46.0 | Beaver | | (372-431) | (113-143) | | (42.0-50.0) | Dam, +------------+-----------+-----------+--------+-------------+ Wisc. | 4 ad. [F] | 326 | 99 | 43% | 35.6 | | | (303-338) | (86-108) | | (34.6-38.0) | ----------+------------+-----------+-----------+--------+-------------+ | 10 ad. [M] | 371 | 130 | 54% | 45.0 | Washtenaw | | (350-405) | (115-140) | | (40.0-50.0) | Co., +------------+-----------+-----------+--------+-------------+ Mich. | 10 ad. [F] | 306 | 97 | 46% | 34.0 | | | (290-335) | (90-120) | | (30.0-40.0) | ----------+------------+-----------+-----------+--------+-------------+
The length of the hind foot averages more than the basal length in males whereas the reverse is true in females. The tail, relative to the length of the body, is longer in males than in females. The average differences in external measurements of the two sexes in Massachusetts, are: total length, 104; length of tail, 42; length of hind foot, 12.1. In Michigan, where the males are smaller, corresponding differences are only, 65, 33, and 11. Weight of 19 adult males from New York (Hamilton, 1933:294), 225 (196-267) grams and in 13 adult females, 102 (72-126) grams. Weights of 2 adults from Michigan are: [M] 258; [F] 101 grams.
Externals.--Longest facial vibrissae black, brown, or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 19.
Color.--Upper parts, in summer, Vandyke Brown or darker than tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304. Sometimes approaching tone 2 of Warm Sepia of Oberthür and Dauthenay, pl. 305. Underparts, in summer, ranging from white through Napthalene Yellow (Peterboro, N. Y.), Pale Orange Yellow (eastern Mass.), near Primuline Yellow (unusual specimen from Leelanau Co., Mich.) to near (c) Deep Chrome (no. 19053, U. S. Nat. Mus., Roan Mts., N. C). In winter, all white except tip of tail, or upper parts near (12" 1) Rood's Brown and tone 2 of Raw Umber of Oberthür and Dauthenay, pl. 301, with underparts white or sometimes tinged with yellowish. Tip of tail at all times black. Upper parts of uniform color except for occasional slight darkening of nose. Color of underparts extends distally on posterior sides of forelegs to foot and sometimes over upper sides of toes and on medial sides of hind limbs only to knees. Least width of color of underparts averaging, in a series of twenty-two males, mostly in full winter pelage, from Liberty Hill, Connecticut, 21 (11-40) per cent of greatest width of color of underparts. In eleven females from the same place, corresponding percentages are 20 (14-29). Black tip of tail in same series of males, most of which are in full winter pelage, 70 (60-75) mm. long; thus longer than hind foot and averaging 50 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on ten adults from Massachusetts): See measurements and plates 16-18; weight, 3.6 (3.3-4.4) grams; basilar length, 44.6 (43.3-46.0); zygomatic breadth less than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more or less than length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less (usually more) than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 6 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa behind or directly below posterior fourth of m1.
Female (based on five adults from Mass.): See measurements and plates 31-33; weight, 1.7 (1.2-2.1) grams; basilar length, 36.5 (35.2-38.1); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more or less than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more or less (usually less) than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 4 to 5-1/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum.
The skull of the female averages 53 per cent lighter than that of the average male.
Comparisons of the skull with those of M. f. olivacea, M. f. spadix, M. f. primulina, and M. f. arthuri, are made in the accounts of those subspecies. As compared with that of M. f. occisor the skull of adult male noveboracensis, is of smaller average size with relatively (to basilar length of Hensel) lesser mastoid and zygomatic breadths. In addition to the zygomatic arches of noveboracensis being less widely bowed outward they seem to be more rounded posteriorly. Comparisons of subadult females indicate that these differences exist in the females as well as in the adult males.
Remarks.--The earliest of the post-Linnaean references to this weasel mostly were under the specific name erminea in the belief that the American animal was the same as the larger of the two common species of weasel in the Old World. The name noveboracensis, now in use for this subspecies, was applied in 1840 and since that time the males usually have borne that name; the females, because they are smaller, were more frequently confused with some other species. Audubon and Bachman in 1853 even proposed the name agilis for the female in the mistaken belief that it was a species distinct from the male. After 1896, when Bangs correctly classified the weasels of the eastern United States, the males have been correctly identified and the females, except by a few authors, likewise have been correctly named. Because many early American naturalists did their first collecting of mammals in the geographic range of noveboracensis, the person who examines labels of specimens of this subspecies can find data written in the hand of Spencer Fullerton Baird, Theodore Roosevelt, and other naturalists famous for their work as scientists or accomplishments otherwise. The material is more nearly adequate than is that of many other subspecies and the number of specimens is exceeded--and only slightly--by that of the subspecies nevadensis, which like noveboracensis has a relatively large geographic range.
Intergradation with Mustela frenata spadix is indicated by subadult males from western Wisconsin, namely, one from Gordon, three from Colfax and one from Meridean. Linear measurements of the teeth of these specimens are exactly intermediate between those of spadix from Elk River, Minnesota, to the west, and noveboracensis from, say, Beaver Dam, Wisconsin, to the east. The specimens from western Wisconsin show approach to spadix also in that the length of the tooth-rows and breadth of the rostrum are slightly greater than in noveboracensis from farther east, say, Beaver Dam, Wisconsin.
Indeed, animals from as far east as Beaver Dam itself might be thought of as showing some approach to spadix. Although, along the eastern seaboard, the upper lips, with rare exceptions, are the same color as the underparts, farther west, in Michigan and Wisconsin, the lips more often than not are white. Animals from Beaver Dam have slightly shorter black tips on the tails, broader extent of the light color of the underparts and females average slightly larger than typical noveboracensis, say, those from Massachusetts. Each of these differences reflects characters found better developed in the spadix-longicauda stock to the west.
Toward the southern part of its range where noveboracensis meets M. f. olivacea there is a marked increase in yellowness of the underparts. This coloration of the underparts, since it is not so well marked in the northern part of the range of noveboracensis, might be regarded as showing intergradation with olivacea and primulina, each of which has far more intensely colored underparts than does noveboracensis. Excepting this increase of yellow on the underparts, however, there are few if any characters of noveboracensis which undergo marked change as approach to the range of olivacea is made. Indeed, the characters of noveboracensis remain constant to within a relatively short distance of the geographic range of olivacea.
Notwithstanding the state of affairs described above, intergradation seems to take place. Three specimens referred to noveboracensis but which at the same time are regarded as intergrades with olivacea are as follows: No. 28.300, Charleston Museum, from five miles east of York, South Carolina, is an adult female with a badly crushed skull. In external measurements the specimen agrees with noveboracensis. The underparts, as regards color and width, are intermediate. The general proportions of the skull and tympanic bullae agree with those of noveboracensis but the skull is larger than in any female of true noveboracensis and approaches that of olivacea. The same can be said of a young female, no. 80, Ohio State Museum, from Roswell, Georgia.
Another female, no. 171559, U. S. Nat. Mus., from Lookout Mountain, 1500 ft., Fort Payne, Alabama, is barely subadult. The external measurements are nearer those of olivacea. The color and narrowness of the underparts are typical of noveboracensis. The proportions and especially size of the skull show approach to olivacea, though they are nearer to noveboracensis when all features are taken into account. In the northern part of its range individuals of noveboracensis attain larger size than farther south. This tendency reaches its extreme, in males at least, in M. f. occisor of Maine. Specimens of noveboracensis from the Adirondacks of New York average larger (see cranial measurements on page 418) than those from farther south, and thus approach occisor in size as well as in geographic position. Also, occasional individuals which strongly show characters of occisor are found even farther south than the Adirondacks of New York. This is true of no. 96518, U. S. Nat. Mus., [M] ad., from Lunenburg; Massachusetts. The animal has a large skull of relatively great width much as in occisor, although its external measurements, relative length of tail and long, terminal, black brush place it with noveboracensis rather than with occisor. Of a pair of specimens from Ossipee, New Hampshire, the male, no. 77108, U. S. Nat. Mus., has a long (175 mm.) tail, and short (60 mm.) black pencil as in occisor, although otherwise it is referable to noveboracensis. Still another specimen, a subadult male, no. 4193, Mus. Comp. Zoöl., from Upton, Maine, has a longer (51 mm.) hind foot than noveboracensis although it otherwise agrees with that subspecies. As remarked by Bangs (1899:55), other than fully adult specimens from the range of occisor are "troublesome," and would not be selected as distinct from noveboracensis if placed in a series of that subspecies, say, from New York State. In view of the facts that several specimens from intermediate localities combine the characters of noveboracensis and occisor, that noveboracensis in the northern part of its range averages larger than it does farther south and thus approaches occisor in size, and that occasional large specimens resembling occisor in several, but not all, features sometimes crop up in the northern part of the range of noveboracensis, it appears that noveboracensis and occisor intergrade. Therefore they are treated as two subspecies of the single species, Mustela frenata.
Intergradation with M. f. primulina has been commented on in the discussion of that subspecies. Female, no. 159980, U. S. Nat. Mus., from Golconda, Illinois, has many characters of primulina but two young males from there agree better with noveboracensis.
Examination of 283 adult and subadult skulls for malformation of the frontal sinuses revealed only ten that were not obviously malformed. Two were from New York, one from Massachusetts, one from Pennsylvania, and six from the 52 specimens from Michigan and Wisconsin. In addition, skulls of many young and even juveniles were malformed.
Specimens examined.--Total number, 555, arranged alphabetically by states and provinces and, unless otherwise noted, from north to south by counties in each state. Except as otherwise noted specimens are in the United States National Museum.
=Alabama.= DeKalb County: Fort Payne, 1.
=Connecticut.= Litchfield County: Riverton, 1[5]; Gaylordsville, 1. Hartford County: East Hartford, 4 (3[5]); Glastonbury, 2[5]; South Glastonbury, 4[5]. Windham County: Plainfield, 2 (1[14]). Fairfield County: Greenwich, 2[2]. New London County: Liberty Hill, 35 (33[75], 2[7]).
=District of Columbia.= Washington, 3; near Washington, 1; Eastern Branch, 1; Congress Heights, 1; Benning, 1; no definite locality, 1.
=Georgia.= Towns County: Young Harris, 1. Cherokee County: Canton, 1. Cobb County: Roswell, 1[81].
=Indiana.= St. Joseph County: Notre Dame, 2[99]. _Porter County_: Hebron, 1. Miami County: Denver, 5 (4[75], 1[4]). Wells County: Bluffton, 1. Howard County: Russiaville, 1. _Jay County_: Salamonia, 1[2]. Boone County, 1[2]. Knox County: Bicknell, 3.
=Illinois.= Lake County: Camp Logan, 3[60]; Fort Sheridan, 1[60]. Cook County: W Northfield, 2; Flossmoor, 1[60]; no locality more definite than county, 1. Du Page County: Bloomingdale Spg., 1[60]. Carroll County: Savanna, 1[87]. McLean County: Normal, 1[7]. Champaign County: Harwood Township, 1[7]. Pike County?: Milton Spring, 1[60]. _Pope County_: Golconda, 3.
=Kentucky.= Woodford County: Midway, 1. Hancock County: Hawesville, 1.
=Maine.= Oxford County: Upton, 1[75]; Bethel, 1[74].
=Maryland.= Howard County: Long Corner, 1; Hanover, 1. Montgomery County: Gaithersburg, 1; Garret Park, 1; Chevy Chase, 1; Bethesda, 1. Prince Georges County: Laurel, 18; Plummer Island, 3; Oxon Hill, 1. Talbot County: Easton, 1. _Dorchester County_: Cambridge, 5[40].
=Massachusetts.= Middlesex County: Wilmington, 6; Burlington, 6; Lexington, 1[75]; Wayland, 2[75]. Berkshire County: New Marlboro, 1[5]. Worcester County: Lunenburg, 2; Lancaster, 1[75]; Princeton, 2[75]. Norfolk County: So. Weymouth, 1[75]. Plymouth County: Wareham, 5[75].
=Michigan.= Marquette County: Michigamme, 1. _Charlevoix County_: Thumb Lake, 1[76]; 1/2 mi. N Thumb Lake, 1[76]. _Leelanau County_: Leland, 3[76]; Duck Lake, 2 mi. S Leland, 1[76]; Lost Pond, 8-1/2 mi. S Leland, 1[76]. Osceola County: Le Roy, 2[76]. Huron County: Rush Lake, 1[76]. Saginaw County: East Saginaw, 1. Oakland County: Royal Oak, 4[76]; South Lyon, 1[76]. Livingston County: Portage Lake, 1[76]. Washtenaw County: Portage Lake, 6[76]; Waterloo, 2[14]; Lima, 1[76]; Ann Arbor, 11[76]; 3 mi. E Ann Arbor, 1[76]; 2 mi. SE Ann Arbor, 1[76]; 2 mi. S Ann Arbor, 1[76]; 3 mi. S Ann Arbor, 1[76]; Dixboro, 1[76]; Pittsfield, 3 (2[76]); Saline, 1[76]; near Saline, 2[76]; 1 mi. S Saline, 2[76]; York, 2[76]; Manchester, 2[76]. Lenawee County: Morenci, 1[76]. Cass County: Marcellus Township, 1[76]. _Berrien County_: Harbert, 1[76]; Warren Wood Preserve, 1[76]; Warren Woods, 1[76].
=New Hampshire.= Grafton County: Franconia, 1[2]. _Carroll County_: South Chatham, 4 (3[5]); Ossipee, 2; Intervale, 1[5]. Merrimack County: Webster, 2[75].
=New Jersey.= Morris County: Morristown, 1. Essex County: West Orange, 1[2]. Mercer County: Princeton, 1[1]. Ocean County: Point Pleasant, 1[2]. Camden County: Haddonfield, 1[1]. Cumberland County: Millville, 2[74].
=New York.= St. Lawrence County: Ogdensburg, 1[74]. _Clinton County_: Rouses Point, 1[80]. County?: Adirondacks, 12. _Essex County_: Elizabethtown, 1; Schroon Lake, 1; no locality more definite than county, 1. Lewis? County: Locust Grove, 4; Lyons Falls, 1. Warren County: Lake George, 6; Caldwell, 1. _Hamilton County_: Beaver Brook, 1/2 mi. above mouth Indian Lake, 1[80]. Oswego County: Scriba, 2[74]; Palermo, 1[74]. Monroe County: Penfield, 3. Madison County: Peterboro, 6 (2[75]). _Schoharie County_: Schoharie, 1[2]. Rensselaer County: East Shodack, 1[80]. Tompkins County: Taughannock Falls, 2[58]; Ithaca, 4 (3[58]); Glenside, Ithaca, 1[58]; 6 mi. Creek, Ithaca, 1[58]. Green County: Lanesville, 1[2]. Orange County: Poplopen's Pond, 1[2]; Highland Falls, 1[2]. Putnam County, 1[19]. Westchester County: Sing Sing, 4; Armonk, 1[2]; Hastings, 3 (2[2], 1[19]). Nassau County: Flushing Meadows, 1[2]; Flushing, 1[58]; near Flushing, 1[2]; Oyster Bay, 2. Long Island: Cold Spring Harbor, 1; Bridgehampton, 1[2]. County in question: Severance, 3; Lake Grove (Long Island?), 1.
=North Carolina= (east to west by counties). Wake County: Raleigh, 4 (1[2], 1[75], 2[76]). Mitchell County: Magnetic City, foot of Roan Mountain, 6; Roan Mt., 1; Roan Mt., 6000 ft., 3. Buncombe County?: Valley of Black Mts., 1[2]. Madison County, 2[11].
=Ohio.= Trumbull County: Warren, 1[93]. Seneca County: Tiffin, 1[81]. Summit County: Ira, 2[81]. Crawford County: Galion, 1[81]. Ashland County: Loudonville, 1[76]. Auglaize County: New Bremen, 3[81]. Franklin County: 3 mi. N Columbus, 1[81]; Minerva Park, Columbus, 5[81]. Fairfield County: Sec. 32, Pleasant Twp., 1[81]; Lancaster, 1[81]. Clinton County: Reesville, 1; 1/2 mi. S and 1/2 mi. W Wilmington, 2[74]. _Pike County_: Waverly, 1[81].
=Ontario.= Sudbury District: Metagama, 2[86]. Carleton County: Ottawa, 2[77]. Muskoka County: Lake of Bays, 1; Bracebridge, 1. Haliburton County: Gooderham, 1[60]. Simcoe County: Orillia, 4 (2[2], 2[60].) Prince Edward County: Bloomfield, 1[77]. _York County_: Toronto, 1[2]. Waterloo County: Branchton, 3[60]; Preston, 2[77]; no locality save county, 1[60]. Welland County: Ridgway, 1[14]. Elgin County: St. Thomas, 1[77]. Essex County: Kingsville, 1[77]; Point Pelee, 1[77].
=Pennsylvania= (east to west by counties). Crawford County: Pymatuning Swamp, 3-1/2 mi. W Linesville, 1[9]; Meadville, 2[9]. Beaver County: Beaver, 1[9]; Raccoon Creek, 1[9]. _Butler County_: Mars, 1[9]; Leasuresville, 4[9]. Allegheny County: Allegheny, 1. Warren County: Bear Lake, 2[9]. _Westmoreland County_: Kingston, 1[9]; Laughlinstown, 2[9]. Somerset County: Confluence, 1[9]; Tub Mill Run, 2 mi. N Springs, 1[9]. _Jefferson County_: Siegel, 1[9]. Clearfield? County: Penfield, 1[9]. Cambria County: Cresson, 1[9]. Fulton County: Well's Tannery, 1[9]. Clinton County: near Round Island, 2[1]. _Cumberland County_: Carlisle, 1. Snyder County: 5 mi. S Selinsgrove, 1. Northumberland County: Pottsgrove, 1. Union County: Mifflinburg, 1. Sullivan County: Lopez, 7 (4[1], 3[74]). Chester County: Westtown, 1[1]; Valley Forge, 1[1]; W Bradford Township, 1[1]; no locality more definite than county, 3. Philadelphia County: Holmsburg, 2[1]. Bucks County: 1[1]. Pike County: Milford, 1.
=Rhode Island.= Providence County: Chepachet, 1. _Washington County_: Lake Warden, 2.
=Quebec.= Megantic County: Black Lake 1[77]. County in question: Meach Lake, 1[77].
=South Carolina.= York County: 5 mi. E York, 1[11]. _Laurens County_: Laurens, 1[39].
=Tennessee.= Campbell County: Highcliff, 1. Carter? County: Roan Mts., 1[2]. Hamilton County: Walden Ridge, near Soddy, 3.
=Vermont.= Windsor County: Hartland, 1[2].
=Virginia.= Shenandoah County: Toms Brook, 1. Arlington County: Arlington, 1; Ballston, 1; Alexandria, 1. Fairfax County: Falls Church, 3; Mt. Vernon, 2; no locality more definite than county, 1. Prince William County: Occoquan, 1. Essex County: Montague, 1. Prince George County, 1. Norfolk County: Wallaceton, 1. Grayson County: Mt Rogers, 3. County in question: Dismal Swamp, 1; Massanutten Mt., 1.
=West Virginia.= Hardy County, 1. Pendleton County: radius of 2 mi. Smoke Hole, 1[74]. Greenbriar County: White Sulphur, 2[60].
=Wisconsin.= Douglas County: Gordon, 1[104]. Vilas County: Mamie Lake, 4. Dunn County: Colfax, 4[104]; Meridean, 1[104]. Door County: state game farm, 17[104]; no locality more definite than county, 1[104]. Dodge County: Rolling Prairie, 1[50]; Beaver Dam, 52[50]. Dane County: Wingra Lake, 1[104]. _Waukesha County_: Pewaukee, 2[104]. Racine County: Waterford Township, 2[104]. Rock County: Milton, 1[104]; Bowers Lake, 1[104]. Walworth County: Lane's Mill, 8 mi. N Elkhorn, 7 (1[104], 6[54]); Delavan, 7.
=Mustela frenata occisor= (Bangs)
Long-tailed Weasel
Plates 16, 17, 18, 31, 32 and 33
Putorius occisor Bangs, Proc. New England Zool. Club, 1:54, June 9, 1899.
Mustela occisor, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912.
Mustela frenata occisor, Hall, Carnegie Instit. Washington Publ. 473:104, November 20, 1936.
Type.--Male, adult, skull and skin; no. 9102, coll. of E. A. and O. Bangs in Mus. Comp. Zoöl.; Bucksport, Hancock County, Maine; January 15, 1899; obtained by Alvah G. Door but measured and sexed by O. Bangs.
The skin is well made and in good condition. It is in full, white winter-dress with black-tipped tail. The skull has the posterior half of the left zygomatic arch broken away; otherwise the skull is unbroken and complete. Left I3 and right P3 are missing. The teeth otherwise all are present and entire.
Range.--Maine; possibly north locally to south side of St. Lawrence River in Quebec and possibly occurring in western New Brunswick. Zonal range Canadian and probably Transition. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. noveboracensis by a number of average differences including larger size, relatively longer tail and relatively wider skull (see p. 225, and measurements on pp. 418, 419).
Description.--Size.--Male: Five adults yield average and extreme measurements as follows: Total length, 443 (430-465); length of tail, 163 (154-175); length of hind foot, 50 (47-54). Tail averages 58 per cent as long as head and body. Length of hind foot averages more than basal length.
Female: Measurements of two subadult female topotypes are as follows: Total length, 346, 318; length of tail, 116, 110; length of hind foot, 39, 35.5.
Tail amounts to 50 per cent and 54 per cent of body-length respectively. Length of hind foot more or less than (about equal to) basal length.
The average differences in external measurements of the two sexes are: Total length, 111; length of tail, 50; length of hind foot, 12.5.
Externals.--As described in Mustela frenata noveboracensis.
Color.--As described in Mustela frenata noveboracensis except that black tail-tip in series of 10 males in full winter pelage 60 (45-80) mm. long; thus averaging 39 per cent of length of tail vertebrae.
Skull and teeth.--Male (based on 3 adults): See measurements and plates 16-18. As described in Mustela frenata noveboracensis except that: Weight, 4.2 (4.1-4.3) grams; basilar length, 45.7 (44.9-46.9); zygomatic breadth more or less than (about equal to) distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate rarely less than length of P4; anterior margin of masseteric fossa behind or directly below posterior half of m2.
Female (based on 2 subadults): See measurements and plates 31-33. As described in Mustela frenata noveboracensis except that: Weight, 2.0 (1.9-2.1) grams; basilar length, 37.3, 38.2.
Comparison of the skull with that of M. f. noveboracensis is made in the account of that subspecies.
Remarks.--Excepting a specimen in the Academy of Natural Sciences of Philadelphia, obtained in 1893, and two in the Boston Society of Natural History, obtained in 1925, I have seen no material of this subspecies in addition to that examined by Bangs at the time he prepared the original description in 1899.
Anderson (1945:56, 57) records a specimen, Canadian National Museum Catalogue Number 18426, from Kamouraska County, Quebec, as of this subspecies and thinks that occisor occurs north of Maine "locally to south side of lower St. Lawrence River in Quebec; probably also in western New Brunswick."
So far as the available material of occisor permits one to judge, it is distinguished from noveboracensis by a combination of characters no one of which invariably can be relied upon as diagnostic. Employing adult males, average differences indicate that M. f. occisor is larger in each of the external and cranial measurements; tail relatively longer; black tip of tail relatively shorter; mastoid and zygomatic breadth relatively greater and zygomatic arches more nearly square posteriorly.
Considering the large number of specimens of noveboracensis which are available in comparison with the few of occisor it is not surprising that some noveboracensis should be found which exceed in size those of occisor. This is the case as regards the basilar length of a very old male, no. 96518, U. S. Nat. Mus., from Lunenburg, Massachusetts. Also, the skull is actually broader than any of those of occisor. However, this specimen is much older than any occisor examined. In a female, no. 4260, Mus. Comp. Zoöl., from Liberty Hill, Connecticut, the skull is longer (but narrower) than in either of the two available females of occisor.
The average differences pointed out above which characterize this extreme northern population of noveboracensis-like weasel in comparison with true noveboracensis without much question are geographic variations. Whether or not these variations are of a degree sufficient to warrant nomenclatural recognition is debatable. With equally scanty material from other regions I have not named variations seemingly as great as those shown by occisor. The paucity of material of occisor is a handicap in making a decision in this instance.
Each of the adult and subadult specimens, except the one from Perry, shows malformation resulting from the infestation of the frontal sinuses with parasites.
Specimens examined.--Total number, 18, listed by counties from west to east and unless otherwise indicated in the Museum of Comparative Zoölogy.
=Maine.= Oxford County: South Andover, 2 (Boston Soc. Nat. Hist.); Umbagog Lake, 1. Franklin County: Seven Pd. Township, 1. Piscataquis County: Grenville, [= Greenville?], 1. _Hancock County_: Bucksport, 10. Washington County: 3rd Mopang Lake, 1 (Acad. Nat. Sci. Phila.); Perry, 1 (Boston Soc. Nat. Hist.). County in question: Moosehead Lake, 1.
=Mustela frenata primulina= Jackson
Long-tailed Weasel
Plates 16, 17, 18, 31, 32 and 33
Mustela primulina Jackson, Proc. Biol. Soc. Washington, 26:123, May 21, 1913.
Putorius noveboracensis, Baird, Mamm. N. Amer., p. 166, 1858 (part).
Mustela longicauda longicauda, Dice, Journ. Mamm., 4:108, May 9, 1923.
Mustela longicauda primulina, Linsdale, Journ. Mamm., 9:141, May 9, 1928.
Mustela frenata primulina, Hall, Carnegie Instit. Washington Publ. 473:104, November 20, 1936.
Mustela frenata, Leopold and Hall, Journ. Mamm., 26:143, July 19, 1945.
Type.--Male?, young, skull and skin; no. 168006, U. S. Nat. Mus., Biol. Surv. Coll.; 5 miles northeast of Avilla, Jasper County, Missouri; May 11, 1905; obtained by Hartley H. T. Jackson; original no. 7869X.
The skin lacks the distal part of the tail--the part which bears the black tip. Otherwise the skin is complete and well preserved. The teeth of the permanent dentition all are present and entire. The lower jaws are complete and unbroken. The skull is broken transversely through the interorbital region, transversely through the braincase and longitudinally through the basioccipital. Both zygomatic arches are gone. The type is judged to be a male rather than a female as stated by the original describer, Jackson (1913:123), whose measurements of hind foot, interorbital constriction, maxillary tooth-row, and mandibular tooth-row agree with those of males and are larger than those of any female seen of this subspecies.
Range.--Upper and Lower Austral life-zones west of the Mississippi River in Missouri and Arkansas, the southeastern half of Iowa, eastern half of Kansas and Oklahoma, northern Louisiana and northeastern Texas. Southern and southwestern limits of range undetermined. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. noveboracensis in males by interorbital breadth averaging less than 24 per cent of basilar length, orbitonasal length averaging less than 34 per cent of basilar length or 64 per cent of mastoid breadth, tympanic bullae as much inflated anteromedially as posteromedially, and in females by orbitonasal length amounting to less than two-thirds of mastoid breadth, by zygomatic breadth averaging more than 21 mm., and by anterolateral margin of tympanic bulla projecting below squamosal; from M. f. spadix by least width of color of under parts amounting to less than 40 per cent of greatest width of color of upper parts, by absence of color of underparts on hind leg below knee, and by smaller size (hind foot less than 50 in males and 40 in females; orbitonasal length less than 15.5 in males and 13.5 in females; length of tooth-rows less than 18 in males and 15.7 in females; mastoid breadth less than 25.5 in males and 22 in females); from M. f. longicauda by Brussels Brown rather than near (h) Clay Color of upper parts, least width of underparts less than 40 per cent of greatest width of color of upper parts, absence of color of underparts on hind leg below knee, zygomatic breadth less than 28.8 in males and 24.1 in females; from M. f. neomexicana by Brussels Brown rather than Buckthorn Brown color of upper parts, in absence of white frontal spot and broad white bands on sides of head, in anterolaterally rounded, rather than "square," tympanic bullae and in zygomatic breadth of less than 30 in males and 24 in females; from M. f. frenata and M. f. texensis by absence of white facial markings and smaller size (basilar length of adult males less than 47, tail-length less than 155 in males, and hind foot less than 40 in females); from M. f. arthuri by less evenly spreading zygomatic arches (see pls. 16, 17 and 18), greater inflation of the tympanic bullae anteromedially and more nearly straight (less convex) dorsal outline of skull.
Description.--Size.--Male: Eighteen adults and subadults from Douglas County, Kansas, yield average and extreme measurements as follows: Total length, 397 (371-440); length of tail, 133 (120-147); length of hind foot, 43 (40-47). Tail averages 50 per cent as long as head and body. Length of hind foot averages less than basal length. Corresponding measurements, originally taken in inches and fractions thereof, of 9 adults and subadults from Boone County, Arkansas, are as follows: 413 (384-438); 138 (127-155); 41 (38-44).
Female: Six adults and subadults from Douglas County, Kansas, yield average and extreme measurements as follows: Total length, 339 (317-355); length of tail, 107 (95-115); length of hind foot, 35 (34-37). Tail averages 46 per cent of length of head and body. Length of hind foot less than basal length. Corresponding measurements, originally taken in inches and fractions thereof, of 5 adults and subadults from Boone County, Arkansas, are as follows: 355 (332-397); 113 (101-127); 33 (29-38).
The average differences in external measurements of the two sexes, in Douglas County, Kansas, are: Total length, 58; length of tail, 26; length of hind foot, 8. An adult male from Boone Co., Iowa, weighed 293 grams.
Externals.--Longest facial vibrissae black or dark brown (often both colors in same specimen) and extending beyond ear; carpal vibrissae colored either like underparts or upper parts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 20.
Color.--Upper parts, in summer, Brussels Brown to near (14 n) Brussels Brown, or tones 2 to 4 of Raw Umber of Oberthür and Dauthenay, pl. 301. Chin and rarely upper lips white. Remainder of underparts Picric Yellow to Primuline Yellow. In winter, color essentially the same except for smoke-gray effect in upper parts and more whitish in underparts. Tip of tail at all times black. Upper parts of uniform color except for occasional darkening of nose and mid-dorsal region. Color of underparts extends distally on posterior sides of forelegs onto antipalmar faces of toes, on medial sides of hind legs only to a point between knee and ankle. Least width of color of underparts averaging, in a series of 21 males from Lawrence, Kansas, 23 (9-35) per cent of greatest width of color of upper parts. Black tip of tail in same series, most of which are in full winter pelage, 52 (40-70) mm. long; thus longer than hind foot and averaging 39 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on ten adults from Douglas County, Kansas): See measurements and plates 16-18; weight, 3.7 (3.3-4.2) grams; basilar length, 44.8 (43.8-46.0); zygomatic breadth more or less (less in 80 per cent) than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior end of tympanic bulla (less in 70 per cent); mastoid breadth more or less (less in 80 per cent) than postpalatal length; postorbital breadth less than length of upper premolars and, except in one specimen, more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less (less in 70 per cent) than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than outside length of P4 (except in one specimen); anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2-1/2 to 5 upper incisors; height of tympanic bulla more (except in one specimen) than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer (except in one specimen) than rostrum; anterior margin of masseteric fossa behind or just below posterior eighth of m1.
Female (based on 5 adults and subadults from Douglas County, Kansas): See measurements and plates 31-33; weight, 2.2 (2.0-2.4) grams; basilar length, 38.9 (37.6-40.7); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and except in one specimen, more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum.
The skull of the female averages 41 per cent lighter than that of the male.
Compared with the skull of M. f. noveboracensis from Massachusetts, that of the male of primulina, in dorsal view, is seen to be shorter anteriorly to the postorbital processes and to have a more marked postorbital constriction. In lateral view the dorsal outline of the skull of primulina is less concave in the postorbital region. In ventral view the skull of primulina is seen to be wider across the mastoid processes and zygomatic arches but the most pronounced difference is in the tympanic bullae. In noveboracensis each bulla is scooped out on the anterior part of the medial face and appears to be narrower anteriorly than posteriorly whereas in primulina the anterior part of the medial face is not scooped out but is moderately inflated and the bulla appears to be of uniform breadth anteriorly and posteriorly. By actual measurement the breadth of the bulla averages 59 per cent of its length in primulina but only 50 per cent in noveboracensis. Other respects in which the skull of the male of primulina differs from that of noveboracensis are as follows: Linear measurements of teeth more; relative to the basilar length, the length of the tooth-rows averages more, whereas the interorbital breadth and orbitonasal length are less.
When skulls of females are compared, each of the differences mentioned above is found to apply, except that the degree of difference is in some parts greater, for example, in the tympanic bullae. In primulina, the bulla is in general like that of the male noveboracensis, whereas in the female noveboracensis it is less inflated, especially anteromedially, shorter, relatively narrower, and in ventral view projects little or none below the squamosal floor of the braincase. The breadth of the bulla averages 51 per cent of its length in primulina but only 47 per cent in noveboracensis. The bullae project below the basioccipital on the average, for a distance of 2.9 millimeters in female primulina and only 2.3 millimeters in female noveboracensis. In primulina the temporal ridges are well developed and fuse to form a low sagittal crest, but in noveboracensis the ridges are absent. Also, in primulina the mastoid processes project farther laterally beyond the braincase. The skull of female noveboracensis is much lighter than that of primulina. Average weights of the two are 1.7 and 2.2 grams. The skulls of females of primulina and noveboracensis differ more than do the skulls of males.
Compared with the skull of spadix, that of the male, and the female, of primulina averages smaller in every part measured. Expressed in percentages of the basilar length, the two depth measurements of the skulls are not significantly different, but, excluding the measurements of the bullae and teeth, the other cranial measurements are less. The main difference in relative proportions is in the tympanic bullae which average only a half millimeter shorter in males of primulina and one and one-tenth millimeters shorter in females. The bullae are, therefore, relative to the basilar length, longer in primulina. The skull of primulina, then, differs from that of spadix mainly in smaller size and relatively longer tympanic bullae, especially in males.
Compared with the skull of M. f. longicauda, that of both sexes of primulina averages smaller in every part measured, except in males where the length of the tympanic bulla, and breadth and length of M1 are the same or slightly larger in primulina. Relative to the basilar length, the length of the tympanic bullae, and in females only, the depth measurements are greater in primulina but all the others, in both sexes, are less. These ratios reflect the relative narrowness of the skull of primulina. Upon direct comparison the narrowness is especially noticeable in the interorbital region, mastoid region, tympanic bullae, and across the zygomata.
Compared with the skull of M. f. neomexicana that of both sexes of primulina averages smaller in every part measured. Excepting the measurements of the teeth, most of the other measurements are constantly larger. Relative to the basilar length, the length of tooth-rows and length of tympanic bulla are more, but excepting the depth measurements, the others are less. Still other differences are, in primulina, less well-developed sagittal crest, anterolateral corner of bulla rounded rather than "square," and in males a transversely convex rather than flat interorbital region.
Compared with M. f. frenata and M. f. texensis, the skulls of males of primulina differ in being smaller in every part measured but relative to the basilar length, have longer tooth-rows, a lesser zygomatic breadth and are less constricted interorbitally.
Compared with the skull of M. f. olivacea, those of both sexes of primulina average smaller in every part measured, have shallower (dorsoventrally) tympanic bullae, a lower sagittal crest and slightly weaker postorbital processes on the frontals. Relative to the basilar length, the several cranial measurements are about the same.
Comparison of the skull with that of M. f. arthuri has been made in the account of that subspecies.
Remarks.--The first specimens of this race known to have been preserved in study collections are one in the United States National Museum, taken at Bridge, Carroll County, Missouri, many years ago by J. Burbage, and less than a dozen specimens preserved before 1900 from eastern Kansas in the University of Kansas Museum of Natural History. In 1913 Hartley H. T. Jackson bestowed a name on this animal on the basis of two specimens taken by him in southwestern Missouri. Later, through the efforts of Charles D. Bunker, and his associates at the University of Kansas, nearly 100 specimens were saved from eastern Kansas, principally from Douglas County. In the course of the present study, Lawrence V. Compton obtained a topotype for the California Museum of Vertebrate Zoölogy, and with the assistance of Mr. B. G. Roberts, a good series of specimens from Boone County, Arkansas, was preserved in the same museum. In the early years of the 20th Century, the late B. H. Bailey at Coe College, Iowa, collected specimens from that state. The specimens from these several sources suffice to give a relatively clear idea of the characters of this subspecies.
Mustela frenata primulina is closely related to M. f. noveboracensis, from which, on the average, it differs in the lighter color of the upper parts of the summer coat, in the more intense coloring of the underparts, and in certain cranial features pointed out above. In the southern part of its range, however, noveboracensis has the underparts only a little less intensely colored with yellow than primulina. Also, the skull of the one topotype from 7-1/2 miles southeast of Carthage, a subadult male in brown, winter pelage, is almost exactly intermediate between that of noveboracensis from Massachusetts and primulina of Douglas County, Kansas, and Boone County, Arkansas. M. f. primulina often has the underparts white in winter, as does this topotype which agrees with the average of noveboracensis in small size of teeth and narrowness across the mastoid processes and zygomatic arches. However, it agrees with primulina in shape and relative size of the rostrum. It is almost exactly intermediate in shape and width of the tympanic bullae.
Three other males, but no females, all in winter pelage, are available from eastern Missouri. Of the two from Silex, Lincoln County, one is nearer noveboracensis and the other nearer primulina on the basis of cranial characters. The third specimen, from four miles south of Lesterville, so far as I can determine by examination of individual cranial characters and tabulation of results, is exactly intermediate. Final decision on the proper allocation of specimens from the parts of Missouri represented can best be made when skulls of females are available. From the fact that the skull of the female referred to noveboracensis from Golconda, Illinois, shows almost as many characters of primulina as of noveboracensis, it is judged that females from as far west as Silex and Lesterville, Missouri, will show even more characters of primulina and so be referable to that form. If this supposition be correct, the present reference of the almost exactly intermediate males, from eastern Missouri, will stand; otherwise, it may not.
Additional intergrades with noveboracensis are available from eastern Iowa. Of five specimens from Hillsboro, Iowa, two males and a female have tympanic bullae like those of primulina but the other two males have bullae like those of noveboracensis. The female is smaller than primulina and in this small size and in general configuration of the skull, viewed dorsally, is more nearly like noveboracensis. As a whole, the population averages almost exactly intermediate. The same is true of 3 males and one female from Muscatine. The subadult male from Keosaqua, to my eye, resembles noveboracensis in the greater length of the skull anteriorly to the postorbital processes, and in the relative narrowness across the mastoidal region, but otherwise is more like that of primulina. Two males and one female from Tipton, although in each instance variously intermediate, are as a whole nearer primulina, No. 2865, Coe College, male adult, from Cedar Rapids, has characters of the three races, spadix, noveboracensis and primulina. In the skull, the width suggests spadix, the narrow mastoid region, noveboracensis, and the tympanic bullae are as in spadix or primulina. One male, no. 12, Coe College, from Dubuque, is as narrow across the mastoid region as is noveboracensis although the bullae are well inflated as in primulina. The skull, without corresponding skin, of a female, no. 140a, Iowa State College, from Green's Island, also resembles noveboracensis in narrowness of the mastoidal region, and in small size of skull, but in larger teeth, broader tympanic bullae, and sagittal crest is referable to primulina. Of two females from Vinton, one adult is typical of primulina but the other, a subadult, is practically indistinguishable from female noveboracensis, from Ann Arbor, Michigan. Three males from Vinton agree well with primulina except that the interorbital region is wider than average and thereby suggests spadix or noveboracensis. An adult female from New Hartford also is typical of primulina except for the broader interorbital region. Three males from Fayette are typical of primulina.
Other specimens from Iowa are intergrades with spadix, or if not with spadix, with the animal of northwestern Iowa which in some ways combines the characters of longicauda and spadix. For example, no. 2665, Coe College, an adult male from Davenport, has the anterior part of the skull (all that is preserved) heavily ridged as in spadix and in addition, the underparts are marked with the shade of reddish displayed by topotypes of spadix and with some yellowish as seen in longicauda. The color pattern, however, is as in primulina. A young male, no. C-51, Iowa State College, from Kelley, Story County, has anteriorly truncate bullae as are more frequently found in the longicauda-spadix stock of northwestern Iowa, than in primulina. In other respects, the animal, in so far as can be judged from the broken skull, agrees with primulina as it certainly does in color, color pattern, and external measurements. An adult male, no. 499a, Iowa State College, from 2 miles east of Ledges St. Park, in Boone County, in short body, size of teeth, and size of skull, in so far as the broken parts can be measured, resembles primulina more closely than it does any other subspecies. The long tail, long hind foot, wide extent of the light-colored underparts, and extension of the color of the underparts onto the hind feet are more as in spadix. Other intergrades with spadix from Iowa are mentioned in the account of spadix.
The specimen from Swartz, Louisiana, suggests intergradation with arthuri in that the anteromedial part of the tympanic bulla is less inflated than in typical primulina.
Intergrades with longicauda are available from Riley and Pratt counties, Kansas. No. 7182, Univ. Kans., subadult male in winter pelage, from near Winkler, has a skull of larger size as in longicauda with which race it seems to agree in large size of body, tail and hind foot, although the collector's measurements are lacking. Color pattern and relative proportions of the skull throughout are as in primulina. The young male, no. 3495, Univ. Kans., from Pratt, Kansas, agrees in external measurements and large size of skull with longicauda, but has the color and color pattern precisely as in primulina. The teeth are smaller as in primulina. Immaturity prevents judging of its relationships on the basis of relative proportions of the skull.
The two specimens, skins only, available from Oklahoma, are provisionally referred to primulina. These are remarkable for the restriction of the color of the underparts and for the intensity of the yellow coloration of the underparts. The specimen from Norman has the color of the underparts entirely absent from the hind legs and not extending posteriorly to the penis. On the chest and lower throat, large spots of color of the upper parts are present and the yellow area of the underparts on the belly is narrower than in any other specimen of primulina examined. The specimen from 8 miles northwest of Stillwater has the color of the underparts only a little less restricted although this color does extend over the inguinal region almost to the knees. The skin of the posterior part of the body of a weasel is available from 10 miles south of Sulphur Springs, Texas. It, likewise, is only provisionally referred to primulina. The coloration is about as in the specimens from Oklahoma but the distribution of the color of the underparts cannot be made out.
The dark color of the upper parts occurs far westward in animals which otherwise display characters of longicauda. Among these intergrades, the larger size of longicauda generally is combined with this dark color. This geographic behavior of the dark color of the upper parts is analogous to the condition described in M. f. spadix. Stated in another way, the dark color of the upper parts is the character, of the eastern animal, last to disappear as one goes westward across the Mississippi Valley toward the range of longicauda which is a subspecies of markedly different size, shape of skull, and coloration.
Only two of 29 specimens from Kansas show infestation of the frontal sinuses. All four of the specimens from Missouri have the frontal sinuses malformed as do 9 of the 14 from Arkansas examined in this respect.
An adult female from Boone County, Iowa, bears the date May 9, 1938, and the annotation by T. G. Scott, "Fox-killed."
Specimens examined.--Total number, 131, arranged alphabetically by states and from north to south by counties in each state. Except as otherwise indicated, specimens are in the University of Kansas, Museum of Natural History.
=Arkansas.= Boone County: 3 mi. E Bergman, 4[74]; 3 mi. SE Bergman, 1[74]; 3 mi. S Bergman, 1[74]; 3 mi. SW Bergman, 1[74]; 4 mi. SE Bergman, 2[74]; 5 mi. SE Bergman, 1[74]; 4-1/2 mi. SE Bergman, 3[74]; 5 mi. SE Bergman, 1[74]; 5 mi. S Bergman, 2[74]; 5 mi. SW Bergman, 2[74]. Washington County: Fayetteville, 1[96]. Crawford County: 10 mi. S Winslow, 1. Sebastian County: Fort Smith, 1[91].
=Iowa.= Fayette County: Fayette, 3[12]. Dubuque County: Dubuque, 1[12]; Green's Island, 1[65]. Butler County: New Hartford, 1[12]. Hardin County: Union, 1[65]. Benton County: Vinton, 5[12]. Linn County: Cedar Rapids, 1[12]. Boone County: Worth Township, Sec. 21, 1[65]; 2 mi. E Ledges St. Park, 1[65]. Story County: Kelley, 1[65]. Cedar County: Tipton, 3[12]. Scott County: Davenport, 2[12]. Muscatine County: Muscatine, 4[12]. Henry County: Hillsboro, 5[91]. Van Buren County: Keosaqua, 1[65]; no locality more definite than county, 1[50]. Taylor County, 1.
=Kansas.= Riley County: near Winkler, 1. Pottawatomie County: Onaga, 1[83]. Atchison County: Doniphan Lake, 1; 5 mi. NE Muscotah, 1; no locality more definite than county, 1. _Douglas County_: Lawrence, 8; 6 mi. NW Lawrence, 1; 1-1/2 mi. W Lawrence, 1; 6 mi. S Lawrence, 1; 7 to 7-1/2 mi. SW Lawrence, 14; 10 mi. W Lawrence, 1; Clinton, 4; Baldwin, 1; no locality more than county, 29 (2[74]). Woodson County: 1-1/2 mi. S Neosho Falls, 1[59]. Greenwood County: 8 mi. SW Toronto, 2. Pratt County: Pratt, 1.
=Louisiana.= Quachita Parish: Swartz, 1[71].
=Missouri.= Carroll County: Bridge Creek, 1[91]. Lincoln County: Silex, 1[74]; 1 mi. E Silex, 1[74]. Reynolds County: 4 mi. S Lesterville, 1[74]. Jasper County: 5 mi. NE Avilla, 1[91]; 7-1/2 mi. SE Carthage, 1[74].
=Oklahoma.= Payne County: 8 mi. NW Stillwater, 1[82]. _Cleveland County_: Norman, 1[100].
=Texas.= Hopkins County: 10 mi. S Sulphur Springs, 1[43].
=Mustela frenata arthuri= Hall
Long-tailed Weasel
Plates 16, 17 and 18
Mustela noveboracensis arthuri Hall, Proc. Biol. Soc. Washington, 40:193, December 2, 1927.
Mustela frenata arthuri Hall, Carnegie Instit. Washington Publ. 473:105, November 20, 1936.
Type.--Male, subadult, skull and skin; no. 37515, Mus. Vert. Zoöl.; Remy, St. James Parish, Louisiana; December 15, 1926; obtained by Stanley C. Arthur.
The skin is stuffed and well preserved. The skull (plates 16-18) is unbroken. The teeth all are present and entire. The presence of a well-developed scrotal pouch shows the specimen to be a male. Contrary to what was stated in the original description the specimen was taken in 1926 and not in 1925.
Range.--Lower Austral Life-zone of southeastern Texas, Louisiana, and into Mississippi. See figure 29 on page 221.
Characters for ready recognition (of males).--Differs from M. f. olivacea in smaller size (adult males with hind foot and basilar length less than 45), depth of skull at anterior margin of basioccipital, ignoring sagittal crest, amounting to more than 63 per cent of mastoid breadth, and greater convexity of dorsal outline of skull in longitudinal axis (see pls. 16-18); from M. f. noveboracensis, in males, by zygomatic breadth not less than distance between anterior palatine foramen and anterior margin of tympanic bulla and by convex dorsal outline of skull in longitudinal axis; from M. f. primulina by evenly spreading zygomatic arches, lesser inflation of tympanic bullae anteromedially than posteromedially, and convex dorsal outline of skull in longitudinal axis; from M. f. texensis and M. f. frenata by absence of white facial markings and postorbital breadth more than distance between posterior borders of P4 and P2.
Description.--Size.--Male: The type, a subadult male, measures (inches and quarter fractions thereof, transposed into millimeters) as follows: Total length, 390; length of tail, 113; length of hind foot, 44. Tail is 41 per cent as long as head and body. Length of hind foot less than basal length.
Typical female unknown.
Externals.--Longest facial vibrissae black, or dark brown (both colors in the type) and extending beyond ear; carpal vibrissae same color as underparts and extending to within 3.5 millimeters of apical pad of fifth digit. Hairiness of foot-soles in type slightly less than shown in figure 20.
Color.--Upper parts in summer tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304; underparts as described in M. f. olivacea. In winter, upper parts (based on type) near (1) Brussels Brown or grayer than tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304, darker on top of head from nose to a line connecting posterior margins of ears. Chin and posterior third of each upper lip white. Remainder of underparts white with wash of Warm Buff. Tip of tail black. Color of underparts extends distally on posterior sides of forelegs over toes but represented on antipalmar faces of feet by only a few scattered hairs. Color of underparts extends distally on medial sides of hind limbs only to knees. Least width of color of underparts amounting to 15 per cent of greatest width of color of upper parts. Black tip of tail 50 mm. long; thus longer than hind foot and 44 per cent as long as tail-vertebrae.
Skull and teeth.--Male (based on type and 2 subadults): See measurements and plates 16-18. As described in M. f. noveboracensis except that: Weight, 4.0 (3.7-4.3) grams; basilar length, 43.5 (43.3-43.6); zygomatic breadth not less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than length of upper premolars; interorbital breadth more than distance between foramen opticum and anterior margin of tympanic bulla; least width of palate more or less than length of P4; tympanic bulla longer than rostrum.
Female: Typical skull unknown. The skull from 12 miles east of Eagle Lake, Texas, lacks the convexity in the dorsal longitudinal axis and the skull agrees with those of larger individuals of primulina except that the anteromedial faces of the tympanic bullae are less inflated, and the mastoid and zygomatic breadths are greater than in any female seen of primulina. Probably this greater breadth is the result of intergradation with M. f. frenata to the westward.
Compared with the skull of M. f. olivacea that of arthuri differs as follows: Averaging smaller in every part measured; basilar length 5 mm. less; by weight a fourth lighter; relative to basilar length, interorbital breadth greater and zygomatic and especially mastoid breadth less; dorsal outline of skull more convex in longitudinal axis; tympanic bullae narrower and less inflated especially on anteromedial faces. Compared with the skull of noveboracensis that of arthuri has the zygomatic breadth equal to or exceeding the distance from the anterior palatine foramen to the anterior margin of the tympanic bulla, whereas the zygomatic breadth is less than this distance in noveboracensis. Also, in arthuri, the rostrum is relatively shorter, the braincase is more inflated anteriorly, the zygomatic arches are more uniformly spreading, and the dorsal outline of the skull is distinctly convex, both transversely and longitudinally, whereas it is transversely more nearly flat in noveboracensis and longitudinally is concave in the interorbital region.
Compared with M. f. primulina, arthuri has narrower bullae, which are much less inflated on their anteromedial faces, a less marked postorbital constriction, a braincase which is narrower across the mastoid region and broader anteriorly, and a skull, which, in longitudinal axis, has the dorsal outline markedly more convex.
Compared with the skull of M. f. texensis that of arthuri is smaller in every part measured; length one-fifth less; one-half as heavy; postorbital constriction less marked; braincase relatively narrower posteriorly and tympanic bullae less inflated especially anteromedially. Compared with the skull of M. f. frenata that of arthuri is smaller in every part measured; basilar length 6 mm. less; a third lighter; postorbital constriction less marked; relative to the basilar length the rostrum is broader, longer and deeper; the zygomatic expanse and breadth of the braincase across the mastoids is less; the dorsal profile of the skull is more convex in longitudinal axis; zygomata evenly spreading rather than abruptly protruding from skull posteriorly; tympanic bullae less inflated anteromedially.
Remarks.--In 1926, Stanley C. Arthur, then Director of the Division of Wild Life, for the Louisiana State Department of Conservation, obtained specimens of this weasel. Some of them were mounted and the remainder were placed in the collections of the United States National Museum and the Museum of Vertebrate Zoölogy. In 1938 to 1940 George H. Lowery saved specimens from Baton Rouge, which showed the color of the summer pelage and revealed that the size of males was more than was indicated by the original materials. In 1940 and 1941 Rollin H. Baker obtained specimens from eastern Texas which greatly extended the known geographic range.
In addition to the localities represented by specimens examined, Arthur (1928:117) has recorded specimens from Greensburg, St. Helena Parish; Braithwaite, Plaquemines Parish; Geismar, Assumption Parish; Laurel Hill, West Feliciana Parish; French Settlement, Livingston Parish; and Kentwood, Tangipahoa Parish. All these localities lie within the eastern half of southern Louisiana. A skin-only, no. 38902, Mus. Vert. Zoöl., obtained from a fur buyer by Stanley C. Arthur, was taken in Mississippi "south of Jackson." Possibly it is of the subspecies arthuri.
Intergradation with M. f. olivacea is indicated by a specimen from Mobile County, Alabama, commented on in the account of olivacea. Intergradation with primulina is indicated by the shape of the anteromedial part of the bullae of the specimen from Swartz, Louisiana, that is referred to primulina. The lack of specimens from the northern two-thirds of Mississippi and from western Tennessee, prevents any definite statement as to the limits of range of arthuri in those areas. In comparison with animals from the type locality, the slightly larger size of the adult male from Baton Rouge, and the still larger size of the adult male of primulina from Swartz, Louisiana, suggests that the olivacea "influence" may extend farther west in the latitude of northern Louisiana than anywhere else.
None of the skulls examined shows malformation of the frontal sinuses such as results from infestation by parasites in some races. Arthur (1928:115) speaks of the ". . . cut-over swamp land, where the tupelo and cypress have been removed, . . ." as constituting suitable habitat for this animal.
Specimens examined.--Total number, 13, as follows:
=Texas.= Colorado County: 12 mi. N Eagle Lake, 1[43]; 5 mi. W Eagle Lake, 1[43]; 3 mi. S Garwood, 1[43].
=Louisiana.= East Baton Rouge Parish: Baton Rouge, 4[71]. Livingston Parish: Springville, 1[74]. Saint James Parish: Convent, 1[91]; Remy, 2 (1[74], 1[45]). Assumption Parish: near Lake Verret, 1[45].
=Mississippi.= Harrison County: Saucier, 1[71].
=Mustela frenata olivacea= Howell
Long-tailed Weasel
Plates 16, 17, 18, 31, 32 and 33
Mustela peninsulae olivacea Howell, Proc. Biol. Soc. Washington, 26:139, May 21, 1913.
Mustela frenata olivacea, Hall, Carnegie Instit. Washington Publ. 473:104, November 20, 1936.
Type.--Male, adult, skull and skin; no. 180802, U. S. Nat. Mus., Biol. Surveys Coll.; Autaugaville, Autauga County, Alabama; December 22, 1912; obtained by L. S. Golsan.
The skull (plates 16-18), although cracked at two places in the interorbital region, is in one piece and not warped out of shape. The teeth all are present and entire. The skin is exceptionally well made and in perfect condition except for the extreme tip of the tail which is broken off.
Range.--Lower and Upper Austral life-zones in eastern Mississippi, Alabama, Georgia, South Carolina, and northern Florida. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. peninsulae in finer, softer pelage and shorter (less than 15.8 in ad. [F]) tympanic bullae; from M. f. noveboracensis, in adult males by wider tympanic bulla which is more than rather than less than 8.5, in adult females by total length which is more than rather than less than 345, and by mastoid breadth which is more than rather than less than distance between articular faces of exoccipital condyle and glenoid fossa; from M. f. arthuri in larger size (adult males with hind foot and basilar length each more than 45); depth of skull at anterior margin of basioccipital, ignoring sagittal crest, amounting to less than 63 per cent of mastoid breadth, and lesser convexity of dorsal outline of skull in longitudinal axis (See pls. 16-18).
Description.--Size.--Male and female: External measurements of adults are available as follows:
Length Length Catalogue Sex Locality Total of of hind no. length tail foot 47165 [M] Box Springs, 454 160 48 Talbot Co., Georgia
47166 [M] Box Springs, 435 147 47 Talbot Co., Georgia
47167 [M] Box Springs, 422 145 45 Talbot Co., Georgia
41023 [M] Thomas Co., 443 140 47 Georgia
41025 [M] Grady Co., 395 142 47 Georgia
223880 [M] Okefinokee Swamp, 416 145 49 Georgia
198 [M] Okefinokee Swamp, 425 140 48 Georgia
Average 7 [M] 427 146 47
49385 [F] Gainesville, 396 124 45* Alachua Co., Florida
41024 [F] Thomas Co., 380 125 41 Georgia
51527 [F] Talbot Co., 376 128 43 Georgia * [not typical]
The length of the hind foot averages less than the basal length in both males and females. The tail averages 52 per cent as long as the head and body in males and 51 per cent in females. Average differences in measurements of the two sexes are: Total length, 49; length of tail, 19; length of hind foot, 5. An adult male, no. 41023, and an adult female, no. 41024, each taken in February, 1929, on the Sinkola Plantation, Thomas County, Georgia, weighed 15 ounces (425 grams) and 7 ounces (198 grams) respectively according to Charles O. Handley.
Externals.--As described in Mustela frenata noveboracensis, except that hairiness of foot-soles slightly less than shown in figure 19.
Color.--Upper parts, in summer, near tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304. In winter lighter, between tones 3 and 4 of Raw Umber of Oberthür and Dauthenay, pl. 301. Dark spot at each angle of mouth present or absent. Underparts ranging from Massicot Yellow to Cream Buff except on chin and upper lips which are white. Tip of tail black. Upper parts of uniform color. Color of underparts extends distally on posterior sides of forelegs over antipalmar faces of toes and on medial sides of hind limbs to ankles. Least width of color of underparts averaging, in a series of five males from Talbot Co., Georgia, 29 (extremes 24-34) per cent of greatest width of color of upper parts. Black tip of tail in same series, averaging 65 (extremes 60-70) mm. long, thus longer than hind foot and averaging 43 per cent of length of tail-vertebrae.
