Systematic Accounts of Species and Subspecies (part 3)

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 -) 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 on page .

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 .

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 -. 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; Wapinita, 1; Antelope, 2; 7 mi. E Antelope, 5. Gilliam County: Condon, 1. Morrow County: 10 mi. S Hardman, 1. Umatilla County: Umatilla, 2; 15 mi. E Ukiah, 4000 ft., 1. Union County: Elgin, 1; 20 mi. E Lehman, 1. Wallowa County: Horse Creek, 15 mi. N Paradise, 1; Enterprise, 1; Wallowa Lake, 1; Wallowa Mts., 8300 ft., 1. Baker County: Haines, 1; Anthony, 3; Bourne, 2. Grant County: Long Creek, 1; Canyon Creek, 1; Strawberry Mts., 2; Silvies, 1. Crook County: Prineville, 4. Deschutes County: Sisters, 2; Bend, 1. Lake County: 3 mi. W Stauffer, 1; Fort Rock, 1. Harney County: 25 mi. NW Burns, 1. Malheur County: 4000 ft., Ironside, 2; 1-1/2 mi. S Vale, 2.

Washington. Walla Walla County: Prescott, 4 (2, 1, 1); Ft. Walla Walla, 2 (1); Wallula, 1. Asotin County: Asotin, 1.

Mustela frenata washingtoni (Merriam)

Long-tailed Weasel

Plates , , , , and

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 on page .

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 .

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 -; 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 -; 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; 5935 ft., Glacier Basin, Mt. Rainier, 5 (1); 5051 to 5100 ft., Owyhigh Lakes, Mt. Rainier, 7 (1), Tahoma Creek, 1; Nisqually entrance, 1; Longmire, 1; Mt. Rainier Nat'l Park, 2. Klickitat County: Trout Lake, S Base Mt. Adams, 35; 3500 ft., Gotchen Creek, Mt. Adams, 1.

Mustela frenata saturata (Merriam)

Long-tailed Weasel

Plates , , and

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 -, ) 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 and 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 .

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 -, . 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 , , , , , and

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 on page .

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 .

Color.—Upper parts, in summer, near (n) Argus Brown or tone 4 of Brownish Drab of Oberthür and Dauthenay, pl. 302. Dark spot at each angle of mouth well developed; often fused with color of upper parts which sometimes covers lower lips. Chin white. Remainder of underparts near (14 a´ to 16 c´) Ochraceous-Buff. In winter, upper parts near (14) Argus Brown with smoked effect and Warm Buff to Naples Yellow below. Tip of tail at all times black. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and usually all of wrists, on medial side of hind legs typically only to knee but sometimes to ankle. Tips of toes of hind feet almost always marked with color of underparts. Least width of color of underparts averaging in a series of 14 males from Blaine, Oregon, 23 (14-36) per cent of greatest width of color of upper parts. Black tip of tail in 8 adult males from Blaine, Oregon, averaging 59 (47-70) mm. long; thus longer than hind foot and averaging 37 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on 9 adults from Blaine, Tillamook Co., Oregon): See measurements and plates -; weight, 4.4 (3.3-5.3) grams; basilar length, 45.6 (42.4-47.7); zygomatic breadth more or less (usually more) than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less (usually more) than postpalatal length; postorbital breadth less (except in some instances of malformations of frontal sinuses which result from infestation by parasites) than length of upper premolars and more or less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 to 4 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than orbitonasal length; anterior margin of masseteric fossa directly below m2.

Female (based on 4 adults): See measurements and plates -; weight, 2.2 (2.2-2.3) grams; basilar length, 38.1 (37.8-39.7); zygomatic breadth more or less (less in three of four specimens) than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; relation of postorbital breadth to other measurements in doubt because of malformation of frontal sinuses by parasites; least width of palate not less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3-1/2 to 5-1/2 upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.

Compared with the skull of M. f. washingtoni that of each sex of altifrontalis averages slightly larger in every measurement taken, except measurements of teeth which are approximately the same, and is relatively deeper through the frontal region and through the braincase as measured at the anterior margin of the basioccipital. Skulls of females of altifrontalis have a relatively broader interorbital region. Skulls of males of altifrontalis further differ in having relatively, as well as actually, longer tympanic bullae, relatively lesser orbitonasal length and a greater relative breadth across the mastoids and across zygomata. Compared with M. f. nevadensis, the skull of the male of altifrontalis averages slightly larger and heavier although the skulls of females are of approximately the same size and weight. Relative to the basilar length, the skulls of both sexes are deeper through the braincase and narrower across the mastoids; the rostrum is broader, especially in males; the tooth-rows are shorter and the interorbital breadth less, especially in females. Comparison with the skull of oregonensis is made in the account of that subspecies.

Remarks.—Until the present study was begun, animals of this race have gone under the name Mustela saturata (Merriam). The United States National Museum has a juvenile taken, in 1858, by Wayne at Astoria, O. T.; the Samuel N. Rhoads collection contained one specimen taken in 1891, at Tacoma, Washington; one in the Bangs' collection was taken at Chilliwack, British Columbia, in 1895, and the Field Museum has one taken on the Olympic Peninsula in 1898. The best material is that collected by Alex Walker, at Tillamook, Oregon.

Intergradation with nevadensis is indicated by several specimens. The coloration of the one adult female, no. 90, Chas. R. Conner Mus., from Swamp Creek, Washington, has the color of the underparts extended down the hind legs over the feet, and over the proximal third of the ventral face of the tail as in nevadensis although the other two specimens from the same place have the color pattern of altifrontalis. Of the four specimens from British Columbia referred to this subspecies, only the specimen from Chilliwack is typical as regards color pattern. The one from Cultus Lake has the color pattern of nevadensis and might be referred to that race almost as well as to altifrontalis. The two specimens from Lihumption Park are intermediate between the two races in tone of color. Neither has the color of the underparts extended onto the tail or continuously over the hind feet as in nevadensis but each does have the color of the underparts less restricted and of lighter hue than in altifrontalis. Only one of the specimens, no. 7848 Canad. Nat. Mus., from Lihumption Park is adult and it has a skull which agrees with that of altifrontalis rather than nevadensis.

After writing the above, a good representation of the weasel population along the eastern side of Puget Sound was made available by friends in that area. Study of the weasels from there shows that their color is intermediate between that of altifrontalis and nevadensis. On the whole, they (specimens from Bellingham, for example) resemble one subspecies about as much as the other. In cranial characters some specimens, in certain features, approach nevadensis but most specimens agree with altifrontalis and all are more nearly like altifrontalis to which race all are referred.

The color of these animals is to me indistinguishable from that of washingtoni. The color of washingtoni is merely intermediate between that of nevadensis and altifrontalis. Nevertheless, the race washingtoni has cranial characters (long narrow skull) which set it off from both altifrontalis and nevadensis. This shape of skull is not found in the specimens from along the eastern side of Puget Sound; these animals have skulls like that of altifrontalis and when departures from this occur they are in the direction of nevadensis and not washingtoni.

The above, then, explains why specimens which are colored like those of washingtoni are not referred to that race but instead to the race altifrontalis.

Of 23 adult skulls examined, 19 have the frontal sinuses malformed as the result of infestation by parasites.

Specimens examined.—Total number, 80, arranged within states by counties from north to south. Unless otherwise indicated specimens are in the United States National Museum.

British Columbia. Chilliwack, 1, Lihumption Park, 4750 ft., 2; Cultus Lake, 1.

Oregon. Clatsop County: Old Fort Clatsop, 1; Astoria, 1. Tillamook County: Tillamook, 12 (7, 2, 2, 1); Netarts, 1; Blaine, 16 (13, 1, 1, 1). Lane County: Reed, 1; Mercer, 1. Curry County: Langlois, 1.

Washington. Whatcom County: Nooksack River, 2000 ft., 14 mi. E Glacier, 1; Swamp Creek, 2050 ft., Nooksack River, 3; Lookout, 4800 ft., Mt. Baker, 2; Bellingham, 8; 5 mi. S Bellingham, 1. Skagit County: Rockport, 300 ft., 1. King County: Bothell, 2; N Seattle 1; Seattle, 1; Tye, 1, 2 mi. E Skykomish, 1; 7 mi. E Kent, 1; Auburn, 3. Pierce County: Tacoma, 1. Clallam County: Sequim, 1; Soleduc Riv., near [sic.] Sappho, 1; Happy Lake, 1; mouth of Boulder Creek, Elwha River, 560 ft., Olympic Mts., 1; Hume's Ranch, 1000 ft., Elwha River, 1; Bogachiel Riv., 1. Mason County: Lake Cushman, 2; 4 mi. N Shelton, 1. Thurston County: Olympia, 2; Tenino, 1. Pacific County: 2-1/2 mi. SE Chinook, 3.

Mustela frenata oregonensis (Merriam)

Long-tailed Weasel

Plates , , , , , and

Putorius xanthogenys oregonensis Merriam, N. Amer. Fauna, 11:25, June 30, 1896; Bangs, Proc. New England Zoöl. Club, 1:57, June 9, 1899.

Mustela xanthogenys oregonensis, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.

Mustela xanthogenys munda, Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933 (part).

Mustela frenata oregonensis, Hall, Carnegie Instit. Washington Publ. 473:107, November 20, 1936.

Type.—Male, subadult, skull and skin; no. 32019/43828, U. S. Nat. Mus., Biol. Surv. Coll.; Grants Pass, Rogue River Valley, Josephine County, Oregon; December 19, 1891; obtained by C. P. Streator; original no. 1404.

The skull (plates -, ) is complete and unbroken. P3 on the left side is missing. Otherwise the teeth all are present although worn probably as a result of gnawing at the trap which captured the specimen. The skin, in brown, winter pelage, is fairly well made.

Although the label on the skin and the label in the skull vial each give the sex of the specimen as female, and although Merriam (1896:25) regarded the specimen as a female, the present writer regards the specimen as a male.

It is as large as other undoubted males and larger than any known female of this subspecies. The labels with the skull and skin give the locality as "Rogue River Valley, Oregon." The listing here of the more restricted locality, Grants Pass, is made on the basis of Merriam's (1896:25) original description of the subspecies.

Range.—Transition and Canadian life-zones along coast of northern California and southern Oregon from Humboldt County, California, north through Curry County, Oregon, thence inland, west of the Cascades, north to the Columbia River. See figures 29 and 30 on pages 221 and 314.

Characters for ready recognition.—Differs from M. f. altifrontalis in presence of frontonasal white patch, lighter color above (tone 2 to 3 of Raw Umber, pl. 301, rather than tone 4 of Brownish Drab, pl. 302, Oberthür and Dauthenay), wider extent of light color of underparts which is extended distally beyond knee, and in females, longer tooth-row which amounts to more than 38 per cent of basilar length; from M. f. munda in shorter hind foot of males which is less than 50, and in both sexes, smaller, less rugose skull (see measurements and plates); from M. f. saturata in presence of frontonasal white patch, in having color of underparts extended uninterruptedly over ankle onto foot; from M. f. nevadensis in presence of frontonasal white patch, lack of light color of underparts on ventral face of tail, and longer skull, which relative to its length in males, is shallower through braincase; from M. f. effera in presence of frontonasal white patch, lack of light color of underparts on ventral face of tail, and larger skull with basilar length averaging more than 41.7 in males; from M. f. washingtoni in presence of frontonasal white patch, shorter skull in males, which in percentage of basilar length has, on the average, orbitonasal length amounting to less than 35, mastoid breadth more than 55, and zygomatic breadth more than 63; and in females larger skull with least width of palate more than length of P4, upper tooth-rows more than 38-1/2 per cent of basilar length, bullae larger and averaging more than 13.4 long.

Description.—Size.—Male: Five males (3 adults and 2 subadults from Eureka, Ferndale, and Carlotta, California) yield average and extreme measurements as follows: Total length, 392 (347-430); length of tail, 138 (110-160); length of hind foot, 46 (43-50). Tail averages 54 (46-61) per cent as long as head and body. Length of hind foot more or less than basal length. The type specimen, and an adult from Goldbeach measure, respectively, as follows: Total length, 412, 386; length of tail, 155, 137; length of hind foot, 44, 46.

Female: Three adults (2 from Fortuna and 1 from Carlotta, California) yield average and extreme measurements as follows: Total length, 367 (360-374); length of tail, 130 (123-134); length of hind foot, 40 (39-40). Tail averages 55 (52-57) per cent as long as head and body. Length of hind foot less than basal length. A subadult from Goldbeach, an adult from 13 mi. SW Grants Pass, and an adult from Medford, measure, respectively, as follows: Total length, 316, 344, 294; length of tail, 114, 120, 122; length of hind foot, 36, 40, 38.

The average differences in external measurements of the two sexes in the vicinity of Carlotta, are: Total length, 25; length of tail, 8; length of hind foot, 6. Corresponding differences, at Goldbeach, are: 70, 23, 10. Probably the females at Fortuna reflect the large size of munda more than do the males at Carlotta and the differences between the measurements of the two sexes probably, therefore, are actually more than are indicated by the figures above.

Externals.—Longest facial vibrissae black, brown or white (often all three colors in same specimen) and extending beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles, in summer pelage, as shown in figure .

Color.—Upper parts, in summer, near (16 l) Brussels Brown or tone 2 of Raw Umber of Oberthür and Dauthenay, pl. 301, to slightly darker than tone 3 of same plate. Darker on nose and top of head, usually with frontonasal white patch but lacking white bar in front of each ear, except in the type and 2 specimens from Salem. Chin, lower lips, angle of mouth, and usually posterior seventh of upper lip white. Remainder of underparts Pale Orange-Yellow. In winter usually lighter above with underparts Warm Buff to Straw Yellow. Tip of tail at all times black. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial side of hind leg, typically over ankle in extremely narrow line which widens out over distal phalanges of antiplantar faces of toes but sometimes interrupted at ankle. Least width of color of underparts averaging, in twenty available specimens, 39 (27-54) per cent of greatest width of color of upper parts. Black tip of tail in five adults averaging 50 (43-60) mm. long; thus averaging longer than hind foot and 33 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on 4 adults and subadults from Eureka, Requa, Goldbeach, and Grant Pass): See measurements and plates -, . As described in Mustela frenata nevadensis except that: Weight, 3.5 (3.5-4.1) grams; basilar length, 42.9 (41.8-44.0); least width of palate more or less than medial length of P4.

Female (based on 2 adults, one from Carlotta and one from 13 mi. SW Grants Pass): See measurements and plates -. As described in Mustela frenata nevadensis except that: Weight, 2.4 (2.2-2.6) grams; basilar length, 37.7 and 39.5; zygomatic breadth less than distance between condylar foramen and M1 and less than distance between anterior palatine foramen and anterior margin of tympanic bulla. See under "Remarks" for additional data on variation in size of skulls of females.

The skulls of the female averages 31 per cent lighter than that of the average male.

Because there is much geographic variation between specimens here referred to oregonensis, the person who is guided by the present account should keep in mind that results, here reported, of comparisons of the skull with those of other races, were obtained by employing specimens of oregonensis from Carlotta and Eureka, California. These specimens from California are judged to have more of the characters of the subspecies munda than do specimens of oregonensis from more northern localities.

Compared with that of M. f. washingtoni the skull of the male is shorter, especially in the preorbital region and is relatively broader across the mastoidal processes and zygomatic arches. The skull of the female is longer in the preorbital region, has a less cylindrical braincase and differs less from the male skull than is the case in M. f. washingtoni. Compared with M. f. effera, the skull of the male is smaller in every part measured and relative to the basilar length is broader across the mastoids and has relatively shorter tympanic bullae. From M. f. nevadensis the skull of the male differs in the same way except that size is about the same. The skull of the female oregonensis is more heavily ridged and is relatively broader across the mastoids than that of effera. From M. f. saturata, oregonensis is not surely known to differ in cranial characters. From M. f. munda, oregonensis differs in having the skull of both sexes smaller, and on the average, in all parts measured, has a less marked postorbital constriction, relatively narrower interorbital region and relatively more expanded zygomata. From M. f. altifrontalis, males of oregonensis differ on the average, in having larger teeth, and relative to the basilar length, a greater mastoid breadth and a shallower braincase as measured at the anterior margin of the basioccipital. Females of oregonensis differ in larger average size of skull, except for breadth of rostrum and interorbital breadth which, therefore, are relatively less in oregonensis, as also is the relative depth of the skull measured at the posterior borders of the upper molars and at the anterior margin of the basioccipital. However, skulls of females of oregonensis have relatively longer tooth-rows and are relatively broader across the zygomata and mastoidal processes.

Remarks.—In 1896, Merriam named oregonensis as a subspecies of the California bridled weasel on the basis of a single specimen taken by Clark P. Streator. Three additional specimens were acquired in later years, by workers of Dr. Merriam's bureau, from near the type locality and specimens from farther north in Oregon have been accumulated at the University of Oregon. The most satisfactory material is that saved from Humboldt County by the late H. E. Wilder, which, when brought together, is adequate to give some idea of the range of variation that can be expected in a given population.

Of two specimens from Goldbeach, one shows approach to altifrontalis in that the color of the underparts stops at the ankle, and in one, the angle of the mouth is dark colored. Specimens from Eugene and vicinity lack the white facial markings, and in this feature approach the adjoining washingtoni-effera-nevadensis stock. A specimen from 6 miles south of Medford shows approach to saturata in the interruption, on the ankle and lower tibial region, of the color of the underparts. One adult female, no. 1413, Univ. Oregon, from the Rogue River Valley, 13 miles southwest of Grants Pass, stands out prominently, among the other specimens from extreme southern Oregon and northwestern California, by reason of the near (18) Apricot Yellow color of the underparts, but this same color occurs in specimens from the more northerly localities of Buchanan, Eugene, Vida Fish Hatchery, and McKenzie Bridge, as well as in no. 2178, Univ. Oregon, from Cresent Lake. The last mentioned specimen is here referred to nevadensis.

Two females referred to oregonensis from southern Oregon differ so greatly in size of skull that they challenge one's imagination in any attempt to provide an explanation for so wide a range of variation in one subspecies. One of these, no. 244520, U. S. Nat. Mus., is an adult female from Medford. The other, no. 224034, U. S. Nat. Mus., is a subadult female (though labeled male) from 43 miles northeast of Grants Pass. The skull of the adult from Medford has a basilar length of 41.5, upper tooth-rows, 16.1 in length, and a weight of 2.75 grams, whereas corresponding figures for the subadult are only 33.8, 12.9, and 1.4. Two other adult females are intermediate in size: No. 1413, Univ. Oregon, from 13 miles southwest of Grants Pass, Oregon, approaches the specimen from Medford in size, and the second specimen, no. 34325, Mus. Vert. Zoöl., from Carlotta, California, is smaller.

Not only is there a difference in length between the skulls of the two extremes of the females but this difference extends to all other dimensions of their skulls, and is most pronounced in the preorbital region. The differences in breadth of the braincase and other parts of the skull are relatively less than the differences in length. Differences of the same nature, although of lesser degree than found in the females, are to be seen in two males. The skull of an adult no. 51590, Mus. Vert. Zoöl., from 6 miles south of Medford, has a basilar length of 46.4, upper tooth-rows, 17.6 mm. long, and a weight of 4.0 grams, whereas corresponding figures for the subadult type specimen from Grants Pass, are only 43.0, 16.2, and 3.3.

The wide range of variation in size of skull of both sexes, together with the considerable variation in color pattern of the specimens here referred to oregonensis raises the suspicion that we are using the name in a composite sense; nevertheless, to recognize more than one subspecies with the material now available would be unwise.

A subadult female, of abnormal color, no. 47149, Mus. Vert. Zoöl., taken by Mr. H. E. Wilder at Carlotta, California, on December 20, 1930, in a region where weasels do not turn white in winter, is white, except for the black tip of the tail, but has a suffusion of orange. This specimen, discussed at greater length on page , is instructive in that it suggests that there are separate determiners for the brown and red elements of the pelage. It is interesting also as suggesting how natural selection may tend to eliminate from the population a conspicuous color-variation of this kind. At any rate, Mr. Wilder (Ms.) states: "This specimen was picked up in a field, where it evidently had been dropped by a hawk or an owl." The braincase of the skull is crushed in three places as though by a raptor's beak. None of the several other weasels, all normally colored, saved by Mr. Wilder from this general locality gives evidence of having fallen a victim to a raptor.

Only 2 skulls of the 12 adults and subadults examined show malformation of the frontal sinuses such as results from the presence of parasites.

Specimens examined.—Total number, 29, arranged within states from north to south by counties. Unless otherwise indicated specimens are in the collection of the United States National Museum.

California. Del Norte County: Requa, 1. Humholdt County: Eureka, 2 (1, 1); Ferndale, 1; Fortuna, 2; Carlotta, 6 (3, 3); 12 mi. E Bridgeville, 1; 2 mi. W Bridgeville, 1.

Oregon. Washington County: Forest Grove, 1. Marion County: Salem, 2. Benton County: Buchanan, 1. Lane County: McKenzie Bridge, 1; Vida Fish Hatchery, 1; Eugene, 1. Douglas County: Anchor, 1. Curry County: Gold Beach, 2. Josephine County: Rogue River Valley (Grants Pass), 1; 13 mi. SW Grants Pass, 1. Jackson County: Medford, 2; 6 mi. S Medford, 1.

Mustela frenata munda (Bangs)

Long-tailed Weasel

Plates , , , , , , , , , and

Putorius xanthogenys mundus Bangs, Proc. New England Zoöl. Club, 1:56, June 9, 1899; Stephens, California mammals, p. 247, 1906.

Mustela frenata, Audubon and Bachman, Journ. Acad. Nat. Sci. Philadelphia, 8 (Pt. 2):291, 1842 (North California about 40° latitude).

Mustela xanthogenys munda, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.

Mustela frenata munda, Hall, Carnegie Instit. Washington Publ. 473:107, November 20, 1936.

Type.—Male, adult, skull, os penis and skin; no. 5459, collection of E. A. and O. Bangs, but now in collection of Mus. Comp. Zoöl.; Point Reyes, Marin County, California; June 19, 1896; obtained by C. A. Allen; original no. 931. (See comments under "Remarks," below, on places in California to which the name Point Reyes has been applied.)

The skull (pls. 19-21, 30) is complete and unbroken. I1 on each side and right I2 are broken away; p2 and p3 on each side have been aborted and the only alveoli remaining are two for the right p3. Otherwise all teeth are present and entire. The skin is fairly well made and in good condition.

Cranially, the type is a "runt"; its small size and the circumstance that the tympanic bulla is longer than the lower molar and premolar tooth-row and longer than the rostrum are features which differentiate the type from any other specimen seen of this race.

Range.—Sea level to at least 6,000 feet (South Yolla Bolly Mountain, Trinity County, California); Upper Sonoran and Transition life-zones of the coast and Coast Range of northwestern California from the Golden Gate northward into southern Humboldt and Trinity counties. See figures and on pages 221 and 314.

Characters for ready recognition.—Differs from M. f. oregonensis in longer hind foot of males which is more than 50 mm., and in both sexes, larger, more prominently ridged skull (see measurements and plates); from M. f. saturata by presence of nasofrontal white spot, larger and relatively shallower skull of males and larger skull of female; from M. f. nevadensis by presence of well-developed, white, facial markings; absence of color of underparts on ventral face of proximal third of tail; and hind foot of males more than 50; from M. f. xanthogenys by near (l) Sudan Brown to near (l) Antique Brown rather than Buckthorn Brown colors of upper parts and greater size, and in adult male basilar length more than 45 and hind foot more than 47; from M. f. nigriauris by having inside of ears same color as back rather than much darker than back.

Description.—Size.—Male: Three adults and two young from Point Arena and Gualala, Mendocino County, yield average and extreme measurements as follows: Total length, 447 (434-470); length of tail, 167 (150-185); length of hind foot, 53 (50-60). Corresponding measurements of three adults from 5 and 6 miles west of Inverness, Marin County, are: 430 (420-440), 154 (141-160), 48 (48-49). Corresponding measurements of four individuals (3 adults and 1 young of large size) from South Yolla Bolly Mountain, Trinity County, are: 383 (374-400), 134 (130-138); 44 (43-44). The tail averages 60 per cent as long as the head and body in the series from Point Arena, 56 per cent in the series from Point Reyes, and 53 per cent in the series from South Yolla Bolly Mountain. In every specimen except two, length of hind foot less than basal length. The two exceptions are no. 19720, M.V.Z., male adult from Point Arena in which the hind foot is recorded as 60 (probably an error in measurement), and no. 19721, M.V.Z., from the same place, in which the skull has not yet attained its full growth.

Female: One adult from Point Arena measures as follows: Total length, 383; length of tail, 134; length of hind foot, 43. Corresponding measurements of an adult from seven miles north of Laytonville, Mendocino County, are: 336, 121, 33 (= 36 on dried skin). Corresponding measurements of an adult from South Yolla Bolly Mountain, Trinity County, are, 326, 113, 37. In these three specimens, the tail is, in the order given, 54, 56, and 53 per cent as long as the head and body. Length of hind foot more than basal length.

Differences in external measurements of the two sexes as indicated by the five males and one female from Point Arena, are: Total length, 64; length of tail, 33; length of hind foot, 10. Weights of 2 adult males are 265 and 221 grams and of one adult female 155 grams.

Externals.—As described in Mustela frenata nigriauris.

