features regarded as of diagnostic value for one or the other of the
two races, _longicauda_ and _noveboracensis_, behave differently. For
example, the dark coloration of the upper parts, which is
characteristic of _noveboracensis_, manifests itself far westward
within the range of _spadix_ whereas the wider extent of the
light-colored underparts, which is characteristic of _longicauda_, and
the Olive Ocher, rather than Pale Orange Yellow, color of these
underparts, are seen in varying degree all the way across the range of
_spadix_. Thus, these animals are colored above like _noveboracensis_
and below like _longicauda_, but not _vice versa_. In these animals,
then, the _longicauda_ type of underparts is dominant, in one sense of
the word, over the _noveboracensis_ type of underparts, and the
_noveboracensis_ type of upper parts is dominant over the _longicauda_
type of upper parts. Each of these features is subject to actual
intergradation and does not always behave as a "unit character," that
is to say, one which is either present or absent. However, the
_noveboracensis_ type of upper parts is carried much farther west
before being diluted than is the _noveboracensis_ type of underparts.
Indeed, within the range of _noveboracensis_ itself, the broad extent
of the _longicauda_ type of underparts is manifest. This is, of course,
near the western margin of the range of _noveboracensis_.
The large size of males of _spadix_, as exemplified by specimens from
Elk River (see measurements on p. 421), seems to be retained across the
northern part of the range here assigned to the subspecies. This larger
size than is found in _longicauda_ from Alberta, is shown also by some
specimens from eastern North Dakota which are assigned to _longicauda_.
However, the average of these Dakotan specimens, all characters
considered, is nearer to my concept of _longicauda_.
Inspection of the cranial measurements of _spadix_ shows also that in
addition to its large size it is distinguishable from any one of the
geographically adjoining races by its relatively (to basilar length)
greater, as well as actually greater, mastoidal breadth. This might be
included with size as a unique character distinguishing _spadix_ from
_longicauda_ and _noveboracensis_. However, it is not clear whether or
not this greater mastoidal breadth is more than a function of the large
size.
Excepting the greater mastoidal breadth and generally larger size of
the skull, the cranial features distinguishing males of _spadix_ from
_longicauda_ are features in which _spadix_ shows approach to
_noveboracensis_. This is true, in _spadix_, of the relatively longer
(in comparison with _longicauda_) rostrum, relatively lesser zygomatic
breadth, relatively shallower braincase measured at the anterior end of
the basioccipital, and relatively deeper skull as measured at the
posterior borders of the last upper molars. This same approach to
_noveboracensis_ already has been pointed out with respect to color of
the upper parts and is evident also in the relative shortness of the
tail which averages only 51 per cent of the length of the head and body
rather than 55 per cent as in _longicauda_.
Because the _longicauda_ type of animal previously has been regarded as
specifically distinct from the _noveboracensis_ type of animal, comment
is offered below on selected specimens, referred to _spadix_, which are
regarded as intergrades with _noveboracensis_ or with other subspecies.
No. 8722, Univ. Wisconsin, adult male, in the white winter coat, from
north central Itasca County, Minnesota, obviously has characters of _M.
f. spadix_ or _longicauda_ that occur to the west and _M. f.
noveboracensis_ of the east. Selected outstanding characters of
_longicauda_ are its long tail, anteriorly truncate tympanic bullae
and large teeth. Characters indicating its affinities with
_noveboracensis_ are smaller size of skull, general narrowness of
skull, and relatively low tympanic bullae. The skull is intermediate as
regards several individual structural features. For example, although
long and narrow and in this feature more nearly approaching
_noveboracensis_, the skull is wider than usual in that subspecies and
thus approaches that of _longicauda_ or _spadix_. The hind foot, in the
dried state, measures 47 millimeters. This large hind foot, obviously
long tail (the specimen lacks external measurements), and anteriorly
truncate bullae constitute basis for here referring the specimen to
_spadix_. However, the seemingly small size of the body and the narrow
skull clearly show relationship to _noveboracensis_.
