Hist.); Umbagog Lake, 1. _Franklin County_: Seven Pd. Township, 1.
_Piscataquis County_: Grenville, [= Greenville?], 1. _Hancock
County_: Bucksport, 10. _Washington County_: 3rd Mopang Lake, 1
(Acad. Nat. Sci. Phila.); Perry, 1 (Boston Soc. Nat. Hist.).
_County_ in question: Moosehead Lake, 1.
=Mustela frenata primulina= Jackson
Long-tailed Weasel
Plates 16, 17, 18, 31, 32 and 33
_Mustela primulina_ Jackson, Proc. Biol. Soc. Washington, 26:123,
May 21, 1913.
_Putorius noveboracensis_, Baird, Mamm. N. Amer., p. 166, 1858
(part).
_Mustela longicauda longicauda_, Dice, Journ. Mamm., 4:108, May 9,
1923.
_Mustela longicauda primulina_, Linsdale, Journ. Mamm., 9:141, May
9, 1928.
_Mustela frenata primulina_, Hall, Carnegie Instit. Washington
Publ. 473:104, November 20, 1936.
_Mustela frenata_, Leopold and Hall, Journ. Mamm., 26:143, July 19,
1945.
_Type._--Male?, young, skull and skin; no. 168006, U. S. Nat.
Mus., Biol. Surv. Coll.; 5 miles northeast of Avilla, Jasper
County, Missouri; May 11, 1905; obtained by Hartley H. T. Jackson;
original no. 7869X.
The skin lacks the distal part of the tail--the part which bears
the black tip. Otherwise the skin is complete and well preserved.
The teeth of the permanent dentition all are present and entire.
The lower jaws are complete and unbroken. The skull is broken
transversely through the interorbital region, transversely through
the braincase and longitudinally through the basioccipital. Both
zygomatic arches are gone. The type is judged to be a male rather
than a female as stated by the original describer, Jackson
(1913:123), whose measurements of hind foot, interorbital
constriction, maxillary tooth-row, and mandibular tooth-row agree
with those of males and are larger than those of any female seen
of this subspecies.
_Range._--Upper and Lower Austral life-zones west of the
Mississippi River in Missouri and Arkansas, the southeastern half
of Iowa, eastern half of Kansas and Oklahoma, northern Louisiana
and northeastern Texas. Southern and southwestern limits of range
undetermined. See figure 29 on page 221.
_Characters for ready recognition._--Differs from _M. f.
noveboracensis_ in males by interorbital breadth averaging less
than 24 per cent of basilar length, orbitonasal length averaging
less than 34 per cent of basilar length or 64 per cent of mastoid
breadth, tympanic bullae as much inflated anteromedially as
posteromedially, and in females by orbitonasal length amounting to
less than two-thirds of mastoid breadth, by zygomatic breadth
averaging more than 21 mm., and by anterolateral margin of
tympanic bulla projecting below squamosal; from _M. f. spadix_ by
least width of color of under parts amounting to less than 40 per
cent of greatest width of color of upper parts, by absence of
color of underparts on hind leg below knee, and by smaller size
(hind foot less than 50 in males and 40 in females; orbitonasal
length less than 15.5 in males and 13.5 in females; length of
tooth-rows less than 18 in males and 15.7 in females; mastoid
breadth less than 25.5 in males and 22 in females); from _M. f.
longicauda_ by Brussels Brown rather than near (_h_) Clay Color of
upper parts, least width of underparts less than 40 per cent of
greatest width of color of upper parts, absence of color of
underparts on hind leg below knee, zygomatic breadth less than
28.8 in males and 24.1 in females; from _M. f. neomexicana_ by
Brussels Brown rather than Buckthorn Brown color of upper parts,
in absence of white frontal spot and broad white bands on sides of
head, in anterolaterally rounded, rather than "square," tympanic
bullae and in zygomatic breadth of less than 30 in males and 24 in
females; from _M. f. frenata_ and _M. f. texensis_ by absence of
white facial markings and smaller size (basilar length of adult
males less than 47, tail-length less than 155 in males, and hind
foot less than 40 in females); from _M. f. arthuri_ by less evenly
spreading zygomatic arches (see pls. 16, 17 and 18), greater
inflation of the tympanic bullae anteromedially and more nearly
straight (less convex) dorsal outline of skull.