Color.—Spot between eyes, narrow band or spot confluent with color of underparts on each side of head anterior to each ear, chin, lower lips, and rarely posterior third or less of each upper lip white; dark spot posterior to each angle of mouth uniformly present and of large size; tip of tail black; remainder of upper parts near (14 l) Sudan Brown and tone 4 of Raw Umber of Oberthür and Dauthenay, pl. 301; occasionally, slightly darker brown on forehead, nose, and about eyes. Underparts near (a to c) Ochraceous-Buff and sometimes Orange-Buff. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind limbs over antiplantar faces of toes. Least width of color of underparts averaging, in a series of 5 males from Mendocino County, 57 (46-67) per cent of greatest width of color of upper parts; 38 (35-40) in 3 males from Point Reyes, Marin County. Black tip of tail in Mendocino County series averaging 53 (46-60) mm., which is same length as hind foot and 32 per cent of length of tail. In Point Reyes males, black tip of tail averages 44 (34-52) mm., which is less than length of hind foot and 45 per cent as long as tail-vertebrae.

Several specimens of the smaller, inland variant (see under "Remarks") are near (l) Antique Brown rather than near (14 l) Sudan Brown above and hence do not differ in this respect from nigriauris.

Skull and teeth.—Male (based on 3 adults from Mendocino County): See measurements and plates -, . As described in Mustela frenata nigriauris except that: Weight, 6.0 (5.4-6.3) grams; basilar length, 47.6 (46.5-48.2); length of tympanic bulla more than length of lower molar and premolar tooth-row.

Female (based on no. 19723, M.V.Z., from Point Arena): See measurements and plates -, . As described in M. f. nigriauris except that: Weight, 3.0 grams; basilar length, 42.3.

The skull of the female is 50 per cent lighter than that of the average male.

Compared with the skull of the male of nevadensis that of munda averages larger in every part measured and specimens from Point Arena are nearly as heavy again, have relatively more expanded zygomata and mastoid processes but are relatively narrower anteriorly as shown by the breadth of the rostrum, interorbital breadth and postorbital breadth. Also the braincase is less inflated anteriorly, the tympanic bullae are lower and the skull is more angular. Females show the same differences although in different degree. Compared with the skull of the male of M. f. nigriauris, that of munda from Point Arena averages larger in every part measured except for the length of the upper tooth-rows. Relative to the basilar length, the skull of munda averages broader across the mastoids and across the zygomata, is deeper through the braincase at the anterior end of the basioccipital, and has a greater development of the lambdoidal crest.

Remarks.—The skin and part skull, no. 536/1849, U. S. Nat. Mus., taken by Lieutenant W. P. Trowbridge at San Pablo Bay, is the first specimen known to have been saved of this subspecies. Since 1899 when O. Bangs diagnosed munda as of small size, the weasel of the humid costal belt north of San Francisco Bay has been regarded as smaller than bridled weasels from farther south in the State. Actually, however, the weasel of the humid costal belt shares with M. f. pulchra the distinction of being one of the two largest weasels in California.

M. f. munda may be a composite subspecies, for the variation in facial markings, in coloration otherwise, in external measurements and in size and shape of skull is great. At one time in the course of the present study, manuscript accounts of two subspecies were prepared for the animals now all called munda and there is still much justification for recognizing two subspecies, one, along the coast proper, the larger, darker-colored animal with reduced white facial markings and large, wide, heavily ridged skull from Point Arena, and 6 miles south of Laytonville, Mendocino County, along with the specimens from 5 and 6 miles west of Inverness, Marin County, and the other, an inland race, which is a smaller, lighter-colored animal with more extensive white facial markings and a smaller, narrower, skull, known by specimens from Point Reyes [station?], Nicasio, 15 mi. north of San Rafael, Freestone, Vallejo, and Mount Sanhedrin. The differences between these two lots of specimens are of great degree. However, a female from Fort Bragg proves to be no larger than three females labeled as from Point Reyes. Also, a male from 2 miles south and one mile east of Stewarts Point on the coast has a skull no larger than the animal from Vallejo, whereas the skin alone of an adult female from 3 miles south of Stewarts Point is large and agrees with the specimens from Point Arena. Consequently, no logical ranges can be worked out for the two variants with the material now available.

Finally, the type specimen of munda is a "runt," smaller than any other male seen. This specimen, purchased by E. A. and O. Bangs from C. A. Allen, who collected and sold specimens widely, was labeled as from Point Reyes. So far as this place-name is concerned, it might refer to: (1) The point of land by that name which projects out into the Pacific Ocean, (2) an abandoned ranch house bearing that name at the head of Drakes Bay, 6 miles north and 3-3/4 miles east of the actual point, or (3) the railway station by the same name at the head of Tomales Bay, 12 miles east and 4-3/4 miles north of the actual point. Allen, himself, lived near San Geronimo (then Nicasio) about nine miles southeast of the Point Reyes railway station. All these places are in Marin County, but differ markedly as regards climate and flora. The first two are treeless, windswept and have much fog, whereas Point Reyes Station is more often sunny, and is situated in a shallow valley, inland, where the open grass-covered west-facing slopes meet the east-facing wooded ones. From which one of these three places the type specimen came, I do not know. The same may be said of the three female specimens labeled Point Reyes; two of these are in the United States National Museum and one in the Field Museum.

The specimens in the Museum of Vertebrate Zoölogy from 5 and 6 miles west of Inverness and those from near the same place in the collection of John Cushing come from within a couple of miles or less of the Point Reyes represented by the abandoned ranch house. These specimens, as remarked above, agree with those from Point Arena in large size, reduced facial markings and wide skull. These are points of difference from the smaller variant suspected of being a recognizable subspecies. It is the smaller variant which the type specimen approaches in size, and with which it agrees in relatively well-developed white facial markings. This suggests that the type specimen came from Point Reyes Station rather than from either of the two other places bearing the name "Point Reyes," from one of which, as just stated, the variant of large size is known. The three females labeled "Point Reyes" also have well-developed white facial markings and are of lesser size than the female of similar age from Point Arena, Mendocino County. The presumption is that these three females also came from Point Reyes Station.

The smaller, inland variant seems to agree in size, cranial characters, and coloration with M. f. nigriauris to the southward of San Francisco Bay, but lacks the black on the head which characterizes nigriauris. The larger variant, on which the description here used for munda is based, comprises animals which differ from nigriauris in larger size, darker color, reduced white facial markings, and larger, relatively wider skull. Both of the variants mentioned above are sharply distinct from nigriauris on the basis of coloration of the inside of the ear which is blackish in nigriauris like the dark facial markings, and in munda is colored like the back. M. f. munda lacks the dark facial markings; an occasional specimen has at most, a trace of the markings but this does not extend back so far as the ears. This difference, blackish versus non-blackish face, persists eastward of San Francisco Bay to at least as far as the Carquinez Straits, where a specimen of munda is available from 4 miles north of Vallejo and one of nigriauris from Glen Frazer Station on the south shore opposite Vallejo.

Intergradation with M. f. nevadensis and possibly with M. f. saturata is indicated by specimens from South Yolla Bolly Mountain, Trinity County. In them the external measurements and measurements of the skull are intermediate. Also the white frontal spot is much reduced in size. The white bars in front of the ears are absent in three specimens, and weakly developed in the other two. The relative proportions of the skulls as a whole are nearer those of nevadensis or saturata than munda. The skull of one of the three adult males and the skull of the adult female suggests M. f. oregonensis in certain features; for example, the dorsal outline of the skull in longitudinal axis is slightly convex as it is in oregonensis.

None of the specimens shows malformation of the frontal sinuses such as results from infestation by parasites.

Specimens examined.—Total number, 37, arranged by counties from north to south. Unless otherwise indicated specimens are in the Museum of Vertebrate Zoölogy.

California. Trinity County: S. Yolla Bolly Mt., 3; 1/2 mi. S S. Yolla Bolly Mt., 1. Tehama County: 2 mi. S S. Yolla Bolly Mt., 1. Mendocino County: 6 mi. N Laytonville, 1; Mt. Sanhedrin, 1; Ft. Bragg, 1; Gualala, 1; Point Arena, 5. Sonoma County: 2 mi. S and 1 mi. E Stewarts Point, 1; 3 mi. S Stewarts Point P. O., 1; Freestone, 1. Napa County: 6 mi. SSW, Napa, 1; 4 mi. N Vallejo, 1. County in question: San Pablo Bay, 1. Marin County: 6 mi. W Inverness, 2; 5 mi. W Inverness, 2(1); Point Reyes, 4 (2 1, 1); Nicasio, 2 (1, 1); Kehoes Ranch, Pierce Point, 1; Drakes Bay, 1; Tomales Point, about 1/2 mi. SW White Gulch, 1; Point Reyes School, 3-3/4 mi. W Inverness, 1; 15 mi. (by road) N San Rafael, 1; Hurley Ranch, 2 mi. W Tomales, 1. No locality more definite than California, 1.

Mustela frenata xanthogenys Gray

Long-tailed Weasel

Plates , , , , , , and

Mustela xanthogenys Gray, Ann. and Mag. Nat. Hist., 11:118, 1843.

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Putorius xanthogenys, Merriam, N. Amer. Fauna, 11:25, June 30, 1896; Bangs, Proc. New England Zoöl. Club, 1:56, June 9, 1899.

Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.

Mustela frenata xanthogenys, Hall, Carnegie Instit. Washington Publ. 473:107, November 20, 1936.

Type.—Male, adult, skull and skin; skull no. 197a-43.6.4.55, skin no. 234a-42.11.21.4, British Museum (Nat. Hist.); from the bank of Sacramento River below mouth of Feather River, or from north shore of San Francisco Bay, California; taken in "1837 or 1838"; presented by Captain Edward Belcher.

The skull (plate ) lacks the occiput, the right mandible posterior to m1, and the right pterygoid; the right zygomatic arch is fractured. The teeth are not greatly worn. The skin was originally mounted for exhibition (R. I. Pocock in Litt.) but in 1937 when I saw the skin, it was prepared as a conventional study skin. The skin is in fairly good condition; some hair is missing on the hind quarters and the skin of the tail is torn at one place.

Range.—Altitudinally, less than 600 feet (Fair Oaks); Lower Sonoran and Upper Sonoran life-zones of all but southern end of the San Joaquin Valley, and probably Sacramento Valley, California. See figures and on pages 221 and 314.

Characters for ready recognition.—Differs from M. f. nevadensis by presence of light facial markings and Buckthorn Brown rather than near (14n to l) Brussels Brown color of upper parts; from M. f. munda by Buckthorn Brown rather than near (l) Sudan Brown, or near (l) Antique Brown color of upper parts and lesser size, in adult males basilar length less than 45 and hind foot less than 47; from M. f. nigriauris by lighter color in same way as from munda and also by having inside of ears same color as back rather than much darker than back; from M. f. pulchra in hind foot of males less than 46 and narrower skull, in males having breadth of rostrum less than 13.9 and mastoid breadth less than 26.0, see comparison of skulls in the account of pulchra.

Description.—Size.—Male: Three adults, from Fresno, Selma and Los Banos, measure, respectively as follows: Total length, 425, 417, 450; length of tail, 152, 154, 180; length of hind foot,—, 43, 44. Tail averages 61 per cent as long as head and body. Length of hind foot less than basal length.

Female: Adults from Selma, Los Banos, and 4 mi. SW Turlock, measure respectively as follows: Total length, 357, 365, 395; length of tail, 133, 132, 145; length of hind foot, 40, 38, 41. Tail averages 58 per cent as long as head and body. Length of hind foot less than basal length.

The average differences in external measurements between the two sexes, as represented by these six specimens, are: Total length, 65; length of tail, 25; length of hind foot, 3.5. One adult male weighs 274 grams and 2 adult females 182 and 214 grams.

Externals.—As described in Mustela frenata nigriauris.

Color.—Spot between eyes, band confluent with color of underparts on each side of head extending anterodorsally anterior to each ear, and posterior half to third of each upper lip white, or whitish tinged with some shade of yellowish; chin and lower lip white; dark spot posterior to each angle of mouth of varying size but uniformly present; tip of tail black; remainder of upper parts Buckthorn Brown of Ridgway or a trifle browner than tone 4 of Brown Pink of Oberthür and Dauthenay, pl. 297. Upper parts of uniform color except for slight darkening of head-markings anterior to ears. Underparts Ochraceous-Buff to Warm Buff. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, on medial sides of hind limbs over antiplantar faces of toes and sometimes tarsal region. Least width of color of underparts averaging, in 9 specimens from Fresno, Selma and Los Banos, 54 (32-74) per cent of greatest width of color of upper parts. Black tip of tail in three males (one subadult and 3 adults) averages 55 (50-60) mm. long. Thus longer than hind foot and averaging 34 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on 2 adults from Fresno and one from Selma): See measurements and plates -, . As described in M. f. nigriauris except that: Weight 3.8 grams; basilar length, 43.7 (43.4-43.9); least width of palate more or less than lateral length of P4; length of tympanic bulla more than length of lower molar and premolar tooth-rows.

Female (no. 2626 W. E. Snyder, from Selma): See measurements and plates -. As described in M. f. nigriauris except that: Weight, 2.5 grams; basilar length, 39.4.

The skull of the female is 34 per cent lighter than the average for the three males.

Compared with skulls of nevadensis from the Sierra Nevada, those of the two adult males from Fresno differ as follows: M1 wider (transversely); tympanic bullae narrower; preorbital part of skull smaller. Comparison with pulchra is made in the account of that subspecies. Compared with skulls of adult males of nigriauris, from Santa Clara County, the two skulls from Fresno are generally smaller and in basilar length, length of tooth-rows and measurements of the teeth fall below the minimum for nigriauris. Relative proportions of the skulls are approximately the same. Comparison with munda reveals essentially the same differences as does comparison with nigriauris except that the difference in size is greater.

Remarks.—The name Mustela xanthogenys Gray was long applied to all the weasels of the interior valleys of California and of the coast of that state south of San Francisco Bay. Gray, when he named the species and when referring to it in later accounts, never defined the locality whence the specimen came more definitely than "California." In 1896, Merriam (1896:25) gave the type locality as "Southern California, probably vicinity of San Diego" and later writers have not contradicted him. The type specimen was obtained in the course of the voyage of the British ship Sulphur, under command of Sir Edward Belcher. Examination of Belcher's (1843, vol. 1, p. 129) narrative of the voyage indicates the following places in California at which the specimen of weasel, described by Gray, could have been obtained: Fort Ross, Bodega, vicinity of San Francisco Bay and up Sacramento River to the mouth of the Feather River, Monterey, Santa Barbara, Buenaventura, San Pedro, San Juan, and San Diego.

Reginald I. Pocock has kindly compared the type specimen in the British Museum with several specimens sent for that purpose. In the first place, comparison of skulls shows that the type specimen is a member of one of the races north of San Diego. In the second place, comparison of skins shows that the inside of the ears are not blackish but similar in color to the back. In fact, Pocock writes under date of February 12, 1929, regarding the type specimen, that "It is practically uniformly colored from the snout to the base of the tail, there being scarcely a trace of the darkening of the head, or muzzle, observable in your specimens [those sent for comparison]." This character of coloration of the ear excludes all the weasels of the Coast region of California from San Francisco Bay southward, namely, M. f. latirostra and M. f. nigriauris. My own examination of this type specimen at a date later than that on which Pocock compared it satisfies me as to the accuracy of his statement above.

Accordingly, the name xanthogenys would seem to apply to one of the two subspecies here called munda and xanthogenys. Perusal of Belcher's narrative of the voyage (loc. cit.) shows that little, if any, opportunity was afforded to obtain vertebrate specimens at Fort Ross or Bodega, both localities within the range of the subspecies here called munda. Furthermore, the type specimen is smaller than individuals of munda from 5 to 6 miles west of Inverness and from Point Arena with which the animals from Fort Ross and Bodega would be expected to agree in size. Weasels from along the north shore of San Francisco Bay are smaller than those on the coast north of the bay. Possibly the type specimen of xanthogenys came from the north side of San Francisco Bay but probably it came from the bank of the Sacramento River and almost certainly not farther up stream from San Francisco Bay than the junction of the Sacramento and Feather rivers. The statement of Belcher (1843, vol. 1, p. 129), regarding the trip up the Sacramento River as far as Point Victoria, lat. 38°46´47" north, and return to San Francisco Bay, that "Cuyote or jackal—fox, racoon, land otter, weasel, and squirrel were obtained" lends strong probability to the idea that this type specimen was taken along the Sacramento River, possibly in the vicinity of the existing city of Sacramento. Unfortunately no specimens are available from the Sacramento Valley. If some were available, a comparison of them and specimens of munda from along the north side of San Francisco Bay and Carquinez Straits with the type specimen of xanthogenys should determine the correct application of the name. For the present it seems best to retain the name munda and apply the name xanthogenys to the weasels inhabiting the northern part of the San Joaquin Valley and presumably the southern part of the Sacramento Valley.

Efforts to obtain specimens of weasels from the Sacramento Valley have been in vain. A juvenal specimen taken five miles south of Fair Oaks, Sacramento County, by Mr. John Fitzgerald, Jr., in December, 1927, was examined at his home and found to agree in coloration with specimens from farther south. Geographically, this specimen probably is more nearly a topotype than any other examined.

Most of the specimens examined are immature and adequate adult cranial material has not been seen. Two adults, one of each sex, from Los Banos have skulls of large size which agree with those of nigriauris. The same is true of one adult and one young female from 4 miles southwest of Turlock, which, unlike the animals from Los Banos, show a darkening of the head extending in reduced degree even to the inside of the ears, as in nigriauris. The slightly darker than average (for xanthogenys) color on the back may indicate intergradation with nevadensis. Intergradation with M. f. nevadensis is shown by specimens, from the southern part of the Sierra Nevada, mentioned in the account of nevadensis.

None of the skulls shows malformation of the frontal sinuses such as result from infestation by parasites.

Specimens examined.—Total number 30, arranged by counties from north to south.

California. Sacramento County: Bank of Sacramento River, 1; 5 mi. S Fair Oaks, 1. San Joaquin County: 4 mi. W Stockton, 1. Merced County: Tegner School, 4 mi. SW Turlock, 2; Los Banos, 4 (2, 1 1). Fresno County: Mendota, 1; Biola, 1; Clovis, 1; Fresno, 5 (1, 1, 2, 1); 5 mi. W Fresno, 1; Selma, 3 (2, 1); 4 mi. NW Sanger, 1; 5 mi. S Selma, 1. Tulare County: Monson, 1; 1-1/2 mi. N Goshen, 1; Milo, 1; 2 mi. N Tipton, 1; Poplar, 2. No locality more definite than California, 1.

Mustela frenata nigriauris Hall

Long-tailed Weasel

Plates , , , , , and

Mustela frenata nigriauris Hall, Carnegie Instit. Washington Publ. 473:95, November 20, 1936.

Putorius xanthogenys, Baird, Mamm. N. Amer., 1858, p. 176 (part).

Mustela xanthogenys Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):375, 1874 (part?).

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Putorius xanthogenys xanthogenys, Grinnel, Proc. California Acad. Sciences, fourth series, 3:292, August 28, 1913.

Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912; Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933.

Type.—Male, adult, skeleton and skin; no. 32820, Mus. Vert. Zoöl.; Half Moon Bay, San Mateo County, California; received at Museum of Vertebrate Zoölogy, May 4, 1922, through A. L. Hagedoorn, after having been in captivity a few days where death occurred owing to injuries received in trap; original no. 1590.

The skull has each of the zygomatic arches and the anterior end of the nasals broken through. The only part missing is the central two millimeters of the left zygomatic arch. The teeth all are present and entire. The skeleton appears to be complete except for the bones of the feet, which are preserved within the skin. The skin is well made and in good condition.

Range.—Altitudinally, sea level to more than 4000 feet; Sonoran and Transition life-zones of Coast Range and coast of California from San Francisco Bay south to Point Conception, Santa Barbara County, California. See figures and on pages 221 and 314.

Characters for ready recognition.—Differs from M. f. munda, xanthogenys, and pulchra by having inside of ears darker than back rather than same color as back, and from xanthogenys and pulchra in near (l) Antique Brown color of upper parts rather than Buckthorn Brown or near (16 j) Buckthorn Brown to near (h) Yellow Ocher respectively; from M. f. latirostra by postorbital breadth, of adult males and females, less, rather than more, than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite.

Description.—Size.—Male: Five adults from Palo Alto, Santa Clara County, yield average and extreme measurements as follows: Total length, 447 (412-465); length of tail, 167 (147-175); length of hind foot, 46 (45-47). Corresponding measurements of four adults from San Francisco are: 412 (394-435); 153 (145-160); 43.5 (41-46). Corresponding measurements of five adults and subadults from Berkeley, Alameda County, are: 419 (390-448); 148 (135-160); 44 (42-47). Tail averages 59 per cent as long as head and body in series from Palo Alto and in one from San Francisco. The average of 55 for the Berkeley series probably reflects a lesser average age. Length of hind foot less than basal length. The type specimen measures, 415, 150, 43. It is smaller than the mean.

Female: A subadult from Palo Alto measures: Total length, 368; length of tail, 126; length of hind foot, 39. An adult and two subadults from Berkeley measure, respectively, as follows: Total length, 347, 365, 340; length of tail, 134, 123, 125; length of hind foot, 37, 38.4, 36.5. In these four females the tail averages 55 per cent as long as head and body. Length of hind foot less than basal length.

The average differences in external measurements of the two sexes, as represented by specimens from Berkeley, Alameda County, are: Total length, 68; length of tail, 21; length of hind foot, 7. Eight adult males weigh 249 (217-335) grams and one adult female 123 grams.

Externals.—Longest facial vibrissae brownish like dark color of head and extending beyond ear; carpal vibrissae mostly color of underparts and extending to apical pad of fifth digit; hairiness of foot-soles slightly more than shown in figure .

Color.—Spot between eyes, band, confluent with color of underparts, on each side of head extending anterodorsally anterior to ear, and posterior third of each upper lip tinged with color of underparts or, less often, pure white; chin and lower lips white; remainder of sides and top of head posteriorly to, or a little behind, a line connecting posterior margins of ears, blackish; inside of pinna of ear, and sometimes outside of pinna, blackish; dark spot posterior to each angle of mouth present on each side in three-fourths of specimens; tip of tail black; remainder of upper parts near (l) Antique Brown, and with more yellow than tone 3 of Raw Umber of Oberthür and Dauthenay, pl. 301. Often with more blackish and red in winter. Underparts near (a to c) Ochraceous-Buff or Ochraceous-Salmon. Ochraceous-Salmon in some juveniles. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists, and on medial sides of hind limbs over antiplantar faces of toes. Least width of color of underparts averaging, in 17 adult males (Berkeley, 5; San Francisco, 5; Palo Alto, 7), 55 (40-73) per cent of greatest width of color of upper parts. Black tip of tail in same series of males averaging 51 (35-60) mm., thus averaging longer than hind foot and 33 per cent of length of tail (Palo Alto and San Francisco, 31 per cent; Berkeley, 35 per cent). In 8 adult females, least color of underparts amounts to 55 (47-62) per cent of greatest width of color of upper parts. Black tip of tail averages 41.5 (28-50) mm., thus averaging longer than hind foot and 32 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on six adults from Stanford Univ. and vicinity): See measurements and plates -; weight (four adults), 5.4 (5.0-5.9) grams; basilar length, 47 (46.1-48.1); zygomatic breadth more than distance between condylar foramen and M1, or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more than postpalatal length; postorbital breadth less than length of upper premolars (less than distance between posterior borders of P4 and P2) and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate less than lateral length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 or 4 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more or less than (about equal to) length of lower molar and premolar tooth-row and longer or shorter (usually shorter) than rostrum; anterior margin of masseteric fossa below anterior half of m2.

Female (based on three adults, Hayward, Palo Alto, and Morro): See measurements and plates -; weight (no. 43574, from Morro) 2.7 grams; basilar length, 41.2 (40.2-42.2); zygomatic breadth more or less than distance between condylar foramen and M1 and more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than lateral length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.

The skull of the female averages 50 per cent lighter than that of the average male.

Comparisons of the skull of the male with those of M. f. latirostra, pulchra, xanthogenys, and munda are made in the accounts of those subspecies.

Remarks.—Like M. f. latirostra, nigriauris long bore the name xanthogenys. The fairly adequate lot of specimens is divided between the collections of several institutions. The most satisfactory material in any one collection is in the Stanford University Natural History Museum where local specimens have been accumulated over a period of many years.

No actual intergrade between nigriauris and xanthogenys has been seen, although the specimens from Los Banos, referred to xanthogenys, have large skulls as in nigriauris. Intergradation with latirostra is shown by specimens, referred to latirostra, from the Los Angeles area. Also the one adult male from 5 miles southeast of Santa Margarita, San Luis Obispo County, is of small size and in this respect approaches latirostra. The range of nigriauris is separated from that of munda by San Francisco Bay, Carquinez Straits, and I suppose by the lower part of the San Joaquin River. On the basis of color of the inside of the pinna of the ear, the two subspecies are uniformly distinct. Intergradation is assumed to occur through the subspecies xanthogenys.

None of the 26 adult and subadult specimens examined for evidences of infestation of the frontal sinuses by parasites shows malformation of the sinuses.

Specimens examined.—Total number, 103, arranged by counties from north to south. Unless otherwise indicated specimens are in the Museum of Vertebrate Zoölogy.