The spot at the angle of the mouth is absent in one-third of the specimens examined. The upper lips are white in specimens from the southern part of the range of olivacea but in the northern part of the range of the subspecies the upper lips are dark colored as in noveboracensis.
Skull and teeth.--Male (based on 5 adults from Talbot Co., Georgia): See measurements and plates 16-18; weight, 5.3 (5.0-6.4) grams; basilar length, 48.3 (45.8-50.1); zygomatic breadth more or less (usually less) than distance between condylar foramen and M1 and more or less (usually more) than distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less than (averaging about equal to) postpalatal length; postorbital breadth less than length of upper premolars and more or less than (about equal to) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than (about equal to) distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than greatest length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla not less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum (one exception); anterior margin of masseteric fossa below posterior half of m2.
Female (based on 2 adults from Thomas Co., Ga., and one from Talbot Co., Ga.): See measurements and plates 31-33; weight, 3.8 (3.5-4.0) grams; basilar length, 43.4 (42.7-44.0); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less (usually more) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including I3) upper incisors; height of tympanic bulla not less (usually more) than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum.
The skull of the female averages 28 per cent lighter than that of the male.
Compared with the skull of M. f. peninsulae, of which only one good skull, and that a female, is available, that of M. f. olivacea averages smaller and has relatively and actually smaller and less inflated bullae. As compared with the skull of M. f. noveboracensis, that of olivacea in the case of males is larger in every part measured and relative to the basilar length is broader across the zygomatic arches and mastoids. However, the rostrum and interorbital region are relatively narrower. The orbitonasal length is relatively less. The tympanic bullae are broader and more inflated. The same differences hold as between females of noveboracensis and olivacea. Indeed, the females of these two races differ more than do the males. Additional, selected differential cranial characters in the females are, in olivacea, as follows: Weight averaging 3.8 grams rather than 1.7 grams; braincase with, rather than without, sagittal crest; anterior border of tympanic bulla separated from foramen ovale by breadth of less than, rather than breadth of more than, 4 upper incisors (including I3); height of tympanic bulla not less than, rather than less than, distance from its anterior margin to foramen ovale; squamosal bone, between anterior margin of tympanic bulla and foramen ovale, ventrally concave rather than ventrally convex. Comparisons of the skulls with those of M. f. arthuri and M. f. primulina are made in the accounts of those subspecies.
Remarks.--Excepting two young specimens from South Carolina in the Charleston Museum, no specimens of this race of large weasel seem to have been preserved until Arthur H. Howell, in the course of his study of the mammals of Alabama, procured specimens on which his name, olivacea, was based. Later, Francis Harper obtained three instructive specimens from Okefinokee Swamp. Really adequate material, for the localities represented, owes its preservation to the alertness of Charles O. Handley, when he resided at Thomasville, Georgia, and to Hallie E. Fuller of Geneva, Talbot Co., Georgia.
The distinctness of M. f. olivacea from M. f. peninsulae is not satisfactorily established due to inadequate material of peninsulae. Differences shown by the specimens seen indicate that, as compared with olivacea, peninsulae is larger, has transversely wider light-colored underparts which possess more yellow, and a larger skull with more inflated tympanic bullae. In each of these characters, olivacea is intermediate between noveboracensis on the north and peninsulae on the south. The question arises, therefore, whether the animals here recognized under the name olivacea really constitute a recognizable subspecies or instead are only representatives of a subspecies which reaches its extreme development in Florida. In the latter event, the name peninsulae would apply to all. Examination of more material from Florida, especially from the southern half of Florida, will be necessary to answer this question.
This large weasel of the southeastern United States is remarkably different from noveboracensis. Indeed, were it not for actual intergrades such as the two from Fort Payne, Alabama, and York, South Carolina, which are described in the account of M. f. noveboracensis, and the six specimens from northwestern Alabama, which are referred to olivacea, the systematist, I believe, would have little or no hesitancy in designating the two as distinct species, especially on the basis of differences to be seen in the skull.
Not only are the two forms structurally more different than usually is the case but between two geographically, adjacent subspecies of the same species of mammal, but the belt where intergradation occurs appears to be narrow. Nevertheless, when material of the two races is laid out in geographic order, and examined in mass, certain features are seen to undergo gradual change as a person's eye travels from specimens from, say, the center of the range of noveboracensis to specimens from southern localities adjoining the territory occupied by olivacea. One of these features subject to gradual change is the color of the underparts. Beginning at the Adirondacks of New York where a large number of the specimens have white underparts, the underparts become more intensely yellowish southward through the range of noveboracensis into that of olivacea. Indeed, this progressive trend seems to continue right on southward through the range of olivacea into that of peninsulae. Turning in the opposite direction we find that the least width of the underparts decreases gradually northward toward the range of noveboracensis. There is, likewise, a decrease to the northward in length of the skull and relative, as well as actual, narrowing of the braincase and tympanic bullae. However, in least width of color of underparts and the mentioned cranial features, the trend stops relatively abruptly at the southern boundary of the geographic range of noveboracensis and does not continue on, northward, into the range of noveboracensis as is the case with the change in intensity of yellowness of the underparts.
Two males, in the United States National Museum, Biological Surveys Collection, from near Leighton, Alabama, no. 178386 from the Tennessee River nine miles north [of Leighton?] and no. 180240 from La Grange Mountain, although clearly referable to olivacea on the basis of cranial characters, show some approach to noveboracensis in lesser size of the skull and agree with noveboracensis in the narrowness of the color of the underparts. Also, these specimens, like others from the northern part of the range of olivacea, for instance, no 31.227, Charleston Museum, from Mayesville, South Carolina, have the color of the underparts extended only part way out on the hind limb toward the foot. In specimens of olivacea from the southern part of its range the color of the underparts is extended onto the hind feet and this trend reaches its extreme in peninsulae, specimens of which have the feet and larger parts of the limbs marked with the light color of the underparts.
An adult female, no. 32.32, Charleston Museum, although typical of olivacea in most respects, is nevertheless an intergrade. The teeth are as small as in some specimens of noveboracensis. The size of the skull is only slightly nearer that of olivacea than it is to that of noveboracensis. The proportions of the skull, however, are distinctly those of olivacea.
Five other specimens, from northwestern Alabama, namely two from eight miles north of Nauvoo, two from Shoal Creek, and one from White Creek, also show intergradation between noveboracensis and olivacea. The remarks concerning color and color pattern of the specimens from Leighton apply equally well to the five from northwestern Alabama. In cranial characters, no. 51658 from Shoal Creek is referable to olivacea, as also is no. 51677 from the same place, providing it is a female rather than a male as sexed by the collector. No. 57146 from White Creek also is referable to olivacea although the skull shows some approach to that of noveboracensis. Of the two males from near Nauvoo, no. 51652 is to me indistinguishable from noveboracensis, but no. 51653 does have some characters of olivacea, although on the whole, the latter, too, seems to be a little nearer noveboracensis than olivacea. However, because the mean of these seven specimens from northwestern Alabama is nearer olivacea than noveboracensis the former name may be applied.
Another specimen from "Souinlonie" Creek, Clark County, Mississippi, has the coloration and rostral configuration of primulina, narrow mastoidal breadth and smaller teeth of noveboracensis and skull of large size with "full" braincase as in olivacea. No. 235364, U. S. Nat. Mus., from the Mobile River at the "L. and N. RR. Crossing," Mobile County, Alabama, although definitely olivacea, shows approach to arthuri in that the dorsal outline of the skull is longitudinally more convex and the tympanic bullae are less inflated than in olivacea and in that the color of the underparts is almost exactly as in the type specimen of arthuri. The young specimen labeled as from "Silver Springs," Florida, has large tympanic bullae (17 mm. long) and several characters that show its relationship to peninsulae as that race is now understood. Because the sex is unknown the identification as olivacea is tentative and is made on the assumption that the specimen is a male. If it is instead a female, the animal is referable to peninsulae.
An adult, female specimen in the Charleston Museum, no. 27.239.1, taken at St. Matthews, South Carolina, on December 8, 1927, contained four embryos which averaged 19 mm. in length and 47.75 centigrams in weight. Another adult female, in the Charleston Museum, no. 32.32, taken on February 21, 1932, at the same place, has prominent mammae, and the collector has noted that two were slightly active.
Sixteen of twenty-nine adults examined show infestation of the frontal sinuses by parasites. However, in none is the malformation of the frontal region so great as frequently occurs in M. f. noveboracensis.
Specimens examined.--Total number, 52, arranged alphabetically by states and from north to south by counties in each state. Except as otherwise indicated specimens are in the University of California Museum of Vertebrate Zoölogy.
=Alabama.= Lawrence County: White Creek, 1; Little Sand Mt., Shoal Creek, 2. Winston County: 7-1/2 mi. N Nauvoo, 1; 8 mi. N Nauvoo, 1. Lauderdale County: near Leighton, 9 mi. N Tennessee River, 1[91]. Colbert County: Leighton, 1[91]. Autauga County: Autaugaville, 1[91]. Dale County: Midland City, 1[91]. _Mobile County_: Mobile River, 12 mi. NE Mobile, 1[91].
=Florida.= Alachua County: Gainesville, 4[61]. Marion County: "Silver Springs," 1.
=Georgia.= Spalding County, 1. Lamar County, 1. Talbot County: southwest part of county, 1; Box Springs, near Geneva, 3; Upatoie Creek, 1 mi. SW Box Springs, 2; 3 mi. SE Geneva, 1; 4 mi. W Geneva, 1; 5 mi. W Geneva, 1; 2 mi. E Geneva, 1. Chattahoochee County, 2. Grady County: Beachton, 3[91]; locality no more definite than county, 4. Thomas County: Sinkola Plantation, 2; locality no more definite than county, 2. Charlton County: 1/2 mi. E Chesser's Island, Okefinokee Swamp, 1[58]. County in question: Billy's Island, Okefinokee Swamp, 1[91]; Okefinokee Swamp, 1[58].
=Mississippi.= Clark County: Souinlonie Creek, 1.
=South Carolina.= Darlington County: Society Hill, 1[91]. Sumter County: Mayesville, 1[11]. Calhoun County: St. Matthews, 2[11]. Georgetown County: Sampit, 1[11]. Charleston County: Rantowles, 1[11]; 8 mi. N Charleston, 1[11]. _Beaufort County_: Yemassee, 1[2].
=Mustela frenata peninsulae= (Rhoads)
Long-tailed Weasel
Plates 16, 17 and 18
Putorius peninsulae Rhoads, Proc. Acad. Nat. Sci. Philadelphia, 1894:152, June 19, 1894; Bangs, Proc. Biol. Soc. Washington, 10:10, February 25, 1896.
Mustela peninsulae, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912.
Mustela p. peninsulae, Bailey, Bailey Mus. and Library Nat. Hist., 1(no. 5):1, December 1, 1930.
Mustela frenata peninsulae, Hall, Carnegie Instit. Washington Publ. 473:105, September 20, 1936.
Type.--Female, young, part skull and skin; no. 8515, Acad. Nat. Sci. Philadelphia; Hudson's, Pasco County [14 miles north of Tarpon Springs], Florida; before 1895; obtained by W. S. Dickinson.
The skull has been cut vertically in two at the plane of the glenoid fossae. These fossae and all the cranium posterior to them are missing. In addition to the part of the cranium anterior to the glenoid fossae, the lower jaws are preserved complete. The teeth all are present and entire. The prominent sutures on the rostrum and palate show the specimen to be young and its small size leaves but little doubt that the animal was a female. The light facial markings are more extensive than in any of the referred specimens. In the type these light facial markings consist of a median isolated spot immediately in front of the ears, a larger one on the nose, with an interrupted bar on each side extending posteroventrally in front of and anterior to the eye, a wider bar, on each side, extending anterodorsally between the ear and eye and finally an isolated spot at the anterior border of each ear. The skin is stuffed and in fair condition except that the vertebrae remain in the tail.
Range.--Austral and probably Tropical life-zones of Florida south of latitude 29°. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. olivacea in coarser pelage and larger tympanic bullae.
Description.--Size.--Male: No external measurements available. Female: The type a young animal and no. 2379, an adult from Tarpon Springs, measure respectively as follows: Total length, 375, 378; length of tail, 100, 130; length of hind foot, 40, 44.5.
Externals.--As described in Mustela frenata noveboracensis except that hairiness of foot-soles as shown in figure 20.
Color.--Upper parts (in winter) near tone 3 of Burnt Umber of Oberthür and Dauthenay, pl. 304. Dark spot at each angle of mouth present or absent. Tip of tail black. Underparts Reed Yellow except on chin and usually on legs where white. Upper lips white entirely around. Upper parts of uniform color. Color of underparts extends distally on legs over both sides of feet and on front legs over wrists. Proximal part of tail slightly lighter below than above. Least width of color of underparts, in seven specimens, averaging 41 (extremes 31-52) per cent of greatest width of color of upper parts. Black tip of tail, in each of two females, 45 mm. long; thus slightly longer than hind foot and amounting to 36 per cent of length of tail-vertebrae.
The spot at the angle of the mouth is absent in four of the ten specimens and is present on both sides in the other six.
Skull and teeth.--Male (based on an adult from Apopka and the anterior part of an adult from Enterprise): See measurements and plates 16-18. As described in Mustela frenata olivacea except that: Weight, 7.0 grams; basilar length, 49.8.
Female (based on an adult from Tarpon Springs, Florida): See measurements. As described in Mustela frenata olivacea except that: Weight, 4.7 grams; basilar length, 44.2; zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla.
In comparison with M. f. olivacea, the insufficient material of M. f. peninsulae suggests that its skull averages larger and has relatively as well as actually larger and more inflated tympanic bullae.
Remarks.--The first published mention of this weasel seems to have been the original description which appeared in 1894. This description was based on a single specimen sent to Samuel N. Rhoads by W. S. Dickinson, who, in the following year, procured another specimen at Tarpon Springs. So far as known only eight other specimens, as listed under "Specimens examined," have found their way into collections of study specimens.
H. H. Bailey (1930:1) credits the range of this subspecies as extending south "to the shores of Florida Bay and the Gulf of Mexico, where ever high ground occurs."
Evidence of intergradation between M. f. peninsulae and M. f. olivacea is provided by specimens of olivacea from Gainesville, Florida, and the Okefinokee Swamp, Georgia. These specimens, on the average, have the color of the underparts wider, the skull larger, and the tympanic bullae relatively larger than do specimens of olivacea from farther north. In these features, approach to M. f. peninsulae is shown.
Light facial markings occur in this subspecies. They are similar to those possessed by weasels which occur at the same latitude and under corresponding climatic conditions on the Pacific Coast. The type specimen and one from Tarpon Springs have white facial markings. Two of the three specimens from Apopka also show white facial markings, although in reduced amount. One of the four specimens of M. f. olivacea from Gainesville, Florida, has well-developed light (white) facial markings. Also of the four specimens of M. f. olivacea examined from Okefinokee Swamp, Georgia, one has prominent white facial markings. However, in it the pattern is so unusual as to suggest that it is an instance of partial albinism rather than an outcropping of a racial tendency, or a pattern of coloration induced by climatic factors.
None of the eight available skulls show any infestation of the frontal sinuses by parasites.
Specimens examined.--Total number, 10, arranged by counties from west to east.
=Florida.= Pasco County: Hudson's, 1[1]. Pinellas County: Tarpon Springs, 1[1]. Hernando County, 1[91]. Polk County: Auburndale, 1[91]; no locality more definite than county, 1[91]. Orange County: Apopka, 3[61]. Volusia County: Enterprise, 1[60]. Seminole County: Osceola, 1[2].
=Mustela frenata spadix= (Bangs)
Long-tailed Weasel
Plates 16, 17, 18, 31, 32 and 33
Putorius longicauda spadix Bangs, Proc. Biol. Soc. Washington, 10:8, February 25, 1896; Merriam, N. Amer. Fauna, 11:21, figs. 10, 11, June 30, 1896; Cory, Mamm. Illinois and Wisconsin, p. 374, 1912.
Mustela longicauda spadix, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912; Bailey, Journ. Mamm., 10:156, May 9, 1929.
Mustela longicauda, Johnson, Journ. Mamm., 11:439, November 11, 1930.
Mustela noveboracensis, Murie, Journ. Mamm., 16:321, November 15, 1935.
Mustela frenata spadix, Hall, Carnegie Instit. Washington Publ. 473:105, November 20, 1936.
Type.--Male, young, skull and skin; no. 3265/1786, Amer. Mus. Nat. Hist.; Fort Snelling, Hennepin County, Minnesota; June 25, 1889; obtained by Edgar A. Mearns; original no. 812.
The skull is complete although there are fractures on the top of the braincase, on the right side of the braincase and at the middle of the right zygomatic arch. The teeth all are present and entire. The skin, although overstuffed, is complete, well preserved, and in summer pelage.
Range.--Upper Austral and Transition life-zones of Minnesota, northern and western Iowa, southeastern North Dakota, eastern part of South Dakota, and northeastern Nebraska. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. noveboracensis and M. f. primulina in that specimens of all ages have least width of color of underparts amounting to more than 41 per cent of greatest width of color of upper parts, and have light color of underparts extended onto hind foot rather than stopped short of ankle; adults with hind feet more than 50 in males and 40 in females; orbitonasal length more than 15.5 in males and 13.5 in females; length of tooth-rows more than 18.0 in males and 15.7 in females; mastoid breadth more than 25.5 in males and 22.0 in females. From M. f. longicauda by color darker than near (h) Clay Color, in males by a flattened occiput in which the depth of the skull, exclusive of the sagittal crest and taken at the anterior border of the basioccipital, amounts to less than 58 per cent of the mastoid breadth.
Description.--Size.--Male: Three adults from Elk River, Minnesota, yield average and extreme measurements as follows: Total length, 458 (444-467); length of tail, 154 (140-165); length of hind foot, 55 (52-59). Tail averages 51 per cent as long as head and body. Length of hind foot averages more than basal length. Corresponding measurements of three subadults from Madison, Minnesota, are as follows: 453 (438-469); 157 (152-165); 50 (47-51). Tail averages 53 per cent as long as head and body.
Female: Three adults from Elk River, Minnesota, yield average and extreme measurements as follows: Total length, 387 (380-391); length of tail, 131 (121-138); length of hind foot, 44 (43-46). Tail averages 51 per cent as long as head and body. Length of hind foot more or less than (approximately equal to) basal length. Corresponding measurements of two adults and one subadult from Madison, Minnesota, are as follows: 385 (379-396); 137 (119-159); 42 (38-44). Tail averages 55 per cent as long as head and body.
The average differences in external measurements of the two sexes from Elk River, are: Total length, 71; length of tail, 23; length of hind foot, 11. At Madison, corresponding differences are 68, 20, and 8. Two adult females from Elk River, Minnesota, weigh 205 and 210 grams.
Externals.--Longest facial vibrissae black, brown, or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) as shown in figure 19.
Color.--Winter pelage all white except tip of tail. In southern part of range sometimes assumes a brown winter coat. Summer pelage with upper parts ranging from near (16´) Cinnamon Brown to Vandyke Brown. Chin and upper lips white. Remainder of underparts ranging from near (a) Olive Ocher to Ochraceous Buff and Pale Orange Yellow. Tip of tail at all times black. Upper parts of uniform color except for occasional slight darkening of nose. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and ankles, on medial sides of hind limbs to ankle, over antiplantar faces of toes and distomedial fourth of each tarsus, and over proximal fifth to third of under side of tail. Least width of color of underparts averaging (in 3 specimens from Elk River) 54 (47-59) per cent of greatest width of color of upper parts. Black tip of tail averaging same length as hind foot and 28 per cent of length of tail-vertebrae. Save for the greater width of the light-colored underparts and relatively short black tip of the tail, both features of M. f. longicauda, spadix is variously intermediate, depending on locality, as between noveboracensis and longicauda.
Skull and teeth.--Male (based on 3 adults from Elk River, Minn.): See measurements and plates 16-18. As described in Mustela frenata longicauda except that: Weight, 5.6 (5.0-6.5); basilar length, 49.0 (48.7-49.2); zygomatic breadth sometimes less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more or less (about equal to) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 5 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of rostrum; anterior margin of masseteric fossa below talonid of m1.
Female (based on 4 adults from Elk River, Minn.): See measurements and plates 31-33. As described in Mustela frenata longicauda except that: Weight, 3.5 (3.3-4.0) grams; basilar length, 42.9 (42.3-43.2); least width of palate more or less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors.
The skull of the female averages 33 per cent lighter than that of the male.
Skulls of adult males of spadix from Elk River, Minnesota, as compared with those of longicauda from Alberta, are larger in every part measured. Relative to the basilar length these skulls of spadix are broader across the mastoid region, narrower across the zygomata, deeper through the plane of the postorbital processes, shallower through the braincase and have relatively shorter tympanic bullae. Whereas the tympanic bullae of longicauda are, on the average, approximately as long as the rostrum (orbitonasal length), in spadix the rostrum is longer than the bulla. Viewed posteriorly, the braincase of spadix is seen to be much shallower and wider than that of longicauda. Indeed, the depth of the braincase, measured at the anterior end of the basioccipital, amounts to only 56 per cent of the mastoid breadth in spadix as against 61 per cent in longicauda. The longer, waistlike, postorbital constriction, relatively smaller braincase, and especially the relatively narrower zygomatic expanse in spadix imparts to its skull a more slender appearance than has the skull of longicauda. These differences are not shown by the skulls of females. To be sure, spadix, in most of its cranial measurements, averages slightly larger, has a relatively shallower braincase and is relatively deeper through the postorbital processes, but these differences are so slight that inclusion of one more specimen, of slightly different proportions, in the average might cause the average measurements to read as they do in longicauda.
Compared with noveboracensis, from Massachusetts, adult skulls of spadix, taking sex into account, are larger in every part measured and are relatively as well as actually wider and deeper throughout. Also, in spadix: Sagittal and lambdoidal crests higher, especially in females; anterior margin of tympanic bulla projecting up sharply from squamosal; occiput more flattened in posterior view; tooth-rows relatively and actually longer but orbitonasal length relatively shorter; postorbital processes more robust; zygomatic arches widely bowed outward rather than evenly rounded; canines larger; squamosal less swollen ventrally, especially in females. Between noveboracensis and spadix, the differential cranial characters are greater in number and degree between females than between males. Comparison of the skull with that of M. f. primulina is made in discussion of that subspecies.
Remarks.--Edgar A. Mearns in 1889 and the early nineties took several specimens of this weasel and it was principally on these that Bangs in 1896 (p. 8) based his description. The best material, however, is that from Elk River, Minnesota, collected in later years by Bernard Bailey, and supplemented by one specimen taken in 1885 by Vernon Bailey and another by his sister Anna Bailey in 1891 at the same place.
Mustela frenata spadix has just one structural feature of a "unique" kind which serves to differentiate it from the geographically adjoining subspecies. This feature is large size. The other diagnostic characters ascribed to spadix are of an intermediate sort--intermediate as between two extremes, one found to the westward in longicauda and the other to the eastward in noveboracensis. For example, the dark-colored upper parts are merely darker than in longicauda and merely lighter than in noveboracensis. The color is not "different"; it is only "intermediate." Furthermore, each of the characters ascribed to spadix, including large size itself, undergoes change from one part of its geographic range to another; the characters are not constant over a wide area. Indeed, excepting the large size which remains relatively uniform over the northern two-thirds of the range, no two localities have been found from which the specimens can be said really to agree in characters.
By way of illustration, the coloration of the upper parts may be cited. Near the range of noveboracensis the average coloration of individuals from one locality is only a little lighter than in noveboracensis. Farther westward the average coloration is a little lighter and farther westward yet, toward the range of the extremely light colored longicauda, the average coloration is lighter still. Although all these animals are darker than longicauda and lighter than noveboracensis, those from the three places do not agree among themselves. Because of the lack of more than one character of a "unique" kind and because of the inconstancy, geographically, of other characters, and for that matter, lack of constancy geographically in combination of characters, the writer regards spadix as a barely recognizable subspecies.
Examination of the specimens of spadix shows that the individual variation in a single species is greater in a region of intergradation than it is some distance inside the borders of the geographic range of a well-marked subspecies. This is illustrated by three specimens of M. f. spadix in fresh summer pelage from the single locality, Elk River, Minnesota. In these, the color of the upper parts varies from a little darker than Cinnamon Brown to Vandyke Brown. At any one locality well within the range of longicauda, or noveboracensis, there is nowhere nearly so much variation in color, even in much larger series of specimens.
Study of the specimens here assigned to spadix reveals that some features regarded as of diagnostic value for one or the other of the two races, longicauda and noveboracensis, behave differently. For example, the dark coloration of the upper parts, which is characteristic of noveboracensis, manifests itself far westward within the range of spadix whereas the wider extent of the light-colored underparts, which is characteristic of longicauda, and the Olive Ocher, rather than Pale Orange Yellow, color of these underparts, are seen in varying degree all the way across the range of spadix. Thus, these animals are colored above like noveboracensis and below like longicauda, but not vice versa. In these animals, then, the longicauda type of underparts is dominant, in one sense of the word, over the noveboracensis type of underparts, and the noveboracensis type of upper parts is dominant over the longicauda type of upper parts. Each of these features is subject to actual intergradation and does not always behave as a "unit character," that is to say, one which is either present or absent. However, the noveboracensis type of upper parts is carried much farther west before being diluted than is the noveboracensis type of underparts. Indeed, within the range of noveboracensis itself, the broad extent of the longicauda type of underparts is manifest. This is, of course, near the western margin of the range of noveboracensis.
The large size of males of spadix, as exemplified by specimens from Elk River (see measurements on p. 421), seems to be retained across the northern part of the range here assigned to the subspecies. This larger size than is found in longicauda from Alberta, is shown also by some specimens from eastern North Dakota which are assigned to longicauda. However, the average of these Dakotan specimens, all characters considered, is nearer to my concept of longicauda.
Inspection of the cranial measurements of spadix shows also that in addition to its large size it is distinguishable from any one of the geographically adjoining races by its relatively (to basilar length) greater, as well as actually greater, mastoidal breadth. This might be included with size as a unique character distinguishing spadix from longicauda and noveboracensis. However, it is not clear whether or not this greater mastoidal breadth is more than a function of the large size.
Excepting the greater mastoidal breadth and generally larger size of the skull, the cranial features distinguishing males of spadix from longicauda are features in which spadix shows approach to noveboracensis. This is true, in spadix, of the relatively longer (in comparison with longicauda) rostrum, relatively lesser zygomatic breadth, relatively shallower braincase measured at the anterior end of the basioccipital, and relatively deeper skull as measured at the posterior borders of the last upper molars. This same approach to noveboracensis already has been pointed out with respect to color of the upper parts and is evident also in the relative shortness of the tail which averages only 51 per cent of the length of the head and body rather than 55 per cent as in longicauda.
Because the longicauda type of animal previously has been regarded as specifically distinct from the noveboracensis type of animal, comment is offered below on selected specimens, referred to spadix, which are regarded as intergrades with noveboracensis or with other subspecies.
No. 8722, Univ. Wisconsin, adult male, in the white winter coat, from north central Itasca County, Minnesota, obviously has characters of M. f. spadix or longicauda that occur to the west and M. f. noveboracensis of the east. Selected outstanding characters of longicauda are its long tail, anteriorly truncate tympanic bullae and large teeth. Characters indicating its affinities with noveboracensis are smaller size of skull, general narrowness of skull, and relatively low tympanic bullae. The skull is intermediate as regards several individual structural features. For example, although long and narrow and in this feature more nearly approaching noveboracensis, the skull is wider than usual in that subspecies and thus approaches that of longicauda or spadix. The hind foot, in the dried state, measures 47 millimeters. This large hind foot, obviously long tail (the specimen lacks external measurements), and anteriorly truncate bullae constitute basis for here referring the specimen to spadix. However, the seemingly small size of the body and the narrow skull clearly show relationship to noveboracensis.
Specimens, referred to spadix, from northern Iowa, are instructive as showing what happens where the ranges of noveboracensis, primulina, spadix, and perhaps longicauda, meet. No. 47167, Univ. Mich. Mus. Zoöl., a nearly adult female, taken on November 22, 1915, at Island, Clay County, and in process of assuming a brown winter pelage, retains enough of the dark summer pelage to show that the color was slightly lighter than average for spadix. The color pattern, white lips, and extension of light color of the underparts onto the feet, agrees with spadix or longicauda as does also the long tooth-row. The overall length of the skull is intermediate between that of spadix and primulina. The proportions of the anterior part of the skull and of the tympanic bullae resemble those found in primulina. A subadult male skull only, no. 123846, American Museum of Natural History, from Webb, Clay County, shows approach to primulina in the narrowness of the rostrum.
A young male from Ruthven, Iowa, no. 48340, Univ. Michigan, has a large skull approaching in size that of spadix, has the longicauda-spadix type of light-colored underparts and color pattern, and is slightly darker above than true longicauda. Another subadult male in the white winter coat from Palo Alto County, no. 35756, Univ. Michigan, has a large skull, which shows approach to primulina in its narrowness anteriorly and in some other features. Although the tail is of moderate length, the body is large as in spadix or longicauda, and the length of the hind foot suggests spadix or longicauda.
A subadult male, no. 425a, Iowa State College, from Manson, Iowa, in brown winter pelage, agrees with primulina in the restriction of the area of the light color of the underparts and in less expanded zygomatic arches. The teeth are intermediate in size between those of noveboracensis and primulina on the one hand and those of spadix and longicauda on the other. In other respects it agrees with, or is more nearly like, spadix.
An adult female, no. 426a, Iowa State College, from Barnum, in the brown winter coat, agrees with primulina except that the orbitonasal length of the skull is more as in spadix and the presence of some light color on the lower part of the hind legs suggests spadix. The skull only, no. 440a, Iowa State College, labeled merely Webster County, Iowa, is almost a duplicate of no. 426a. A subadult male, no. 427a, Iowa State College, from Moorland, Iowa, only about six miles southeast of Barnum, likewise is indistinguishable from primulina except for having a white winter coat and in being relatively broad in the mastoidal region. Nevertheless, both of these animals are here referred to spadix because the average of specimens from this general area is nearer that of spadix. No. 497a, Iowa State College, an adult female in white winter pelage, from Ames, approaches primulina in the narrow rostrum and smaller teeth but otherwise approaches or even agrees with spadix.
Two adult males, without external measurements, from Pilot Mound, Iowa, have skulls quite like males of longicauda from Alberta. The only approach noted to eastern forms is the restricted color of the underparts on no. 2856, Coe College, which has a brown winter coat. The color of the underparts is not extended so far out on the feet as in longicauda. Also the tympanic bullae of this specimen are a trifle narrower. The other male, no. 2652, is in the white winter coat. The one female from the same place, no. 2660, Coe College, in brown winter pelage, has a skull notably unlike that of longicauda or spadix; the skull is narrower and practically indistinguishable from that of the largest female skull of primulina available from Lawrence, Kansas, save that the tooth-row is much longer. The color pattern also agrees with that of primulina or noveboracensis in that the color of the underparts extends only as far as the knee on the hind legs and is narrow on the belly. Nevertheless, another adult female, no. 120a from Amaqua Township, some 6 miles southwest of Pilot Mound, is in all respects typical of spadix. This is the more remarkable because another comparable specimen from less than 20 miles to the southwest in Worth Township is equally typical of primulina.
Two young females from Chester, Iowa, nos. 2656 and 2874/2873, Coe College, have skulls larger than those of corresponding age of primulina or noveboracensis. The color is as in spadix. The color pattern of the underparts also is as in spadix or longicauda except that the width of the area of light color on the belly is restricted somewhat although not so much as in noveboracensis or primulina. Of four males from the same place, also in the collection of Coe College, no. A2874 is a white skin only and does not provide diagnostic characters. The three other males, each in summer pelage, are marked and colored as are the two females from the same place except that male no. 2861 has the color of the underparts so much attenuated on the hind legs that it barely, uninterruptedly, extends to the feet. No. 2658 is young, or perhaps barely subadult. The skull is large and referable to spadix. The two adults, nos. 2861 and 2657, differ cranially from typical (Elk River, Minn.) spadix only in being slightly narrower across the mastoids and in having the bullae a little narrower. In these departures they show some approach to primulina and to noveboracensis. Another male, subadult, no. 2867, Coe College, from Decorah, which has acquired half of the white winter coat, agrees with the males from Chester except that the preorbital part of the skull is shortened much as in some specimens of primulina.
From Lansing, in extreme northeastern Iowa, a large subadult male, no. 2864, Coe College, of 453 mm. in total length and half through with acquiring the white winter coat, agrees with the males previously described from Chester except in having the palate narrower as in noveboracensis. The adult female available from Lansing, no. 2863/2862, Coe College, in white winter pelage except for the top of the head, although a large skin, has a skull smaller than that of any spadix or longicauda and of about the same size as that of no. 3838, Univ. Kansas Mus. Nat. Hist., of primulina, from Lawrence, Kansas, except that the skull of no. 2863/2862 is much narrower across the mastoids. This specimen, then, shows approach to noveboracensis in narrowness of the mastoidal region, to primulina in other respects and to spadix.
Many of these instructive specimens from Iowa, made available to the present writer by Mr. W. F. Kubichek, were brought together at the Coe College Museum by the late B. H. Bailey. Most of them were obtained from trappers who did not supply the conventional external measurements taken in the flesh. Even though these are lacking, the specimens clearly show that actual intergradation occurs where the ranges of M. f. longicauda, spadix, noveboracensis and primulina meet.
The dark color of the upper parts, restriction of the color of the underparts on the ankles with the result that the color reaches the toes in interrupted fashion, and large skull, of no. 18912 of the Museum of the University of South Dakota, from Roberts County, South Dakota, clearly place this specimen with spadix, rather than with longicauda. Likewise, male, subadult, no. 11376, Univ. South Dakota, from Clay County, South Dakota, is referable to spadix. Although without external measurements, the specimen obviously is large. The patch of summer pelage on its head and neck is darker than the summer pelage of longicauda, and the orbitonasal length is greater than the length of the tympanic bullae; all these features are characters of spadix. The adult male from Fort Sisseton, South Dakota, no. 188407, United States National Museum, figured by Merriam (1896, p. 20, figs. 7-9), is almost exactly intermediate between longicauda and spadix, although here referred to the latter.
Five specimens, nos. 147375, 147432, 147762, 148720 and 148721, U. S. Nat. Mus., including 3 skulls only from Beemer, Cuming County, Nebraska, are intergrades between M. f. longicauda, M. f. primulina and M. f. spadix. One skin is in white winter pelage and the other, a female, is in summer pelage which in coloration and color pattern agrees with that of spadix. External measurements of the male agree with those of longicauda. Measurements of the female agree with those of spadix except that the tail is shorter as in primulina. The skulls are as long as in longicauda but are more slender than in either longicauda or spadix although nearer the latter in this respect. In dorsal aspect, the skulls especially posteriorly to the orbital region, resemble primulina. All points considered, the animals seem best referred to spadix.
Although the degree of development of certain morphological features has been settled upon as indicative of the race spadix, some doubt remains as to where the western boundary of its range should be shown. This results from the fact that color has been taken into account as one diagnostic feature and this feature is lacking in the white winter specimens which, from the following places, are all that are available: Kittson County, Minnesota; Moorhead, Minnesota; Casselton and Valley City in North Dakota; Armour, South Dakota and Clay County, South Dakota. In summary, more specimens in the summer coat will be required to establish definitely the boundary between the ranges of longicauda and spadix.
Surber (1932:49) has referred to additional specimens of this weasel in the University of Minnesota Museum as from Winona, Hennepin and Isanti counties of that state.
At Elk River, Minnesota, B. Bailey (1929:156) found this species to be about half as abundant as Mustela cicognanii and that it is "more often found in the open timber and about the dry ridges and fields." Of seventeen adult or subadult skulls of this race from Minnesota, ten have obvious marks of infestation of the frontal sinuses. In no skull, however, has the infestation resulted in so much malformation, as occurs in noveboracensis.
Specimens examined.--Total number, 76, arranged alphabetically by states and from north to south by counties in each state.
=Iowa.= Lyon County: Granite, 1[65]. Howard County: Chester, 6[12]. Winneshiek County: Decorah, 1[12]; 8 mi. NE Ossian, 1[76]. Allamakee County: Lansing, 2[12]. Clay County: Island, 1[76]; Webb, 1[2]. Palo Alto County: Ruthven, 1[76]; no locality more definite than county, 1[76]. Calhoun County: Manson, 1[65]. Webster County: Barnum, 1[65]; Moorland, 1[65]; no locality more definite than county, 1[65]. Boone County: Pilot Mound, 3[12]; Amaqua Township, Sec. 19, 1[65]. Story County: Ames, 1[65].
=Minnesota.= Kittson County, 1[2]. Roseau County: 2-1/2 mi. SW Roseau, Jadis Township, 1[14]. Itasca County: T. 61N, R. 26W, 1[104]. Clay County: Moorhead, 2[9]. Atkin County: Atkin, 1[50]. Otter Tail County: Lake Lizzie, 1[9]; Parkers Prairie, 1[57]. Grant County: 3 mi. NW Barrett, 1[76]. Benton (now Mille Lacs?) County: Princeton, 1[91]. Sherburne County: Elk River, 14 (6[59], 4[14], 3[91], 1[74]). Hennepin County: Fort Snelling, 6 (5[2], 1[91]). Carver? County: Chaska, 1[60]. Lac qui Parle County: Madison, 5 (3[91], 2[1]); no locality more definite than county, 2 (1[68], 1[75]). Yellow Medicine County: Wood Lake, 1[2]. Blue Earth County: Rapidan, 1[64]. County in question: Moore Lake, 1[91].
=Nebraska.= Cuming County: Beemer, 5[91].
=North Dakota.= Cass County: Fargo, 1[91]; Casselton, 1[91]. Dickey County: Oakes, 1[91].
=South Dakota.= Roberts County, 1[102]. Marshall County: Fort Sisseton, 1[91]. Douglas County: Armour, 1[14]. Clay County, 1[102].
=Mustela frenata longicauda= Bonaparte
Long-tailed Weasel
Plates 16, 17, 18, 31, 32 and 33
Mustela longicauda Bonaparte, Charlesworth's Mag. Nat. Hist., 2:38, 1838.
Putorius longicauda, Baird, Mamm. N. Amer., p. 169, 1858; Coues, Fur-bearing animals, p. 136, 1877; Bangs, Proc. Biol. Soc. Washington, 10:7, figs. 1, 1a of pls. 1, 2 and 3, February 25, 1896; Merriam, N. Amer. Fauna, 11:19, pl. 3, figs. 3, 3a, 4, 4a, pl. 5, figs. 1, 1a, text figs. 7-9, June 30, 1896.
Mustela longicauda longicauda, Bailey, N. Amer. Fauna, 49:166, January 8, 1927.
Mustela frenata longicauda, Hall, Carnegie Instit. Washington Publ. 473:105, November 20, 1936; Hall, Canadian Field-Nat., 52:108, October, 1938.
Mustela frenata, Sowls, Journ. Mamm., 29:126, May 14, 1948.
Type.--Possibly not in existence. No. 43.3.3.3 [from Carlton House, Saskatchewan] in the British Museum of Natural History has been regarded by several zoölogists as the type. It is a subadult female, skull and skin, from North America. See the account of M. erminea cicognanii for reasons for and reasons against regarding this specimen as the holotype.
No. 43.3.3.3 from the collection of Dr. John Richardson is in the white winter coat and now (Sept. 24, 1937) is prepared as a study skin. Evidences of its once having been mounted are: holes in the soles of the hind feet for supporting-wires, large straight wire in the tail, folds in the skin of the now backward-projecting hind feet, and unevenness of the skin on the back resulting from straightening out the specimen. The tip of the tail and some skin from the middle of the belly are missing. Otherwise the skin is intact. The skull is that of an animal in its first year, lacks the zygomatic arch on each side, but otherwise is complete and unbroken. The teeth all are present and entire except that p2 on the right side is missing from its alveolus.
Range.--Transition and Upper Sonoran life-zones of the Great Plains, southward from central Alberta, Saskatchewan and southern Manitoba through eastern Montana, the Dakotas and Nebraska into southeastern Wyoming, northeastern Colorado and western Kansas. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. primulina in near (h) Clay Color rather than Brussels Brown of upper parts, least width of color of underparts more than 40 per cent of greatest width of color of upper parts, color of underparts extended onto hind foot rather than stopped short of ankle, zygomatic breadth more than 28.8 in adult males and more than 24.1 in adult females; from M. f. spadix in lighter color being near (h) Clay Color, in males by deeper occiput in which the depth of the skull, exclusive of the sagittal crest and taken at the anterior border of the basioccipital amounts to more than 59 per cent of the mastoid breadth; from M. f. oribasus in near (h) Clay Color rather than near (14_n_) Brussels Brown color of the upper parts and in males by deeper occiput in which the depth of the skull, exclusive of the sagittal crest and taken at the anterior border of the basioccipital, amounts to more than 59 per cent of the mastoid breadth; from M. f. alleni in larger size, adult males having a total length of more than 400 millimeters, hind foot more than 45, basilar length more than 43.5, and females having a total length of more than 375 and basilar length not less than 40.0; from M. f. nevadensis in near (h) Clay Color rather than near (14_n_ to 1) Brussels Brown of upper parts, basilar length more than 40 in females and averaging more than 45 in males; from M. f. neomexicana by near (h) Clay Color rather than Buckthorn Brown color of upper parts, absence of white and Argus Brown facial markings, and length of tooth-rows amounting to more than 37 per cent of basilar length.
Description.--Size.--Male: Five adults from Alberta yield average and extreme measurements as follows: Total length, 438 (418-473); length of tail, 158 (140-193); length of hind foot, 50 (46-54). Tail averages 56 per cent as long as head and body. Length of hind foot averaging more than basal length. Corresponding measurements of five adults and subadults from North Dakota are as follows: 465 (445-516); 164 (150-179); 51 (50-54). Tail averages 55 per cent as long as head and body.
Female: Six adults (Alberta, 4; Saskatchewan, 1; Manitoba, 1) yield average and extreme measurements as follows: Total length, 401 (383-425); length of tail, 145 (141-159); length of hind foot, 43 (41-44). Tail averages 57 per cent as long as head and body. Length of hind foot more or less than (approximately equal to) basal length.
The average differences in external measurements of the two sexes are: Total length, 37; length of tail, 13; length of hind foot, 7. General comparisons indicate that the Alberta-taken males may not attain so large a size as those from some other areas. Thus the differences in external measurements might be some greater elsewhere, say, in North Dakota.
Externals.--Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) only slightly greater than shown in figure 20.
Color.--Winter pelage all white except tip of tail. Summer pelage with upper parts near (h) Clay Color or near tone 3 and 4 of Snuff Brown of Oberthür and Dauthenay, pl. 303. Chin and upper lips white. Remainder of underparts ranging from near (a) Olive Ocher to near (16´) Ochraceous Buff. Upper parts of uniform color except for occasional darkening of head in front of ears. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind limbs to ankles over antiplantar faces of toes and distomedial third of each tarsus, and over proximal fourth to third of under side of tail. Least width of color of underparts averaging, in a series of 10 males from Alberta, 58 (45-60) per cent of greatest width of color of upper parts. Corresponding figures for 10 females from the same place are 57 (50-74). Black tip of tail in same series of males, most of which are in full summer pelage, averaging 43 (35-60) mm. long. Thus, averaging shorter than hind foot and 27 per cent of length of tail-vertebrae.
As compared with M. f. neomexicana, longicauda lacks the white facial markings, black ears, black forehead and nose, but otherwise is similarly colored. As compared with M. f. nevadensis, M. f. oribasus and M. f. spadix, each of color pattern similar to longicauda, selected differences of longicauda are its much lighter color, especially of the upper parts, with less conspicuous darkening on the nose. From M. f. primulina, longicauda differs in lighter color of upper parts, reddish rather than yellowish underparts, and light rather than dark-colored hind feet.
Skull and teeth.--Male (based on 5 adults from Alberta): See measurements and plates 16-18; weight, 4.7 (4.6-4.9) grams; basilar length, 46.0 (44.7-46.8); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more than postpalatal length; postorbital breadth less than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum more or less (usually less) than length of tympanic bulla; least width of palate less than greatest length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below talonid of m1 or anterior half of m2.
Female (based on 5 adults: Alberta, 3; N. D., 1; Sask., 1.): See measurements and plates 31-33; weight, 3.1 (2.8-3.5) grams; basilar length, 42.3 (40.0-43.7); zygomatic breadth more or less (approximately equal to) than distance between condylar foramen and M1 or that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate not more than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including I3) upper incisors; height of tympanic bulla not less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-rows and longer or shorter than rostrum.
The skull of the female averages 34 per cent lighter than that of the male.
Comparisons of the skull with those of M. f. primulina, M. f. spadix, M. f. oribasus, M. f. alleni, M. f. nevadensis, and M. f. neomexicana are made in accounts of those subspecies.
Remarks.--Richardson's (1829:47) account on which Bonaparte may be said to have based his name, records measurements in inches and lines which I transpose into millimeters as follows: Total length, 440 mm.; length of head and body, 305; length of tail (vertebrae), 135; length of tail (including fur), 164 mm. Specimen no. 43.3.3.3 in the British Museum, which has by some persons been regarded as the type, yields measurements as follows: Total length, 408 (which allows for 15 mm. loss of the fleshy part of the end of the tail); length of head and body, 272; length of tail (vertebrae), 136 (= 121 + 15); length of tail (including fur), 162 (142 + 20 mm. that appears to have been lost). Richardson's specimen would appear to have been of unusual proportions and to have been larger than no. 43.3.3.3. Some reasons for and reasons against regarding this specimen as the holotype are given in the account of M. erminea cicognanii.
The name longicauda was applied to practically all long-tailed weasels of the western United States at one time but as one after another of the geographic variants in the mountainous regions were designated as separable, the name longicauda came to be restricted to the light-colored, relatively large, animal of the Great Plains.
The intergradation of longicauda with spadix and oribasus has been commented on in the discussions of those subspecies. The larger size and darker color of specimens referred to longicauda from Devils Lake and Grafton, North Dakota, are features indicative of intergradation there with spadix. Two young females from Waterton Lake Park, Alberta, by their darker than average color, suggest intergradation with oribasus, as, for that matter, does the specimen from Waterton Lake [= Chief Mountain Lake, in Montana] itself, which, however, is even darker than the two specimens taken on the Canadian side of the line and hence is referred to oribasus. An adult female, no. 175586, U. S. Nat. Mus., from Moose Pass, Alberta, examined after the above was written, is larger than any other female seen of longicauda and in this respect may show approach to oribasus, which in the northern part of its range is of large size as judged by males from the Bowron Lake region.
One male, no. 8564, Nat. Mus. Canada, from Max Lake, Turtle Mountain, Manitoba, presents puzzling characters. The external measurements of 465, 170, and 57, are in keeping with the great length of the skull which has a basilar length of 48.8. The tooth-rows are 19.3 in length and the mastoid breadth, 25.4. The relative narrowness indicated by the mastoid breadth is maintained throughout the skull. The only other specimens relating to the Turtle Mountains that have been seen are two male, skins without measurements or corresponding skulls, nos. 38902 and 38903, Amer. Mus. Nat. Hist., labeled as from either "Stump Lake or Turtle Mts.," North Dakota. One of these, no. 38902, is much darker than the other. Possibly it is from the Turtle Mountains and the other, lighter-colored one, is from Stump Lake. Study of additional specimens from the Turtle Mountains might show the existence there of a distinct race.
Four specimens, in the collection of Myron Swenk, from Inland, Clay County, Nebraska, are instructive as showing how intergradation occurs between primulina and longicauda. A subadult male, no. 10, is intermediate in external measurements and in color but in each instance is nearer primulina. The same is true of the least width of the color of the underparts. The color of the underparts extends uninterruptedly over the hind legs to the toes as in longicauda, but is absent from the underside of the tail as in primulina. In the skull, the basilar length, breadth of bulla, and size of teeth are nearer longicauda, as are also the ratios to the basilar length of the length of tooth-rows, breadth of the rostrum, length of the tympanic bulla, and depth of the braincase at the anterior margin of the basioccipital. Ratios to the basilar length of the interorbital breadth, mastoid breadth, zygomatic breadth, and depth of the skull at the posterior borders of the upper molars are nearer to those of primulina. The relatively long rostrum, as represented by the orbitonasal length, is nearest to that of spadix. A young, almost subadult, female, no. 7, agrees with primulina in color, color pattern, and length of hind foot. The other external measurements are intermediate, but nearer those of primulina. Size of skull and teeth are as in longicauda. Relative proportions of parts of the skull are not diagnostic in specimens as young as this female. An adult female, skull only, no. 8, agrees with, or approaches nearer to, longicauda in size of skull and teeth and in relative proportion of every part studied. A juvenile, skull only, of questionable sex, no. 9, provides no diagnostic characters. On the basis of color, these specimens from Inland are distinctly nearer primulina. On the basis of cranial characters they are distinctly nearer longicauda. External measurements are intermediate and are a little nearer those of primulina. By placing the most weight on the cranial characters, the animals may be referred to longicauda. The same may be said of 2 skins, one skin with a skull, from Hastings, Nebraska. In each skin the color-pattern is as in primulina; in one the under side of the tail is nevertheless lighter-colored more as in longicauda and the skull, adult male 121651 American Museum of Natural History, approaches nearer to primulina in narrowness but has the large teeth of longicauda.
Intergradation with neomexicana is suggested by one specimen, no. 7936, Univ. Kans., from Thomas County, Kansas, which has well-developed white facial markings.
The specimen, no. 180, Kansas Agric. College, from Glasco, is mounted, of large size, in white winter pelage, and lacks external measurements. On the basis of its obvious large size, and a hind foot measurement of 49 millimeters obtained from the mounted skin, the animal is provisionally referred to longicauda rather than to primulina.
Putorius culbertsoni is a name now credited to Coues (1877:136). Although Coues probably intended only to indicate that Baird wrote this name on the labels of two specimens in the mammal collection of the Smithsonian Institution, Coues gave an "indication" of the application of the name by publishing at the same time the catalogue numbers of specimens whose labels bore the name and thus, in accordance with article 21 of the International Rules of Zoölogical Nomenclature, himself becomes the author of the name. Of the two specimens mentioned by Coues, only the first recorded by him, no. 4320 (with skull no. 37995, U. S. Nat. Mus.), can now be found.
Fortunately, the skull of this specimen labeled (see Lyon and Osgood, 1909:218) as taken at Fort Laramie, Wyoming, is well preserved. Its only defects are a fracture in the left zygomatic arch and the absence of parts of each of the first lower molars. In deciding on the subspecific application of the name Putorius culbertsoni Coues, the skull of the type must be principally relied upon, for there is available only one other specimen, a skin only (no. 12596, U. S. Nat. Mus.), from the same place, and it, like the type, is in white winter pelage and lacks flesh measurements.
The ranges as now known of three subspecies of Mustela frenata approach near to Fort Laramie. These are M. f. longicauda, M. f. alleni, and M. f. nevadensis. The skull of the type of culbertsoni is not typical of any one of the three mentioned races. The small size of its teeth and relative (to basilar length) shallowness of the frontal region of the skull through the postorbital processes of the frontal are as in nevadensis. The zygomatic arches are not so greatly expanded as in some specimens of longicauda and are more like the average for nevadensis or alleni, as also is the relatively (to basilar length) long orbitonasal length. However, each of these characters is subject to variation and alone is not surely diagnostic, especially toward the margin of the range of any one of the subspecies concerned. The same may be said of the relatively great breadth of the skull interorbitally--a feature typically found in longicauda. More important, in my estimation, is the large size of the skull; all parts measured (excepting the teeth, the depth at the posterior border of the last upper molars, the zygomatic breadth, and the depth of the tympanic bullae) equal or approach nearest to the average for males of longicauda of similar age.
The small size of alleni prevents its identification with culbertsoni. The question of application lies between nevadensis and longicauda. If the long-tailed weasel at Fort Laramie is found to be referable to the race earlier named longicauda, no change in current nomenclature will be effected. If, on the other hand, the long-tailed weasel from Fort Laramie is found to be referable to nevadensis this name will have to fall before the earlier proposed name culbertsoni. There is, however, a third possibility, namely, that the long-tailed weasel of the Transition and Upper Sonoran zones of southern Wyoming and northern Colorado, as for example, at Lay, Colorado, may represent a recognizable race characterized by size about as in longicauda, relative proportions of skull about as in nevadensis and coloration intermediate, to which the name culbertsoni may apply. For more detailed discussion of this possibility, see remarks under M. f. nevadensis.
Satisfactory application of the name Putorius culbertsoni Coues requires an adequate series of adult specimens, of both sexes in the summer coat with external measurements taken in the flesh, from the type locality and like material from elsewhere in southern Wyoming. On the evidence furnished by the skull of the type of culbertsoni, that name tentatively is placed in the synonomy of longicauda.
Only 2 of 25 adults examined for malformation of the frontal sinuses by parasites showed evidence of disease.
Specimens examined.--Total number, 138, arranged alphabetically by provinces and states and further by districts or counties from north to south except as otherwise indicated. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=Alberta.= St. Albert, 1; S. Edmonton, 3; Islay, 4[77]; Battle River, south of Camrose, 1[77]; Daysland, 1[77]; Moose Pass, 1; Blindman River, 2 (1[75], 1[2]); Red Deer, 3 (2[2], 1[60]); Bearberry Creek near Sundre, 1[77]; Canad. Nat. Park, N.W. Territory, 1[60]; Red Deer River, Didsbury, 1; Canmore, 1; Calgary, 11 (6[60], 2[1], 1[86]); Red Deer River, 3[2]; Little Sandhill Creek, Red Deer River, 1[77]; Waterton Lake Park, 2[77]; Sweetgrass Hills, 1[77]; Alberta, 1[14].
=Colorado.= Yuma County: Wray 4 (1[88], 3[74]).
=Kansas.= Rawlins County: 7 mi. N, 3 mi. W Beardsley, 1[74]; 6 mi. S and 2 mi. E Atwood, 1[74]; 15 mi. SE Atwood, 1[74]. Thomas County: near Brewster, 2[93]; no locality more definite than county, 2[93]. Trego County, 2 (1[2]). Cloud County: Glasco, 1[67].
=Manitoba.= Portage la Prairie, 3[75]; Carberry, 2 (1[2], 1[1]); Carman, 1[60]; Max Lake, Turtle Mt., 1[77].
=Montana.= Glacier County: St. Marys Lake, 1; Blackfoot, 1: Blackfoot Agency, 1. Blaine County: 6 mi. east Chinook, 1[74]. Pondera County: 1/2 mi. SE Conrad, 1[74]. Toole County: Shelby Junction, 1. Hill County: Havre, 1. Fergus County: Moccasin Mts., 5 mi. NW Hilger, 1; 7 mi. NE Hilger, 1. Rosebud County: 3/4 mi. N Ingomar, 1. County in question, Milk River, 2.
=Nebraska.= Dawes County: Chadron, 2[35]. Cherry County: Kennedy, 1; no locality more definite than county, 1. _Brown County_: Long Pine, 1[68]. Antelope County: Neligh, 1[35]. Adams County: Hastings, 2[2]. Clay County: Inland, 4[35].
=North Dakota= (arranged by counties from west to east). _Divide County_: Crosby, 1. Mountrail County: Lostwood, 1. Little Missouri River, 1. Golden Valley County: Sentinel Butte, 1. Billings County: Medora, 1[60]. McLean County: 3 mi. W Elbowoods, 1. Oliver County: Ft. Clark, 2. Morton County: Mandan, 1. Sioux County: 3 mi. N Cannonball, 1. Logan County: 6 mi. SW Napoleon, 1. Rolette County: Turtle Mts., 1[76]; Fish Lake, 1. Benson County: Ft. Totten, 3[14]; Sully Hill Nat. Park, 1. Ramsey County: Devils Lake, 2. Stump Lake or Turtle Mts., 2[2]. Nelson County: Stump Lake, 1. Grand County: Larimore, 1. Walsh County: Grafton, 11 (4[76], 3[74], 2[2]). _Stutsman County_: Jamestown, 1. Barnes County: Valley City, 1.
=Saskatchewan.= Wingard, 5; Osier, 2[75]; Simpson, 1[2]; Touchwood Hills, 4[7]; South arm Last Mountain Lake, 1[77]; Rush Lake (Assiniboia, N.W.T.), 2[75].
=South Dakota.= Pennington County: Rapid City, 1.
=Wyoming.= Goshen County: Fort Laramie, 2.
=Mustela frenata oribasus= (Bangs)
Long-tailed Weasel
Plates 16, 17, 18, 31, 32, 33 and 40
Putorius (Arctogale) longicauda oribasus Bangs, Proc. New England Zoöl. Club, 1:81, December 27, 1899.
Putorius longicauda, Coues, Fur-bearing animals, p. 136, 1877 (part).
Mustela longicauda oribasus, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912.
Mustela longicauda oribasa, Hall, Univ. California Publ. Zoöl., 40:368, November 5, 1934.
Mustela frenata oribasa, Hall, Carnegie Instit. Washington Publ. 437:105, November 20, 1936.
Type.--Female, adult, skull and skin; no. 9058, collection of E. A. and O. Bangs, but now in collection of Mus. Comp. Zoöl.; source of Kettle River, 7500 feet [the summit between middle fork of Kettle River and Cherry Creek at Pinnacles--oral information from the collector, Feb. 12, 1936], British Columbia; September 10, 1898; obtained by Allan Brooks; original no. 1368.
The skull (plate 40) is complete and unbroken. The teeth all are present and entire except right I^3 which has the anterior half broken away. The skin is complete, fairly well made, and in summer pelage.