_Description._--_Size._--Male: Eighteen adults and subadults from
Douglas County, Kansas, yield average and extreme measurements as
follows: Total length, 397 (371-440); length of tail, 133
(120-147); length of hind foot, 43 (40-47). Tail averages 50 per
cent as long as head and body. Length of hind foot averages less
than basal length. Corresponding measurements, originally taken in
inches and fractions thereof, of 9 adults and subadults from Boone
County, Arkansas, are as follows: 413 (384-438); 138 (127-155); 41
(38-44).
Female: Six adults and subadults from Douglas County, Kansas,
yield average and extreme measurements as follows: Total length,
339 (317-355); length of tail, 107 (95-115); length of hind foot,
35 (34-37). Tail averages 46 per cent of length of head and body.
Length of hind foot less than basal length. Corresponding
measurements, originally taken in inches and fractions thereof, of
5 adults and subadults from Boone County, Arkansas, are as
follows: 355 (332-397); 113 (101-127); 33 (29-38).
The average differences in external measurements of the two sexes,
in Douglas County, Kansas, are: Total length, 58; length of tail,
26; length of hind foot, 8. An adult male from Boone Co., Iowa,
weighed 293 grams.
_Externals._--Longest facial vibrissae black or dark brown (often
both colors in same specimen) and extending beyond ear; carpal
vibrissae colored either like underparts or upper parts and
extending to apical pad of fifth digit; hairiness of foot-soles as
shown in figure 20.
_Color._--Upper parts, in summer, Brussels Brown to near (14 _n_)
Brussels Brown, or tones 2 to 4 of Raw Umber of Oberthür and
Dauthenay, pl. 301. Chin and rarely upper lips white. Remainder of
underparts Picric Yellow to Primuline Yellow. In winter, color
essentially the same except for smoke-gray effect in upper parts
and more whitish in underparts. Tip of tail at all times black.
Upper parts of uniform color except for occasional darkening of
nose and mid-dorsal region. Color of underparts extends distally
on posterior sides of forelegs onto antipalmar faces of toes, on
medial sides of hind legs only to a point between knee and ankle.
Least width of color of underparts averaging, in a series of 21
males from Lawrence, Kansas, 23 (9-35) per cent of greatest width
of color of upper parts. Black tip of tail in same series, most of
which are in full winter pelage, 52 (40-70) mm. long; thus longer
than hind foot and averaging 39 per cent of length of
tail-vertebrae.
_Skull and teeth._--Male (based on ten adults from Douglas County,
Kansas): See measurements and plates 16-18; weight, 3.7 (3.3-4.2)
grams; basilar length, 44.8 (43.8-46.0); zygomatic breadth more or
less (less in 80 per cent) than distance between condylar foramen
and M1 or than between anterior palatine foramen and anterior end
of tympanic bulla (less in 70 per cent); mastoid breadth more or
less (less in 80 per cent) than postpalatal length; postorbital
breadth less than length of upper premolars and, except in one
specimen, more than width of basioccipital measured from medial
margin of one foramen lacerum posterior to its opposite;
interorbital breadth more or less (less in 70 per cent) than
distance between foramen opticum and anterior margin of tympanic
bulla; breadth of rostrum less than length of tympanic bulla;
least width of palate less than outside length of P4 (except in
one specimen); anterior margin of tympanic bulla as far posterior
to foramen ovale as width of 2-1/2 to 5 upper incisors; height of
tympanic bulla more (except in one specimen) than distance from
its anterior margin to foramen ovale; length of tympanic bulla
more than length of lower molar and premolar tooth-row and longer
(except in one specimen) than rostrum; anterior margin of
masseteric fossa behind or just below posterior eighth of m1.