California. Contra Costa County: Glen Frazer Station, 1; 2 mi. W Pinole, 1; 1 mi. E Pinole, 1; Richmond, 1; Lafayette, 1; 7 mi. E Clayton, 1; Moraga Valley, 1; Pinehurst, Redwood Canyon, 1; Concord, 1. Alameda County: Berkeley, 11; Oakland, 1; Piedmont, 1; Haywards, 2; near Haywards, 2; 10 mi. E Haywards, 1; Redwood Canyon, 1; Calaveras Dam, 1. San Francisco County: San Francisco, 11 (5, 2, 1, 1); Ocean View, 1; Visitation Valley, 1. San Mateo County: Moss Beach, 1; Half Moon Bay, 1; Redwood City, 1; Menlo Park, 9 (5, 2); no locality more definite than county, 1. Santa Clara County: 1/4 mi. N Milpitas, 1; 1/4 mi. S Milpitas, 1; Stanford University, 6 (4, 2); Palo Alto, 11 (6, 2, 1, 1). Santa Cruz County: 3 mi. E Santa Cruz, 1; 2-1/2 mi. E Santa Cruz, 1; Santa Cruz, 6 (2, 1, 1). Monterey County: 1 mi. E mouth Salinas River, 10 ft., 1; Pacific Grove, 1; Monterey, 2 (1); Carmel, 1; Carmel Valley, 1; Point Lobos, 1; Gonzales, 1. San Luis Obispo County: 5 mi. SE Santa Margarita, 1; Morro, 1; 3-1/2 mi. S Oceano, 6. Santa Barbara County: Santa Maria, 1; 5 mi. N Las Cruces, 1; 7 mi. W Gaviota, 1; Gaviota, 1.

Mustela frenata latirostra Hall

Long-tailed Weasel

Plates , , , , , and 36

Mustela frenata latirostra Hall, Carnegie Instit. Washington Publ. 473:96, November 20, 1936.

Putorius xanthogenys, Baird, Mamm. N. Amer., p. 176, 1858 (part); Stephens, California mammals, p. 246, 1906; Merriam, N. Amer. Fauna, 11:25, June 30, 1896 (part).

Putorius (Gale) brasilianus frenatus, Coues, Fur-bearing animals, p. 142 (part).

Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912; Grinnell, Univ. California Publ. Zoöl., 40:102, September 26, 1933.

Mustela arizonensis, Grinnell and Swarth, Univ. California Publ. Zoöl. 10, 376, October 31, 1913.

Type.—Male, adult, skull and skin; no. 3257, Mus. Vert. Zoöl.; San Diego, San Diego County, California; May 20, 1907; obtained by Frank X. Holzner.

Right M1 is missing and the part of the jaw bearing this tooth is broken away. With this exception the skull is complete and unbroken and the teeth are all present and entire. The skin is fairly well made and in good condition except that it is slightly soiled.

Range.—Altitudinally sea level to 8000 feet (Tahquitz Valley, San Jacinto Mountains); Sonoran and Transition life-zones of coast and mountains west of Mohave and Imperial deserts of southern California from Point Conception and Cuyama Valley southward at least to Mexican boundary. See figures and on pages 221 and 314.

Characters for ready recognition.—Differs from M. f. nigriauris by having postorbital breadth of adult males and females, more, rather than less, than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; from M. f. pulchra by having tympanic bulla longer than rostrum (orbitonasal length) and by near (l) Antique Brown rather than near (16 j) Buckthorn Brown to near (h) Yellow Ocher color of upper parts.

Description.—Size.—Male: Six adults and subadults from San Diego yield average and extreme measurements as follows: Total length, 439 (428-449); length of tail, 153 (142-160); length of hind foot, 45 (40-47). Corresponding measurements for a series of eight adult males from the vicinity of Los Angeles are: 416 (394-428); 158 (151-166); 44 (40-47). In the series from San Diego the tail averages 54 per cent as long as head and body. In the series from Los Angeles the average is 61 per cent. Length of hind foot in each series, less than basal length. The type specimen measures, 435, 142, 42.

Female: No. 5070, adult, from San Diego, measures 367, 141, 38. Nos. 22 and 6748 from Santa Ysabel, measure: 359, 380; 130, 140; 39, 35. No. 7194 from Jamacha measures, 358, 125, 35. Three adult females from Los Angeles yield the following: Total length, 373, 345, 368; length of tail, 150, 122, 134; length of hind foot,—, 41, 41. In no. 5070 the tail is 62 per cent as long as the head and body and in the three from Los Angeles it averages 60 (55-67) per cent. Length of hind foot, in each case, less than basal length.

The average differences in external measurements of the two sexes as shown by the six males from San Diego and the four females from San Diego County are: Total length, 73; length of tail, 19; length of hind foot, 8. Corresponding differences shown by the eight males and three females from Los Angeles are: 54, 23, 3.

Externals.—Longest facial vibrissae brownish, like dark color of head and extending beyond ear; carpal vibrissae mostly color of underparts and extending to apical pad of fifth digit; hairiness of foot-soles slightly more than shown in figure .

Color.—Spot between eyes, band confluent with color of underparts on each side of head extending anterodorsally anterior to ear, and posterior third of each upper lip tinged with color of underparts or, less often, white; chin and lower lips white; remainder of sides and top of head posteriorly to near line connecting posterior margins of ears, blackish; inside of pinna of ear, and sometimes outside of pinna, blackish; dark spot posterior to each angle of mouth present on each side in three-fourths of specimens; tip of tail black; remainder of upper parts near (l) Antique Brown, and with more yellow than tone 3 of Raw Umber of Oberthür and Dauthenay, pl. 301. Underparts Ochraceous-Buff to Warm Buff and in some specimens from Los Angeles and Ventura counties Ochraceous-Orange, especially in young and juveniles. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists and on medial sides of hind limbs over antiplantar faces of toes. Least width of color of underparts averaging, in 15 adult and subadult males from San Diego County, 54 (35-75) per cent of greatest width of color of upper parts. Black tip of tail in same series of males averaging 54.5 (46-60) mm. long. Thus averaging longer than hind foot and 35 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on 6 adults from San Diego County). See measurements and plates -. As described in M. f. nigriauris except that: Weight (4 specimens), 3.9 (3.8-4.0) grams; basilar length 43.8 (41.9-47.0); postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2 to 2-1/2 (including I3) upper incisors; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum; anterior margin of masseteric fossa below m2.

Female (based on 4 adults from San Diego County): See measurements. As described in M. f. nigriauris except that: Weight, 2.6 (2.2-2.8) grams; basilar length, 40.0 and 40.1; postorbital breadth more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; length of tympanic bulla more than length of rostrum.

The skull of the female averages 34 per cent lighter than that of the average male.

The skull of the male of latirostra, compared with that of nigriauris, is by weight, more than one-fourth lighter, has a lesser basilar length, a lesser mastoid breadth, a lesser zygomatic breadth and a narrower M1. In these features no overlap has been observed between adults from the general vicinities of the type localities of the two forms. In adult males of latirostra the postorbital breadth, with one exception, is more than the combined length of P4 and P3 whereas the reverse is true in adult males of nigriauris. Both males and females of latirostra have a generally smaller skull with relatively broader interorbital and postorbital parts and the tympanic bullae are relatively larger, rounder and more inflated.

Compared with the skull of the male of pulchra that of latirostra is, by weight, more than one-fourth lighter, has a lesser basilar and orbitonasal length, lesser zygomatic and mastoid breadth and a more nearly flat braincase. In these features no overlap has been observed between adults from the general vicinities of the type localities of the two subspecies. Also, in latirostra the tympanic bulla is longer than the rostrum whereas the opposite is true in pulchra. The skull of latirostra is generally smaller and relatively, on the average, has the preorbital part of the skull deeper and broader with longer tooth-rows, although with shorter rostrum, while the zygomatic and mastoid breadths are less. Study of skulls of subadult females of pulchra indicate that females of latirostra and pulchra differ in the same fashion as do males.

Remarks.—This subspecies long has gone by the name M. xanthogenys and the type locality was generally supposed to be in the vicinity of San Diego. This supposition seems to have originated with Merriam's (1896:25) statement that the type locality was "Southern California, probably vicinity of San Diego." Nevertheless, as set forth in the account of M. f. xanthogenys the type specimen concerned now is thought to have come from much farther north.

Although 76 Recent specimens are available from southern California, additional adults are needed to understand the geographic variation there. M. f. latirostra may be a composite—made up of more than one geographic race. Specimens from San Diego County differ so much in relative length of the tail that at one stage in the present study it was thought that a difference in this respect existed between the coastal animals and those from farther inland. Material received later did not wholly substantiate this view and because of the uniformly small size of all of the skulls from that county, the animals were later regarded as of the same subspecies. Eventually, even this supposed common feature proved to be inconstant for an adult male from Jamacha, no. 7098, of the San Diego Society of Natural History, and another adult male from San Marcos, no. 8869, collection of Ralph Ellis, were later examined and found to have skulls as large as those of average-sized, adult males of nigriauris.

Despite these puzzling local variations, it is established that the long-tailed weasels of southern California are smaller than those from farther north. Also, the southern animal averages smaller in weight and size of skull, and the skull is differently proportioned. Specimens in series from Los Angeles County definitely are intermediate in size and shape of skull between latirostra from San Diego County and nigriauris from, say, Santa Clara County, but definitely more closely resemble latirostra from San Diego County than they do nigriauris. A skull of a young animal, not here identified to subspecies, from Potholes, in the Colorado River Valley, 10 miles northeast of Bard, Imperial County, California, may have closest relationship to M. f. latirostra. Additional comment on this specimen is offered in the account of M. f. neomexicana.

From the asphalt pits of Rancho La Brea, in Los Angeles County, a total of 57 skulls have been examined, more than half of which are reasonably complete. I have been unable to learn whether these came from pits regarded by students of the deposit as wholly Recent, from pits regarded as of Pleistocene age, or from both. Suffice to say that only two specimens were found which could be distinguished from skulls of the subspecies of weasel living in that area today.

These two specimens, lent to me by Professor Chester Stock, were with other skulls received from the Los Angeles Museum of History, Art and Science and bore identifying numbers as follows: 16/20-27, the anterior part of the skull of an adult, and 16, the skull posterior to the cribiform plate of a subadult or possibly young individual. The latter has a mastoid breadth of 28.0 millimeters, a tympanic bulla 16.1 long and other measurements in proportion. It is larger than any specimen of weasel, of any subspecies, seen from California and in the subgenus Mustela seems to be exceeded in size only by certain individuals of M. f. texensis. M. f. neomexicana attains relatively large size and comparisons were made with individuals of that subspecies. However, the young specimen from Rancho La Brea differs from neomexicana in that the tympanic bullae rise less steeply on the medial sides and the inferior lip of the external auditory meatus is less developed laterally. Age considered, the sagittal crest is less developed and the mastoid processes project more abruptly from the skull. The anterior part of the skull of the adult, no. 16/20-27 is larger than any specimen seen of M. f. latirostra or adjoining subspecies, and among California-taken specimens is equaled in size only by the largest males of M. f. munda from the northwest coastal district in Mendocino County. This adult from Rancho La Brea differs from neomexicana, sex and age taken into account, in greater postorbital breadth, lesser rostral width in comparison with the interorbital breadth, and in having the temporal ridges at the anterior end of the sagittal crest spread out into a Y-shaped, rather than a T-shaped, pattern. All these differences from neomexicana are features of agreement with the California bridled weasels of the subspecies latirostra, nigriauris, and munda. The same is true of the characters which set apart the young specimen from neomexicana. In summary: of 57 weasel skulls examined from the asphalt pits at Rancho La Brea, Los Angeles County, all but two are indistinguishable from the skulls of the Recent weasel living in that region today. These two skulls agree in qualitative characters with animals of the California coastal subspecies now living from Los Angeles northward to Humboldt County, but are larger. For the time being these two may be thought of as giants of the same type of animal inhabiting the Los Angeles region today.

Only one of 41 adult and subadult skulls examined for malformation of the frontal sinuses shows infestation by parasites.

Specimens examined.—Total number, 142, listed by counties from north to south. Unless otherwise indicated specimens are in the Museum of Vertebrate Zoölogy.

California. Santa Barbara County: Rincon Point, 1. Ventura County: Cuyama Valley, 2200 ft., 1; Nordhoff, 3; Santa Paula, 1; Ventura, 7. Los Angeles County: near Owensmouth, 1; Cahuenga, 1; Llano, 10 mi. E Littlerock, 1; Flint Ridge, Pasadena, 1; Pasadena, 3; Lankershim, 1; 1 mi. S Lankershim, 1; Duarte, 1; Covina, 1; Claremont, 1; El Monte, 4 (2, 1); Montebello, 1; Alhambra, 6 (5, 1); El Nogal, 2; Gardena, 1; Palos Verdes Estate, 3; Rancho La Brea asphalt deposits, 57 and . San Bernardino County: San Bernardino Valley, 1; San Bernardino, 4 (2, 1); Redlands, 2 (1); Bluff Lake, 2 (1, 1). Riverside County: West Riverside, 1; Arlington, 800 ft., 1; 3-1/2 mi. E and 1/2 mi. N Beaumont, 2600 ft., 1; Banning, 1; Cabazon, 1; San Jacinto Plain, 1; Tahquitz Valley, 8000 ft., 1; Elsinore, 1. San Diego County: Twin Oaks, 1; San Marcos, 2 (1, 1); Escondido, 1; Witch Creek, 1; Ballena, 1; Santa Ysabel, 3 (2, 1); Julián, 1; La Jolla, 1; Lakeside, 1; El Cajon, 1; El Vido (not found on map), 1; San Diego, 9 (1, 1, 1, 1); Jamacha, 2; Chula Vista, 1.

Mustela frenata pulchra Hall

Long-tailed Weasel

Plates , and

Mustela frenata pulchra Hall, Carnegie Instit. Washington Publ. 473:98, November 20, 1936.

Type.—Male, adult, skeleton and skin; no. 16668, Mus. Vert. Zoöl.; Buttonwillow, Kern County, California; April 30, 1912; obtained by J. Grinnell; original no. 1953.

The skull (plates -) is complete and unbroken (a fracture in the right jugal has healed). All teeth are present and entire. The skeleton lacks the os penis, left fibula, shaft of left tibia and the distal three or four caudal vertebrae. Some of the bones of the feet distal to the radius and tibia are with the skeleton, and the remainder probably are in the skin. The skin is fairly well made and in good condition, except for the left hind leg which was torn when the animal was captured. A well-developed scrotal pouch shows the specimen to have been a male.

Range.—Altitudinally around 300 feet in San Joaquin Valley to 2500 feet at Isabella; Upper Sonoran and Lower Sonoran life-zones of southern end of San Joaquin Valley and in mountains at southern end of Valley, California. See figures and on pages 221 and 314.

Characters for ready recognition.—Differs from M. f. nevadensis in presence of light facial markings, and from M. f. nevadensis and M. f. inyoensis in near (16 j) Buckthorn Brown to near (h) Yellow Ocher rather than near (14 n to l) Brussels Brown color of upper parts, and greater size with hind foot more than 40 in females and basilar length averaging more than 46.0 in males; from M. f. latirostra in having rostrum (orbitonasal length) longer than tympanic bulla and from M. f. latirostra and M. f. nigriauris by color of upper parts as stated above rather than near (l) Antique Brown, and by having inside of ears same color as back rather than much darker than back; from M. f. xanthogenys in hind foot of males more than 46 and broader skull which in males has breadth of rostrum more than 13.9 and mastoid breadth more than 26.0.

Description.—Size.—Male: The type specimen and five other adults yield average and extreme measurements as follows: Total length, 454 (428-477); length of tail, 178 (153-184); length of hind foot, 50 (47-55). Tail averages 65 per cent as long as head and body. Length of hind foot approximately equal to basal length. The type specimen measures, 460, 184, 49.

Female: Three subadult topotypes yield average and extreme measurements as follows: Total length, 399 (383-411); length of tail, 154 (140-161); length of hind foot, 42 (41-42). Tail averages 63 per cent as long as head and body. Length of hind foot less than basal length.

The average differences in external measurements of the two sexes are: Total length, 55; length of tail, 24; length of hind foot, 8.

Externals.—As described in Mustela frenata nigriauris.

Color.—Spot between eyes, band confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, posterior third of each upper lip, lower lips and chin white or more often darker than Ochraceous-Buff and therefore same color as belly; dark spot posterior to each angle of mouth present but small; tip of tail black; remainder of upper parts near (16 j) Buckthorn Brown to near (h) Yellow Ocher and from tone 2 to 4 of Brown Pink of Oberthür and Dauthenay, pl. 297, but with a trifle more reddish brown. Upper parts of uniform color except for occasional slight darkening of nose, forehead, and areas around eyes. Underparts darker (a) than Ochraceous-Buff. Color of underparts extends distally on posterior sides of forelegs over toes, onto antipalmar faces of feet and wrists, on medial sides of hind limbs over antiplantar faces of toes, tarsal region and sometimes in diluted fashion on proximal third of underside of tail. Least width of color of underparts averaging, in 6 male topotypes, 55 (43-81) per cent of greatest width of color of upper parts. Black tip of tail in same series of males averaging 58 (53-63) mm. long; thus averaging longer than hind foot and 33 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on 6 ads., type and 5 topotypes): See measurements and plates -. As described in M. f. nigriauris except that: Weight (6 ads.), 5.3 (4.5-6.1) grams; basilar length, 47.6 (46.0-48.6); (one skull, no. 335, with postorbital breadth more than distance between posterior borders of P4 and P2); interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 2 to 3-1/2 (including I3) upper incisors; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than rostrum.

Female: Adult skull of typical female not seen.

As compared with the skull of the type specimen of inyoensis, skulls of adult males of pulchra are larger throughout, relatively broader, especially in the preorbital part of the skull, have more inflated tympanic bullae, and are less convex in dorsal outline. Comparison of the skull with that of latirostra has been made in discussion of that subspecies. Comparison of skulls of adult males of nigriauris and pulchra shows that those of pulchra average larger in every measurement taken except those of m1, M1, P4, and depth of skull at posterior borders of upper molars. The basilar length is only slightly more and it follows that, relative to this length, other measurements of the skull are relatively, as well as actually, larger. In no one measurement is there an entire lack of overlap, but the skulls of adult males, and probably adult females, may be distinguished from those of nigriauris by the combination of the following mentioned, average differences: Tympanic bullae larger in each of three dimensions; preorbital and interorbital parts of skull broader and notably heavier; interorbital breadth greater; zygomatic arches more expanded laterally; mastoid processes more prominent. As compared with xanthogenys, differences of similar nature, but of greater degree, distinguish pulchra. As compared with those of nevadensis (represented by specimens from Mono Co., Calif.), skulls of adult males of pulchra average larger in every measurement taken and no overlap exists in basilar length, orbitonasal length, mastoid breadth, zygomatic breadth, length of tympanic bulla, or depth of skull at either the anterior margin of the basioccipital or at the posterior margins of the upper molars. Relatively, the preorbital portion is about the same size in the two forms.

Remarks.—The best material of this big weasel was obtained in 1910 and 1911 by John Wimmer and forwarded to the California Academy of Sciences through John R. Rowley, although in 1905, one specimen had been obtained by A. S. Bunnell for the collections of the United States Bureau of Biological Survey, another by J. Grinnell for the Museum of Vertebrate Zoölogy in 1912, and in 1933, another by L. M. Huey, for the San Diego Society of Natural History.

The males from the type locality are relatively uniform in size and shape of skull. The one exception is no. 137935, U. S. Nat. Mus., slightly younger than the others. Its skull is relatively more slender than any of the others and does not display several of the differential characters. The male, no. 127566, U. S. Nat. Mus., from Alila (= Earlimart) is intermediate in cranial features between pulchra and xanthogenys as known from specimens taken in the vicinity of Fresno. The skull of the female, no. 127565, from the same place, is too young to provide diagnostic characters. Since the skull of an adult female of topotypical pulchra is unknown, doubt attaches to the identification of the adult, female specimen, no. 115895, U. S. Nat. Mus., from Delano. It has a relatively broad skull in comparison with the adult female of xanthogenys from Los Banos. The adult female, no. 9998, San Diego Soc. Nat. Hist., from 2 mi. SW Simmler, shows approach to nigriauris in slightly reduced size. The skin alone from Coalinga, a male, taken on April 10, 1935, measures 462, 179, 47. The adult female, with crushed skull, from 4 miles east of Coalinga, measures 350, 129, 40. In size, these specimens agree better with pulchra than with xanthogenys. The skin alone from 3 miles south of Coalinga is unsexed and without external measurements. Skulls of adults from Coalinga are needed to permit of more positive identification of the subspecies found there. The female from 4 miles east of Coalinga, taken on February 21, 1936, is in process of molt on the underparts, and the longer hair which is near (20´) Naples Yellow contrasts strongly with the incoming shorter hair which is near (10 c) Salmon-Orange. The skin alone, no. 16270, Mus. Vert. Zoöl., from Isabella, was made up from a decayed animal and is of but little use. It is referred to pulchra purely because of geographic nearness of Isabella to the type locality of pulchra. The most that can be told from the specimen is that it is a relatively light-colored, bridled weasel. The fact that the color is slightly darker than in pulchra may or may not indicate intergradation with nevadensis. No. 54103/41042, U. S. Nat. Mus., consisting of crushed bits of skull and the skin of the head, is from Willow Spring, Kern County. This marginal locality is really in the Mojave Desert rather than in the San Joaquin Valley. The light color of the skin of the head suggests pulchra, but it is realized that a complete specimen might show the animal there to be unlike pulchra.

None of the skulls shows evidence of having had the frontal sinuses infested by parasites.

Specimens examined.—Total number, 18, listed by counties from north to south. Unless otherwise indicated, specimens are in the Museum of Vertebrate Zoölogy.

California. Fresno County: Coalinga, 1; 4 mi. E Coalinga, 1; 3 mi. S Coalinga, 1. Tulare County: Alila (= Earlimart), 2. Kern County: Delano, 1; Buttonwillow, 9 (6, 2); Isabella, 1; Willow Spring, 1 San Luis Obispo County: 2 mi. SW Simmler, 1.

Mustela frenata inyoensis Hall

Long-tailed Weasel

Plates , and

Mustela frenata inyoensis Hall, Carnegie Instit. Washington Publ. 473:99, November 20, 1936.

Putorius xanthogenys, Merriam, N. Amer. Fauna, 11:25, June 30, 1896 (part).

Mustela xanthogenys xanthogenys, Miller, U. S. Nat. Mus. Bull., 79:99, December 31, 1912.

Type.—Male, adult, skull (with skeleton) and skin; no. 25907, Mus. Vert. Zoöl.; Carl Walter's Ranch, 2 mi. N Independence, Inyo County, California; June 26, 1917; obtained by A. C. Shelton; original no. 3143.

The skull (plates -) is complete and unbroken. All teeth are present and entire. The skin is well made and in good condition.

Range.—From 3700 feet (Lone Pine) to at least 4000 feet (Alvord); Lower Sonoran Life-zone of the floor of Owens Valley in Inyo County, California. See figures and on pages 221 and 314.

Characters for ready recognition.—Differs from M. f. nevadensis in presence of white facial markings; from M. f. pulchra in near (l) Brussels Brown rather than near (16 j) Buckthorn Brown to near (h) Yellow Ocher color of upper parts and basilar length of less than 45 in males; from M. f. latirostra in brownish rather than blackish color of inside of ear and orbitonasal length of more than 15.

Description.—Size.—Male: Two adults, the type specimen and no. 25392/32805, measure, respectively, as follows: Total length, 423 and 390; length of tail, 170 and 145; length of hind foot, 42 and 44. Tail is 67 and 59 per cent as long as head and body. Length of hind foot less than basal length.

Female: No. 12400, Field Mus. Nat. Hist., which is young, has the following measurements: Total length, 390; length of tail, 150; length of hind foot, 39. Tail is 63 per cent as long as head and body. Length of hind foot less than basal length.

The differences in external measurements between the two sexes, as represented by the male type specimen and by the young female, are: Total length, 33, length of tail, 20; length of hind foot, 3.

Externals.—Longest facial vibrissae black or dark brown and reaching beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles (in summer pelage) slightly less than shown in figure .

Color.—Large spot between eyes, band confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, upper throat, chin, lower lips and in some specimens part or all of upper lips white; patch between eyes and bars in front of ears tinged with some shade of yellowish in one specimen; dark spot posterior to each angle of mouth present in four of five specimens; tip of tail black; remainder of upper parts, in summer, near (l) Brussels Brown or tones 1 to 2 of Raw Umber of Oberthür and Dauthenay, pl. 301; slightly darker brown on forehead, nose and about eyes. In winter near (j) Snuff Brown or lighter than Brussels Brown with a smoked effect. Underparts Buff-Yellow, winter and summer. Color of underparts extends distally on posterior sides of forelegs over toes onto antipalmar faces of feet and wrists and on medial sides of hind legs over antiplantar faces of toes. Least width of color of underparts averaging, in 5 available specimens 34 (24-42) per cent of greatest width of color of upper parts. Black tip of tail, in two adult males, averaging 53 (45 and 60) mm. Thus longer than hind foot and averaging 34 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on the type): See measurements and plates -. As described in M. f. nigriauris except that: Weight, 4.4 grams; basilar length, 44.7; postorbital breadth not less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; length of tympanic bulla less than length of lower molar and premolar tooth-row.

Female: Adult unknown.

Compared with the skull of the male of nevadensis, no single difference not covered by individual variation in nevadensis has been detected. Selected differences of inyoensis in comparison with latirostra are larger size, less inflated tympanic bullae and relative narrowness of the postorbital, interorbital and preorbital parts of the skull. Comparison of the skull with that of M. f. pulchra is made in the account of that subspecies.

Remarks.—Although two specimens of this subspecies were taken during the Death Valley Survey conducted by Dr. C. Hart Merriam, only three additional individuals are known to have been saved as study specimens since that time.

M. f. inyoensis as now known may be thought of as closely similar to M. f. nevadensis except for the presence of well-developed white facial markings like those found in the weasels of the San Joaquin Valley and coastal region of California south of San Francisco Bay. The nonwhite areas of the head are almost the same color as the back and not distinctly blackish as in M. f. latirostra and M. f. nigriauris. The one specimen in the winter coat, no. 25392/32805, U. S. Nat. Mus., from Lone Pine, is brown rather than white. The brown has the pale smoke-tinge common in the winter pelage of subspecies whose members are either brown or white in winter. The range of this subspecies is thought to include the floor and lower elevations of Owens Valley although it may occur in limited numbers southwestward along the base of the Sierra Nevada and through the mountains in places of low elevation like Walker Pass its range may meet that of pulchra.