Specimens, referred to _spadix_, from northern Iowa, are instructive as
showing what happens where the ranges of _noveboracensis_, _primulina_,
_spadix_, and perhaps _longicauda_, meet. No. 47167, Univ. Mich. Mus.
Zoöl., a nearly adult female, taken on November 22, 1915, at Island,
Clay County, and in process of assuming a brown winter pelage, retains
enough of the dark summer pelage to show that the color was slightly
lighter than average for _spadix_. The color pattern, white lips, and
extension of light color of the underparts onto the feet, agrees with
_spadix_ or _longicauda_ as does also the long tooth-row. The overall
length of the skull is intermediate between that of _spadix_ and
_primulina_. The proportions of the anterior part of the skull and of
the tympanic bullae resemble those found in _primulina_. A subadult
male skull only, no. 123846, American Museum of Natural History, from
Webb, Clay County, shows approach to _primulina_ in the narrowness of
the rostrum.
A young male from Ruthven, Iowa, no. 48340, Univ. Michigan, has a large
skull approaching in size that of _spadix_, has the _longicauda-spadix_
type of light-colored underparts and color pattern, and is slightly
darker above than true _longicauda_. Another subadult male in the white
winter coat from Palo Alto County, no. 35756, Univ. Michigan, has a
large skull, which shows approach to _primulina_ in its narrowness
anteriorly and in some other features. Although the tail is of moderate
length, the body is large as in _spadix_ or _longicauda_, and the
length of the hind foot suggests _spadix_ or _longicauda_.
A subadult male, no. 425a, Iowa State College, from Manson, Iowa, in
brown winter pelage, agrees with _primulina_ in the restriction of the
area of the light color of the underparts and in less expanded
zygomatic arches. The teeth are intermediate in size between those of
_noveboracensis_ and _primulina_ on the one hand and those of _spadix_
and _longicauda_ on the other. In other respects it agrees with, or is
more nearly like, _spadix_.
An adult female, no. 426a, Iowa State College, from Barnum, in the
brown winter coat, agrees with _primulina_ except that the orbitonasal
length of the skull is more as in _spadix_ and the presence of some
light color on the lower part of the hind legs suggests _spadix_. The
skull only, no. 440a, Iowa State College, labeled merely Webster
County, Iowa, is almost a duplicate of no. 426a. A subadult male, no.
427a, Iowa State College, from Moorland, Iowa, only about six miles
southeast of Barnum, likewise is indistinguishable from _primulina_
except for having a white winter coat and in being relatively broad in
the mastoidal region. Nevertheless, both of these animals are here
referred to _spadix_ because the average of specimens from this general
area is nearer that of _spadix_. No. 497a, Iowa State College, an adult
female in white winter pelage, from Ames, approaches _primulina_ in the
narrow rostrum and smaller teeth but otherwise approaches or even
agrees with _spadix_.
Two adult males, without external measurements, from Pilot Mound, Iowa,
have skulls quite like males of _longicauda_ from Alberta. The only
approach noted to eastern forms is the restricted color of the
underparts on no. 2856, Coe College, which has a brown winter coat. The
color of the underparts is not extended so far out on the feet as in
_longicauda_. Also the tympanic bullae of this specimen are a trifle
narrower. The other male, no. 2652, is in the white winter coat. The
one female from the same place, no. 2660, Coe College, in brown winter
pelage, has a skull notably unlike that of _longicauda_ or _spadix_;
the skull is narrower and practically indistinguishable from that of
the largest female skull of _primulina_ available from Lawrence,
Kansas, save that the tooth-row is much longer. The color pattern also
agrees with that of _primulina_ or _noveboracensis_ in that the color
of the underparts extends only as far as the knee on the hind legs and
is narrow on the belly. Nevertheless, another adult female, no. 120a
from Amaqua Township, some 6 miles southwest of Pilot Mound, is in all
respects typical of _spadix_. This is the more remarkable because
another comparable specimen from less than 20 miles to the southwest in
Worth Township is equally typical of _primulina_.
