measurements and plates 19-21; weight, 3.5 (2.8-4.7) grams;
basilar length, 43.7 (40.0-47.7); zygomatic breadth more or less
than distance between condylar foramen and M1 or than between
anterior palatine foramen and anterior margin of tympanic bulla;
mastoid breadth more or less than postpalatal length; postorbital
breadth less than length of upper premolars and greater than width
of basioccipital measured from medial margin of one foramen
lacerum posterior to its opposite; interorbital breadth more or
less than distance between foramen opticum and anterior margin of
tympanic bullae; breadth of rostrum less (except in no. 82180)
than length of tympanic bulla; least width of palate more (except
in no. 81954) than length of P4; anterior margin of tympanic bulla
as far posterior to foramen ovale as width of 2 to 5 upper
incisors; height of tympanic bulla more or less than distance from
its anterior margin to foramen ovale; length of tympanic bulla
more (except in two instances) than length of lower molar and
premolar tooth-row and shorter (except in two instances) than
rostrum; anterior margin of masseteric fossa below m2.
Female (based on 11 ad. topotypes): See measurements and plates
34-36; weight, 2.0 (1.8-2.2) grams; basilar length, 37.6
(37.0-38.9); zygomatic breadth less (except in no. 70945) than
distance between condylar foramen and M1 or than between anterior
palatine foramen and anterior margin of tympanic bulla;
postorbital breadth less than length of upper premolars and more
than width of basioccipital measured from medial margin of one
foramen lacerum posterior to its opposite; least width of palate
less (except in one specimen) than greatest length of P4; tympanic
bulla as far posterior to foramen ovale as width of 3-1/2 to 5-1/2
upper incisors; height of tympanic bulla more or less than
distance from its anterior margin to foramen ovale; length of
tympanic bulla more than length of lower molar and premolar
tooth-row and longer or shorter than rostrum.
Compared with _M. f. nevadensis_, the skull of the male of
_washingtoni_ averages more slender, as shown by the mastoid and
zygomatic breadths and has the preorbital part longer, on the average,
as shown by the greater ratio (to the basilar length) of the length of
the tooth-rows and orbitonasal length. Also, on the average, the
postorbital constriction is longer than in _nevadensis_ and the
tympanic bullae are smaller. In females, the skull is lighter, the
tooth-rows are shorter, the tympanic bullae are smaller, and the
preorbital part of the skull is shorter and narrower as shown by the
orbitonasal length and interorbital breadth. Except that the tympanic
bullae are actually, although not relatively, smaller in males of
_effera_, it differs from _washingtoni_ in the same way as does
_nevadensis_ as regards relative proportions, but, of course, the
actual difference in size is greater since _effera_ is smaller than
_nevadensis_. Comparison of the skull with that of _oregonensis_ is
made in the account of that subspecies.
_Remarks._--_M. f. washingtoni_ was described and named in 1896 by
Merriam as a distinct species. Subsequently, specimens which here are
regarded as intergrades between _altifrontalis_ and _nevadensis_, were
classified as _washingtoni_.
The external measurements given for the specimens from Mount Adams are
those recorded on the labels in inches and fractions thereof. Instead
of total length there sometimes is written "tip to tip." In the series
of 19 winter-taken topotypes the hairs project beyond the end of the
caudal vertebrae for an average distance of 28 (19-40) millimeters. If
the hairs on the end of the tail were included in the measurements, 28
millimeters should be subtracted from the averages. Probably the
measurements should stand as given, since an adult male topotype, no.
226758, U. S. Nat. Mus., taken subsequently by Walter P. Taylor
measures 405; 152; 51.
_Mustela frenata washingtoni_ is not a strongly marked geographic race.