Range.--Canadian and Hudsonian life-zones from near 56°N in the Rocky Mountains of British Columbia and Alberta and Ootsa Lake along the Fraser and Chilcotin rivers south to Alta Lake, in the Caribou and Monashee mountains, probably in the Selkirks and Rockies, and through the Rockies of Montana into extreme northern Wyoming. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. longicauda by near (14 n) Brussels Brown rather than near (h) Clay Color of upper parts and in males by relatively shallower occiput in which the depth of the skull, exclusive of the sagittal crest and taken at the anterior border of the basioccipital, amounts to less than 59 per cent of the mastoid breadth; from M. f. nevadensis by greater average size, see measurements.
Description.--Size.--Male: Two adults from Florence, Montana, measure as follows: Total length, 440, 440; length of tail, 165, 161; length of hind foot, 47, 49. Corresponding measurements of an adult male from Quesnel, British Columbia, are: 443; 168; 55. Tail amounts to 60, 58, and 61 per cent as long as head and body. Length of hind foot averages more than basal length.
Female: The type specimen, the only typical adult or subadult specimen of this sex of which external measurements are available, measures: Total length, 392, length of tail, 150, length of hind foot, 46. Tail is 63 per cent as long as head and body. Length of hind foot amounts to more than basal length.
The differences in external measurements, between the one female and the average of the three males are: Total length, 49; length of tail, 15; length of hind foot, 4.
Externals.--Longest facial vibrissae brown or white (often both colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to or beyond apical pad of fifth digit; hairiness of foot-soles (in summer pelage) slightly less than shown in figure 19.
Color.--Upper parts, in summer, near (14 n) Brussels Brown, more blackish and less reddish than tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304; in type near tone 4, pl. 301 of Oberthür and Dauthenay. Underparts, in summer, Buff Yellow or near (20 c) Amber Yellow. In winter, all white except tip of tail which is at all times black. Upper parts of uniform color except for occasional slight darkening of top of head and along mid-dorsal line of back. Color of underparts extends distally on posterior sides of forelegs over feet, on medial sides of hind limbs over antiplantar faces of toes and over proximal two-thirds of ventral side of tail. Least width of color of underparts amounting to 43 per cent of greatest width of color of upper parts, 75 per cent in male from 4 miles northeast of Quesnel, British Columbia, and 52 (33-66) in four males from Montana. Black tip of tail in four males from Montana averaging 50 (44-60) mm. long. Thus averaging approximately as long as hind foot and 33 per cent of length of tail-vertebrae.
Color not different than in many specimens of M. f. nevadensis. Color comparison with M. f. longicauda has been made in the account of that subspecies.
Skull and teeth.--Male (based on 5 adults and 2 subadults from British Columbia and 4 adults from Montana): See measurements and plates 16-18. As described in Mustela frenata longicauda except that: Weight, 5.0 (3.8-6.0) grams; basilar length, 46.7 (43.6-48.8); postorbital breadth in one of nine instances less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2-1/2 to 5 upper incisors; length of tympanic bulla not less than length of lower molar and premolar tooth-row and shorter than rostrum.
Female (based on the type, specimen): See measurements and plates 31-33, 40. As described in Mustela frenata longicauda except that: Weight, 3.5 grams; basilar length, 41.6 mm.; zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than outside length of P^{4}; tympanic bulla as far posterior to foramen ovale as width of 4-1/2 upper incisors; height of tympanic bulla less than distance from anterior margin of tympanic bulla to foramen ovale; length of tympanic bulla less than length of rostrum. If more than one skull were available of the female of oribasus it is believed that the description would agree with that of longicauda in nearly all features.
The skull of the female is 30 per cent lighter than that of the average male.
Comparison with longicauda reveals that, on the average, skulls of males are larger, relative to the basilar length broader across the mastoids, shallower through the braincase as measured at the anterior end of the basioccipital exclusive of the sagittal crest, with longer rostrum. Compared with nevadensis, the skull averages larger in all measurements taken, and has a relatively broader rostrum, relatively greater mastoid breadth and a braincase which is shallower relative to the basilar length. By weight, the skull of nevadensis is a fourth lighter, and in linear measurements 5 to 18 per cent smaller.
Remarks.--Some of the specimens from Montana, which here are referred to oribasus, more than half a century ago were listed by Coues (1877:138) under the name longicauda. It was not until 1899 that this race was given a name by Bangs, who at that time (1899B:81) accurately made out the distinctive color features. Distinctive cranial characters cannot be described with assurance even now because there still are too few specimens.
The type specimen, at one time examined by the present writer, has on the stuffed skin no well-developed mammae, scrotal pouch, or other visible sexual part. Probably the collector's sex mark for female is correct.
As judged by the two skulls of subadult males from the Barkerville region, individuals of this race attain larger size than do those of longicauda. On the basis of larger size than either longicauda or nevadensis, the specimens from the Rocky Mountains of Montana and two from northern Wyoming are referred to this race. The short, wide, flat, tympanic bullae, relatively great mastoidal breadth, and some other features of the specimen from Donovan, Montana, point toward oribasus, whereas nearly as many more cranial features, in this instance mainly differences in size, are indicative of nevadensis to which race the specimen might almost equally well be referred. Another male from Darby, in the Bitterroot Valley of Montana, has a slightly longer hind foot than those from Florence, but a female from Hamilton, agrees more nearly with nevadensis. The average of all the specimens from the Bitterroot Valley is a little nearer oribasus. Four skulls from Buffalo, Wyoming, here referred to nevadensis show approach to oribasus in size of skull. The specimens from Big Snowy Mountains, and the Highwood Mountains of Montana are too young clearly to show size of the adult skull, but are distinctly darker colored than longicauda of the plains country proper. Of two subadult females from Tacy, Montana, the color of the one in summer pelage is distinctly nearer that of oribasus and nevadensis than it is to that of longicauda to which some approach in color might be expected. The reduced size of both of the specimens is further suggestive of nevadensis and it may be that adult specimens from these more eastern mountainous areas in Montana will show that nevadensis is the name proper to apply to animals of this region.
Intergradation with nevadensis is suggested by specimens collected from along the upper reaches of Okanagan Lake, British Columbia, by Major Allan Brooks and Mr. J. A. Munro and by a series of skulls from Ione, Pend Orielle County, Washington, lent me by Mr. Walter Dalquest. At each place, the average of all specimens is nearest to that of nevadensis.
Specimens from near Waterton Lake show several steps in the transition from the light-colored longicauda type of coloration to the darker coloration characterizing oribasus. One taken here, at a time when the body of water referred to seems to have been known as Chief Mountain Lake, is barely dark enough to be placed with oribasus. Two other specimens from across the Canadian Border labeled as "Waterton Lake Park" are slightly lighter colored above, and on this account are placed with longicauda.
The two adult males from Lillooet, British Columbia, are referable to oribasus although neither is quite typical. One has a saturated coloration suggestive of that of altifrontalis and the skull is shorter and broader than in other specimens of oribasus. The female from Lillooet, skin alone, no. 916, Prov. Mus., B. C. is small for oribasus. The female, no. 1539, collection of Kenneth Racey, from Alta Lake, in brown winter pelage, in almost every measurement falls nearly midway between altifrontalis and oribasus but slightly nearer the latter. The skull from Chezacut and 3 animals from Wistaria, British Columbia, probably are females and show a greater average size than specimens from farther to the southeast. For example, the basilar length of the skull, 44.8 (44.3 to 45.1), exceeds that of the type specimen. The animals from Wistaria on Ootsa Lake furnish the northwesternmost station of occurrence of which I have record for this subspecies.
The northernmost records of occurrence, at "Clearwater River, Peace River, B. C," and at Little Prairie, are furnished by a white skin without skull, no. 257450, U. S. Nat. Mus., purchased on August 2, 1932, at the place mentioned by W. H. Sheldon and Richard Borden, and a skull with white winter skin, no. 3585, Provincial Museum, British Columbia, respectively. The characters distinguishing longicauda and oribasus are not shown by white winter skins; the skull shows some features of longicauda, and the reference of these specimens to oribasus rather than longicauda is tentative.
Only the skull from Little Prairie shows evidence of infestation of the frontal sinuses by parasites. In the Barkerville area of British Columbia, Mr. and Mrs. Thomas T. McCabe obtained only 2 skulls of this subspecies from a total of 238 weasel skulls gathered by local trappers. The others were Mustela erminea.
Specimens examined.--Total number, 46, listed by localities from north to south and unless otherwise indicated, in the United States National Museum.
=British Columbia.= West of Hudson Hope, 1[7]; Clearwater River, tributary to Peace River, 1; Little Prairie, a few miles south of Peace River and about 40 miles west of the main highway between Dawson Creek and Fort St. John, 1[85]; Wistaria, 3[85]; Four Mile Creek, 4 mi. NE Quesnel, 1[21]; Isaacs Lake, 3200 ft., 1[74]; Barkerville region, 1[74]; Clear River, 4800 ft., 1[74]; Chezacut, 1[31]; Lillooet 3 (2[77], 1[85]); Alta Lake, 1[31]; source of Kettle River, 7500 ft., 1[75]; E side Beaverfoot Range, 4000 to 4500 ft. between Fraser Creek and 6 mi. SE of Fraser Creek, 1[74]; Cranbrook, 1[86]; head of Cross River, 10 mi. below Assiniboine Pass, 1[7]; camp east of "Kootanie," 1[7]; camp east of Kootanie River, 1[7].
=Alberta.= Thoral Creek, 7000 ft., 50 mi. NE Jasper, 1[2].
=Montana.= Glacier? County: Chief Mt. Lake (= Waterton Lake), 1. Flathead County: Columbia Falls, 1. Chouteau? County: Highwood Mts., 1. Fergus? County: Big Snowy Mts., 1. Wheatland County: Harlowton, 1[74]. Ravalli County: Florence, 2; Hamilton, 1[56]; Darby, 1[56]; Carlos [= Charlos] Heights, 2[74]; Tin Cup District, 2[74]; no locality more definite than county, 2[74]. _Beaverhead County_: Donovan, 1. Madison County: Sheridan, 1[74]. _Gallatin County_: Ranch 7-11, Eldridge, 1[60]. Stillwater County: Tacy, 2[76]. County in question: Gallatin Valley, 1; Yellowstone Park, 1[75].
=Wyoming.= Glen Creek, Mammoth Hot Springs, 1. Park County: Four Bears, 1[2].
=Mustela frenata alleni= (Merriam)
Long-tailed Weasel
Plates 18, 19, 20, 31, 32 and 33
Putorius alleni Merriam, N. Amer. Fauna, 11:24, June 30, 1896.
Mustela alleni, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.
Mustela frenata alleni, Hall, Carnegie Instit. Washington Publ. 473:106, November 20, 1936.
Type.--Male, adult, skull and skin; no. 186451, U. S. Nat. Mus. (formerly 4485/5120, collection of Dr. C. Hart Merriam); Custer, South Dakota; obtained by Vernon Bailey; original no. 90.
The skull is complete and unbroken. The upper incisors are missing. All the other teeth are present although the premolars, and especially the canines, are much worn, possibly as the result of the animal's efforts to free itself from a trap. The skin is fairly well made, in a good state of preservation, and entire.
Range.--Canadian, Transition and Upper Sonoran life-zones of the Black Hills of South Dakota and adjacent semi-bad-land territory of Wyoming and Nebraska southward to Mitchell, Scottsbluff County. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. longicauda in smaller size, adult males having a total length of less than 400, hind foot less than 45, basilar length less than 43.5, and in adult females total length less than 375, and basilar length less than 40; from M. f. nevadensis in near Clay Color rather than near (14 n to l) Brussels Brown of upper parts in summer.
Description.--Size.--Male: External measurements of the type specimen are: Total length, 372; length of tail, 137; length of hind foot, 44. Tail is 58 per cent as long as head and body. Length of hind foot more than basal length.
Female: No external measurements for typical adults are available. No. M1 #41 from Mitchell, Scottsbluff Co., Nebraska, an adult female which is an intergrade with the larger M. f. longicauda, measures as follows: Total length, 367; length of tail, 120; length of hind foot, 41.
Externals.--Longest facial vibrissae dark brown or white and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) as shown in figure 20.
Color.--Winter pelage unknown; probably white except, of course, tip of tail. Summer pelage as described in Mustela frenata longicauda except that: Least width of color of underparts averaging, in 3 males from Black Hills, 54 (38-62) per cent of greatest width of color of upper parts. Black tip of tail averaging 43 (40-45) mm. long. Thus, averaging approximately same length as hind foot and in type specimen amounting to 33 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on the type and no. 7440 Amer. Mus. Nat. Hist., from Hill City, S. Dak.): See measurements and plates 18-20. As described in Mustela frenata longicauda except that: Weight, 3.1 (3.0-3.2) grams; basilar length, 41.0 (40.9-41.0); mastoid breadth not less than postpalatal length; breadth of rostrum more than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 5 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale.
Female (based on no. 7441, American Mus. Nat. Hist., from Black Hills, S. Dak.): See measurements and plates 31-33. As described in Mustela frenata longicauda except that: Weight, 2.0 grams; basilar length 37.6. The skull of the female is 35 per cent lighter than the average for the two males.
Comparison with M. f. longicauda and M. f. nevadensis reveals that the tympanic bullae average more nearly flat and that the skull is smaller.
Remarks.--Animals of this subspecies were described and named by Merriam in 1896 as a distinct species on the basis of two or possibly three specimens from the Black Hills of South Dakota and the name seems never to have been applied to specimens from other regions. Vernon Bailey obtained only the one specimen, the type, on his trip in 1888, but two more were obtained for the American Museum of Natural History by Walter Granger in 1894.
Mustela frenata alleni combines the light coloration of M. f. longicauda with the small size of M. f. nevadensis. Indeed, the size may average less than that of nevadensis. M. f. alleni seems to reach its extreme of small size in the Black Hills of South Dakota. Specimens from Mitchell, Scottsbluff County, Nebraska, here referred to alleni are of larger size and in this respect are intermediate between the subspecies alleni and longicauda. Of the two specimens available from Chadron, Nebraska, and here referred to as longicauda, the female, M1 #6, is almost exactly intermediate in size between alleni and longicauda, whereas the male, Ml #11, is as large as the average-sized longicauda.
None of the nine skulls (5 adults) shows malformation resulting from the infestation of the frontal sinuses with parasites.
Specimens examined.--Total number, 10, as follows.
=Wyoming.= Crook County: Sundance, 1[91].
=South Dakota.= Pennington County: Hill City, 1[2]; 20 mi. N Elk Mt, 1[91]. County in question: Black Hills, 1[2]. _Custer County_: Custer, 2 (1[91], 1[2]).
=Nebraska.= Scottsbluff County: Mitchell, 4[35].
=Mustela frenata arizonensis= (Mearns)
Long-tailed Weasel
Plates 19, 20, 21, 31, 32 and 33
Putorius arizonensis Mearns, Bull. Amer. Mus. Nat. Hist., 3:234, June, 1891; Merriam, N. Amer. Fauna, 11:22, fig. 12, June 30, 1896.
Mustela arizonensis, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.
Mustela frenata arizonensis, Hall, Carnegie Instit. Washington Publ. 473:106, November 20, 1936.
Type.--Female, adult, skull and skin; no. 2490/1886, Amer. Mus. Nat. Hist.; San Francisco Forest [then (1886?), Yavapai County], Arizona; June 20, 1886; obtained by Edgar A. Mearns.
The skull (plates 31-33) is complete and unbroken save for a small puncture in the right squamosal. The incisors above and below and M^2 and P^2 on each side are missing. Four canines are preserved separately. Otherwise the teeth are in place. The skin has been taken down from a mount. Some hair has been lost from in front of the ears. Seven mammae are evident and show the animal to have been nursing young. The slightly faded color was mentioned by Mearns in the original description. He says (1891:234): "The memorandum of the colors was made before skinning, the specimen having been subsequently preserved in a solution of alum and salt, which extracted much of the coloring matter."
Range.--Transition to Hudsonian life-zones of Arizona and extreme western New Mexico, along the Colorado River, and south of the Little Colorado River, from San Francisco Mountain region along Mogollon Plateau to extreme western New Mexico. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. neomexicana by near (14 n) Brussels Brown rather than Buckthorn Brown color of upper parts, in absence rather than presence of white frontal spot continuous with color of underparts, in basilar length of less than 44 in males and 39.3 in females; from M. f. nevadensis in that total length averages less than 375 in males and 330 in females, basilar length averaging less than 41 in males and less than 36.7 in females.
Description.--Size.--Male: No. 24679/32071, from Springerville, and no. 248993 from the Kaibab Plateau, measure respectively, as follows: Total length, 363, 367; length of tail, 140, 143; length of hind foot, 41.5, 41.0. Tail is 63, and 64 per cent as long as head and body. These males, the only specimens of that sex of which external measurements are available, probably are grading toward nevadensis and therefore are nontypical.
Female: Three specimens, one young from Little Spring, a subadult from Deadmans Flat and the type specimen, measure respectively as follows: Total length, 323, 296, 302; length of tail, 110, 101, 109; length of hind foot, 38, 33, 36. These average, 307, 107, 36. Tail averages 53 per cent as long as head and body.
Differences in external measurements of the two sexes are: Total length, 56; length of tail, 39; hind foot, 5.5.
Externals.--Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot soles (in summer pelage) about as shown in figure 19.
Color.--Winter pelage unknown. Summer pelage with upper parts near (14 n) Brussels Brown or tone 2 of Raw Umber of Oberthür and Dauthenay, Pl. 301, darker on top of head from nose to line connecting posterior margins of ears. Tip of tail always black. Chin and upper lips white. Remainder of underparts Buff Yellow to Straw Yellow and rarely Ochraceous Buff. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind legs to ankles and over antiplantar faces of toes, medial third of tarsus, and over proximal fifth to fourth of ventral side of tail. Least width of color of underparts averaging, in 8 specimens, 44 (29-54) per cent of greatest width of color of upper parts. Black tip of tail, in four females averaging 35 (33-38) mm. long. Thus, averaging shorter than hind foot and 32 per cent of length of tail-vertebrae. Three of the eight specimens before me (no. 242671 from 25 mi. SE Flagstaff, not available at time of this accounting) have the dark spot near the angle of the mouth faintly indicated, whereas the other five lack the spots. The color is as in M. f. nevadensis.
Skull and teeth.--Male (based on 55211, 65231, and 248993; see p. 422): See measurements and plates 19-21; weight 2.7 and 3.1 grams; basilar length, 40.4; zygomatic breadth more than distance between condylar foramen and Ml or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more than postpalatal length; postorbital breadth less than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than medial length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3-1/2 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar-premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below talonid of m1.
Female (based on the type specimen): See measurements and plates 31-33; weight, 1.6 grams; basilar length, 35.5; zygomatic breadth less than distance between condylar foramen and M1 and more than distance between anterior palatine foramen and anterior margin of tympanic bulla (nearly equal in each instance); postorbital breadth less than length of upper premolars and greater (7.1-8.4) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate equal to inside length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar-premolar tooth-row and greater than length of rostrum.
The skull of the female averages 41 per cent lighter than that of the male.
Compared with the skull of M. f. nevadensis, that of arizonensis is smaller, less heavily ridged and has more inflated tympanic bullae and a relatively greater mastoid breadth. Comparison with the skull of M. f. neomexicana is made in the account of that subspecies.
Remarks.--In 1891 Mearns (234-235) named this weasel as a full species on the basis of two individuals taken by him in 1886 and 1887. Since that time only a few additional specimens have been preserved. Only four are adults. Although this material does not permit of a definition of the subspecies as precise as could be wished, still, it clearly shows that the animals from the plateau region of Arizona are recognizably different from those farther north in the Sierra Nevada of California and those of the Rocky Mountains and Great Basin region northward to the Canadian border. These more northern animals have gone by Mearns' name, arizonensis, since the date of its proposal until 1939 when the name nevadensis was proposed.
The smaller size, especially of the skull, and the greater inflation of the tympanic bullae are the outstanding characters which distinguish arizonensis from the similarly marked nevadensis. The bullae are relatively much inflated throughout but especially so on the posteromedial parts.
Although the three adult males and two subadult females available of this subspecies are smaller in most parts measured than any of the scores of nevadensis of similar age that have been measured, overlap in size probably will be found as additional specimens of arizonensis become available. A young female, no. 18513, coll. D. R. Dickey, from Little Spring, does have certain cranial measurements as large as are found in the minimum-sized nevadensis from farther north.
Intergradation with the two subspecies whose geographic ranges adjoin that of arizonensis is indicated by specimens at hand. One of these is the adult male from 25 miles southeast of Flagstaff, which shows decided approach to neomexicana, in color and in possessing white facial markings less well developed than in neomexicana. Even better developed white facial markings, with intervening blackish coloration, are displayed by no. 148271, U. S. Nat. Mus., from 8500 feet altitude on Willow Creek, New Mexico. This subadult female shows approach to neomexicana also in larger size of the skull and entire animal. The great inflation of the posterior part of each of its bullae and the dark color of the upper parts are characters of arizonensis. The color of the underparts stops at the ankles leaving the hind feet dark colored, in which respect the specimen is unlike either neomexicana or arizonensis. If additional specimens showing the same characters as this one be found at other nearby localities they probably should be given recognition as a separate subspecies. For the present it seems best to regard the specimen merely as an intergrade. Although it might, with almost equal propriety, be referred to either neomexicana or arizonensis, the specimen is here placed with the latter. The subadult male from Springerville, Arizona, is of larger size than the topotypical male of arizonensis and in this respect shows slight approach to nevadensis. The narrower mastoidal breadth and slightly less inflated tympanic bullae of the male from the Kaibab Plateau may reflect merely individual variation or may represent intergradation in these features with nevadensis.
The statement made by Merriam (1896:22) that, "The type specimen . . . is an immature female and is of unusually small size. A male obtained by him [Mearns] near the same place is of the normal size, as is another male in the Department collection from Springerville, Ariz., collected by E. W. Nelson," needs correction. The female is not immature. The specimen obtained by Mearns near the same place probably refers to Amer. Mus. No. 2489, from Quaking Asp Settlement, which lacks both the skull and external measurements. As stuffed it is of small size for a male. The male from Springerville, as shown by the external and cranial measurements, is not of normal (i. e. average) size, but is smaller than the average for the other populations of similarly colored weasels referred to by Merriam (op. cit.) as arizonensis but here described under the name nevadensis.
None of the skulls shows signs of infestation of the frontal sinuses by parasites.
Specimens examined.--Total number, 17, arranged alphabetically by states and from north to south by counties in each state. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=Arizona.= Coconino County: VT Park, Kaibab Plateau, 1; Deadman Flat, 6400 ft., 1[74]; Little Spring, 1[59]; Government Prairie, near Parks, 1[74]; Coconino? County: San Francisco Forest (Yavapai Co., in 1886), 1[2]; 25 mi. SE Flagstaff, 1; Quaking Asp Settlement, 1[2]. Apache County: Springerville, 1; North Fork White River, White Mts., 8200 ft., 4[87]; head San Francisco River, Judd Ranch, Alpine, 1[74]; 2 mi. SE Big Lake Knoll, 8700 ft., 24 mi. S Springerville, 1[74]. Greenlee County: S end Blue Range, 9000 ft., Prieto Plateau, 1; Beaver Creek, 7000 ft., 1[74].
=New Mexico.= Grant County: Mogollon Mts., Willow Creek, 8500 ft., 1.
=Mustela frenata nevadensis= Hall
Long-tailed Weasel
Plates 19, 20, 21, 33, 34, 35 and 39
Mustela frenata nevadensis Hall, Carnegie Instit. Washington Publ. 473:91, November 20, 1936.
Putorius longicauda, Coues, Fur-bearing animals, p. 136, 1877 (part); Merriam, N. Amer. Fauna, 5:83, July 30, 1891.
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Putorius arizonensis, Merriam, N. Amer. Fauna, 11:22, figs. 13, 14, June 30, 1896 (part); Stephens, Mammals of California, p. 247, 1906.
Mustela arizonensis, Grinnell and Swarth, Univ. California Publ. Zoöl., 10:376, October 31, 1913; Whitlow and Hall, Univ. California Publ. Zoöl., 40:247, September 30, 1933.
Mustela arizonensis arizonensis, Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933.
Mustela frenata, Boyer, Journ. Mamm., 24:99, February 20, 1943.
Type.--Female, adult, skull and skin; no. 41053, Mus. Vert. Zoöl.; three miles east Baker, White Pine County, Nevada; May 30, 1929; obtained by E. R. Hall and W. C. Russell; original no. 2674, E. R. H.
The skull (plates 33-35) is complete and unbroken. The teeth all are present and entire. The skin is fairly well made. Eight mammae are evident and show the animal to have been nursing young.
Range.--Altitudinally, 700 feet at Wenatchee, Washington, to the highest parts of the mountains of the western United States; Upper Sonoran Life-zone to Arctic Alpine Life-zone; southern British Columbia in the Cascades and territory west to Monashee Mountains, and Nelson, southward in the Cascades of northern Washington, over western Washington, Idaho, Utah, and Nevada to northeastern Arizona and northern New Mexico; westward from the eastern base of the Rocky Mountains in Colorado to the western base of the Sierra Nevada and Cascades of California and to the Cascades of southern Oregon. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. oribasus by smaller average size, see measurements; from M. f. longicauda by near (14 n to l) Brussels Brown rather than near (h) Clay Color of the upper parts, and in males by a shallower occiput in which the depth of the skull, exclusive of the sagittal crest, and taken at the anterior border of the basioccipital, amounts to less than 59 per cent of the mastoid breadth; from M. f. alleni by near (14 n to l) Brussels Brown rather than near (h) Clay Color of upper parts in summer; from M. f. neomexicana by near (14 n to l) Brussels Brown rather than Buckthorn Brown color of upper parts, in absence of white frontal spot continuous with color of underparts, in basilar length of less than 46 in males and 40 in females; from M. f. arizonensis by total length averaging more than 375 in males and 330 in females, basilar length averaging more than 41 in males and 36.7 in females; from M. f. inyoensis by absence of white facial markings; from M. f. pulchra by absence of light facial markings, near (14 n to l) Brussels Brown rather than near (16 j) Buckthorn Brown color of upper parts, and lesser size, hind foot less than 40 in females and basilar length averaging less than 46.0 in males; from M. f. xanthogenys by absence of light facial markings and near (14 n to l) Brussels Brown rather than Buckthorn Brown color of upper parts; from M. f. munda by absence of white facial markings, presence of color of underparts on ventral face of proximal third of tail, and hind foot of less than 50 in males; from M. f. saturata by presence of light color of underparts on tail and ankle and in lesser average breadth across mastoid processes of skull (see measurements); from M. f. oregonensis by absence of nasofrontal white patch, presence of light color of underparts on ventral face of tail, and shorter skull, which, relative to its length in males, is deeper through the braincase; from M. f. washingtoni by presence of light color of underparts on ventral face of tail, by skull which in male relative to basilar length is shorter in the preorbital region and wider across the zygomata and mastoid processes, and in female has longer preorbital region and larger bullae (see measurements); from M. f. altifrontalis by lighter colored upper parts which are tones 1 to 3 of Raw Umber, pl. 301, rather than tone 4 of Brownish Drab, pl. 302, of Oberthür and Dauthenay, by Buff-Yellow to Straw Yellow rather than near (14´ a to 16´ c) Ochraceous-Buff color of underparts, by least width of color of underparts amounting to more than 37 per cent of greatest width of color of upper parts, by presence of color of underparts on ventral side of tail and on hind leg over ankle, and by lesser depth of skull through frontal region; from M. f. effera by larger size, males averaging 12-1/2 per cent larger in external measurements, 8 per cent larger in linear measurements of skull, and 22 per cent heavier in weight of skull, total length averaging 400 rather than 360, basilar length averaging 43.6 rather than 40.5.
Description.--Size.--Male: Twenty-one adults from the southern half of the Sierra Nevada of California yield average and extreme measurements as follows: Total length, 400 (356-428); length of tail, 150 (125-178); length of hind foot, 46.1 (42-50). Tail averages 60 per cent as long as head and body. Length of hind foot averaging more than basal length. Corresponding measurements of twelve adults from extreme southern and southwestern Colorado are as follows: 407 (355-431); 150 (133-170); 46.0 (42-49).
Female: Ten adults from the Sierra Nevada of California yield average and extreme measurements as follows: Total length, 349 (336-362); length of tail, 127 (120-133); length of hind foot, 36.3 (32-39). Tail averages 57 per cent as long as head and body. Length of hind foot less than basal length. Corresponding measurements of ten adults from the Rocky Mountains of central Colorado are as follows: 347 (325-375); 123 (111-141); 40 (32-43).
The average differences in external measurements of the two sexes, in the Sierras of California are: Total length, 51; length of tail, 23; length of hind foot, 9.8. Weight of 7 adult males from California is 267 (226-345) grams. Two adult females from there weigh 148 and 115 grams and 3 from White Pine County, Nevada, 134, 122 and 124, giving an average of 129 grams.
Externals.--Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) about as shown in figure 19.
Color.--Upper parts, in summer, near (14 n to l) Brussels Brown or tones 1 to 3 of Raw Umber of Oberthür and Dauthenay, pl. 301, darker on top of head from nose to line connecting posterior margins of ears. Chin and upper lips white. Remainder of underparts Buff-Yellow to Straw Yellow and sometimes Ochraceous-Buff especially in young, and in some adults from southern Colorado. In winter, all white, except tip of tail, or upper parts near (j) Snuff Brown or lighter than Brussels Brown with a smoked effect, and underparts white. Tip of tail at all times black. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind legs to ankles, over antiplantar faces of toes, medial third of tarsus and usually over proximal tenth to three-fourths of ventral side of tail. Least width of color of underparts averaging, in a series of twenty males from the southern half of the Sierra Nevada of California, 59 (37-76) per cent of greatest width of color of upper parts. In seven males from southern Colorado corresponding percentages are 55 (37-71). Black tip of tail in series from Sierra Nevada averaging 50 (40-60) mm. long; thus longer than hind foot and averaging 33-1/3 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 25 adults, from Sierra Nevada of California): See measurements and plates 19-21; weight, 3.7 (2.9-4.9) grams; basilar length, 43.6 (40.6-46.1); zygomatic breadth more than distance between condylar foramen and M1 (save in four instances) and more than distance between anterior palatine foramen and anterior margin of tympanic bulla (save in two specimens); mastoid breadth more (80 per cent of specimens) or less (20 per cent) than postpalatal length; postorbital breadth less than length of upper premolars and more or less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than medial length of P4 (except in two specimens); anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa not carried farther forward than point directly below hypoconid of m1.
Female (based on ten adults from Sierra Nevada of California): See measurements and plates 33-35; weight, 2.2 (1.8-2.4) grams; basilar length, 38.2 (36.7-39.5); zygomatic breadth more (except in one specimen) than distance between condylar foramen and M1 and more (save in two specimens) than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and less than (except in one specimen) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more or less than either outside or inside length of P4 but generally less than inside length; tympanic bulla as far posterior to foramen ovale as width of 3 to 5-1/2 upper incisors; height of tympanic bulla more or less (usually more) than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than length of rostrum.
The skull of the female averages 41 per cent lighter than that of the average male.
Compared with the skull of M. f. longicauda, that of both sexes averages smaller in every measurement taken. Males of nevadensis, on the average, relative to the basilar length, are narrower in the interorbital region and across the zygomata but have the orbitonasal length greater. Stated in another way, the rostrum of longicauda appears to be shorter and broader and the zygomata are more expanded. Females of nevadensis, on the average, relative to the basilar length are narrower across the mastoid processes and zygomata and have the braincase deeper at the anterior margin of the basioccipital. Also in nevadensis the mastoid processes do not project so far laterally beyond the braincase, the lambdoidal crest and postorbital processes are less well developed and except in the interparietal region, the temporal ridges hardly meet and they form a sagittal furrow rather than a low sagittal crest which characterizes adult females of longicauda. Each of these differences separating the females of longicauda from those of nevadensis are of the same nature, although not necessarily of the same degree, as those which appear in longicauda with increasing age. The differences mentioned above are readily appreciable when series of specimens are compared. However, none of the differences is of great degree, and most parts of the skulls of the two subspecies are of similar relative proportions. Even so, there is but little overlap in actual size. Comparisons with the skulls of M. f. oribasus, alleni, neomexicana, arizonensis, inyoensis, pulchra, xanthogenys, munda, saturata, oregonensis, washingtoni, altifrontalis, and effera are made in the accounts of those subspecies.
Remarks.--The populations to which the name nevadensis at present is assigned have gone by the name arizonensis since Mearns proposed this name in 1891. Before that time Coues (1877:141) had included individuals of this race under the name Putorius longicauda.
Among the populations here assigned to M. f. nevadensis, there is some geographic variation but it is of lesser degree than in most other species of mammals which range over the same region. Comparison of 20 adult males from the Rocky Mountains of Colorado with 25 adult males from a place as far distant as the Sierra Nevada of California shows that the two populations closely resemble each other. The specimens from Colorado average a trifle wider across the zygomata, have a longer body and therefore relatively shorter tail, and, except in southern Colorado, a slightly longer hind foot. Comparison of ten adult females from each of the two areas reveals that those from Colorado have a markedly longer hind foot, and a tail somewhat shorter relative to the length of the body. The mentioned differences are the only ones found among the great number of points investigated, except that as remarked by Merriam (1896:23) the Sierran animal has the yellow of the underparts reaching farther up under the chin, the underside of the tail on the average is more suffused with yellowish and the white on the upper lip is more extensive. As regards the last mentioned feature, my check of 34 skins from Colorado reveals that the white extends all the way around the upper lip in every specimen but one, whereas in 69 specimens from the Sierra Nevada the white extends all the way around the upper lip in only 39. However, as further remarked by Merriam (loc. cit.), not only this but the other color features are inconstant in addition to being slight. When the occurrence of the dark spots near the angles of the mouth are tabulated, it is found that in 33 Colorado-taken specimens they are absent in 19, faintly indicated in 13, and well developed in 1. In 62 California-taken specimens they are absent in 37, faintly indicated in 20, and well developed in 5.
In northwestern Colorado, southern Wyoming, and possibly through the Bear River Divide into southeastern Idaho, long-tailed weasels here referred to nevadensis approach longicauda in large size and occasionally in other features, more closely than do specimens of nevadensis from most other places in its range. This tendency is thought to be significant for much of the area in question lies in or below the Transition Life-zone, the same life zones in which farther to the eastward true longicauda occurs.
One specimen that illustrates this approach to longicauda is an adult male, no. 2334, collection of E. R. Warren, from 6160 feet, Lay, Routt [now Moffat] County, Colorado. In large size and, relative to the basilar length, shorter rostrum and shorter tympanic bullae, it agrees with longicauda but the darker color and, relative to the basilar length, narrowness of the rostrum, interorbital region, zygomatic expanse and the shallowness through the region of the postorbital processes place it with nevadensis. Of two other specimens from Steamboat Springs, Routt County, a young male, no. 4010, in the collection of E. R. Warren, has a hind foot (50 mm.) as long as in longicauda; and the other, no. 138195, U. S. Nat. Mus., an adult male, agrees well enough in size and proportions with nevadensis but has the coloration typical of longicauda.
From Wyoming, one subadult female, no. 177553, U. S. Nat. Mus., from Garrett, is intermediate in size and coloration but is nearer to nevadensis in these particulars, as it is in all other points considered except size of the molar teeth which are as large as in longicauda and larger than in any female nevadensis from Colorado or California. Another female, an adult, no. 179304, U. S. Nat. Mus., from Lonetree, Wyoming, agrees with longicauda in size of skull. Indeed, ten of seventeen cranial measurements exceed the maximum for Colorado-taken nevadensis. Where differences exist in relative proportions of the skull as expressed in percentages of the basilar length, the specimen approaches nevadensis in 5 instances and longicauda in only 3. The color is intermediate but much nearer that of nevadensis with which the animal agrees also in external measurements. Ten subadults (5 of each sex) from within 12 miles of Laramie (not Fort Laramie) show greater resemblance to nevadensis but definitely approach longicauda. Average external measurements are: [M], 408, 155, 44; [F], 361, 134, 40. The two other specimens examined from this general locality, a young female, no. 2711, Mus. Vert. Zoöl., from Fort Bridger, and a subadult female, no. 188377, U. S. Nat. Mus., from Bridger Pass, show no departures from nevadensis of similar age.
The specimens from scattered localities in the Transition Life-zone of northwestern Colorado and southern Wyoming are larger than nevadensis is elsewhere, and also in certain other features resemble longicauda of the plains to the eastward. Everything considered, the animals in question are much more like nevadensis than longicauda. Study of more specimens, especially from Wyoming, might provide grounds for recognizing as a different subspecies the animals in this large area comprising parts of Colorado and Wyoming from which so few specimens now are available. Possibly the name Putorius culbertsoni Coues would apply. Decision on that point will require adequate material from the type locality, Fort Laramie. See discussion of this name under M. f. longicauda.
In southeastern Idaho males are larger than they are at most other places within the range of nevadensis. An average of 7 adults and subadults from Pegram, Montpelier, Springfield, and the vicinity of Pocatello, reveals, when compared with the average of nevadensis from Colorado and that of longicauda from the Great Plains, that this population from southeastern Idaho is nearest to longicauda in linear measurements of the orbitonasal length, mastoid breadth, length of tympanic bullae, and as expressed in percentage of the basilar length, length of tooth-row, breadth of rostrum, and zygomatic breadth. In all other points of size, relative proportions and color, the animals approach nearer to, or actually agree with, nevadensis.
The specimens commented upon clearly show intergradation between nevadensis and longicauda. Similarly, the specimens from Scottsbluff County, Nebraska, here referred to M. f. alleni, by their larger size suggest intergradation of that subspecies with the larger nevadensis-longicauda stock although the approach is more toward longicauda than nevadensis. Between oribasus and nevadensis, however, there is no lack of material showing intergradation. As set forth in the account of oribasus, specimens from Montana are truly intermediate structurally as well as geographically.
Intergradation with washingtoni is shown by specimens from the northern part of the Cascade Range in Chelan and Okanogan counties, Washington. The adult male, U. S. Nat. Mus., no. 235183, from Bald Mountain, is referable to washingtoni on the basis of cranial characters but all the other adult and subadult specimens examined from Chelan and Okanogan counties are nearer nevadensis on the basis of cranial characters. Indeed, some show no approach to washingtoni in cranial characters. As might be expected on geographic grounds, the specimen from Easton, U. S. Nat. Mus., male subadult, no. 116870, shows approach to washingtoni. This is true of the coloration of the hind limbs, small size of the tympanic bullae, and relatively greater length of the preorbital part of the skull. However, the greater width of the light color of the underparts and relatively great breadth across the mastoid processes and zygomatic arches are points of agreement with nevadensis. Similarly, a series of 7 specimens from the Entait River, 20 miles above its mouth, in tone of color is nearer to washingtoni, as is one of the two skulls of adult males in length of the preorbital region. However, in greater breadth of the skull otherwise, and in the relatively great width of the light color of the underparts, the animals are nearer to nevadensis, to which they are here referred. Some of these characters mentioned above in which departure is shown from typical nevadensis are characters that show approach to altifrontalis. This is especially true of the more intense coloration and restriction of the color of the underparts.
Complete intergradation with effera is shown by specimens from southern Oregon. The change from small effera to the larger nevadensis here is gradual; consequently in northeastern California and southern Oregon the size increases gradually to the northward. Specimens showing complete intergradation with oregonensis and saturata are wanting. However, one specimen from Crescent Lake suggests oregonensis in having near (18) apricot yellow underparts such as occur frequently in oregonensis. Also some specimens from northern California approach saturata in having the color of the underparts reduced in the extent to which it reaches out on the under side of the tail. This fact and the consideration that the two races are less different from one another than are other kinds which definitely are known to intergrade leave no doubt but that material from the intervening localities would show complete intergradation.
Intergradation between nevadensis and munda is indicated by specimens from South Yolla Bolly Mountain, Trinity County, which are commented on at greater length in the account of M. f. munda. M. f. inyoensis is so closely related to nevadensis as to leave no doubt that specimens from suitable localities will show actual intergradation. That intergradation occurs directly with the bridled weasel of the interior valleys of California, M. f. xanthogenys, is shown by specimens from along the west-facing flank of the southern part of the Sierra Nevada. Probably intergradation occurs all along the Sierra Nevada on the western slope but specimens are lacking to show this. Weasels are known to occur in the foothill territory and the lesser attention given to this region by mammal collectors than to the higher parts of the mountains may explain the lack of preserved specimens. Individual specimens, here referred to nevadensis, but, showing varying degrees of approach to xanthogenys are as follows: A female from Hume; a male and a female from 8000 feet elevation, Monache Meadows; a male from 9800 feet elevation on the east fork of the Kaweah River; and 7 specimens, probably one family, from one-half mile south of Mineral King, 7850 feet. Of the specimens from 7850 feet, the adult male has no light facial markings and the head is only slightly darker than the back. The adult female has much restricted, light facial markings and the intervening areas are darker than in the male. The five juveniles trapped in the same burrow as the female, each has more extensive light facial markings than the adult female although the area of this varies from only slightly more than in the female to as much as in typical specimens of xanthogenys. Also, the dark color of the head in these five specimens averages darker than in nevadensis and more as in weasels to the southwestward especially latirostra. One of the five juveniles is lighter colored over all of the upper parts than nevadensis and is suggestive of xanthogenys in this respect. Finally, the adult male has on the underparts small spots of ochraceous orange suggestive of latirostra and some individuals of pulchra. No. 30655/42628, U. S. Nat. Mus., taken on Mount Whitney, also shows white facial markings and some other features of the valley-inhabiting xanthogenys. A suggestion of intergradation with arizonensis is furnished by specimens, referred to that race, from Springerville and the Kaibab Plateau. No specimens happen to be available from the region in which intergradation would be expected between nevadensis and neomexicana. Since neomexicana and arizonensis intergrade it is probable that nevadensis also will be found to intergrade with neomexicana. In summary, nevadensis is judged to intergrade with each of the subspecies of Mustela frenata whose range adjoins that of nevadensis.
This subspecies is remarkably free from injury to the frontal sinuses such as result from the presence of parasites. In 98 adults from Oregon, California, Nevada, and Colorado, no malformation was noted. Only 1 of the 26 specimens from Washington was malformed and it was an intergrade with washingtoni. The single adult from New Mexico was diseased, as were 3 of the 6 from British Columbia, 1 of the 20 from Idaho, and 1 of the 7 from Utah.
Specimens examined.--Total number, 568, arranged alphabetically by provinces and states and from north to south by counties in each state. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=Arizona.= Apache County: 15 mi. E Luka Chu Kai Navajo School, 8000 ft., 2.
=British Columbia.= Monte Cr., 20 mi. E Kamloops, 1[21]; Sicamous, 2; Okanagan, 18 (7[2], 6[85], 1[75], 1[86]); Monashee Pass, 1[31]; Swan Lake, near Okanagan Landing, 1[22]; Okanagan Landing, 11 (2[74], 3[31], 3[86], 3[22]); Vernon, 1[74]; Hope-Princeton Summit, 5600 ft., 1[77]; Hope, 1[20]; Similkameen, 1[77]; Osoyoos-Bridesville Summit, 1[77]; Anarchist Mt., Osoyoos, 1[31]; Myer's Creek, 1[77]; Rossland, Mt. Glory, 7000 ft, 1[77]; Cascade, 1[77]; Nelson, 1.
=California.= Siskiyou County: Hornbrook, 1; Tule Lake Refuge, 5[74]; Upper Mud Creek, 6700 ft., Mt. Shasta, 3; Mt. Shasta, 1. Modoc County: Goose Lake, 1[20]; Joseph Creek, 1[74]; 5280 ft., Parker Creek, near Alturas, 1[74]; Warner Mts., near Alturas, 1[8]; 5 mi. NW Eagle Peak, 7000 ft., 2[74]; Shields Creek, 5000 ft., 1[74]; Jess Valley, 1[8]. Shasta County: Cassel, 1. _Lassen County_: 3 mi. W Eagle Lake, 5800 ft., 1[74]; 4 mi. S Eagle Lake, 6000 ft., 2[74]; Mill Creek, 5000 ft., S base Mt. Lassen, 1; 6 mi. SW Calneva, 1. Tehama County: Dale's, 600 ft., on Paines Creek, 1[74]. Plumas County: Kelly's, 2 mi. S Willow Lake, 5200 ft., 3[74]; Quincy, 4[68]; Beckwith, Sierra Valley, 1. Butte County: Jonesville, 1[74]. Sierra County: Little Truckee River, 6500 ft., 3 mi. N Independence Lake, 2[42]. Nevada County: Independence Lake, 1[74]. Placer County: Donner, 3; 2 mi. W Soda Springs Station, 6500 ft., 1[74]; Blue Canyon, 5000 ft., 2 (1[74]); 4 mi. S Tahoe City, 1[74]. Eldorado County: 5 mi. S Tallac, 6300 ft., 1; Gilmore Lake, Mt. Tallac, 2[74]; Mt. Tallac, 1[68]; Phillips, 1[59]. Alpine County: 8000 ft., Hope Valley, 1; 8000 ft., Silver Creek, 1. Tuolumne County: Strawberry, 5200 ft., 1[74]; 9300 ft., Ten Lakes, Yosemite Park, 1[74]; Tuolumne Meadows, 8600 ft., Yosemite Park, 1[74]; Tuolumne Meadows (Soda Springs), 1; Tuolumne Meadows, 8500 ft., Yosemite Park, 1[74]; Sequoia, 1. Mariposa County: Chinquapin, 6256 ft., 2[74]; Merced Grove Big Trees, 5400 ft., 1[74]; Wawona, 1; no locality more definite than county, 1. Madera County: Bass Lake, 1[74]. _Mono County_: Tioga Crest, near Tioga Pass, 4[74]; Warren Creek, 1[74]; Tioga Lake, 1[74]; Ellery Lake, 9600 ft., 1[74]; Mono Lake P. O., Mono Lake, 1[74]; Walker Lake, 8000 ft., 2[74]; Pine City, 1; Mammoth, 13 (12[59], 1[14]); 10300 ft., near Big Prospector Meadow, White Mts., 2[74]. Inyo County: Little Onion Valley, 7500 ft., 1[74]; N Fork Bishop Cr., 10500 ft., 1[74]; S fork Bishop Cr., Andrews Camp, 8000 ft., 1[74]; South Lake, S Fk. Bishop Cr., 9750 ft., 1[74]; Lamarck Cr., 9900 ft., 15 mi. SW Bishop, 1[74]. Fresno County: Hume, 1. Tulare County: Mt. Whitney, 2; Whitney Meadow, 9800 ft., 1[74]; Monache Meadow, 8000 ft., 3[74]; E fork Kaweah River, 9800 ft., 1; 1/2 mi. S Mineral King, 7850 ft., 7[52]; Quaking Aspen Meadow, 7500 ft., 1[52].
=Colorado.= Moffat County: Lay, 1[19]. Routt County: Steamboat Springs, 2 (1[19]); no locality more definite than county, 1[57]. Jackson County: Higho, North Park, 8400 ft., 1; Buffalo or Illinois Creek, "near Rand," 6[74]. Washington County: 6 mi. NE Hillrose, 1[74]. Larimer County: Estes Park, 2 (1[2], 1[7]); Pinewood, 1; Loveland, 2 (1[57]); no locality more definite than county, 1[7]. Rio Blanco County: Compass Creek, 9000 ft., 1[2]; White River, 6200 ft., 1[21]; Piceance Creek, 6200 ft., 1[2]; Dry Fork, 6200-6600 ft., 4[2]; Meeker, 1; Marvine, 1[74]. _Grand County_: Crembling [= Kremmling?], 1[50]; Middle Park, 1[57]. Boulder County: Foot Mt. Meeker, 8700 ft., 1[2]; Silver Lake Mine, 1[60]; Boulder, 1[60]; Dixie Lake, 2 (1[2], 1[57]); Caribou, 1[2]; no locality more definite than county, 1. _Clear Creek County_?: Grays Peak, 1[93]. Jefferson County: 7000 ft., Mt. Parks, 1[57]; 6 mi. W Denver, 1[57]. Adams County: Barr, 1[2]; near East Lake, 2[57]. Denver County: Denver, 2 (1[2], 1[74]). Arapahoe County: Littleton, 1[19]. Summit County: Breckenridge, 1[57]. Eagle County: Eagle, 9500 ft., 1[104]. Park County: Jefferson, 4 (1[2]); 12800 ft., Mt. Bross, 1[57]. Mesa County: Tunnel, 1. Montrose County: near Crawford, Clear Fork of Smiths Fork, 1[19]; Coventry, 3 (1[19]); Naturita, 1; Paradox, 1[94]; West Paradox Valley, 1[57]. Pitkin County: Placita, 2[26]. Gunnison County: Marble, 1[26], Crested Butte, 2[19]; Deckers Ranch, Crested Butte, 2[19]; Sapinero, 7245 ft., 1[19]. Chaffee County: Buena Vista, 1[76]; Hancock, 1[16]; Salida, 5[19]. Teller County: Glencore, Pikes Peak, 1[76]. _El Paso County_: Monument, 1[76]; Seven Lakes, 1[19]; Lake Moraine, 10250 ft., 1[19]; Colorado Springs, 6000 ft., 1[19]; 5 mi. E Sand Creek, Colorado Springs, 1[19]; no locality more definite than county, 1[50]. Saguache County: Villa Grove, 1[19]; Pierce Place, Cochetopa Nat. Forest, 1; Houselog Creek, Cochetopa Nat. Forest, 1; P. Tevebaugh's Ranch, near Cochetopa Pass, 1; P. Tevebaugh's Ranch, 9 mi. S Cochetopa Pass, 1. Rio Grande County: between Monte Vista and Del Norte, 1[88]. Archuleta County: Upper Navajo River, 2[57]; Navajo River, 5 (4[57], 1[2]); Chromo, 2[57]. Conejos County: Osier, 3[57]. Montezuma County: Ure Peak, 1[57]. County in question: Del Norte Peak, 1[76]; no locality more definite than state, 4[75].
=Idaho.= Latah County: Cedar Mt., 4000 ft., 12 mi. NE Moscow, 1[55]; Moscow and 1/2 mi. W, 2[97]. Idaho County: Lochsa River (= Locksaw Fork), 1; between Selway Riv., and S Fork Clearwater Riv., 8[74]; Selway Divide, 8[74]; Pilot Creek, 2[74]; Newsome Cr., 1[74]. Lemhi County: Salmon River Mts., (now Lemhi Mts.), 8000 ft., 5; Leadore, 3. Adams County: summit Smith Mt., 7500 ft., 1[41]. Washington County: Midvale, 2. Custer County: Pahsimeroi Mts., 1; Double Springs, 16 mi. NE Dickey, 1[74]; Mackay?, 1; Stanley Lake, 1. Payette County: 2 mi. S Payette, 1[74]. Fremont County: 17 mi. E, 4 mi. N Ashton, 6275 ft., 2[74]. Teton County: 3 mi. S Victor, 1[74]. Jefferson County: 20 mi. W Camas, 1. Blaine County: Sawtooth City, 1; Ketchum, 5 (3[50], 2[75]). Canyon County: Nampa 3. Clark County: Dry Creek, Targhee Nat. Forest, 1[2]; Birch Creek, 2. County in question: North fork of Teton River, 1. Bingham County: Shelley, 1; Alridge, 2; Springfield, 1. Lincoln County: Shoshone, 1. Minidoka County: 1/4 mi. E Heyburn Bridge, 1[74]. _Power County_: 4 mi. NW American Falls, 1[74]. Bannock County: 3 mi. N Schutt's Mine, Ross Creek, 1[74]; 3 mi. N Pocatello, 1[74]; near (within 10 miles of) Pocatello, 1[74]; 3 mi. S Pocatello, 1[74]; 1 mi. E Portneuf, 1[74]; 2 mi. up Mink Creek, 2 (1[74], 1[41]); Inkom, 2; Swan Lake, 1. Owyhee County: 5 mi. SE Riddle, 1; Three Creek, 2. Cassia County: Elba, 1[52]. Bear Lake County: Geneva, 6171 ft., 1[74]; Montpelier, 1; Paris, 6000 ft., 1[6]; Pegram, 2.
=Nevada.= Humboldt County: Alder Creek, 7000 ft., Pine Forest Mts., 1[74]; head of Big Creek, 8000 ft., Pine Forest Mts., 1[74]; Cottonwood Range, 1; Calico Mt., Little Owyhee R., 1; Mahogany, Little Owyhee R., 2; Sulphur, 1. Pershing County: Lovelocks, 1. Elko County: Mountain City, 3; Three Lakes, Ruby Mts., 1[41]. Washoe County: Pyramid Lake, 1; 3 mi. E Reno, 1[74]; Incline Creek, 7100 ft., 1[74]; 2-1/2 mi. S Incline, 6250 ft., 1[74]; E side Marlette Lake, 8000 ft., 1[74]; Marlette Lake, 8000 ft., 1[74]. Ormsby County: 1/2 mi. S Marlette Lake, 8150 ft., 1[74]. Churchill County: 4 mi. W Fallon, 1[74]; 3 mi. W Fallon, 1[74]; 2 mi. W Fallon, 1[74]; Fallon, 3970 ft., 1[74]. 5 mi. S Fallon, 4000 ft., 1[74]; 8 mi. S and 3 mi. E Fallon, 1[74]. _Douglas County_: Mt. Siegel, 1[60]. Mineral County: Lapon Cañon, 8900 ft., Mt. Grant, 1. Nye County: Arc Dome, 1; 10700 ft., 1/2 mi. SW Jefferson Peak, Toquima Range, 1[74]. White Pine County: 3 mi. E Baker, 1[74]; Baker Creek, 6600 ft., 4[74]; Baker Creek, 8400 to 8450 ft., 4[74]; Gleason Creek, 7500 ft., 1[74]. Esmeralda County: Arlemont, 4850 ft., Fish Lake Valley, 1[74]. Lincoln County: 3 mi. S Crystal Spring, 3900 ft., Pahranagat Valley, 1[27].
=New Mexico.= Taos County: 2 mi. N Twining, 10500 ft., 1; Taos, 2. Santa Fe County: 11600 ft., Pecos Baldy, 1. _San Miguel County_: 8000 ft., above Willis, Pecos River, Forest Reserve, 2[75]; Ribera, 1.
=Oregon= (by counties from west to east). Jackson County: Rustler Peak, Crater Nat. Forest, 1[46]; Siskiyou (probably south of), 2. Klamath County: 20 mi. W Crescent, 1[101]; Anna Creek, Mt. Mazama, 2; S Boundary Crater Lake Nat. Park, 1[74]; Fort Klamath, 15; Upper Klamath Lake, 2[4]; Klamath Falls, 1[75]. _Lake County_: Dog Lake Ranger Station, 30 mi. SW Lakeview, 1. _Harney County_: Camp Harney, 2[75]; Burns, 2 (1[101]); 20 mi. S Burns, 1[46]; Narrows, 1[59]; Voltage, 1; Shirk P. O., 2; Keiger Gorge, Steen Mts., 4. Malheur County: Riverside, 1; 2 mi. NW Riverside, 2; Barren Valley, Cord, 1; Cedar Mts., 2; Cow Creek Lake, 1; Jordan Valley, 1. County in question: Sageview, 1.
=Utah.= Cache County: Logan, 1[74]. Rich County: 8000 ft., near Laketown, 1. Boxelder County: Willard, 1[103]. _Salt Lake County_: Salt Lake City, 1[74]; Barclay, 6500 ft., Wasatch Mts., 1; Mill Creek, 1[103]. Utah County: Provo Bench, 2[6]; Aspen Grove, Mt. Timpanogos, 1[6]; Payson, 1[6]. Juab County: between Santaquin and Starr, 1[103]. Uinta County: Dry Fork Canyon, 20 mi. NW Vernal, 1[9]. Carbon County: Sunnyside, 1[44]; Range Creek, 1[44]. Millard County: Deseret, 1[74]. Sevier? County: Fish Lake Plateau, 1. Grand County: Warner Ranger Station, La Sal Mts., 1[6]. Beaver County: Britts Meadows, 11000 ft., Beaver Range, 1[2]; Britts Meadows, Beaver Range, 1; Puffer Lake, 1[44]. Garfield County: Boulder, 2[6]. Washington County: Pine Valley, 1[44]; St. George, 1. San Juan County: Geyser Pass, La Sal Mts., 2[6]. County in question: Salt Lake, 2; Wasatch Mts., 1; La Sal Mts., 11000 ft., 1.
=Washington.= Okanogan County: Bald Mt., 6800 ft., 1; Bauerman Ridge, 6800 ft., Tungsten Mine, 1; Hart Pass, Methow River Trail, 1[46]; Conconully, 2 (1[51], 1[49]); 5 mi. NW Loomis, 1; Molson, 3800 ft., 1; Tunk Mt., 3500 ft., 1. Whatcom County: Barron, 5000 ft., 1. Stevens County: Colville, 1; Orin, 11[51]. _Pend Oreille County_: Ione, 6[51]. Chelan County: Chelan Mts., 1[2]; Lake Chelan, 1[46]; Manson, 1; Entiat River, 1680 ft., 20 mi. from mouth, 7; Dryden, 2[49]; Wenatchee, 1. Kittitas County: Easton, 2 (1[51]); Ellensburg, 1[51]; 4 mi. E Ellensburg, 1[51]. _Grant County_: Neppel, 1[51]. Lincoln County: Sprague, 1. _Spokane County_: Spokane, 1[94]; Cheney 2[89]. Whitman County: Pullman, 11 (6[55], 1[68], 1[10]); 6 mi. S Pullman, 1. Garfield County: Snake River, 1. Yakima County: Yakima, 1[74]; 1 mi. W Moxee, 1[74].