Female (based on 5 adults and subadults from Douglas County,
Kansas): See measurements and plates 31-33; weight, 2.2 (2.0-2.4)
grams; basilar length, 38.9 (37.6-40.7); zygomatic breadth less
than distance between condylar foramen and M1 or than between
anterior palatine foramen and anterior margin of tympanic bulla;
postorbital breadth less than length of upper premolars and except
in one specimen, more than width of basioccipital measured from
medial margin of one foramen lacerum posterior to its opposite;
least width of palate less than greatest length of P4; tympanic
bulla as far posterior to foramen ovale as width of 3 to 5 upper
incisors; height of tympanic bulla more or less than distance from
its anterior margin to foramen ovale; length of tympanic bulla
more than length of lower molar and premolar tooth-row and longer
than rostrum.
The skull of the female averages 41 per cent lighter than that of
the male.
Compared with the skull of _M. f. noveboracensis_ from Massachusetts,
that of the male of _primulina_, in dorsal view, is seen to be shorter
anteriorly to the postorbital processes and to have a more marked
postorbital constriction. In lateral view the dorsal outline of the
skull of _primulina_ is less concave in the postorbital region. In
ventral view the skull of _primulina_ is seen to be wider across the
mastoid processes and zygomatic arches but the most pronounced
difference is in the tympanic bullae. In _noveboracensis_ each bulla is
scooped out on the anterior part of the medial face and appears to be
narrower anteriorly than posteriorly whereas in _primulina_ the
anterior part of the medial face is not scooped out but is moderately
inflated and the bulla appears to be of uniform breadth anteriorly and
posteriorly. By actual measurement the breadth of the bulla averages 59
per cent of its length in _primulina_ but only 50 per cent in
_noveboracensis_. Other respects in which the skull of the male of
_primulina_ differs from that of _noveboracensis_ are as follows:
Linear measurements of teeth more; relative to the basilar length, the
length of the tooth-rows averages more, whereas the interorbital
breadth and orbitonasal length are less.
When skulls of females are compared, each of the differences mentioned
above is found to apply, except that the degree of difference is in
some parts greater, for example, in the tympanic bullae. In
_primulina_, the bulla is in general like that of the male
_noveboracensis_, whereas in the female _noveboracensis_ it is less
inflated, especially anteromedially, shorter, relatively narrower, and
in ventral view projects little or none below the squamosal floor of
the braincase. The breadth of the bulla averages 51 per cent of its
length in _primulina_ but only 47 per cent in _noveboracensis_. The
bullae project below the basioccipital on the average, for a distance
of 2.9 millimeters in female _primulina_ and only 2.3 millimeters in
female _noveboracensis_. In _primulina_ the temporal ridges are well
developed and fuse to form a low sagittal crest, but in
_noveboracensis_ the ridges are absent. Also, in _primulina_ the
mastoid processes project farther laterally beyond the braincase. The
skull of female _noveboracensis_ is much lighter than that of
_primulina_. Average weights of the two are 1.7 and 2.2 grams. The
skulls of females of _primulina_ and _noveboracensis_ differ more than
do the skulls of males.
Compared with the skull of _spadix_, that of the male, and the female,
of _primulina_ averages smaller in every part measured. Expressed in
percentages of the basilar length, the two depth measurements of the
skulls are not significantly different, but, excluding the measurements
of the bullae and teeth, the other cranial measurements are less. The
main difference in relative proportions is in the tympanic bullae which
average only a half millimeter shorter in males of _primulina_ and one
and one-tenth millimeters shorter in females. The bullae are,
therefore, relative to the basilar length, longer in _primulina_. The
skull of _primulina_, then, differs from that of _spadix_ mainly in
smaller size and relatively longer tympanic bullae, especially in
males.