In many features it is intermediate between _M. f. altifrontalis_ and
_M. f. nevadensis_. This is especially true of coloration. In the
series from Mount Adams and that from Mount Rainier, some individuals
have the light color of the underparts extended down the hind legs over
the feet and over the proximal face of the ventral third of the tail as
in _nevadensis_, whereas others from the same place have the light
color of the underparts absent from the tail and extending no farther
down the hind limbs than the knees. The light color of the underparts
in the series of topotypes is so restricted that the transverse extent
at the narrowest place amounts to only 24 (10-37) per cent of the
greatest width of the color of the upper parts. This narrowness of the
color of the underparts has been likened by Merriam (1896:18) to the
condition in _Mustela frenata noveboracensis_. So it is, but it is
similar to the condition found also in the geographically adjoining _M.
f. altifrontalis_.
Of the 37 skulls of subadults and a few adults, 11 had the frontal
sinuses malformed as a result of infestation by parasites.
_Specimens examined._--Total number, 56, arranged within each
state by localities from north to south. Unless otherwise
indicated specimens are in the collection of the United States
National Museum.
=Oregon.= Mt. Jefferson, Permilia Lake, 1.
=Washington.= _Pierce County_: 5500 ft., Spray Park, Mt. Rainier,
1; Spray Park, 1[74]; 5935 ft., Glacier Basin, Mt. Rainier, 5
(1[10]); 5051 to 5100 ft., Owyhigh Lakes, Mt. Rainier, 7 (1[10]),
Tahoma Creek, 1[72]; Nisqually entrance, 1[72]; Longmire, 1[72];
Mt. Rainier Nat'l Park, 2[72]. _Klickitat County_: Trout Lake, S
Base Mt. Adams, 35; 3500 ft., Gotchen Creek, Mt. Adams, 1.
=Mustela frenata saturata= (Merriam)
Long-tailed Weasel
Plates 19, 20, 21 and 30
_Putorius saturatus_ Merriam, N. Amer. Fauna, 11:21, June 30, 1896.
_Mustela saturata_, Miller, U. S. Nat. Mus. Bull., 79:98, December
31, 1912.
_Mustela arizonensis saturata_, Grinnell, Univ. California Publ.
Zoöl., 40:102, September 26, 1933.
_Mustela frenata saturata_, Hall, Carnegie Instit. Washington Publ.
473:106, November 20, 1936.
_Type._--Male, adult, skull and skin; no. 65930, U. S. Nat. Mus.,
Biol. Surv. Coll.; Siskiyou, Jackson County, Oregon; June 6, 1894;
obtained by C. P. Streator; original no. 3905.
The skull (plates 19-21, 30) lacks the middle part of each
zygomatic arch. The teeth all are present although much worn,
probably from gnawing at the trap which captured the animal. The
skin, in fresh summer pelage, is fairly well made.
_Range._--Transition and Boreal life-zones of Siskiyou and Trinity
mountains in southern Oregon and northwestern California. See
figures 29 and 30 on pages 221 and 314.
_Characters for ready recognition._--Differs from _M. f.
nevadensis_ in lacking light color of underparts on tail and ankle
and in greater average breadth across mastoid processes of skull
(see measurements); from _M. f. oregonensis_ in lacking white
nasofrontal spot, in having color of underparts interrupted at
ankle; from _M. f. munda_ in lacking white nasofrontal spot, in
smaller and relatively deeper skull of males and smaller skull of
the female.
_Description._--_Size._--Male: Four adult males (the type, 1 from
Mt. Ashland and 2 from Jackson Lake) yield average and extreme
measurements as follows: Total length, 414 (402-437); length of
tail, 150 (136-160); length of hind foot, 46 (43-50). Tail
averages 57 (49-62) per cent as long as head and body. Length of
hind foot more or less than basal length.
Female: One young from the summit of the Trinity Mountains east of
Hoopa and one nontypical adult from 5500 feet elevation on South
Fork Mountain, Humboldt County, measure respectively as follows:
Total length, 330, 325; length of tail, 115, 123; length of hind
foot, 37, 37. Tail is 53 and 61 per cent as long as head and body.
Length of hind foot less than basal length.
Average differences in external measurements between the two
sexes, indicated by the unsatisfactory material available, are:
Total length, 86; length of tail, 31; length of hind foot, 9.