Two young females from Chester, Iowa, nos. 2656 and 2874/2873, Coe
College, have skulls larger than those of corresponding age of
_primulina_ or _noveboracensis_. The color is as in spadix. The color
pattern of the underparts also is as in _spadix_ or _longicauda_ except
that the width of the area of light color on the belly is restricted
somewhat although not so much as in _noveboracensis_ or _primulina_.
Of four males from the same place, also in the collection of Coe
College, no. A2874 is a white skin only and does not provide diagnostic
characters. The three other males, each in summer pelage, are marked
and colored as are the two females from the same place except that male
no. 2861 has the color of the underparts so much attenuated on the hind
legs that it barely, uninterruptedly, extends to the feet. No. 2658 is
young, or perhaps barely subadult. The skull is large and referable to
_spadix_. The two adults, nos. 2861 and 2657, differ cranially from
typical (Elk River, Minn.) _spadix_ only in being slightly narrower
across the mastoids and in having the bullae a little narrower. In
these departures they show some approach to _primulina_ and to
_noveboracensis_. Another male, subadult, no. 2867, Coe College, from
Decorah, which has acquired half of the white winter coat, agrees with
the males from Chester except that the preorbital part of the skull is
shortened much as in some specimens of _primulina_.
From Lansing, in extreme northeastern Iowa, a large subadult male, no.
2864, Coe College, of 453 mm. in total length and half through with
acquiring the white winter coat, agrees with the males previously
described from Chester except in having the palate narrower as in
_noveboracensis_. The adult female available from Lansing, no.
2863/2862, Coe College, in white winter pelage except for the top of
the head, although a large skin, has a skull smaller than that of any
_spadix_ or _longicauda_ and of about the same size as that of no.
3838, Univ. Kansas Mus. Nat. Hist., of _primulina_, from Lawrence,
Kansas, except that the skull of no. 2863/2862 is much narrower across
the mastoids. This specimen, then, shows approach to _noveboracensis_
in narrowness of the mastoidal region, to _primulina_ in other respects
and to _spadix_.
Many of these instructive specimens from Iowa, made available to the
present writer by Mr. W. F. Kubichek, were brought together at the Coe
College Museum by the late B. H. Bailey. Most of them were obtained
from trappers who did not supply the conventional external measurements
taken in the flesh. Even though these are lacking, the specimens
clearly show that actual intergradation occurs where the ranges of _M.
f. longicauda_, _spadix_, _noveboracensis_ and _primulina_ meet.
The dark color of the upper parts, restriction of the color of the
underparts on the ankles with the result that the color reaches the
toes in interrupted fashion, and large skull, of no. 18912 of the
Museum of the University of South Dakota, from Roberts County, South
Dakota, clearly place this specimen with _spadix_, rather than with
_longicauda_. Likewise, male, subadult, no. 11376, Univ. South Dakota,
from Clay County, South Dakota, is referable to _spadix_. Although
without external measurements, the specimen obviously is large. The
patch of summer pelage on its head and neck is darker than the summer
pelage of _longicauda_, and the orbitonasal length is greater than the
length of the tympanic bullae; all these features are characters of
_spadix_. The adult male from Fort Sisseton, South Dakota, no. 188407,
United States National Museum, figured by Merriam (1896, p. 20, figs.
7-9), is almost exactly intermediate between _longicauda_ and _spadix_,
although here referred to the latter.
Five specimens, nos. 147375, 147432, 147762, 148720 and 148721, U. S.
Nat. Mus., including 3 skulls only from Beemer, Cuming County,
Nebraska, are intergrades between _M. f. longicauda_, _M. f. primulina_
and _M. f. spadix_. One skin is in white winter pelage and the other, a
female, is in summer pelage which in coloration and color pattern
agrees with that of _spadix_. External measurements of the male agree
with those of _longicauda_. Measurements of the female agree with those
of _spadix_ except that the tail is shorter as in _primulina_. The
skulls are as long as in _longicauda_ but are more slender than in
either _longicauda_ or _spadix_ although nearer the latter in this
respect. In dorsal aspect, the skulls especially posteriorly to the
orbital region, resemble _primulina_. All points considered, the
animals seem best referred to _spadix_.