Compared with the skull of _M. f. longicauda_, that of both sexes of
_primulina_ averages smaller in every part measured, except in males
where the length of the tympanic bulla, and breadth and length of M1
are the same or slightly larger in _primulina_. Relative to the basilar
length, the length of the tympanic bullae, and in females only, the
depth measurements are greater in _primulina_ but all the others, in
both sexes, are less. These ratios reflect the relative narrowness of
the skull of _primulina_. Upon direct comparison the narrowness is
especially noticeable in the interorbital region, mastoid region,
tympanic bullae, and across the zygomata.
Compared with the skull of _M. f. neomexicana_ that of both sexes of
_primulina_ averages smaller in every part measured. Excepting the
measurements of the teeth, most of the other measurements are
constantly larger. Relative to the basilar length, the length of
tooth-rows and length of tympanic bulla are more, but excepting the
depth measurements, the others are less. Still other differences are,
in _primulina_, less well-developed sagittal crest, anterolateral
corner of bulla rounded rather than "square," and in males a
transversely convex rather than flat interorbital region.
Compared with _M. f. frenata_ and _M. f. texensis_, the skulls of males
of _primulina_ differ in being smaller in every part measured but
relative to the basilar length, have longer tooth-rows, a lesser
zygomatic breadth and are less constricted interorbitally.
Compared with the skull of _M. f. olivacea_, those of both sexes of
_primulina_ average smaller in every part measured, have shallower
(dorsoventrally) tympanic bullae, a lower sagittal crest and slightly
weaker postorbital processes on the frontals. Relative to the basilar
length, the several cranial measurements are about the same.
Comparison of the skull with that of _M. f. arthuri_ has been made in
the account of that subspecies.
_Remarks._--The first specimens of this race known to have been
preserved in study collections are one in the United States National
Museum, taken at Bridge, Carroll County, Missouri, many years ago by J.
Burbage, and less than a dozen specimens preserved before 1900 from
eastern Kansas in the University of Kansas Museum of Natural History.
In 1913 Hartley H. T. Jackson bestowed a name on this animal on the
basis of two specimens taken by him in southwestern Missouri. Later,
through the efforts of Charles D. Bunker, and his associates at the
University of Kansas, nearly 100 specimens were saved from eastern
Kansas, principally from Douglas County. In the course of the present
study, Lawrence V. Compton obtained a topotype for the California
Museum of Vertebrate Zoölogy, and with the assistance of Mr. B. G.
Roberts, a good series of specimens from Boone County, Arkansas, was
preserved in the same museum. In the early years of the 20th Century,
the late B. H. Bailey at Coe College, Iowa, collected specimens from
that state. The specimens from these several sources suffice to give a
relatively clear idea of the characters of this subspecies.
_Mustela frenata primulina_ is closely related to _M. f.
noveboracensis_, from which, on the average, it differs in the lighter
color of the upper parts of the summer coat, in the more intense
coloring of the underparts, and in certain cranial features pointed out
above. In the southern part of its range, however, _noveboracensis_ has
the underparts only a little less intensely colored with yellow than
_primulina_. Also, the skull of the one topotype from 7-1/2 miles
southeast of Carthage, a subadult male in brown, winter pelage, is
almost exactly intermediate between that of _noveboracensis_ from
Massachusetts and _primulina_ of Douglas County, Kansas, and Boone
County, Arkansas. _M. f. primulina_ often has the underparts white in
winter, as does this topotype which agrees with the average of
_noveboracensis_ in small size of teeth and narrowness across the
mastoid processes and zygomatic arches. However, it agrees with
_primulina_ in shape and relative size of the rostrum. It is almost
exactly intermediate in shape and width of the tympanic bullae.
Three other males, but no females, all in winter pelage, are available
from eastern Missouri. Of the two from Silex, Lincoln County, one is
nearer _noveboracensis_ and the other nearer _primulina_ on the basis
of cranial characters. The third specimen, from four miles south of
Lesterville, so far as I can determine by examination of individual
cranial characters and tabulation of results, is exactly intermediate.