_Externals._--Longest facial vibrissae black or dark brown and
extending beyond ear; carpal vibrissae same color as underparts
and extending as far as apical pad of fifth digit; hairiness of
foot-soles, in summer pelage, as shown in figure 19.
_Color._--Upper parts, in summer, Brussels Brown to near (_n_)
Brussels Brown or lighter than tone 3 of Raw Umber of Oberthür and
Dauthenay, pl. 301, usually darkest on nose and forehead. Chin
white. Remainder of underparts Buff-Yellow to Warm Buff. Tip of
tail black. Winter pelage unknown. Color of underparts extends
distally on posterior sides of forelegs over toes onto antipalmar
faces of feet and sometimes wrists, on medial sides of hind legs
only to ankles, but toes sometimes with isolated white markings.
Least width of color of underparts in the type and 2 adults from
Jackson Lake averaging 35 (30-40) per cent of greatest width of
color of upper parts. Black tip of tail averaging 54 (53-55) mm.
long; thus longer than hind foot and averaging 37 per cent of
length of tail-vertebrae.
_Skull and teeth._--Male (based on 4 adults: Type, Mt. Ashland, 1;
Jackson Lake, 2): See measurements and plates 19-21, 30. As
described in _Mustela frenata nevadensis_ except that: Weight, 3.8
(3.5-4.3) grams; basilar length, 44.4 (42.6-45.8); zygomatic
breadth more or less than distance between condylar foramen and M1
or than distance between anterior palatine foramen and anterior
margin of tympanic bulla; mastoid breadth more than postpalatal
length; least width of palate less than medial length of P4
(except in one specimen).
Female (based on one adult possibly not typical, from 5500 ft.,
South Fork Mt.): See measurements. As described in _Mustela
frenata nevadensis_ except that: Weight, 2.2 grams; basilar
length, 38.1; zygomatic breadth less than distance between
condylar foramen and M1 and less than distance between anterior
palatine foramen and anterior margin of tympanic bulla;
postorbital breadth more than width of basioccipital measured from
medial margin of one foramen lacerum posterior to its opposite.
The skull of the male of _saturata_, relative to the basilar length, is
broader across the mastoids and narrower across the rostrum and
interorbital region than that of _nevadensis_. Skull not known
certainly to differ from that of _oregonensis_. Compared with the skull
of _munda_, that of the male of _saturata_ is smaller in every part
measured except depth of tympanic bullae which averages 3.6
millimeters, rather than 3.5 as in _munda_. Also, the skull of
_saturata_ has a less-marked postorbital constriction, is less heavily
ridged, less angular, does not have the impressions of the temporal
muscles carried so far forward on the frontal bones and is relatively
much narrower across the zygomatic arches.
_Remarks._--In 1896, Merriam named _M. f. saturata_ as a distinct
species on the basis of one specimen, taken by Clark P. Streator at
Siskiyou, Oregon, and a second specimen taken the year previously by
Allan C. Brooks at Chilliwack, British Columbia. On the basis of these
two specimens, Merriam (1896:22) ascribed to the race a range ". . . on
the Cascade and Siskiyou mountains of Oregon and Washington, reaching a
short distance into British Columbia." Since that time, this name,
_saturata_, has been employed for the dark-colored weasels, of the
coastal region of Oregon, Washington, and extreme southwestern British
Columbia, which here are arranged under the name _M. f. altifrontalis_.
_M. f. saturata_ proves to be restricted to the humid mountainous
region inland from the coast in northern California and in the Siskiyou
Mountains of southern Oregon. Its range is separated by that of _M. f.
oregonensis_ from the range of the darker-colored, deeper-skulled, _M.
f. altifrontalis_ of the humid costal region proper.
On May 5, 1933, Mr. Clark P. Streator, informed the writer that he
remembered taking the type specimen of _Mustela frenata saturata_
(Merriam) in the town of Siskiyou, Oregon. The exact place, he said,
was reached, at the time of his work there, by going one or two blocks
east of the depot, then through a garden into the thick woods where
there were springs and numerous burrows of the rodent, _Aplodontia_.