Although the degree of development of certain morphological features
has been settled upon as indicative of the race _spadix_, some doubt
remains as to where the western boundary of its range should be shown.
This results from the fact that color has been taken into account as
one diagnostic feature and this feature is lacking in the white winter
specimens which, from the following places, are all that are available:
Kittson County, Minnesota; Moorhead, Minnesota; Casselton and Valley
City in North Dakota; Armour, South Dakota and Clay County, South
Dakota. In summary, more specimens in the summer coat will be required
to establish definitely the boundary between the ranges of _longicauda_
and _spadix_.
Surber (1932:49) has referred to additional specimens of this weasel in
the University of Minnesota Museum as from Winona, Hennepin and Isanti
counties of that state.
At Elk River, Minnesota, B. Bailey (1929:156) found this species to be
about half as abundant as _Mustela cicognanii_ and that it is "more
often found in the open timber and about the dry ridges and fields."
Of seventeen adult or subadult skulls of this race from Minnesota, ten
have obvious marks of infestation of the frontal sinuses. In no skull,
however, has the infestation resulted in so much malformation, as
occurs in _noveboracensis_.
_Specimens examined._--Total number, 76, arranged alphabetically
by states and from north to south by counties in each state.
=Iowa.= _Lyon County_: Granite, 1[65]. _Howard County_: Chester,
6[12]. _Winneshiek County_: Decorah, 1[12]; 8 mi. NE Ossian,
1[76]. _Allamakee County_: Lansing, 2[12]. _Clay County_: Island,
1[76]; Webb, 1[2]. _Palo Alto County_: Ruthven, 1[76]; no locality
more definite than county, 1[76]. _Calhoun County_: Manson, 1[65].
_Webster County_: Barnum, 1[65]; Moorland, 1[65]; no locality more
definite than county, 1[65]. _Boone County_: Pilot Mound, 3[12];
Amaqua Township, Sec. 19, 1[65]. _Story County_: Ames, 1[65].
=Minnesota.= _Kittson County_, 1[2]. _Roseau County_: 2-1/2 mi. SW
Roseau, Jadis Township, 1[14]. _Itasca County_: T. 61N, R. 26W,
1[104]. _Clay County_: Moorhead, 2[9]. _Atkin County_: Atkin,
1[50]. _Otter Tail County_: Lake Lizzie, 1[9]; Parkers Prairie,
1[57]. _Grant County_: 3 mi. NW Barrett, 1[76]. _Benton_ (now
Mille Lacs?) _County_: Princeton, 1[91]. _Sherburne County_: Elk
River, 14 (6[59], 4[14], 3[91], 1[74]). _Hennepin County_: Fort
Snelling, 6 (5[2], 1[91]). _Carver? County_: Chaska, 1[60]. _Lac
qui Parle County_: Madison, 5 (3[91], 2[1]); no locality more
definite than county, 2 (1[68], 1[75]). _Yellow Medicine County_:
Wood Lake, 1[2]. _Blue Earth County_: Rapidan, 1[64]. _County_ in
question: Moore Lake, 1[91].
=Nebraska.= _Cuming County_: Beemer, 5[91].
=North Dakota.= _Cass County_: Fargo, 1[91]; Casselton, 1[91].
_Dickey County_: Oakes, 1[91].
=South Dakota.= _Roberts County_, 1[102]. _Marshall County_: Fort
Sisseton, 1[91]. _Douglas County_: Armour, 1[14]. _Clay County_,
1[102].
=Mustela frenata longicauda= Bonaparte
Long-tailed Weasel
Plates 16, 17, 18, 31, 32 and 33
_Mustela longicauda_ Bonaparte, Charlesworth's Mag. Nat. Hist.,
2:38, 1838.