=Wyoming.= NW Wyoming, 1[75]. Yellowstone National Park: Lamar River, 1; Yellowstone Lake, 1. Park County: Greybull River, 1[80]. Teton County: Crystal Creek, 2; Jackson, 1; Whetstone Creek, 2[76]. Johnson County: Buffalo, 4 (2[93]). _Fremont County_: Continental Divide, 20 mi. NW Dubois, 1[75]. _Sublette County_: Bronx, 1[75]. Carbon County: Medicine Bow Mts., 1[75]; 15 mi. SE Parco, 1[74]. Albany County: Garrett, 1; 12 mi. W Laramie, 1[74]; 7 mi. W Laramie, 2[74]; 5 mi. W Laramie, 4[74]; "near" Laramie, 1[74]; 3 mi. SW Laramie, 1[74]; 12 mi. S Laramie, 1[74]. Uinta County: Fort Bridger, 6800 ft., 1[74]; Lonetree, 1; Bridger Pass, 1. County in question: Laramie River, 2. No locality more definite than state, 1.
=Mustela frenata effera= Hall
Long-tailed Weasel
Plates 19, 20 and 21
Mustela frenata effera Hall, Carnegie Instit. Washington Publ. 473:93, November 20, 1936.
Mustela arizonensis, Dice, Journ. Mamm., 1:12, November 28, 1919.
Type.--Male, adult, skull and skin; no. 33637, Amer. Mus. Nat. Hist.; Ironside, 4000 ft., Malheur County, Oregon; September 8, 1912; obtained by H. E. Anthony; original no. 267.
The skull (plates 19-21) is complete and unbroken. The teeth all are present and entire. The skin, in summer pelage, is well made.
Range.--Upper Sonoran to Arctic Alpine life-zones of northern two-thirds of Oregon east of the Cascades, and southeastern Washington, south of the Snake River. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. nevadensis in small size, males averaging 12-1/2 per cent smaller in external measurements, 8 per cent smaller in linear measurements of skull, and 22 per cent in weight of skull, total length averaging 360 rather than 400, condylobasal length averaging 40.5 rather than 43.6; from M. f. oregonensis in absence of frontonasal white patch, presence of light color of underparts on ventral face of tail and smaller skull with basilar length averaging less than 41.7 in males; from M. f. washingtoni in presence of light color of underparts on ventral face of tail, in male skull by linear measurements averaging 7 (5-12) per cent shorter and relative to basilar length shorter in preorbital region and broader across mastoid processes and zygomatic arches.
Description.--Size.--Male: Eight (6 adult and 2 subadult) males from northeastern Oregon yield average and extreme measurements as follows: Total length, 360 (340-378); length of tail, 129 (122-136); length of hind foot, 42 (40-44). Tail averages 56 (52-59) per cent as long as head and body. Length of hind foot more or less than (about same as) basal length.
Female: No. 212423 from Vale, and no. 566 V. B. Scheffer, from 15 mi. E Ukiah, measure, respectively: Total length, 312, 306; length of tail, 113, 114; length of hind foot, 35, 35. Tail averages 57 per cent as long as head and body.
Differences in external measurements between the one adult female and the average of the males are: Total length, 51; length of tail, 16; length of hind foot, 7.
Externals.--Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) about as shown in stage 4 of figure 19.
Color.--Upper parts, in summer, near (14 n to l) Brussels Brown or tones 1 to 3 of Raw Umber of Oberthür and Dauthenay, pl. 301, darker on top of head from nose to, or slightly behind, line connecting posterior margins of ears. Chin and usually all of upper lips white. Remainder of underparts Buff-Yellow to Straw Yellow. In winter all white except tip of tail or upper parts near (j) Snuff Brown or lighter than Brussels Brown with a smoked effect, with underparts white. Tip of tail at all times black. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of toes and wrists, on medial sides of hind legs to ankles over antiplantar faces of toes, distomedial third of tarsus and usually over proximal fourth to three-fourths of ventral side of tail. Least width of color of underparts averaging, in 15 males, 53 (36-69) per cent of greatest width of color of upper parts. Black tip of tail averaging 47 (38-67) mm. long. Thus averaging longer than hind foot, and 36 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 6 adults from northeastern Oregon): See measurements and plates 19-21. As described in Mustela frenata nevadensis except that: Weight, 2.9 (2.5-3.4) grams; basilar length, 40.5 (39.3-41.8).
Female (based on no. 212423, adult from Vale): In so far as parts of the broken skull permit a person to judge, the skull is as described in M. f. nevadensis except that: Smaller; lighter; postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite.
As compared with the skull of M. f. nevadensis that of effera seems, on the average, to have the preorbital part relatively smaller. Otherwise, the skull is a miniature of the skull of nevadensis, averaging about eight per cent smaller in linear measurements and weighs twenty-two per cent less. Comparisons of the skull with those of M. f. washingtoni and M. f. oregonensis are made in accounts of those subspecies.
Remarks.--This geographic race has long borne the name of Mustela arizonensis (Mearns). Small size differentiates effera from nevadensis and specimens have been allocated to one or the other subspecies on the basis of size, or average size when several individuals are available from one locality. Complete intergradation with each adjoining subspecies is indicated by numerous specimens, more of which are assigned to these adjoining subspecies than to effera itself.
The minimum of size in M. f. effera is found in the Blue Mountain region of northeastern Oregon. Specimens from the area intervening between these mountains and the Cascades average larger but are nearer the mean of typical effera than they are to the means of washingtoni, oregonensis or nevadensis.
Two males, nos. 204883, adult, and 204884, young, from Sisters, Oregon, near the eastern base of the Cascades, show approach structurally to M. f. washingtoni as it is represented at the nearby locality, Permilia Lake, at the west base of Mount Jefferson. Everything considered, however, the two specimens from Sisters are nearer to effera. A male from Condon, Oregon, shows approach to the Cascade race in slightly increased size.
No perfect skulls of adult females are available from the part of northwestern Oregon in which effera reaches its typical state of development as judged by the small size of the skull of the adult male. Skulls of adult females are available, however, from more nearly marginal localities. These, though smaller than in nevadensis, show relatively less difference in size when compared with nevadensis than do skulls of males. Even so the females at these marginal localities are smaller than those of nevadensis of comparable age and adequate material of adult female effera from the region where the males attain their extreme of small size probably will show about the same relative difference in size between nevadensis and effera as is known to exist between the adult males of these two subspecies. The small size of a subadult female, no 74631, U. S. Nat. Mus., from Asotin, Washington, constitutes partial basis for this opinion.
Of 14 adults examined none showed malformation of the frontal sinuses due to infestation by parasites.
Specimens examined.--Total number, 53, arranged within each state by counties from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=Oregon.= Wasco County: 4 mi. S The Dalles, 1[74]; Wapinita, 1; Antelope, 2; 7 mi. E Antelope, 5. Gilliam County: Condon, 1[46]. Morrow County: 10 mi. S Hardman, 1. Umatilla County: Umatilla, 2; 15 mi. E Ukiah, 4000 ft., 1[49]. Union County: Elgin, 1; 20 mi. E Lehman, 1[46]. Wallowa County: Horse Creek, 15 mi. N Paradise, 1; Enterprise, 1[46]; Wallowa Lake, 1[46]; Wallowa Mts., 8300 ft., 1. Baker County: Haines, 1[49]; Anthony, 3[2]; Bourne, 2. Grant County: Long Creek, 1[46]; Canyon Creek, 1[46]; Strawberry Mts., 2; Silvies, 1[14]. Crook County: Prineville, 4. Deschutes County: Sisters, 2; Bend, 1. Lake County: 3 mi. W Stauffer, 1; Fort Rock, 1[46]. Harney County: 25 mi. NW Burns, 1. Malheur County: 4000 ft., Ironside, 2[2]; 1-1/2 mi. S Vale, 2.
=Washington.= Walla Walla County: Prescott, 4 (2[76], 1[60], 1[74]); Ft. Walla Walla, 2 (1[75]); Wallula, 1[76]. _Asotin County_: Asotin, 1.
=Mustela frenata washingtoni= (Merriam)
Long-tailed Weasel
Plates 19, 20, 21, 34, 35 and 36
Putorius washingtoni Merriam, N. Amer. Fauna, 11:18, pl. 4, figs. 3, 3a, 4, 4a, June 30, 1896.
Mustela washingtoni, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912.
Mustela frenata washingtoni, Hall, Carnegie Instit. Washington Publ. 473:106, November 20, 1936.
Type.--Male, adult, skin and skull; no. 76322, U. S. Nat. Mus., Biol. Surv. Coll.; Trout Lake, Mt. Adams, Klickitat (?) County, Washington; December 15, 1895; obtained by D. N. Kaegi; original no. 2.
The skull is unbroken. The left incisors above are missing. Otherwise the teeth are present and entire. The skin is well made, in brown winter pelage, lacks collector's measurements, has no bones in the feet, but by large size is judged to be a male.
Range.--Altitudinally from near 2000 feet at Trout Lake up to the highest parts of the Cascade Range from Mount Jefferson, Oregon, north to Mount Rainier, Washington; Upper Sonoran Life-zone to Arctic Alpine Life-zone. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. altifrontalis in lighter color of upper parts and underparts, latter ranging from Buff-Yellow to Naples Yellow rather than near (14 a to 16 c) Ochraceous-Buff, in shallower skull in both sexes (see measurements), in males, a longer preorbital region, narrower skull with shorter bullae, and in females, a smaller skull with interorbital breadth averaging less than 24 per cent of basilar length; from M. f. nevadensis in absence of light color of underparts on ventral face of tail, in skulls of males, by longer preorbital region and narrower skull across mastoid processes and zygomatic arches, in skulls of females, by shorter preorbital region, and smaller bullae (see measurements); from M. f. effera in absence of light color of underparts on ventral face of tail, in skulls of males, by linear measurements averaging 7 (5-12) per cent larger, and relative to basilar length, longer in the preorbital region and narrower across mastoid processes and zygomatic arches; from M. f. oregonensis in absence of frontonasal white patch, longer skull in males, which in percentage of basilar length has, on the average, orbitonasal length amounting to more than 35, mastoid breadth less than 55, and zygomatic breadth less than 63, and in females, smaller skull with least width of palate less than length of P4, upper tooth-rows less than 38-1/2 per cent of basilar length, bullae smaller, averaging less than 13.4 in length.
Description.--Size.--Male: Fifteen subadult topotypes yield average and extreme measurements as follows: Total length, 400 (357-437); length of tail, 149 (122-171); length of hind foot, 47.6 (42-59). Tail averages 59 per cent as long as head and body. Length of hind foot averaging more than basal length. Corresponding measurements of one adult and 3 young from Mount Rainier are: 415 (405-423); 155 (145-164); 51 (50-53).
Female: Five adult topotypes yield average and extreme measurements as follows: Total length, 349 (330-393); length of tail, 124 (114-133); length of hind foot, 38 (36-39). Tail averages 55 per cent as long as head and body. Length of hind foot averaging about same as basal length. Corresponding measurements of two adults and 6 young from Mount Rainier are: 338 (320-360); 121 (115-132); 36 (34-40).
The average differences in external measurements of the two sexes, from Mount Adams, are: Total length, 51; length of tail, 25; length of hind foot, 9.6. Corresponding differences between the specimens from Mount Rainier are: 77; 34; 15.
Externals.--Longest facial vibrissae black or brown (often both colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to or beyond apical pad of fifth digit; hairiness of foot-soles slightly less than shown in figure 19.
Color.--Upper parts in summer near (14 n) Argus Brown or tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304; one topotype Buckthorn Brown or tone 3 to 4 of Snuff Brown of Oberthür and Dauthenay, pl. 303. Dark spot at each angle of mouth present or absent, and when present, often fused with color of upper parts, which rarely covers lower lips. Chin, and usually lower lips, white. Remainder of underparts Buff-Yellow to Naples Yellow. In winter, all white except tip of tail which is at all times black, or upper parts near (14) Brussels Brown to near (j) Snuff Brown with smoked effect and underparts white, rarely with trace of yellowish. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and usually all of wrists, on medial sides of hind legs anywhere from knee to tips of toes. Least width of color of underparts averaging in ten topotypes, 24 (10-37) per cent of greatest width of color of upper parts. Black tip of tail in same series averaging 55 (45-60) mm. long, thus longer than hind foot and averaging 37 per cent of length of tail-vertebrae.
The color of the underparts is not so narrow in the specimens from Mount Rainier and it is believed that the slender bodies used in stuffing the topotypes has accentuated in them the appearance of narrowness of the light-colored underparts.
Skull and teeth.--Male (based on 22 adult topotypes): See measurements and plates 19-21; weight, 3.5 (2.8-4.7) grams; basilar length, 43.7 (40.0-47.7); zygomatic breadth more or less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less than postpalatal length; postorbital breadth less than length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bullae; breadth of rostrum less (except in no. 82180) than length of tympanic bulla; least width of palate more (except in no. 81954) than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2 to 5 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; length of tympanic bulla more (except in two instances) than length of lower molar and premolar tooth-row and shorter (except in two instances) than rostrum; anterior margin of masseteric fossa below m2.
Female (based on 11 ad. topotypes): See measurements and plates 34-36; weight, 2.0 (1.8-2.2) grams; basilar length, 37.6 (37.0-38.9); zygomatic breadth less (except in no. 70945) than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less (except in one specimen) than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3-1/2 to 5-1/2 upper incisors; height of tympanic bulla more or less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.
Compared with M. f. nevadensis, the skull of the male of washingtoni averages more slender, as shown by the mastoid and zygomatic breadths and has the preorbital part longer, on the average, as shown by the greater ratio (to the basilar length) of the length of the tooth-rows and orbitonasal length. Also, on the average, the postorbital constriction is longer than in nevadensis and the tympanic bullae are smaller. In females, the skull is lighter, the tooth-rows are shorter, the tympanic bullae are smaller, and the preorbital part of the skull is shorter and narrower as shown by the orbitonasal length and interorbital breadth. Except that the tympanic bullae are actually, although not relatively, smaller in males of effera, it differs from washingtoni in the same way as does nevadensis as regards relative proportions, but, of course, the actual difference in size is greater since effera is smaller than nevadensis. Comparison of the skull with that of oregonensis is made in the account of that subspecies.
Remarks.--M. f. washingtoni was described and named in 1896 by Merriam as a distinct species. Subsequently, specimens which here are regarded as intergrades between altifrontalis and nevadensis, were classified as washingtoni.
The external measurements given for the specimens from Mount Adams are those recorded on the labels in inches and fractions thereof. Instead of total length there sometimes is written "tip to tip." In the series of 19 winter-taken topotypes the hairs project beyond the end of the caudal vertebrae for an average distance of 28 (19-40) millimeters. If the hairs on the end of the tail were included in the measurements, 28 millimeters should be subtracted from the averages. Probably the measurements should stand as given, since an adult male topotype, no. 226758, U. S. Nat. Mus., taken subsequently by Walter P. Taylor measures 405; 152; 51.
Mustela frenata washingtoni is not a strongly marked geographic race. In many features it is intermediate between M. f. altifrontalis and M. f. nevadensis. This is especially true of coloration. In the series from Mount Adams and that from Mount Rainier, some individuals have the light color of the underparts extended down the hind legs over the feet and over the proximal face of the ventral third of the tail as in nevadensis, whereas others from the same place have the light color of the underparts absent from the tail and extending no farther down the hind limbs than the knees. The light color of the underparts in the series of topotypes is so restricted that the transverse extent at the narrowest place amounts to only 24 (10-37) per cent of the greatest width of the color of the upper parts. This narrowness of the color of the underparts has been likened by Merriam (1896:18) to the condition in Mustela frenata noveboracensis. So it is, but it is similar to the condition found also in the geographically adjoining M. f. altifrontalis.
Of the 37 skulls of subadults and a few adults, 11 had the frontal sinuses malformed as a result of infestation by parasites.
Specimens examined.--Total number, 56, arranged within each state by localities from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=Oregon.= Mt. Jefferson, Permilia Lake, 1.
=Washington.= Pierce County: 5500 ft., Spray Park, Mt. Rainier, 1; Spray Park, 1[74]; 5935 ft., Glacier Basin, Mt. Rainier, 5 (1[10]); 5051 to 5100 ft., Owyhigh Lakes, Mt. Rainier, 7 (1[10]), Tahoma Creek, 1[72]; Nisqually entrance, 1[72]; Longmire, 1[72]; Mt. Rainier Nat'l Park, 2[72]. Klickitat County: Trout Lake, S Base Mt. Adams, 35; 3500 ft., Gotchen Creek, Mt. Adams, 1.
=Mustela frenata saturata= (Merriam)
Long-tailed Weasel
Plates 19, 20, 21 and 30
Putorius saturatus Merriam, N. Amer. Fauna, 11:21, June 30, 1896.
Mustela saturata, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912.
Mustela arizonensis saturata, Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933.
Mustela frenata saturata, Hall, Carnegie Instit. Washington Publ. 473:106, November 20, 1936.
Type.--Male, adult, skull and skin; no. 65930, U. S. Nat. Mus., Biol. Surv. Coll.; Siskiyou, Jackson County, Oregon; June 6, 1894; obtained by C. P. Streator; original no. 3905.
The skull (plates 19-21, 30) lacks the middle part of each zygomatic arch. The teeth all are present although much worn, probably from gnawing at the trap which captured the animal. The skin, in fresh summer pelage, is fairly well made.
Range.--Transition and Boreal life-zones of Siskiyou and Trinity mountains in southern Oregon and northwestern California. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. nevadensis in lacking light color of underparts on tail and ankle and in greater average breadth across mastoid processes of skull (see measurements); from M. f. oregonensis in lacking white nasofrontal spot, in having color of underparts interrupted at ankle; from M. f. munda in lacking white nasofrontal spot, in smaller and relatively deeper skull of males and smaller skull of the female.
Description.--Size.--Male: Four adult males (the type, 1 from Mt. Ashland and 2 from Jackson Lake) yield average and extreme measurements as follows: Total length, 414 (402-437); length of tail, 150 (136-160); length of hind foot, 46 (43-50). Tail averages 57 (49-62) per cent as long as head and body. Length of hind foot more or less than basal length.
Female: One young from the summit of the Trinity Mountains east of Hoopa and one nontypical adult from 5500 feet elevation on South Fork Mountain, Humboldt County, measure respectively as follows: Total length, 330, 325; length of tail, 115, 123; length of hind foot, 37, 37. Tail is 53 and 61 per cent as long as head and body. Length of hind foot less than basal length.
Average differences in external measurements between the two sexes, indicated by the unsatisfactory material available, are: Total length, 86; length of tail, 31; length of hind foot, 9.
Externals.--Longest facial vibrissae black or dark brown and extending beyond ear; carpal vibrissae same color as underparts and extending as far as apical pad of fifth digit; hairiness of foot-soles, in summer pelage, as shown in figure 19.
Color.--Upper parts, in summer, Brussels Brown to near (n) Brussels Brown or lighter than tone 3 of Raw Umber of Oberthür and Dauthenay, pl. 301, usually darkest on nose and forehead. Chin white. Remainder of underparts Buff-Yellow to Warm Buff. Tip of tail black. Winter pelage unknown. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and sometimes wrists, on medial sides of hind legs only to ankles, but toes sometimes with isolated white markings. Least width of color of underparts in the type and 2 adults from Jackson Lake averaging 35 (30-40) per cent of greatest width of color of upper parts. Black tip of tail averaging 54 (53-55) mm. long; thus longer than hind foot and averaging 37 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 4 adults: Type, Mt. Ashland, 1; Jackson Lake, 2): See measurements and plates 19-21, 30. As described in Mustela frenata nevadensis except that: Weight, 3.8 (3.5-4.3) grams; basilar length, 44.4 (42.6-45.8); zygomatic breadth more or less than distance between condylar foramen and M1 or than distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more than postpalatal length; least width of palate less than medial length of P4 (except in one specimen).
Female (based on one adult possibly not typical, from 5500 ft., South Fork Mt.): See measurements. As described in Mustela frenata nevadensis except that: Weight, 2.2 grams; basilar length, 38.1; zygomatic breadth less than distance between condylar foramen and M1 and less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite.
The skull of the male of saturata, relative to the basilar length, is broader across the mastoids and narrower across the rostrum and interorbital region than that of nevadensis. Skull not known certainly to differ from that of oregonensis. Compared with the skull of munda, that of the male of saturata is smaller in every part measured except depth of tympanic bullae which averages 3.6 millimeters, rather than 3.5 as in munda. Also, the skull of saturata has a less-marked postorbital constriction, is less heavily ridged, less angular, does not have the impressions of the temporal muscles carried so far forward on the frontal bones and is relatively much narrower across the zygomatic arches.
Remarks.--In 1896, Merriam named M. f. saturata as a distinct species on the basis of one specimen, taken by Clark P. Streator at Siskiyou, Oregon, and a second specimen taken the year previously by Allan C. Brooks at Chilliwack, British Columbia. On the basis of these two specimens, Merriam (1896:22) ascribed to the race a range ". . . on the Cascade and Siskiyou mountains of Oregon and Washington, reaching a short distance into British Columbia." Since that time, this name, saturata, has been employed for the dark-colored weasels, of the coastal region of Oregon, Washington, and extreme southwestern British Columbia, which here are arranged under the name M. f. altifrontalis. M. f. saturata proves to be restricted to the humid mountainous region inland from the coast in northern California and in the Siskiyou Mountains of southern Oregon. Its range is separated by that of M. f. oregonensis from the range of the darker-colored, deeper-skulled, M. f. altifrontalis of the humid costal region proper.
On May 5, 1933, Mr. Clark P. Streator, informed the writer that he remembered taking the type specimen of Mustela frenata saturata (Merriam) in the town of Siskiyou, Oregon. The exact place, he said, was reached, at the time of his work there, by going one or two blocks east of the depot, then through a garden into the thick woods where there were springs and numerous burrows of the rodent, Aplodontia. Two other weasels labeled as taken at Siskiyou, on September 28 and 29, 1893, by Mr. Streator, are much lighter colored than the type of saturata and have the color of the underparts extended distally on the hind legs to the tips of the toes and in other features of coloration are more like nevadensis, the subspecies to which they are referred, than saturata. Probably these did not come from exactly the same place that the type specimen of saturata did. Although Mr. Streator does not remember the taking of these particular specimens in 1893, he does remember that on this visit to Siskiyou, he walked southward through the railroad tunnel and collected on the opposite side of the ridge from Siskiyou. Here on more southern exposures, the country was markedly different than in the thick forest at Siskiyou. Probably these two specimens taken in 1893, and referred to nevadensis, came from a little way south of Siskiyou and from a different habitat and life-zone than the type specimen of M. f. saturata.
Of the 6 specimens examined, only one, the type, shows malformation of the frontal sinuses such as result from infestation by parasites.
Specimens examined.--Total number, 6, as follows:
=California.= Siskiyou County: Jackson Lake, 5900 ft., 2, Mus. Vert. Zoöl. Humboldt County: South Fork Mt., 5500 ft., 1, Mus. Vert. Zoöl. County in question, Trinity Mts., summit east of Hoopa, 5800 ft., 1, U. S. Nat. Mus.
=Oregon.= Jackson County: Mt. Ashland, 1, Univ. Oreg.; Siskiyou, 1, U. S. Nat. Mus.
=Mustela frenata altifrontalis= Hall
Long-tailed Weasel
Plates 1, 19, 20, 21, 34, 35 and 36
Mustela frenata altifrontalis Hall, Carnegie Instit. Washington Publ. 473:94, November 20, 1936.
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Putorius saturatus Merriam, N. Amer. Fauna, 11:21, June 30, 1896 (part).
Mustela saturata, Miller, U. S. Nat. Mus. Bull., 79:98, December 31, 1912.
Type.--Male, adult, skull and skin; no. 42093, Mus. Vert. Zoöl.; Tillamook, Tillamook County, Oregon; July 10, 1928; obtained by Alex Walker; original no. 717.
The skull is complete and unbroken. P3 on the left side is missing; otherwise the teeth all are present and entire. The skin is well made and the enlarged scrotal pouch shows the collector's sexing of the specimen to have been correct.
Range.--Altitudinally from sea level up to at least 4800 feet (Mount Baker) in the Transition Life-zone of the humid, coastal region of Oregon, Washington and extreme southwestern British Columbia. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. nevadensis in tone 4 of Brownish Drab, pl. 302, rather than tones 1-3, of Raw Umber, pl. 301, of Oberthür and Dauthenay of upper parts, in near (14 a´ to 16 c´) Ochraceous-Buff rather than Buff-Yellow to Straw Yellow of underparts, in that least width of color of underparts amounts to less than 37 per cent of greatest width of color of upper parts, in absence of color of underparts on ventral side of tail and on hind leg distal to knee, and in greater depth of skull through frontal region; from M. f. washingtoni in darker color of upper parts and underparts, latter near (14 a´ to 16 c´) Ochraceous-Buff rather than ranging from Buff-Yellow to Naples Yellow, in deeper skull in both sexes (see measurements), in males a shorter preorbital region, broader skull with longer bullae and in females a larger skull with interorbital breadth averaging more than 24 per cent of basilar length; from M. f. oregonensis in frontonasal white patch absent, color above darker (tone 4 of Brownish Drab, pl. 302, rather than tone 2 to 3 of Raw Umber, pl. 301 of Oberthür and Dauthenay), light-colored underparts narrower and not extended distally beyond knee, in females tooth-row shorter, amounting to less than 38 per cent of basilar length.
Description.--Size.--Male: Eight adult topotypes yield average and extreme measurements as follows: Total length, 426 (392-445); length of tail, 160 (148-170); length of hind foot, 47 (42-53). Tail averages 60 per cent as long as head and body. Length of hind foot averages more than basal length.
Female: Five adults from Tillamook and Blaine, Oregon, yield average and extreme measurements as follows: Total length, 347 (320-370); length of tail, 125 (114-131); length of hind foot, 38 (35-44). Tail averages 56 per cent as long as head and body. Length of hind foot less than basal length.
The average differences in the external measurements are: Total length, 79; length of tail, 35; length of hind foot, 9.
Externals.--Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to or beyond apical pad of fifth digit; hairiness of foot-soles (in summer pledge) slightly less than shown in figure 19.
Color.--Upper parts, in summer, near (n) Argus Brown or tone 4 of Brownish Drab of Oberthür and Dauthenay, pl. 302. Dark spot at each angle of mouth well developed; often fused with color of upper parts which sometimes covers lower lips. Chin white. Remainder of underparts near (14 a´ to 16 c´) Ochraceous-Buff. In winter, upper parts near (14) Argus Brown with smoked effect and Warm Buff to Naples Yellow below. Tip of tail at all times black. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and usually all of wrists, on medial side of hind legs typically only to knee but sometimes to ankle. Tips of toes of hind feet almost always marked with color of underparts. Least width of color of underparts averaging in a series of 14 males from Blaine, Oregon, 23 (14-36) per cent of greatest width of color of upper parts. Black tip of tail in 8 adult males from Blaine, Oregon, averaging 59 (47-70) mm. long; thus longer than hind foot and averaging 37 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 9 adults from Blaine, Tillamook Co., Oregon): See measurements and plates 19-21; weight, 4.4 (3.3-5.3) grams; basilar length, 45.6 (42.4-47.7); zygomatic breadth more or less (usually more) than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less (usually more) than postpalatal length; postorbital breadth less (except in some instances of malformations of frontal sinuses which result from infestation by parasites) than length of upper premolars and more or less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than orbitonasal length; anterior margin of masseteric fossa directly below m2.
Female (based on 4 adults): See measurements and plates 34-36; weight, 2.2 (2.2-2.3) grams; basilar length, 38.1 (37.8-39.7); zygomatic breadth more or less (less in three of four specimens) than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; relation of postorbital breadth to other measurements in doubt because of malformation of frontal sinuses by parasites; least width of palate not less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3-1/2 to 5-1/2 upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.
Compared with the skull of M. f. washingtoni that of each sex of altifrontalis averages slightly larger in every measurement taken, except measurements of teeth which are approximately the same, and is relatively deeper through the frontal region and through the braincase as measured at the anterior margin of the basioccipital. Skulls of females of altifrontalis have a relatively broader interorbital region. Skulls of males of altifrontalis further differ in having relatively, as well as actually, longer tympanic bullae, relatively lesser orbitonasal length and a greater relative breadth across the mastoids and across zygomata. Compared with M. f. nevadensis, the skull of the male of altifrontalis averages slightly larger and heavier although the skulls of females are of approximately the same size and weight. Relative to the basilar length, the skulls of both sexes are deeper through the braincase and narrower across the mastoids; the rostrum is broader, especially in males; the tooth-rows are shorter and the interorbital breadth less, especially in females. Comparison with the skull of oregonensis is made in the account of that subspecies.
Remarks.--Until the present study was begun, animals of this race have gone under the name Mustela saturata (Merriam). The United States National Museum has a juvenile taken, in 1858, by Wayne at Astoria, O. T.; the Samuel N. Rhoads collection contained one specimen taken in 1891, at Tacoma, Washington; one in the Bangs' collection was taken at Chilliwack, British Columbia, in 1895, and the Field Museum has one taken on the Olympic Peninsula in 1898. The best material is that collected by Alex Walker, at Tillamook, Oregon.
Intergradation with nevadensis is indicated by several specimens. The coloration of the one adult female, no. 90, Chas. R. Conner Mus., from Swamp Creek, Washington, has the color of the underparts extended down the hind legs over the feet, and over the proximal third of the ventral face of the tail as in nevadensis although the other two specimens from the same place have the color pattern of altifrontalis. Of the four specimens from British Columbia referred to this subspecies, only the specimen from Chilliwack is typical as regards color pattern. The one from Cultus Lake has the color pattern of nevadensis and might be referred to that race almost as well as to altifrontalis. The two specimens from Lihumption Park are intermediate between the two races in tone of color. Neither has the color of the underparts extended onto the tail or continuously over the hind feet as in nevadensis but each does have the color of the underparts less restricted and of lighter hue than in altifrontalis. Only one of the specimens, no. 7848 Canad. Nat. Mus., from Lihumption Park is adult and it has a skull which agrees with that of altifrontalis rather than nevadensis.
After writing the above, a good representation of the weasel population along the eastern side of Puget Sound was made available by friends in that area. Study of the weasels from there shows that their color is intermediate between that of altifrontalis and nevadensis. On the whole, they (specimens from Bellingham, for example) resemble one subspecies about as much as the other. In cranial characters some specimens, in certain features, approach nevadensis but most specimens agree with altifrontalis and all are more nearly like altifrontalis to which race all are referred.
The color of these animals is to me indistinguishable from that of washingtoni. The color of washingtoni is merely intermediate between that of nevadensis and altifrontalis. Nevertheless, the race washingtoni has cranial characters (long narrow skull) which set it off from both altifrontalis and nevadensis. This shape of skull is not found in the specimens from along the eastern side of Puget Sound; these animals have skulls like that of altifrontalis and when departures from this occur they are in the direction of nevadensis and not washingtoni.
The above, then, explains why specimens which are colored like those of washingtoni are not referred to that race but instead to the race altifrontalis.
Of 23 adult skulls examined, 19 have the frontal sinuses malformed as the result of infestation by parasites.
Specimens examined.--Total number, 80, arranged within states by counties from north to south. Unless otherwise indicated specimens are in the United States National Museum.
=British Columbia.= Chilliwack, 1[74], Lihumption Park, 4750 ft., 2[77]; Cultus Lake, 1[77].
=Oregon.= Clatsop County: Old Fort Clatsop, 1[74]; Astoria, 1. Tillamook County: Tillamook, 12 (7[14], 2[74], 2[2], 1[46]); Netarts, 1[46]; Blaine, 16 (13[14], 1[93], 1[76], 1[59]). _Lane County_: Reed, 1; Mercer, 1[46]. Curry County: Langlois, 1[46].
=Washington.= Whatcom County: Nooksack River, 2000 ft., 14 mi. E Glacier, 1; Swamp Creek, 2050 ft., Nooksack River, 3[10]; Lookout, 4800 ft., Mt. Baker, 2[10]; Bellingham, 8[25]; 5 mi. S Bellingham, 1[49]. Skagit County: Rockport, 300 ft., 1. King County: Bothell, 2[94]; N Seattle 1[51]; Seattle, 1[49]; Tye, 1[51], 2 mi. E Skykomish, 1[51]; 7 mi. E Kent, 1[76]; Auburn, 3[94]. _Pierce County_: Tacoma, 1[1]. Clallam County: Sequim, 1[49]; Soleduc Riv., near [sic.] Sappho, 1[49]; Happy Lake, 1[60]; mouth of Boulder Creek, Elwha River, 560 ft., Olympic Mts., 1; Hume's Ranch, 1000 ft., Elwha River, 1; Bogachiel Riv., 1[49]. _Mason County_: Lake Cushman, 2; 4 mi. N Shelton, 1[51]. _Thurston County_: Olympia, 2[49]; Tenino, 1[51]. Pacific County: 2-1/2 mi. SE Chinook, 3[74].
=Mustela frenata oregonensis= (Merriam)
Long-tailed Weasel
Plates 19, 20, 21, 30, 34, 35 and 36
Putorius xanthogenys oregonensis Merriam, N. Amer. Fauna, 11:25, June 30, 1896; Bangs, Proc. New England Zoöl. Club, 1:57, June 9, 1899.
Mustela xanthogenys oregonensis, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.
Mustela xanthogenys munda, Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933 (part).
Mustela frenata oregonensis, Hall, Carnegie Instit. Washington Publ. 473:107, November 20, 1936.
Type.--Male, subadult, skull and skin; no. 32019/43828, U. S. Nat. Mus., Biol. Surv. Coll.; Grants Pass, Rogue River Valley, Josephine County, Oregon; December 19, 1891; obtained by C. P. Streator; original no. 1404.
The skull (plates 19-21, 30) is complete and unbroken. P3 on the left side is missing. Otherwise the teeth all are present although worn probably as a result of gnawing at the trap which captured the specimen. The skin, in brown, winter pelage, is fairly well made.
Although the label on the skin and the label in the skull vial each give the sex of the specimen as female, and although Merriam (1896:25) regarded the specimen as a female, the present writer regards the specimen as a male.
It is as large as other undoubted males and larger than any known female of this subspecies. The labels with the skull and skin give the locality as "Rogue River Valley, Oregon." The listing here of the more restricted locality, Grants Pass, is made on the basis of Merriam's (1896:25) original description of the subspecies.
Range.--Transition and Canadian life-zones along coast of northern California and southern Oregon from Humboldt County, California, north through Curry County, Oregon, thence inland, west of the Cascades, north to the Columbia River. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. altifrontalis in presence of frontonasal white patch, lighter color above (tone 2 to 3 of Raw Umber, pl. 301, rather than tone 4 of Brownish Drab, pl. 302, Oberthür and Dauthenay), wider extent of light color of underparts which is extended distally beyond knee, and in females, longer tooth-row which amounts to more than 38 per cent of basilar length; from M. f. munda in shorter hind foot of males which is less than 50, and in both sexes, smaller, less rugose skull (see measurements and plates); from M. f. saturata in presence of frontonasal white patch, in having color of underparts extended uninterruptedly over ankle onto foot; from M. f. nevadensis in presence of frontonasal white patch, lack of light color of underparts on ventral face of tail, and longer skull, which relative to its length in males, is shallower through braincase; from M. f. effera in presence of frontonasal white patch, lack of light color of underparts on ventral face of tail, and larger skull with basilar length averaging more than 41.7 in males; from M. f. washingtoni in presence of frontonasal white patch, shorter skull in males, which in percentage of basilar length has, on the average, orbitonasal length amounting to less than 35, mastoid breadth more than 55, and zygomatic breadth more than 63; and in females larger skull with least width of palate more than length of P4, upper tooth-rows more than 38-1/2 per cent of basilar length, bullae larger and averaging more than 13.4 long.
Description.--Size.--Male: Five males (3 adults and 2 subadults from Eureka, Ferndale, and Carlotta, California) yield average and extreme measurements as follows: Total length, 392 (347-430); length of tail, 138 (110-160); length of hind foot, 46 (43-50). Tail averages 54 (46-61) per cent as long as head and body. Length of hind foot more or less than basal length. The type specimen, and an adult from Goldbeach measure, respectively, as follows: Total length, 412, 386; length of tail, 155, 137; length of hind foot, 44, 46.
Female: Three adults (2 from Fortuna and 1 from Carlotta, California) yield average and extreme measurements as follows: Total length, 367 (360-374); length of tail, 130 (123-134); length of hind foot, 40 (39-40). Tail averages 55 (52-57) per cent as long as head and body. Length of hind foot less than basal length. A subadult from Goldbeach, an adult from 13 mi. SW Grants Pass, and an adult from Medford, measure, respectively, as follows: Total length, 316, 344, 294; length of tail, 114, 120, 122; length of hind foot, 36, 40, 38.
The average differences in external measurements of the two sexes in the vicinity of Carlotta, are: Total length, 25; length of tail, 8; length of hind foot, 6. Corresponding differences, at Goldbeach, are: 70, 23, 10. Probably the females at Fortuna reflect the large size of munda more than do the males at Carlotta and the differences between the measurements of the two sexes probably, therefore, are actually more than are indicated by the figures above.
Externals.--Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles, in summer pelage, as shown in figure 20.
Color.--Upper parts, in summer, near (16 l) Brussels Brown or tone 2 of Raw Umber of Oberthür and Dauthenay, pl. 301, to slightly darker than tone 3 of same plate. Darker on nose and top of head, usually with frontonasal white patch but lacking white bar in front of each ear, except in the type and 2 specimens from Salem. Chin, lower lips, angle of mouth, and usually posterior seventh of upper lip white. Remainder of underparts Pale Orange-Yellow. In winter usually lighter above with underparts Warm Buff to Straw Yellow. Tip of tail at all times black. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial side of hind leg, typically over ankle in extremely narrow line which widens out over distal phalanges of antiplantar faces of toes but sometimes interrupted at ankle. Least width of color of underparts averaging, in twenty available specimens, 39 (27-54) per cent of greatest width of color of upper parts. Black tip of tail in five adults averaging 50 (43-60) mm. long; thus averaging longer than hind foot and 33 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 4 adults and subadults from Eureka, Requa, Goldbeach, and Grant Pass): See measurements and plates 19-21, 30. As described in Mustela frenata nevadensis except that: Weight, 3.5 (3.5-4.1) grams; basilar length, 42.9 (41.8-44.0); least width of palate more or less than medial length of P4.
Female (based on 2 adults, one from Carlotta and one from 13 mi. SW Grants Pass): See measurements and plates 34-36. As described in Mustela frenata nevadensis except that: Weight, 2.4 (2.2-2.6) grams; basilar length, 37.7 and 39.5; zygomatic breadth less than distance between condylar foramen and M1 and less than distance between anterior palatine foramen and anterior margin of tympanic bulla. See under "Remarks" for additional data on variation in size of skulls of females.
The skulls of the female averages 31 per cent lighter than that of the average male.
Because there is much geographic variation between specimens here referred to oregonensis, the person who is guided by the present account should keep in mind that results, here reported, of comparisons of the skull with those of other races, were obtained by employing specimens of oregonensis from Carlotta and Eureka, California. These specimens from California are judged to have more of the characters of the subspecies munda than do specimens of oregonensis from more northern localities.
Compared with that of M. f. washingtoni the skull of the male is shorter, especially in the preorbital region and is relatively broader across the mastoidal processes and zygomatic arches. The skull of the female is longer in the preorbital region, has a less cylindrical braincase and differs less from the male skull than is the case in M. f. washingtoni. Compared with M. f. effera, the skull of the male is smaller in every part measured and relative to the basilar length is broader across the mastoids and has relatively shorter tympanic bullae. From M. f. nevadensis the skull of the male differs in the same way except that size is about the same. The skull of the female oregonensis is more heavily ridged and is relatively broader across the mastoids than that of effera. From M. f. saturata, oregonensis is not surely known to differ in cranial characters. From M. f. munda, oregonensis differs in having the skull of both sexes smaller, and on the average, in all parts measured, has a less marked postorbital constriction, relatively narrower interorbital region and relatively more expanded zygomata. From M. f. altifrontalis, males of oregonensis differ on the average, in having larger teeth, and relative to the basilar length, a greater mastoid breadth and a shallower braincase as measured at the anterior margin of the basioccipital. Females of oregonensis differ in larger average size of skull, except for breadth of rostrum and interorbital breadth which, therefore, are relatively less in oregonensis, as also is the relative depth of the skull measured at the posterior borders of the upper molars and at the anterior margin of the basioccipital. However, skulls of females of oregonensis have relatively longer tooth-rows and are relatively broader across the zygomata and mastoidal processes.
Remarks.--In 1896, Merriam named oregonensis as a subspecies of the California bridled weasel on the basis of a single specimen taken by Clark P. Streator. Three additional specimens were acquired in later years, by workers of Dr. Merriam's bureau, from near the type locality and specimens from farther north in Oregon have been accumulated at the University of Oregon. The most satisfactory material is that saved from Humboldt County by the late H. E. Wilder, which, when brought together, is adequate to give some idea of the range of variation that can be expected in a given population.
Of two specimens from Goldbeach, one shows approach to altifrontalis in that the color of the underparts stops at the ankle, and in one, the angle of the mouth is dark colored. Specimens from Eugene and vicinity lack the white facial markings, and in this feature approach the adjoining washingtoni-effera-nevadensis stock. A specimen from 6 miles south of Medford shows approach to saturata in the interruption, on the ankle and lower tibial region, of the color of the underparts. One adult female, no. 1413, Univ. Oregon, from the Rogue River Valley, 13 miles southwest of Grants Pass, stands out prominently, among the other specimens from extreme southern Oregon and northwestern California, by reason of the near (18) Apricot Yellow color of the underparts, but this same color occurs in specimens from the more northerly localities of Buchanan, Eugene, Vida Fish Hatchery, and McKenzie Bridge, as well as in no. 2178, Univ. Oregon, from Cresent Lake. The last mentioned specimen is here referred to nevadensis.
Two females referred to oregonensis from southern Oregon differ so greatly in size of skull that they challenge one's imagination in any attempt to provide an explanation for so wide a range of variation in one subspecies. One of these, no. 244520, U. S. Nat. Mus., is an adult female from Medford. The other, no. 224034, U. S. Nat. Mus., is a subadult female (though labeled male) from 43 miles northeast of Grants Pass. The skull of the adult from Medford has a basilar length of 41.5, upper tooth-rows, 16.1 in length, and a weight of 2.75 grams, whereas corresponding figures for the subadult are only 33.8, 12.9, and 1.4. Two other adult females are intermediate in size: No. 1413, Univ. Oregon, from 13 miles southwest of Grants Pass, Oregon, approaches the specimen from Medford in size, and the second specimen, no. 34325, Mus. Vert. Zoöl., from Carlotta, California, is smaller.
Not only is there a difference in length between the skulls of the two extremes of the females but this difference extends to all other dimensions of their skulls, and is most pronounced in the preorbital region. The differences in breadth of the braincase and other parts of the skull are relatively less than the differences in length. Differences of the same nature, although of lesser degree than found in the females, are to be seen in two males. The skull of an adult no. 51590, Mus. Vert. Zoöl., from 6 miles south of Medford, has a basilar length of 46.4, upper tooth-rows, 17.6 mm. long, and a weight of 4.0 grams, whereas corresponding figures for the subadult type specimen from Grants Pass, are only 43.0, 16.2, and 3.3.
The wide range of variation in size of skull of both sexes, together with the considerable variation in color pattern of the specimens here referred to oregonensis raises the suspicion that we are using the name in a composite sense; nevertheless, to recognize more than one subspecies with the material now available would be unwise.
A subadult female, of abnormal color, no. 47149, Mus. Vert. Zoöl., taken by Mr. H. E. Wilder at Carlotta, California, on December 20, 1930, in a region where weasels do not turn white in winter, is white, except for the black tip of the tail, but has a suffusion of orange. This specimen, discussed at greater length on page 43, is instructive in that it suggests that there are separate determiners for the brown and red elements of the pelage. It is interesting also as suggesting how natural selection may tend to eliminate from the population a conspicuous color-variation of this kind. At any rate, Mr. Wilder (Ms.) states: "This specimen was picked up in a field, where it evidently had been dropped by a hawk or an owl." The braincase of the skull is crushed in three places as though by a raptor's beak. None of the several other weasels, all normally colored, saved by Mr. Wilder from this general locality gives evidence of having fallen a victim to a raptor.
Only 2 skulls of the 12 adults and subadults examined show malformation of the frontal sinuses such as results from the presence of parasites.
Specimens examined.--Total number, 29, arranged within states from north to south by counties. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=California.= Del Norte County: Requa, 1[8]. Humholdt County: Eureka, 2 (1[74], 1[75]); Ferndale, 1[74]; Fortuna, 2[63]; Carlotta, 6 (3[74], 3[59]); 12 mi. E Bridgeville, 1[59]; 2 mi. W Bridgeville, 1[59].
=Oregon.= Washington County: Forest Grove, 1. Marion County: Salem, 2. Benton County: Buchanan, 1. Lane County: McKenzie Bridge, 1[101]; Vida Fish Hatchery, 1[101]; Eugene, 1[101]. Douglas County: Anchor, 1. Curry County: Gold Beach, 2[60]. Josephine County: Rogue River Valley (Grants Pass), 1; 13 mi. SW Grants Pass, 1[101]. Jackson County: Medford, 2; 6 mi. S Medford, 1[74].
=Mustela frenata munda= (Bangs)
Long-tailed Weasel
Plates 1, 19, 20, 21, 22, 23, 30, 34, 35, 36 and 40
Putorius xanthogenys mundus Bangs, Proc. New England Zoöl. Club, 1:56, June 9, 1899; Stephens, California mammals, p. 247, 1906.
Mustela frenata, Audubon and Bachman, Journ. Acad. Nat. Sci. Philadelphia, 8 (Pt. 2):291, 1842 (North California about 40° latitude).
Mustela xanthogenys munda, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.
Mustela frenata munda, Hall, Carnegie Instit. Washington Publ. 473:107, November 20, 1936.
Type.--Male, adult, skull, os penis and skin; no. 5459, collection of E. A. and O. Bangs, but now in collection of Mus. Comp. Zoöl.; Point Reyes, Marin County, California; June 19, 1896; obtained by C. A. Allen; original no. 931. (See comments under "Remarks," below, on places in California to which the name Point Reyes has been applied.)
The skull (pls. 19-21, 30) is complete and unbroken. I1 on each side and right I2 are broken away; p2 and p3 on each side have been aborted and the only alveoli remaining are two for the right p3. Otherwise all teeth are present and entire. The skin is fairly well made and in good condition.
Cranially, the type is a "runt"; its small size and the circumstance that the tympanic bulla is longer than the lower molar and premolar tooth-row and longer than the rostrum are features which differentiate the type from any other specimen seen of this race.
Range.--Sea level to at least 6,000 feet (South Yolla Bolly Mountain, Trinity County, California); Upper Sonoran and Transition life-zones of the coast and Coast Range of northwestern California from the Golden Gate northward into southern Humboldt and Trinity counties. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. oregonensis in longer hind foot of males which is more than 50 mm., and in both sexes, larger, more prominently ridged skull (see measurements and plates); from M. f. saturata by presence of nasofrontal white spot, larger and relatively shallower skull of males and larger skull of female; from M. f. nevadensis by presence of well-developed, white, facial markings; absence of color of underparts on ventral face of proximal third of tail; and hind foot of males more than 50; from M. f. xanthogenys by near (l) Sudan Brown to near (l) Antique Brown rather than Buckthorn Brown colors of upper parts and greater size, and in adult male basilar length more than 45 and hind foot more than 47; from M. f. nigriauris by having inside of ears same color as back rather than much darker than back.
Description.--Size.--Male: Three adults and two young from Point Arena and Gualala, Mendocino County, yield average and extreme measurements as follows: Total length, 447 (434-470); length of tail, 167 (150-185); length of hind foot, 53 (50-60). Corresponding measurements of three adults from 5 and 6 miles west of Inverness, Marin County, are: 430 (420-440), 154 (141-160), 48 (48-49). Corresponding measurements of four individuals (3 adults and 1 young of large size) from South Yolla Bolly Mountain, Trinity County, are: 383 (374-400), 134 (130-138); 44 (43-44). The tail averages 60 per cent as long as the head and body in the series from Point Arena, 56 per cent in the series from Point Reyes, and 53 per cent in the series from South Yolla Bolly Mountain. In every specimen except two, length of hind foot less than basal length. The two exceptions are no. 19720, M.V.Z., male adult from Point Arena in which the hind foot is recorded as 60 (probably an error in measurement), and no. 19721, M.V.Z., from the same place, in which the skull has not yet attained its full growth.
Female: One adult from Point Arena measures as follows: Total length, 383; length of tail, 134; length of hind foot, 43. Corresponding measurements of an adult from seven miles north of Laytonville, Mendocino County, are: 336, 121, 33 (= 36 on dried skin). Corresponding measurements of an adult from South Yolla Bolly Mountain, Trinity County, are, 326, 113, 37. In these three specimens, the tail is, in the order given, 54, 56, and 53 per cent as long as the head and body. Length of hind foot more than basal length.
Differences in external measurements of the two sexes as indicated by the five males and one female from Point Arena, are: Total length, 64; length of tail, 33; length of hind foot, 10. Weights of 2 adult males are 265 and 221 grams and of one adult female 155 grams.
Externals.--As described in Mustela frenata nigriauris.
Color.--Spot between eyes, narrow band or spot confluent with color of underparts on each side of head anterior to each ear, chin, lower lips, and rarely posterior third or less of each upper lip white; dark spot posterior to each angle of mouth uniformly present and of large size; tip of tail black; remainder of upper parts near (14 l) Sudan Brown and tone 4 of Raw Umber of Oberthür and Dauthenay, pl. 301; occasionally, slightly darker brown on forehead, nose, and about eyes. Underparts near (a to c) Ochraceous-Buff and sometimes Orange-Buff. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind limbs over antiplantar faces of toes. Least width of color of underparts averaging, in a series of 5 males from Mendocino County, 57 (46-67) per cent of greatest width of color of upper parts; 38 (35-40) in 3 males from Point Reyes, Marin County. Black tip of tail in Mendocino County series averaging 53 (46-60) mm., which is same length as hind foot and 32 per cent of length of tail. In Point Reyes males, black tip of tail averages 44 (34-52) mm., which is less than length of hind foot and 45 per cent as long as tail-vertebrae.
Several specimens of the smaller, inland variant (see under "Remarks") are near (l) Antique Brown rather than near (14 l) Sudan Brown above and hence do not differ in this respect from nigriauris.
Skull and teeth.--Male (based on 3 adults from Mendocino County): See measurements and plates 19-23, 30. As described in Mustela frenata nigriauris except that: Weight, 6.0 (5.4-6.3) grams; basilar length, 47.6 (46.5-48.2); length of tympanic bulla more than length of lower molar and premolar tooth-row.
Female (based on no. 19723, M.V.Z., from Point Arena): See measurements and plates 34-36, 40. As described in M. f. nigriauris except that: Weight, 3.0 grams; basilar length, 42.3.
The skull of the female is 50 per cent lighter than that of the average male.
Compared with the skull of the male of nevadensis that of munda averages larger in every part measured and specimens from Point Arena are nearly as heavy again, have relatively more expanded zygomata and mastoid processes but are relatively narrower anteriorly as shown by the breadth of the rostrum, interorbital breadth and postorbital breadth. Also the braincase is less inflated anteriorly, the tympanic bullae are lower and the skull is more angular. Females show the same differences although in different degree. Compared with the skull of the male of M. f. nigriauris, that of munda from Point Arena averages larger in every part measured except for the length of the upper tooth-rows. Relative to the basilar length, the skull of munda averages broader across the mastoids and across the zygomata, is deeper through the braincase at the anterior end of the basioccipital, and has a greater development of the lambdoidal crest.
Remarks.--The skin and part skull, no. 536/1849, U. S. Nat. Mus., taken by Lieutenant W. P. Trowbridge at San Pablo Bay, is the first specimen known to have been saved of this subspecies. Since 1899 when O. Bangs diagnosed munda as of small size, the weasel of the humid costal belt north of San Francisco Bay has been regarded as smaller than bridled weasels from farther south in the State. Actually, however, the weasel of the humid costal belt shares with M. f. pulchra the distinction of being one of the two largest weasels in California.
M. f. munda may be a composite subspecies, for the variation in facial markings, in coloration otherwise, in external measurements and in size and shape of skull is great. At one time in the course of the present study, manuscript accounts of two subspecies were prepared for the animals now all called munda and there is still much justification for recognizing two subspecies, one, along the coast proper, the larger, darker-colored animal with reduced white facial markings and large, wide, heavily ridged skull from Point Arena, and 6 miles south of Laytonville, Mendocino County, along with the specimens from 5 and 6 miles west of Inverness, Marin County, and the other, an inland race, which is a smaller, lighter-colored animal with more extensive white facial markings and a smaller, narrower, skull, known by specimens from Point Reyes [station?], Nicasio, 15 mi. north of San Rafael, Freestone, Vallejo, and Mount Sanhedrin. The differences between these two lots of specimens are of great degree. However, a female from Fort Bragg proves to be no larger than three females labeled as from Point Reyes. Also, a male from 2 miles south and one mile east of Stewarts Point on the coast has a skull no larger than the animal from Vallejo, whereas the skin alone of an adult female from 3 miles south of Stewarts Point is large and agrees with the specimens from Point Arena. Consequently, no logical ranges can be worked out for the two variants with the material now available.
Finally, the type specimen of munda is a "runt," smaller than any other male seen. This specimen, purchased by E. A. and O. Bangs from C. A. Allen, who collected and sold specimens widely, was labeled as from Point Reyes. So far as this place-name is concerned, it might refer to: (1) The point of land by that name which projects out into the Pacific Ocean, (2) an abandoned ranch house bearing that name at the head of Drakes Bay, 6 miles north and 3-3/4 miles east of the actual point, or (3) the railway station by the same name at the head of Tomales Bay, 12 miles east and 4-3/4 miles north of the actual point. Allen, himself, lived near San Geronimo (then Nicasio) about nine miles southeast of the Point Reyes railway station. All these places are in Marin County, but differ markedly as regards climate and flora. The first two are treeless, windswept and have much fog, whereas Point Reyes Station is more often sunny, and is situated in a shallow valley, inland, where the open grass-covered west-facing slopes meet the east-facing wooded ones. From which one of these three places the type specimen came, I do not know. The same may be said of the three female specimens labeled Point Reyes; two of these are in the United States National Museum and one in the Field Museum.
The specimens in the Museum of Vertebrate Zoölogy from 5 and 6 miles west of Inverness and those from near the same place in the collection of John Cushing come from within a couple of miles or less of the Point Reyes represented by the abandoned ranch house. These specimens, as remarked above, agree with those from Point Arena in large size, reduced facial markings and wide skull. These are points of difference from the smaller variant suspected of being a recognizable subspecies. It is the smaller variant which the type specimen approaches in size, and with which it agrees in relatively well-developed white facial markings. This suggests that the type specimen came from Point Reyes Station rather than from either of the two other places bearing the name "Point Reyes," from one of which, as just stated, the variant of large size is known. The three females labeled "Point Reyes" also have well-developed white facial markings and are of lesser size than the female of similar age from Point Arena, Mendocino County. The presumption is that these three females also came from Point Reyes Station.
The smaller, inland variant seems to agree in size, cranial characters, and coloration with M. f. nigriauris to the southward of San Francisco Bay, but lacks the black on the head which characterizes nigriauris. The larger variant, on which the description here used for munda is based, comprises animals which differ from nigriauris in larger size, darker color, reduced white facial markings, and larger, relatively wider skull. Both of the variants mentioned above are sharply distinct from nigriauris on the basis of coloration of the inside of the ear which is blackish in nigriauris like the dark facial markings, and in munda is colored like the back. M. f. munda lacks the dark facial markings; an occasional specimen has at most, a trace of the markings but this does not extend back so far as the ears. This difference, blackish versus non-blackish face, persists eastward of San Francisco Bay to at least as far as the Carquinez Straits, where a specimen of munda is available from 4 miles north of Vallejo and one of nigriauris from Glen Frazer Station on the south shore opposite Vallejo.
Intergradation with M. f. nevadensis and possibly with M. f. saturata is indicated by specimens from South Yolla Bolly Mountain, Trinity County. In them the external measurements and measurements of the skull are intermediate. Also the white frontal spot is much reduced in size. The white bars in front of the ears are absent in three specimens, and weakly developed in the other two. The relative proportions of the skulls as a whole are nearer those of nevadensis or saturata than munda. The skull of one of the three adult males and the skull of the adult female suggests M. f. oregonensis in certain features; for example, the dorsal outline of the skull in longitudinal axis is slightly convex as it is in oregonensis.
None of the specimens shows malformation of the frontal sinuses such as results from infestation by parasites.
Specimens examined.--Total number, 37, arranged by counties from north to south. Unless otherwise indicated specimens are in the Museum of Vertebrate Zoölogy.
=California.= Trinity County: S. Yolla Bolly Mt., 3[91]; 1/2 mi. S S. Yolla Bolly Mt., 1. Tehama County: 2 mi. S S. Yolla Bolly Mt., 1. Mendocino County: 6 mi. N Laytonville, 1; Mt. Sanhedrin, 1[87]; Ft. Bragg, 1; Gualala, 1; Point Arena, 5. Sonoma County: 2 mi. S and 1 mi. E Stewarts Point, 1; 3 mi. S Stewarts Point P. O., 1; Freestone, 1. Napa County: 6 mi. SSW, Napa, 1; 4 mi. N Vallejo, 1. County in question: San Pablo Bay, 1[91]. _Marin County_: 6 mi. W Inverness, 2; 5 mi. W Inverness, 2(1[28]); Point Reyes, 4 (2[91] 1[60], 1[75]); Nicasio, 2 (1[60], 1[75]); Kehoes Ranch, Pierce Point, 1[28]; Drakes Bay, 1[28]; Tomales Point, about 1/2 mi. SW White Gulch, 1; Point Reyes School, 3-3/4 mi. W Inverness, 1; 15 mi. (by road) N San Rafael, 1[52]; Hurley Ranch, 2 mi. W Tomales, 1. No locality more definite than California, 1[7].