Two other weasels labeled as taken at Siskiyou, on September 28 and
29, 1893, by Mr. Streator, are much lighter colored than the type of
_saturata_ and have the color of the underparts extended distally on
the hind legs to the tips of the toes and in other features of
coloration are more like _nevadensis_, the subspecies to which they are
referred, than _saturata_. Probably these did not come from exactly the
same place that the type specimen of _saturata_ did. Although Mr.
Streator does not remember the taking of these particular specimens in
1893, he does remember that on this visit to Siskiyou, he walked
southward through the railroad tunnel and collected on the opposite
side of the ridge from Siskiyou. Here on more southern exposures, the
country was markedly different than in the thick forest at Siskiyou.
Probably these two specimens taken in 1893, and referred to
_nevadensis_, came from a little way south of Siskiyou and from a
different habitat and life-zone than the type specimen of _M. f.
saturata_.
Of the 6 specimens examined, only one, the type, shows malformation of
the frontal sinuses such as result from infestation by parasites.
_Specimens examined._--Total number, 6, as follows:
=California.= _Siskiyou County_: Jackson Lake, 5900 ft., 2, Mus.
Vert. Zoöl. _Humboldt County_: South Fork Mt., 5500 ft., 1, Mus.
Vert. Zoöl. _County_ in question, Trinity Mts., summit east of
Hoopa, 5800 ft., 1, U. S. Nat. Mus.
=Oregon.= _Jackson County_: Mt. Ashland, 1, Univ. Oreg.; Siskiyou,
1, U. S. Nat. Mus.
=Mustela frenata altifrontalis= Hall
Long-tailed Weasel
Plates 1, 19, 20, 21, 34, 35 and 36
_Mustela frenata altifrontalis_ Hall, Carnegie Instit. Washington
Publ. 473:94, November 20, 1936.
_Putorius (Gale) brasiliensis frenatus_, Coues, Fur-bearing
animals, p. 142, 1877 (part).
_Putorius saturatus_ Merriam, N. Amer. Fauna, 11:21, June 30, 1896
(part).
_Mustela saturata_, Miller, U. S. Nat. Mus. Bull., 79:98, December
31, 1912.
_Type._--Male, adult, skull and skin; no. 42093, Mus. Vert. Zoöl.;
Tillamook, Tillamook County, Oregon; July 10, 1928; obtained by
Alex Walker; original no. 717.
The skull is complete and unbroken. P3 on the left side is
missing; otherwise the teeth all are present and entire. The skin
is well made and the enlarged scrotal pouch shows the collector's
sexing of the specimen to have been correct.
_Range._--Altitudinally from sea level up to at least 4800 feet
(Mount Baker) in the Transition Life-zone of the humid, coastal
region of Oregon, Washington and extreme southwestern British
Columbia. See figure 29 on page 221.
_Characters for ready recognition._--Differs from _M. f.
nevadensis_ in tone 4 of Brownish Drab, pl. 302, rather than tones
1-3, of Raw Umber, pl. 301, of Oberthür and Dauthenay of upper
parts, in near (14 _a´_ to 16 _c´_) Ochraceous-Buff rather than
Buff-Yellow to Straw Yellow of underparts, in that least width of
color of underparts amounts to less than 37 per cent of greatest
width of color of upper parts, in absence of color of underparts
on ventral side of tail and on hind leg distal to knee, and in
greater depth of skull through frontal region; from _M. f.
washingtoni_ in darker color of upper parts and underparts, latter
near (14 _a_´ to 16 _c_´) Ochraceous-Buff rather than ranging from
Buff-Yellow to Naples Yellow, in deeper skull in both sexes (see
measurements), in males a shorter preorbital region, broader skull
with longer bullae and in females a larger skull with interorbital
breadth averaging more than 24 per cent of basilar length; from
_M. f. oregonensis_ in frontonasal white patch absent, color above
darker (tone 4 of Brownish Drab, pl. 302, rather than tone 2 to 3
of Raw Umber, pl. 301 of Oberthür and Dauthenay), light-colored
underparts narrower and not extended distally beyond knee, in
females tooth-row shorter, amounting to less than 38 per cent of
basilar length.