_Putorius longicauda_, Baird, Mamm. N. Amer., p. 169, 1858; Coues,
Fur-bearing animals, p. 136, 1877; Bangs, Proc. Biol. Soc.
Washington, 10:7, figs. 1, 1a of pls. 1, 2 and 3, February 25,
1896; Merriam, N. Amer. Fauna, 11:19, pl. 3, figs. 3, 3a, 4, 4a,
pl. 5, figs. 1, 1a, text figs. 7-9, June 30, 1896.
_Mustela longicauda longicauda_, Bailey, N. Amer. Fauna, 49:166,
January 8, 1927.
_Mustela frenata longicauda_, Hall, Carnegie Instit. Washington
Publ. 473:105, November 20, 1936; Hall, Canadian Field-Nat.,
52:108, October, 1938.
_Mustela frenata_, Sowls, Journ. Mamm., 29:126, May 14, 1948.
_Type._--Possibly not in existence. No. 43.3.3.3 [from Carlton
House, Saskatchewan] in the British Museum of Natural History has
been regarded by several zoölogists as the type. It is a subadult
female, skull and skin, from North America. See the account of _M.
erminea cicognanii_ for reasons for and reasons against regarding
this specimen as the holotype.
No. 43.3.3.3 from the collection of Dr. John Richardson is in the
white winter coat and now (Sept. 24, 1937) is prepared as a study
skin. Evidences of its once having been mounted are: holes in the
soles of the hind feet for supporting-wires, large straight wire
in the tail, folds in the skin of the now backward-projecting hind
feet, and unevenness of the skin on the back resulting from
straightening out the specimen. The tip of the tail and some skin
from the middle of the belly are missing. Otherwise the skin is
intact. The skull is that of an animal in its first year, lacks
the zygomatic arch on each side, but otherwise is complete and
unbroken. The teeth all are present and entire except that p2 on
the right side is missing from its alveolus.
_Range._--Transition and Upper Sonoran life-zones of the Great
Plains, southward from central Alberta, Saskatchewan and southern
Manitoba through eastern Montana, the Dakotas and Nebraska into
southeastern Wyoming, northeastern Colorado and western Kansas.
See figure 29 on page 221.
_Characters for ready recognition._--Differs from _M. f.
primulina_ in near (_h_) Clay Color rather than Brussels Brown of
upper parts, least width of color of underparts more than 40 per
cent of greatest width of color of upper parts, color of
underparts extended onto hind foot rather than stopped short of
ankle, zygomatic breadth more than 28.8 in adult males and more
than 24.1 in adult females; from _M. f. spadix_ in lighter color
being near (_h_) Clay Color, in males by deeper occiput in which
the depth of the skull, exclusive of the sagittal crest and taken
at the anterior border of the basioccipital amounts to more than
59 per cent of the mastoid breadth; from _M. f. oribasus_ in near
(_h_) Clay Color rather than near (14_n_) Brussels Brown color of
the upper parts and in males by deeper occiput in which the depth
of the skull, exclusive of the sagittal crest and taken at the
anterior border of the basioccipital, amounts to more than 59 per
cent of the mastoid breadth; from _M. f. alleni_ in larger size,
adult males having a total length of more than 400 millimeters,
hind foot more than 45, basilar length more than 43.5, and females
having a total length of more than 375 and basilar length not less
than 40.0; from _M. f. nevadensis_ in near (_h_) Clay Color rather
than near (14_n_ to 1) Brussels Brown of upper parts, basilar
length more than 40 in females and averaging more than 45 in
males; from _M. f. neomexicana_ by near (_h_) Clay Color rather
than Buckthorn Brown color of upper parts, absence of white and
Argus Brown facial markings, and length of tooth-rows amounting to
more than 37 per cent of basilar length.
_Description._--_Size._--Male: Five adults from Alberta yield
average and extreme measurements as follows: Total length, 438
(418-473); length of tail, 158 (140-193); length of hind foot, 50
(46-54). Tail averages 56 per cent as long as head and body.