=Mustela frenata xanthogenys= Gray
Long-tailed Weasel
Plates 21, 22, 23, 28, 30, 34, 35 and 36
Mustela xanthogenys Gray, Ann. and Mag. Nat. Hist., 11:118, 1843.
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Putorius xanthogenys, Merriam, N. Amer. Fauna, 11:25, June 30, 1896; Bangs, Proc. New England Zoöl. Club, 1:56, June 9, 1899.
Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.
Mustela frenata xanthogenys, Hall, Carnegie Instit. Washington Publ. 473:107, November 20, 1936.
Type.--Male, adult, skull and skin; skull no. 197a-43.6.4.55, skin no. 234a-42.11.21.4, British Museum (Nat. Hist.); from the bank of Sacramento River below mouth of Feather River, or from north shore of San Francisco Bay, California; taken in "1837 or 1838"; presented by Captain Edward Belcher.
The skull (plate 28) lacks the occiput, the right mandible posterior to m1, and the right pterygoid; the right zygomatic arch is fractured. The teeth are not greatly worn. The skin was originally mounted for exhibition (R. I. Pocock in Litt.) but in 1937 when I saw the skin, it was prepared as a conventional study skin. The skin is in fairly good condition; some hair is missing on the hind quarters and the skin of the tail is torn at one place.
Range.--Altitudinally, less than 600 feet (Fair Oaks); Lower Sonoran and Upper Sonoran life-zones of all but southern end of the San Joaquin Valley, and probably Sacramento Valley, California. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. nevadensis by presence of light facial markings and Buckthorn Brown rather than near (14_n_ to l) Brussels Brown color of upper parts; from M. f. munda by Buckthorn Brown rather than near (l) Sudan Brown, or near (l) Antique Brown color of upper parts and lesser size, in adult males basilar length less than 45 and hind foot less than 47; from M. f. nigriauris by lighter color in same way as from munda and also by having inside of ears same color as back rather than much darker than back; from M. f. pulchra in hind foot of males less than 46 and narrower skull, in males having breadth of rostrum less than 13.9 and mastoid breadth less than 26.0, see comparison of skulls in the account of pulchra.
Description.--Size.--Male: Three adults, from Fresno, Selma and Los Banos, measure, respectively as follows: Total length, 425, 417, 450; length of tail, 152, 154, 180; length of hind foot,--, 43, 44. Tail averages 61 per cent as long as head and body. Length of hind foot less than basal length.
Female: Adults from Selma, Los Banos, and 4 mi. SW Turlock, measure respectively as follows: Total length, 357, 365, 395; length of tail, 133, 132, 145; length of hind foot, 40, 38, 41. Tail averages 58 per cent as long as head and body. Length of hind foot less than basal length.
The average differences in external measurements between the two sexes, as represented by these six specimens, are: Total length, 65; length of tail, 25; length of hind foot, 3.5. One adult male weighs 274 grams and 2 adult females 182 and 214 grams.
Externals.--As described in Mustela frenata nigriauris.
Color.--Spot between eyes, band confluent with color of underparts on each side of head extending anterodorsally anterior to each ear, and posterior half to third of each upper lip white, or whitish tinged with some shade of yellowish; chin and lower lip white; dark spot posterior to each angle of mouth of varying size but uniformly present; tip of tail black; remainder of upper parts Buckthorn Brown of Ridgway or a trifle browner than tone 4 of Brown Pink of Oberthür and Dauthenay, pl. 297. Upper parts of uniform color except for slight darkening of head-markings anterior to ears. Underparts Ochraceous-Buff to Warm Buff. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind limbs over antiplantar faces of toes and sometimes tarsal region. Least width of color of underparts averaging, in 9 specimens from Fresno, Selma and Los Banos, 54 (32-74) per cent of greatest width of color of upper parts. Black tip of tail in three males (one subadult and 3 adults) averages 55 (50-60) mm. long. Thus longer than hind foot and averaging 34 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 2 adults from Fresno and one from Selma): See measurements and plates 21-23, 30. As described in M. f. nigriauris except that: Weight 3.8 grams; basilar length, 43.7 (43.4-43.9); least width of palate more or less than lateral length of P4; length of tympanic bulla more than length of lower molar and premolar tooth-rows.
Female (no. 2626 W. E. Snyder, from Selma): See measurements and plates 34-36. As described in M. f. nigriauris except that: Weight, 2.5 grams; basilar length, 39.4.
The skull of the female is 34 per cent lighter than the average for the three males.
Compared with skulls of nevadensis from the Sierra Nevada, those of the two adult males from Fresno differ as follows: M1 wider (transversely); tympanic bullae narrower; preorbital part of skull smaller. Comparison with pulchra is made in the account of that subspecies. Compared with skulls of adult males of nigriauris, from Santa Clara County, the two skulls from Fresno are generally smaller and in basilar length, length of tooth-rows and measurements of the teeth fall below the minimum for nigriauris. Relative proportions of the skulls are approximately the same. Comparison with munda reveals essentially the same differences as does comparison with nigriauris except that the difference in size is greater.
Remarks.--The name Mustela xanthogenys Gray was long applied to all the weasels of the interior valleys of California and of the coast of that state south of San Francisco Bay. Gray, when he named the species and when referring to it in later accounts, never defined the locality whence the specimen came more definitely than "California." In 1896, Merriam (1896:25) gave the type locality as "Southern California, probably vicinity of San Diego" and later writers have not contradicted him. The type specimen was obtained in the course of the voyage of the British ship Sulphur, under command of Sir Edward Belcher. Examination of Belcher's (1843, vol. 1, p. 129) narrative of the voyage indicates the following places in California at which the specimen of weasel, described by Gray, could have been obtained: Fort Ross, Bodega, vicinity of San Francisco Bay and up Sacramento River to the mouth of the Feather River, Monterey, Santa Barbara, Buenaventura, San Pedro, San Juan, and San Diego.
Reginald I. Pocock has kindly compared the type specimen in the British Museum with several specimens sent for that purpose. In the first place, comparison of skulls shows that the type specimen is a member of one of the races north of San Diego. In the second place, comparison of skins shows that the inside of the ears are not blackish but similar in color to the back. In fact, Pocock writes under date of February 12, 1929, regarding the type specimen, that "It is practically uniformly colored from the snout to the base of the tail, there being scarcely a trace of the darkening of the head, or muzzle, observable in your specimens [those sent for comparison]." This character of coloration of the ear excludes all the weasels of the Coast region of California from San Francisco Bay southward, namely, M. f. latirostra and M. f. nigriauris. My own examination of this type specimen at a date later than that on which Pocock compared it satisfies me as to the accuracy of his statement above.
Accordingly, the name xanthogenys would seem to apply to one of the two subspecies here called munda and xanthogenys. Perusal of Belcher's narrative of the voyage (loc. cit.) shows that little, if any, opportunity was afforded to obtain vertebrate specimens at Fort Ross or Bodega, both localities within the range of the subspecies here called munda. Furthermore, the type specimen is smaller than individuals of munda from 5 to 6 miles west of Inverness and from Point Arena with which the animals from Fort Ross and Bodega would be expected to agree in size. Weasels from along the north shore of San Francisco Bay are smaller than those on the coast north of the bay. Possibly the type specimen of xanthogenys came from the north side of San Francisco Bay but probably it came from the bank of the Sacramento River and almost certainly not farther up stream from San Francisco Bay than the junction of the Sacramento and Feather rivers. The statement of Belcher (1843, vol. 1, p. 129), regarding the trip up the Sacramento River as far as Point Victoria, lat. 38°46´47" north, and return to San Francisco Bay, that "Cuyote or jackal--fox, racoon, land otter, weasel, and squirrel were obtained" lends strong probability to the idea that this type specimen was taken along the Sacramento River, possibly in the vicinity of the existing city of Sacramento. Unfortunately no specimens are available from the Sacramento Valley. If some were available, a comparison of them and specimens of munda from along the north side of San Francisco Bay and Carquinez Straits with the type specimen of xanthogenys should determine the correct application of the name. For the present it seems best to retain the name munda and apply the name xanthogenys to the weasels inhabiting the northern part of the San Joaquin Valley and presumably the southern part of the Sacramento Valley.
Efforts to obtain specimens of weasels from the Sacramento Valley have been in vain. A juvenal specimen taken five miles south of Fair Oaks, Sacramento County, by Mr. John Fitzgerald, Jr., in December, 1927, was examined at his home and found to agree in coloration with specimens from farther south. Geographically, this specimen probably is more nearly a topotype than any other examined.
Most of the specimens examined are immature and adequate adult cranial material has not been seen. Two adults, one of each sex, from Los Banos have skulls of large size which agree with those of nigriauris. The same is true of one adult and one young female from 4 miles southwest of Turlock, which, unlike the animals from Los Banos, show a darkening of the head extending in reduced degree even to the inside of the ears, as in nigriauris. The slightly darker than average (for xanthogenys) color on the back may indicate intergradation with nevadensis. Intergradation with M. f. nevadensis is shown by specimens, from the southern part of the Sierra Nevada, mentioned in the account of nevadensis.
None of the skulls shows malformation of the frontal sinuses such as result from infestation by parasites.
Specimens examined.--Total number 30, arranged by counties from north to south.
=California.= Sacramento County: Bank of Sacramento River, 1[7]; 5 mi. S Fair Oaks, 1[29]. San Joaquin County: 4 mi. W Stockton, 1[74]. Merced County: Tegner School, 4 mi. SW Turlock, 2; Los Banos, 4 (2[74], 1[91] 1[87]). Fresno County: Mendota, 1[74]; Biola, 1[30]; Clovis, 1[55]; Fresno, 5 (1[74], 1[91], 2[55], 1[1]); 5 mi. W Fresno, 1[14]; Selma, 3 (2[50], 1[104]); 4 mi. NW Sanger, 1[55]; 5 mi. S Selma, 1[62]. Tulare County: Monson, 1[74]; 1-1/2 mi. N Goshen, 1[74]; Milo, 1[91]; 2 mi. N Tipton, 1[74]; Poplar, 2[53]. No locality more definite than California, 1[4].
=Mustela frenata nigriauris= Hall
Long-tailed Weasel
Plates 22, 23, 24, 34, 35, 36 and 41
Mustela frenata nigriauris Hall, Carnegie Instit. Washington Publ. 473:95, November 20, 1936.
Putorius xanthogenys, Baird, Mamm. N. Amer., 1858, p. 176 (part).
Mustela xanthogenys Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):375, 1874 (part?).
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Putorius xanthogenys xanthogenys, Grinnel, Proc. California Acad. Sciences, fourth series, 3:292, August 28, 1913.
Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912; Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933.
Type.--Male, adult, skeleton and skin; no. 32820, Mus. Vert. Zoöl.; Half Moon Bay, San Mateo County, California; received at Museum of Vertebrate Zoölogy, May 4, 1922, through A. L. Hagedoorn, after having been in captivity a few days where death occurred owing to injuries received in trap; original no. 1590.
The skull has each of the zygomatic arches and the anterior end of the nasals broken through. The only part missing is the central two millimeters of the left zygomatic arch. The teeth all are present and entire. The skeleton appears to be complete except for the bones of the feet, which are preserved within the skin. The skin is well made and in good condition.
Range.--Altitudinally, sea level to more than 4000 feet; Sonoran and Transition life-zones of Coast Range and coast of California from San Francisco Bay south to Point Conception, Santa Barbara County, California. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. munda, xanthogenys, and pulchra by having inside of ears darker than back rather than same color as back, and from xanthogenys and pulchra in near (l) Antique Brown color of upper parts rather than Buckthorn Brown or near (16 j) Buckthorn Brown to near (h) Yellow Ocher respectively; from M. f. latirostra by postorbital breadth, of adult males and females, less, rather than more, than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite.
Description.--Size.--Male: Five adults from Palo Alto, Santa Clara County, yield average and extreme measurements as follows: Total length, 447 (412-465); length of tail, 167 (147-175); length of hind foot, 46 (45-47). Corresponding measurements of four adults from San Francisco are: 412 (394-435); 153 (145-160); 43.5 (41-46). Corresponding measurements of five adults and subadults from Berkeley, Alameda County, are: 419 (390-448); 148 (135-160); 44 (42-47). Tail averages 59 per cent as long as head and body in series from Palo Alto and in one from San Francisco. The average of 55 for the Berkeley series probably reflects a lesser average age. Length of hind foot less than basal length. The type specimen measures, 415, 150, 43. It is smaller than the mean.
Female: A subadult from Palo Alto measures: Total length, 368; length of tail, 126; length of hind foot, 39. An adult and two subadults from Berkeley measure, respectively, as follows: Total length, 347, 365, 340; length of tail, 134, 123, 125; length of hind foot, 37, 38.4, 36.5. In these four females the tail averages 55 per cent as long as head and body. Length of hind foot less than basal length.
The average differences in external measurements of the two sexes, as represented by specimens from Berkeley, Alameda County, are: Total length, 68; length of tail, 21; length of hind foot, 7. Eight adult males weigh 249 (217-335) grams and one adult female 123 grams.
Externals.--Longest facial vibrissae brownish like dark color of head and extending beyond ear; carpal vibrissae mostly color of underparts and extending to apical pad of fifth digit; hairiness of foot-soles slightly more than shown in figure 20.
Color.--Spot between eyes, band, confluent with color of underparts, on each side of head extending anterodorsally anterior to ear, and posterior third of each upper lip tinged with color of underparts or, less often, pure white; chin and lower lips white; remainder of sides and top of head posteriorly to, or a little behind, a line connecting posterior margins of ears, blackish; inside of pinna of ear, and sometimes outside of pinna, blackish; dark spot posterior to each angle of mouth present on each side in three-fourths of specimens; tip of tail black; remainder of upper parts near (l) Antique Brown, and with more yellow than tone 3 of Raw Umber of Oberthür and Dauthenay, pl. 301. Often with more blackish and red in winter. Underparts near (a to c) Ochraceous-Buff or Ochraceous-Salmon. Ochraceous-Salmon in some juveniles. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, and on medial sides of hind limbs over antiplantar faces of toes. Least width of color of underparts averaging, in 17 adult males (Berkeley, 5; San Francisco, 5; Palo Alto, 7), 55 (40-73) per cent of greatest width of color of upper parts. Black tip of tail in same series of males averaging 51 (35-60) mm., thus averaging longer than hind foot and 33 per cent of length of tail (Palo Alto and San Francisco, 31 per cent; Berkeley, 35 per cent). In 8 adult females, least color of underparts amounts to 55 (47-62) per cent of greatest width of color of upper parts. Black tip of tail averages 41.5 (28-50) mm., thus averaging longer than hind foot and 32 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on six adults from Stanford Univ. and vicinity): See measurements and plates 22-24; weight (four adults), 5.4 (5.0-5.9) grams; basilar length, 47 (46.1-48.1); zygomatic breadth more than distance between condylar foramen and M1, or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more than postpalatal length; postorbital breadth less than length of upper premolars (less than distance between posterior borders of P4 and P2) and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than lateral length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 or 4 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more or less than (about equal to) length of lower molar and premolar tooth-row and longer or shorter (usually shorter) than rostrum; anterior margin of masseteric fossa below anterior half of m2.
Female (based on three adults, Hayward, Palo Alto, and Morro): See measurements and plates 34-36; weight (no. 43574, from Morro) 2.7 grams; basilar length, 41.2 (40.2-42.2); zygomatic breadth more or less than distance between condylar foramen and M1 and more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than lateral length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.
The skull of the female averages 50 per cent lighter than that of the average male.
Comparisons of the skull of the male with those of M. f. latirostra, pulchra, xanthogenys, and munda are made in the accounts of those subspecies.
Remarks.--Like M. f. latirostra, nigriauris long bore the name xanthogenys. The fairly adequate lot of specimens is divided between the collections of several institutions. The most satisfactory material in any one collection is in the Stanford University Natural History Museum where local specimens have been accumulated over a period of many years.
No actual intergrade between nigriauris and xanthogenys has been seen, although the specimens from Los Banos, referred to xanthogenys, have large skulls as in nigriauris. Intergradation with latirostra is shown by specimens, referred to latirostra, from the Los Angeles area. Also the one adult male from 5 miles southeast of Santa Margarita, San Luis Obispo County, is of small size and in this respect approaches latirostra. The range of nigriauris is separated from that of munda by San Francisco Bay, Carquinez Straits, and I suppose by the lower part of the San Joaquin River. On the basis of color of the inside of the pinna of the ear, the two subspecies are uniformly distinct. Intergradation is assumed to occur through the subspecies xanthogenys.
None of the 26 adult and subadult specimens examined for evidences of infestation of the frontal sinuses by parasites shows malformation of the sinuses.
Specimens examined.--Total number, 103, arranged by counties from north to south. Unless otherwise indicated specimens are in the Museum of Vertebrate Zoölogy.
=California.= Contra Costa County: Glen Frazer Station, 1; 2 mi. W Pinole, 1[13]; 1 mi. E Pinole, 1; Richmond, 1[13]; Lafayette, 1; 7 mi. E Clayton, 1; Moraga Valley, 1; Pinehurst, Redwood Canyon, 1; Concord, 1. Alameda County: Berkeley, 11; Oakland, 1; Piedmont, 1; Haywards, 2; near Haywards, 2; 10 mi. E Haywards, 1[91]; Redwood Canyon, 1; Calaveras Dam, 1. _San Francisco County_: San Francisco, 11 (5[8], 2[91], 1[60], 1[7]); Ocean View, 1[68]; Visitation Valley, 1. San Mateo County: Moss Beach, 1; Half Moon Bay, 1; Redwood City, 1[87]; Menlo Park, 9 (5[87], 2[68]); no locality more definite than county, 1[8]. _Santa Clara County_: 1/4 mi. N Milpitas, 1; 1/4 mi. S Milpitas, 1; Stanford University, 6 (4[68], 2[91]); Palo Alto, 11 (6[41], 2[60], 1[75], 1[87]). Santa Cruz County: 3 mi. E Santa Cruz, 1; 2-1/2 mi. E Santa Cruz, 1; Santa Cruz, 6 (2[91], 1[68], 1[4]). _Monterey County_: 1 mi. E mouth Salinas River, 10 ft., 1[37]; Pacific Grove, 1[8]; Monterey, 2 (1[7]); Carmel, 1[8]; Carmel Valley, 1[68]; Point Lobos, 1; Gonzales, 1. San Luis Obispo County: 5 mi. SE Santa Margarita, 1; Morro, 1[91]; 3-1/2 mi. S Oceano, 6. Santa Barbara County: Santa Maria, 1[87]; 5 mi. N Las Cruces, 1; 7 mi. W Gaviota, 1; Gaviota, 1.
=Mustela frenata latirostra= Hall
Long-tailed Weasel
Plates 1, 22, 23, 24, 34, 35 and 36
Mustela frenata latirostra Hall, Carnegie Instit. Washington Publ. 473:96, November 20, 1936.
Putorius xanthogenys, Baird, Mamm. N. Amer., p. 176, 1858 (part); Stephens, California mammals, p. 246, 1906; Merriam, N. Amer. Fauna, 11:25, June 30, 1896 (part).
Putorius (Gale) brasilianus frenatus, Coues, Fur-bearing animals, p. 142 (part).
Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912; Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933.
Mustela arizonensis, Grinnell and Swarth, Univ. California Publ. Zoöl. 10, 376, October 31, 1913.
Type.--Male, adult, skull and skin; no. 3257, Mus. Vert. Zoöl.; San Diego, San Diego County, California; May 20, 1907; obtained by Frank X. Holzner.
Right M1 is missing and the part of the jaw bearing this tooth is broken away. With this exception the skull is complete and unbroken and the teeth are all present and entire. The skin is fairly well made and in good condition except that it is slightly soiled.
Range.--Altitudinally sea level to 8000 feet (Tahquitz Valley, San Jacinto Mountains); Sonoran and Transition life-zones of coast and mountains west of Mohave and Imperial deserts of southern California from Point Conception and Cuyama Valley southward at least to Mexican boundary. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. nigriauris by having postorbital breadth of adult males and females, more, rather than less, than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; from M. f. pulchra by having tympanic bulla longer than rostrum (orbitonasal length) and by near (l) Antique Brown rather than near (16 j) Buckthorn Brown to near (h) Yellow Ocher color of upper parts.
Description.--Size.--Male: Six adults and subadults from San Diego yield average and extreme measurements as follows: Total length, 439 (428-449); length of tail, 153 (142-160); length of hind foot, 45 (40-47). Corresponding measurements for a series of eight adult males from the vicinity of Los Angeles are: 416 (394-428); 158 (151-166); 44 (40-47). In the series from San Diego the tail averages 54 per cent as long as head and body. In the series from Los Angeles the average is 61 per cent. Length of hind foot in each series, less than basal length. The type specimen measures, 435, 142, 42.
Female: No. 5070, adult, from San Diego, measures 367, 141, 38. Nos. 22 and 6748 from Santa Ysabel, measure: 359, 380; 130, 140; 39, 35. No. 7194 from Jamacha measures, 358, 125, 35. Three adult females from Los Angeles yield the following: Total length, 373, 345, 368; length of tail, 150, 122, 134; length of hind foot,--, 41, 41. In no. 5070 the tail is 62 per cent as long as the head and body and in the three from Los Angeles it averages 60 (55-67) per cent. Length of hind foot, in each case, less than basal length.
The average differences in external measurements of the two sexes as shown by the six males from San Diego and the four females from San Diego County are: Total length, 73; length of tail, 19; length of hind foot, 8. Corresponding differences shown by the eight males and three females from Los Angeles are: 54, 23, 3.
Externals.--Longest facial vibrissae brownish, like dark color of head and extending beyond ear; carpal vibrissae mostly color of underparts and extending to apical pad of fifth digit; hairiness of foot-soles slightly more than shown in figure 20.
Color.--Spot between eyes, band confluent with color of underparts on each side of head extending anterodorsally anterior to ear, and posterior third of each upper lip tinged with color of underparts or, less often, white; chin and lower lips white; remainder of sides and top of head posteriorly to near line connecting posterior margins of ears, blackish; inside of pinna of ear, and sometimes outside of pinna, blackish; dark spot posterior to each angle of mouth present on each side in three-fourths of specimens; tip of tail black; remainder of upper parts near (l) Antique Brown, and with more yellow than tone 3 of Raw Umber of Oberthür and Dauthenay, pl. 301. Underparts Ochraceous-Buff to Warm Buff and in some specimens from Los Angeles and Ventura counties Ochraceous-Orange, especially in young and juveniles. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists and on medial sides of hind limbs over antiplantar faces of toes. Least width of color of underparts averaging, in 15 adult and subadult males from San Diego County, 54 (35-75) per cent of greatest width of color of upper parts. Black tip of tail in same series of males averaging 54.5 (46-60) mm. long. Thus averaging longer than hind foot and 35 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 6 adults from San Diego County). See measurements and plates 22-24. As described in M. f. nigriauris except that: Weight (4 specimens), 3.9 (3.8-4.0) grams; basilar length 43.8 (41.9-47.0); postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2 to 2-1/2 (including I3) upper incisors; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum; anterior margin of masseteric fossa below m2.
Female (based on 4 adults from San Diego County): See measurements. As described in M. f. nigriauris except that: Weight, 2.6 (2.2-2.8) grams; basilar length, 40.0 and 40.1; postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; length of tympanic bulla more than length of rostrum.
The skull of the female averages 34 per cent lighter than that of the average male.
The skull of the male of latirostra, compared with that of nigriauris, is by weight, more than one-fourth lighter, has a lesser basilar length, a lesser mastoid breadth, a lesser zygomatic breadth and a narrower M1. In these features no overlap has been observed between adults from the general vicinities of the type localities of the two forms. In adult males of latirostra the postorbital breadth, with one exception, is more than the combined length of P4 and P3 whereas the reverse is true in adult males of nigriauris. Both males and females of latirostra have a generally smaller skull with relatively broader interorbital and postorbital parts and the tympanic bullae are relatively larger, rounder and more inflated.
Compared with the skull of the male of pulchra that of latirostra is, by weight, more than one-fourth lighter, has a lesser basilar and orbitonasal length, lesser zygomatic and mastoid breadth and a more nearly flat braincase. In these features no overlap has been observed between adults from the general vicinities of the type localities of the two subspecies. Also, in latirostra the tympanic bulla is longer than the rostrum whereas the opposite is true in pulchra. The skull of latirostra is generally smaller and relatively, on the average, has the preorbital part of the skull deeper and broader with longer tooth-rows, although with shorter rostrum, while the zygomatic and mastoid breadths are less. Study of skulls of subadult females of pulchra indicate that females of latirostra and pulchra differ in the same fashion as do males.
Remarks.--This subspecies long has gone by the name M. xanthogenys and the type locality was generally supposed to be in the vicinity of San Diego. This supposition seems to have originated with Merriam's (1896:25) statement that the type locality was "Southern California, probably vicinity of San Diego." Nevertheless, as set forth in the account of M. f. xanthogenys the type specimen concerned now is thought to have come from much farther north.
Although 76 Recent specimens are available from southern California, additional adults are needed to understand the geographic variation there. M. f. latirostra may be a composite--made up of more than one geographic race. Specimens from San Diego County differ so much in relative length of the tail that at one stage in the present study it was thought that a difference in this respect existed between the coastal animals and those from farther inland. Material received later did not wholly substantiate this view and because of the uniformly small size of all of the skulls from that county, the animals were later regarded as of the same subspecies. Eventually, even this supposed common feature proved to be inconstant for an adult male from Jamacha, no. 7098, of the San Diego Society of Natural History, and another adult male from San Marcos, no. 8869, collection of Ralph Ellis, were later examined and found to have skulls as large as those of average-sized, adult males of nigriauris.
Despite these puzzling local variations, it is established that the long-tailed weasels of southern California are smaller than those from farther north. Also, the southern animal averages smaller in weight and size of skull, and the skull is differently proportioned. Specimens in series from Los Angeles County definitely are intermediate in size and shape of skull between latirostra from San Diego County and nigriauris from, say, Santa Clara County, but definitely more closely resemble latirostra from San Diego County than they do nigriauris. A skull of a young animal, not here identified to subspecies, from Potholes, in the Colorado River Valley, 10 miles northeast of Bard, Imperial County, California, may have closest relationship to M. f. latirostra. Additional comment on this specimen is offered in the account of M. f. neomexicana.
From the asphalt pits of Rancho La Brea, in Los Angeles County, a total of 57 skulls have been examined, more than half of which are reasonably complete. I have been unable to learn whether these came from pits regarded by students of the deposit as wholly Recent, from pits regarded as of Pleistocene age, or from both. Suffice to say that only two specimens were found which could be distinguished from skulls of the subspecies of weasel living in that area today.
These two specimens, lent to me by Professor Chester Stock, were with other skulls received from the Los Angeles Museum of History, Art and Science and bore identifying numbers as follows: 16/20-27, the anterior part of the skull of an adult, and 16, the skull posterior to the cribiform plate of a subadult or possibly young individual. The latter has a mastoid breadth of 28.0 millimeters, a tympanic bulla 16.1 long and other measurements in proportion. It is larger than any specimen of weasel, of any subspecies, seen from California and in the subgenus Mustela seems to be exceeded in size only by certain individuals of M. f. texensis. M. f. neomexicana attains relatively large size and comparisons were made with individuals of that subspecies. However, the young specimen from Rancho La Brea differs from neomexicana in that the tympanic bullae rise less steeply on the medial sides and the inferior lip of the external auditory meatus is less developed laterally. Age considered, the sagittal crest is less developed and the mastoid processes project more abruptly from the skull. The anterior part of the skull of the adult, no. 16/20-27 is larger than any specimen seen of M. f. latirostra or adjoining subspecies, and among California-taken specimens is equaled in size only by the largest males of M. f. munda from the northwest coastal district in Mendocino County. This adult from Rancho La Brea differs from neomexicana, sex and age taken into account, in greater postorbital breadth, lesser rostral width in comparison with the interorbital breadth, and in having the temporal ridges at the anterior end of the sagittal crest spread out into a Y-shaped, rather than a T-shaped, pattern. All these differences from neomexicana are features of agreement with the California bridled weasels of the subspecies latirostra, nigriauris, and munda. The same is true of the characters which set apart the young specimen from neomexicana. In summary: of 57 weasel skulls examined from the asphalt pits at Rancho La Brea, Los Angeles County, all but two are indistinguishable from the skulls of the Recent weasel living in that region today. These two skulls agree in qualitative characters with animals of the California coastal subspecies now living from Los Angeles northward to Humboldt County, but are larger. For the time being these two may be thought of as giants of the same type of animal inhabiting the Los Angeles region today.
Only one of 41 adult and subadult skulls examined for malformation of the frontal sinuses shows infestation by parasites.
Specimens examined.--Total number, 142, listed by counties from north to south. Unless otherwise indicated specimens are in the Museum of Vertebrate Zoölogy.
=California.= Santa Barbara County: Rincon Point, 1. _Ventura County_: Cuyama Valley, 2200 ft., 1[91]; Nordhoff, 3[59]; Santa Paula, 1[59]; Ventura, 7. Los Angeles County: near Owensmouth, 1[24]; Cahuenga, 1[91]; Llano, 10 mi. E Littlerock, 1; Flint Ridge, Pasadena, 1[59]; Pasadena, 3; Lankershim, 1[24]; 1 mi. S Lankershim, 1[24]; Duarte, 1[59]; Covina, 1[59]; Claremont, 1[91]; El Monte, 4 (2[75], 1[24]); Montebello, 1; Alhambra, 6 (5[2], 1[91]); El Nogal, 2[8]; Gardena, 1[26]; Palos Verdes Estate, 3; Rancho La Brea asphalt deposits, 57[70]^{ and }[92]. _San Bernardino County_: San Bernardino Valley, 1[75]; San Bernardino, 4 (2[20], 1[91]); Redlands, 2 (1[38]); Bluff Lake, 2 (1[59], 1[33]). Riverside County: West Riverside, 1; Arlington, 800 ft., 1[17]; 3-1/2 mi. E and 1/2 mi. N Beaumont, 2600 ft., 1; Banning, 1[91]; Cabazon, 1[91]; San Jacinto Plain, 1[20]; Tahquitz Valley, 8000 ft., 1; Elsinore, 1[1]. San Diego County: Twin Oaks, 1[91]; San Marcos, 2 (1[87], 1[41]); Escondido, 1; Witch Creek, 1[91]; Ballena, 1[20]; Santa Ysabel, 3 (2[20], 1[87]); Julián, 1; La Jolla, 1; Lakeside, 1[91]; El Cajon, 1[91]; El Vido (not found on map), 1[91]; San Diego, 9 (1[91], 1[20], 1[87], 1[32]); Jamacha, 2[87]; Chula Vista, 1[20].
=Mustela frenata pulchra= Hall
Long-tailed Weasel
Plates 22, 23 and 24
Mustela frenata pulchra Hall, Carnegie Instit. Washington Publ. 473:98, November 20, 1936.
Type.--Male, adult, skeleton and skin; no. 16668, Mus. Vert. Zoöl.; Buttonwillow, Kern County, California; April 30, 1912; obtained by J. Grinnell; original no. 1953.
The skull (plates 22-24) is complete and unbroken (a fracture in the right jugal has healed). All teeth are present and entire. The skeleton lacks the os penis, left fibula, shaft of left tibia and the distal three or four caudal vertebrae. Some of the bones of the feet distal to the radius and tibia are with the skeleton, and the remainder probably are in the skin. The skin is fairly well made and in good condition, except for the left hind leg which was torn when the animal was captured. A well-developed scrotal pouch shows the specimen to have been a male.
Range.--Altitudinally around 300 feet in San Joaquin Valley to 2500 feet at Isabella; Upper Sonoran and Lower Sonoran life-zones of southern end of San Joaquin Valley and in mountains at southern end of Valley, California. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. nevadensis in presence of light facial markings, and from M. f. nevadensis and M. f. inyoensis in near (16 j) Buckthorn Brown to near (h) Yellow Ocher rather than near (14 n to l) Brussels Brown color of upper parts, and greater size with hind foot more than 40 in females and basilar length averaging more than 46.0 in males; from M. f. latirostra in having rostrum (orbitonasal length) longer than tympanic bulla and from M. f. latirostra and M. f. nigriauris by color of upper parts as stated above rather than near (l) Antique Brown, and by having inside of ears same color as back rather than much darker than back; from M. f. xanthogenys in hind foot of males more than 46 and broader skull which in males has breadth of rostrum more than 13.9 and mastoid breadth more than 26.0.
Description.--Size.--Male: The type specimen and five other adults yield average and extreme measurements as follows: Total length, 454 (428-477); length of tail, 178 (153-184); length of hind foot, 50 (47-55). Tail averages 65 per cent as long as head and body. Length of hind foot approximately equal to basal length. The type specimen measures, 460, 184, 49.
Female: Three subadult topotypes yield average and extreme measurements as follows: Total length, 399 (383-411); length of tail, 154 (140-161); length of hind foot, 42 (41-42). Tail averages 63 per cent as long as head and body. Length of hind foot less than basal length.
The average differences in external measurements of the two sexes are: Total length, 55; length of tail, 24; length of hind foot, 8.
Externals.--As described in Mustela frenata nigriauris.
Color.--Spot between eyes, band confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, posterior third of each upper lip, lower lips and chin white or more often darker than Ochraceous-Buff and therefore same color as belly; dark spot posterior to each angle of mouth present but small; tip of tail black; remainder of upper parts near (16 j) Buckthorn Brown to near (h) Yellow Ocher and from tone 2 to 4 of Brown Pink of Oberthür and Dauthenay, pl. 297, but with a trifle more reddish brown. Upper parts of uniform color except for occasional slight darkening of nose, forehead, and areas around eyes. Underparts darker (a) than Ochraceous-Buff. Color of underparts extends distally on posterior sides of forelegs over toes, onto antipalmar faces of feet and wrists, on medial sides of hind limbs over antiplantar faces of toes, tarsal region and sometimes in diluted fashion on proximal third of underside of tail. Least width of color of underparts averaging, in 6 male topotypes, 55 (43-81) per cent of greatest width of color of upper parts. Black tip of tail in same series of males averaging 58 (53-63) mm. long; thus averaging longer than hind foot and 33 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 6 ads., type and 5 topotypes): See measurements and plates 22-24. As described in M. f. nigriauris except that: Weight (6 ads.), 5.3 (4.5-6.1) grams; basilar length, 47.6 (46.0-48.6); (one skull, no. 335, with postorbital breadth more than distance between posterior borders of P4 and P2); interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2 to 3-1/2 (including I3) upper incisors; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than rostrum.
Female: Adult skull of typical female not seen.
As compared with the skull of the type specimen of inyoensis, skulls of adult males of pulchra are larger throughout, relatively broader, especially in the preorbital part of the skull, have more inflated tympanic bullae, and are less convex in dorsal outline. Comparison of the skull with that of latirostra has been made in discussion of that subspecies. Comparison of skulls of adult males of nigriauris and pulchra shows that those of pulchra average larger in every measurement taken except those of m1, M1, P4, and depth of skull at posterior borders of upper molars. The basilar length is only slightly more and it follows that, relative to this length, other measurements of the skull are relatively, as well as actually, larger. In no one measurement is there an entire lack of overlap, but the skulls of adult males, and probably adult females, may be distinguished from those of nigriauris by the combination of the following mentioned, average differences: Tympanic bullae larger in each of three dimensions; preorbital and interorbital parts of skull broader and notably heavier; interorbital breadth greater; zygomatic arches more expanded laterally; mastoid processes more prominent. As compared with xanthogenys, differences of similar nature, but of greater degree, distinguish pulchra. As compared with those of nevadensis (represented by specimens from Mono Co., Calif.), skulls of adult males of pulchra average larger in every measurement taken and no overlap exists in basilar length, orbitonasal length, mastoid breadth, zygomatic breadth, length of tympanic bulla, or depth of skull at either the anterior margin of the basioccipital or at the posterior margins of the upper molars. Relatively, the preorbital portion is about the same size in the two forms.
Remarks.--The best material of this big weasel was obtained in 1910 and 1911 by John Wimmer and forwarded to the California Academy of Sciences through John R. Rowley, although in 1905, one specimen had been obtained by A. S. Bunnell for the collections of the United States Bureau of Biological Survey, another by J. Grinnell for the Museum of Vertebrate Zoölogy in 1912, and in 1933, another by L. M. Huey, for the San Diego Society of Natural History.
The males from the type locality are relatively uniform in size and shape of skull. The one exception is no. 137935, U. S. Nat. Mus., slightly younger than the others. Its skull is relatively more slender than any of the others and does not display several of the differential characters. The male, no. 127566, U. S. Nat. Mus., from Alila (= Earlimart) is intermediate in cranial features between pulchra and xanthogenys as known from specimens taken in the vicinity of Fresno. The skull of the female, no. 127565, from the same place, is too young to provide diagnostic characters. Since the skull of an adult female of topotypical pulchra is unknown, doubt attaches to the identification of the adult, female specimen, no. 115895, U. S. Nat. Mus., from Delano. It has a relatively broad skull in comparison with the adult female of xanthogenys from Los Banos. The adult female, no. 9998, San Diego Soc. Nat. Hist., from 2 mi. SW Simmler, shows approach to nigriauris in slightly reduced size. The skin alone from Coalinga, a male, taken on April 10, 1935, measures 462, 179, 47. The adult female, with crushed skull, from 4 miles east of Coalinga, measures 350, 129, 40. In size, these specimens agree better with pulchra than with xanthogenys. The skin alone from 3 miles south of Coalinga is unsexed and without external measurements. Skulls of adults from Coalinga are needed to permit of more positive identification of the subspecies found there. The female from 4 miles east of Coalinga, taken on February 21, 1936, is in process of molt on the underparts, and the longer hair which is near (20´) Naples Yellow contrasts strongly with the incoming shorter hair which is near (10 c) Salmon-Orange. The skin alone, no. 16270, Mus. Vert. Zoöl., from Isabella, was made up from a decayed animal and is of but little use. It is referred to pulchra purely because of geographic nearness of Isabella to the type locality of pulchra. The most that can be told from the specimen is that it is a relatively light-colored, bridled weasel. The fact that the color is slightly darker than in pulchra may or may not indicate intergradation with nevadensis. No. 54103/41042, U. S. Nat. Mus., consisting of crushed bits of skull and the skin of the head, is from Willow Spring, Kern County. This marginal locality is really in the Mojave Desert rather than in the San Joaquin Valley. The light color of the skin of the head suggests pulchra, but it is realized that a complete specimen might show the animal there to be unlike pulchra.
None of the skulls shows evidence of having had the frontal sinuses infested by parasites.
Specimens examined.--Total number, 18, listed by counties from north to south. Unless otherwise indicated, specimens are in the Museum of Vertebrate Zoölogy.
=California.= Fresno County: Coalinga, 1[23]; 4 mi. E Coalinga, 1; 3 mi. S Coalinga, 1[8]. Tulare County: Alila (= Earlimart), 2[91]. Kern County: Delano, 1[91]; Buttonwillow, 9 (6[8], 2[91]); Isabella, 1; Willow Spring, 1[91] _San Luis Obispo County_: 2 mi. SW Simmler, 1[87].
=Mustela frenata inyoensis= Hall
Long-tailed Weasel
Plates 22, 23 and 24
Mustela frenata inyoensis Hall, Carnegie Instit. Washington Publ. 473:99, November 20, 1936.
Putorius xanthogenys, Merriam, N. Amer. Fauna, 11:25, June 30, 1896 (part).
Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.
Type.--Male, adult, skull (with skeleton) and skin; no. 25907, Mus. Vert. Zoöl.; Carl Walter's Ranch, 2 mi. N Independence, Inyo County, California; June 26, 1917; obtained by A. C. Shelton; original no. 3143.
The skull (plates 22-24) is complete and unbroken. All teeth are present and entire. The skin is well made and in good condition.
Range.--From 3700 feet (Lone Pine) to at least 4000 feet (Alvord); Lower Sonoran Life-zone of the floor of Owens Valley in Inyo County, California. See figures 29 and 30 on pages 221 and 314.
Characters for ready recognition.--Differs from M. f. nevadensis in presence of white facial markings; from M. f. pulchra in near (l) Brussels Brown rather than near (16 j) Buckthorn Brown to near (h) Yellow Ocher color of upper parts and basilar length of less than 45 in males; from M. f. latirostra in brownish rather than blackish color of inside of ear and orbitonasal length of more than 15.
Description.--Size.--Male: Two adults, the type specimen and no. 25392/32805, measure, respectively, as follows: Total length, 423 and 390; length of tail, 170 and 145; length of hind foot, 42 and 44. Tail is 67 and 59 per cent as long as head and body. Length of hind foot less than basal length.
Female: No. 12400, Field Mus. Nat. Hist., which is young, has the following measurements: Total length, 390; length of tail, 150; length of hind foot, 39. Tail is 63 per cent as long as head and body. Length of hind foot less than basal length.
The differences in external measurements between the two sexes, as represented by the male type specimen and by the young female, are: Total length, 33, length of tail, 20; length of hind foot, 3.
Externals.--Longest facial vibrissae black or dark brown and reaching beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) slightly less than shown in figure 19.
Color.--Large spot between eyes, band confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, upper throat, chin, lower lips and in some specimens part or all of upper lips white; patch between eyes and bars in front of ears tinged with some shade of yellowish in one specimen; dark spot posterior to each angle of mouth present in four of five specimens; tip of tail black; remainder of upper parts, in summer, near (l) Brussels Brown or tones 1 to 2 of Raw Umber of Oberthür and Dauthenay, pl. 301; slightly darker brown on forehead, nose and about eyes. In winter near (j) Snuff Brown or lighter than Brussels Brown with a smoked effect. Underparts Buff-Yellow, winter and summer. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists and on medial sides of hind legs over antiplantar faces of toes. Least width of color of underparts averaging, in 5 available specimens 34 (24-42) per cent of greatest width of color of upper parts. Black tip of tail, in two adult males, averaging 53 (45 and 60) mm. Thus longer than hind foot and averaging 34 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on the type): See measurements and plates 22-24. As described in M. f. nigriauris except that: Weight, 4.4 grams; basilar length, 44.7; postorbital breadth not less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; length of tympanic bulla less than length of lower molar and premolar tooth-row.
Female: Adult unknown.
Compared with the skull of the male of nevadensis, no single difference not covered by individual variation in nevadensis has been detected. Selected differences of inyoensis in comparison with latirostra are larger size, less inflated tympanic bullae and relative narrowness of the postorbital, interorbital and preorbital parts of the skull. Comparison of the skull with that of M. f. pulchra is made in the account of that subspecies.
Remarks.--Although two specimens of this subspecies were taken during the Death Valley Survey conducted by Dr. C. Hart Merriam, only three additional individuals are known to have been saved as study specimens since that time.
M. f. inyoensis as now known may be thought of as closely similar to M. f. nevadensis except for the presence of well-developed white facial markings like those found in the weasels of the San Joaquin Valley and coastal region of California south of San Francisco Bay. The nonwhite areas of the head are almost the same color as the back and not distinctly blackish as in M. f. latirostra and M. f. nigriauris. The one specimen in the winter coat, no. 25392/32805, U. S. Nat. Mus., from Lone Pine, is brown rather than white. The brown has the pale smoke-tinge common in the winter pelage of subspecies whose members are either brown or white in winter. The range of this subspecies is thought to include the floor and lower elevations of Owens Valley although it may occur in limited numbers southwestward along the base of the Sierra Nevada and through the mountains in places of low elevation like Walker Pass its range may meet that of pulchra.
The type specimen was taken in an alfalfa field by ranch hands, who, according to A. C. Shelton (MS), stated that the species was common at the type locality. None of the five specimens shows infestation of the frontal sinuses by parasites.
Specimens examined.--Total number, 5, listed by localities from north to south.
=California.= Inyo County: Alvord, 4000 ft., 1 (U. S. Nat. Mus.); 2 mi. N Independence, 1 (Mus. Vert. Zoöl.); Lone Pine, 3 (2 in Field Mus. Nat. Hist. and 1 in U. S. Nat. Mus.).
=Mustela frenata neomexicana= (Barber and Cockerell)
Long-tailed Weasel
Plates 1, 22, 23, 24, 34, 35 and 36
Putorius frenatus neomexicanus Barber and Cockerell, Proc. Acad. Nat. Sci. Philadelphia, 1898:188; Lantz, Trans. Kansas Acad. Sci., 19:178, 1905.
Mustela frenata neomexicana, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Bailey, Animal Life of Carlsbad Cavern, p. 97, 1928; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.
Mustela frenatus neomexicanus, Bailey, N. Amer. Fauna, 35:19, September 5, 1913.
Type.--Male, adult, skull and skin; no. 10475, Mus. Comp. Zoöl.; Armstrong's Lake, Mesilla Park, Dona Ana County, New Mexico; February 1, 1898; obtained by A. C. Tryson; original no. 58 of C. M. Barber.
The skull is imperfectly cleaned but unbroken. The right upper incisors, right P2 and left p3 are broken away. The skin is indifferently stuffed but in a good state of preservation except that the distal part of the tail is missing. The animal's coat is ragged, and this imperfect appearance is heightened by injury to the posterior part of the body, probably at the time of capture.
Range.--From 3800 feet (type locality) to 9000 feet (Cloudcroft, N. Mex.); Upper Sonoran and Lower Sonoran life-zones of northern México, southeastern Arizona, New Mexico and western Texas, panhandle of Oklahoma, southeastern Colorado and southwestern Kansas. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. frenata and M. f. texensis by Buckthorn Brown rather than Brussels Brown color of upper parts, mastoid breadth of adult males ordinarily more, rather than less, than postpalatal length; from M. f. leucoparia by Buckthorn Brown rather than Argus Brown color of upper parts, distance from anterior margin of tympanic bulla to foramen ovale less, rather than more, than four-fifths height of tympanic bulla; from M. f. arizonensis and M. f. nevadensis by Buckthorn Brown, rather than near (14 n) Brussels Brown or, in winter, white color of upper parts, in presence of white frontal spot continuous with color of underparts, in basilar length of more than 46 mm. in males and 40 mm. in females; from M. f. longicauda by Buckthorn Brown rather than near (h) Clay Color of upper parts, by presence of white facial markings on Argus Brown head, and by length of tooth-rows amounting to less than 37 per cent of basilar length; from M. f. primulina by Buckthorn Brown rather than Brussels Brown color of upper parts, in presence of white frontal spot and broad white bands on side of head, in anteriorly truncate rather than anterolaterally rounded bullae and zygomatic breadth of more than 30 in males and 24 in females.
Description.--Size.--Male: The type specimen (see Barber and Cockerell, 1898:188) measured: Total length, 500; length of tail, 205; length of hind foot, 50. Tail 70 per cent as long as head and body. Length of hind foot less than basal length.
Female: No. 21779 from Tombstone, Arizona, measured: Total length, 419; length of tail, 165; length of hind foot from dried skin, 41 (probably 43 in flesh). Tail 65 per cent as long as head and body. Length of hind foot less than basal length.
The average differences in external measurements of the two sexes, as known from these two individuals, are: Total length, 81; length of tail, 40; length of hind foot, 7.
Compared with M. f. frenata, the size, proportions of parts and difference in size of the two sexes, appears to be about the same.
Externals.--Longest facial vibrissae colored like upper parts [in the type specimen some of the "long bristles of the upper lip" are white as pointed out by Barber and Cockerell (op. cit.: 188)] and extending beyond ear; carpal vibrissae colored like underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 20.
Color.--Broad white bands on sides of head, extending anterodorsally anterior to each ear, confluent with white spot between eyes and with color of underparts; posterior half or all of each upper lip edged with white; usually few white hairs on top of head between ears; remainder of top of head near Argus Brown of Ridgway and Chocolate, tone 4, of Oberthür and Dauthenay; dark spot posterior to each angle of mouth usually absent; tip of tail black; remainder of upper parts varying, in different specimens, from Buckthorn Brown to Dresden Brown of Ridgway, and Brown Pink (tones 3 to 4, pl. 297, of Oberthür and Dauthenay); underparts Antimony Yellow or near (c) Warm Buff of Ridgway, and Brown Pink (tone 1, pl. 297, of Oberthür and Dauthenay); color of underparts extends distally on legs over forefeet and hind feet. Least width of color of underparts averaging 46 (41-55) per cent of greatest width of color of upper parts; black tip of tail 35 to 45 mm. long in females; 43 to 68 mm. long in males and averaging 21 (20-36) per cent as long as tail-vertebrae.
No specimen of this subspecies in the white winter coat has been seen. Animals taken in midwinter are available from Mesilla Park, Willcox, and 10 miles east of Roswell.
From M. f. frenata, neomexicana differs in: upper parts and underparts much lighter colored; white facial markings more extensive; color of underparts more extended onto feet. From M. f. leucoparia, neomexicana differs as follows: above and below, much lighter colored, but white facial markings less extensive and color of underparts less extended onto feet and legs.
Skull and teeth.--Male (based on adults: the type; no. 131582 from Berino, New Mexico; and no. 1485 from Seward Co., Kansas): See measurements and plates 22-24. As described in Mustela frenata frenata except that: Weight, 6.2 (4.9 and 7.5); basilar length, 49.3 (48 and 50.5); mastoid breadth more than postpalatal length; least width of palate less than length of P4; anterior margin of masseteric fossa directly below m2 or heel of ml.
Female (based on three adults): See measurements and plates 34-36. As described in Mustela frenata frenata except that: Weight, 3.1 (2.6-3.5) grams; basilar length, 42.7 (40.8-45.5); zygomatic breadth less than distance between condylar foramen and M1 and more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla.
The skull of the female averages 50 per cent lighter than that of the male.
As compared with the skull of the male of M. f. frenata, that of neomexicana is decidedly more angular and ridged. The postorbital constriction is narrower, the mastoid breadth greater (it is less than the postpalatal length in some subadult males), the sagittal crest much higher with impressions of the temporal and masseter muscles carried farther forward on the frontals, rostrum shorter and tympanic bullae wider and more inflated. Similar, though less marked, differences exist between the females. As compared with M. f. leucoparia and perotae, the same differences as noted above between frenata and neomexicana exist. In addition the tympanic bullae are so far removed from the foramen ovale that the distance from the anterior end of each bulla to the foramen ovale, instead of being less than the height of tympanic bullae, is in leucoparia more than four-fifths this height and in perotae more than the entire height. Also, in perotae, the squamosal, anterior to each tympanic bulla, is ventrally convex rather than ventrally concave as in neomexicana. Compared with M. f. longicauda, neomexicana is relatively narrower in the interorbital region, has relatively shorter tooth-rows, a V-shaped rather than a U-shaped interpterygoid space and in males has the interorbital region flat rather than convex and the sagittal crest is higher. The same differences are to be noted in comparison with nevadensis but here the difference in relative length of tooth-row is less. The same differences exist also in comparison with M. f. arizonensis except that its interorbital breadth, relative to the rest of the skull, is about the same. Difference in size is especially marked here; even females of neomexicana average larger than males of arizonensis.
Remarks.--When Barber and Cockerell named this subspecies in 1898, they had three specimens. Only two others are known to have been taken before this time. These are a skeleton, without corresponding skin, taken at Lozier, Texas, in 1890 by Wm. Lloyd, and no. 21779/36482, U. S. Nat. Mus., taken on April 6, 1893, by R. D. Lusk at Tombstone, Arizona. On the back of a label recently attached to the last mentioned specimen the name C. K. Worthen appears and probably signifies that the specimen was purchased from this dealer in vertebrate specimens.
M. f. neomexicana has a large geographic range. The old male from Liberal, Seward County, Kansas, extends the known range far to the northeast. Geographically, this occurrence is logical for the southwestern desertlike conditions extend to this part of Kansas. Probably the subspecies occurs in southeastern Colorado and in the panhandle of Oklahoma where conditions are similar. Bailey (1905:198) lists neomexicana as a member of the mammalian fauna of Texas. As stated by him (loc. cit.:198) this inclusion is based on geographic grounds and not on actual specimens. Strecker (1926:13) also includes neomexicana in his list of Texas mammals but writes me, under date of January 9, 1928, that "I included Mustela frenata neomexicana as a Texas mammal on the strength of its being mentioned by Bailey. . . ." On better ground, Bailey (1928:97) lists the subspecies as occurring in southeastern New Mexico at Carlsbad Cavern. However, Bailey (loc. cit.) knew of the existence of weasels just below El Paso and at Langtry, Texas. An unsexed skeleton, no. 167891, in the United States National Museum, from Lozier, Texas, is not certainly identifiable to subspecies. If, as I think, the animal is a female, its skull is intermediate between that of frenata and neomexicana although when all features are considered it is seen to be nearest the latter. The large size (basilar length of 46.5 mm.) may reflect some relationship to texensis. The field notes of the collector furnished me by Dr. H. H. T. Jackson (MS), describe the color as brownish yellow above and sulphur below. The admission of this subspecies to the list of mammals of Texas is made certain by the female (no. 1572, Texas Cooperative Research Collection) taken on July 28, 1940, 1-1/2 mi. NW Kent, Texas, by C. E. Scull.
The skull alone from Durango (City of), extends the known range far to the south. This skull is typical of neomexicana. Skins from the same place would be especially interesting as showing the approach, if any, in color, of neomexicana to M. f. leucoparia.
Mr. D. D. Stone of Casa Grande, Arizona, writes, under date of February 2, 1927, that a weasel was seen by an acquaintance of his in a field near Chandler, Maricopa County, Arizona. Probably this was neomexicana. If so, its range extends much farther west than collected specimens show.
Actual intergradation with M. f. frenata is not shown by the material at hand. The two females from Albuquerque, although typically neomexicana as regards color, have smaller, less prominently ridged skulls than females of neomexicana of the same age from farther south and approach M. f. nevadensis.
Probably the geographic ranges of M. f. neomexicana and M. f. latirostra do not meet; the only evidence of the existence of weasels in all of the large area, comprising western Arizona and the deserts of eastern California, which intervenes between the ranges of the two subspecies is the skull of a young individual, no. 68842, Mus. Vert. Zoöl., from 10 miles northeast of Bard, Imperial County, California. There, on December 29, 1932, Jack C. vonBloeker, Jr., retrieved the weathered skull with some of the vertebrae attached, from the top of a wood rat's nest beneath a mesquite tree near the west bank of the Colorado River.
The idea that the carcass may have been washed down the river from far upstream gains no support from a comparison of the specimen itself for the tympanic bullae are larger than in nevadensis and the skull is larger than the largest males seen of arizonensis, the two upriver races. On the basis of size the skull could be either a male of latirostra or a female of neomexicana. These two subspecies, like arizonensis and the skull in question, have much inflated bullae. However, the immaturity of the specimen conceals any other diagnostic cranial features, and prevents referring it certainly to either neomexicana or latirostra. In any event the specimen provides no evidence of intergradation between the two forms last mentioned. Speculating on its identity, I should say that it might be either an intergrade between arizonensis and nevadensis, from southern Utah or northwestern Arizona, or a member of an unnamed race resident in the lower part of the valley of the Colorado River.
Whereas M. f. panamensis and M. f. aureoventris are the darkest-colored weasels and occur in regions of heavy rainfall, M. f. neomexicana is the lightest-colored American weasel and occurs in an extremely arid region where the rainfall and humidity are slight.
According to Barber and Cockerell (1898:189) "The type specimen was shot in the grass on the shore of Armstrong's Lake. . . ." Bailey (1928:97) found the tracks of one of these animals "in the great pit at the west entrance to" Carlsbad Cavern and supposes they "hunt the cave walls for mice and other small game." Data on the label attached to no. 230973 states that the specimen was taken, two miles west of Willcox, Arizona, in a prairie dog town.
Only two of the 23 skulls show evidence of infestation of the frontal sinuses by parasites.
Specimens examined.--Total number, 28, arranged alphabetically by states and from north to south by counties in each state. Unless otherwise indicated specimens are in the United States National Museum.
=Arizona.= Graham County: Safford, 1. Cochise County: 2 mi. W Willcox, 1; Willcox, 1; 8000 ft., Chiricahua Mts., 1; 6000 ft., Pinery Canyon, Chiricahua Mts., 1[33]; Tombstone, 1; Sulphur Spring Valley, 1[74].
=Durango.= "Durango City," 1.
=Kansas.= Seward County: Liberal, 1[93].
=New Mexico.= Bernalillo County: 3 mi. NW Albuquerque, 2. Lincoln County: 7800 ft., South Fork Eagle Creek, White Mts., 1. Chaves County: Pecos River, 10 mi. E Roswell, 8[74]; Dexter, 1[74]. Otero County: Cloudcroft, 9000 ft., 1[90]. _Dona Ana County_: Mesilla Park, 2 (1[75], 1[7]); Berino, 2.
=Texas.= Culberson County: 1-1/2 mi. NW Kent, 1[90]. _Terrel County_: Lozier, 1.
=Mustela frenata texensis= Hall
Long-tailed Weasel
Plates 22, 23 and 24
Mustela frenata texensis Hall, Carnegie Instit. Washington Publ. 473:99, November 20, 1936.
Mustela frenata, Strecker, The Baylor Bull., 27:14, September, 1924.
Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926 (part).
Type.--Male, adult, skull with skin of head, neck and tail; no. 14821, Amer. Mus. Nat. Hist.; Kerr County, Texas; September 17, 1897; obtained by H. P. Attwater.