_Description._--_Size._--Male: Eight adult topotypes yield average
and extreme measurements as follows: Total length, 426 (392-445);
length of tail, 160 (148-170); length of hind foot, 47 (42-53).
Tail averages 60 per cent as long as head and body. Length of hind
foot averages more than basal length.
Female: Five adults from Tillamook and Blaine, Oregon, yield
average and extreme measurements as follows: Total length, 347
(320-370); length of tail, 125 (114-131); length of hind foot, 38
(35-44). Tail averages 56 per cent as long as head and body.
Length of hind foot less than basal length.
The average differences in the external measurements are: Total
length, 79; length of tail, 35; length of hind foot, 9.
_Externals._--Longest facial vibrissae black, brown or white
(often all three colors in same specimen) and extending beyond
ear; carpal vibrissae same color as underparts and extending to or
beyond apical pad of fifth digit; hairiness of foot-soles (in
summer pledge) slightly less than shown in figure 19.
_Color._--Upper parts, in summer, near (_n_) Argus Brown or tone 4
of Brownish Drab of Oberthür and Dauthenay, pl. 302. Dark spot at
each angle of mouth well developed; often fused with color of
upper parts which sometimes covers lower lips. Chin white.
Remainder of underparts near (14 _a´_ to 16 _c´_) Ochraceous-Buff.
In winter, upper parts near (14) Argus Brown with smoked effect
and Warm Buff to Naples Yellow below. Tip of tail at all times
black. Color of underparts extends distally on posterior sides of
forelegs over toes onto antipalmar faces of feet and usually all
of wrists, on medial side of hind legs typically only to knee but
sometimes to ankle. Tips of toes of hind feet almost always marked
with color of underparts. Least width of color of underparts
averaging in a series of 14 males from Blaine, Oregon, 23 (14-36)
per cent of greatest width of color of upper parts. Black tip of
tail in 8 adult males from Blaine, Oregon, averaging 59 (47-70)
mm. long; thus longer than hind foot and averaging 37 per cent of
length of tail-vertebrae.
_Skull and teeth._--Male (based on 9 adults from Blaine, Tillamook
Co., Oregon): See measurements and plates 19-21; weight, 4.4
(3.3-5.3) grams; basilar length, 45.6 (42.4-47.7); zygomatic
breadth more or less (usually more) than distance between condylar
foramen and M1 or than between anterior palatine foramen and
anterior margin of tympanic bulla; mastoid breadth more or less
(usually more) than postpalatal length; postorbital breadth less
(except in some instances of malformations of frontal sinuses
which result from infestation by parasites) than length of upper
premolars and more or less than width of basioccipital measured
from medial margin of one foramen lacerum posterior to its
opposite; interorbital breadth more or less than distance between
foramen opticum and anterior margin of tympanic bulla; breadth of
rostrum less than length of tympanic bulla; least width of palate
more or less than length of P4; anterior margin of tympanic bulla
as far posterior to foramen ovale as width of 3 to 4 (including
I3) upper incisors; height of tympanic bulla more than distance
from its anterior margin to foramen ovale; length of tympanic
bulla more than length of lower molar and premolar tooth-row and
more or less than orbitonasal length; anterior margin of
masseteric fossa directly below m2.
Female (based on 4 adults): See measurements and plates 34-36;
weight, 2.2 (2.2-2.3) grams; basilar length, 38.1 (37.8-39.7);
zygomatic breadth more or less (less in three of four specimens)
than distance between condylar foramen and M1 or than between
anterior palatine foramen and anterior margin of tympanic bulla;
relation of postorbital breadth to other measurements in doubt
because of malformation of frontal sinuses by parasites; least
width of palate not less than greatest length of P4; tympanic
bulla as far posterior to foramen ovale as width of 3-1/2 to 5-1/2
upper incisors; height of tympanic bulla more than distance from
its anterior margin to foramen ovale; length of tympanic bulla
more than length of lower molar and premolar tooth-row and longer
or shorter than rostrum.