Length of hind foot averaging more than basal length.
Corresponding measurements of five adults and subadults from North
Dakota are as follows: 465 (445-516); 164 (150-179); 51 (50-54).
Tail averages 55 per cent as long as head and body.
Female: Six adults (Alberta, 4; Saskatchewan, 1; Manitoba, 1)
yield average and extreme measurements as follows: Total length,
401 (383-425); length of tail, 145 (141-159); length of hind foot,
43 (41-44). Tail averages 57 per cent as long as head and body.
Length of hind foot more or less than (approximately equal to)
basal length.
The average differences in external measurements of the two sexes
are: Total length, 37; length of tail, 13; length of hind foot, 7.
General comparisons indicate that the Alberta-taken males may not
attain so large a size as those from some other areas. Thus the
differences in external measurements might be some greater
elsewhere, say, in North Dakota.
_Externals._--Longest facial vibrissae black, brown or white
(often all three colors in same specimen) and extending beyond
ear; carpal vibrissae same color as underparts and extending to
apical pad of fifth digit; hairiness of foot-soles (in summer
pelage) only slightly greater than shown in figure 20.
_Color._--Winter pelage all white except tip of tail. Summer
pelage with upper parts near (_h_) Clay Color or near tone 3 and 4
of Snuff Brown of Oberthür and Dauthenay, pl. 303. Chin and upper
lips white. Remainder of underparts ranging from near (_a_) Olive
Ocher to near (16´) Ochraceous Buff. Upper parts of uniform color
except for occasional darkening of head in front of ears. Color of
underparts extends distally on posterior sides of forelegs over
toes onto antipalmar faces of feet and wrists, on medial sides of
hind limbs to ankles over antiplantar faces of toes and
distomedial third of each tarsus, and over proximal fourth to
third of under side of tail. Least width of color of underparts
averaging, in a series of 10 males from Alberta, 58 (45-60) per
cent of greatest width of color of upper parts. Corresponding
figures for 10 females from the same place are 57 (50-74). Black
tip of tail in same series of males, most of which are in full
summer pelage, averaging 43 (35-60) mm. long. Thus, averaging
shorter than hind foot and 27 per cent of length of
tail-vertebrae.
As compared with _M. f. neomexicana_, _longicauda_ lacks the white
facial markings, black ears, black forehead and nose, but
otherwise is similarly colored. As compared with _M. f.
nevadensis_, _M. f. oribasus_ and _M. f. spadix_, each of color
pattern similar to _longicauda_, selected differences of
_longicauda_ are its much lighter color, especially of the upper
parts, with less conspicuous darkening on the nose. From _M. f.
primulina_, _longicauda_ differs in lighter color of upper parts,
reddish rather than yellowish underparts, and light rather than
dark-colored hind feet.
_Skull and teeth._--Male (based on 5 adults from Alberta): See
measurements and plates 16-18; weight, 4.7 (4.6-4.9) grams;
basilar length, 46.0 (44.7-46.8); zygomatic breadth more than
distance between condylar foramen and M1 or than between anterior
palatine foramen and anterior margin of tympanic bulla; mastoid
breadth more than postpalatal length; postorbital breadth less
than length of upper premolars and more than width of
basioccipital measured from medial margin of one foramen lacerum
posterior to its opposite; interorbital breadth greater than
distance between foramen opticum and anterior margin of tympanic
bulla; breadth of rostrum more or less (usually less) than length
of tympanic bulla; least width of palate less than greatest length
of P4; anterior margin of tympanic bulla as far posterior to
foramen ovale as width of 3 to 4 (including I3) upper incisors;
height of tympanic bulla more than distance from its anterior
margin to foramen ovale; length of tympanic bulla more than length
of lower molar and premolar tooth-row and longer or shorter than
rostrum; anterior margin of masseteric fossa below talonid of m1
or anterior half of m2.