The type specimen was taken in an alfalfa field by ranch hands, who, according to A. C. Shelton (MS), stated that the species was common at the type locality. None of the five specimens shows infestation of the frontal sinuses by parasites.

Specimens examined.—Total number, 5, listed by localities from north to south.

California. Inyo County: Alvord, 4000 ft., 1 (U. S. Nat. Mus.); 2 mi. N Independence, 1 (Mus. Vert. Zoöl.); Lone Pine, 3 (2 in Field Mus. Nat. Hist. and 1 in U. S. Nat. Mus.).

Mustela frenata neomexicana (Barber and Cockerell)

Long-tailed Weasel

Plates , , , , , and

Putorius frenatus neomexicanus Barber and Cockerell, Proc. Acad. Nat. Sci. Philadelphia, 1898:188; Lantz, Trans. Kansas Acad. Sci., 19:178, 1905.

Mustela frenata neomexicana, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Bailey, Animal Life of Carlsbad Cavern, p. 97, 1928; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.

Mustela frenatus neomexicanus, Bailey, N. Amer. Fauna, 35:19, September 5, 1913.

Type.—Male, adult, skull and skin; no. 10475, Mus. Comp. Zoöl.; Armstrongs Lake, Mesilla Park, Dona Ana County, New Mexico; February 1, 1898; obtained by A. C. Tryson; original no. 58 of C. M. Barber.

The skull is imperfectly cleaned but unbroken. The right upper incisors, right P2 and left p3 are broken away. The skin is indifferently stuffed but in a good state of preservation except that the distal part of the tail is missing. The animal's coat is ragged, and this imperfect appearance is heightened by injury to the posterior part of the body, probably at the time of capture.

Range.—From 3800 feet (type locality) to 9000 feet (Cloudcroft, N. Mex.); Upper Sonoran and Lower Sonoran life-zones of northern México, southeastern Arizona, New Mexico and western Texas, panhandle of Oklahoma, southeastern Colorado and southwestern Kansas. See figure on page .

Characters for ready recognition.—Differs from M. f. frenata and M. f. texensis by Buckthorn Brown rather than Brussels Brown color of upper parts, mastoid breadth of adult males ordinarily more, rather than less, than postpalatal length; from M. f. leucoparia by Buckthorn Brown rather than Argus Brown color of upper parts, distance from anterior margin of tympanic bulla to foramen ovale less, rather than more, than four-fifths height of tympanic bulla; from M. f. arizonensis and M. f. nevadensis by Buckthorn Brown, rather than near (14 n) Brussels Brown or, in winter, white color of upper parts, in presence of white frontal spot continuous with color of underparts, in basilar length of more than 46 mm. in males and 40 mm. in females; from M. f. longicauda by Buckthorn Brown rather than near (h) Clay Color of upper parts, by presence of white facial markings on Argus Brown head, and by length of tooth-rows amounting to less than 37 per cent of basilar length; from M. f. primulina by Buckthorn Brown rather than Brussels Brown color of upper parts, in presence of white frontal spot and broad white bands on side of head, in anteriorly truncate rather than anterolaterally rounded bullae and zygomatic breadth of more than 30 in males and 24 in females.

Description.—Size.—Male: The type specimen (see Barber and Cockerell, 1898:188) measured: Total length, 500; length of tail, 205; length of hind foot, 50. Tail 70 per cent as long as head and body. Length of hind foot less than basal length.

Female: No. 21779 from Tombstone, Arizona, measured: Total length, 419; length of tail, 165; length of hind foot from dried skin, 41 (probably 43 in flesh). Tail 65 per cent as long as head and body. Length of hind foot less than basal length.

The average differences in external measurements of the two sexes, as known from these two individuals, are: Total length, 81; length of tail, 40; length of hind foot, 7.

Compared with M. f. frenata, the size, proportions of parts and difference in size of the two sexes, appears to be about the same.

Externals.—Longest facial vibrissae colored like upper parts [in the type specimen some of the "long bristles of the upper lip" are white as pointed out by Barber and Cockerell (op. cit.: 188)] and extending beyond ear; carpal vibrissae colored like underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure .

Color.—Broad white bands on sides of head, extending anterodorsally anterior to each ear, confluent with white spot between eyes and with color of underparts; posterior half or all of each upper lip edged with white; usually few white hairs on top of head between ears; remainder of top of head near Argus Brown of Ridgway and Chocolate, tone 4, of Oberthür and Dauthenay; dark spot posterior to each angle of mouth usually absent; tip of tail black; remainder of upper parts varying, in different specimens, from Buckthorn Brown to Dresden Brown of Ridgway, and Brown Pink (tones 3 to 4, pl. 297, of Oberthür and Dauthenay); underparts Antimony Yellow or near (c) Warm Buff of Ridgway, and Brown Pink (tone 1, pl. 297, of Oberthür and Dauthenay); color of underparts extends distally on legs over forefeet and hind feet. Least width of color of underparts averaging 46 (41-55) per cent of greatest width of color of upper parts; black tip of tail 35 to 45 mm. long in females; 43 to 68 mm. long in males and averaging 21 (20-36) per cent as long as tail-vertebrae.

No specimen of this subspecies in the white winter coat has been seen. Animals taken in midwinter are available from Mesilla Park, Willcox, and 10 miles east of Roswell.

From M. f. frenata, neomexicana differs in: upper parts and underparts much lighter colored; white facial markings more extensive; color of underparts more extended onto feet. From M. f. leucoparia, neomexicana differs as follows: above and below, much lighter colored, but white facial markings less extensive and color of underparts less extended onto feet and legs.

Skull and teeth.—Male (based on adults: the type; no. 131582 from Berino, New Mexico; and no. 1485 from Seward Co., Kansas): See measurements and plates -. As described in Mustela frenata frenata except that: Weight, 6.2 (4.9 and 7.5); basilar length, 49.3 (48 and 50.5); mastoid breadth more than postpalatal length; least width of palate less than length of P4; anterior margin of masseteric fossa directly below m2 or heel of ml.

Female (based on three adults): See measurements and plates -. As described in Mustela frenata frenata except that: Weight, 3.1 (2.6-3.5) grams; basilar length, 42.7 (40.8-45.5); zygomatic breadth less than distance between condylar foramen and M1 and more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla.

The skull of the female averages 50 per cent lighter than that of the male.

As compared with the skull of the male of M. f. frenata, that of neomexicana is decidedly more angular and ridged. The postorbital constriction is narrower, the mastoid breadth greater (it is less than the postpalatal length in some subadult males), the sagittal crest much higher with impressions of the temporal and masseter muscles carried farther forward on the frontals, rostrum shorter and tympanic bullae wider and more inflated. Similar, though less marked, differences exist between the females. As compared with M. f. leucoparia and perotae, the same differences as noted above between frenata and neomexicana exist. In addition the tympanic bullae are so far removed from the foramen ovale that the distance from the anterior end of each bulla to the foramen ovale, instead of being less than the height of tympanic bullae, is in leucoparia more than four-fifths this height and in perotae more than the entire height. Also, in perotae, the squamosal, anterior to each tympanic bulla, is ventrally convex rather than ventrally concave as in neomexicana. Compared with M. f. longicauda, neomexicana is relatively narrower in the interorbital region, has relatively shorter tooth-rows, a V-shaped rather than a U-shaped interpterygoid space and in males has the interorbital region flat rather than convex and the sagittal crest is higher. The same differences are to be noted in comparison with nevadensis but here the difference in relative length of tooth-row is less. The same differences exist also in comparison with M. f. arizonensis except that its interorbital breadth, relative to the rest of the skull, is about the same. Difference in size is especially marked here; even females of neomexicana average larger than males of arizonensis.

Remarks.—When Barber and Cockerell named this subspecies in 1898, they had three specimens. Only two others are known to have been taken before this time. These are a skeleton, without corresponding skin, taken at Lozier, Texas, in 1890 by Wm. Lloyd, and no. 21779/36482, U. S. Nat. Mus., taken on April 6, 1893, by R. D. Lusk at Tombstone, Arizona. On the back of a label recently attached to the last mentioned specimen the name C. K. Worthen appears and probably signifies that the specimen was purchased from this dealer in vertebrate specimens.

M. f. neomexicana has a large geographic range. The old male from Liberal, Seward County, Kansas, extends the known range far to the northeast. Geographically, this occurrence is logical for the southwestern desertlike conditions extend to this part of Kansas. Probably the subspecies occurs in southeastern Colorado and in the panhandle of Oklahoma where conditions are similar. Bailey (1905:198) lists neomexicana as a member of the mammalian fauna of Texas. As stated by him (loc. cit.:198) this inclusion is based on geographic grounds and not on actual specimens. Strecker (1926:13) also includes neomexicana in his list of Texas mammals but writes me, under date of January 9, 1928, that "I included Mustela frenata neomexicana as a Texas mammal on the strength of its being mentioned by Bailey. . . ." On better ground, Bailey (1928:97) lists the subspecies as occurring in southeastern New Mexico at Carlsbad Cavern. However, Bailey (loc. cit.) knew of the existence of weasels just below El Paso and at Langtry, Texas. An unsexed skeleton, no. 167891, in the United States National Museum, from Lozier, Texas, is not certainly identifiable to subspecies. If, as I think, the animal is a female, its skull is intermediate between that of frenata and neomexicana although when all features are considered it is seen to be nearest the latter. The large size (basilar length of 46.5 mm.) may reflect some relationship to texensis. The field notes of the collector furnished me by Dr. H. H. T. Jackson (MS), describe the color as brownish yellow above and sulphur below. The admission of this subspecies to the list of mammals of Texas is made certain by the female (no. 1572, Texas Cooperative Research Collection) taken on July 28, 1940, 1-1/2 mi. NW Kent, Texas, by C. E. Scull.

The skull alone from Durango (City of), extends the known range far to the south. This skull is typical of neomexicana. Skins from the same place would be especially interesting as showing the approach, if any, in color, of neomexicana to M. f. leucoparia.

Mr. D. D. Stone of Casa Grande, Arizona, writes, under date of February 2, 1927, that a weasel was seen by an acquaintance of his in a field near Chandler, Maricopa County, Arizona. Probably this was neomexicana. If so, its range extends much farther west than collected specimens show.

Actual intergradation with M. f. frenata is not shown by the material at hand. The two females from Albuquerque, although typically neomexicana as regards color, have smaller, less prominently ridged skulls than females of neomexicana of the same age from farther south and approach M. f. nevadensis.

Probably the geographic ranges of M. f. neomexicana and M. f. latirostra do not meet; the only evidence of the existence of weasels in all of the large area, comprising western Arizona and the deserts of eastern California, which intervenes between the ranges of the two subspecies is the skull of a young individual, no. 68842, Mus. Vert. Zoöl., from 10 miles northeast of Bard, Imperial County, California. There, on December 29, 1932, Jack C. vonBloeker, Jr., retrieved the weathered skull with some of the vertebrae attached, from the top of a wood rat's nest beneath a mesquite tree near the west bank of the Colorado River.

The idea that the carcass may have been washed down the river from far upstream gains no support from a comparison of the specimen itself for the tympanic bullae are larger than in nevadensis and the skull is larger than the largest males seen of arizonensis, the two upriver races. On the basis of size the skull could be either a male of latirostra or a female of neomexicana. These two subspecies, like arizonensis and the skull in question, have much inflated bullae. However, the immaturity of the specimen conceals any other diagnostic cranial features, and prevents referring it certainly to either neomexicana or latirostra. In any event the specimen provides no evidence of intergradation between the two forms last mentioned. Speculating on its identity, I should say that it might be either an intergrade between arizonensis and nevadensis, from southern Utah or northwestern Arizona, or a member of an unnamed race resident in the lower part of the valley of the Colorado River.

Whereas M. f. panamensis and M. f. aureoventris are the darkest-colored weasels and occur in regions of heavy rainfall, M. f. neomexicana is the lightest-colored American weasel and occurs in an extremely arid region where the rainfall and humidity are slight.

According to Barber and Cockerell (1898:189) "The type specimen was shot in the grass on the shore of Armstrong's Lake. . . ." Bailey (1928:97) found the tracks of one of these animals "in the great pit at the west entrance to" Carlsbad Cavern and supposes they "hunt the cave walls for mice and other small game." Data on the label attached to no. 230973 states that the specimen was taken, two miles west of Willcox, Arizona, in a prairie dog town.

Only two of the 23 skulls show evidence of infestation of the frontal sinuses by parasites.

Specimens examined.—Total number, 28, arranged alphabetically by states and from north to south by counties in each state. Unless otherwise indicated specimens are in the United States National Museum.

Arizona. Graham County: Safford, 1. Cochise County: 2 mi. W Willcox, 1; Willcox, 1; 8000 ft., Chiricahua Mts., 1; 6000 ft., Pinery Canyon, Chiricahua Mts., 1; Tombstone, 1; Sulphur Spring Valley, 1.

Durango. "Durango City," 1.

Kansas. Seward County: Liberal, 1.

New Mexico. Bernalillo County: 3 mi. NW Albuquerque, 2. Lincoln County: 7800 ft., South Fork Eagle Creek, White Mts., 1. Chaves County: Pecos River, 10 mi. E Roswell, 8; Dexter, 1. Otero County: Cloudcroft, 9000 ft., 1. Dona Ana County: Mesilla Park, 2 (1, 1); Berino, 2.

Texas. Culberson County: 1-1/2 mi. NW Kent, 1. Terrel County: Lozier, 1.

Mustela frenata texensis Hall

Long-tailed Weasel

Plates , and

Mustela frenata texensis Hall, Carnegie Instit. Washington Publ. 473:99, November 20, 1936.

Mustela frenata, Strecker, The Baylor Bull., 27:14, September, 1924.

Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926 (part).

Type.—Male, adult, skull with skin of head, neck and tail; no. 14821, Amer. Mus. Nat. Hist.; Kerr County, Texas; September 17, 1897; obtained by H. P. Attwater.

The skull (plates -) and dentition are complete and unbroken. The preserved parts of the skin are not stuffed.

Range.—Lower Sonoran and possibly Upper Sonoran life-zones of central Texas. See figure on page .

Characters for ready recognition.—Differs from Mustela frenata arthuri in possessing white facial markings and postorbital breadth less than distance between posterior borders of P4 and P2; from M. f. frenata in larger size of body and skull, the basilar length of which in adult males is more than 52.5; from M. f. neomexicana in Brussels Brown rather than Buckthorn Brown color of upper parts and basilar length of skull more than 52.5.

Description.—Size.—Male: Measurements taken from the dried skins of a young male, no. 15476, Mus. Comp. Zoöl., from Kerr County, Texas, and a subadult male, no. 2017, Baylor Univ. Mus., from 5 mi. N Waco, Texas, are, respectively, as follows: Total length, 600 and more than 510; length of tail, 200 and 225; length of hind foot, 52 and 52.

Female: Skins unknown.

Externals.—As described in Mustela frenata frenata.

Color.—As described in Mustela frenata frenata.

Skull and teeth.—Adult male: See measurements and plates -. As described in Mustela frenata frenata except that: Weight, 8 grams; basilar length 54; least width of palate less than length of P4; anterior margin of masseteric fossa anterior to middle of m2.

Female: Skull unknown.

Remarks.—The type specimen, taken by the veteran collector of Texan mammals, H. P. Attwater, appears to have been the first one of these animals to find its way into the collection of any museum or naturalist. The second specimen from Kerr County was secured by, or through, the well-known commercial collector, F. B. Armstrong. Two trade skins, from Kerr County, taken on December 10, 1938, are in the Texas Cooperative Research Collection, as is also the skeleton of a young animal from Fredericksburg. The two other specimens from McLennan County (both males contrary to the statement of Strecker, 1924:14), owe their preservation to the alertness of John K. Strecker, Curator of the Baylor University Museum, who has given a complete account of their history.

The range of this subspecies is thought to include much of central Texas.

The preserved parts of the skin of the type specimen show it to have been generally large. The part of the tail preserved measures 226 millimeters and the skin of the head and neck is correspondingly large. The skin alone, no. 427, from near Waco, Texas, has, as now stuffed, a body 365 millimeters long. Individuals of this race attain larger size than those of any other American member of the subgenus Mustela with the possible exception of Mustela frenata macrophonius from Veracruz, México. In addition to large size, texensis and macrophonius are analogous in that each has a small geographic range at the northern end of an extensive range of its similarly colored southern relative from which it differs mainly in size. Each of the two groups, goldmani and macrophonius on the one hand and perotae, frenata and texensis on the other, has relatively uniform color, color pattern and body proportions over a large region but at its northern extremity develops a "giant" population, M. f. macrophonius and M. f. texensis, respectively. The skull of the type specimen of M. f. texensis is the largest one seen of any American weasel. The type specimen of M. f. macrophonius has a basilar length that is greater by one-tenth of a millimeter but in every other measurement taken the skull of M. f. texensis is the larger. Its weight, 8 grams, also shows it to be larger.

The broad, white bands in front of the ears are confluent with the white patch between the eyes on both sides in two specimens and on one side only in one other specimen. A white patch between the ears is present in four specimens. The dark spot at each angle of the mouth is absent on both sides in four specimens and on one side only in one other specimen. Thus out of a possible twelve cases, the broad bands in front of the ears are confluent with the spot between the eyes in five cases. Four of the six specimens have a white spot between the ears. The dark spot at each angle of the mouth is present three out of a possible twelve times.

The skull of no. 2017, from five miles north of Waco, is smaller than either of the two skulls seen from Kerr County and in this respect approaches M. f. frenata. There is no actual evidence of intergradation with any other subspecies but intergradation probably does take place with M. f. neomexicana and possibly with M. f. arthuri and M. f. primulina.

Strecker (1924:14) remarks that of the two specimens obtained near Waco, one was taken in a trap baited for mink and the other was shot in a hen house. None of the four skulls had the frontal sinuses infested with parasites.

Specimens examined.—Total number, 7, arranged by counties from north to south.

Texas. McLennan County: 5 mi. N Waco, 1; Erath, 1. Gillespie County: Fredericksburg, 1. Kerr County: 4; 1; and 2 trade skins.

Mustela frenata frenata Lichtenstein

Long-tailed Weasel

Plates , , , , , , and

Mustela frenata Lichtenstein, Darstellung neuer oder wenig bekannter Säugethier, 1832, pl. 42, and corresponding text, unpaged; Seton, Lives of game animals, 2:576, 1929.

Mustela brasiliensis Sevastianoff, Mem. de l'Acad. Imp. Sci. St. Petersburg, 4:356-363, tab. 4, 1813, name on plate only, the description being in the text (not of Gmelin, 1788); Gray, Proc. Zoöl. Soc. London, 1865:114.

Putorius frenatus, Baird Mamms. N. Amer., p. 173, 1858; Merriam, N. Amer. Fauna, 11:26, pl. 3, figs. 1, 1a, 1b, June 30, 1896; Bailey, N. Amer. Fauna, 25:198, October 24, 1905.

Putorius (Gale) brasiliensis aequatorialis Coues, Fur-bearing animals, p. 142, 1877, part? ("merely as a substitute for Gray's [supposedly] preoccupied name" that is, aureoventris).

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Putorius mexicanus Coues, Fur-bearing animals, p. 142, 1877, [nomen nudum, cited by Coues in synonymy as "Putorius mexicanus, Berlandier, MSS. ic. ined. 4 (Tamaulipas and Matamoros)"].

Putorius brasiliensis frenata, Allen, Bull. Amer. Mus. Nat. Hist., 3:219, April 17, 1891.

Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 6:197, May 31, 1894; Bangs, Proc. Biol. Soc. Washington, 10:9, February 25, 1896; Allen, Bull. Amer. Mus. Nat. Hist., 8:74, April 22, 1896.

Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.

Type.—Female, adult, skull and skin; no. 991, Berlin Zool. Mus., México City, México; June, 1829; obtained by F. Deppe.

The specimen once mounted, now is remade into a study skin and lacks the distal part of the tail. The skull (plates -, ) lacks the basicranial region.

Range.—Altitudinally, sea level (Brownsville, Texas) to 7600 feet (Tlalpam, México); from southern Texas as far south as México City; Lower Sonoran to at least Transition life-zone. See figure 29 on page .

Characters for ready recognition.—Differs from M. f. perotae in nonextension of blackish over anterior fourth of neck, least width of color of underparts more than 37 per cent of greatest width of color of upper parts; height of tympanic bulla more than distance from its anterior margin to foramen ovale; from M. f. leucoparia by restricted white facial markings that cover less than half surface of head in front of ears, by nonextension of black of head onto anterior half of neck and by wider (more than 7.8) tympanic bullae; from M. f. neomexicana by Brussels Brown rather than Buckthorn Brown color of upper parts and mastoid breadth less than postpalatal length; from M. f. texensis by smaller size of body and skull (basilar length in adult males less than 52.5); from M. f. arthuri by white facial markings and postorbital breadth less than distance between posterior borders of P4 and P2; from M. f. tropicalis by nonextension of blackish over anterior fourth of neck, least width of underparts more than 37 per cent of greatest width of upper parts, postorbital breadth of adult males less than distance between posterior borders of P4 and P2.

Description.—Size.—Male: Fifteen adults and subadults from Brownsville, Texas, yield average and extreme measurements as follows: Total length, 485 (430-556); length of tail, 202 (165-250); length of hind foot, 48 (40-55). Averages believed to be reliable but extremes probably are not. Tail averages 71 per cent as long as head and body. Length of hind foot less than basal length. Corresponding measurements of an adult male (topotype, no. 50826) from Tlalpam, México, are: 505, 203, 53. Another adult male, from Miquihuana, Tamaulipas, México, measures: 520, 215, 52.

Female: Six adults, subadults and young from Brownsville, Texas, yield average and extreme measurements as follows: Total length, 420 (362-456); length of tail, 173 (126-200); length of hind foot, 41 (40-46). Tail averages 70 per cent as long as head and body. Length of hind foot more (with possible exception of no. 36362/48732 U. S. Nat. Mus.) than basal length.

The average differences in external measurements of the two sexes are: Total length, 65; length of tail, 29; length of hind foot, 7.

Externals.—Longest facial vibrissae black and reaching beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure .

Color.—Spot between eyes, broad band, confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, and posterior two-thirds to one-half of each upper lip, white; remainder of sides and top of head, posteriorly to line connecting posterior margins of ears, blackish; dark spot posterior to angle of mouth present on both sides in about half the specimens; tip of tail black; remainder of upper parts Brussels Brown; chin white; remainder of underparts near (16´a) Ochraceous-Buff (same color in juveniles and young), which color extends distally on posterior sides of forelegs over forefeet and on medial sides of hind legs to feet and sometimes onto upper sides of toes. Least width of color of underparts averaging, in a series of seventeen males from Brownsville, Texas, 47 (extremes 38-53) per cent of greatest width of color of upper parts. Black tip of tail, in same series, averaging 49 (extremes 40-55) mm. long, thus about equal to length of hind foot and averaging 24 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on ten adults from Brownsville): See measurements and plates -; weight (three adults, one topotype and two from Brownsville, Texas), 6.2 (5.3-7.2) grams; basilar length, 49.8 (48.2-51.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoidal breadth less than postpalatal length; postorbital breadth less than length of upper premolars (less than distance between posterior borders of P4 and P2) and not greater (usually less) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 3 or 4 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter (usually longer) than rostrum; anterior margin of masseteric fossa just behind m2.

Female (based on two adults from Brownsville, Texas): See measurements and plates -, ; weight, 3.4 (3.3-3.5) grams; basilar length (six, adult to young) 43.3 (41.3-47.3); zygomatic breadth more or less than distance between condylar foramen and M1 and more than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less than (about equal to) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than outside length of P4; tympanic bulla as far posterior to foramen ovale as width of 2 to 3-1/2 (including I3) upper incisors; height of tympanic bulla more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum.

The skull of the female averages 45 per cent lighter than that of the average male.

Comparison of the skull with those of M. f. arthuri, tropicalis, perotae, leucoparia and neomexicana has been made in accounts of those subspecies. As compared with M. f. texensis (known only from males), the only difference detected is smaller size.

Remarks.—As Merriam (1896:27) has said: "In 1813 a Russian naturalist, Sevastianoff, gave the name 'Mustela brasiliensis' to a weasel brought to St. Petersburg by Capt. A. J. Krusenstern on his return from a voyage around the world. The animal was said to have come from Brazil, but no definite locality was given." This name was long applied by many European naturalists to American weasels which had white facial markings, and several American naturalists adopted the name. However, Lichtenstein in 1832 applied the name Mustela frenata to the weasels of the vicinity of México City and that name was used for bridled weasels from México and the southwestern United States by most subsequent German writers and by several Americans. In 1896 Merriam (1896:27) showed that Sevastianoff's Mustela brasiliensis, 1813, although probably the same as Mustela frenata, was preoccupied by Gmelin's Mustela brasiliensis, 1788, applied to an otter and that Lichtenstein's name must be used as the next available one. Since that time, 1896, frenata has been the name applied to the large bridled-weasels of Texas and the high table land of México south to México City. It may be added that in 1937 search by the writer among the specimens and records at the Russian Academy of Sciences, in Leningrad, failed to reveal any trace of the type specimen of Sevastianoff's Mustela brasiliensis.