Compared with the skull of _M. f. washingtoni_ that of each sex of
_altifrontalis_ averages slightly larger in every measurement taken,
except measurements of teeth which are approximately the same, and is
relatively deeper through the frontal region and through the braincase
as measured at the anterior margin of the basioccipital. Skulls of
females of _altifrontalis_ have a relatively broader interorbital
region. Skulls of males of _altifrontalis_ further differ in having
relatively, as well as actually, longer tympanic bullae, relatively
lesser orbitonasal length and a greater relative breadth across the
mastoids and across zygomata. Compared with _M. f. nevadensis_, the
skull of the male of _altifrontalis_ averages slightly larger and
heavier although the skulls of females are of approximately the same
size and weight. Relative to the basilar length, the skulls of both
sexes are deeper through the braincase and narrower across the
mastoids; the rostrum is broader, especially in males; the tooth-rows
are shorter and the interorbital breadth less, especially in females.
Comparison with the skull of _oregonensis_ is made in the account of
that subspecies.
_Remarks._--Until the present study was begun, animals of this race
have gone under the name _Mustela saturata_ (Merriam). The United
States National Museum has a juvenile taken, in 1858, by Wayne at
Astoria, O. T.; the Samuel N. Rhoads collection contained one specimen
taken in 1891, at Tacoma, Washington; one in the Bangs' collection was
taken at Chilliwack, British Columbia, in 1895, and the Field Museum
has one taken on the Olympic Peninsula in 1898. The best material is
that collected by Alex Walker, at Tillamook, Oregon.
Intergradation with _nevadensis_ is indicated by several specimens. The
coloration of the one adult female, no. 90, Chas. R. Conner Mus., from
Swamp Creek, Washington, has the color of the underparts extended down
the hind legs over the feet, and over the proximal third of the ventral
face of the tail as in _nevadensis_ although the other two specimens
from the same place have the color pattern of _altifrontalis_. Of the
four specimens from British Columbia referred to this subspecies, only
the specimen from Chilliwack is typical as regards color pattern. The
one from Cultus Lake has the color pattern of _nevadensis_ and might be
referred to that race almost as well as to _altifrontalis_. The two
specimens from Lihumption Park are intermediate between the two races
in tone of color. Neither has the color of the underparts extended onto
the tail or continuously over the hind feet as in _nevadensis_ but each
does have the color of the underparts less restricted and of lighter
hue than in _altifrontalis_. Only one of the specimens, no. 7848 Canad.
Nat. Mus., from Lihumption Park is adult and it has a skull which
agrees with that of _altifrontalis_ rather than _nevadensis_.
After writing the above, a good representation of the weasel population
along the eastern side of Puget Sound was made available by friends in
that area. Study of the weasels from there shows that their color is
intermediate between that of _altifrontalis_ and _nevadensis_. On the
whole, they (specimens from Bellingham, for example) resemble one
subspecies about as much as the other. In cranial characters some
specimens, in certain features, approach _nevadensis_ but most
specimens agree with _altifrontalis_ and all are more nearly like
_altifrontalis_ to which race all are referred.
The color of these animals is to me indistinguishable from that of
_washingtoni_. The color of _washingtoni_ is merely intermediate
between that of _nevadensis_ and _altifrontalis_. Nevertheless, the
race _washingtoni_ has cranial characters (long narrow skull) which set
it off from both _altifrontalis_ and _nevadensis_. This shape of skull
is not found in the specimens from along the eastern side of Puget
Sound; these animals have skulls like that of _altifrontalis_ and when
departures from this occur they are in the direction of _nevadensis_
and not _washingtoni_.
The above, then, explains why specimens which are colored like those
of _washingtoni_ are not referred to that race but instead to the race
_altifrontalis_.