SYSTEMATIC ACCOUNTS OF SPECIES AND SUBSPECIES (continued 6)
Reading edition
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 I3 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 P4; 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, M1 #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 M2 and P2 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 page 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 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 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: ♂, 408, 155, 44; ♀, 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[102]. 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[101]. Salt Lake County: Salt Lake City, 1[74]; Barclay, 6500 ft., Wasatch Mts., 1; Mill Creek, 1[101]. 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.