The geographic range of this subspecies is relatively large and, as might therefore be expected, specimens show geographic variation. The specimens from Tlalpam, which Merriam (op. cit.:27) regards as topotypes, differ in certain respects from specimens from Texas. The skull of the adult male "topotype," no. 50826, differs from any other adult male seen in that the basilar length, the length of the upper tooth-rows, the orbitonasal length, the ratio of the same to the basilar length, the mastoidal breadth, the zygomatic breadth, the depth of the skull at the posterior margins of the upper molars, and the length and breadth of M1, are greater. The height of the tympanic bullae is less than the average height for these structures in more northern specimens. The specimens from Tlalpam have also larger external measurements than the average of more northern specimens. All of these features show an approach to the subspecies of more southern distribution. On the other hand, the blackish of the head is not more intense or more extended posteriorly onto the neck than in specimens from Brownsville, Texas. The skin, with skull crushed, no. 767, in the Paris Museum, from 3200 meters elevation near Toluca, does have the black color of the head extended 30 millimeters posteriorly to the ears. In this feature, and also in the extensively white face on which the white bar in front of each ear connects with the frontal spot, as well as with the color of the underparts, the specimen resembles leucoparia. Better material from the western part of the state of México may show the range of leucoparia to extend eastward almost or quite to Toluca.

An adult male, taken on July 15 at Miquihuana, Tamaulipas, is unique in several respects. The top of its head is black, rather than blackish, and this color extends posteriorly on the top and sides of the neck almost halfway to the shoulders. All of the upper parts are much more darkly colored than in other specimens of this race. The least width of the color of the underparts is 63 per cent of the greatest width of the color of the upper parts; thus the color of the underparts is considerably more extensive than in any other specimen seen. The underparts are more intensely colored than in the average specimen. The mastoidal breadth is greater than in any other adult male and amounts to more than the postpalatal length. On available maps the elevation of Miquihuana is given as 1892 meters (about 6200 feet). Thus the dark colors can hardly be ascribed to more tropical conditions than those under which animals from Brownsville, Texas, live. Brownsville is only a few feet above sea level and only 235 miles farther north. The difference noted, therefore, seems to be of geographic significance. However, there is from Alvarez, San Luis Potosí, approximately 115 miles south of Miquihuana, a young (nearly subadult) female, no. 21968, which is as light colored as specimens from Brownsville, Texas, or Tlalpam, México. The only distinctive feature of this specimen is the much greater extent of its white facial markings; they are more extensive even than in the specimen from Miquihuana.

Finally, the series from Brownsville, Texas, indicates that the animal there is smaller than frenata from the vicinity of México (city). The skull is similarly proportioned except that relative to the basilar length the orbitonasal length is more. Several other measurements of the skull of the adult male from Tlalpam, as pointed out above, are actually, although not relatively, greater than in any specimen from Brownsville. The similarities between specimens from the two localities, Tlalpam and Brownsville, are striking; since the two localities lie at opposite, extreme ends of the range more geographic variation would be expected. All that is known of the characters of populations from intermediate localities is that the one specimen from Alvarez shows no peculiarities whereas the one from Miquihuana suggests the existence there of a geographic variant.

None of the specimens seen shows actual intergradation with M. f. neomexicana or with M. f. arthuri but it is supposed that frenata intergrades with each of these subspecies. The difference between frenata and arthuri is greater than between frenata and neomexicana. Bailey (1905:198) records tracks of a weasel seen just below El Paso which he supposed had been made by a weasel of the neomexicana type. He also cited the taking of a weasel at Langtry which suggested to him (op. cit.) ". . . a continuous range from the country of frenatus up the Rio Grande to the type locality of neomexicanus at Mesilla Valley," New Mexico. Other records of occurrence in Texas cited by Bailey, in addition to those provided by specimens examined by the writer, are San Diego, Beeville, and Port Lavaca. The Port Lavaca record is the easternmost one assigned to the subspecies frenata; possibly specimens from there would be referable to arthuri.

The series of thirty-four specimens from Brownsville, Texas, permits measuring the amount of individual and age variation in several features. For instance, the material is sufficient to show that external measurements of subadults and those that fall in the upper part of the category designated as "young" may be included with the measurements of adults, because the mentioned measurements are not appreciably greater in adults. The series of skulls, although not providing more than six of any one age, shows the range of variation in size and proportion of certain parts and enables the student the better to evaluate cranial characters of nearby races known from only a few specimens. For example, not one of the twenty skulls of males from Brownsville and immediate vicinity is as large as either of the two specimens of texensis from Kerr County.

The white facial markings vary much in size and shape. In the series of thirty-four skins from Brownsville the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in three specimens and on one side only in six other specimens. These bands are confluent with the color of the underparts in all but two specimens. In one specimen the connection is lacking on both sides and in the other on one side only. A white patch between the ears is present in two specimens. The dark spot at each angle of the mouth is absent on both sides in eleven specimens and absent on one side only in ten others.

In six other specimens from parts of Texas north of Brownsville, the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in one specimen. A white spot between the ears is present in one specimen. The dark spot at each angle of the mouth is absent on both sides in six specimens and on one side only in three other specimens.

In eleven specimens from México, the broad white bands in front of the ears are confluent with the white spot between the eyes on both sides in two specimens and on one side only in one other specimen. The white spot between the ears is present in one specimen. The dark spot at each angle of the mouth is absent on both sides, in six specimens, and on one side only in one other specimen.

Thus, in 51 specimens the broad bands (one in front of each ear) are confluent with the white patch between the eyes in nineteen out of 100 instances, and not with the color of the underparts in three instances. A white spot between the ears is present in four specimens. The dark spot at each angle of the mouth is present 47 out of a possible 98 times.

Four juvenal specimens from Brownsville, Texas, with their dates of capture and probable age, are as follows: no. 58574, ♀, three weeks old, taken on February 15; no. 17318/24239, ♂, four weeks old, taken on March 16; no. 45899, ♀, forty days old, taken on May 21; no. 21778/36481, ♂, thirty days old, taken on October 20. In the order given, the dates of birth of these four juveniles would be approximately as follows: January 25, February 15, April 1, and September 20. The dates of birth of other specimens less than three months old as judged by the stage of development of the skull, and reckoning backward from the dates of capture, are as follows: April 1, April 30, May 25, October 12, and December 21. Thus, young appear to be brought forth at Brownsville, Texas, in the fall, winter and spring, that is to say from the latter part of September until the latter part of May.

Mustela frenata frenata is either free of the parasites that infest the frontal sinuses of most weasels, or withstands their presence remarkably well, for only one skull shows a definite pathological condition of the frontal sinuses.

Allen (1896:74) quotes H. P. Attwater, with respect to this species in Bexar County, Texas, as follows: "Not common, but occasionally met within the chaparral and cactus lands, where Wood Rats, Rabbits and Quail abound. They were frequently met with around San Antonio during the great 'Tramp Rat' [= Sigmodon hispidus texianus, see Bailey (1905:116)] invasion of 1889-90."

Specimens examined.—Total number, 63, arranged by counties, and in México by states, from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.

Texas. Bexar County: San Antonio, 2 (1). Goliad County: Charco, 1. Nueces County: Corpus Christi, 1. San Diego County (not found), 1. Hidalgo County: La Hacienda, 1. Duval County: San Diego, 2. County in question: Lower Rio Grande, 1. Cameron County: Brownsville, 34 (3, 4, 3, 2, 1, 1, 1); no locality more definite than county, 2.

Nuevo León. Río Ramis, 20 mi. NW Montemorelos, 1.

Tamaulipas. Matamoros, 6; Miquihuana (now in Nuevo León), 1.

San Luis Potosí. Alvarez, 1.

México: Region montagneuse des environs de Toluca, Nevada Toluca, 3200 M., 1

Distrito Federal. City of México, 2 (1); Tlalpam, 2. No locality more definite than México, 4 (1, 3).

Mustela frenata leucoparia (Merriam)

Long-tailed Weasel

Plates , , , , , , , and

Putorius frenatus leucoparia Merriam, N. Amer. Fauna, 11:28, June 30, 1896.

Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 2:165, October 21, 1889.

Putorius frenatus frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 22:259, July 25, 1906.

Mustela frenata leucoparia, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.

Type.—Male, adult, skull and skin; no. 34914/47179, U. S. Nat. Mus., Biol. Surv. Coll.; Pátzcuaro, Michoacán, México; July 27, 1892; obtained by E. W. Nelson; original no. 2960.

The skull (plates and ) lacks most of the braincase; a fragment, consisting of the supraoccipital and the coalesced frontals and parietals remains. The rostrum, left zygomatic arch, palate, left pterygoid, left glenoid fossa and right postorbital process are intact. The teeth all are present and entire. The lower jaw lacks the right coronoid process and the lateral part of the articular condyle. The skin is well made and in good condition. It differs from an adult male topotype (36855, U. S. Nat. Mus.) and other referred specimens in having: the black of the head extended farther posteriorly on the neck, the maximum amount of white on the head, and a white stripe 50 mm. long extending down the middle of the nape from a point between the ears more than half way to the shoulders.

Range.—Sonoran and Transition life-zones of mountains west of México (city) in Michoacán and Nayarit. See figure on page .

Characters for ready recognition.—Differs from M. f. goldmani in least width of color of underparts more than 47 per cent of greatest width of color of upper parts, hind feet colored like underparts rather than like upper parts; postorbital constriction less than, rather than more than, combined length of upper premolars; from M. f. macrophonius by same details of coloration as from goldmani and by ventrally concave rather than ventrally convex pretympanic part of squamosal; from M. f. perotae by least width of color of underparts more than 40 per cent of greatest width of color up upper parts; height of tympanic bulla more than three-fifths distance from its anterior margin to foramen ovale; from M. f. frenata by white facial markings that cover half of surface of head in front of ears, by extension of black of head onto neck halfway to shoulders and by narrower (less than 7.8) tympanic bullae; from M. f. neomexicana by Argus Brown rather than Buckthorn Brown color of upper parts and distance from anterior margin of tympanic bulla to foramen ovale more, rather than less, than four-fifths of height of tympanic bulla.

Description.—Size.—Male: Two adults and one young from Los Reyes and Pátzcuaro, Michoacán, yield average and extreme measurements as follows: Total length, 514 (510-523); length of tail, 206 (196-215); length of hind foot, 55 (52-58). Tail averages 67 per cent as long as head and body. Length of hind foot more than basal length.

Female: One adult from Artenkiki, Jalisco, and one subadult from Pátzcuaro, Michoacán, measure, respectively, as follows: Total length, 412, 400; length of tail, 159, 159; length of hind foot, 41, 42. Tail averages 64 per cent as long as head and body. Length of hind foot equal to or greater than basal length.

The average differences in external measurements of the two sexes are: Total length, 108; length of tail, 47; length of hind foot, 13.

Mustela frenata leucoparia has a greater total length and length of tail than either M. f. frenata or goldmani. The hind foot is longer than that of frenata and approximately the same as in goldmani. Relative to the body length, the tail averages longer than that of goldmani and shorter than that of frenata.

Externals.—As described in Mustela frenata frenata.

Color.—Broad white bands on sides of head, extending anterodorsally anterior to each ear, confluent with white spot between eyes and with color of underparts; posterior third of each upper lip white; remainder of sides and top of head, and neck posteriorly to point halfway to shoulders from ears, black; no dark spots at angles of mouth; tip of tail black; remainder of upper parts Argus Brown; chin white and sometimes also chest, neck and medial sides of hind legs; remainder of underparts near (16´) Ochraceous-Buff (near (a) Ochraceous-Buff in juvenal female), which color extends distally over all of each foreleg (except its lateral face proximally from about middle of forearm) and on medial side of hind leg and over most of upper side of each foot. Least width of color of underparts averaging, in eight specimens, 54 (extremes 44-61) per cent of greatest width of color of upper parts; black tip of tail averaging, in four males, 52 (extremes 38-78) mm. long, thus averaging 25 per cent of length of tail-vertebrae.

As compared with M. f. frenata and goldmani: white facial markings more extensive; color of underparts less restricted and more extended on legs; black tip of tail relatively of about same extent as in frenata and thus much less than in goldmani; black color of head extending farther posteriorly than in frenata but not so far as in goldmani.

Skull and teeth.—Male (adult): See measurements and plates -, 29, 30. As described in Mustela frenata frenata except that: Weight (no. 128972) 6.3 grams; basilar length, 51.2; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla more or less than (about equal to) distance from its anterior margin to foramen ovale; anterior margin of masseteric fossa anywhere from slightly anterior, to slightly posterior, to m2.

Female (based on no. 26153): See measurements and plates -. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 44.5; zygomatic breadth less than distance between condylar foramen and M1, or than between anterior palatine foramen and anterior margin of tympanic bulla; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla not more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row or than length of rostrum.

The skull of the female is 43 per cent lighter than that of the male.

Comparison of the skull with those of M. f. perotae, goldmani and neomexicana has been made in the accounts of those subspecies. As compared with that of frenata the main difference is the less inflated tympanic bulla, the height of which is approximately equal to, rather than decidedly more than, distance from its anterior margin to foramen ovale.

Remarks.—The first specimen known to have been preserved is the alcoholic in the British Museum of Natural History, taken in September, 1891, on the Río Santiago in Jalisco, by D. A. C. Buller. The other known specimens of this white-faced weasel are divided between the American Museum and the United States National Museum. The two referred specimens from Jalisco were the last of several helpful ones collected in México and Central America by J. H. Batty, and these two were taken less than three months before Batty's tragic death in Chiapas (see Allen, J. A., 1906:191). The five specimens from Michoacán were taken by Nelson or Nelson and Goldman together. Merriam had only three of these when he named the subspecies and remarked (1896:29) that "This form is the poorest subspecies described in the present paper." Although the form is not strongly marked, the two additional specimens from Michoacán and better comparative material than Merriam had confirm several of the differential characters ascribed to it by him and indicate the existence of still other characters.

M. f. leucoparia occurs in the Sonoran and Transition life-zones. No. 27258 from Los Masos, and no. 26153 from Artenkiki (see specimens examined for other spellings) approach true frenata in coloration. Each of these specimens has a few white hairs between the ears and the white patch between the eyes is confluent on one side only with the lateral white bands on the side of the head. No. 27258 from Los Masos has a dark spot at each angle of the mouth. The 7 other specimens are relatively uniform in coloration. Each has the white spot between the eyes confluent on both sides with the extensive white areas on each side of the face. None has a dark spot at either angle of the mouth. Of these 7 specimens, the type specimen and three others have white hairs forming a median line between the ears and a fifth specimen has a white spot behind each ear.

M. f. leucoparia is most like M. f. frenata. Unlike frenata, leucoparia has tympanic bullae that are less inflated, narrower and less projected, at their anterior margins, from the cranium. In these characters leucoparia is intermediate between M. f. frenata and M. f. goldmani. The latter subspecies has the least inflated, narrowest and least projecting tympanic bullae of the three. The black color of the head extends, on the average, farther posteriorly than in M. f. frenata but not so far as in M. f. goldmani. The general color, too, is intermediate between that of M. f. frenata and that of the much darker M. f. goldmani. The white facial markings are more extensive than in either M. f. frenata or M. f. goldmani. This applies to both the white area between the eyes and the one on each side of the head between the ear and eye. M. f. neomexicana, whose range possibly meets that of M. f. leucoparia, also has more extensive white facial markings than M. f. frenata but less extensive markings than M. f. leucoparia.

On the basis of skulls alone, specimens of frenata from Tlalpam and those of leucoparia from Los Reyes can hardly be distinguished. This fact, and the circumstance that the specimens from the northern part of the range of leucoparia closely resemble frenata in color, constitute sufficient evidence for regarding the two as only subspecifically distinct. The female, no. 26153 from Artenkiki, as mentioned above, approaches true frenata in coloration. On this account it is not to be regarded as typical and it was because no other skulls of adult females were available that this one was used for comparison with females of allied races.

M. f. leucoparia is, then, a subspecies of the large, temperate-zone group and is unique in possessing the maximum extent of white facial markings.

None of the seven skulls shows signs of having had the frontal sinuses infested with parasites.

Specimens examined.—Total number, 8, all from México. Localities are listed by states from north to south. Specimens from Michoacán are in the United States National Museum; one from Río Santiago is in the British Museum of Natural History; all others are in the American Museum of Natural History.

Nayarit. Tepic, 1.

Jalisco. Río Santiago, 1; Los Masos, 1; "Artenkiki" (J. A. Allen, 1906, p. 238, writes "Artenkikil" and, on p. 259, "Artenkiki."), 1.

Michoacán. Zamora, 1; Los Reyes, 1; Pátzcuaro, 3.

Mustela frenata perotae Hall

Long-tailed Weasel

Plates , and

Mustela frenata perotae Hall, Carnegie Instit. Washington Publ. 473:100, November 20, 1936.

Putorius frenatus, Merriam, N. Amer. Fauna, 11: pl. 3, fig. 2, June 30, 1896.

Type.—Female, adult, skull and skin; no. 54278, U. S. Nat. Mus., Biol. Surv. Coll.; 12,500 feet, Cofre de Perote, Veracruz, México; May 26, 1893; obtained by E. W. Nelson; original no. 4864.

The skull (plates, 37-39) lacks the right zygomatic arch. Left p2 is missing. The skin is fairly well made and in good condition except that the extreme tip of the tail has been broken off and there are two holes in the right hind leg. The underparts show the beginning of a spring molt.

Range.—From 7500 (?) feet (Perote) to 13,500 feet (Popocatépetl), Upper Sonoran, Transition and Boreal life-zones of mountains along Puebla-México boundary, eastward to western central Veracruz and south into Oaxaca. See figure on page .

Characters for ready recognition.—Differs from M. f. frenata, its nearest relative, in extension from head of blackish onto anterior fourth of neck; restriction of color of underparts (least width of same less than 37 per cent of greatest width of color of upper parts), height of tympanic bulla less than distance from its anterior margin to foramen ovale; from M. f. macrophonius and M. f. goldmani in presence of, rather than absence of, color of underparts on hind feet; upper parts (black) Brussels Brown rather than Argus Brown or darker; from M. f. tropicalis in larger size (adult female with total length more than 400, basilar length more than 40, weight of skull more than 3 grams); postorbital breadth less than combined length of upper premolars; m1 more than 5.4 long; from M. f. leucoparia in white facial markings so restricted that spot between eyes is not confluent with white stripe in front of ear, or, if so, narrowly (less than 4 wide) confluent; color of upper parts extending onto antipalmar face of forefoot, least width of color of underparts not more than 40 per cent of greatest width of color of upper parts; height of tympanic bulla not more than three-fifths distance from its anterior margin to foramen ovale.

Description.—Size.—Male: A nontypical specimen from Cerro San Felipe, Oaxaca, measures: Total length, 500; length of tail, 205; length of hind foot, 52.

Female: The type specimen, measures: Total length, 418; length of tail, 160; length of hind foot, 45.

In this male the tail is 70, and in the female, 62 per cent as long as the head and body. In each the hind foot is longer than the basal length.

The differences in external measurements between these two specimens, representing the two sexes, are: Total length, 82; length of tail, 45; length of hind foot, 7.

Externals.—As described in Mustela frenata frenata.

Color (based on type specimen).—Color and color pattern as described in Mustela frenata frenata except that: blackish of sides and top of head extends one-fourth of way back to shoulders from ears; throat and breast as well as chin white; remainder of underparts near (16´ c) Ochraceous-Buff; least width of color of underparts equals 36 per cent of greatest width of color of upper parts; black tip of tail equal to 28 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on a referred specimen from Cerro San Felipe which certainly is nontypical): See measurements. As described in Mustela frenata frenata except that: Weight, 4.9 grams; basilar length, 49.2; postorbital breadth more than distance between posterior borders of P4 and P2; tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla.

Female (based on type specimen, an adult): See measurements and plates -. As described in Mustela frenata frenata except that: Weight 3.4 grams; basilar length, 43.5; zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla one-half to three-fifths distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer than rostrum.

The skull of the female is 33 per cent lighter than that of the nontypical (and smaller than average) male from Cerro San Felipe.

Comparison of the skull with that of M. f. tropicalis is made in the account of that subspecies. Compared with the skull of M. f. macrophonius, that of the female of perotae is more flattened, has the longitudinal dorsal outline distinctly concave rather than flat just behind the postorbital processes, and much wider tympanic bullae. Accordingly, the basioccipital is slightly narrower in perotae. The more marked postorbital constriction of the type specimen of perotae possibly is due to its relatively greater age. As compared with the skull of M. f. leucoparia, that of the female of perotae has less inflated tympanic bullae, the height of each being half as great as distance from its anterior margin to foramen ovale, whereas, in leucoparia (as represented by no. 26153) the two distances are equal. As compared with that of M. f. frenata, the skull of the female of perotae differs mainly in the lesser inflation of the tympanic bullae and their relative position. The height of each bulla is in perotae only half as much as, but in frenata more than, the distance from its anterior margin to foramen ovale. The anterior margin of the bulla is much less projected from the floor of the braincase in perotae. The squamosal anterior to each bulla is convex ventrally in perotae but flat or concave ventrally in frenata.

Remarks.—The type specimen and a juvenal female from the town of Perote were taken in the spring of 1893 by E. W. Nelson. Of these two, the type specimen was mentioned and figured by Merriam (1898:30, fig. 16 [= fig. 15], pl. 3, fig. 2) as Putorius frenatus. The referred nontypical specimen from Cerro San Felipe, Oaxaca, was referred by Merriam (op. cit.:29) to Putorius frenatus goldmani with the comment that it was intermediate ". . . both in coloration and cranial characters, between typical frenatus and goldmani;. . . ." No other published references to this subspecies, or specimens of it, have been seen. In 1941 and 1942, W. B. Davis and associates took four specimens along the boundary between the states of Puebla and México.

Although the specimen from Cerro San Felipe, Oaxaca, is referred to Mustela frenata perotae, to the description of which it answers best, that specimen, on account of its structural characters and geographic position relative to adjacent races, is in reality an intergrade between several of the adjacent races. Some of its intermediate characters are pointed out in the discussion of M. f. goldmani. In the specimens from 45 and 55 kilometers ESE of México (city) the black color of the top of the head does not extend so far behind the ears as in the holotype of M. f. perotae and in this feature the two specimens show intergradation between the two subspecies, perotae and frenata.

The type specimen taken on May 26, is acquiring new hair on the belly and lower sides which appears to be the result of a normal molt.

As would be expected from its geographic position, M. f. perotae resembles M. f. frenata of northern México and the high mountain forms of southern México more than it does the lowland tropical forms. This is true as regards size of entire animal, proportions of its parts, and size, general angularity and major proportions of its skull. The marked postorbital constriction, convex supralacrymal face of rostrum, width of tympanic bullae and angularity of the braincase place it nearest M. f. frenata as does also the color and color pattern. The ventrally convex squamosal anterior to each tympanic bulla and the slight degree of projection from the cranium of the anterior margin of each tympanic bulla are intermediate in degree between the condition in M. f. macrophonius and that in M. f. frenata. Thus M. f. perotae combines several characters of M. f. frenata on the one hand with some of M. f. macrophonius on the other and in some features, for instance in the size, shape and degree of inflation of the tympanic bullae, presents intermediate stages of development.

On the eastern plain below the high mountain, Cofre de Perote, there ranges the similarly colored, smaller, tropical weasel, Mustela frenata tropicalis. Between M. f. perotae and M. f. tropicalis there is marked differentiation in the skulls with much less differentiation in coloration. The differences in typical skulls of the two subspecies are so pronounced that one would, at first glance, hardly believe it possible for direct intergradation to occur between them on the sides of this mountain. Merriam (1896:30) thought that it did not. The two skulls figured by him (op. cit.:31) are a topotype of M. f. tropicalis from Jico and the one which now is the type specimen of M. f. perotae. They show the great difference in size and proportions and are females of comparable ages, not of different ages as I suspected before examining the skulls. However, despite this marked difference in the skulls, there is some, although not conclusive, evidence of intergradation furnished by a young female from Xuchil, Veracruz. This specimen is described in connection with M. f. tropicalis (see page ).

None of the seven skulls shows marked deformity of the interorbital region, but two of the three adults appear to have had these parts infested with nematodes.

Specimens examined.—Total number, 7, all from México, listed by localities from north to south. Specimens from Veracruz and Oaxaca in the United States National Museum; remainder in Texas Cooperative Research Collection.

México: Monte Río Frío, 45 Km. ESE México City, 1; 55 Km. ESE México City, 1; N slope Mt. Popocatépetl, 13,555 ft., 1.

Puebla. Río Otlati, 8700 ft., 1.

Veracruz. Cofre de Perote, 12,500 ft., 1; Perote, 1.

Oaxaca. Cerro San Felipe, 10,000 ft., 1.

Mustela frenata goldmani (Merriam)

Long-tailed Weasel

Plates , , , and

Putorius frenatus goldmani Merriam, N. Amer. Fauna, 11:28, June 30, 1896; Elliot, Proc. Biol. Soc. Washington, 18:236, December 9, 1905.

Mustela frenata goldmani, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, adult, skull and skin; no. 77519, U. S. Nat. Mus., Biol. Surv. Coll.; Pinabete, Chiapas, México; February 10, 1896; obtained by E. A. Goldman (on attached label collectors recorded as Nelson and Goldman); original no. 9279.

The skull (plates and ) has the rostrum badly injured. All the right, and part of the left nasal, the upper part of the right maxilla, the postorbital process and intervening area of frontals are missing. Each zygomatic arch is broken but the parts are present and attached to the skull. The frontal and interorbital regions are greatly malformed owing to parasites that infested the sinuses. Right I2 and I3, right and left i3, and the medial parts of the paraconid and protoconid of right m1 are missing. The light facial markings are less extensive than in any of the referred specimens. These markings consist of a separate spot between the eyes and a white line, confluent with the color of the underparts, on each side of the head, that extends from the base of the ear to above the eye. The dark color of the underparts is represented at the angles of the mouth by a spot on the left side and a similar dark area, confluent with the dark color of the face, on the right side. The large size, characters of the skull, and scrotal pouch on the skin prove the specimen to be a male as stated on the label.