The skull (plates 22-24) and dentition are complete and unbroken. The preserved parts of the skin are not stuffed.
Range.--Lower Sonoran and possibly Upper Sonoran life-zones of central Texas. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela frenata arthuri in possessing white facial markings and postorbital breadth less than distance between posterior borders of P4 and P2; from M. f. frenata in larger size of body and skull, the basilar length of which in adult males is more than 52.5; from M. f. neomexicana in Brussels Brown rather than Buckthorn Brown color of upper parts and basilar length of skull more than 52.5.
Description.--Size.--Male: Measurements taken from the dried skins of a young male, no. 15476, Mus. Comp. Zoöl., from Kerr County, Texas, and a subadult male, no. 2017, Baylor Univ. Mus., from 5 mi. N Waco, Texas, are, respectively, as follows: Total length, 600 and more than 510; length of tail, 200 and 225; length of hind foot, 52 and 52.
Female: Skins unknown.
Externals.--As described in Mustela frenata frenata.
Color.--As described in Mustela frenata frenata.
Skull and teeth.--Adult male: See measurements and plates 22-24. As described in Mustela frenata frenata except that: Weight, 8 grams; basilar length 54; least width of palate less than length of P4; anterior margin of masseteric fossa anterior to middle of m2.
Female: Skull unknown.
Remarks.--The type specimen, taken by the veteran collector of Texan mammals, H. P. Attwater, appears to have been the first one of these animals to find its way into the collection of any museum or naturalist. The second specimen from Kerr County was secured by, or through, the well-known commercial collector, F. B. Armstrong. Two trade skins, from Kerr County, taken on December 10, 1938, are in the Texas Cooperative Research Collection, as is also the skeleton of a young animal from Fredericksburg. The two other specimens from McLennan County (both males contrary to the statement of Strecker, 1924:14), owe their preservation to the alertness of John K. Strecker, Curator of the Baylor University Museum, who has given a complete account of their history.
The range of this subspecies is thought to include much of central Texas.
The preserved parts of the skin of the type specimen show it to have been generally large. The part of the tail preserved measures 226 millimeters and the skin of the head and neck is correspondingly large. The skin alone, no. 427, from near Waco, Texas, has, as now stuffed, a body 365 millimeters long. Individuals of this race attain larger size than those of any other American member of the subgenus Mustela with the possible exception of Mustela frenata macrophonius from Veracruz, México. In addition to large size, texensis and macrophonius are analogous in that each has a small geographic range at the northern end of an extensive range of its similarly colored southern relative from which it differs mainly in size. Each of the two groups, goldmani and macrophonius on the one hand and perotae, frenata and texensis on the other, has relatively uniform color, color pattern and body proportions over a large region but at its northern extremity develops a "giant" population, M. f. macrophonius and M. f. texensis, respectively. The skull of the type specimen of M. f. texensis is the largest one seen of any American weasel. The type specimen of M. f. macrophonius has a basilar length that is greater by one-tenth of a millimeter but in every other measurement taken the skull of M. f. texensis is the larger. Its weight, 8 grams, also shows it to be larger.
The broad, white bands in front of the ears are confluent with the white patch between the eyes on both sides in two specimens and on one side only in one other specimen. A white patch between the ears is present in four specimens. The dark spot at each angle of the mouth is absent on both sides in four specimens and on one side only in one other specimen. Thus out of a possible twelve cases, the broad bands in front of the ears are confluent with the spot between the eyes in five cases. Four of the six specimens have a white spot between the ears. The dark spot at each angle of the mouth is present three out of a possible twelve times.
The skull of no. 2017, from five miles north of Waco, is smaller than either of the two skulls seen from Kerr County and in this respect approaches M. f. frenata. There is no actual evidence of intergradation with any other subspecies but intergradation probably does take place with M. f. neomexicana and possibly with M. f. arthuri and M. f. primulina.
Strecker (1924:14) remarks that of the two specimens obtained near Waco, one was taken in a trap baited for mink and the other was shot in a hen house. None of the four skulls had the frontal sinuses infested with parasites.
Specimens examined.--Total number, 7, arranged by counties from north to south.
=Texas.= McLennan County: 5 mi. N Waco, 1[3]; Erath, 1[3]. Gillespie County: Fredericksburg, 1[90]. Kerr County: 4[75]; 1[2]; and 2[90] trade skins.
=Mustela frenata frenata= Lichtenstein
Long-tailed Weasel
Plates 1, 22, 23, 24, 36, 37, 38 and 40
Mustela frenata Lichtenstein, Darstellung neuer oder wenig bekannter Säugethier, 1832, pl. 42, and corresponding text, unpaged; Seton, Lives of game animals, 2:576, 1929.
Mustela brasiliensis Sevastianoff, Mem. de l'Acad. Imp. Sci. St. Petersburg, 4:356-363, tab. 4, 1813, name on plate only, the description being in the text (not of Gmelin, 1788); Gray, Proc. Zoöl. Soc. London, 1865:114.
Putorius frenatus, Baird Mamms. N. Amer., p. 173, 1858; Merriam, N. Amer. Fauna, 11:26, pl. 3, figs. 1, 1a, 1b, June 30, 1896; Bailey, N. Amer. Fauna, 25:198, October 24, 1905.
Putorius (Gale) brasiliensis aequatorialis Coues, Fur-bearing animals, p. 142, 1877, part? ("merely as a substitute for Gray's [supposedly] preoccupied name" that is, aureoventris).
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Putorius mexicanus Coues, Fur-bearing animals, p. 142, 1877, [nomen nudum, cited by Coues in synonymy as "_Putorius mexicanus_, Berlandier, MSS. ic. ined. 4 (Tamaulipas and Matamoros)"].
Putorius brasiliensis frenata, Allen, Bull. Amer. Mus. Nat. Hist., 3:219, April 17, 1891.
Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 6:197, May 31, 1894; Bangs, Proc. Biol. Soc. Washington, 10:9, February 25, 1896; Allen, Bull. Amer. Mus. Nat. Hist., 8:74, April 22, 1896.
Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.
Type.--Female, adult, skull and skin; no. 991, Berlin Zool. Mus., México City, México; June, 1829; obtained by F. Deppe.
The specimen once mounted, now is remade into a study skin and lacks the distal part of the tail. The skull (plates 36-38, 40) lacks the basicranial region.
Range.--Altitudinally, sea level (Brownsville, Texas) to 7600 feet (Tlalpam, México); from southern Texas as far south as México City; Lower Sonoran to at least Transition life-zone. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. perotae in nonextension of blackish over anterior fourth of neck, least width of color of underparts more than 37 per cent of greatest width of color of upper parts; height of tympanic bulla more than distance from its anterior margin to foramen ovale; from M. f. leucoparia by restricted white facial markings that cover less than half surface of head in front of ears, by nonextension of black of head onto anterior half of neck and by wider (more than 7.8) tympanic bullae; from M. f. neomexicana by Brussels Brown rather than Buckthorn Brown color of upper parts and mastoid breadth less than postpalatal length; from M. f. texensis by smaller size of body and skull (basilar length in adult males less than 52.5); from M. f. arthuri by white facial markings and postorbital breadth less than distance between posterior borders of P4 and P2; from M. f. tropicalis by nonextension of blackish over anterior fourth of neck, least width of underparts more than 37 per cent of greatest width of upper parts, postorbital breadth of adult males less than distance between posterior borders of P4 and P2.
Description.--Size.--Male: Fifteen adults and subadults from Brownsville, Texas, yield average and extreme measurements as follows: Total length, 485 (430-556); length of tail, 202 (165-250); length of hind foot, 48 (40-55). Averages believed to be reliable but extremes probably are not. Tail averages 71 per cent as long as head and body. Length of hind foot less than basal length. Corresponding measurements of an adult male (topotype, no. 50826) from Tlalpam, México, are: 505, 203, 53. Another adult male, from Miquihuana, Tamaulipas, México, measures: 520, 215, 52.
Female: Six adults, subadults and young from Brownsville, Texas, yield average and extreme measurements as follows: Total length, 420 (362-456); length of tail, 173 (126-200); length of hind foot, 41 (40-46). Tail averages 70 per cent as long as head and body. Length of hind foot more (with possible exception of no. 36362/48732 U. S. Nat. Mus.) than basal length.
The average differences in external measurements of the two sexes are: Total length, 65; length of tail, 29; length of hind foot, 7.
Externals.--Longest facial vibrissae black and reaching beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 20.
Color.--Spot between eyes, broad band, confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, and posterior two-thirds to one-half of each upper lip, white; remainder of sides and top of head, posteriorly to line connecting posterior margins of ears, blackish; dark spot posterior to angle of mouth present on both sides in about half the specimens; tip of tail black; remainder of upper parts Brussels Brown; chin white; remainder of underparts near (16´a) Ochraceous-Buff (same color in juveniles and young), which color extends distally on posterior sides of forelegs over forefeet and on medial sides of hind legs to feet and sometimes onto upper sides of toes. Least width of color of underparts averaging, in a series of seventeen males from Brownsville, Texas, 47 (extremes 38-53) per cent of greatest width of color of upper parts. Black tip of tail, in same series, averaging 49 (extremes 40-55) mm. long, thus about equal to length of hind foot and averaging 24 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on ten adults from Brownsville): See measurements and plates 22-24; weight (three adults, one topotype and two from Brownsville, Texas), 6.2 (5.3-7.2) grams; basilar length, 49.8 (48.2-51.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoidal breadth less than postpalatal length; postorbital breadth less than length of upper premolars (less than distance between posterior borders of P4 and P2) and not greater (usually less) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 or 4 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter (usually longer) than rostrum; anterior margin of masseteric fossa just behind m2.
Female (based on two adults from Brownsville, Texas): See measurements and plates 36-38, 40; weight, 3.4 (3.3-3.5) grams; basilar length (six, adult to young) 43.3 (41.3-47.3); zygomatic breadth more or less than distance between condylar foramen and M1 and more than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less than (about equal to) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than outside length of P4; tympanic bulla as far posterior to foramen ovale as width of 2 to 3-1/2 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.
The skull of the female averages 45 per cent lighter than that of the average male.
Comparison of the skull with those of M. f. arthuri, tropicalis, perotae, leucoparia and neomexicana has been made in accounts of those subspecies. As compared with M. f. texensis (known only from males), the only difference detected is smaller size.
Remarks.--As Merriam (1896:27) has said: "In 1813 a Russian naturalist, Sevastianoff, gave the name 'Mustela brasiliensis' to a weasel brought to St. Petersburg by Capt. A. J. Krusenstern on his return from a voyage around the world. The animal was said to have come from Brazil, but no definite locality was given." This name was long applied by many European naturalists to American weasels which had white facial markings, and several American naturalists adopted the name. However, Lichtenstein in 1832 applied the name Mustela frenata to the weasels of the vicinity of México City and that name was used for bridled weasels from México and the southwestern United States by most subsequent German writers and by several Americans. In 1896 Merriam (1896:27) showed that Sevastianoff's Mustela brasiliensis, 1813, although probably the same as Mustela frenata, was preoccupied by Gmelin's Mustela brasiliensis, 1788, applied to an otter and that Lichtenstein's name must be used as the next available one. Since that time, 1896, frenata has been the name applied to the large bridled-weasels of Texas and the high table land of México south to México City. It may be added that in 1937 search by the writer among the specimens and records at the Russian Academy of Sciences, in Leningrad, failed to reveal any trace of the type specimen of Sevastianoff's Mustela brasiliensis.
The geographic range of this subspecies is relatively large and, as might therefore be expected, specimens show geographic variation. The specimens from Tlalpam, which Merriam (op. cit.:27) regards as topotypes, differ in certain respects from specimens from Texas. The skull of the adult male "topotype," no. 50826, differs from any other adult male seen in that the basilar length, the length of the upper tooth-rows, the orbitonasal length, the ratio of the same to the basilar length, the mastoidal breadth, the zygomatic breadth, the depth of the skull at the posterior margins of the upper molars, and the length and breadth of M1, are greater. The height of the tympanic bullae is less than the average height for these structures in more northern specimens. The specimens from Tlalpam have also larger external measurements than the average of more northern specimens. All of these features show an approach to the subspecies of more southern distribution. On the other hand, the blackish of the head is not more intense or more extended posteriorly onto the neck than in specimens from Brownsville, Texas. The skin, with skull crushed, no. 767, in the Paris Museum, from 3200 meters elevation near Toluca, does have the black color of the head extended 30 millimeters posteriorly to the ears. In this feature, and also in the extensively white face on which the white bar in front of each ear connects with the frontal spot, as well as with the color of the underparts, the specimen resembles leucoparia. Better material from the western part of the state of México may show the range of leucoparia to extend eastward almost or quite to Toluca.
An adult male, taken on July 15 at Miquihuana, Tamaulipas, is unique in several respects. The top of its head is black, rather than blackish, and this color extends posteriorly on the top and sides of the neck almost halfway to the shoulders. All of the upper parts are much more darkly colored than in other specimens of this race. The least width of the color of the underparts is 63 per cent of the greatest width of the color of the upper parts; thus the color of the underparts is considerably more extensive than in any other specimen seen. The underparts are more intensely colored than in the average specimen. The mastoidal breadth is greater than in any other adult male and amounts to more than the postpalatal length. On available maps the elevation of Miquihuana is given as 1892 meters (about 6200 feet). Thus the dark colors can hardly be ascribed to more tropical conditions than those under which animals from Brownsville, Texas, live. Brownsville is only a few feet above sea level and only 235 miles farther north. The difference noted, therefore, seems to be of geographic significance. However, there is from Alvarez, San Luis Potosí, approximately 115 miles south of Miquihuana, a young (nearly subadult) female, no. 21968, which is as light colored as specimens from Brownsville, Texas, or Tlalpam, México. The only distinctive feature of this specimen is the much greater extent of its white facial markings; they are more extensive even than in the specimen from Miquihuana.
Finally, the series from Brownsville, Texas, indicates that the animal there is smaller than frenata from the vicinity of México (city). The skull is similarly proportioned except that relative to the basilar length the orbitonasal length is more. Several other measurements of the skull of the adult male from Tlalpam, as pointed out above, are actually, although not relatively, greater than in any specimen from Brownsville. The similarities between specimens from the two localities, Tlalpam and Brownsville, are striking; since the two localities lie at opposite, extreme ends of the range more geographic variation would be expected. All that is known of the characters of populations from intermediate localities is that the one specimen from Alvarez shows no peculiarities whereas the one from Miquihuana suggests the existence there of a geographic variant.
None of the specimens seen shows actual intergradation with M. f. neomexicana or with M. f. arthuri but it is supposed that frenata intergrades with each of these subspecies. The difference between frenata and arthuri is greater than between frenata and neomexicana. Bailey (1905:198) records tracks of a weasel seen just below El Paso which he supposed had been made by a weasel of the neomexicana type. He also cited the taking of a weasel at Langtry which suggested to him (op. cit.) ". . . a continuous range from the country of frenatus up the Rio Grande to the type locality of neomexicanus at Mesilla Valley," New Mexico. Other records of occurrence in Texas cited by Bailey, in addition to those provided by specimens examined by the writer, are San Diego, Beeville, and Port Lavaca. The Port Lavaca record is the easternmost one assigned to the subspecies frenata; possibly specimens from there would be referable to arthuri.
The series of thirty-four specimens from Brownsville, Texas, permits measuring the amount of individual and age variation in several features. For instance, the material is sufficient to show that external measurements of subadults and those that fall in the upper part of the category designated as "young" may be included with the measurements of adults, because the mentioned measurements are not appreciably greater in adults. The series of skulls, although not providing more than six of any one age, shows the range of variation in size and proportion of certain parts and enables the student the better to evaluate cranial characters of nearby races known from only a few specimens. For example, not one of the twenty skulls of males from Brownsville and immediate vicinity is as large as either of the two specimens of texensis from Kerr County.
The white facial markings vary much in size and shape. In the series of thirty-four skins from Brownsville the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in three specimens and on one side only in six other specimens. These bands are confluent with the color of the underparts in all but two specimens. In one specimen the connection is lacking on both sides and in the other on one side only. A white patch between the ears is present in two specimens. The dark spot at each angle of the mouth is absent on both sides in eleven specimens and absent on one side only in ten others.
In six other specimens from parts of Texas north of Brownsville, the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in one specimen. A white spot between the ears is present in one specimen. The dark spot at each angle of the mouth is absent on both sides in six specimens and on one side only in three other specimens.
In eleven specimens from México, the broad white bands in front of the ears are confluent with the white spot between the eyes on both sides in two specimens and on one side only in one other specimen. The white spot between the ears is present in one specimen. The dark spot at each angle of the mouth is absent on both sides, in six specimens, and on one side only in one other specimen.
Thus, in 51 specimens the broad bands (one in front of each ear) are confluent with the white patch between the eyes in nineteen out of 100 instances, and not with the color of the underparts in three instances. A white spot between the ears is present in four specimens. The dark spot at each angle of the mouth is present 47 out of a possible 98 times.
Four juvenal specimens from Brownsville, Texas, with their dates of capture and probable age, are as follows: no. 58574, [F], three weeks old, taken on February 15; no. 17318/24239, [M], four weeks old, taken on March 16; no. 45899, [F], forty days old, taken on May 21; no. 21778/36481, [M], thirty days old, taken on October 20. In the order given, the dates of birth of these four juveniles would be approximately as follows: January 25, February 15, April 1, and September 20. The dates of birth of other specimens less than three months old as judged by the stage of development of the skull, and reckoning backward from the dates of capture, are as follows: April 1, April 30, May 25, October 12, and December 21. Thus, young appear to be brought forth at Brownsville, Texas, in the fall, winter and spring, that is to say from the latter part of September until the latter part of May.
Mustela frenata frenata is either free of the parasites that infest the frontal sinuses of most weasels, or withstands their presence remarkably well, for only one skull shows a definite pathological condition of the frontal sinuses.
Allen (1896:74) quotes H. P. Attwater, with respect to this species in Bexar County, Texas, as follows: "Not common, but occasionally met within the chaparral and cactus lands, where Wood Rats, Rabbits and Quail abound. They were frequently met with around San Antonio during the great 'Tramp Rat' [= Sigmodon hispidus texianus, see Bailey (1905:116)] invasion of 1889-90."
Specimens examined.--Total number, 63, arranged by counties, and in México by states, from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=Texas.= Bexar County: San Antonio, 2 (1[2]). Goliad County: Charco, 1. Nueces County: Corpus Christi, 1[2]. _San Diego County_ (not found), 1. Hidalgo County: La Hacienda, 1. _Duval County_: San Diego, 2[7]. County in question: Lower Rio Grande, 1. Cameron County: Brownsville, 34 (3[2], 4[1], 3[93], 2[75], 1[59], 1[60], 1[4]); no locality more definite than county, 2.
=Nuevo León.= Río Ramis, 20 mi. NW Montemorelos, 1[90].
=Tamaulipas.= Matamoros, 6; Miquihuana (now in Nuevo León), 1[75].
=San Luis Potosí.= Alvarez, 1[75].
=México=: Region montagneuse des environs de Toluca, Nevada Toluca, 3200 M., 1[84]
=Distrito Federal.= City of México, 2 (1[4]); Tlalpam, 2. No locality more definite than México, 4 (1[4], 3[7]).
=Mustela frenata leucoparia= (Merriam)
Long-tailed Weasel
Plates 1, 24, 25, 26, 29, 30, 36, 37 and 38
Putorius frenatus leucoparia Merriam, N. Amer. Fauna, 11:28, June 30, 1896.
Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 2:165, October 21, 1889.
Putorius frenatus frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 22:259, July 25, 1906.
Mustela frenata leucoparia, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.
Type.--Male, adult, skull and skin; no. 34914/47179, U. S. Nat. Mus., Biol. Surv. Coll.; Pátzcuaro, Michoacán, México; July 27, 1892; obtained by E. W. Nelson; original no. 2960.
The skull (plates 29 and 30) lacks most of the braincase; a fragment, consisting of the supraoccipital and the coalesced frontals and parietals remains. The rostrum, left zygomatic arch, palate, left pterygoid, left glenoid fossa and right postorbital process are intact. The teeth all are present and entire. The lower jaw lacks the right coronoid process and the lateral part of the articular condyle. The skin is well made and in good condition. It differs from an adult male topotype (36855, U. S. Nat. Mus.) and other referred specimens in having: the black of the head extended farther posteriorly on the neck, the maximum amount of white on the head, and a white stripe 50 mm. long extending down the middle of the nape from a point between the ears more than half way to the shoulders.
Range.--Sonoran and Transition life-zones of mountains west of México (city) in Michoacán and Nayarit. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. goldmani in least width of color of underparts more than 47 per cent of greatest width of color of upper parts, hind feet colored like underparts rather than like upper parts; postorbital constriction less than, rather than more than, combined length of upper premolars; from M. f. macrophonius by same details of coloration as from goldmani and by ventrally concave rather than ventrally convex pretympanic part of squamosal; from M. f. perotae by least width of color of underparts more than 40 per cent of greatest width of color up upper parts; height of tympanic bulla more than three-fifths distance from its anterior margin to foramen ovale; from M. f. frenata by white facial markings that cover half of surface of head in front of ears, by extension of black of head onto neck halfway to shoulders and by narrower (less than 7.8) tympanic bullae; from M. f. neomexicana by Argus Brown rather than Buckthorn Brown color of upper parts and distance from anterior margin of tympanic bulla to foramen ovale more, rather than less, than four-fifths of height of tympanic bulla.
Description.--Size.--Male: Two adults and one young from Los Reyes and Pátzcuaro, Michoacán, yield average and extreme measurements as follows: Total length, 514 (510-523); length of tail, 206 (196-215); length of hind foot, 55 (52-58). Tail averages 67 per cent as long as head and body. Length of hind foot more than basal length.
Female: One adult from Artenkiki, Jalisco, and one subadult from Pátzcuaro, Michoacán, measure, respectively, as follows: Total length, 412, 400; length of tail, 159, 159; length of hind foot, 41, 42. Tail averages 64 per cent as long as head and body. Length of hind foot equal to or greater than basal length.
The average differences in external measurements of the two sexes are: Total length, 108; length of tail, 47; length of hind foot, 13.
Mustela frenata leucoparia has a greater total length and length of tail than either M. f. frenata or goldmani. The hind foot is longer than that of frenata and approximately the same as in goldmani. Relative to the body length, the tail averages longer than that of goldmani and shorter than that of frenata.
Externals.--As described in Mustela frenata frenata.
Color.--Broad white bands on sides of head, extending anterodorsally anterior to each ear, confluent with white spot between eyes and with color of underparts; posterior third of each upper lip white; remainder of sides and top of head, and neck posteriorly to point halfway to shoulders from ears, black; no dark spots at angles of mouth; tip of tail black; remainder of upper parts Argus Brown; chin white and sometimes also chest, neck and medial sides of hind legs; remainder of underparts near (16´) Ochraceous-Buff (near (a) Ochraceous-Buff in juvenal female), which color extends distally over all of each foreleg (except its lateral face proximally from about middle of forearm) and on medial side of hind leg and over most of upper side of each foot. Least width of color of underparts averaging, in eight specimens, 54 (extremes 44-61) per cent of greatest width of color of upper parts; black tip of tail averaging, in four males, 52 (extremes 38-78) mm. long, thus averaging 25 per cent of length of tail-vertebrae.
As compared with M. f. frenata and goldmani: white facial markings more extensive; color of underparts less restricted and more extended on legs; black tip of tail relatively of about same extent as in frenata and thus much less than in goldmani; black color of head extending farther posteriorly than in frenata but not so far as in goldmani.
Skull and teeth.--Male (adult): See measurements and plates 24-26, 29, 30. As described in Mustela frenata frenata except that: Weight (no. 128972) 6.3 grams; basilar length, 51.2; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla more or less than (about equal to) distance from its anterior margin to foramen ovale; anterior margin of masseteric fossa anywhere from slightly anterior, to slightly posterior, to m2.
Female (based on no. 26153): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 44.5; zygomatic breadth less than distance between condylar foramen and M1, or than between anterior palatine foramen and anterior margin of tympanic bulla; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla not more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row or than length of rostrum.
The skull of the female is 43 per cent lighter than that of the male.
Comparison of the skull with those of M. f. perotae, goldmani and neomexicana has been made in the accounts of those subspecies. As compared with that of frenata the main difference is the less inflated tympanic bulla, the height of which is approximately equal to, rather than decidedly more than, distance from its anterior margin to foramen ovale.
Remarks.--The first specimen known to have been preserved is the alcoholic in the British Museum of Natural History, taken in September, 1891, on the Río Santiago in Jalisco, by D. A. C. Buller. The other known specimens of this white-faced weasel are divided between the American Museum and the United States National Museum. The two referred specimens from Jalisco were the last of several helpful ones collected in México and Central America by J. H. Batty, and these two were taken less than three months before Batty's tragic death in Chiapas (see Allen, J. A., 1906:191). The five specimens from Michoacán were taken by Nelson or Nelson and Goldman together. Merriam had only three of these when he named the subspecies and remarked (1896:29) that "This form is the poorest subspecies described in the present paper." Although the form is not strongly marked, the two additional specimens from Michoacán and better comparative material than Merriam had confirm several of the differential characters ascribed to it by him and indicate the existence of still other characters.
M. f. leucoparia occurs in the Sonoran and Transition life-zones. No. 27258 from Los Masos, and no. 26153 from Artenkiki (see specimens examined for other spellings) approach true frenata in coloration. Each of these specimens has a few white hairs between the ears and the white patch between the eyes is confluent on one side only with the lateral white bands on the side of the head. No. 27258 from Los Masos has a dark spot at each angle of the mouth. The 7 other specimens are relatively uniform in coloration. Each has the white spot between the eyes confluent on both sides with the extensive white areas on each side of the face. None has a dark spot at either angle of the mouth. Of these 7 specimens, the type specimen and three others have white hairs forming a median line between the ears and a fifth specimen has a white spot behind each ear.
M. f. leucoparia is most like M. f. frenata. Unlike frenata, leucoparia has tympanic bullae that are less inflated, narrower and less projected, at their anterior margins, from the cranium. In these characters leucoparia is intermediate between M. f. frenata and M. f. goldmani. The latter subspecies has the least inflated, narrowest and least projecting tympanic bullae of the three. The black color of the head extends, on the average, farther posteriorly than in M. f. frenata but not so far as in M. f. goldmani. The general color, too, is intermediate between that of M. f. frenata and that of the much darker M. f. goldmani. The white facial markings are more extensive than in either M. f. frenata or M. f. goldmani. This applies to both the white area between the eyes and the one on each side of the head between the ear and eye. M. f. neomexicana, whose range possibly meets that of M. f. leucoparia, also has more extensive white facial markings than M. f. frenata but less extensive markings than M. f. leucoparia.
On the basis of skulls alone, specimens of frenata from Tlalpam and those of leucoparia from Los Reyes can hardly be distinguished. This fact, and the circumstance that the specimens from the northern part of the range of leucoparia closely resemble frenata in color, constitute sufficient evidence for regarding the two as only subspecifically distinct. The female, no. 26153 from Artenkiki, as mentioned above, approaches true frenata in coloration. On this account it is not to be regarded as typical and it was because no other skulls of adult females were available that this one was used for comparison with females of allied races.
M. f. leucoparia is, then, a subspecies of the large, temperate-zone group and is unique in possessing the maximum extent of white facial markings.
None of the seven skulls shows signs of having had the frontal sinuses infested with parasites.
Specimens examined.--Total number, 8, all from México. Localities are listed by states from north to south. Specimens from Michoacán are in the United States National Museum; one from Río Santiago is in the British Museum of Natural History; all others are in the American Museum of Natural History.
=Nayarit.= Tepic, 1.
=Jalisco.= Río Santiago, 1; Los Masos, 1; "Artenkiki" (J. A. Allen, 1906, p. 238, writes "Artenkikil" and, on p. 259, "Artenkiki."), 1.
=Michoacán.= Zamora, 1; Los Reyes, 1; Pátzcuaro, 3.
=Mustela frenata perotae= Hall
Long-tailed Weasel
Plates 36, 37 and 38
Mustela frenata perotae Hall, Carnegie Instit. Washington Publ. 473:100, November 20, 1936.
Putorius frenatus, Merriam, N. Amer. Fauna, 11: pl. 3, fig. 2, June 30, 1896.
Type.--Female, adult, skull and skin; no. 54278, U. S. Nat. Mus., Biol. Surv. Coll.; 12,500 feet, Cofre de Perote, Veracruz, México; May 26, 1893; obtained by E. W. Nelson; original no. 4864.
The skull (plates, 37-39) lacks the right zygomatic arch. Left p2 is missing. The skin is fairly well made and in good condition except that the extreme tip of the tail has been broken off and there are two holes in the right hind leg. The underparts show the beginning of a spring molt.
Range.--From 7500 (?) feet (Perote) to 13,500 feet (Popocatépetl), Upper Sonoran, Transition and Boreal life-zones of mountains along Puebla-México boundary, eastward to western central Veracruz and south into Oaxaca. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. frenata, its nearest relative, in extension from head of blackish onto anterior fourth of neck; restriction of color of underparts (least width of same less than 37 per cent of greatest width of color of upper parts), height of tympanic bulla less than distance from its anterior margin to foramen ovale; from M. f. macrophonius and M. f. goldmani in presence of, rather than absence of, color of underparts on hind feet; upper parts (black) Brussels Brown rather than Argus Brown or darker; from M. f. tropicalis in larger size (adult female with total length more than 400, basilar length more than 40, weight of skull more than 3 grams); postorbital breadth less than combined length of upper premolars; m1 more than 5.4 long; from M. f. leucoparia in white facial markings so restricted that spot between eyes is not confluent with white stripe in front of ear, or, if so, narrowly (less than 4 wide) confluent; color of upper parts extending onto antipalmar face of forefoot, least width of color of underparts not more than 40 per cent of greatest width of color of upper parts; height of tympanic bulla not more than three-fifths distance from its anterior margin to foramen ovale.
Description.--Size.--Male: A nontypical specimen from Cerro San Felipe, Oaxaca, measures: Total length, 500; length of tail, 205; length of hind foot, 52.
Female: The type specimen, measures: Total length, 418; length of tail, 160; length of hind foot, 45.
In this male the tail is 70, and in the female, 62 per cent as long as the head and body. In each the hind foot is longer than the basal length.
The differences in external measurements between these two specimens, representing the two sexes, are: Total length, 82; length of tail, 45; length of hind foot, 7.
Externals.--As described in Mustela frenata frenata.
Color (based on type specimen).--Color and color pattern as described in Mustela frenata frenata except that: blackish of sides and top of head extends one-fourth of way back to shoulders from ears; throat and breast as well as chin white; remainder of underparts near (16´ c) Ochraceous-Buff; least width of color of underparts equals 36 per cent of greatest width of color of upper parts; black tip of tail equal to 28 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on a referred specimen from Cerro San Felipe which certainly is nontypical): See measurements. As described in Mustela frenata frenata except that: Weight, 4.9 grams; basilar length, 49.2; postorbital breadth more than distance between posterior borders of P4 and P2; tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla.
Female (based on type specimen, an adult): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight 3.4 grams; basilar length, 43.5; zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla one-half to three-fifths distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum.
The skull of the female is 33 per cent lighter than that of the nontypical (and smaller than average) male from Cerro San Felipe.
Comparison of the skull with that of M. f. tropicalis is made in the account of that subspecies. Compared with the skull of M. f. macrophonius, that of the female of perotae is more flattened, has the longitudinal dorsal outline distinctly concave rather than flat just behind the postorbital processes, and much wider tympanic bullae. Accordingly, the basioccipital is slightly narrower in perotae. The more marked postorbital constriction of the type specimen of perotae possibly is due to its relatively greater age. As compared with the skull of M. f. leucoparia, that of the female of perotae has less inflated tympanic bullae, the height of each being half as great as distance from its anterior margin to foramen ovale, whereas, in leucoparia (as represented by no. 26153) the two distances are equal. As compared with that of M. f. frenata, the skull of the female of perotae differs mainly in the lesser inflation of the tympanic bullae and their relative position. The height of each bulla is in perotae only half as much as, but in frenata more than, the distance from its anterior margin to foramen ovale. The anterior margin of the bulla is much less projected from the floor of the braincase in perotae. The squamosal anterior to each bulla is convex ventrally in perotae but flat or concave ventrally in frenata.
Remarks.--The type specimen and a juvenal female from the town of Perote were taken in the spring of 1893 by E. W. Nelson. Of these two, the type specimen was mentioned and figured by Merriam (1898:30, fig. 16 [= fig. 15], pl. 3, fig. 2) as Putorius frenatus. The referred nontypical specimen from Cerro San Felipe, Oaxaca, was referred by Merriam (op. cit.:29) to Putorius frenatus goldmani with the comment that it was intermediate ". . . both in coloration and cranial characters, between typical frenatus and goldmani;. . . ." No other published references to this subspecies, or specimens of it, have been seen. In 1941 and 1942, W. B. Davis and associates took four specimens along the boundary between the states of Puebla and México.
Although the specimen from Cerro San Felipe, Oaxaca, is referred to Mustela frenata perotae, to the description of which it answers best, that specimen, on account of its structural characters and geographic position relative to adjacent races, is in reality an intergrade between several of the adjacent races. Some of its intermediate characters are pointed out in the discussion of M. f. goldmani. In the specimens from 45 and 55 kilometers ESE of México (city) the black color of the top of the head does not extend so far behind the ears as in the holotype of M. f. perotae and in this feature the two specimens show intergradation between the two subspecies, perotae and frenata.
The type specimen taken on May 26, is acquiring new hair on the belly and lower sides which appears to be the result of a normal molt.
As would be expected from its geographic position, M. f. perotae resembles M. f. frenata of northern México and the high mountain forms of southern México more than it does the lowland tropical forms. This is true as regards size of entire animal, proportions of its parts, and size, general angularity and major proportions of its skull. The marked postorbital constriction, convex supralacrymal face of rostrum, width of tympanic bullae and angularity of the braincase place it nearest M. f. frenata as does also the color and color pattern. The ventrally convex squamosal anterior to each tympanic bulla and the slight degree of projection from the cranium of the anterior margin of each tympanic bulla are intermediate in degree between the condition in M. f. macrophonius and that in M. f. frenata. Thus M. f. perotae combines several characters of M. f. frenata on the one hand with some of M. f. macrophonius on the other and in some features, for instance in the size, shape and degree of inflation of the tympanic bullae, presents intermediate stages of development.
On the eastern plain below the high mountain, Cofre de Perote, there ranges the similarly colored, smaller, tropical weasel, Mustela frenata tropicalis. Between M. f. perotae and M. f. tropicalis there is marked differentiation in the skulls with much less differentiation in coloration. The differences in typical skulls of the two subspecies are so pronounced that one would, at first glance, hardly believe it possible for direct intergradation to occur between them on the sides of this mountain. Merriam (1896:30) thought that it did not. The two skulls figured by him (op. cit.:31) are a topotype of M. f. tropicalis from Jico and the one which now is the type specimen of M. f. perotae. They show the great difference in size and proportions and are females of comparable ages, not of different ages as I suspected before examining the skulls. However, despite this marked difference in the skulls, there is some, although not conclusive, evidence of intergradation furnished by a young female from Xuchil, Veracruz. This specimen is described in connection with M. f. tropicalis (see p. 366).
None of the seven skulls shows marked deformity of the interorbital region, but two of the three adults appear to have had these parts infested with nematodes.
Specimens examined.--Total number, 7, all from México, listed by localities from north to south. Specimens from Veracruz and Oaxaca in the United States National Museum; remainder in Texas Cooperative Research Collection.
=México=: Monte Río Frío, 45 Km. ESE México City, 1; 55 Km. ESE México City, 1; N slope Mt. Popocatépetl, 13,555 ft., 1.
=Puebla.= Río Otlati, 8700 ft., 1.
=Veracruz.= Cofre de Perote, 12,500 ft., 1; Perote, 1.
=Oaxaca.= Cerro San Felipe, 10,000 ft., 1.
=Mustela frenata goldmani= (Merriam)
Long-tailed Weasel
Plates 1, 24, 25, 26 and 30
Putorius frenatus goldmani Merriam, N. Amer. Fauna, 11:28, June 30, 1896; Elliot, Proc. Biol. Soc. Washington, 18:236, December 9, 1905.
Mustela frenata goldmani, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.
Type.--Male, adult, skull and skin; no. 77519, U. S. Nat. Mus., Biol. Surv. Coll.; Pinabete, Chiapas, México; February 10, 1896; obtained by E. A. Goldman (on attached label collectors recorded as Nelson and Goldman); original no. 9279.
The skull (plates 24 and 30) has the rostrum badly injured. All the right, and part of the left nasal, the upper part of the right maxilla, the postorbital process and intervening area of frontals are missing. Each zygomatic arch is broken but the parts are present and attached to the skull. The frontal and interorbital regions are greatly malformed owing to parasites that infested the sinuses. Right I2 and I3, right and left i3, and the medial parts of the paraconid and protoconid of right m1 are missing. The light facial markings are less extensive than in any of the referred specimens. These markings consist of a separate spot between the eyes and a white line, confluent with the color of the underparts, on each side of the head, that extends from the base of the ear to above the eye. The dark color of the underparts is represented at the angles of the mouth by a spot on the left side and a similar dark area, confluent with the dark color of the face, on the right side. The large size, characters of the skull, and scrotal pouch on the skin prove the specimen to be a male as stated on the label.
Range.--Two thousand five hundred feet (El Cipres, Guatemala) to 9500 feet (near Tecpám, Guatemala), Upper Tropical Life-zone of mountains and western coasts of southern México, Guatemala and Salvador. See figure 29 on page 221.
Characters for ready recognition (characters based on males).--Differs from M. f. nicaraguae and M. f. perda by larger size (total length of adult males more than 489), least width of color of underparts not less than 26 per cent of greatest width of color of upper parts, weight of skull of adult male more than 5 grams; from M. f. macrophonius by smaller size (total length of adult males less than 540), skull of male with basilar length less than 52.5 and weight less than 6 grams; from M. f. perotae (typical specimens of same sex not available) by darker color of upper parts which are Argus Brown or darker rather than Brussels Brown; nonextension of color of underparts onto hind feet; from M. f. leucoparia in least width of color of underparts not more than 37 per cent of greatest width of color of upper parts; color of underparts not extended onto hind feet; black tip of tail two-fifths rather than one-fourth as long as tail-vertebrae; height of tympanic bulla less than four-fifths distance from its anterior margin to foramen ovale.
Description.--Size.--Male: Four adults yield average and extreme measurements as follows: Total length, 508 (500-512); length of tail, 196 (185-207); length of hind foot, 55.5 (54-58). Tail averages 63 (59-67) per cent as long as head and body. Length of hind foot more than basal length.
Female: Typical specimen unknown.
Externals.--Longest facial vibrissae black and reaching beyond ear; carpal vibrissae wholly or in part of same color as upper parts and reaching as far as hypothenar pad; hairiness of foot-soles distinctly less than that shown in figure 20 on page 60.
Color.--Spot between eyes, band, confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear and posterior third of each upper lip, white; remainder of sides and top of head and neck posteriorly to or slightly behind shoulders, black; dark spots at angles of mouth usually absent; tip of tail black; remainder of upper parts Argus Brown or near (n) Argus Brown; chin, throat and breast white; remainder of underparts near (16' c) Ochraceous-Buff; color of underparts extending distally on posterior sides of forelegs onto medial toes and on hind legs to points between knees and heels. Least width of color of underparts, in five adult males, averaging 28 (extremes 26-33) per cent of greatest width of color of upper parts; black tip of tail, in four adult males, averaging 40 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on five adults): See measurements and plates 24-26, 30; weight, 5.4 (5.3-5.5) grams; basilar length, 49.9 (49.6-51.3); zygomatic breadth (except in no. 12523 from Salvador) more than or equal to distance between condylar foramen and M1 or between anterior palatine foramen and anterior margin of tympanic bulla. Mastoid breadth less than postpalatal length; postorbital breadth more or less than length of upper premolars and greater than width of basioccipital measured from median margin of one foramen lacerum posterior to its opposite; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of five upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than or equal to length of rostrum; anterior margin of masseteric fossa immediately behind m2.
Female: Typical skull unknown.
Comparison of male skull with that of M. f. perda made in discussion of that form. Comparison with that of M. f. nicaraguae shows similar differences, some of which are more pronounced. For example, squamosals anterior to tympanic bullae more convex ventrally and these bullae project less from braincase than in M. f. perda; thus the difference in these features is greater between goldmani and nicaraguae than between goldmani and perda.
As compared with the skull of the male of M. f. macrophonius, each one of the skulls of the adult males of M. f. goldmani is smaller in every measurement taken, with two exceptions. The width of the tympanic bullae was more in three specimens of M. f. goldmani as was also the depth of the same in three specimens. Relative to the basilar length all but two of these measurements average less in goldmani; the exceptions are the zygomatic breadth and depth of the skull at the anterior margin of the tympanic bullae which average more. Relative to the basilar length, the orbitonasal length and depth of the skull at the posterior margin of M1 are less in each skull of goldmani. Thus, excepting the width and height of the tympanic bullae and the relative zygomatic breadth and relative depth of the braincase posteriorly, the skull of goldmani is shorter and relatively as well as actually narrower and lighter throughout.
As compared with the skull of the male of M. f. leucoparia, that of M. f. goldmani averages a trifle shorter and no skull of goldmani equals that of leucoparia in actual or relative zygomatic and mastoid breadths or length or height of tympanic bullae. In depth, the skull of goldmani averages actually and relatively greater. Its teeth are smaller. The squamosal anterior to each tympanic bulla is convex ventrally whereas it is concave ventrally in leucoparia as in frenata.
Remarks.--When Merriam (1896:28) named this subspecies, he had only one specimen but he called attention to the more important diagnostic characters, which additional specimens show pertain to the race as a whole.
M. f. goldmani in typical form occurs in high mountains of the Upper Tropical Life-zone and is most closely related to M. f. frenata and M. f. macrophonius. The altitude at which the two specimens were taken, twenty miles southeast of Teopisca in Chiapas, is not known. Merriam (1896:28) states that the type specimen was obtained at "about 8200 feet." The specimen taken by Stirton in Salvador comes from 8000 feet and the one obtained by Barber in Guatemala from 9500 feet. The specimen from Dueñas, the skin alone of a young animal, is not instructive.
As regards size, goldmani is larger than the immediately adjacent subspecies from the Lower Tropical Life-zone but is smaller than M. f. leucoparia or macrophonius. As compared with M. f. frenata, goldmani is longer, has an actually as well as relatively shorter tail, and a much longer hind foot.
The most outstanding difference in externals from frenata is the naked foot soles.
Molting probably takes place twice each year although actual proof of this is lacking. In number 133254 from twenty miles southeast of Teopisca, taken on May 12, the molt is well advanced. Another specimen from the same place still retains the winter coat.
In color, goldmani is much darker than frenata, has less extensive white facial markings, longer black tip on tail, more restricted color of underparts, and lacks the extension of color of the underparts onto the hind feet.
Of the adult males from the high mountains, the type specimen from Chiapas is lightest, and the one from Salvador is darkest. This progressively darker color to the southward probably is geographic variation.
In total length and relative and actual length of tail, the specimen from Salvador is the smallest of the five adult males from the higher mountains. In addition to its darker color and smaller size, no. 12523 from Salvador shows certain distinctive cranial characters. The zygomatic breadth is less than, rather than more than, or equal to, the distance between the condylar foramen and M1 or than that between the anterior palatine foramen and the anterior margin of the tympanic bulla. This difference appears to be correlated with geographic position, since no. 15953 from Guatemala has the three distances about equal and therefore is intermediate in this respect between the specimen from Salvador and those from Chiapas, in which the zygomatic breadth is greater than the other two measurements. Also in the greater depth of the skull and smaller size of the teeth, this specimen from Salvador approaches the subspecies of the Lower Tropical Life-zone. It has, however, the longest, highest and widest tympanic bullae of any of the five specimens. The amount of ventral convexity of the squamosal in front of each tympanic bulla appears not to be greater than in the other specimens.
As indicative of intergradation with perotae, leucoparia and possibly frenata, there is the specimen from Cerro San Felipe, Oaxaca. The degree of restriction of the color of the underparts is intermediate between that of goldmani and leucoparia. The same is true as regards the amount of projection from the braincase of the anterior margins of the tympanic bullae. The squamosal immediately anterior to each tympanic bulla is flat instead of ventrally convex as in goldmani or ventrally concave as in leucoparia and frenata. In accordance with the custom adopted in this paper of referring every specimen to some one subspecies, this specimen from Cerro San Felipe is referred to Mustela frenata perotae, to the description of which it most nearly answers.
Possibly goldmani, as here constituted, is a composite form. The specimens from the high mountains closely resemble one another. However, a specimen, no. 68541 from "Finca El Cipres," Guatemala, which place Mr. G. Goodwin tells me is at an elevation of 2500 feet, approximately 5 miles north of Retalhuleu, has a basilar length of 47.3 and is correspondingly small in other parts. This suggests the existence of a small, lowland race on the western side of the central divide corresponding to perda and tropicalis on the eastern side. From only a few miles away, at San Sebastian, there is available, the adult skull of a still smaller animal. This skull only, no. 41026, in the Berlin Zoological Museum, has a basilar length of 46.1, zygomatic breadth of 27.4, and other cranial measurements notably smaller than those of specimens from the high mountains. A skin-only, no. 12038, collection of Donald R. Dickey, from La Cebia, altitude 2150 feet, near the city of San Salvador, seemingly represents an animal smaller than typical goldmani. This specimen from La Cebia has the light color of the underparts extended distally on the hind legs to the tips of the toes as in M. f. tropicalis. However, the upper parts are darker and resemble those of M. f. goldmani. A fourth specimen from only 3500 feet elevation, on the south side of Volcano Tajumulco, Guatemala, no. 41768, Field Museum of Natural History, a subadult male, measures only 490 in total length and has the least color of the underparts so restricted as to amount to only 22 per cent of the greatest width of the color of the upper parts. Both these features are suggestive of the lowland races.
These four specimens indicate that the lowland population on the western side of the divide is smaller than the mountain population. The juvenile from Carolina and a young male from Finca Cipres, however, both closely resemble individuals of goldmani from the higher mountains. All these animals here are referred to goldmani. More specimens may reveal an amount and a pattern of geographic variation in weasels of this region that will require application of another subspecific name.
The female, no. 68540, from Puebla agrees remarkably well with the skull of the female, no. 132528, of macrophonius. Differences displayed by the specimen from Puebla are its slightly narrower braincase and longer space between the foramen ovale and anterior end of the tympanic bulla. Considering the far eastern location of Puebla (just north of Río Motagua, at 89° W, according to a sketch map provided by Mr. G. G. Goodwin), this specimen might be expected to show some approach to the small lowland races. Actually, however, it displays the characters of goldmani better than does the subadult female from Volcano San Lucas, which is nearer the metropolis of goldmani, and I assume at a higher elevation than Puebla.
Concerning this weasel Merriam (1896:29) says: "Mr. E. W. Nelson writes me that this fine weasel is found sparingly in the forest about Pinabete, Chiapas, at an altitude of 7000 to 8000 feet (2100 to 2500 meters). The type specimen was shot in the afternoon while hunting on a heavily wooded hill slope. It was heard making long, slow leaps over the dry, crisp leaves. Coming to a log, it stood up and rested its fore feet on the log, in which position it was shot by Mr. Goldman."
The specimen taken by R. A. Stirton in Salvador comes from an elevation of 8000 feet in the rain forest of the Upper Tropical Life-zone. Mr. Stirton tells me that one morning on visiting his traps set for small rodents, he found in one the partly eaten remains of a Heteromys. Leaving these remains as found he placed a steel trap beside them and on the following morning found the male weasel in the trap.
At least three of the ten specimens had the frontal sinuses infested with parasites.
Specimens examined.--Total number, 15, listed by localities from north to south, and unless otherwise indicated in the American Museum of Natural History.
=México=: Chiapas: 20 mi. SE Teopisca, 2[91]; Pinabete, 1[91].
=Guatemala=: Puebla, 1; Finca Porvenir, 3500 ft., S slope Volcan Tajumulco, 1[60]; Sierra [=? Cerro] Santa Elena, 9500 ft. (near Tecpám), 1[60]; Carolina, 1; Volcano San Lucas, 1; "Finca El Cipres," 1; "Finca Cipres," 2500 ft., 1; Finca San Isidro, San Sebastión, Dept. Retalhuleu, 1[4]; Dueñas, 1[7]; no locality more definite than Guatemala, 1[7].
=El Salvador=: Los Esesmiles, 8000 ft., Chalatenango, 1[59]; La Cebia, 2150 ft., near San Salvador, 1[59].
=Mustela frenata macrophonius= (Elliot)
Long-tailed Weasel
Plates 24, 25, 26, 30, 37, 38 and 39
Putorius macrophonius Elliot, Proc. Biol. Soc. Washington, 18:235, December 9, 1905.
Mustela macrophonius, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912.
Mustela frenata macrophonius, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.
Type.--Male, adult, skull and skin; no. 14063, Field Mus. Nat. Hist.; Achotal, Veracruz, México; January 15, 1904; obtained by Edmund Heller and Charles M. Barber; original no. 3424.
The skull (plates 24-26, 30) is complete and unbroken. Excepting right P2, which has been aborted or broken away, all the teeth are present. The skin is well made and in good condition. As shown by the scrotal pouch, the specimen is a male.
Range.--Tropical Life-zone, probably into Boreal life-zones, of mountains along eastern border of southern Veracruz. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. frenata frenata and M. f. perotae and M. f. leucoparia in lacking color of underparts on hind feet and in larger skull (skulls of adult males with basilar length more than 52.5); from M. f. goldmani by larger size of skull (see above) and entire animal and wider tympanic bullae; from M. f. tropicalis and M. f. perda by larger size (total length of adult males more than 510), postorbital breadth amounting to less than combined length of upper premolars.
Description.--Size.--Male: External measurements of the type specimen, an adult, are: Total length, 598; length of tail, 246; length of hind foot, 59. Tail 70 per cent as long as body; length of hind foot more than basal length.
Female: The skin, without field collector's measurements, of an adult female from Pérez, Veracruz, shows this sex to be correspondingly large. Because the skin is understuffed and because the hind feet are skinned out, reliable measurements can not be obtained from the dried skin.
Externals.--As described in Mustela frenata goldmani except that all carpal vibrissae are of same color as upper parts and that hairiness of foot-soles is halfway between that shown in figures 20 and 21.
Color.--As in darkest individuals of M. f. goldmani, thus, color of upper parts on posterior part of back near (n) Argus Brown. Color of underparts near (12) Mikado Orange in a juvenile, extending distally on posterior sides of forelegs onto inner toes and on hind legs to points between knees and heels. Least width of color of underparts 28 per cent of greatest width of color of upper parts. Black tip of tail 34 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on type specimen): See measurements and plates 24-26, 30. As described in Mustela frenata frenata except that: Weight, 6.9 grams; basilar length, 54.1; zygomatic breadth less than distance between condylar foramen and M1 or that between anterior palatine foramen and anterior margin of tympanic bulla; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 6 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; anterior margin of masseteric fossa below posterior half of m2.
Female (based on no. 132528): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 43.5; zygomatic breadth less than distance between condylar foramen and M1 and more or less than (in the single specimen, equal to) that between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more or less than (about equal to) outside length of P4; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale.
The skull of the female is 48 per cent lighter than that of the male.
Comparison of the skull with that of M. f. goldmani is made in the account of that subspecies. Similar differences probably exist between males of perotae and macrophonius. As compared with skulls of males of M. f. tropicalis and perda, the skull of the male of macrophonius is larger in every measurement taken. The postorbital constriction is less, rather than more, than the combined length of the upper premolars. Relative to the basilar length, the following measurements are less than in any specimen of tropicalis or perda: length of tooth-rows; orbitonasal length; depth of skull at posterior border of upper molars; and depth of skull at anterior margin of basioccipital.
Remarks.--This large weasel appears to have escaped the notice of naturalists until the spring of 1903 when J. Friesser obtained an adult female and juvenal male at Pérez for the collection of the United States Bureau of Biological Survey. These specimens were tentatively referred to Mustela tropicalis. In the following January, Edmund Heller and Charles M. Barber obtained the adult male that was made the type specimen by Elliot who did not see, or if he did, did not mention, the specimens from Pérez. He did, however, refer a young female from Xuchil, Veracruz, to his Putorius macrophonius. This young female is here referred to Mustela frenata tropicalis.
The extent of the geographic range of this subspecies is not well known.
Mustela frenata macrophonius and M. f. texensis are the largest American weasels. The basilar length in the type specimen is greater by one-tenth of a millimeter than in the type specimen of M. f. texensis. The other cranial measurements taken are greater in M. f. texensis. The skull of the female from Pérez is one of the largest skulls examined of that sex. The juvenal male has teeth as large as those of the type specimen and the skull is the largest for its age of any seen. Although the skin of the female is understuffed and hence does not provide reliable measurements, it shows that the female is also large.
The white bands in front of the ears are confluent with the white patch between the eyes on one side only in one specimen. It is the juvenal male. These bands are not confluent with the color of the underparts on either side in the female and on one side only in the adult male. None of the specimens has a white patch between the ears. The dark spot at each angle of the mouth is present only in the juvenile where it occurs on each side. Of the three specimens, the juvenile is the darkest and the adult male the lightest. The white facial markings are most extensive in the juvenal male and the least extensive in the adult female.
M. f. macrophonius most closely resembles M. f. goldmani but in the relatively flattened braincase, deep constriction of the postorbital region and general angularity of the skull approaches M. f. perotae and M. f. frenata.
Only one of the three skulls, that of the female, shows evidence of infestation of the frontal sinuses by parasites, and this did not result in malformation of the interorbital region.
Specimens examined.--Total number, 3, all from México, listed by localities from north to south.
=Veracruz.= Achotal, 1 (Field Mus. Nat. Hist.); Pérez, 2 (U. S. Nat. Mus.).
=Mustela frenata tropicalis= (Merriam)
Long-tailed Weasel
Plates 25, 26, 27, 30, 37, 38 and 39
Putorius tropicalis Merriam, N. Amer. Fauna, 11:30, pl. 3, figs. 5, 5a, 6, 6a, text fig. 16, June 30, 1896; Merriam, Proc. Biol. Soc. Washington, 15:68, March 22, 1902.
Putorius frenatus, Merriam, N. Amer. Fauna, 11:27, June 30, 1896.
Mustela tropicalis tropicalis, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916.
Mustela frenata tropicalis, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.
Type.--Male, adult, skull and skin; no. 54994, U. S. Nat. Mus., Biol. Surv. Coll.; Jico, Veracruz, México; July 9, 1893; obtained by E. W. Nelson; original no. 5195.
The skull (plates 25-27, 30) is complete. All the upper incisors, except the second and third on the left side, are missing. The right upper canine is broken. The skin is well made and in good condition.
Range.--Up to 5000 feet (as now known) in Tropical Life-zone of Veracruz, México. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. frenata and M. f. perotae in least width of color of underparts not exceeding 36 per cent of greatest width of color of upper parts and in postorbital breadth exceeding length of upper molar and premolar tooth-rows; from M. f. macrophonius and M. f. perda in least width of color of underparts averaging more than 29 per cent of greatest width of color of upper parts; and from M. f. perda by longer tympanic bullae which in males are more than 14.9; and from M. f. macrophonius by lesser basilar length (not more than 48) and in postorbital breadth exceeding length of upper molar and premolar tooth-row.
Description.--Size.--Male: The type specimen and no. 12764/11058, a subadult, from Jalapa, Veracruz, measure, respectively, as follows: Total length, 444, 442; length of tail, 175, 160; length of hind foot, 50, 47. The tail is 65 and 57 per cent as long as the head and body. The hind foot is more or less than (approximately equal to) the basilar length.
Female: Merriam (1896:31) gives the measurements of a female topotype (probably no. 54993, U. S. Nat. Mus., which has no measurements written on the attached label) as: Total length, 333; length of tail, 121; length of hind foot, 37. The length of the tail amounts to 57 per cent of the length of the body. The length of the hind foot of no. 54993, U. S. Nat. Mus. is the same as the basal length.
The differences in external measurements between the male and the female topotypes are: Total length, 111; length of tail, 54; length of hind foot, 13.
Externals.--As described in Mustela frenata frenata except that carpal vibrissae do not reach apical pad of fifth digit and hairiness of foot soles is less.
Color.--As described in M. f. frenata except that: Blackish of head extends half way or more from ears to shoulders; upper parts near (14) Brussels Brown or slightly faded tone 2 of Maroon of Oberthür and Dauthenay, pl. 341; underparts of juvenal pelage near (a) Ochraceous-Buff. Least width of color of underparts averaging (in three specimens from Jico and one from Jalapa) 34 (extremes 30-37) per cent of greatest width of color of underparts. Black tip of tail, in two male topotypes, 57.5 (55 and 60) mm. long; thus longer than hind foot and in each individual comprising 34 per cent of length of tail-vertebrae.
As compared with M. f. frenata: White facial markings slightly less extensive; blackish (not black) of head extending onto neck; upper parts slightly darker; ventral side of tail noticeably darker; color of underparts more restricted, averaging approximately one-third rather than nearly one-half width color of upper parts; black tip of tail one-third rather than one-fourth length of tail and much longer than hind foot. Similar differences of lesser amount exist between perotae and tropicalis. M. f. perda, macrophonius and goldmani bear the opposite relation to tropicalis. That is to say, in the latter: White facial markings slightly more extensive; blackish of head less extended over neck; upper parts markedly lighter; color of underparts less restricted and black tip of tail shorter.