Range.—Two thousand five hundred feet (El Cipres, Guatemala) to 9500 feet (near Tecpám, Guatemala), Upper Tropical Life-zone of mountains and western coasts of southern México, Guatemala and Salvador. See figure on page .

Characters for ready recognition (characters based on males).—Differs from M. f. nicaraguae and M. f. perda by larger size (total length of adult males more than 489), least width of color of underparts not less than 26 per cent of greatest width of color of upper parts, weight of skull of adult male more than 5 grams; from M. f. macrophonius by smaller size (total length of adult males less than 540), skull of male with basilar length less than 52.5 and weight less than 6 grams; from M. f. perotae (typical specimens of same sex not available) by darker color of upper parts which are Argus Brown or darker rather than Brussels Brown; nonextension of color of underparts onto hind feet; from M. f. leucoparia in least width of color of underparts not more than 37 per cent of greatest width of color of upper parts; color of underparts not extended onto hind feet; black tip of tail two-fifths rather than one-fourth as long as tail-vertebrae; height of tympanic bulla less than four-fifths distance from its anterior margin to foramen ovale.

Description.—Size.—Male: Four adults yield average and extreme measurements as follows: Total length, 508 (500-512); length of tail, 196 (185-207); length of hind foot, 55.5 (54-58). Tail averages 63 (59-67) per cent as long as head and body. Length of hind foot more than basal length.

Female: Typical specimen unknown.

Externals.—Longest facial vibrissae black and reaching beyond ear; carpal vibrissae wholly or in part of same color as upper parts and reaching as far as hypothenar pad; hairiness of foot-soles distinctly less than that shown in figure on page .

Color.—Spot between eyes, band, confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear and posterior third of each upper lip, white; remainder of sides and top of head and neck posteriorly to or slightly behind shoulders, black; dark spots at angles of mouth usually absent; tip of tail black; remainder of upper parts Argus Brown or near (n) Argus Brown; chin, throat and breast white; remainder of underparts near (16' c) Ochraceous-Buff; color of underparts extending distally on posterior sides of forelegs onto medial toes and on hind legs to points between knees and heels. Least width of color of underparts, in five adult males, averaging 28 (extremes 26-33) per cent of greatest width of color of upper parts; black tip of tail, in four adult males, averaging 40 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on five adults): See measurements and plates -, ; weight, 5.4 (5.3-5.5) grams; basilar length, 49.9 (49.6-51.3); zygomatic breadth (except in no. 12523 from Salvador) more than or equal to distance between condylar foramen and M1 or between anterior palatine foramen and anterior margin of tympanic bulla. Mastoid breadth less than postpalatal length; postorbital breadth more or less than length of upper premolars and greater than width of basioccipital measured from median margin of one foramen lacerum posterior to its opposite; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of five upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than or equal to length of rostrum; anterior margin of masseteric fossa immediately behind m2.

Female: Typical skull unknown.

Comparison of male skull with that of M. f. perda made in discussion of that form. Comparison with that of M. f. nicaraguae shows similar differences, some of which are more pronounced. For example, squamosals anterior to tympanic bullae more convex ventrally and these bullae project less from braincase than in M. f. perda; thus the difference in these features is greater between goldmani and nicaraguae than between goldmani and perda.

As compared with the skull of the male of M. f. macrophonius, each one of the skulls of the adult males of M. f. goldmani is smaller in every measurement taken, with two exceptions. The width of the tympanic bullae was more in three specimens of M. f. goldmani as was also the depth of the same in three specimens. Relative to the basilar length all but two of these measurements average less in goldmani; the exceptions are the zygomatic breadth and depth of the skull at the anterior margin of the tympanic bullae which average more. Relative to the basilar length, the orbitonasal length and depth of the skull at the posterior margin of M1 are less in each skull of goldmani. Thus, excepting the width and height of the tympanic bullae and the relative zygomatic breadth and relative depth of the braincase posteriorly, the skull of goldmani is shorter and relatively as well as actually narrower and lighter throughout.

As compared with the skull of the male of M. f. leucoparia, that of M. f. goldmani averages a trifle shorter and no skull of goldmani equals that of leucoparia in actual or relative zygomatic and mastoid breadths or length or height of tympanic bullae. In depth, the skull of goldmani averages actually and relatively greater. Its teeth are smaller. The squamosal anterior to each tympanic bulla is convex ventrally whereas it is concave ventrally in leucoparia as in frenata.

Remarks.—When Merriam (1896:28) named this subspecies, he had only one specimen but he called attention to the more important diagnostic characters, which additional specimens show pertain to the race as a whole.

M. f. goldmani in typical form occurs in high mountains of the Upper Tropical Life-zone and is most closely related to M. f. frenata and M. f. macrophonius. The altitude at which the two specimens were taken, twenty miles southeast of Teopisca in Chiapas, is not known. Merriam (1896:28) states that the type specimen was obtained at "about 8200 feet." The specimen taken by Stirton in Salvador comes from 8000 feet and the one obtained by Barber in Guatemala from 9500 feet. The specimen from Dueñas, the skin alone of a young animal, is not instructive.

As regards size, goldmani is larger than the immediately adjacent subspecies from the Lower Tropical Life-zone but is smaller than M. f. leucoparia or macrophonius. As compared with M. f. frenata, goldmani is longer, has an actually as well as relatively shorter tail, and a much longer hind foot.

The most outstanding difference in externals from frenata is the naked foot soles.

Molting probably takes place twice each year although actual proof of this is lacking. In number 133254 from twenty miles southeast of Teopisca, taken on May 12, the molt is well advanced. Another specimen from the same place still retains the winter coat.

In color, goldmani is much darker than frenata, has less extensive white facial markings, longer black tip on tail, more restricted color of underparts, and lacks the extension of color of the underparts onto the hind feet.

Of the adult males from the high mountains, the type specimen from Chiapas is lightest, and the one from Salvador is darkest. This progressively darker color to the southward probably is geographic variation.

In total length and relative and actual length of tail, the specimen from Salvador is the smallest of the five adult males from the higher mountains. In addition to its darker color and smaller size, no. 12523 from Salvador shows certain distinctive cranial characters. The zygomatic breadth is less than, rather than more than, or equal to, the distance between the condylar foramen and M1 or than that between the anterior palatine foramen and the anterior margin of the tympanic bulla. This difference appears to be correlated with geographic position, since no. 15953 from Guatemala has the three distances about equal and therefore is intermediate in this respect between the specimen from Salvador and those from Chiapas, in which the zygomatic breadth is greater than the other two measurements. Also in the greater depth of the skull and smaller size of the teeth, this specimen from Salvador approaches the subspecies of the Lower Tropical Life-zone. It has, however, the longest, highest and widest tympanic bullae of any of the five specimens. The amount of ventral convexity of the squamosal in front of each tympanic bulla appears not to be greater than in the other specimens.

As indicative of intergradation with perotae, leucoparia and possibly frenata, there is the specimen from Cerro San Felipe, Oaxaca. The degree of restriction of the color of the underparts is intermediate between that of goldmani and leucoparia. The same is true as regards the amount of projection from the braincase of the anterior margins of the tympanic bullae. The squamosal immediately anterior to each tympanic bulla is flat instead of ventrally convex as in goldmani or ventrally concave as in leucoparia and frenata. In accordance with the custom adopted in this paper of referring every specimen to some one subspecies, this specimen from Cerro San Felipe is referred to Mustela frenata perotae, to the description of which it most nearly answers.

Possibly goldmani, as here constituted, is a composite form. The specimens from the high mountains closely resemble one another. However, a specimen, no. 68541 from "Finca El Cipres," Guatemala, which place Mr. G. Goodwin tells me is at an elevation of 2500 feet, approximately 5 miles north of Retalhuleu, has a basilar length of 47.3 and is correspondingly small in other parts. This suggests the existence of a small, lowland race on the western side of the central divide corresponding to perda and tropicalis on the eastern side. From only a few miles away, at San Sebastian, there is available, the adult skull of a still smaller animal. This skull only, no. 41026, in the Berlin Zoological Museum, has a basilar length of 46.1, zygomatic breadth of 27.4, and other cranial measurements notably smaller than those of specimens from the high mountains. A skin-only, no. 12038, collection of Donald R. Dickey, from La Cebia, altitude 2150 feet, near the city of San Salvador, seemingly represents an animal smaller than typical goldmani. This specimen from La Cebia has the light color of the underparts extended distally on the hind legs to the tips of the toes as in M. f. tropicalis. However, the upper parts are darker and resemble those of M. f. goldmani. A fourth specimen from only 3500 feet elevation, on the south side of Volcano Tajumulco, Guatemala, no. 41768, Field Museum of Natural History, a subadult male, measures only 490 in total length and has the least color of the underparts so restricted as to amount to only 22 per cent of the greatest width of the color of the upper parts. Both these features are suggestive of the lowland races.

These four specimens indicate that the lowland population on the western side of the divide is smaller than the mountain population. The juvenile from Carolina and a young male from Finca Cipres, however, both closely resemble individuals of goldmani from the higher mountains. All these animals here are referred to goldmani. More specimens may reveal an amount and a pattern of geographic variation in weasels of this region that will require application of another subspecific name.

The female, no. 68540, from Puebla agrees remarkably well with the skull of the female, no. 132528, of macrophonius. Differences displayed by the specimen from Puebla are its slightly narrower braincase and longer space between the foramen ovale and anterior end of the tympanic bulla. Considering the far eastern location of Puebla (just north of Río Motagua, at 89° W, according to a sketch map provided by Mr. G. G. Goodwin), this specimen might be expected to show some approach to the small lowland races. Actually, however, it displays the characters of goldmani better than does the subadult female from Volcano San Lucas, which is nearer the metropolis of goldmani, and I assume at a higher elevation than Puebla.

Concerning this weasel Merriam (1896:29) says: "Mr. E. W. Nelson writes me that this fine weasel is found sparingly in the forest about Pinabete, Chiapas, at an altitude of 7000 to 8000 feet (2100 to 2500 meters). The type specimen was shot in the afternoon while hunting on a heavily wooded hill slope. It was heard making long, slow leaps over the dry, crisp leaves. Coming to a log, it stood up and rested its fore feet on the log, in which position it was shot by Mr. Goldman."

The specimen taken by R. A. Stirton in Salvador comes from an elevation of 8000 feet in the rain forest of the Upper Tropical Life-zone. Mr. Stirton tells me that one morning on visiting his traps set for small rodents, he found in one the partly eaten remains of a Heteromys. Leaving these remains as found he placed a steel trap beside them and on the following morning found the male weasel in the trap.

At least three of the ten specimens had the frontal sinuses infested with parasites.

Specimens examined.—Total number, 15, listed by localities from north to south, and unless otherwise indicated in the American Museum of Natural History.

México: Chiapas: 20 mi. SE Teopisca, 2; Pinabete, 1.

Guatemala: Puebla, 1; Finca Porvenir, 3500 ft., S slope Volcan Tajumulco, 1; Sierra [= ? Cerro] Santa Elena, 9500 ft. (near Tecpám), 1; Carolina, 1; Volcano San Lucas, 1; "Finca El Cipres," 1; "Finca Cipres," 2500 ft., 1; Finca San Isidro, San Sebastión, Dept. Retalhuleu, 1; Dueñas, 1; no locality more definite than Guatemala, 1.

El Salvador: Los Esesmiles, 8000 ft., Chalatenango, 1; La Cebia, 2150 ft., near San Salvador, 1.

Mustela frenata macrophonius (Elliot)

Long-tailed Weasel

Plates , , , , , and

Putorius macrophonius Elliot, Proc. Biol. Soc. Washington, 18:235, December 9, 1905.

Mustela macrophonius, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912.

Mustela frenata macrophonius, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, adult, skull and skin; no. 14063, Field Mus. Nat. Hist.; Achotal, Veracruz, México; January 15, 1904; obtained by Edmund Heller and Charles M. Barber; original no. 3424.

The skull (plates -, ) is complete and unbroken. Excepting right P2, which has been aborted or broken away, all the teeth are present. The skin is well made and in good condition. As shown by the scrotal pouch, the specimen is a male.

Range.—Tropical Life-zone, probably into Boreal life-zones, of mountains along eastern border of southern Veracruz. See figure on page .

Characters for ready recognition.—Differs from M. frenata frenata and M. f. perotae and M. f. leucoparia in lacking color of underparts on hind feet and in larger skull (skulls of adult males with basilar length more than 52.5); from M. f. goldmani by larger size of skull (see above) and entire animal and wider tympanic bullae; from M. f. tropicalis and M. f. perda by larger size (total length of adult males more than 510), postorbital breadth amounting to less than combined length of upper premolars.

Description.—Size.—Male: External measurements of the type specimen, an adult, are: Total length, 598; length of tail, 246; length of hind foot, 59. Tail 70 per cent as long as body; length of hind foot more than basal length.

Female: The skin, without field collector's measurements, of an adult female from Pérez, Veracruz, shows this sex to be correspondingly large. Because the skin is understuffed and because the hind feet are skinned out, reliable measurements can not be obtained from the dried skin.

Externals.—As described in Mustela frenata goldmani except that all carpal vibrissae are of same color as upper parts and that hairiness of foot-soles is halfway between that shown in figures and .

Color.—As in darkest individuals of M. f. goldmani, thus, color of upper parts on posterior part of back near (n) Argus Brown. Color of underparts near (12) Mikado Orange in a juvenile, extending distally on posterior sides of forelegs onto inner toes and on hind legs to points between knees and heels. Least width of color of underparts 28 per cent of greatest width of color of upper parts. Black tip of tail 34 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on type specimen): See measurements and plates -, . As described in Mustela frenata frenata except that: Weight, 6.9 grams; basilar length, 54.1; zygomatic breadth less than distance between condylar foramen and M1 or that between anterior palatine foramen and anterior margin of tympanic bulla; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 6 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; anterior margin of masseteric fossa below posterior half of m2.

Female (based on no. 132528): See measurements and plates -. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 43.5; zygomatic breadth less than distance between condylar foramen and M1 and more or less than (in the single specimen, equal to) that between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more or less than (about equal to) outside length of P4; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale.

The skull of the female is 48 per cent lighter than that of the male.

Comparison of the skull with that of M. f. goldmani is made in the account of that subspecies. Similar differences probably exist between males of perotae and macrophonius. As compared with skulls of males of M. f. tropicalis and perda, the skull of the male of macrophonius is larger in every measurement taken. The postorbital constriction is less, rather than more, than the combined length of the upper premolars. Relative to the basilar length, the following measurements are less than in any specimen of tropicalis or perda: length of tooth-rows; orbitonasal length; depth of skull at posterior border of upper molars; and depth of skull at anterior margin of basioccipital.

Remarks.—This large weasel appears to have escaped the notice of naturalists until the spring of 1903 when J. Friesser obtained an adult female and juvenal male at Pérez for the collection of the United States Bureau of Biological Survey. These specimens were tentatively referred to Mustela tropicalis. In the following January, Edmund Heller and Charles M. Barber obtained the adult male that was made the type specimen by Elliot who did not see, or if he did, did not mention, the specimens from Pérez. He did, however, refer a young female from Xuchil, Veracruz, to his Putorius macrophonius. This young female is here referred to Mustela frenata tropicalis.

The extent of the geographic range of this subspecies is not well known.

Mustela frenata macrophonius and M. f. texensis are the largest American weasels. The basilar length in the type specimen is greater by one-tenth of a millimeter than in the type specimen of M. f. texensis. The other cranial measurements taken are greater in M. f. texensis. The skull of the female from Pérez is one of the largest skulls examined of that sex. The juvenal male has teeth as large as those of the type specimen and the skull is the largest for its age of any seen. Although the skin of the female is understuffed and hence does not provide reliable measurements, it shows that the female is also large.

The white bands in front of the ears are confluent with the white patch between the eyes on one side only in one specimen. It is the juvenal male. These bands are not confluent with the color of the underparts on either side in the female and on one side only in the adult male. None of the specimens has a white patch between the ears. The dark spot at each angle of the mouth is present only in the juvenile where it occurs on each side. Of the three specimens, the juvenile is the darkest and the adult male the lightest. The white facial markings are most extensive in the juvenal male and the least extensive in the adult female.

M. f. macrophonius most closely resembles M. f. goldmani but in the relatively flattened braincase, deep constriction of the postorbital region and general angularity of the skull approaches M. f. perotae and M. f. frenata.

Only one of the three skulls, that of the female, shows evidence of infestation of the frontal sinuses by parasites, and this did not result in malformation of the interorbital region.

Specimens examined.—Total number, 3, all from México, listed by localities from north to south.

Veracruz. Achotal, 1 (Field Mus. Nat. Hist.); Pérez, 2 (U. S. Nat. Mus.).

Mustela frenata tropicalis (Merriam)

Long-tailed Weasel

Plates , , , , , and

Putorius tropicalis Merriam, N. Amer. Fauna, 11:30, pl. 3, figs. 5, 5a, 6, 6a, text fig. 16, June 30, 1896; Merriam, Proc. Biol. Soc. Washington, 15:68, March 22, 1902.

Putorius frenatus, Merriam, N. Amer. Fauna, 11:27, June 30, 1896.

Mustela tropicalis tropicalis, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916.

Mustela frenata tropicalis, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, adult, skull and skin; no. 54994, U. S. Nat. Mus., Biol. Surv. Coll.; Jico, Veracruz, México; July 9, 1893; obtained by E. W. Nelson; original no. 5195.

The skull (plates -, ) is complete. All the upper incisors, except the second and third on the left side, are missing. The right upper canine is broken. The skin is well made and in good condition.

Range.—Up to 5000 feet (as now known) in Tropical Life-zone of Veracruz, México. See figure on page .

Characters for ready recognition.—Differs from M. f. frenata and M. f. perotae in least width of color of underparts not exceeding 36 per cent of greatest width of color of upper parts and in postorbital breadth exceeding length of upper molar and premolar tooth-rows; from M. f. macrophonius and M. f. perda in least width of color of underparts averaging more than 29 per cent of greatest width of color of upper parts; and from M. f. perda by longer tympanic bullae which in males are more than 14.9; and from M. f. macrophonius by lesser basilar length (not more than 48) and in postorbital breadth exceeding length of upper molar and premolar tooth-row.

Description.—Size.—Male: The type specimen and no. 12764/11058, a subadult, from Jalapa, Veracruz, measure, respectively, as follows: Total length, 444, 442; length of tail, 175, 160; length of hind foot, 50, 47. The tail is 65 and 57 per cent as long as the head and body. The hind foot is more or less than (approximately equal to) the basilar length.

Female: Merriam (1896:31) gives the measurements of a female topotype (probably no. 54993, U. S. Nat. Mus., which has no measurements written on the attached label) as: Total length, 333; length of tail, 121; length of hind foot, 37. The length of the tail amounts to 57 per cent of the length of the body. The length of the hind foot of no. 54993, U. S. Nat. Mus. is the same as the basal length.

The differences in external measurements between the male and the female topotypes are: Total length, 111; length of tail, 54; length of hind foot, 13.

Externals.—As described in Mustela frenata frenata except that carpal vibrissae do not reach apical pad of fifth digit and hairiness of foot soles is less.

Color.—As described in M. f. frenata except that: Blackish of head extends half way or more from ears to shoulders; upper parts near (14) Brussels Brown or slightly faded tone 2 of Maroon of Oberthür and Dauthenay, pl. 341; underparts of juvenal pelage near (a) Ochraceous-Buff. Least width of color of underparts averaging (in three specimens from Jico and one from Jalapa) 34 (extremes 30-37) per cent of greatest width of color of underparts. Black tip of tail, in two male topotypes, 57.5 (55 and 60) mm. long; thus longer than hind foot and in each individual comprising 34 per cent of length of tail-vertebrae.

As compared with M. f. frenata: White facial markings slightly less extensive; blackish (not black) of head extending onto neck; upper parts slightly darker; ventral side of tail noticeably darker; color of underparts more restricted, averaging approximately one-third rather than nearly one-half width color of upper parts; black tip of tail one-third rather than one-fourth length of tail and much longer than hind foot. Similar differences of lesser amount exist between perotae and tropicalis. M. f. perda, macrophonius and goldmani bear the opposite relation to tropicalis. That is to say, in the latter: White facial markings slightly more extensive; blackish of head less extended over neck; upper parts markedly lighter; color of underparts less restricted and black tip of tail shorter.

Skull and teeth.—Male (based on type specimen and a subadult, no. 11058, from Jalapa): See measurements and plates -, . As described in Mustela frenata perda except that: Weight 4.7 (4.6 and 4.7) grams; basilar length 46.7 (45.5 and 47.8); zygomatic breadth more or less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 (including I3) upper incisors; anterior margin of masseteric fossa below middle of m2 or posterior to that tooth.

Female (based on no. 54993 and no. 1060): See measurements and plates -. As described in Mustela frenata perda except that: Weight (of 54993) 2.2 grams; basilar length, 37.5 (36.0-39.0); zygomatic breadth more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more than greatest length of P4; height of tympanic bulla equal to one-third to three-fourths of distance from its anterior margin to foramen ovale.

The skull of the adult female is 53 per cent lighter than that of the type specimen, a male.

Comparison of the skulls of males and females with those of M. f. perda, the nearest relative, has been made in the discussion of that subspecies. Some of the features that readily distinguish skulls of M. f. tropicalis from those of M. f. frenata, perotae and macrophonius are as follows: Weight less than 4.8 grams; basilar length less than 48; postorbital breadth more than length of upper M-Pm tooth-row. The skulls of male frenata, perotae and macrophonius are much larger, heavier, and are decidedly more angular with more constricted postorbital region the least width of which is less than the length of the upper premolars. In frenata the anterior margins of the tympanic bullae are protruded much farther from the braincase. The skull of the female of M. f. tropicalis is smaller, weighing less than 3 grams; basilar length less than 41; postorbital breadth more than length of upper molar and premolar tooth-row.

Remarks.—This subspecies was originally described by Merriam as a full species. Later he described Putorius tropicalis perdus as another subspecies. Allen (1916) placed P. t. perdus in synonymy but named Mustela tropicalis nicaraguae as new. In the present paper all three forms are recognized but are regarded as only subspecifically distinct from the other bridled weasels of México and Central America.

The limits of the geographic range of tropicalis are fairly well known on the south and west but the only specimen available from the tropical coastal region north of Jico, is a young female from a point 50 miles south of Victoria. Thus, how far north along the coast it ranges toward Matamoros, where M. f. frenata occurs, is not known. The three specimens from Jico, a young female from Jalapa and another adult collected by J. Potts and labeled as coming from México City, are assumed to be typical. The latter specimen certainly came from an elevation lower than that of México City because M. f. frenata occurs there. Although the female from Jalapa, agrees well with specimens from Jico, a male, no. 12764/11058, from Jalapa, has a relatively broader skull, as in perda, although the tympanic bullae are short as in tropicalis. The resemblances to perda in features of coloration are: slightly darker upper parts, and the termination just below the knees of the color of the underparts. There are three specimens labeled as from Orizaba that indicate intergradation with perotae as does also the coloration of the juvenal female from 5 kilometers north of Jalapa. The specimens labeled as from Orizaba are old, poorly-prepared skins, only two of which have partial skulls. The size and coloration of the skins suggest perotae as do also the partial skulls in some respects although the skulls show greater resemblance to those of tropicalis.

The topotype, female, no. 54993, was figured by Merriam (1896, fig. 16, p. 31) along with that of what now is the type specimen of M. f. perotae. Merriam called attention to the great difference in size between the skulls of the two sexes of M. f. tropicalis and compared the condition to that found in noveboracensis. Although the skull of the female from Jico is fully adult, it probably is exceptionally small.

The young female from Xuchil is indistinguishable in coloration from the juvenal female of M. f. perotae from Perote, but in size of skull and size of teeth is intermediate between the female of tropicalis from Jalapa and the females from Cofre de Perote.

There is then, indication of intergradation with M. f. perotae as well as with M. f. perda. M. f. tropicalis differs from M. f. perotae and M. f. frenata in about the same way that M. f. perda differs from M. f. goldmani and M. f. macrophonius. M. f. tropicalis and perda each is smaller and more intensely colored than goldmani and macrophonius, and inhabits the lowland to the east of their highland relative.

At least five of the nine skulls have the frontal sinuses infested by parasites.

Specimens examined.—Total number, 13, all from México, listed by localities from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.

Tamaulipas. 50 mi. S Victoria, 1. Near? México City, 1.

Veracruz. Jico, 3; 5 km. N Jalapa, 1; Jalapa, 2 (1, 1); Xuchil, 1; Orizaba, 4 (2, 1).

Mustela frenata perda (Merriam)

Long-tailed Weasel

Plates , , , , , and

Putorius tropicalis perdus Merriam, Proc. Biol. Soc. Washington, 15:67, March 22, 1902.

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Mustela tropicalis perda, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912.

Mustela tropicalis tropicalis, Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916.

Mustela frenata perda, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, subadult, skull and skin; no. 100041, U. S. Nat. Mus., Biol. Surv. Coll.; Teapa, Tabasco, México; March 31, 1900; obtained by E. W. Nelson and E. A. Goldman; original no., 14074.

The skull (plates -, ) is unbroken and all the teeth are present and entire. The skin is well made and in good condition.

Range.—Fifty feet (Chichén Itzá) to 4000 feet (San Vicente) in Lower Tropical Life-zone south from southern Veracruz through southern México into Guatemala. See figure on page .

Characters for ready recognition.—Differs from M. f. nicaraguae in lesser extent of color of underparts (not more than 22 per cent of greatest extent of color of upper parts), black tip of tail more than 38 per cent of length of tail, and broader skull (in adult males, mastoid breadth more than 23.9 and zygomatic breadth more than 27.4); from M. f. tropicalis in more restricted color of underparts (least width of color of underparts less than 28 per cent of greatest width of color of upper parts) and shorter tympanic bullae, which in males are less than 15; from M. f. goldmani by total length not exceeding 489, least width of color of underparts not exceeding 24 per cent of greatest width of color of upper parts, weight of adult skull less than 5 grams and basilar length less than 48.5.

Description.—Size.—Male: The type specimen and another subadult from San Vicente, Chiapas, measure, respectively, as follows: Total length, 473 and 443; length of tail, 184 and 169; length of hind foot, 51 and 51.5. The tail is 62 and 64 per cent as long as the head and body. The length of the hind foot is greater than the basal length.

Female: Estimates made from the dried skin of no. 218036 are: Total length, 375; length of tail, 140; length of hind foot, 40. The hind foot of no. 65422 from Catemaco also measures 40.

The average differences in external measurements of the two sexes are: Total length, 83; length of tail, 37; length of hind foot, 11.

Externals.—As described in Mustela frenata goldmani except that hairiness of foot soles is slightly less.

Color.—As described in Mustela frenata goldmani except that: back near (n) Argus Brown or Carbon Brown, tone 3, of Oberthür and Dauthenay, pl. 342; underparts Ochraceous-Buff. Least width of color of underparts, in four specimens, averaging 20 (extremes 18-22) per cent of greatest width of color of upper parts; black tip of tail, in two subadult males, averaging 48 (extremes 46-49) per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on type specimen and subadult no. 132997 from San Vicente): See measurements and plates -, ; weight 4.4 grams (same for each); basilar length 45.7 (45.3 and 46.1); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more or less than (approximately equal to) length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum not greater than length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4-1/2 to 5-1/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below middle of m2.

Female (based on two subadults, nos. 65422 and 218036): See measurements and plates -; weight, 2.4 (2.3-2.5) grams; basilar length, 40.5 (40.4-40.6); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than outside length of P4 and less than inside length of same; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla equal to one-third to one-half distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than (about equal to) length of rostrum.

The skull of the female averages 48 per cent lighter than that of the male.

Comparison of the skull of the male with that of M. f. nicaraguae has been made in the account of that subspecies. The skull of the male as compared with that of M. f. tropicalis has shorter tympanic bullae, deeper braincase at anterior margin of basioccipital, lesser zygomatic and palatal breadth and smaller P4 and m1. The skull of the female is larger in every measurement taken except those reflecting width of the preorbital portion. This part is actually narrower but probably mainly because the females of perda are younger than those of tropicalis. Features in which three skulls of subadults of M. f. perda differ from the five adults of M. f. goldmani and show no overlap are: lesser basilar length, lesser weight, greater relative length of upper tooth-rows, greater relative width of rostrum, greater relative length of rostrum, lesser mastoid and zygomatic breadths, lesser width, length and height of tympanic bullae; lesser outside length of P4 and greater relative depth of braincase at anterior margin of basioccipital and at posterior margin of M1. Features in which perda averages less are: length of tooth-rows, interorbital breadth, orbitonasal length, relative zygomatic breadth, length of m1, outside and inside lengths of P4, width and length of M1, and depth of skull at posterior margin of M1. Features in which perda averages more than goldmani are: relative interorbital breadth, relative mastoid breadth and depth of skull at anterior margin of basioccipital. The length of the inner half of M1 averages the same. As compared with goldmani, the skull of the male of perda is shorter, otherwise generally smaller, but relatively broader except across the zygomatic arches, and relatively deeper. The anterior margins of the tympanic bullae project slightly less from the braincase and the squamosals immediately in front of these bullae are slightly more convex ventrally.

Remarks.—Described by Merriam in 1902 as a subspecies of Putorius tropicalis, the form perda was regarded by Allen (1916:99) as not subspecifically distinct from P. t. tropicalis.

This is the eastern, lowland subspecies of the Tropical Life-zone, corresponding to M. f. goldmani of the higher mountains just as M. f. tropicalis corresponds to M. f. frenata and perotae of the high mountains and table land. The difference in size between perda and nicaraguae and between perda and tropicalis is slight. M. f. perda is slightly less richly colored than M. f. nicaraguae but has the color of the underparts more restricted and has a longer black tip on the tail. In these respects it is second only to M. f. panamensis among Central American weasels. Evidence of intergradation with goldmani is furnished by the specimens from Cobán, Guatemala, and the nearby locality San Cristóbal in Verapaz, Guatemala. Reduced size as compared with goldmani suggests affinity with perda but the greater width of the light-colored underparts, which averages 24 (extremes 18-32) per cent of the greatest width of the color of the upper parts, shows approach to goldmani. Farther north, in Chiapas, however, specimens of perda from San Cristóbal and San Vicente are readily distinguishable from those of goldmani taken a few miles away at Pinabete and near Teopisca. The latter two localities are, however, several thousand feet higher than San Cristóbal (Chiapas) and San Vicente.

Two of the nine skulls (only 3 adult) examined for malformation of the frontal sinuses reveal infestation by parasites.

Specimens examined.—Total number, 18, listed by localities from north to south, and unless otherwise indicated in the United States National Museum.

Veracruz. Catemaco, 1.

Tabasco. Teapa, 2 (1).

Chiapas. San Cristóbal, 1; San Vicente, 1; no locality more definite than state, 1.

Yucatán. Chichén-Itzá, 1.

Guatemala: Cobán, 2 (1, 1); Finca la Providenci, S. Cristóbal, Verapaz, 3; central Guatemala, 1; no locality more definite than Guatemala, 5 (2).

Mustela frenata nicaraguae Allen

Long-tailed Weasel

Plates 1, 25, 26, 27 and 30

Mustela tropicalis nicaraguae Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916.

Putorius tropicalis, Allen, Bull. Amer. Mus. Nat. Hist., 24:661, 1908.

Mustela frenata nicaraguae, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, subadult, skull and skin; no. 30754, Amer. Mus. Nat. Hist., Matagalpa, Nicaragua; April 16, 1910; obtained by W. B. Richardson; original no., 712.

The skull (plates -, ) of the type specimen lacks the entire right zygomatic arch. Otherwise it is complete. The teeth all are present and unbroken. The skin is complete and unfaded but only partly stuffed.

Range.—Honduras and Nicaragua. Altitudinal and zonal limits unknown. See figure on page .

Characters for ready recognition.—Differs from M. f. costaricensis and M. f. goldmani in shorter black tip of tail (not more than 35 per cent of length of tail) and lesser width (usually not more than 7 mm.) of tympanic bulla; from M. f. perda in greater extent of color of underparts (22 or more per cent of greatest width of color of upper parts), shorter black tip on tail (not more than 35 per cent as long as tail) and narrower skull, the mastoid breadth in adult males being less than 23.9 and the zygomatic breadth less than 27.

Description.—Size.—Male: Average and extreme measurements of five subadults and one young (four from Matagalpa and one from San Rafel del Norte) are: Total length, 450 (420-480); length of tail, 178 (150-190); length of hind foot, 48 (46-50). Tail averages 65 (extremes 56-69) per cent as long as head and body. Length of hind foot (measurements from dried skins) more than basal length.

Female: Measurements unrecorded.

Externals.—As described in Mustela frenata goldmani, except that hairiness of foot soles (between that shown in figures and ) is less, slightly less even than in M. f. perda.

Color.—As described in Mustela frenata goldmani except that: Back near (n) Argus Brown, or Carbon Brown, tone 4 of Oberthür and Dauthenay, pl. 342. Underparts Ochraceous-Buff. Least width of color of underparts, in four males, young, subadult and adult, 24 (extremes 22-26) per cent of greatest width of color of upper parts; the corresponding per cent in one female is 32; black tip of tail, in two subadult males, averaging 29 (extremes, 28-30) per cent of length of tail-vertebrae; corresponding per cent in one female, 36.

Skull and teeth.—Male (based on type specimen, one adult topotype [?] and one subadult from San Rafel del Norte): See measurements and plates -, . As described in Mustela frenata perda except that: Weight, 4.2 grams (estimated for adults); basilar length 45.0 (44.8-45.5); interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of four to five upper incisors; length of tympanic bulla not less than length of lower molar and premolar tooth-row; anterior margin of masseteric fossa below anterior margin of m2 or posterior to that tooth.

Female: Skull unknown.

Comparison of the skull of the male with that of M. f. costaricensis is made in the account of that subspecies. As compared with that of M. f. perda, which it most closely resembles, the skull of the male has a narrower, shorter rostrum, lesser interorbital breadth, lesser mastoid and zygomatic breadths and slightly shallower braincase, measured at anterior margin of basioccipital. The tympanic bullae are slightly less projected, at their anterior margins, from the braincase and the squamosal, directly anterior to each, is a little more convex ventrally. The skull of M. f. nicaraguae is, then, slightly shorter than that of M. f. perda and relatively narrower.

Remarks.—When naming this form, Allen (1916:100) characterized it as "Similar to M. tropicalis tropicalis but general coloration much darker and the white face markings somewhat reduced in area." In the sentence preceding the one quoted, Putorius tropicalis perdus was placed as a synonym of Putorius tropicalis tropicalis. M. f. nicaraguae and M. f. perda are nearly alike in color and color pattern but differ in cranial characters. M. f. perda and M. f. tropicalis are widely different in color and more especially in color pattern but differ only slightly in cranial characters. The aggregate difference between perda and nicaraguae is less than that between perda and tropicalis. All three are lowland forms and each is smaller than the adjacent highland forms, namely, M. f. goldmani, macrophonius, perotae and frenata.

The weasels from Honduras definitely are not typical of nicaraguae as it is known from the specimens from Nicaragua itself. The specimens from the state of Tegucigalpa, Honduras, are larger. Some are darker than topotypical nicaraguae. The dorsal outline of the skull is more nearly flat (less convex) in some. In these and several other differential features studied, the average of specimens from Tegucigalpa is intermediate toward goldmani, but everything considered the animals seem best placed with nicaraguae rather than with goldmani or perda, to which latter also, they show some resemblance. With better material from Nicaragua and additional specimens from Salvador (here referred to goldmani) a restudy of all the material now referred to the three races named would be profitable. Aims of this restudy might be to determine if a highland race additional to goldmani should be recognized and if the lowland races perda and nicaraguae differ from one another in the way that the existing specimens indicate.

In the five males from Matagalpa, the narrow white band in front of each ear is confluent with the color of the underparts on one side only in one specimen and on both sides in two specimens. None of these bands is confluent with the white patch between the eyes. A dark spot at the angle of the mouth is present on one side in one specimen. The corresponding area is dark colored in all other specimens but not separated from the color of the upper parts. In the specimen from San Rafel del Norte the white bands are not confluent with the color of the underparts. The female from Mambacho has the mentioned bands confluent with the color of the underparts. This female approaches M. f. costaricensis in the dark color of the upper parts but has more extensive white facial markings than some specimens from much farther north. Like a female seen of M. f. costaricensis, this one has a "frosted" nape but the white hairs on the back of the neck are less numerous than in the female of M. f. costaricensis.

M. f. nicaraguae in typical form, then, is thought of as a small, lowland, tropical subspecies only slightly differentiated from M. f. perda. By reason of its intermediate characters, it constitutes a link between the lowland forms, and the larger animals called M. f. goldmani and M. f. costaricensis.

None of the four skulls from Nicaragua shows signs of infestation of the frontal sinuses by parasites.

Specimens examined.—Total number, 16, listed by localities from north to south. Specimens are in the American Museum of Natural History, unless otherwise indicated.

Honduras: Alto Cantoral, 2; Cerro Grande La Paz, 1. La Flor Archaga, 1; Comayagüela, 1; vicinity of Tegucigalpa, 2; no locality more definite than Honduras, 1.

Nicaragua: San Rafel del Norte, 1; Matagalpa, 6; Ma[o]mbacho, 1.

Mustela frenata costaricensis Goldman

Long-tailed Weasel

Plates , , , , and

Mustela costaricensis Goldman, Proc. Biol. Soc. Washington, :9, January 23, 1912.

Mustela brasiliensis, Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):374, 1874.

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Putorius affinis, Merriam, N. Amer. Fauna, 11:31, June 30, 1896 (part).

Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Lönnberg, Arkiv för Zool., 14(no. 4):16, 1921.

Mustela frenata costaricensis, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, young, skull and skin; no. 13770/37149, U. S. Nat. Mus.; San José, Costa Rica; obtained by C. H. Van Patten.

The skull (plates -) is complete and unbroken. All teeth are present and unworn. The skin apparently has been remade. It lacks the distal two-thirds of the tail. The head is somewhat shrunken. The color is possibly faded but if so only to a slight degree. Otherwise, the skin is in good condition. The orange color of the underparts is so intense as to suggest that the full, adult pelage has not been acquired. No white markings are present on the face. There is no sex mark on the label attached to the skin but the size and proportions of the skull and the scrotal pouch on the skin prove that the specimen is a male. The presence of sutures on the dorsal face of the rostrum and the short, wide, and low sagittal crest show the specimen to be young.

Range.—Costa Rica. Altitudinal and zonal range unknown. See figure on page .

Characters for ready recognition.—Differs from M. f. panamensis in lighter color of upper parts (tone 2 rather than tone 4 of Reddish Black of Oberthür and Dauthenay, pl. 344) and longitudinally flat interorbital region of skull; from M. f. nicaraguae in darker color of upper parts (of Oberthür and Dauthenay, tone 2 of pl. 344 rather than tone 4 of pl. 342) and greater width (more than 7) of tympanic bulla.

Description.—Size.—Male: No collector's measurements available of fully grown animals. Estimated measurements of adult males: Total length, 470; length of tail, 165; length of hind foot (taken from dried skins of 3 adults), 52 (50-52). Tail estimated to average 55 per cent as long as head and body. Length of hind foot more or less than (about equal to) basal length.

Female: A subadult or adult, from the Candelaria Mountains, and a subadult from Irazú, measure, respectively: Total length, 370, 385; length of tail, 130, 150; length of hind foot, 40, 31. Tail 59 per cent as long as head and body. Length of hind foot probably about equal to basal length.

The estimated differences in external measurements of the two sexes are: Total length, 92; length of tail, 25; length of hind foot, 16 (probably average difference is less).

Externals.—As described in M. f. panamensis (figure ) except that foot soles are slightly more hairy.

Color.—As described in Mustela frenata panamensis except that: back is near Reddish Black, tone 2 of Oberthür and Dauthenay, pl. 344; chin, lips, and throat white or whitish; remainder of underparts near (c) Ochraceous-Buff; color of underparts rarely extending distally onto toes of forefeet. Least width of color of underparts, in eleven specimens, averaging 23 (10-36) per cent of greatest width of color of upper parts; black tip of tail, in six specimens, averaging 36 (31-38) per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on 2 adults, no. 3.2.1.6. from vicinity of San José and no. 11408, U. S. Nat. Mus., from "Costa Rica"): See measurements and plates -; weight, 5.9 grams; basilar length 49 +; zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth in undiseased skulls less than length of upper premolars (less than distance between posterior borders of P2 and P4) and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum more or less (about equal to) length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; tympanic bulla longer or shorter than (about equal to) lower molar and premolar tooth-row and longer or shorter than (about equal to) rostrum; anterior margin of masseteric fossa directly below posterior border of m2.

Female: Skull of adult unknown.

Comparison of the skull of the male with that of M. f. panamensis has been made in the account of that subspecies. As compared with that of M. f. nicaraguae the skull of M. f. costaricensis is heavier and in every measurement taken is larger. The skull is generally more massive and it follows that most measurements of depth and width are greater in relation to the basilar length as well as actually greater. The individual teeth are larger and the tympanic bullae wider and at their anterior ends are more projected from the braincase. Indeed the skull is more like that of M. f. goldmani than like that of M. f. nicaraguae.

Remarks.—The half dozen ill-prepared skins, with partial skulls inside, of this form in the United States National Museum long were referred either to Mustela brasiliensis or Mustela affinis. It was not until 1912 when Goldman studied these specimens that the distinctive characters of the Costa Rican weasel were recognized and made the basis of the name costaricensis.

M. f. costaricensis is well differentiated from M. f. nicaraguae and M. f. goldmani which occur to the northward and from M. f. panamensis which occurs to the southward and is a large, heavy-skulled, dark-colored animal with white facial markings restricted or absent. In the type specimen and the female from the Candelaria Mountains the white facial markings are only narrow facial bars or a few white hairs, but in the young male from Cervantes there is a well developed bar 6 millimeters wide on each side of the face and a separate nasofrontal spot, 10 x 12 mm. The young female from Cachí has a V-shaped frontonasal spot, on the right side of the face a white bar 5 mm. wide and 17 mm. long connected with the color of the underparts, and on the left side a white spot in front of the ear and another between the ear and eye. White facial markings were not recorded in the other specimens. The color of the upper parts is only a little less dark than those of M. f. panamensis. Owing to the numerous white hairs on the dorsal side of the neck, the nape of the female from the Candelaria Mountains has a frosted appearance not present in other specimens.

M. f. costaricensis is a large animal and among its geographic neighbors is approached in size only by a specimen of panamensis from Boquete, Panamá. Also the young male from Cervantes suggests panamensis in the less flattened interorbital region, but even so is more like costaricensis. The small size of two young males, one from Navarro and the other from the vicinity of San José, is suggestive of M. f. nicaraguae. However, the large size of most of the specimens and the configuration of the skull are more as in M. f. goldmani than in M. f. nicaraguae and thus suggest that the known specimens are of high mountain subspecies. The long black tip of the tail is another point of resemblance to M. f. goldmani, the high mountain subspecies to the north. Perhaps in the lowlands of Costa Rica, there are weasels of another subspecies.

Of the eight skulls examined for malformation of the frontal sinuses, each of the two adults and two subadults shows signs of having the frontal sinuses infested with parasites.

Specimens examined.—Total number, 14, listed by localities from north to south.

Costa Rica: Irazú (Frasu or Irasu on label), 3000 M., 1; Cervantes, 1; San José, 1; vicinity of San José, 2; Azahar Cartago, 1; Tucurrique, 1; Cachí, 1; El Muñco [= Muñeco?] (Río Nivarro [= Navarro?]), 4000 ft., 10 mi. S Cartago, Caribbean Slope, 1; Navarro, 1; Candelaria Mts., 1; no locality more definite than Costa Rica, 3.

Mustela frenata panamensis Hall

Long-tailed Weasel

Plates , , , , , and

Mustela frenata panamensis Hall, Proc. Biol. Soc. Washington, 45:139, September 9, 1932; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Mustela brasiliensis, Alston, Biol. Cent. Amer., Mammalia, p. 78, 1879.

Putorius affinis, Bangs, Bull. Mus. Comp. Zoöl., 39:49, April, 1902; Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.

Mustela affinis, Goldman, Proc. Biol. Soc. Washington, 25:10, January 23, 1912; Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.

Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Goldman, Smithsonian Miscel. Col., 69 (no. 5): 161, 1920.

Type.—Female, subadult, skull and skin; no. 170970, U. S. Nat. Mus., Biol. Surv. Coll.; Río Indio, Canal Zone, near Gatún, Panamá; February 17, 1911; obtained by E. A. Goldman; original no. 20897.

The skull is complete and unbroken. The left lower incisor is broken off but all the other teeth are present and entire. The skin is well made and seems to be in faded, worn, first, adult pelage.

Range.—Sea level (type locality) to 5800 feet (Boquete, see Bangs [1902:49]); Upper Tropical and Lower Tropical life-zones of Panamá. See figure on page .

Characters for ready recognition.—Differs from both M. f. meridana and M. f. costaricensis in darker tone (tone 4 of Oberthür and Dauthenay, pl. 344) of color of upper parts and in convex dorsal outline of skull (Compare figures of mentioned subspecies on plates -).

Description.—Size.—Male: Two adults from Boquete in the Museum of Comparative Zoölogy, nos. 10112 and 10113, measure, respectively, as follows: Total length, 480 and 400; length of tail, 170 and 143; length of hind foot, 52 and 43. Hind feet of two other adult males measure 46 on dried skins. Tail, in two specimens mentioned above, is 55 and 56 per cent as long as head and body. Length of hind foot, in each of three adults, slightly longer than basal length. Corresponding measurements of no. 178970 from Mt. Pirre are: 422, 164, 50. Tail 64 per cent (same per cent as in young male, no. 137514 from Boquete) as long as head and body, and hind foot longer than basal length.

Female: An adult and a young from Chiriquí, nos. 18434 and 18435 (Acad. Nat. Sci. Philadelphia), measure, respectively: Total length, 372, 389; length of tail, 138, 144; length of hind foot, 42, 41. The type specimen measures: Total length, 408; length of tail, 159; length of hind foot, 46.5. Tail 64 per cent as long as head and body, and hind foot longer than basal length.

The average differences in external measurements of the two sexes from the vicinity of Boquete are: Total length, 59; length of tail, 15; length of hind foot, 6.

Externals.—Longest facial vibrissae black and extending beyond posterior border of ear; carpal vibrissae wholly, or in part, black and extending as far as hypothenar pad; hairiness of foot-soles as shown in figure .

Color.—Usually, posterior fourth of each upper lip and sometimes few hairs in front of ear, white; sides and top of head and neck posteriorly to, or behind, shoulders, black; dark areas at angles of mouth confluent with color of upper parts; tip of tail, black; remainder of upper parts near (n) Bay of Ridgway and Reddish Black, tone 4, pl. 344 of Oberthür and Dauthenay; chin and lips, whitish; remainder of underparts Warm Buff or near (16´ c) Ochraceous-Buff; near (12) Salmon-Orange in juveniles and small young; color of underparts extending distally on posterior sides of forelegs to wrists, but not to soles, and on hind legs to or slightly below knees. Least width of color of underparts, in seven specimens, averaging 18 (extremes 11-28) per cent of greatest width of color of upper parts; black tip of tail, in five adults and subadults, averaging 45 (extremes 41-50) per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on three adults from Boquete): See measurements and plates -; weight, 5 (4.5-5.4) grams; basilar length, 45.2 (42.8-48.3); zygomatic breadth more or less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum approximately same (more or less than) length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4-1/2 to 5-1/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or slightly less than (approximately equal to) length of lower molar and premolar tooth-row or length of rostrum; anterior margin of masseteric fossa directly below posterior fourth of m2.

Female (based on subadult, type specimen and one adult from Siola): See measurements; weight, 3.3 and 2.1 grams; basilar length, 41.3 and 39.3; zygomatic breadth more than distance between condylar foramen and M1 and more or less than (about equal to) that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than combined length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than length of P4 (less in the adult); anterior margin of tympanic bulla as far posterior to foramen ovale as width of five upper incisors; height of tympanic bulla less than (about half) distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of lower molar and premolar tooth-row or than rostrum.

The skull of the one adult female from Chiriquí is 58 per cent lighter than the average of the two adult males.

The skull of the male of M. f. panamensis as compared with that of M. f. meridana, is heavier and averages larger in nearly every measurement taken. Relative to basilar length, tooth-rows, orbitonasal length, interorbital breadth and zygomatic breadth averaging narrower. Mastoid breadth always narrower. Tympanic bullae longer, narrower, and usually slightly less protruded. P4 and m1 larger. Dorsal outline of skull, viewed laterally, more convex. Postorbital breadth actually and relatively greater. Postorbital processes, mastoid processes, and sagittal crest not so well developed. Differences between skulls of females, in so far as known, similar to those described between males.

As compared with M. f. costaricensis, M. f. panamensis has a lighter skull averaging smaller in every measurement taken except interorbital breadth, which is greater. Relative to basilar length, width of rostrum, interorbital breadth and depth of skull at plane of upper molars, less. Tympanic bullae shorter, narrower, less protruded. P4, M1, and m1 larger. Dorsal outline of skull, viewed laterally, more convex. Postorbital breadth relatively and actually greater. Postorbital processes, mastoid processes, sagittal crest and lambdoidal crest less developed. No skull of an adult female of M. f. costaricensis is available for comparison.

Remarks.—This subspecies had not been recognized by previous workers because specimens from Panamá were supposed to be Mustela affinis Gray up until 1916, when Allen (1916:100) restricted the type locality of M. affinis to Bogotá, Colombia. At that time Allen referred specimens from Panamá to Mustela affinis costaricensis, and Goldman (1920:161) followed Allen.

The specimens examined show much variation. Part of this is geographic variation. For instance the specimens from Boquete approach M. f. costaricensis in size more than do those from farther south. Too few adult females have been seen to ascertain the amount of secondary sexual variation. Bangs (1902:49) suggested that the sex of no. 10113 was wrongly recorded and that it was not really a male. If so, this would reduce the range of apparent variation in size of males from Boquete by half and bring it into accord with the amount normally existing in adult males from one locality. No. 10113 is adult but the skin shows no mammae which would prove it to be a female instead of a pigmy male. Although even smaller than 10113, the type specimen is so much larger than females of M. f. meridana that I have wondered if it is correctly sexed. However, the fact that it was sexed by E. A. Goldman, a collector of wide experience, lessens the possibility that a mistake was made.

The color of the underparts is more restricted in panamensis than in any other subspecies of the species. Excluding the specimen from Mt. Pirre, the least width of color of the underparts averages 16 (extremes 6-24) per cent of greatest width of the color of the upper parts. This feature, together with the black color, imparts an appearance to the Panamá weasel that is strikingly like that of a mink. M. f. panamensis is one of the two blackest weasels; M. f. aureoventris is the other. Each of these subspecies occurs in a region of heavy rainfall and there clearly is a positive correlation between high humidity and intensity of color. The black tip of the tail, as regards extent, here reaches the maximum attained among Central and South American weasels. The foot soles are less hairy than in any other member of the subgenus Mustela. The tympanic bullae are lower and less inflated than in any other subspecies of the species.