Skull and teeth.--Male (based on type specimen and a subadult, no. 11058, from Jalapa): See measurements and plates 25-27, 30. As described in Mustela frenata perda except that: Weight 4.7 (4.6 and 4.7) grams; basilar length 46.7 (45.5 and 47.8); zygomatic breadth more or less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 (including I3) upper incisors; anterior margin of masseteric fossa below middle of m2 or posterior to that tooth.
Female (based on no. 54993 and no. 1060): See measurements and plates 37-39. As described in Mustela frenata perda except that: Weight (of 54993) 2.2 grams; basilar length, 37.5 (36.0-39.0); zygomatic breadth more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more than greatest length of P4; height of tympanic bulla equal to one-third to three-fourths of distance from its anterior margin to foramen ovale.
The skull of the adult female is 53 per cent lighter than that of the type specimen, a male.
Comparison of the skulls of males and females with those of M. f. perda, the nearest relative, has been made in the discussion of that subspecies. Some of the features that readily distinguish skulls of M. f. tropicalis from those of M. f. frenata, perotae and macrophonius are as follows: Weight less than 4.8 grams; basilar length less than 48; postorbital breadth more than length of upper M-Pm tooth-row. The skulls of male frenata, perotae and macrophonius are much larger, heavier, and are decidedly more angular with more constricted postorbital region the least width of which is less than the length of the upper premolars. In frenata the anterior margins of the tympanic bullae are protruded much farther from the braincase. The skull of the female of M. f. tropicalis is smaller, weighing less than 3 grams; basilar length less than 41; postorbital breadth more than length of upper molar and premolar tooth-row.
Remarks.--This subspecies was originally described by Merriam as a full species. Later he described Putorius tropicalis perdus as another subspecies. Allen (1916) placed P. t. perdus in synonymy but named Mustela tropicalis nicaraguae as new. In the present paper all three forms are recognized but are regarded as only subspecifically distinct from the other bridled weasels of México and Central America.
The limits of the geographic range of tropicalis are fairly well known on the south and west but the only specimen available from the tropical coastal region north of Jico, is a young female from a point 50 miles south of Victoria. Thus, how far north along the coast it ranges toward Matamoros, where M. f. frenata occurs, is not known. The three specimens from Jico, a young female from Jalapa and another adult collected by J. Potts and labeled as coming from México City, are assumed to be typical. The latter specimen certainly came from an elevation lower than that of México City because M. f. frenata occurs there. Although the female from Jalapa, agrees well with specimens from Jico, a male, no. 12764/11058, from Jalapa, has a relatively broader skull, as in perda, although the tympanic bullae are short as in tropicalis. The resemblances to perda in features of coloration are: slightly darker upper parts, and the termination just below the knees of the color of the underparts. There are three specimens labeled as from Orizaba that indicate intergradation with perotae as does also the coloration of the juvenal female from 5 kilometers north of Jalapa. The specimens labeled as from Orizaba are old, poorly-prepared skins, only two of which have partial skulls. The size and coloration of the skins suggest perotae as do also the partial skulls in some respects although the skulls show greater resemblance to those of tropicalis.
The topotype, female, no. 54993, was figured by Merriam (1896, fig. 16, p. 31) along with that of what now is the type specimen of M. f. perotae. Merriam called attention to the great difference in size between the skulls of the two sexes of M. f. tropicalis and compared the condition to that found in noveboracensis. Although the skull of the female from Jico is fully adult, it probably is exceptionally small.
The young female from Xuchil is indistinguishable in coloration from the juvenal female of M. f. perotae from Perote, but in size of skull and size of teeth is intermediate between the female of tropicalis from Jalapa and the females from Cofre de Perote.
There is then, indication of intergradation with M. f. perotae as well as with M. f. perda. M. f. tropicalis differs from M. f. perotae and M. f. frenata in about the same way that M. f. perda differs from M. f. goldmani and M. f. macrophonius. M. f. tropicalis and perda each is smaller and more intensely colored than goldmani and macrophonius, and inhabits the lowland to the east of their highland relative.
At least five of the nine skulls have the frontal sinuses infested by parasites.
Specimens examined.--Total number, 13, all from México, listed by localities from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.
=Tamaulipas.= 50 mi. S Victoria, 1[71]. Near? México City, 1.
=Veracruz.= Jico, 3; 5 km. N Jalapa, 1[90]; Jalapa, 2 (1[2], 1[75]); Xuchil, 1[60]; Orizaba, 4 (2[75], 1[4]).
=Mustela frenata perda= (Merriam)
Long-tailed Weasel
Plates 25, 26, 27, 30, 37, 38 and 39
Putorius tropicalis perdus Merriam, Proc. Biol. Soc. Washington, 15:67, March 22, 1902.
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Mustela tropicalis perda, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912.
Mustela tropicalis tropicalis, Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916.
Mustela frenata perda, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.
Type.--Male, subadult, skull and skin; no. 100041, U. S. Nat. Mus., Biol. Surv. Coll.; Teapa, Tabasco, México; March 31, 1900; obtained by E. W. Nelson and E. A. Goldman; original no., 14074.
The skull (plates 25-27, 30) is unbroken and all the teeth are present and entire. The skin is well made and in good condition.
Range.--Fifty feet (Chichén Itzá) to 4000 feet (San Vicente) in Lower Tropical Life-zone south from southern Veracruz through southern México into Guatemala. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. nicaraguae in lesser extent of color of underparts (not more than 22 per cent of greatest extent of color of upper parts), black tip of tail more than 38 per cent of length of tail, and broader skull (in adult males, mastoid breadth more than 23.9 and zygomatic breadth more than 27.4); from M. f. tropicalis in more restricted color of underparts (least width of color of underparts less than 28 per cent of greatest width of color of upper parts) and shorter tympanic bullae, which in males are less than 15; from M. f. goldmani by total length not exceeding 489, least width of color of underparts not exceeding 24 per cent of greatest width of color of upper parts, weight of adult skull less than 5 grams and basilar length less than 48.5.
Description.--Size.--Male: The type specimen and another subadult from San Vicente, Chiapas, measure, respectively, as follows: Total length, 473 and 443; length of tail, 184 and 169; length of hind foot, 51 and 51.5. The tail is 62 and 64 per cent as long as the head and body. The length of the hind foot is greater than the basal length.
Female: Estimates made from the dried skin of no. 218036 are: Total length, 375; length of tail, 140; length of hind foot, 40. The hind foot of no. 65422 from Catemaco also measures 40.
The average differences in external measurements of the two sexes are: Total length, 83; length of tail, 37; length of hind foot, 11.
Externals.--As described in Mustela frenata goldmani except that hairiness of foot soles is slightly less.
Color.--As described in Mustela frenata goldmani except that: back near (n) Argus Brown or Carbon Brown, tone 3, of Oberthür and Dauthenay, pl. 342; underparts Ochraceous-Buff. Least width of color of underparts, in four specimens, averaging 20 (extremes 18-22) per cent of greatest width of color of upper parts; black tip of tail, in two subadult males, averaging 48 (extremes 46-49) per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on type specimen and subadult no. 132997 from San Vicente): See measurements and plates 25-27, 30; weight 4.4 grams (same for each); basilar length 45.7 (45.3 and 46.1); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more or less than (approximately equal to) length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum not greater than length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4-1/2 to 5-1/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below middle of m2.
Female (based on two subadults, nos. 65422 and 218036): See measurements and plates 36-39; weight, 2.4 (2.3-2.5) grams; basilar length, 40.5 (40.4-40.6); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than outside length of P4 and less than inside length of same; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla equal to one-third to one-half distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than (about equal to) length of rostrum.
The skull of the female averages 48 per cent lighter than that of the male.
Comparison of the skull of the male with that of M. f. nicaraguae has been made in the account of that subspecies. The skull of the male as compared with that of M. f. tropicalis has shorter tympanic bullae, deeper braincase at anterior margin of basioccipital, lesser zygomatic and palatal breadth and smaller P4 and m1. The skull of the female is larger in every measurement taken except those reflecting width of the preorbital portion. This part is actually narrower but probably mainly because the females of perda are younger than those of tropicalis. Features in which three skulls of subadults of M. f. perda differ from the five adults of M. f. goldmani and show no overlap are: lesser basilar length, lesser weight, greater relative length of upper tooth-rows, greater relative width of rostrum, greater relative length of rostrum, lesser mastoid and zygomatic breadths, lesser width, length and height of tympanic bullae; lesser outside length of P4 and greater relative depth of braincase at anterior margin of basioccipital and at posterior margin of M1. Features in which perda averages less are: length of tooth-rows, interorbital breadth, orbitonasal length, relative zygomatic breadth, length of m1, outside and inside lengths of P4, width and length of M1, and depth of skull at posterior margin of M1. Features in which perda averages more than goldmani are: relative interorbital breadth, relative mastoid breadth and depth of skull at anterior margin of basioccipital. The length of the inner half of M1 averages the same. As compared with goldmani, the skull of the male of perda is shorter, otherwise generally smaller, but relatively broader except across the zygomatic arches, and relatively deeper. The anterior margins of the tympanic bullae project slightly less from the braincase and the squamosals immediately in front of these bullae are slightly more convex ventrally.
Remarks.--Described by Merriam in 1902 as a subspecies of Putorius tropicalis, the form perda was regarded by Allen (1916:99) as not subspecifically distinct from P. t. tropicalis.
This is the eastern, lowland subspecies of the Tropical Life-zone, corresponding to M. f. goldmani of the higher mountains just as M. f. tropicalis corresponds to M. f. frenata and perotae of the high mountains and table land. The difference in size between perda and nicaraguae and between perda and tropicalis is slight. M. f. perda is slightly less richly colored than M. f. nicaraguae but has the color of the underparts more restricted and has a longer black tip on the tail. In these respects it is second only to M. f. panamensis among Central American weasels. Evidence of intergradation with goldmani is furnished by the specimens from Cobán, Guatemala, and the nearby locality San Cristóbal in Verapaz, Guatemala. Reduced size as compared with goldmani suggests affinity with perda but the greater width of the light-colored underparts, which averages 24 (extremes 18-32) per cent of the greatest width of the color of the upper parts, shows approach to goldmani. Farther north, in Chiapas, however, specimens of perda from San Cristóbal and San Vicente are readily distinguishable from those of goldmani taken a few miles away at Pinabete and near Teopisca. The latter two localities are, however, several thousand feet higher than San Cristóbal (Chiapas) and San Vicente.
Two of the nine skulls (only 3 adult) examined for malformation of the frontal sinuses reveal infestation by parasites.
Specimens examined.--Total number, 18, listed by localities from north to south, and unless otherwise indicated in the United States National Museum.
=Veracruz.= Catemaco, 1.
=Tabasco.= Teapa, 2 (1[7]).
=Chiapas.= San Cristóbal, 1; San Vicente, 1; no locality more definite than state, 1.
=Yucatán.= Chichén-Itzá, 1[76].
=Guatemala=: Cobán, 2 (1[7], 1[4]); Finca la Providenci, S. Cristóbal, Verapaz, 3[76]; central Guatemala, 1; no locality more definite than Guatemala, 5 (2[7]).
=Mustela frenata nicaraguae= Allen
Long-tailed Weasel
Plates 1, 25, 26, 27 and 30
Mustela tropicalis nicaraguae Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916.
Putorius tropicalis, Allen, Bull. Amer. Mus. Nat. Hist., 24:661, 1908.
Mustela frenata nicaraguae, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.
Type.--Male, subadult, skull and skin; no. 30754, Amer. Mus. Nat. Hist., Matagalpa, Nicaragua; April 16, 1910; obtained by W. B. Richardson; original no., 712.
The skull (plates 25-27, 30) of the type specimen lacks the entire right zygomatic arch. Otherwise it is complete. The teeth all are present and unbroken. The skin is complete and unfaded but only partly stuffed.
Range.--Honduras and Nicaragua. Altitudinal and zonal limits unknown. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. costaricensis and M. f. goldmani in shorter black tip of tail (not more than 35 per cent of length of tail) and lesser width (usually not more than 7 mm.) of tympanic bulla; from M. f. perda in greater extent of color of underparts (22 or more per cent of greatest width of color of upper parts), shorter black tip on tail (not more than 35 per cent as long as tail) and narrower skull, the mastoid breadth in adult males being less than 23.9 and the zygomatic breadth less than 27.
Description.--Size.--Male: Average and extreme measurements of five subadults and one young (four from Matagalpa and one from San Rafel del Norte) are: Total length, 450 (420-480); length of tail, 178 (150-190); length of hind foot, 48 (46-50). Tail averages 65 (extremes 56-69) per cent as long as head and body. Length of hind foot (measurements from dried skins) more than basal length.
Female: Measurements unrecorded.
Externals.--As described in Mustela frenata goldmani, except that hairiness of foot soles (between that shown in figures 20 and 21) is less, slightly less even than in M. f. perda.
Color.--As described in Mustela frenata goldmani except that: Back near (n) Argus Brown, or Carbon Brown, tone 4 of Oberthür and Dauthenay, pl. 342. Underparts Ochraceous-Buff. Least width of color of underparts, in four males, young, subadult and adult, 24 (extremes 22-26) per cent of greatest width of color of upper parts; the corresponding per cent in one female is 32; black tip of tail, in two subadult males, averaging 29 (extremes, 28-30) per cent of length of tail-vertebrae; corresponding per cent in one female, 36.
Skull and teeth.--Male (based on type specimen, one adult topotype [?] and one subadult from San Rafel del Norte): See measurements and plates 25-27, 30. As described in Mustela frenata perda except that: Weight, 4.2 grams (estimated for adults); basilar length 45.0 (44.8-45.5); interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of four to five upper incisors; length of tympanic bulla not less than length of lower molar and premolar tooth-row; anterior margin of masseteric fossa below anterior margin of m2 or posterior to that tooth.
Female: Skull unknown.
Comparison of the skull of the male with that of M. f. costaricensis is made in the account of that subspecies. As compared with that of M. f. perda, which it most closely resembles, the skull of the male has a narrower, shorter rostrum, lesser interorbital breadth, lesser mastoid and zygomatic breadths and slightly shallower braincase, measured at anterior margin of basioccipital. The tympanic bullae are slightly less projected, at their anterior margins, from the braincase and the squamosal, directly anterior to each, is a little more convex ventrally. The skull of M. f. nicaraguae is, then, slightly shorter than that of M. f. perda and relatively narrower.
Remarks.--When naming this form, Allen (1916:100) characterized it as "Similar to M. tropicalis tropicalis but general coloration much darker and the white face markings somewhat reduced in area." In the sentence preceding the one quoted, Putorius tropicalis perdus was placed as a synonym of Putorius tropicalis tropicalis. M. f. nicaraguae and M. f. perda are nearly alike in color and color pattern but differ in cranial characters. M. f. perda and M. f. tropicalis are widely different in color and more especially in color pattern but differ only slightly in cranial characters. The aggregate difference between perda and nicaraguae is less than that between perda and tropicalis. All three are lowland forms and each is smaller than the adjacent highland forms, namely, M. f. goldmani, macrophonius, perotae and frenata.
The weasels from Honduras definitely are not typical of nicaraguae as it is known from the specimens from Nicaragua itself. The specimens from the state of Tegucigalpa, Honduras, are larger. Some are darker than topotypical nicaraguae. The dorsal outline of the skull is more nearly flat (less convex) in some. In these and several other differential features studied, the average of specimens from Tegucigalpa is intermediate toward goldmani, but everything considered the animals seem best placed with nicaraguae rather than with goldmani or perda, to which latter also, they show some resemblance. With better material from Nicaragua and additional specimens from Salvador (here referred to goldmani) a restudy of all the material now referred to the three races named would be profitable. Aims of this restudy might be to determine if a highland race additional to goldmani should be recognized and if the lowland races perda and nicaraguae differ from one another in the way that the existing specimens indicate.
In the five males from Matagalpa, the narrow white band in front of each ear is confluent with the color of the underparts on one side only in one specimen and on both sides in two specimens. None of these bands is confluent with the white patch between the eyes. A dark spot at the angle of the mouth is present on one side in one specimen. The corresponding area is dark colored in all other specimens but not separated from the color of the upper parts. In the specimen from San Rafel del Norte the white bands are not confluent with the color of the underparts. The female from Mambacho has the mentioned bands confluent with the color of the underparts. This female approaches M. f. costaricensis in the dark color of the upper parts but has more extensive white facial markings than some specimens from much farther north. Like a female seen of M. f. costaricensis, this one has a "frosted" nape but the white hairs on the back of the neck are less numerous than in the female of M. f. costaricensis.
M. f. nicaraguae in typical form, then, is thought of as a small, lowland, tropical subspecies only slightly differentiated from M. f. perda. By reason of its intermediate characters, it constitutes a link between the lowland forms, and the larger animals called M. f. goldmani and M. f. costaricensis.
None of the four skulls from Nicaragua shows signs of infestation of the frontal sinuses by parasites.
Specimens examined.--Total number, 16, listed by localities from north to south. Specimens are in the American Museum of Natural History, unless otherwise indicated.
=Honduras=: Alto Cantoral, 2; Cerro Grande La Paz, 1. La Flor Archaga, 1[75]; Comayagüela, 1[75]; vicinity of Tegucigalpa, 2; no locality more definite than Honduras, 1[4].
=Nicaragua=: San Rafel del Norte, 1; Matagalpa, 6; Ma[o]mbacho, 1.
=Mustela frenata costaricensis= Goldman
Long-tailed Weasel
Plates 25, 26, 27, 28, 29 and 30
Mustela costaricensis Goldman, Proc. Biol. Soc. Washington, 25:9, January 23, 1912.
Mustela brasiliensis, Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):374, 1874.
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Putorius affinis, Merriam, N. Amer. Fauna, 11:31, June 30, 1896 (part).
Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Lönnberg, Arkiv för Zool., 14(no. 4):16, 1921.
Mustela frenata costaricensis, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.
Type.--Male, young, skull and skin; no. 13770/37149, U. S. Nat. Mus.; San José, Costa Rica; obtained by C. H. Van Patten.
The skull (plates 28-30) is complete and unbroken. All teeth are present and unworn. The skin apparently has been remade. It lacks the distal two-thirds of the tail. The head is somewhat shrunken. The color is possibly faded but if so only to a slight degree. Otherwise, the skin is in good condition. The orange color of the underparts is so intense as to suggest that the full, adult pelage has not been acquired. No white markings are present on the face. There is no sex mark on the label attached to the skin but the size and proportions of the skull and the scrotal pouch on the skin prove that the specimen is a male. The presence of sutures on the dorsal face of the rostrum and the short, wide, and low sagittal crest show the specimen to be young.
Range.--Costa Rica. Altitudinal and zonal range unknown. See figure 29 on page 221.
Characters for ready recognition.--Differs from M. f. panamensis in lighter color of upper parts (tone 2 rather than tone 4 of Reddish Black of Oberthür and Dauthenay, pl. 344) and longitudinally flat interorbital region of skull; from M. f. nicaraguae in darker color of upper parts (of Oberthür and Dauthenay, tone 2 of pl. 344 rather than tone 4 of pl. 342) and greater width (more than 7) of tympanic bulla.
Description.--Size.--Male: No collector's measurements available of fully grown animals. Estimated measurements of adult males: Total length, 470; length of tail, 165; length of hind foot (taken from dried skins of 3 adults), 52 (50-52). Tail estimated to average 55 per cent as long as head and body. Length of hind foot more or less than (about equal to) basal length.
Female: A subadult or adult, from the Candelaria Mountains, and a subadult from Irazú, measure, respectively: Total length, 370, 385; length of tail, 130, 150; length of hind foot, 40, 31. Tail 59 per cent as long as head and body. Length of hind foot probably about equal to basal length.
The estimated differences in external measurements of the two sexes are: Total length, 92; length of tail, 25; length of hind foot, 16 (probably average difference is less).
Externals.--As described in M. f. panamensis (figure 21) except that foot soles are slightly more hairy.
Color.--As described in Mustela frenata panamensis except that: back is near Reddish Black, tone 2 of Oberthür and Dauthenay, pl. 344; chin, lips, and throat white or whitish; remainder of underparts near (c) Ochraceous-Buff; color of underparts rarely extending distally onto toes of forefeet. Least width of color of underparts, in eleven specimens, averaging 23 (10-36) per cent of greatest width of color of upper parts; black tip of tail, in six specimens, averaging 36 (31-38) per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on 2 adults, no. 3.2.1.6. from vicinity of San José and no. 11408, U. S. Nat. Mus., from "Costa Rica"): See measurements and plates 25-30; weight, 5.9 grams; basilar length 49 +; zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth in undiseased skulls less than length of upper premolars (less than distance between posterior borders of P2 and P4) and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum more or less (about equal to) length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; tympanic bulla longer or shorter than (about equal to) lower molar and premolar tooth-row and longer or shorter than (about equal to) rostrum; anterior margin of masseteric fossa directly below posterior border of m2.
Female: Skull of adult unknown.
Comparison of the skull of the male with that of M. f. panamensis has been made in the account of that subspecies. As compared with that of M. f. nicaraguae the skull of M. f. costaricensis is heavier and in every measurement taken is larger. The skull is generally more massive and it follows that most measurements of depth and width are greater in relation to the basilar length as well as actually greater. The individual teeth are larger and the tympanic bullae wider and at their anterior ends are more projected from the braincase. Indeed the skull is more like that of M. f. goldmani than like that of M. f. nicaraguae.
Remarks.--The half dozen ill-prepared skins, with partial skulls inside, of this form in the United States National Museum long were referred either to Mustela brasiliensis or Mustela affinis. It was not until 1912 when Goldman studied these specimens that the distinctive characters of the Costa Rican weasel were recognized and made the basis of the name costaricensis.
M. f. costaricensis is well differentiated from M. f. nicaraguae and M. f. goldmani which occur to the northward and from M. f. panamensis which occurs to the southward and is a large, heavy-skulled, dark-colored animal with white facial markings restricted or absent. In the type specimen and the female from the Candelaria Mountains the white facial markings are only narrow facial bars or a few white hairs, but in the young male from Cervantes there is a well developed bar 6 millimeters wide on each side of the face and a separate nasofrontal spot, 10 x 12 mm. The young female from Cachí has a V-shaped frontonasal spot, on the right side of the face a white bar 5 mm. wide and 17 mm. long connected with the color of the underparts, and on the left side a white spot in front of the ear and another between the ear and eye. White facial markings were not recorded in the other specimens. The color of the upper parts is only a little less dark than those of M. f. panamensis. Owing to the numerous white hairs on the dorsal side of the neck, the nape of the female from the Candelaria Mountains has a frosted appearance not present in other specimens.
M. f. costaricensis is a large animal and among its geographic neighbors is approached in size only by a specimen of panamensis from Boquete, Panamá. Also the young male from Cervantes suggests panamensis in the less flattened interorbital region, but even so is more like costaricensis. The small size of two young males, one from Navarro and the other from the vicinity of San José, is suggestive of M. f. nicaraguae. However, the large size of most of the specimens and the configuration of the skull are more as in M. f. goldmani than in M. f. nicaraguae and thus suggest that the known specimens are of high mountain subspecies. The long black tip of the tail is another point of resemblance to M. f. goldmani, the high mountain subspecies to the north. Perhaps in the lowlands of Costa Rica, there are weasels of another subspecies.
Of the eight skulls examined for malformation of the frontal sinuses, each of the two adults and two subadults shows signs of having the frontal sinuses infested with parasites.
Specimens examined.--Total number, 14, listed by localities from north to south.
=Costa Rica=: Irazú (Frasu or Irasu on label), 3000 M., 1[4]; Cervantes, 1[2]; San José, 1[91]; vicinity of San José, 2[7]; Azahar Cartago, 1[78]; Tucurrique, 1[7]; Cachí, 1[7]; El Muñco [= Muñeco?] (Río Nivarro [= Navarro?]), 4000 ft., 10 mi. S Cartago, Caribbean Slope, 1[76]; Navarro, 1[91]; Candelaria Mts., 1[75]; no locality more definite than Costa Rica, 3[91].
=Mustela frenata panamensis= Hall
Long-tailed Weasel
Plates 1, 25, 26, 27, 28, 29 and 30
Mustela frenata panamensis Hall, Proc. Biol. Soc. Washington, 45:139, September 9, 1932; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.
Mustela brasiliensis, Alston, Biol. Cent. Amer., Mammalia, p. 78, 1879.
Putorius affinis, Bangs, Bull. Mus. Comp. Zoöl., 39:49, April, 1902; Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.
Mustela affinis, Goldman, Proc. Biol. Soc. Washington, 25:10, January 23, 1912; Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.
Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Goldman, Smithsonian Miscel. Col., 69 (no. 5): 161, 1920.
Type.--Female, subadult, skull and skin; no. 170970, U. S. Nat. Mus., Biol. Surv. Coll.; Río Indio, Canal Zone, near Gatún, Panamá; February 17, 1911; obtained by E. A. Goldman; original no. 20897.
The skull is complete and unbroken. The left lower incisor is broken off but all the other teeth are present and entire. The skin is well made and seems to be in faded, worn, first, adult pelage.
Range.--Sea level (type locality) to 5800 feet (Boquete, see Bangs [1902:49]); Upper Tropical and Lower Tropical life-zones of Panamá. See figure 29 on page 221.
Characters for ready recognition.--Differs from both M. f. meridana and M. f. costaricensis in darker tone (tone 4 of Oberthür and Dauthenay, pl. 344) of color of upper parts and in convex dorsal outline of skull (Compare figures of mentioned subspecies on plates 25-27).
Description.--Size.--Male: Two adults from Boquete in the Museum of Comparative Zoölogy, nos. 10112 and 10113, measure, respectively, as follows: Total length, 480 and 400; length of tail, 170 and 143; length of hind foot, 52 and 43. Hind feet of two other adult males measure 46 on dried skins. Tail, in two specimens mentioned above, is 55 and 56 per cent as long as head and body. Length of hind foot, in each of three adults, slightly longer than basal length. Corresponding measurements of no. 178970 from Mt. Pirre are: 422, 164, 50. Tail 64 per cent (same per cent as in young male, no. 137514 from Boquete) as long as head and body, and hind foot longer than basal length.
Female: An adult and a young from Chiriquí, nos. 18434 and 18435 (Acad. Nat. Sci. Philadelphia), measure, respectively: Total length, 372, 389; length of tail, 138, 144; length of hind foot, 42, 41. The type specimen measures: Total length, 408; length of tail, 159; length of hind foot, 46.5. Tail 64 per cent as long as head and body, and hind foot longer than basal length.
The average differences in external measurements of the two sexes from the vicinity of Boquete are: Total length, 59; length of tail, 15; length of hind foot, 6.
Externals.--Longest facial vibrissae black and extending beyond posterior border of ear; carpal vibrissae wholly, or in part, black and extending as far as hypothenar pad; hairiness of foot-soles as shown in figure 21.
Color.--Usually, posterior fourth of each upper lip and sometimes few hairs in front of ear, white; sides and top of head and neck posteriorly to, or behind, shoulders, black; dark areas at angles of mouth confluent with color of upper parts; tip of tail, black; remainder of upper parts near (n) Bay of Ridgway and Reddish Black, tone 4, pl. 344 of Oberthür and Dauthenay; chin and lips, whitish; remainder of underparts Warm Buff or near (16´ c) Ochraceous-Buff; near (12) Salmon-Orange in juveniles and small young; color of underparts extending distally on posterior sides of forelegs to wrists, but not to soles, and on hind legs to or slightly below knees. Least width of color of underparts, in seven specimens, averaging 18 (extremes 11-28) per cent of greatest width of color of upper parts; black tip of tail, in five adults and subadults, averaging 45 (extremes 41-50) per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on three adults from Boquete): See measurements and plates 25-30; weight, 5 (4.5-5.4) grams; basilar length, 45.2 (42.8-48.3); zygomatic breadth more or less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum approximately same (more or less than) length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4-1/2 to 5-1/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or slightly less than (approximately equal to) length of lower molar and premolar tooth-row or length of rostrum; anterior margin of masseteric fossa directly below posterior fourth of m2.
Female (based on subadult, type specimen and one adult from Siola): See measurements; weight, 3.3 and 2.1 grams; basilar length, 41.3 and 39.3; zygomatic breadth more than distance between condylar foramen and M1 and more or less than (about equal to) that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than combined length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than length of P4 (less in the adult); anterior margin of tympanic bulla as far posterior to foramen ovale as width of five upper incisors; height of tympanic bulla less than (about half) distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of lower molar and premolar tooth-row or than rostrum.
The skull of the one adult female from Chiriquí is 58 per cent lighter than the average of the two adult males.
The skull of the male of M. f. panamensis as compared with that of M. f. meridana, is heavier and averages larger in nearly every measurement taken. Relative to basilar length, tooth-rows, orbitonasal length, interorbital breadth and zygomatic breadth averaging narrower. Mastoid breadth always narrower. Tympanic bullae longer, narrower, and usually slightly less protruded. P4 and m1 larger. Dorsal outline of skull, viewed laterally, more convex. Postorbital breadth actually and relatively greater. Postorbital processes, mastoid processes, and sagittal crest not so well developed. Differences between skulls of females, in so far as known, similar to those described between males.
As compared with M. f. costaricensis, M. f. panamensis has a lighter skull averaging smaller in every measurement taken except interorbital breadth, which is greater. Relative to basilar length, width of rostrum, interorbital breadth and depth of skull at plane of upper molars, less. Tympanic bullae shorter, narrower, less protruded. P4, M1, and m1 larger. Dorsal outline of skull, viewed laterally, more convex. Postorbital breadth relatively and actually greater. Postorbital processes, mastoid processes, sagittal crest and lambdoidal crest less developed. No skull of an adult female of M. f. costaricensis is available for comparison.
Remarks.--This subspecies had not been recognized by previous workers because specimens from Panamá were supposed to be Mustela affinis Gray up until 1916, when Allen (1916:100) restricted the type locality of M. affinis to Bogotá, Colombia. At that time Allen referred specimens from Panamá to Mustela affinis costaricensis, and Goldman (1920:161) followed Allen.
The specimens examined show much variation. Part of this is geographic variation. For instance the specimens from Boquete approach M. f. costaricensis in size more than do those from farther south. Too few adult females have been seen to ascertain the amount of secondary sexual variation. Bangs (1902:49) suggested that the sex of no. 10113 was wrongly recorded and that it was not really a male. If so, this would reduce the range of apparent variation in size of males from Boquete by half and bring it into accord with the amount normally existing in adult males from one locality. No. 10113 is adult but the skin shows no mammae which would prove it to be a female instead of a pigmy male. Although even smaller than 10113, the type specimen is so much larger than females of M. f. meridana that I have wondered if it is correctly sexed. However, the fact that it was sexed by E. A. Goldman, a collector of wide experience, lessens the possibility that a mistake was made.
The color of the underparts is more restricted in panamensis than in any other subspecies of the species. Excluding the specimen from Mt. Pirre, the least width of color of the underparts averages 16 (extremes 6-24) per cent of greatest width of the color of the upper parts. This feature, together with the black color, imparts an appearance to the Panamá weasel that is strikingly like that of a mink. M. f. panamensis is one of the two blackest weasels; M. f. aureoventris is the other. Each of these subspecies occurs in a region of heavy rainfall and there clearly is a positive correlation between high humidity and intensity of color. The black tip of the tail, as regards extent, here reaches the maximum attained among Central and South American weasels. The foot soles are less hairy than in any other member of the subgenus Mustela. The tympanic bullae are lower and less inflated than in any other subspecies of the species.
Adequate specimens from central and southern Panamá may reveal the existence of one or more additional subspecies since animals from each of the three localities now represented differ from those from the other two and some of these differences are correlated with geographic position. However, specimens from all three localities agree in several features. For example all of them have the dorsal outline of the skull highly convex, transversely, and, more especially, longitudinally. In this respect they are sharply differentiated from any other American weasel. Nevertheless, M. f. panamensis is clearly a link between the North and South American subspecies and panamensis intergrades with the adjacent subspecies. The large size of the skull and teeth and the slightly more ventrally projected tympanic bullae of no. 10112 from Boquete approach features seen in M. f. costaricensis. The smaller size of skull and teeth of no. 178970 from Mt. Pirre are points of resemblance to M. f. meridana.
The type specimen was selected from a region where M. f. panamensis is thought to have its distinctive characters well developed. The specimen is not adult and, therefore, does not show as many differential characters as does a nontypical adult from Boquete. Nevertheless, the majority of the above mentioned differential characters are shown by the type specimen and an adult from the same place would, it is judged, show all the differential characters better than would an adult from Boquete.
Of the 11 skulls examined, 6 show no signs of having had the frontal sinuses infested with parasites.
Specimens examined.--Total number, 19, listed by localities from north to south and unless otherwise indicated in the United States National Museum.
=Panamá=: Boquete, 10 (3[75], 1[8], 1[2], 3[4], 1[7]); Río Gariche [é], 5300 ft., 1[1]; Siola, 1[1]; Chiriquí, 1[7]; Río Indio, near Gatún, 1; Mt. Pirre, 3 (2[1]); Calovébora, 1[7] (locality not found, possibly misspelling of Calovébora); no locality more definite than Panamá, 1[4].
=Mustela frenata meridana= Hollister
Long-tailed Weasel
Plates 25, 26, 27, 37, 38 and 39
Mustela meridana Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.
Putorius affinis, Robinson and Lyon, Proc. U. S. Nat. Mus., 24:147, October, 1901; Allen, Bull. Amer. Mus. Nat. Hist., 30:256, December 2, 1911.
Mustela affinis, Osgood, Field Mus. Nat. Hist. Publ. 155, zoöl. ser. 10:61, January 10, 1912.
Putorius macrurus, Allen, Bull. Amer. Mus. Nat. Hist., 31:92, April 19, 1912.
Mustela affinis affinis, Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916 (part).
Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916 (part).
Mustela frenata meridana, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.
Type.--Male, subadult, skull and skin; no. 123341, U. S. Nat. Mus., 1630 meters elevation, Montes de Mérida, near Mérida, Venezuela; August 14, 1903; obtained by S. Briceno.
The skull (plates 25 and 26) lacks the right exoccipital condyle and posterior half of the right zygomatic arch. The teeth all are present, unworn and entire. The skin is well made and complete.
Range.--Near sea level (San Julián) to 8500 feet (Montes de Culata, Mérida, Venezuela), and 9000 feet (Santa Elena, Colombia). Temperate to Subtropical life-zones of Venezuela and northern and western Colombia. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, presence of p2 and two roots, rather than one root, on P2; from M. frenata panamensis in lighter color of upper parts (tone 3 rather than tone 4, pl. 344, Reddish Black, Oberthür and Dauthenay), flat rather than convex dorsal outline of skull immediately behind postorbital processes (see pl. 27); from M. f. affinis, in males, by lesser average breadth and length of skull and greater actual and relative size (see measurements) of facial part of skull; from M. f. aureoventris, in males, by lighter upper parts (tone 3 rather than tone 4, pl. 344, Reddish Black, Oberthür and Dauthenay) and by smaller skull and teeth (basilar length less than 45, length of m1 less than 6.3, width of M1 less than 4.8, outside length of P4 less than 5.7).
Description.--Size.--Male: Average and extreme measurements of topotypes (as recorded by collectors on labels, and so uniform as to show them not to be accurate to within more than 5 mm.) are as follows: Total length, 434 (410-460); length of tail, 164 (150-180); length of hind foot, 50 (no variation in collectors' measurements). Tail averages 61 per cent as long as head and body. Length of hind foot more than basal length. Corresponding measurements of no. 22191, a young male from Mérida, measured by Osgood or Conover, are 439, 165, 54. The adult male no. 18703, from Páramo de Tama (eastern boundary of Venezuela) has the following measurements written on the label by Osgood: 404, 150, 47.
Female: Average and extreme measurements of topotypes (as recorded by collectors on labels and so uniform as to show them not to be accurate to within more than 5 mm.) are as follows: Total length, 347 (320-370); length of tail, 128 (120-130); length of hind foot, 40 (no variation in collectors' measurements). Tail averages 57 per cent as long as head and body. Length of hind foot more than basal length. Two females, adult no. 11034 and young no. 11033 from Cincinnati, Santa Marta, Colombia, measured by M. A. Carriker, Jr., measure, respectively, as follows: 371, 330; 140, 140; 38, 36. No. 14463, adult, from Río Zapata, Colombia, measured (by J. H. Batty), 315, 138, 39. No. 32182, adult, from Mira Flores, Cauca, Colombia, measured (by W. B. Richardson), 375, 150, 43.
The average differences in external measurements of the two sexes, at Mérida, are: Total length, 87; length of tail, 36; length of hind foot, 10.
Externals.--Longest facial vibrissae black (few rarely white) and extending beyond ear; carpal vibrissae colored like underparts or upper parts, and not extending beyond apical pad of fifth digit; hairiness of foot soles slightly greater than shown in figure 21.
Color.--As described in Mustela frenata panamensis except that: Posterior fourth of each upper lip rarely, and small spot in front of ear usually, white; black of head proper not extending back of ears and grading insensibly into color of upper parts; anterior half of upper parts of adults "frosted" with numerous white hairs (tick bites?), upper parts near (n) Bay or tone 2 of Reddish Black (pl. 344, Oberthür and Dauthenay) or tone 3 in freshest, unfaded pelage. Least width of color of underparts (in ten males from Mérida) 20 (17-23) per cent of greatest width of color of upper parts. Black tip of tail, in same series, 60 to 75 mm. long, thus longer than hind foot and 41 (40-44) per cent as long as tail-vertebrae.
Skull and teeth.--Male (based on type specimen and seven topotypes, five adults and three subadults): See measurements and plates 25-27; weight, 4.1 (3.8-4.3) grams; basilar length, 43.6 (42.3-44.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth greater than length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum greater than length of tympanic bulla; least width of palate greater than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or less than (approximately equal to) alveolar length of lower molar and premolar tooth-row and shorter than rostrum; anterior margin of masseteric fossa posterior to m2 and confined to posterior third (34 per cent average, 32 minimum, 37 maximum) of mandible.
Female (based on four adult topotypes): See measurements and plates 37-39; weight (no. 143665), 2.3 grams; basilar length 37.2 (36.3-38.2); zygomatic breadth more or less than distance between condylar foramen and M1 or than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth (sinuses badly infested with parasites) more than length of upper premolars or width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than length of P4; tympanic bulla as far posterior to foramen ovale as combined width of 4 to 5 upper incisors; height of tympanic bulla less than (one half to three fourths of) distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row.
The skull of the female is 44 per cent lighter than that of the average male.
Comparisons of the skull with those of M. f. panamensis and affinis have been made in the accounts of those subspecies. As compared with the skull of the male of M. f. aureoventris, that of meridana averages smaller in every measurement taken. Indeed, none of the skulls of meridana equals that of aureoventris in basilar length, length of tooth-rows, length of tympanic bulla, depth of skull at anterior margin of basioccipital or at posterior margin of upper molars, or measurements of teeth. Relative to the basilar length, most of the measurements are greater in meridana. Exceptions are the relative length of the tooth-rows, and the two measurements of depth of the skull which average less.
Remarks.--In 1914 when Hollister named this weasel he compared it with M. f. affinis and most of the differential characters which he ascribed to meridana were merely "more than" or "less than" in affinis. In affinis, Hollister included specimens from Chiriquí, Panamá, and the coast of Venezuela. The specimens from these three places were referred by Allen (1916:101) to Mustela affinis costaricensis, and he restricted (op. cit.:100) the type locality of Mustela affinis affinis to Bogotá, Colombia, and synonymized Mustela meridana with M. a. affinis. Hollister probably would not have named meridana had he had specimens from Bogotá for comparison and had he regarded them as topotypes of affinis for the difference is slight. Nevertheless, within the large geographic range of M. f. meridana there is some geographic variation. There is more of such variation in the color of the pelage than in shape and size of the skull. The specimen from San Julián is darker than the average and in this respect approaches true panamensis. San Julián is situated at a relatively low elevation on the coast of Venezuela.
M. f. meridana so closely resembles M. f. affinis that the writer has no quarrel with anyone who would synonymize meridana. However, as represented by topotypes, the two races unquestionably are, on the average, different, and specimens from the southeastern part of the range of affinis probably are individually distinguishable from topotypes of meridana.
Variation in the skulls of the series from Mérida is relatively small. This applies to both males and females. The external measurements recorded by native collectors are not accurate to within more than five millimeters but, considering this, variation in external measurements also seems to be slight. The difference in size of the two sexes appears to be uniformly greater than in weasels from Central America. The twenty-six topotypes show that the color and color pattern are relatively uniform. All are of nearly the same tone except juveniles or young which are, as in the case of panamensis, much brighter colored on the underparts. Also, the young have darker-colored upper parts. The adults, without exception, have numerous white hairs scattered over the back of the head, neck and between the shoulders. I have no trustworthy evidence to support the suggestion that these white hairs are the results of tick bites or that they are caused by other parasites which damage the hair follicles. The white facial markings vary relatively little in the 45 specimens carefully examined in this regard. Also, the variation in color pattern of the two sides of the head is small. Indeed, within rather narrow limits, the color of the two sides of the head is the same in every specimen except two. In these two the white spots anterior to the ears are confluent with the color of the underparts. Only one specimen, no. 21342, has a white spot between the eyes and this spot is small. Ten of the twenty-six specimens have a definite white spot or band in front of each ear. Two specimens have such a spot on one side only. The dark spots at the angles of the mouth are present on two sides in three specimens and on one side only in three others. The mentioned spots are, then, present nine out of a possible fifty-two times. When the spots are absent, dark color usually is present in the required area but is confluent with the color of the upper parts.
A young male from San Julián, Robinson and Lyon (1901:147) state ". . . was shot . . . as it ran over some bowlders in a ravine. Its eyes shone with the same greenish light as do the eyes of our common weasel, and it emitted the same strong odor." No. 14463, Am. Mus. Nat. Hist., from Río Zapata, Colombia, according to data on the label, was "taken in timber belt in valley in balk hills" and the native name is Cosonebi. Two specimens taken on the Páramo de Tama, head of Tachira River, Venezuela and Colombia are commented on by Osgood (1912:61) as follows: "One . . . was caught in a steel trap baited with birds and set by the side of a rushing mountain stream. . . . The other was shot in midday as it came prowling about our 'house' in the clearing. . . ."
Of the thirty-three skulls before me, twelve have the frontal sinuses malformed by parasites. These twelve include most of the adults for few of the subadults and fewer of the young show pathologic conditions in the frontal region.
Note on localities.--Several of the localities in Colombia mentioned in "Specimens examined" are described and located by Chapman (1917:640-656, pl. 41) in his "Distribution of Bird-life in Colombia." Place names for Colombia on labels, not found on any map, or duplicated names of which I can not certainly select one, are Río Barrotow, Río Oscuro, Río Zapata, Río Japata, Guasca and El Baldro. Sonson may or may not be the town of that name situated some eighty miles northwest of Bogotá and on the east flank of the Central Andes west of the Magdalena River on the drainage of the Cauca River. In Venezuela most of the specimens from Mérida are labeled 1630 meters, Montes de Mérida. San Julián is some seven miles east of La Guaira (see Robinson and Lyon, 1901:136). San Esteban is located a little way back from the coast between Puerto Cabello and Valencia. Páramo de Tama is on the Venezuelan-Colombian border near the source of the Tachira River (see Osgood, 1912:35). Mt. Duida is shown as at 3° 30´ N and 65° 40´ W by Chapman (1931:13) and Mt. Auyán-tepui as near 5° 15´ N and 62° 50´ W by Chapman (1937:760).
Specimens examined.--Total number, 78, arranged by localities from north to south and unless otherwise indicated in the British Museum of Natural History.
=Venezuela=: San Julián, 1[91]; Carácas, 2; Galipare, Cerro del Avila, 6500 feet, 1; San Esteban, 1[2]; Mérida, 45 (10[91], 14[2], 10[4], 2[60], 2[14], 1[78]); Páramo de Tama, 1[60].
=Colombia=: Páramo de Tama, 1[60]; Cincinnati, 3[9]; Valdiva, 3800 ft., 1; Medellín, 2; 7200 ft., Barro Blanco, 1[2]; Santa Elena, 9000 ft., 1[2]; Santa Elena, 1[2]; Sonson, 2 (1[91], 1[2]); Mt. Auyan-tepui, 1[2]; Pueblo Rico, 5200 ft., 1[91]; Mira Flores, 1[2]; Jerico, near Cauca River, 1; Tornel, 20 mi. NE Quitichao, 1; Mt. Duida, 1[2]; El Tambo, Cauca, 1[78]; El Baldro, 1[2]; Río Japata, 2[2]; Río Zapata, 4500 ft., 1; Río Oscuro, 3300 ft., 1; Río Barrotow, 3300 ft., 1; Guasca, 1[75]; no locality more definite than Colombia, 1.
=Mustela frenata affinis= Gray
Long-tailed Weasel
Plate 30
Mustela affinis Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):375, 1874.
Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).
Putorius affinis, Merriam, N. Amer. Fauna, 11:31, June 30, 1896.
Mustela affinis affinis, Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916; Allen, Bull. Amer. Mus. Nat. Hist., 35:220, May 31, 1916.
Mustela frenata affinis, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.
Type.--Male, adult, skull with skin; no. 54.1.11.3 (skull originally numbered 195d, later 54.6.3.4), Brit. Mus. Nat. Hist.; Colombia [given as new Granada in original description]; purchased from Mr. S. Stevens. Type locality restricted by Allen (1916:99) to Bogotá, Colombia.
The skin is in a good state of preservation and has been made over into a conventional study specimen from a mount on exhibition. Exposure to light when mounted probably accounts for the faded color. The skull (plate 30) lacks the middle 9 mm. of the right zygomatic arch, occiput, basioccipital and posterior two-thirds of the left tympanic bulla. The teeth all are present and entire.
Range.--Four thousand six hundred feet (Quetame) to 9154 feet (El Carmen), Tropical to Temperate life-zones of eastern Andes of Colombia. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, by presence of p2 and by two roots rather than one root on P2; from M. frenata meridana, in case of males, by, on average, greater breadth and length of skull and lesser actual and relative size (see measurements) of facial part of skull; from M. f. aureoventris by lighter-colored upper parts (tone 2 rather than tone 4, pl. 344, Reddish Black of Oberthür and Dauthenay); from M. f. macrura by darker color (Reddish Black, tone 2, pl. 344, Ober. and Dauth., rather than Chocolate, tone 3, pl. 343, Ober. and Dauth.).
Description.--Size.--Male: Measurements in life, estimated from dried skins, are: Total length, 455; length of tail, 175; length of hind foot, 52. Proportions of parts supposedly as described in Mustela frenata meridana.
Female: Estimates from two dried skins: Total length, 365; length of tail, 135; length of hind foot, 43. Proportions of parts supposedly as described in Mustela frenata meridana.
The estimated differences in external measurements of the two sexes are: Total length, 90; length of tail, 40; length of hind foot, 9.
Externals.--As described in Mustela frenata meridana.
Color.--As described in Mustela frenata panamensis except that: posterior fourth of each upper lip and spot in front of each ear white in approximately half of the specimens; black of head proper not extending back of ears and grading insensibly into color of upper parts; upper parts near (n) Bay, or tone 2 of Reddish Black (pl. 344, Oberthür and Dauthenay). Least width of color of underparts (in five males from vicinity of Bogotá) 24 (15-29) per cent of greatest width of color of upper parts. Black tip of tail, in same series, 60 to 75 mm. long, thus longer than hind foot and averaging 38 per cent as long as tail-vertebrae.
Skull and teeth.--Male (based on three adults and two subadult topotypes): See measurements and plate 30. As described in Mustela frenata meridana except that: Weight, 4.5 grams (estimated); basilar length 45.8±; interorbital breadth not greater than distance between foramen opticum and anterior margin of tympanic bulla (type as in meridana where interorbital breadth is more than distance between foramen opticum and anterior margin of tympanic bulla); least width of palate not less than length of P4; masseteric fossa confined to posterior two-fifths (38 to 40 per cent; average 39 per cent) of mandible and not extended anteriorly to middle of m2.
Female: No adults examined.
As compared with M. f. meridana the skull of the male is larger, to the average amount of 2.2 mm. in basilar length and 1.2 mm. in zygomatic breadth of adults; length of tooth-rows and mastoid breadth average greater but relatively less; breadth of rostrum, interorbital breadth and orbitonasal length average actually and relatively less. Thus the skull of affinis is longer and broader, but the facial region is actually, as well as relatively, smaller. As compared with the skull of the male of M. f. aureoventris, that of M. f. affinis is about the same in basilar length. However, in no specimen of affinis are the measurements of length of tooth-rows or breadth of rostrum, actually, or relatively, as great as in aureoventris. The same is true of all measurements taken of M1, P4 and m1. The specimens from the vicinity of Quito and north of there, although referred to macrura, are nearly as dark as typical affinis, approach affinis in cranial characters, and indicate intergradation of affinis with macrura.
Remarks.--Mustela affinis was named by John Edward Gray in 1874 (p. 375) on the basis of a specimen from New Granada. Although usually synonymized with Mustela brasiliensis by later authors until 1896 when Merriam (1896:31) applied the name to weasels from Costa Rica, nearly all the South American and several of the Central American weasels have, at one time or another, had Gray's name, affinis, applied to them. Gray, in 1865 (p. 115) when giving measurements of Mustela aureoventris, probably mentioned the specimen, that later became the holotype. In 1916 (p. 98) Allen restricted the type locality to Bogotá, Colombia. Allen's action was a necessary procedure in clearing up the systematics of South American weasels and was based on good grounds. As set forth by Allen (loc. cit.), and more in detail by Chapman (1917:642), Bogotá has long been the shipping point for Colombian vertebrate specimens, many of which were obtained in the mountains to the east. Allen (1916A:220) quotes Thomas as saying that the type specimen was purchased from Stevens at about the same time that a number of Colombian birds were purchased from the same dealer. Also, specimens from Bogotá agree with Gray's description of the type specimen.
Mustela frenata affinis, as here defined, constitutes one of the several slight geographic variants met with, on the sides of, and between, the three north and south mountain chains of Colombia. The others are lumped under the name Mustela frenata meridana. M. f. affinis, in common with specimens from the northern part of the range of macrura has large teeth. Weasels of all of the region from Quito to Bogotá have large teeth. To the north there is the smaller-toothed meridana and to the south the smaller-toothed macrura grading into the still smaller-toothed agilis, and boliviensis.
Two skins, without corresponding skulls, from Caqueta are lighter colored than any others of affinis; possibly the skins are faded by exposure to light. Since they probably come from an elevation of less than 1000 feet in the Amazonian region, they may pertain to another subspecies.
Complete, unbroken, skulls of affinis are needed to ascertain the degree to which affinis and meridana differ in cranial features. The several specimens from the immediate region of Bogotá show well the color and the color pattern but lack collectors' measurements.
None of the ten skulls examined shows malformation of the frontal region due to infestation of the frontal sinuses by parasites. Possibly three of the four adults were infested, although not severely.
Specimens examined.--Total number, 27, arranged by localities from north to south and unless otherwise indicated in the United States National Museum.
=Colombia=: El Carmen, 1[2]; W. Cundinamarca, 1[7]; Muzzo [= Muzo?], 1[4]; Bogotá, 1; Castillo, near Bogotá, 1[7]; Fambrias, near Bogotá, 1[75]; Bogotá district, 1[2]; Choachí, 9 (1[75], 2[7], 1[84]); Páramo de Choachí, 2 (1[2], 1[84]); Laguna del Verjón (= City of Bogotá), 1[75]; Quetame, 2[2]; Fusagasuga, 1; Caqueta, 2[2]; no locality more definite than Colombia, 3 (1[7]).
=Mustela frenata aureoventris= Gray
Long-tailed Weasel
Plates 27, 28 and 29
Mustela aureoventris Gray, Proc. Zoöl. Soc. London, 1864:55, pl. 8, 1864; Gray, Proc. Zoöl. Soc. London, 1865:115, 1865.
Putorius (Gale) brasiliensis var. aequatorialis Coues, Fur-bearing animals, p. 142, 1877, part? ("merely as a substitute for Gray's [supposedly] preoccupied name," that is, aureoventris).
Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916 (part).
Mustela macrura, Lönnberg, Arkiv för Zool., 14 (no. 4):11, 1921 (part ?).
Mustela frenata aureoventris, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.
Type.--Probably female, juvenile, skull with skin, no. 64.6.6.3 (formerly 1432a), British Mus. Nat. Hist.; probably Subtropical Life-zone of western Ecuador (locality given as Quito, probably because received from that place).
The skin, once exhibited as a mount, has lost some hair from the back and other parts of the body and is not suitable for remaking into a conventional study specimen. The skull lacks the occiput, basioccipital, premaxillae, upper incisors, two of the lower incisors, all of the canines, premolars 2/2 on both sides, right P3, left p3, and has the left jugal mesially defective. The premolars present are not all fully emerged.
Range.--Pacific coastal regions of Ecuador and Colombia; Subtropical and Tropical life-zones. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, by presence of p2 and by two rather than one root on P2; from Mustela frenata macrura by Reddish Black, tone 4, plate 344 rather than Chocolate, tone 3, pl. 343 (of Oberthür and Dauthenay), or slightly darker color of upper parts; from M. f. affinis and M. f. meridana by darker color (tone 4 rather than tone 2, Reddish Black of Ober. and Dauth.) of upper parts and larger size of teeth (M1 with length more than 2.4 and breadth more than 4.7; P4 with outside length more than 5.9; length of m1 more than 6.2).
Description.--Unless otherwise stated, information concerning this subspecies is derived from the one referred specimen available, a young male, no. 34677, Amer. Mus. Nat. Hist.
Size.--Male: Total length, 470; length of tail, 160; length of hind foot, 50. Tail 51 per cent as long as head and body.
Female: Not known.
Externals.--Longest facial vibrissae black and reaching beyond ear. Carpal vibrissae reaching to or beyond apical pad of fifth digit; hairiness of foot soles slightly less than shown in figure 20.
Color.--Sides and top of head and neck posteriorly to shoulders black; white facial markings represented by only five white hairs anterior to right ear, one anterior to left ear and three far back on forehead; dark areas at angles of mouth confluent with color of upper parts; tip of tail black; remainder of upper parts near (n) Bay or Reddish Black, tone 4 of Oberthür and Dauthenay, pl. 344; chin whitish; remainder of underparts Warm Buff, deep orange in juvenile, type specimen, according to Gray (1864, pl. 8); color of underparts extending distally on posterior sides of forelegs to wrists but not reaching foot soles and on hind legs to or slightly below knees. Least width of color of underparts equal to 15 per cent of greatest width of color of upper parts. Black tip of tail equal to 27 per cent of length of tail-vertebrae.
In color, no. 34677 is, to me as it was to Allen (1916:101), indistinguishable from the darkest specimens (nos. 178970 and 10112) of M. f. panamensis. Therefore, M. f. aureoventris is one of the two darkest subspecies of weasels.
Skull and teeth.--Male: See measurements and plates 27-29; weight, 4.3 grams; basilar length, 45.8; zygomatic breadth approximately equal to distance between condylar foramen and M1 and to distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more than length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less (at least in young specimen) than length of tympanic bulla; least width of palate seldom if ever greater than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of three (including I3) upper incisors; height of tympanic bulla not greater than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than orbitonasal length; anterior margin of masseteric fossa below anterior half of m2.
Skulls of males of M. f. aureoventris, and Mustela frenata macrura from the vicinity of Quito so closely resemble one another as not to be distinguished with the material now available, although the teeth of aureoventris are larger. Comparisons of the skulls of males with those of M. f. meridana and affinis, which are readily distinguishable from those of aureoventris, have been made in the accounts of those subspecies.
Female: Skull of adult unknown.
Remarks.--This subspecies of the Tropical Life-zone, or at least the Subtropical Life-zone, of Ecuador, in certain cranial characters resembles Mustela frenata macrura of the Temperate Life-zone. The two differ markedly in color. Nevertheless, a large number of the specimens collected in Ecuador are intermediate in color as well as in zonal distribution.
The type specimen is young or a juvenile. The measurements of no. 34677 from Gualea indicate an animal similar in size to M. f. affinis. Gray (1864:55) states that the type specimen measures "Length of body and head 6 inches, of tail 4-1/2 inches." The plate (pl. 8) accompanying Gray's original description (loc. cit.) is marked one-half natural size and represents the animal as having a head and body length of eight and one-half inches. One year later Gray (1865:115) gives the measurements of this species as "Length of body and head 12, tail 8 inches." Since he had at this time another specimen, larger than the type specimen (which specimen later, probably, became the type of Mustela affinis Gray), the larger measurements probably were taken from it.
Geographically, and as regards cranial characters, Mustela frenata aureoventris is most closely related to M. f. affinis and to the northern section of M. f. macrura, but in color to M. f. panamensis. M. f. aureoventris and M. f. panamensis are the two darkest-colored subspecies and each occurs in a region of extremely heavy rainfall. There is a skin only, no. 32620, Amer. Mus. Nat. Hist., from Munchique, obtained on June 1, 1911, which is appreciably darker than specimens of M. f. affinis in corresponding pelage and is intermediate between M. f. affinis and M. f. aureoventris in color as it is geographically. The specimen measures 495, 202, 52.
The name Mustela aureoventris Gray has been regarded by most authors as preoccupied by Mustela auriventer Hodgson (1841:909). However, the writer is not of this opinion and agrees with Thomas (1920:224) that "The name aureoventris is not invalidated by the auriventer of Hodgson, as, apart from 'one-letterist' differences, its first half comes from the adjective aureus, while Hodgson's name is based on the substantive aurum, so that not only the spellings but the derivations are different." The spelling of Gray's name should be aureoventris for this is the spelling in the original description which in pagination precedes the colored plate of the animal that is labeled Mustela aureoventris. Putorius brasiliensis var. aequatorialis Coues (1877:142) is the only name known to the writer that has been proposed as a substitute for Mustela aureoventris Gray.
Thomas (1920:224) treats Mustela macrura Taczanowski as a synonym of Mustela aureoventris Gray. Allen (1916:101) also treats the two names as applying to the same kind of weasel but regards aureoventris as preoccupied and therefore uses the name macrura. Taczanowski's original description (1874:311) and plate of Mustela macrura indicate an animal that is lighter colored than M. f. affinis. Gray's original description (1864:55) and plate of aureoventris indicate an animal that is darker colored than M. f. affinis. Indeed Gray (1865:115) in speaking of the type of aureoventris as compared with an adult from New Granada [= Colombia] that probably later became the type specimen of Mustela affinis, states: "The young from Quito is much darker than the adult;. . . ." Comparison of the plates accompanying the original descriptions of aureoventris and macrura well illustrate the difference stated in the written descriptions. My examination of the type specimens of M. macrura and M. f. aureoventris shows them to have been fairly accurately portrayed in the plates accompanying the original descriptions. Accordingly the two names are used for the two kinds of animals which appear, however, to be only subspecifically distinct.
Comparison of Gray's plate (1864, pl. 8) with the available specimens from South America indicates that the name aureoventris is based on an individual that is lighter colored than no. 34677 Amer. Mus. Nat. Hist., from Gualea, Ecuador, but on one which resembles no. 34677 more than it does the lighter-colored specimens from the Temperate Zone of Ecuador and northern Perú. Because Quito, Ecuador, is in the Temperate Life-zone and because the available specimens from this zone in Ecuador and northern Perú are distinctly lighter colored than Gray's plate representing the type of aureoventris shows this specimen to be, it is judged to have come from an altitude lower than that of Quito (9350 feet, according to Chapman, 1926:717); probably it came from the Subtropical Life-zone of Ecuador. Indeed Gray (1864:55) did not say that the specimen was collected or obtained at Quito but that it was ". . . received from Quito. . . ." Chapman (1926:717) has pointed out that Quito, since 1846 has been the distributing point for bird skins which specimens ". . . come from the vicinity of the city, from the 'Napo' region on the Amazonian slopes of the Andes, and from Nanegal, Gualea, and other localities on the Pacific side rarely below the Subtropical Zone." It is also pointed out that only some of the specimens are labeled with their approximate place of capture and that even then these localities cannot be accepted as definite; they indicate mainly whether the specimen is from the eastern or western side of the Andes.
The above mentioned considerations and information gained by study of the specimens cause me to think that the type is an intergrade tending toward the lighter-colored Mustela f. macrura of the Temperate Zone although sufficiently dark to be referred to the dark subspecies represented by no. 34677 Amer. Mus. Nat. Hist., from Gualea, Ecuador.
The skull of no. 34677 shows no infestation of the frontal sinuses by parasites.
Specimens examined.--Total number, 3, as follows:
=Ecuador=: Gualea, 1, Amer. Mus. Nat. Hist.
=Colombia=: 8325 ft., Munchique, 1, Amer. Mus. Nat. Hist. In the British Museum of Natural History, the type, (1).
=Mustela frenata helleri= Hall
Long-tailed Weasel
Plates 27, 28 and 29
Mustela frenata helleri Hall, Proc. Biol. Soc. Washington, 48:143, August 22, 1935; Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.
Type.--Male, adult, skull and skin; no. 24133, Field Mus. Nat. Hist.; 3000 feet, Hacienda San Antonio, Río Chinchao, Perú; August 22, 1922. Obtained by Edmund Heller. Original no. 6589.
The skull (plates 27-29) is complete and unbroken. The teeth all are present, entire and but slightly worn. The skin is well made, unfaded, and in good condition.
Range.--Three thousand feet (type locality) to 6700 feet (Ambo), Tropical and Subtropical life-zones of eastern Perú. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, presence of p2 and two roots rather than one root on P2; from Mustela frenata macrura by darker color (Carbon Brown, tone 3, pl. 342 rather than Chocolate, tone 3, pl. 343, Oberthür and Dauthenay) of upper parts.
Description.--Size.--Male: Measurements of the type specimen and topotype, no. 24132, are, respectively, as follows: Total length, 382, 418; length of tail, 152, 164; length of hind foot, 52, 48. Tail 66 and 65 per cent as long as head and body. Hind foot more than basal length.
Female: Measurements of two referred females, no. 24134 from Ambo and no. 24136 from Huanuco, are, respectively, as follows: Total length, 328 and 303; length of tail, 118 and 103; length of hind foot, 39 and 38.5. Tail 56 and 51 per cent as long as head and body. Hind foot shorter than basal length.
The average differences in external measurements of the two sexes are: Total length, 85; length of tail, 49; length of hind foot, 11.
Externals.--Longest facial vibrissae black and extending beyond ear; carpal vibrissae same color as upper parts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 20.
Color.--Rarely a few white hairs anterior to each ear; posterior fifth of each upper lip white; top of head, posteriorly to slightly behind ears, black, grading into color of upper parts of body; dark spots at angles of mouth absent; tip of tail black; remainder of upper parts near (n) Argus Brown and Carbon Brown, tone 3 (pl. 342, Oberthür and Dauthenay); chin whitish; remainder of underparts Warm Buff; color of underparts extends distally on posterior sides of forelegs to wrists but not reaching foot-soles and on hind legs to slightly below knees. Least width of color of underparts 24 per cent of greatest width of color of upper parts in each of two males and 19 to 30 per cent in three females. Black tip of tail longer than hind foot and averaging 40 (39-42) per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on type specimen and adult no. 24132): See measurements and plates 27-29. As described in Mustela frenata macrura except that: Weight, 4.5 (4.2 and 4.8); basilar length, 44.6 (44.0-45.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; breadth of rostrum more or less than (approximately equal to) length of tympanic bulla; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of rostrum; anterior margin of masseteric fossa posterior to m2 by length of that tooth.
Female (based on nos. 24134 to 24136): See measurements. As described in Mustela frenata macrura except that: Weight, 1.7 (1.5-1.9) grams; basilar length, 36.5 (35.3-38.1); zygomatic breadth less than distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more or less than (approximately equal to) outside length of P4; length of tympanic bulla less than length of rostrum.
The skull of the female averages 62 per cent lighter than that of the male.
The skull of the male is generally large and heavy as are the teeth. Comparison with macrura is made in the account of that subspecies. From males of affinis those of helleri differ in: skull shorter; breadth of rostrum and interorbital breadth actually and relatively greater.
Remarks.--The five specimens examined of this subspecies were taken by Edmund Heller for the Field Museum of Natural History in 1922 and 1923. It is to honor his contributions to mammalogy that the subspecies is named helleri. No. 24135 is the specimen carried as a pet for some time by Mr. and Mrs. Heller and of which Mrs. Heller (1924:481) has given an account.
This subspecies is insufficiently known, especially as to geographic range; probably it occupies a considerable range in the Tropical Life-zone along the eastern base of the Andes. The three females, two from Ambo and one from Huanuco, come from a much higher altitude than do the two males and the climate is said to be arid at Ambo and Huanuco. The skulls of the females are 62 per cent lighter and correspondingly smaller in measurements, than those of males. This difference is more than that found in any other South American weasel and it may be that the females are of a subspecies other than helleri.
The type specimen has a broad skull with major proportions strikingly like those of Mustela stolzmanni. Possibly the similar climatic conditions under which the two live have left their impress in similar fashion in this part of each of the two species. The teeth, tympanic bullae, and certain other parts of the skull are, however, so differently proportioned as to show that the skulls represent two species. The referred male has a much longer skull than the type specimen and the relative proportions of breadth and depth of the two skulls differ widely. Judging from large series of weasels examined from localities outside the range of M. f. helleri, the two skulls probably represent almost the maximum of individual variation occurring in one subspecies.
The dark color is as might be expected since helleri inhabits the humid Tropical Zone.
None of the five skulls shows signs of having had the frontal sinuses infested by parasites.
Specimens examined.--Total number, 5, all in the Field Museum of Natural History.
=Perú=: 3500 ft., Hacienda Buena Vista, Río Chinchao, 1; 3000 ft., Hacienda San Antonio, Río Chinchao, 1; Huanuco, 1; Ambo, 2.
=Mustela frenata agilis= Tschudi
Long-tailed Weasel
Plates 27, 28, 29, 39 and 40
Mustela agilis Tschudi, Fauna Peruana, p. 110, 1844; Gray, Proc. Zoöl. Soc. London, 1865:113, 1865; Taczanowski, Proc. Zoöl. Soc. London, 1874:311, 1874; Taczanowski, Proc. Zoöl. Soc. London, 1881:648, 1881; Allen, Bull. Amer. Mus. Nat. Hist., 35:104; April 28, 1916; Thomas, Proc. U. S. Nat. Mus., 58-224, 1920.
Mustela macrura, Allen, Bull. Amer. Mus. Nat. Hist., 35:103, April 28, 1916.
Mustela frenata agilis, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939.
Type.--No type specimen, or type locality more restricted than cold, barren highlands of the Cordillera [referring to Perú] designated.
Range.--High, barren Cordillera of Perú (see Tschudi, orig. descr.); as here restricted, Temperate Life-zone and higher in western Andes and intermountain valleys of Perú. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela frenata macrura by lighter color (Chocolate, tone 2 rather than 3, pl. 343, Oberthür and Dauthenay) of upper parts; length of upper tooth-rows, in females, less than 13; inside length of P4 more than 4.6; from M. f. aureoventris by smaller teeth (maximum size just given for agilis); from M. f. boliviensis by lighter color, upper parts being Chocolate, tone 2, pl. 343, rather than tone 4 or darker of Carbon Brown, pl. 342 (Oberthür and Dauthenay).
Description.--Size.--Male: The stuffed skin of an adult, from Lima, measures: Total length, 460; length of tail, 125; length of hind foot, 45.7. A skin alone from Huarochirí has a body, as now stuffed, 277 mm. long. The tail is missing and the bones of the hind feet have been removed.
Female: The mounted specimen, no. 565, Mus. Polonais d'Hist. Nat., yields measurements, taken by me, as follows: Total length, 250; length of tail, 75; length of hind foot, 32.5. The female, no. 21147, from Macate, measures, 300, 102, 34.
Externals.--Longest facial vibrissae, either dark-or light-colored and extending beyond ear; carpal vibrissae either dark-or light-colored and extending to apical pad of fifth digit; hairiness of foot soles as shown in figure 20.
Color.--Tschudi's description of the color is, in substance, as follows: Head, back and tail reddish gray; base of hair gray, followed by broader grayish-yellow ring and then reddish-brown tip; nose simply dark brown or upper lips edged with white; throat, breast, belly and higher parts of inner sides of extremities whitish gray, at times wholly gray, bases of hairs always gray; feet darker than body, almost chestnut brown; tail darker on tip than at base; ears externally dark brown, internally whitish.
No. 565 possibly somewhat faded from exposure to light, has all the upper parts near (14´ j) Ochraceous-Tawny or Cinnamon, and tone 4 of Oberthür and Dauthenay, plate 323; posterior half of each upper lip white; no other white facial markings present; dark spot at each angle of mouth (one spot confluent with color of upper parts); tip of tail probably black (tip missing); underparts white, belly probably originally with slight tinge of yellow or allied color; color of underparts extending distally on forelegs to feet and onto upper sides of toes and on hind legs to just above heels. Least width of color of underparts equal to about one-fourth of greatest width of color of upper parts.
No. 21147, subadult, from Macate, has a white band confluent with the underparts extending anterodorsally anterior to each ear and the posterior third of each upper lip white. Top of head near (n) Mars Brown, and Carbon Brown, tone 3 (pl. 342, Ober. and Dauth.); tip of tail black; remainder of upper parts near (16" j) Tawny-Olive, and Chocolate (tone 2, of pl. 343 of Ober. and Dauth.) or Raw Umber (tone 3 of pl. 301 of Ober. and Dauth.); anterior half of underparts, including posterior sides of forelegs and antipalmar faces of forefeet, white; remainder of underparts tinged with Warm Buff and extended on posterior legs almost to ankles.
No. 8.1.10.1., male adult, from Lima, is also light colored, and as described in no. 21147, except that left side of head has a white spot rather than bar; posterior eighth of each upper lip white; white frontonasal spot present, 11 x 11 mm.; antipalmar faces of forefeet spotted with brown color of upper parts; color of underparts extending distally on hind legs along medial side of foot to point halfway between heel and tip of inner toe.
No. 13257 from Huarochirí in color and color pattern closely resembles no. 21147. It differs from no. 21147 in slightly lighter color of upper parts, entirely white underparts, less extension of color of underparts onto forefeet, few white hairs instead of white band in front of each ear; color of underparts more restricted.
In each of the four specimens, the least width of the underparts, expressed as a percentage of the upper parts, is as follows: no. 13257, 11 per cent; no. 21147, 29 per cent; no. 565, 31 per cent; no. 8.1.10.1., nineteen per cent.
Skull and teeth.--Male (based on no. 8.1.10.1.): See measurements and plates 27-29. As described in Mustela frenata macrura except that: Weight 4.1 grams; basilar length, 42.5; zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; tympanic bullae shorter than rostrum.
Female (based on no. 21147): See measurements and plates 39 and 40. As described in Mustela frenata macrura except that: Weight (no. 21147, subadult), 1.5 grams; basilar length, 35.2; least width of palate less than outside length of P4; tympanic bulla as far posterior to foramen ovale as combined width of five upper incisors; no. 565 answers to the same description but differs from no. 21147 in greater basilar length and larger tympanic bullae which are slightly more projected, at their anterior margins, from the braincase.
To judge from the skull of the female from Macate and the skull of the male from Lima, the skull and teeth of agilis are smaller than in any other South American subspecies of Mustela frenata, except M. f. boliviensis.
Remarks.--Tschudi almost certainly used the name Mustela agilis in a composite sense. His statement (see quoted matter below) about the marked variation in color of this species, as represented by the skins carried by the Indian women as purses, indicates that the forms here designated as Mustela macrura, M. helleri and possibly others additional to the one here called agilis were included by him under the name Mustela agilis. Taczanowski took account of Mustela agilis when he described other species from Perú. Allen (1916:104) and Thomas (1920:224) were not convinced that Mustela agilis and Mustela macrura were distinct species or subspecies.
Search on August 28, 1937, in the Musée d'Histoire Naturelle, at Neuchatel, Switzerland, by Mr. Théodore Delachaux, assistant there, and the writer, revealed no trace of weasels from Tschudi's collection, although some other specimens of mammals that he figured in the "Fauna Peruana" are preserved in that Museum. Not only were the collections of specimens examined but the new catalogue and old catalogue of mammals were vainly searched for mention of weasels deposited by Tschudi. Later, at the British Museum of Natural History, on page 105 of a personal notebook, of the late Mr. Oldfield Thomas, record was found of his fruitless search for the same specimens of Mustela in May, 1902, at Neuchatel.
Although Tschudi certainly used the name Mustela agilis in a composite sense, as subspecies are at present understood, his description most nearly applies to the light-colored animals from western Perú--the lightest colored of any South American weasels seen. They are of approximately the same color as North American subspecies inhabiting semiarid regions, for example Mustela frenata longicauda of the Great Plains.
Another, but in my opinion less weighty, justification for applying Tschudi's name agilis to these light-colored weasels of western Perú is that by one line of reasoning, Taczanowski in naming macrura (jelskii is a synonym of it) from farther eastward in Perú, and that Hall in naming helleri from still farther eastward, and boliviensis to the southeastward, geographically restricted the application of the name agilis. Hall's action did this because he recognized geographic variation and employed the subspecies concept. Taczanowski, however, proposed his name macrura for a kind of animal which he indicated was specifically (as opposed to subspecifically) distinct from agilis and his account (1881:649) of jelskii indicates that he thought Mustela agilis Tschudi might occur in the same place as the animals which he named as new kinds. Thus, we can not credit Taczanowski with intent to restrict the name agilis geographically, even though later authors may choose to rule that his naming of macrura in effect did so restrict the application of Mustela agilis Tschudi.
The equivalents in millimeters given by Allen (1916:104) for Tschudi's measurements of 9 to 10 inches entire length, and tails of 4 inches to 4 inches and 4 lines, apparently are based on the London scale in use today. If Tschudi employed the Rhine scale also of eight lines to the inch, but one which has the foot longer by an amount of 20 millimeters, or the Leipzig scale in which the foot is 22 millimeters shorter than the London foot, the measurements recorded by Tschudi differ in one direction or the other from those computed by Allen. However, knowledge of which scale Tschudi employed would not help much, if any, in more precise application of the name agilis because he does not indicate whether his measurements are of male or female animals; animals of the two sexes of the same subspecies differ more in external measurements than animals of the same sex of different subspecies of Peruvian weasels.
Specimen no. 565, in the Polish Museum of Natural History, without definite locality, is provisionally referred to this subspecies. The specimen is intermediate in several respects between the female from Macate and the one of macrura from Cutervo.
Tschudi (1844:111-112) has given the following account: "Lebensweise und geographische Verbreitung. Das peruanische Wiesel lebt auf den kalten, öden Hochebenen der Cordillera an sonnigen Steinhaufen und Felsen gewöhnlich in Gesellschaft von 8-12 Stücken. Diese Thierchen sind so ausserordentlich behende und scheu, dass bei dem leisesten Geräusche die ganze Schaar mit Blitzesschnelle verschwindet. Es ist uns auch nie gelungen, eines derselben zu erlegen. Die Indianer aber verstehen es, dieselben lebendig einzufangen und zu zähmen. Ein sehr zahmes sahen wir bei einer uns befreundeten Dame in Tarma; gegen alle Fremden biss es mit Wuth und liess sich nicht anfassen, während es sich von seiner Herrin Alles gefallen liess; sie öffnete ihm den Mund und steckte ihm den Finger hinein, ohne dass es eine böse Miene dazu machte, während es bei der geringsten Bewegung, die wir machten, es zu ergreifen, grimmig auf uns lossprang. Wenn es eingeschüchtert wurde, versteckte es sich in den Busen seiner Gebieterin und kroch ihr bald nachher zum Aermel heraus. An den Wänden und Meublen kletterte es mit grosser Behendigkeit und schlüpfte durch so kleine Ritzen und Löcher, dass wir fast an der Möglichkeit dieses Hindurchdringens gezweifelt haben würden, wenn wir es nicht selbst mit angesehen hätten. Wenn es unartig war, wurde es mit einer Schnur an seinem kleinen Halsbande festgebunden; dadurch vermehrte sich sein Zorn, so dass es zuweilen gegen die Dame auffuhr. Mehrmals verschwand es während 8-10 Tagen und kam dann plötzlich wieder zum Vorschein. Seine Nahrung bestand in Gemüse und Fleisch, besonders aber liebte es Zuckerbrod in Milch aufgeweicht; einmal machte es sich an einen Kanarienvogel, den es auch tödtete. Es erhielt seine Strafe und verschwand dann für immer. Die Indianer sollen dieses Wiesel zum Fange der Viscacha abrichten (davon weiter unten). Sie nennen es Comadreja, auch Ardilla. ([footnote] Ardilla ist spanisch und heisst Eichhörnchen. Mit diesem Namen werden sehr verschiedene Thiere bezeichnet; ausser dem Sc. variabilis und der Galictis agilis auch noch mehrere Nager und einige Didelphysarten.) Die Indianerinnen verfertigen sich aus dem kleinen Felle Geldbeutel. Des Sonntags trifft man unter den vielen tausend Punaindianerinnen die nach den grossen Dörfern der Sierra kommen, um ihre Einkäufe zu machen, kaum ein halbes Dutzend, die nicht solche Börsen mit sich führten, und dann kann man auch die verschiedensten Farbennuancen, die bei dieser Species vorkommen, beobachten."
None of the three skulls referred to this subspecies shows infestation of the frontal sinuses by parasites.
Specimens examined.--Total number, 4.
=Perú=: Macate, 1 (Field Mus. Nat. Hist.); Huarochirí, 1 (Mus. Comp. Zool.); Lima, 1 (British Mus. Nat. Hist.); no locality more definite than Perú, 1 (Mus. Polonais d'Hist. Nat.).
=Mustela frenata macrura= Taczanowski
Long-tailed Weasel
Plates 1, 27, 28, 29, 30, 37, 38, 39 and 40
Mustela macrura Taczanowski, Proc. Zoöl. Soc. London, 1874:311, pl. 48, May 19, 1874; ibid., 1881:647, May 17, 1881; ibid., 835, November 15, 1881; Lönnberg, Arkiv för Zool., 8 (no. 1):21, 1913 (?); Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914; Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Lönnberg, Arkiv för Zool., 14 (no. 4):11, 1921.
Putorius (Gale) braziliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877.
Mustela jelskii Taczanowski, Proc. Zoöl. Soc. London, 1881:647, May 17, 1881.
Mustela affinis, Lönnberg, Arkiv för Zool., 8 (no. 1):21, July 12, 1913.
Mustela aureoventris, Thomas, Proc. U. S. Nat. Mus., 58:224, 1920.
Mustela frenata macrura, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939.
Type.--Male, adult, mounted skin, with skull separate; no. 561, Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Lake Junín, central Perú; 1873; obtained by M. Jelski.
The skull (plates 27-29, 30), mounted with the skin but removed by me for study, lacks the right jugal, the basisphenoid, the basioccipital and parts of each exoccipital bearing the exoccipital condyles. The right tympanic bulla, although detached from the skull, is preserved separately. The teeth all are present and entire. The skin is fairly-well mounted, in a good state of preservation, and shows no fading due to exposure to light.
Range.--Altitudinally, 3200 (Guainche) to at least 12000 feet (Pichincha); Upper Subtropical and Temperate life-zones of central Perú and Ecuador north from the states of Apurimac and Cuzco, Perú, to San Antonio, northern Ecuador. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts; presence of p2 and two roots rather than one root on P2; from Mustela frenata helleri, M. f. affinis and M. f. aureoventris by lighter color of upper parts which are Chocolate tone 3, pl. 343, Oberthür and Dauthenay, whereas, with reference to the same color standard, the colors are: in helleri, Carbon Brown, tone 3, pl. 342; in affinis, Reddish Black, tone 2, pl. 344; in aureoventris, Reddish Black, tone 4, pl. 344; from M. f. agilis by darker color (Chocolate, tone 3 rather than 2, pl. 343, Oberthür and Dauthenay) of upper parts, length of upper tooth-rows, in females, more than 13, inside length of P4 more than 4.6; from M. f. boliviensis by lighter color of upper parts which are as above rather than tone 4 of Carbon Brown, pl. 342 of Oberthür and Dauthenay, and larger size (in males, hind foot more than 45 and m1 more than 5.6).
Description.--Size.--Male (measurements as recorded by Taczanowski in the original description, for two specimens, type and topotype, with correction of the length of tail of his "female" [= male]): Total length, 420, 415; length of tail, 150, 145; length of hind foot, 51, 51. An adult from Yana Mayo, Río Tarma, was measured by Hendee as 394, 134. Hind foot relaxed measures, 47. Tail 55 per cent as long as head and body. Length of hind foot more than basal length.
Female (based on measurements given by Taczanowski (1881:647) of no. 564): Total length, 323; length of tail, 120; length of hind foot, 37. Tail 59 per cent as long as head and body. Length of hind foot approximately equal to basal length.
Differences in external measurements of the two sexes are: Total length, 87; length of tail, 23; length of hind foot, 13.
Externals.--Longest facial vibrissae extending beyond ear; carpal vibrissae color of either upper parts or underparts; hairiness of foot-soles as shown in figure 20.
Color.--(Based on specimens from Cutervo and south thereof). Rarely few white hairs between eyes and in front of ears; top of head posteriorly to slightly behind eyes, near (n) Chestnut-Brown (Ridgway) and Carbon Brown, tone 2 or darker (pl. 342, Oberthür and Dauthenay); posterior half of upper lip rarely white; dark spots at angles of mouth absent; tip of tail black; remainder of upper parts near (l) Russet (Ridgway) and Chocolate, tone 3 (pl. 343, Ober. and Dauth.); underparts white or whitish on medial sides of forelegs, otherwise cream color with tinge of Ochraceous-Buff; color of underparts extended distally on posterior sides of forelegs to just below elbow (in type specimen) or onto forefeet (in specimen from Yana Mayo) and on medial sides of hind legs to points between knees and ankles. Least width of color of underparts averages (in six skins) 17 (14-21) per cent of greatest width of color of upper parts. Black tip of tail longer than hind foot and averaging 36 (32-49) per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on type specimen and no. 562): See measurements and plates 27-30; weight, not known; basilar length, 43.2 (40.8 and 45.5); zygomatic breadth more or less than distance between condylar foramen and M1 and more than that between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less than postpalatal length; postorbital breadth more than length of upper premolars and greater than width of basioccipital, measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more than inside length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 (including I3) upper incisors; height of tympanic bulla not more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below or behind m2.
Female (based on no. 564, from Cutervo, Perú, type specimen of Mustela jelskii Taczanowski): See measurements and plates 37-40; weight, not known; basilar length, 38±; zygomatic breadth less than distance between condylar foramen and Ml and not greater than that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than alveolar length of upper premolars and (probably) more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than inside length of P4; tympanic bulla as far posterior to foramen ovale as width of at least 5-1/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.
As compared with that of helleri, the skull of the male of macrura from Junín southward has a lesser mastoid breadth, notably smaller teeth, and a flatter skull which averages lighter throughout. The skulls of females available indicate that the skull and teeth are larger than in agilis.
Remarks.--Seven years after Taczanowski named this subspecies, he applied the name jelskii to a female taken farther north than the original examples of macrura. As indicated in synonymy, various other names have been applied to animals included by the present author in this subspecies.
Mustela frenata macrura intergrades with M. f. affinis as shown by practically all the referred specimens from north of Junín. As one proceeds northward the color of the weasels becomes progressively darker and the teeth become larger until the conditions found in affinis are met with near the northern border of Ecuador. From the material available it appears that the light-colored upper parts found in macrura characterize weasels of, at least, the Temperate Zone, from Marcapata, Perú, to near Quito, Ecuador. West of the range of macrura there exists the still lighter-colored subspecies, M. f. agilis. Immediately adjacent on the north, east, and south, darker-colored weasels occur. So far as color is concerned, the geographic range of the subspecies M. f. macrura is not difficult to define. However, the small size of the teeth characterizes only that part of this light-colored subspecies from Junín southward including the subspecies boliviensis at the southern extremity of the range of the species. From Cutervo northward the light-colored weasels of the Temperate Zone have teeth similar in size to those of the darker, more northern affinis. To designate the slightly larger-toothed, light-colored animals from Ecuador as a subspecies distinct from affinis and macrura is one solution but at present it seems best to refer all of these light-colored animals to macrura.
The type specimen and topotype no. 562 differ more in the amount of inflation of the tympanic bullae than adult males of comparable ages from a given locality usually do. In other respects, the differences between the two skulls are not greater than those ordinarily found in specimens from the same locality. No. 562 has the tympanic bullae greatly, relative to the other South American weasels, inflated posteriorly. Otherwise, the bullae agree with those of the type specimen.
Specimens from southwestern Ecuador, average large, and include the largest specimens of the species Mustela frenata seen from South America. A subadult male, no. 61406, in the American Museum of Natural History, is the largest. Its external measurements are 482, 191, 56. The basilar length of the skull is 48.2 and the zygomatic breadth is 30.3. Although not so large as this specimen, the corresponding measurements of specimens from Alamor, El Chiral, and even from as far away as Sigsig also are distinctly large.
The skull of the female from Ollantaytambo and that of the male from Marcapata have teeth equally as small as do the specimens from Lake Junín.
The skin alone, no. 194328, from Ollantaytambo has the color of the underparts extended over the entire upper sides of the forefeet. The male from Marcapata has less of this color on the forefeet and is in this respect intermediate between the specimens from Lake Junín and the one from Ollantaytambo.
In size of teeth the female, type specimen of M. jelskii, from Cutervo, shows an approach to the larger-toothed weasels of the northern part of the range of macrura.
The specimens in the Riksmuseum from the vicinity of Quito, Ecuador, have been rather fully described by Lönnberg (1921:11-17) and need little comment here, except to say that they show, as he suggested, that the weasel of the Temperate Zone of Ecuador is an intermediate link between M. f. macrura and M. f. affinis.
The adult female and juvenal male labeled as from Ambato have little left of the skulls except some of the teeth and the assignment of the specimens to the subspecies macrura is made mainly on geographic grounds. These two specimens probably are part of the shipment of birds and mammals of which Chapman (1926:703) speaks as follows: "A small collection of native-made skins purchased by the American Museum from a commission merchant in New York City as from 'Ambato' proved to be from the eastern slope of the Andes." Another skin in the same Museum, labeled by a native collector as from "Baeza arriba" [= above Baeza] is so dark colored and has the color of the underparts so much restricted, as to suggest that it belongs to the race aureoventris. Possibly, therefore, it was taken not at Baeza, Ecuador, which I find to the eastward of Quito at 77° 55' W and O° 25' S, but at some place of the same name on the Pacific Slope, unless the locality has been altogether wrongly recorded on the label. If the specimen was taken near the Baeza above referred to, then it gives evidence of an unnamed race of Mustela on the eastern slope of the Andes, characterized by its dark color. Unfortunately the specimen is young and its skull therefore offers insufficient basis for the judging of its subspecific relationships.
Other specimens, in the British Museum of Natural History, recorded as taken "near Quito" and here tentatively listed under macrura, mostly, include specimens so dark colored as to lead me to think they came from country, lower than Quito, adjacent to the range of aureoventris.
Nematodes taken from the right frontal sinus of no. 562 from Junín proved to belong to the superfamily Oxyuriodea according to Professor W. B. Herms and Mr. O. L. Williams, who have independently identified them. Because these worms had been dried fifty-five years in the mounted specimen and were later boiled in cleaning the skull, a more accurate determination was impossible and whether or not they pertain to the same species found in North American weasels cannot be said. Of 18 adult skulls examined for this type of infestation, 13 were found affected as judged by the evident malformation of the frontal region.
Specimens examined.--Total number, 74, arranged by localities from north to south and unless otherwise indicated in the American Museum of Natural History.
=Ecuador=: Ibarra, 6600 ft., and 7500 ft., 2[7]; San Antonio, 8000-8500 ft., 5 mi. N Quito, 4 (2[7], 2[78]); Nono, 10000 ft., 1; Mindo, 1[78]; Zambiza, 8000-8100 ft., NE Quito, 4 (2[78], 2[95]); Carapungo, 8500 ft, NE Quito, 1[78]; Panecillo, 10000 ft., near Quito, 2[78]; Guapulo, 8800 ft., 3 mi. E of Quito, 1[78]; Pichincha, 10500 ft., and 12000 ft., 2 (1[78], 1[95]); San Ignacio, 11500 ft., Pichincha, 1; Santa Rosa, 9600 ft., Río Pita, 2; near Santa Rosa, 9000 ft., 1; Río San Rafel, 9000 ft., 1; N side Quito, 9000 ft., 1[78]; Quito, 1[4]; near Quito, 5[7]; Nára Papallacta, 11000 ft., 1[78]; below Papallacta, 9000 ft., 1[78]; Chillo Valley, 1[78]; "Hacienda Hda," 10000 ft., Pintag, Valencia, 1; Baeza arriba, 1; Ambato, 2; San Francisco, 8000 ft., E of Ambato, 1; Chunchí, Pagma Forest, 6400 ft., 1[1]; Canar, 2600 M., 1[7]; Malletura, 7600 ft., 1; Contrayerbas, 11000 ft., 1; Sisig, 8500 ft., 3[7]; El Chiral, 1; Almor, 1; Guainche, 3200 ft., 1; no locality more definite than Ecuador, 4[95]; "Received from Quito," 1[7]; Quisaya, 6000 ft. (locality not found), 1[7]; La Carolina (locality not found), 1[78].
=Perú=: La Lejía, 1; Huancabamba, 4 (2[75]); Cutervo, 9000 ft., 1[73]; Condechacha, 7000 ft., Río Utcubamba, 1[7]; San Pedro, 8600-9400 ft., S of Chachapoyas, 1; Celendín, 1[7]; Junín, 2[73]; Yana Mayo, Río Tarma, 1[7]; Ollantaytambo, 9000 ft., 3 (1[7], 2[91]); Ocabamba, 1[7]; Anta Cuzco, 3400 and 3500 M., 2[4]; Marcapata, 1[91].
=Mustela frenata boliviensis= Hall
Long-tailed Weasel
Plates 28, 29 and 30
Mustela frenata boliviensis Hall, Proc. Biol. Soc. Washington, 51:67, March 18, 1938.
Mustela frenata macrura, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939 (part).
Type.--Male, adult, skull and skin; no. 72587, Amer. Mus. Nat. Hist.; Nequejahuira, 8000 feet, Bolivia; May 19, 1926; obtained by G. H. H. Tate; original no. 4135 (see plates 28-30).
Range.--As now known 8000 to 9500 feet in the Andes from Limbaní, Perú, south to Nequejahuira, Bolivia; upper Subtropical and Temperate life-zones. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts; presence of p2 and two roots rather than one root on P2; from Mustela frenata macrura by darker color of upper parts (tone 4 or darker of Carbon Brown, pl. 342 rather than tone 3 of Chocolate, pl. 343, Oberthür and Dauthenay) and lesser size (in males hind foot less than 45 and m1 less than 5.6); from Mustela frenata agilis by darker color of upper parts (as given above rather than tone 2 of Chocolate, pl. 343, of Oberthür and Dauthenay).
Description.--Size.--Male: The type and two young specimens from Limbaní, Perú, measure respectively, as follows: Total length, 383, 368, 304; length of tail, 140, 132, 115; length of hind foot, 43, 44, 41. Tail 55 per cent as long as head and body. Length of hind foot approximately equal to basal length.
Female: Unknown.
Externals.--As described in Mustela frenata macrura.
Color.--Top of head blackish posteriorly to behind ears; upper lips same color as upper parts of head; dark area at angle of mouth not separated from upper parts as a distinct spot; tip of tail black; remainder of upper parts near (n) Mars Brown of Ridgway and tone 4 or darker of Carbon Brown (pl. 342, Oberthür and Dauthenay); underparts Cream-Colored with strong wash of Ochraceous-Buff; whitish on insides of forelegs to just below elbow; color of underparts extended distally on forelegs over ankles onto antipalmar faces of inner toes, and on hind legs to knees. Least width of color of underparts averages 15 (11-19) per cent of greatest width of color of upper parts. Black tip of tail in type longer than hind foot and amounting to 36 per cent of length of tail-vertebrae.
Skull and teeth.--Male (based on the type): See measurements and plates 28-30. As described in Mustela frenata macrura except that: Weight, 2.8 grams; basilar length, 41.6; zygomatic breadth less than distance between anterior palatine foramen and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors.
Female: Skull unknown.
Remarks.--Apparently the first specimens of this race to find their way into a zoölogical collection were the two young males taken on February 17, 1904, at Limbaní, by Geo. Ockenden (sic).
M. f. boliviensis is smaller than any other South American weasel except possibly agilis. Better material of the two races probably will show even agilis to be larger.
Early in my study of Mustela after examination of the one young specimen, from Limbaní, in the United States National Museum, an account of this race was drawn up, but the account was discarded for want of satisfactory material and the animal was referred to macrura. Then, in 1937, when the two other specimens were studied, the race was formally characterized as different from previously recognized kinds.
The collector has noted on the labels of the two young from Limbaní that they were shot in the afternoon when running together beneath bushes. The frontal sinuses of the type are malformed as a result of infestation by parasites.
Specimens examined.--Total number, 3, as follows:
=Perú=: Carabaya, Limbaní, 2 (one in U. S. Nat. Mus. and one in Berlin Zool. Mus.).
=Bolivia=: Nequejahuira, 1 Amer. Mus. Nat. Hist.
=Mustela frenata (?) gracilis= (Brown)
Plates 39 and 40
Putorius gracilis Brown, Mem. Amer. Mus. Nat. Hist., 9(pt. 4):182, pl. 17, 1908.
Mustela gracilis, Hay, Iowa Geol. Surv. Bull., 23:32, 1914; Hay, Carnegie Instit. Washington Publ. 322A:252, October 15, 1924; Hay, Carnegie Instit. Washington Publ. 390(vol. 2):528, 1930.
Mustela frenata gracilis, Hall, Carnegie Instit. Washington Publ. 473:112, November 20, 1936.
Type.--Adult skull without lower jaws, probably of a female, no. 12431, Amer. Mus. Nat. Hist.; from Conard Fissure, four miles west of Willcockson, Newton County, Arkansas; obtained sometime in the period 1903 to 1905 inclusive. (See plates 39 and 40.)
Range.--Known only from the Pleistocene deposit in Conard Fissure, at the type locality in northern Arkansas.
Description.--Skull. Probably female (based on the type): See measurements and plates 39 and 40; weight unknown; basilar length, 38.1; least width of palate less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of rostrum.
Comparison and remarks.--The type specimen was the only individual referred by Brown (1908) to this species. The remaining material of weasels from this deposit was referred by Brown to his Putorius cicognanii angustidens. Examination of the original materials convinces the writer, too, that the specimens, except no. 12431, are of the species erminea [= cicognanii of Brown]. No. 12431 itself may possibly be erminea but is far more probably of the species frenata. The uncertainty is due to the fact that an occasional skull alone of a subadult male erminea is extremely difficult certainly to distinguish from a skull alone of an adult female frenata. This is true among Recent specimens in the northern Mississippi Valley today; more exactly in Iowa and southern Minnesota the females of frenata, oftentimes intergrades between the subspecies Mustela frenata longicauda, M. f. noveboracensis and M. f. primulina, by only the skulls are next to indistinguishable from certain, unusually slender skulls of male erminea. At other places where the ranges of the two species meet, this difficulty is not so often encountered. Also, the type of gracilis has the skull broken in such a way that the postglenoid length in relation to the length of the skull as a whole could not be accurately determined in this particular skull.
The type specimen of gracilis surely is an adult and because of its small size is thought to be a female. Of known long-tailed weasels of the species frenata, gracilis is structurally nearest to M. f. primulina which occurs in the same region today and to M. f. noveboracensis, the long-tailed weasel of the eastern United States. M. gracilis differs from noveboracensis and agrees with primulina in possessing well-marked temporal ridges which fuse to form a low sagittal crest, in having the mastoid processes projecting farther, laterally, beyond the braincase, in having the anterior ends of the tympanic bullae produced below the squamosal rather than on the same plane with the squamosal, and in having the bullae more inflated anteromedially. M. gracilis differs from both noveboracensis (97 [M] and 56 [F] with skulls of comparable age) and primulina (64 [M] and 24 [F] with skulls of comparable age) in that the zygomatic breadth amounts to less than 58 per cent of the basilar length. Another difference from any one of the skulls of females of primulina is the longer rostrum, which, when measured from the posterior base of the postorbital process of the frontal to the anterior end of the nasal on the same side, amounts to more than 35 per cent of the basilar length. As pointed out by Brown (1908:182) this specimen represents the extreme of slender skull among known kinds of American weasels.
Selected measurements of no. 12431, the type specimen of Mustela gracilis, are as follows: Basilar length of Hensel, 38.1 mm.; length of upper tooth-rows, 14.3 to 14.4; breadth of rostrum, 11.0; interorbital breadth, 8.5; orbitonasal length, 13.6; mastoid breadth, 18.2; length of tympanic bulla, 13.0; breadth of tympanic bulla, 6.3; depth of tympanic bulla, 3.25; outside length of P4, 4.5; inside length of P4, 4.7; breadth of M1, 3.4; length of inner moiety of M1, 1.8; depth of skull at anterior margin of basioccipital, 12.2; depth of skull at posterior borders of last upper molars, 11.3; distance from foramen ovale to tympanic bulla, 3.6 mm.
=MUSTELA AFRICANA= Desmarest
Tropical Weasel
(Synonymy under subspecies)
Type.--Mustela africana Desmarest, Nouv. Dict. d'Hist. Nat., vol. 19, p. 376. 1818.
Range.--Known from the headwaters of the Amazon in eastern Perú and from near the mouth of the same river, on its southern side in Brazil, all within the Tropical Life-zone. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela frenata, the only geographically adjacent species of the genus, by: presence of thenar pad on forefoot; presence of a longitudinal, median, abdominal stripe of same color as upper parts; upper lips being broadly edged, entirely round, with color of underparts; failure of longest facial vibrissae to reach posterior margin of ear; absence of p2; relative flatness (see pl. 29, fig. i and pl. 39, fig. h) of tympanic bullae.
Characters of the species.--Size large (total length of adults approximately 500 mm.); foot-soles naked; thenar pad present on forefoot; length of claws, measured on concave sides, less than one and one-fourth times depth of claws measured at bases; longest facial vibrissae not reaching posterior margin of ear; tail relatively long haired; tail at all ages terminating in point as is characteristic of only juveniles and very young of Mustela frenata and M. erminea; tip of tail, and muzzle, only slightly darker than remainder of upper parts; upper lips broadly edged with color of underparts; pelage coarse, harsh and sparse; longitudinal, median, abdominal stripe of same color as upper parts present; skull broad and deep; braincase large, rounded, and much inflated anteriorly; palatal region wide; tympanic bullae less inflated than in any other American species of the subgenus; angle of lower jaw reduced; dental formula
3 1 2-3 1 -, -, ---, -; teeth heavy; medial lobe of M1 but slightly larger than 3 1 2 2
lateral lobe. See plates 28, 29, 30, 39 and cranial measurements.
Geographic variation.--The reddish versus chocolate color of the upper parts constitutes the only variation of a geographic nature so far detected.
Remarks.--One of the most noteworthy of the several unique characters of this large, tropical weasel is the longitudinal, median, abdominal band. The species exhibits the minimum degree of development of certain features that become progressively less apparent as one proceeds southward from Central America. The relative uniformity of the coloration of the upper parts (reduction in intensity of black color on the muzzle and tip of the tail) and the reduction of the tympanic bullae are two cases in point. Viewed dorsally the general outline of the skull is most nearly matched by that of the skull of Mustela frenata meridana from Venezuela or that of M. f. helleri from Perú. However, the resemblance is not close. The tympanic bullae, although unique among American weasels, are more like those of M. f. meridana from Venezuela than like those of any other kind. The great postorbital width (relatively less in M. africana than in several South American subspecies of Mustela frenata) and small angular process of the mandible are characters, in varying degrees, also common to all South American weasels. Structurally M. africana clearly is more nearly like other subspecies of M. frenata from South America than it is like any species or subspecies from North America.
Mustela africana is the most primitive of the American weasels. The distinctive cranial and dental characters, excepting the reduction in number of premolars, are of a primitive nature. For example, the relatively wide postorbital region, the large braincase that is inflated anteriorly, and the flattened, tympanic bullae, are points of resemblance to the holarctic Mustela erminea, which species is regarded as nearest the original stem form; also the mentioned characters correspond to ontogenetic stages passed through by other weasels. Mostly on these accounts, one is led to look upon M. africana as a migrant from North America. It may have become isolated from its original stock, by a water barrier in the Central American region, for a length of time sufficient to permit of a degree of differentiation to develop between it and the North American weasels which prevented crossbreeding with the frenata stock when that stock, at a later time, reached South America. This assumption is suggested only by evidence from the Recent specimens. No remains of true weasels (subgenus Mustela) have been recorded from deposits in South America older than the Recent period. The alternate possibility, that M. africana intergrades with some race of M. frenata in western or northern South America, has been considered and regarded as highly improbable.
Cabrera (1940:15) has made the distinctive structural characters of Mustela africana basis of the generic name Grammogale to include the one species africana. I am inclined to accord Grammogale only subgeneric rank.
It is possibly significant that Mustela africana is intermediate in several respects between Lyncodon and typical Mustela. The median, longitudinal, abdominal band of the same color as the upper parts in M. africana and the relative uniformity of the coloration of its upper parts might be considered as an intermediate stage between the dark, bicolored (black muzzle and tail tip and brown body) upper parts and light-colored underparts of the North American weasels on the one hand and the light, unicolored upper parts and dark-colored underparts of the Patagonian weasel (Lyncodon) on the other hand. The number of premolars,
2-3 ---, is also intermediate between the numbers 2
3 2 - and - of the North American Mustela and the Patagonian 3 2
Lyncodon, respectively. The American Mustela and the Patagonian Lyncodon, respectively. The more medially, as opposed to anteriorly, directed medial cusp of P4 (characteristic of approximately half of the specimens examined), and the structure of the skull in general of M. africana also seem to be morphologically intermediate between those parts of Mustela and Lyncodon.
On the chance that Lyncodon is closely enough related to Mustela, to be included in a group with Mustela rather than in a group with Grisonella, it is worth noting that Lyncodon lujanensis Ameghino (1889:324, 325), from the villa of Lujan and at the city of Córdoba, at each place in the Pampean [= Pleistocene] formation of Argentine (see also Cabrera, 1928:263) is the first and only fossil form of this group recorded from the whole of South America. Actually, however, Lyncodon seems to me to be as nearly related to Grisonella, if not more so, than to Mustela. If Lyncodon is more closely related to Grisonella and Grison than to Mustela, then the above remarked intermediacy in characters of M. africana has more of interest as a tendency to parallelism than it has of phylogenetic import. Appraisal of phylogenetic relationships would require appraisal of the ancestral stem forms of the Grison stock and the Mustela stock. None of either is known from deposits of the Pliocene, the period of time immediately preceding the Pleistocene.
None of the skulls of Mustela africana seen or figured has the nasomaxillary sutures entirely obliterated and the specimens would, judged on this character alone and by analogy with North American species, be regarded as young and subadult. However, the sutures close at what seems to be a later age than in M. frenata and M. erminea. The condition of the mammae in the type specimen of M. stolzmanni and in the specimen from Moyobamba, indicate that they have borne young. North American weasels old enough to bear young lack visible traces of the nasomaxillary sutures. I have examined no skulls of africana with greatly worn teeth and hence cannot say if the sutures are obliterated in advanced age.
If available data be correct, this species is unique within the genus in that the two sexes are of approximately the same actual size and of the same relative proportions in the body and in the skull. There was no difference between individuals said to be of different sexes from Pará, described and figured by Goeldi (1904:61-62, pls. 1, 2). The undoubted female, type specimen of Mustela africana stolzmanni, is as large as the undoubted male, no. 37475, of the same species, but of a different subspecies, from Pará, Brazil. All the specimens of M. a. africana that I have handled are labeled male and those of M. a. stolzmanni female. More material may show that the female is smaller than the male, as is the case in all near relatives of M. africana.
Little has been recorded concerning the habits of this species. Tate (1931:254) states that a live individual which he saw in a cage at Pará had been captured "swimming in the salt water of the estuary about half a mile away from the shore." On the label of the specimen from Moyobamba, there appears: "caught in Willow tree."
Subspecies examined.--All described forms, of which there are two.
=Mustela africana africana= Desmarest
Tropical Weasel
Plates 28, 29, 30 and 41
Mustela africana Desmarest, Nouv. Diction. d'Hist. Nat., 19:376, 1818; Cabrera, Bol. Real Soc. Españ. de Hist. Nat., 13:429, November, 1913; Cabrera, Bol. Real Soc. Españ. de Hist. Nat., 14:175, pl. 1, March, 1914.
Putorius (Mustela) brasiliensis paraensis, Goeldi, Zool. Jahrb. abt. f. systematik, geogr. u., Biol., 10:556, pl. 21, September 15, 1897, type from Pará, Brazil, near Pará, Ward of Marco da Legoa, Brazil; Goeldi, Bol. do Mus. Paraense, 3:195 [translation of orig. descr.], August, 1901.
Putorius paraensis, Goeldi, Bol. do Museu Goeldi, 4:61, pls. 1, 2, 1904.
Mustela affinis paraensis, Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.
Mustela paraensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:105, April 28, 1916; Tate, Journ. Mamm., 12:253, August 24, 1931.
Mustela stolzmanni paraensis, Hall, Carnegie Instit. Washington Publ. 473:111, November 20, 1936; Hall, Physis, 16:167, pl. 1, figs. 1-4, 1939.
Type.--Male, adult or subadult, mounted; no. 848, Paris Museum; from the "Cabinet de Lisbonne 1808," originally from South America as determined from the characters of the animal; probably came from Brazil, and for the present assumed to be from Pará.
On August 25, 1937, the skull was in the mounted skin and the specimen was in the position shown in the figure published by Cabrera (1914, pl. 1). Except for the loss of the distal part of the tail, and fading because of exposure to light, the specimen was in good condition. See also under remarks.
Range.--Known from the south side of the Amazon River, near its mouth at Pará and Cametá, Río Tocantins, in the Tropical Life-zone of Brazil. See figure 29 on page 221.
Characters for ready recognition.--Differs from Mustela frenata, the only other geographically adjacent species of the genus, in presence of median, longitudinal, abdominal stripe of same color as upper parts and naked foot-soles, in absence of p2 and in reduced size of tympanic bullae (see pls. 29 and 30) and from Mustela africana stolzmanni by lighter color of upper parts which although near Chestnut-Brown are in adults 10' l (darker in yg. M. C. Z., no. 30802), instead of 11' n as in M. a. stolzmanni.
Description.--Size.--This is a relatively large weasel. Goeldi (1897:559) gives the total length of the type specimen of his P. b. paraensis, a female, as 520 mm. (495 in the flesh) and, by computation from his figures, the length of the tail as 200 (205 in the flesh). These measurements probably include the hairs on the tip of the tail as probably also do the measurements given of two other specimens (see Goeldi, 1904:62). One of these specimens, a female, measured: Total length, 520; length of tail, 200. The other specimen, a male, measured: Total length, 510; length of tail, 200. The skin of no. 37475, Amer. Mus. Nat. Hist., a male, has the following measurements written on the attached label: Total length, 548; length of tail, 234; length of hind foot, 56. The hairs project 20 mm. beyond the tip of the last vertebra of the tail and probably are included in the measurements of total length and length of tail. Collectors' measurements of a young male from Cametá, and a subadult labeled as male, from Pará Murutucu, are respectively as follows: 500, 430; 210, 190; 50 and 54.
Externals.--Foot-soles naked, except for a few scattered hairs on ventral sides of interdigital membranes; length of claws, measured on concave sides, not more than one and one-fifth times depth of claws measured at bases; carpal vibrissae not extending beyond apical pad of first digit (not beyond hypothenar pad except in one young specimen); longest facial vibrissae not extending to posterior margin of ear; superior genal tuft not found, hairiness of foot-soles as shown in figure 22.
Color.--Upper parts near (10 l) Chestnut-Brown and relatively uniform since tip of tail and muzzle are only slightly darker than remainder; underparts with longitudinal stripe of same color as upper parts extending along median line of belly from throat or breast posteriorly to within 40 to 50 millimeters of anus. Underparts otherwise near (20" a) Olive-Ocher (lips and chin whiter in one young specimen). Color of underparts extends distally on median sides of forelegs to bare foot-soles and on median side of hind legs two-thirds of distance from knee to ankle. Upper lips broadly edged with whitish, which color passes posteriorly below and not touching eye to ventral margin of concha of ear. An inverted, basally broad, V-shaped extension passes upward 4 millimeters, just posterior to the eye.
Skull and teeth.--See measurements (plates 28-30). Male: (based on 3 adult and subadult topotypes and figures and descriptions published by Goeldi, 1897 and 1904.) Weight, 7.0 grams; basilar length, 45.8 (44.6-47.8); skull broad and deep; braincase large, rounded, and much inflated anteriorly; distance from postorbital process to anterior, nasal notch approximately equal to breadth across exoccipital condyles; palatal region wide; tympanic bullae less inflated than in any other species; mastoid bone, laterally, concave; length of upper tooth-rows in adults and subadults less than breadth of palate measured between two outer margins of fourth upper premolars; alveolar distance between C1 and P4 less than length of P4; teeth heavy; medial lobe of M1 only slightly larger than lateral lobe; deuterocone of P4 heavy and often inclined mesially; p2 absent (P2 present above on both sides in only one of seven specimens seen or described); lower jaw heavy; masseteric fossa not extending anteriorly to posterior fourth of talonid of m1; paraconid of m1 low and base of cleft between it and protoconid relatively low on tooth.
Female: No skull examined but from figures published by Goeldi (1904, pl. 2), apparently as described in the male.
Remarks.--Desmarest in 1818 gave a remarkably good description of this animal which he named as a new species, Mustela africana, but mistakenly indicated that the single specimen known to him came originally from Africa. Until 1913 the name was applied, wrongly, to weasels of northern Africa or to those of the Azores Islands and St. Thomas Island. In that year Cabrera (1913:429) identified the species with the one later named Putorius (Mustela) brasiliensis paraensis by Goeldi (1897:556, pl. 21) from Pará, Brazil. Despite Cabrera's clear identification in 1913, and his later mention of the correct application of the name Mustela africana, it was not correctly employed by other authors, including myself who even as late as 1936 (p. 111) instead used Goeldi's name. In 1937 Mr. Cabrera called my attention to his published account of Mustela africana and so permitted me to examine the type specimen in the Paris Museum, whither I was bound when I received Mr. Cabrera's letter. My own examination of the specimen fully confirmed the conclusions published by Cabrera (1913:429).
As a matter of historical interest, however, it is worth noting that Cabrera (1913) originally supposed the type specimen to have been taken as booty of war from Portugal by the French and that Cabrera later, at the request of P. Trouessart, pointed out (1914:176) that the specimen had been acquired in exchange ("a cambio") since according to Dr. Trouessart the Museum register showed that offer had been made to Portugal to return this and other specimens but that Portugal had replied that it had nothing to reclaim. Dr. P. Rode in August, 1937, at the Paris Museum, gave it to me as his opinion that the specimen had been an outright gift from the "Cabinet de Lisbonne" to E. Geoffroy St.-Hilaire on his trip to Portugal in 1808 when he was given also from the same cabinet several primates, all from Brazil. Of the labels attached to the pedestal on which the specimen is mounted, that of most ancient appearance is glued to the bottom of the stand and bears in a hand apparently written before Trouessart's entries on the same label, the information "Du Cabinet de Lisbonne 1808" and "J. H. S. 1809."
The opened mouth of the mounted specimen permits one to determine that P2 is absent on each side above. The stuffed scrotal pouch and hair projecting downward about the preputial opening clearly show the animal to have been a male. The least faded portions of the mounted specimen, its sides, are of the same reddish color as characterizes adults from Pará and not of the darker chocolate color of specimens of M. stolzmanni from Perú. The specimen is indistinguishable from topotypes of P. paraensis of Goeldi and his name will have to fall as a synonym of Mustela africana Desmarest.
Goeldi gave an extended description, with figures of the skull, head, and entire animal, when he named paraensis. As his account shows, he was unaware that Taczanowski had described a similar weasel from the headwaters of the Amazon, or for that matter that any weasel excepting Mustela affinis Gray, had been found in South America. Goeldi's later account of additional specimens (1904:61, pls. 1, 2) gives much useful information about the animal. Photographs of several specimens and photographs and detailed measurements of several skulls are presented by him.
Pará, and Cametá, Brazil, places from which Mustela africana africana is known, are nearly 2000 miles from the localities in eastern Perú and eastern Ecuador from which M. a. stolzmanni is known, and no specimens, from intermediate localities, are available to show actual intergradation of the two. However, the similarity in structure of the two weasels is so great as to indicate close affinity. Furthermore, it is understood that environmental conditions at and between the two localities are similar. These considerations, in the light of our knowledge of actual intergradation of geographic races of weasels in other places, cause me to treat, with a feeling of assurance, M. africana [= P. paraensis Goeldi] and M. stolzmanni Taczanowski as subspecies of a single species. M. Rodolpho Legueira Rodríguez writes me, under date of June 16, 1928, that the type specimen of Putorius (Mustela) brasiliensis paraensis Goeldi is stuffed and preserved in a "vitrine" at the Museum Goeldi (Museum Paraense) De Historia Natural e Ethnographia, Pará, Brazil.
The one young specimen seen, that from Cametá, is darker colored than any of the four older specimens examined. It is almost exactly the Chestnut Brown of Ridgway (1912) and therefore approaches closely in color the adult specimens of M. a. stolzmanni. This same tendency to greater richness of color in young than in adults is seen also in Mustela frenata.
Specimens examined.--Total number, 5, all from eastern Brazil, as follows: Pará, 2 (1[2], 1[7]); Pará Murutucu, 1[7]; Río Tocantins, Cametá, 1[75]; type specimen, 1[84].
=Mustela africana stolzmanni= Taczanowski
Tropical Weasel
Plates 39 and 40
Mustela stolzmanni Taczanowski, Proc. Zoöl. Soc. London, 1881:835, November 15, 1881; Allen, Bull. Amer. Mus. Nat. Hist., 35:105, April 28, 1916.
Mustela stolzmanni stolzmanni, Hall, Carnegie Instit. Washington Publ. 473:111, November 20, 1936; Hall, Physis, 16:167, pl. 1, figs. 5, 6, 1939.
Type.--Female, adult, mounted skin, with skull separate; no. 563, Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Yurimaguas, Perú; 1880; obtained by J. Stolzmann.
The skull (plate 40), mounted with the skin but removed by me for study, consists of the premaxillae, maxillae, two halves of the lower jaw and dentition. Of these parts, right m2, left coronoid process, right P4 and M1 and adjacent part of maxilla are lost. The skin is well mounted, in a good state of preservation and shows no fading due to exposure to light. Inguinal mammae are distinctly shown on the skin and prove that the specimen is a female. Except for a few scattered hairs on the lower throat, a spot six by eight millimeters on the medial side of the region of the olecranon of the left foreleg and another of similar size in the left axilla, the underparts are, excepting the ventral longitudinal, abdominal stripe, unmarked by color of the upper parts.