Adequate specimens from central and southern Panamá may reveal the existence of one or more additional subspecies since animals from each of the three localities now represented differ from those from the other two and some of these differences are correlated with geographic position. However, specimens from all three localities agree in several features. For example all of them have the dorsal outline of the skull highly convex, transversely, and, more especially, longitudinally. In this respect they are sharply differentiated from any other American weasel. Nevertheless, M. f. panamensis is clearly a link between the North and South American subspecies and panamensis intergrades with the adjacent subspecies. The large size of the skull and teeth and the slightly more ventrally projected tympanic bullae of no. 10112 from Boquete approach features seen in M. f. costaricensis. The smaller size of skull and teeth of no. 178970 from Mt. Pirre are points of resemblance to M. f. meridana.

The type specimen was selected from a region where M. f. panamensis is thought to have its distinctive characters well developed. The specimen is not adult and, therefore, does not show as many differential characters as does a nontypical adult from Boquete. Nevertheless, the majority of the above mentioned differential characters are shown by the type specimen and an adult from the same place would, it is judged, show all the differential characters better than would an adult from Boquete.

Of the 11 skulls examined, 6 show no signs of having had the frontal sinuses infested with parasites.

Specimens examined.—Total number, 19, listed by localities from north to south and unless otherwise indicated in the United States National Museum.

Panamá: Boquete, 10 (3, 1, 1, 3, 1); Río Gariche [é], 5300 ft., 1; Siola, 1; Chiriquí, 1; Río Indio, near Gatún, 1; Mt. Pirre, 3 (2); Calovebora, 1 (locality not found, possibly misspelling of Calovébora); no locality more definite than Panamá, 1.

Mustela frenata meridana Hollister

Long-tailed Weasel

Plates , , , , and

Mustela meridana Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.

Putorius affinis, Robinson and Lyon, Proc. U. S. Nat. Mus., 24:147, October, 1901; Allen, Bull. Amer. Mus. Nat. Hist., 30:256, December 2, 1911.

Mustela affinis, Osgood, Field Mus. Nat. Hist. Publ. 155, zoöl. ser. 10:61, January 10, 1912.

Putorius macrurus, Allen, Bull. Amer. Mus. Nat. Hist., 31:92, April 19, 1912.

Mustela affinis affinis, Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916 (part).

Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916 (part).

Mustela frenata meridana, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.

Type.—Male, subadult, skull and skin; no. 123341, U. S. Nat. Mus., 1630 meters elevation, Montes de Mérida, near Mérida, Venezuela; August 14, 1903; obtained by S. Briceno.

The skull (plates and ) lacks the right exoccipital condyle and posterior half of the right zygomatic arch. The teeth all are present, unworn and entire. The skin is well made and complete.

Range.—Near sea level (San Julián) to 8500 feet (Montes de Culata, Mérida, Venezuela), and 9000 feet (Santa Elena, Colombia). Temperate to Subtropical life-zones of Venezuela and northern and western Colombia. See figure on page .

Characters for ready recognition.—Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, presence of p2 and two roots, rather than one root, on P2; from M. frenata panamensis in lighter color of upper parts (tone 3 rather than tone 4, pl. 344, Reddish Black, Oberthür and Dauthenay), flat rather than convex dorsal outline of skull immediately behind postorbital processes (see pl. 27); from M. f. affinis, in males, by lesser average breadth and length of skull and greater actual and relative size (see measurements) of facial part of skull; from M. f. aureoventris, in males, by lighter upper parts (tone 3 rather than tone 4, pl. 344, Reddish Black, Oberthür and Dauthenay) and by smaller skull and teeth (basilar length less than 45, length of m1 less than 6.3, width of M1 less than 4.8, outside length of P4 less than 5.7).

Description.—Size.—Male: Average and extreme measurements of topotypes (as recorded by collectors on labels, and so uniform as to show them not to be accurate to within more than 5 mm.) are as follows: Total length, 434 (410-460); length of tail, 164 (150-180); length of hind foot, 50 (no variation in collectors' measurements). Tail averages 61 per cent as long as head and body. Length of hind foot more than basal length. Corresponding measurements of no. 22191, a young male from Mérida, measured by Osgood or Conover, are 439, 165, 54. The adult male no. 18703, from Páramo de Tama (eastern boundary of Venezuela) has the following measurements written on the label by Osgood: 404, 150, 47.

Female: Average and extreme measurements of topotypes (as recorded by collectors on labels and so uniform as to show them not to be accurate to within more than 5 mm.) are as follows: Total length, 347 (320-370); length of tail, 128 (120-130); length of hind foot, 40 (no variation in collectors' measurements). Tail averages 57 per cent as long as head and body. Length of hind foot more than basal length. Two females, adult no. 11034 and young no. 11033 from Cincinnati, Santa Marta, Colombia, measured by M. A. Carriker, Jr., measure, respectively, as follows: 371, 330; 140, 140; 38, 36. No. 14463, adult, from Río Zapata, Colombia, measured (by J. H. Batty), 315, 138, 39. No. 32182, adult, from Mira Flores, Cauca, Colombia, measured (by W. B. Richardson), 375, 150, 43.

The average differences in external measurements of the two sexes, at Mérida, are: Total length, 87; length of tail, 36; length of hind foot, 10.

Externals.—Longest facial vibrissae black (few rarely white) and extending beyond ear; carpal vibrissae colored like underparts or upper parts, and not extending beyond apical pad of fifth digit; hairiness of foot soles slightly greater than shown in figure .

Color.—As described in Mustela frenata panamensis except that: Posterior fourth of each upper lip rarely, and small spot in front of ear usually, white; black of head proper not extending back of ears and grading insensibly into color of upper parts; anterior half of upper parts of adults "frosted" with numerous white hairs (tick bites?), upper parts near (n) Bay or tone 2 of Reddish Black (pl. 344, Oberthür and Dauthenay) or tone 3 in freshest, unfaded pelage. Least width of color of underparts (in ten males from Mérida) 20 (17-23) per cent of greatest width of color of upper parts. Black tip of tail, in same series, 60 to 75 mm. long, thus longer than hind foot and 41 (40-44) per cent as long as tail-vertebrae.

Skull and teeth.—Male (based on type specimen and seven topotypes, five adults and three subadults): See measurements and plates -; weight, 4.1 (3.8-4.3) grams; basilar length, 43.6 (42.3-44.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth greater than length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum greater than length of tympanic bulla; least width of palate greater than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or less than (approximately equal to) alveolar length of lower molar and premolar tooth-row and shorter than rostrum; anterior margin of masseteric fossa posterior to m2 and confined to posterior third (34 per cent average, 32 minimum, 37 maximum) of mandible.

Female (based on four adult topotypes): See measurements and plates -; weight (no. 143665), 2.3 grams; basilar length 37.2 (36.3-38.2); zygomatic breadth more or less than distance between condylar foramen and M1 or than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth (sinuses badly infested with parasites) more than length of upper premolars or width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than length of P4; tympanic bulla as far posterior to foramen ovale as combined width of 4 to 5 upper incisors; height of tympanic bulla less than (one half to three fourths of) distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row.

The skull of the female is 44 per cent lighter than that of the average male.

Comparisons of the skull with those of M. f. panamensis and affinis have been made in the accounts of those subspecies. As compared with the skull of the male of M. f. aureoventris, that of meridana averages smaller in every measurement taken. Indeed, none of the skulls of meridana equals that of aureoventris in basilar length, length of tooth-rows, length of tympanic bulla, depth of skull at anterior margin of basioccipital or at posterior margin of upper molars, or measurements of teeth. Relative to the basilar length, most of the measurements are greater in meridana. Exceptions are the relative length of the tooth-rows, and the two measurements of depth of the skull which average less.

Remarks.—In 1914 when Hollister named this weasel he compared it with M. f. affinis and most of the differential characters which he ascribed to meridana were merely "more than" or "less than" in affinis. In affinis, Hollister included specimens from Chiriquí, Panamá, and the coast of Venezuela. The specimens from these three places were referred by Allen (1916:101) to Mustela affinis costaricensis, and he restricted (op. cit.:100) the type locality of Mustela affinis affinis to Bogotá, Colombia, and synonymized Mustela meridana with M. a. affinis. Hollister probably would not have named meridana had he had specimens from Bogotá for comparison and had he regarded them as topotypes of affinis for the difference is slight. Nevertheless, within the large geographic range of M. f. meridana there is some geographic variation. There is more of such variation in the color of the pelage than in shape and size of the skull. The specimen from San Julián is darker than the average and in this respect approaches true panamensis. San Julián is situated at a relatively low elevation on the coast of Venezuela.

M. f. meridana so closely resembles M. f. affinis that the writer has no quarrel with anyone who would synonymize meridana. However, as represented by topotypes, the two races unquestionably are, on the average, different, and specimens from the southeastern part of the range of affinis probably are individually distinguishable from topotypes of meridana.

Variation in the skulls of the series from Mérida is relatively small. This applies to both males and females. The external measurements recorded by native collectors are not accurate to within more than five millimeters but, considering this, variation in external measurements also seems to be slight. The difference in size of the two sexes appears to be uniformly greater than in weasels from Central America. The twenty-six topotypes show that the color and color pattern are relatively uniform. All are of nearly the same tone except juveniles or young which are, as in the case of panamensis, much brighter colored on the underparts. Also, the young have darker-colored upper parts. The adults, without exception, have numerous white hairs scattered over the back of the head, neck and between the shoulders. I have no trustworthy evidence to support the suggestion that these white hairs are the results of tick bites or that they are caused by other parasites which damage the hair follicles. The white facial markings vary relatively little in the 45 specimens carefully examined in this regard. Also, the variation in color pattern of the two sides of the head is small. Indeed, within rather narrow limits, the color of the two sides of the head is the same in every specimen except two. In these two the white spots anterior to the ears are confluent with the color of the underparts. Only one specimen, no. 21342, has a white spot between the eyes and this spot is small. Ten of the twenty-six specimens have a definite white spot or band in front of each ear. Two specimens have such a spot on one side only. The dark spots at the angles of the mouth are present on two sides in three specimens and on one side only in three others. The mentioned spots are, then, present nine out of a possible fifty-two times. When the spots are absent, dark color usually is present in the required area but is confluent with the color of the upper parts.

A young male from San Julián, Robinson and Lyon (1901:147) state ". . . was shot . . . as it ran over some bowlders in a ravine. Its eyes shone with the same greenish light as do the eyes of our common weasel, and it emitted the same strong odor." No. 14463, Am. Mus. Nat. Hist., from Río Zapata, Colombia, according to data on the label, was "taken in timber belt in valley in balk hills" and the native name is Cosonebi. Two specimens taken on the Páramo de Tama, head of Tachira River, Venezuela and Colombia are commented on by Osgood (1912:61) as follows: "One . . . was caught in a steel trap baited with birds and set by the side of a rushing mountain stream. . . . The other was shot in midday as it came prowling about our 'house' in the clearing. . . ."

Of the thirty-three skulls before me, twelve have the frontal sinuses malformed by parasites. These twelve include most of the adults for few of the subadults and fewer of the young show pathologic conditions in the frontal region.

Note on localities.—Several of the localities in Colombia mentioned in "Specimens examined" are described and located by Chapman (1917:640-656, pl. 41) in his "Distribution of Bird-life in Colombia." Place names for Colombia on labels, not found on any map, or duplicated names of which I can not certainly select one, are Río Barrotow, Río Oscuro, Río Zapata, Río Japata, Guasca and El Baldro. Sonson may or may not be the town of that name situated some eighty miles northwest of Bogotá and on the east flank of the Central Andes west of the Magdalena River on the drainage of the Cauca River. In Venezuela most of the specimens from Mérida are labeled 1630 meters, Montes de Mérida. San Julián is some seven miles east of La Guaira (see Robinson and Lyon, 1901:136). San Esteban is located a little way back from the coast between Puerto Cabello and Valencia. Páramo de Tama is on the Venezuelan-Colombian border near the source of the Tachira River (see Osgood, 1912:35). Mt. Duida is shown as at 3° 30´ N and 65° 40´ W by Chapman (1931:13) and Mt. Auyán-tepui as near 5° 15´ N and 62° 50´ W by Chapman (1937:760).

Specimens examined.—Total number, 78, arranged by localities from north to south and unless otherwise indicated in the British Museum of Natural History.

Venezuela: San Julián, 1; Carácas, 2; Galipare, Cerro del Avila, 6500 feet, 1; San Esteban, 1; Mérida, 45 (10, 14, 10, 2, 2, 1); Páramo de Tama, 1.

Colombia: Páramo de Tama, 1; Cincinnati, 3; Valdiva, 3800 ft., 1; Medellín, 2; 7200 ft., Barro Blanco, 1; Santa Elena, 9000 ft., 1; Santa Elena, 1; Sonson, 2 (1, 1); Mt. Auyan-tepuy, 1; Pueblo Rico, 5200 ft., 1; Mira Flores, 1; Jerico, near Cauca River, 1; Tornel, 20 mi. NE Quitichao, 1; Mt. Duida, 1; El Tambo, Cauca, 1; El Baldro, 1; Río Japata, 2; Río Zapata, 4500 ft., 1; Río Oscuro, 3300 ft., 1; Río Barrotow, 3300 ft., 1; Guasca, 1; no locality more definite than Colombia, 1.

Mustela frenata affinis Gray

Long-tailed Weasel

Plate

Mustela affinis Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):375, 1874.

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Putorius affinis, Merriam, N. Amer. Fauna, 11:31, June 30, 1896.

Mustela affinis affinis, Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916; Allen, Bull. Amer. Mus. Nat. Hist., 35:220, May 31, 1916.

Mustela frenata affinis, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.

Type.—Male, adult, skull with skin; no. 54.1.11.3 (skull originally numbered 195d, later 54.6.3.4), Brit. Mus. Nat. Hist.; Colombia [given as new Granada in original description]; purchased from Mr. S. Stevens. Type locality restricted by Allen (1916:99) to Bogotá, Colombia.

The skin is in a good state of preservation and has been made over into a conventional study specimen from a mount on exhibition. Exposure to light when mounted probably accounts for the faded color. The skull (plate ) lacks the middle 9 mm. of the right zygomatic arch, occiput, basioccipital and posterior two-thirds of the left tympanic bulla. The teeth all are present and entire.

Range.—Four thousand six hundred feet (Quetame) to 9154 feet (El Carmen), Tropical to Temperate life-zones of eastern Andes of Colombia. See figure on page .

Characters for ready recognition.—Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, by presence of p2 and by two roots rather than one root on P2; from M. frenata meridana, in case of males, by, on average, greater breadth and length of skull and lesser actual and relative size (see measurements) of facial part of skull; from M. f. aureoventris by lighter-colored upper parts (tone 2 rather than tone 4, pl. 344, Reddish Black of Oberthür and Dauthenay); from M. f. macrura by darker color (Reddish Black, tone 2, pl. 344, Ober. and Dauth., rather than Chocolate, tone 3, pl. 343, Ober. and Dauth.).

Description.—Size.—Male: Measurements in life, estimated from dried skins, are: Total length, 455; length of tail, 175; length of hind foot, 52. Proportions of parts supposedly as described in Mustela frenata meridana.

Female: Estimates from two dried skins: Total length, 365; length of tail, 135; length of hind foot, 43. Proportions of parts supposedly as described in Mustela frenata meridana.

The estimated differences in external measurements of the two sexes are: Total length, 90; length of tail, 40; length of hind foot, 9.

Externals.—As described in Mustela frenata meridana.

Color.—As described in Mustela frenata panamensis except that: posterior fourth of each upper lip and spot in front of each ear white in approximately half of the specimens; black of head proper not extending back of ears and grading insensibly into color of upper parts; upper parts near (n) Bay, or tone 2 of Reddish Black (pl. 344, Oberthür and Dauthenay). Least width of color of underparts (in five males from vicinity of Bogotá) 24 (15-29) per cent of greatest width of color of upper parts. Black tip of tail, in same series, 60 to 75 mm. long, thus longer than hind foot and averaging 38 per cent as long as tail-vertebrae.

Skull and teeth.—Male (based on three adults and two subadult topotypes): See measurements and plate . As described in Mustela frenata meridana except that: Weight, 4.5 grams (estimated); basilar length 45.8±; interorbital breadth not greater than distance between foramen opticum and anterior margin of tympanic bulla (type as in meridana where interorbital breadth is more than distance between foramen opticum and anterior margin of tympanic bulla); least width of palate not less than length of P4; masseteric fossa confined to posterior two-fifths (38 to 40 per cent; average 39 per cent) of mandible and not extended anteriorly to middle of m2.

Female: No adults examined.

As compared with M. f. meridana the skull of the male is larger, to the average amount of 2.2 mm. in basilar length and 1.2 mm. in zygomatic breadth of adults; length of tooth-rows and mastoid breadth average greater but relatively less; breadth of rostrum, interorbital breadth and orbitonasal length average actually and relatively less. Thus the skull of affinis is longer and broader, but the facial region is actually, as well as relatively, smaller. As compared with the skull of the male of M. f. aureoventris, that of M. f. affinis is about the same in basilar length. However, in no specimen of affinis are the measurements of length of tooth-rows or breadth of rostrum, actually, or relatively, as great as in aureoventris. The same is true of all measurements taken of M1, P4 and m1. The specimens from the vicinity of Quito and north of there, although referred to macrura, are nearly as dark as typical affinis, approach affinis in cranial characters, and indicate intergradation of affinis with macrura.

Remarks.—Mustela affinis was named by John Edward Gray in 1874 (p. 375) on the basis of a specimen from New Granada. Although usually synonymized with Mustela brasiliensis by later authors until 1896 when Merriam (1896:31) applied the name to weasels from Costa Rica, nearly all the South American and several of the Central American weasels have, at one time or another, had Gray's name, affinis, applied to them. Gray, in 1865 (p. 115) when giving measurements of Mustela aureoventris, probably mentioned the specimen, that later became the holotype. In 1916 (p. 98) Allen restricted the type locality to Bogotá, Colombia. Allen's action was a necessary procedure in clearing up the systematics of South American weasels and was based on good grounds. As set forth by Allen (loc. cit.), and more in detail by Chapman (1917:642), Bogotá has long been the shipping point for Colombian vertebrate specimens, many of which were obtained in the mountains to the east. Allen (1916A:220) quotes Thomas as saying that the type specimen was purchased from Stevens at about the same time that a number of Colombian birds were purchased from the same dealer. Also, specimens from Bogotá agree with Gray's description of the type specimen.

Mustela frenata affinis, as here defined, constitutes one of the several slight geographic variants met with, on the sides of, and between, the three north and south mountain chains of Colombia. The others are lumped under the name Mustela frenata meridana. M. f. affinis, in common with specimens from the northern part of the range of macrura has large teeth. Weasels of all of the region from Quito to Bogotá have large teeth. To the north there is the smaller-toothed meridana and to the south the smaller-toothed macrura grading into the still smaller-toothed agilis, and boliviensis.

Two skins, without corresponding skulls, from Caqueta are lighter colored than any others of affinis; possibly the skins are faded by exposure to light. Since they probably come from an elevation of less than 1000 feet in the Amazonian region, they may pertain to another subspecies.

Complete, unbroken, skulls of affinis are needed to ascertain the degree to which affinis and meridana differ in cranial features. The several specimens from the immediate region of Bogotá show well the color and the color pattern but lack collectors' measurements.

None of the ten skulls examined shows malformation of the frontal region due to infestation of the frontal sinuses by parasites. Possibly three of the four adults were infested, although not severely.

Specimens examined.—Total number, 27, arranged by localities from north to south and unless otherwise indicated in the United States National Museum.

Colombia: El Carmen, 1; W. Cundinamarca, 1; Muzzo [= Muzo?], 1; Bogotá, 1; Castillo, near Bogotá, 1; Fambrias, near Bogotá, 1; Bogotá district, 1; Choachí, 9 (1, 2, 1); Páramo de Choachí, 2 (1, 1); Laguna del Verjón (= City of Bogotá), 1; Quetame, 2; Fusagasuga, 1; Caqueta, 2; no locality more definite than Colombia, 3 (1).

Mustela frenata aureoventris Gray

Long-tailed Weasel

Plates , and

Mustela aureoventris Gray, Proc. Zoöl. Soc. London, 1864:55, pl. 8, 1864; Gray, Proc. Zoöl. Soc. London, 1865:115, 1865.

Putorius (Gale) brasiliensis var. aequatorialis Coues, Fur-bearing animals, p. 142, 1877, part? ("merely as a substitute for Gray's [supposedly] preoccupied name," that is, aureoventris).

Mustela affinis costaricensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916 (part).

Mustela macrura, Lönnberg, Arkiv för Zool., 14 (no. 4):11, 1921 (part ?).

Mustela frenata aureoventris, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.

Type.—Probably female, juvenile, skull with skin, no. 64.6.6.3 (formerly 1432a), British Mus. Nat. Hist.; probably Subtropical Life-zone of western Ecuador (locality given as Quito, probably because received from that place).

The skin, once exhibited as a mount, has lost some hair from the back and other parts of the body and is not suitable for remaking into a conventional study specimen. The skull lacks the occiput, basioccipital, premaxillae, upper incisors, two of the lower incisors, all of the canines, premolars 2/2 on both sides, right P3, left p3, and has the left jugal mesially defective. The premolars present are not all fully emerged.

Range.—Pacific coastal regions of Ecuador and Colombia; Subtropical and Tropical life-zones. See figure on page .

Characters for ready recognition.—Differs from Mustela africana stolzmanni by absence of median, longitudinal, abdominal stripe of same color as upper parts, by presence of p2 and by two rather than one root on P2; from Mustela frenata macrura by Reddish Black, tone 4, plate 344 rather than Chocolate, tone 3, pl. 343 (of Oberthür and Dauthenay), or slightly darker color of upper parts; from M. f. affinis and M. f. meridana by darker color (tone 4 rather than tone 2, Reddish Black of Ober. and Dauth.) of upper parts and larger size of teeth (M1 with length more than 2.4 and breadth more than 4.7; P4 with outside length more than 5.9; length of m1 more than 6.2).

Description.—Unless otherwise stated, information concerning this subspecies is derived from the one referred specimen available, a young male, no. 34677, Amer. Mus. Nat. Hist.

Size.—Male: Total length, 470; length of tail, 160; length of hind foot, 50. Tail 51 per cent as long as head and body.

Female: Not known.

Externals.—Longest facial vibrissae black and reaching beyond ear. Carpal vibrissae reaching to or beyond apical pad of fifth digit; hairiness of foot soles slightly less than shown in figure 20.

Color.—Sides and top of head and neck posteriorly to shoulders black; white facial markings represented by only five white hairs anterior to right ear, one anterior to left ear and three far back on forehead; dark areas at angles of mouth confluent with color of upper parts; tip of tail black; remainder of upper parts near (n) Bay or Reddish Black, tone 4 of Oberthür and Dauthenay, pl. 344; chin whitish; remainder of underparts Warm Buff, deep orange in juvenile, type specimen, according to Gray (1864, pl. 8); color of underparts extending distally on posterior sides of forelegs to wrists but not reaching foot soles and on hind legs to or slightly below knees. Least width of color of underparts equal to 15 per cent of greatest width of color of upper parts. Black tip of tail equal to 27 per cent of length of tail-vertebrae.

In color, no. 34677 is, to me as it was to Allen (1916:101), indistinguishable from the darkest specimens (nos. 178970 and 10112) of M. f. panamensis. Therefore, M. f. aureoventris is one of the two darkest subspecies of weasels.

Skull and teeth.—Male: See measurements and plates -; weight, 4.3 grams; basilar length, 45.8; zygomatic breadth approximately equal to distance between condylar foramen and M1 and to distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more than length of upper premolars and greater than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth greater than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less (at least in young specimen) than length of tympanic bulla; least width of palate seldom if ever greater than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of three (including I3) upper incisors; height of tympanic bulla not greater than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and shorter than orbitonasal length; anterior margin of masseteric fossa below anterior half of m2.

Skulls of males of M. f. aureoventris, and Mustela frenata macrura from the vicinity of Quito so closely resemble one another as not to be distinguished with the material now available, although the teeth of aureoventris are larger. Comparisons of the skulls of males with those of M. f. meridana and affinis, which are readily distinguishable from those of aureoventris, have been made in the accounts of those subspecies.

Female: Skull of adult unknown.

Remarks.—This subspecies of the Tropical Life-zone, or at least the Subtropical Life-zone, of Ecuador, in certain cranial characters resembles Mustela frenata macrura of the Temperate Life-zone. The two differ markedly in color. Nevertheless, a large number of the specimens collected in Ecuador are intermediate in color as well as in zonal distribution.

The type specimen is young or a juvenile. The measurements of no. 34677 from Gualea indicate an animal similar in size to M. f. affinis. Gray (1864:55) states that the type specimen measures "Length of body and head 6 inches, of tail 4-1/2 inches." The plate (pl. 8) accompanying Gray's original description (loc. cit.) is marked one-half natural size and represents the animal as having a head and body length of eight and one-half inches. One year later Gray (1865:115) gives the measurements of this species as "Length of body and head 12, tail 8 inches." Since he had at this time another specimen, larger than the type specimen (which specimen later, probably, became the type of Mustela affinis Gray), the larger measurements probably were taken from it.

Geographically, and as regards cranial characters, Mustela frenata aureoventris is most closely related to M. f. affinis and to the northern section of M. f. macrura, but in color to M. f. panamensis. M. f. aureoventris and M. f. panamensis are the two darkest-colored subspecies and each occurs in a region of extremely heavy rainfall. There is a skin only, no. 32620, Amer. Mus. Nat. Hist., from Munchique, obtained on June 1, 1911, which is appreciably darker than specimens of M. f. affinis in corresponding pelage and is intermediate between M. f. affinis and M. f. aureoventris in color as it is geographically. The specimen measures 495, 202, 52.