Final decision on the proper allocation of specimens from the parts of
Missouri represented can best be made when skulls of females are
available. From the fact that the skull of the female referred to
_noveboracensis_ from Golconda, Illinois, shows almost as many
characters of _primulina_ as of _noveboracensis_, it is judged that
females from as far west as Silex and Lesterville, Missouri, will show
even more characters of _primulina_ and so be referable to that form.
If this supposition be correct, the present reference of the almost
exactly intermediate males, from eastern Missouri, will stand;
otherwise, it may not.
Additional intergrades with _noveboracensis_ are available from eastern
Iowa. Of five specimens from Hillsboro, Iowa, two males and a female
have tympanic bullae like those of _primulina_ but the other two males
have bullae like those of _noveboracensis_. The female is smaller than
_primulina_ and in this small size and in general configuration of the
skull, viewed dorsally, is more nearly like _noveboracensis_. As a
whole, the population averages almost exactly intermediate. The same is
true of 3 males and one female from Muscatine. The subadult male from
Keosaqua, to my eye, resembles _noveboracensis_ in the greater length
of the skull anteriorly to the postorbital processes, and in the
relative narrowness across the mastoidal region, but otherwise is more
like that of _primulina_. Two males and one female from Tipton,
although in each instance variously intermediate, are as a whole nearer
_primulina_, No. 2865, Coe College, male adult, from Cedar Rapids, has
characters of the three races, _spadix_, _noveboracensis_ and
_primulina_. In the skull, the width suggests _spadix_, the narrow
mastoid region, _noveboracensis_, and the tympanic bullae are as in
_spadix_ or _primulina_. One male, no. 12, Coe College, from Dubuque,
is as narrow across the mastoid region as is _noveboracensis_ although
the bullae are well inflated as in _primulina_. The skull, without
corresponding skin, of a female, no. 140a, Iowa State College, from
Green's Island, also resembles _noveboracensis_ in narrowness of the
mastoidal region, and in small size of skull, but in larger teeth,
broader tympanic bullae, and sagittal crest is referable to
_primulina_. Of two females from Vinton, one adult is typical of
_primulina_ but the other, a subadult, is practically indistinguishable
from female _noveboracensis_, from Ann Arbor, Michigan. Three males
from Vinton agree well with _primulina_ except that the interorbital
region is wider than average and thereby suggests _spadix_ or
_noveboracensis_. An adult female from New Hartford also is typical of
_primulina_ except for the broader interorbital region. Three males
from Fayette are typical of _primulina_.
Other specimens from Iowa are intergrades with _spadix_, or if not with
_spadix_, with the animal of northwestern Iowa which in some ways
combines the characters of _longicauda_ and _spadix_. For example, no.
2665, Coe College, an adult male from Davenport, has the anterior part
of the skull (all that is preserved) heavily ridged as in _spadix_ and
in addition, the underparts are marked with the shade of reddish
displayed by topotypes of _spadix_ and with some yellowish as seen in
_longicauda_. The color pattern, however, is as in _primulina_. A young
male, no. C-51, Iowa State College, from Kelley, Story County, has
anteriorly truncate bullae as are more frequently found in the
_longicauda-spadix_ stock of northwestern Iowa, than in _primulina_. In
other respects, the animal, in so far as can be judged from the broken
skull, agrees with _primulina_ as it certainly does in color, color
pattern, and external measurements. An adult male, no. 499a, Iowa State
College, from 2 miles east of Ledges St. Park, in Boone County, in
short body, size of teeth, and size of skull, in so far as the broken
parts can be measured, resembles _primulina_ more closely than it does
any other subspecies. The long tail, long hind foot, wide extent of the
light-colored underparts, and extension of the color of the underparts
onto the hind feet are more as in _spadix_. Other intergrades with
_spadix_ from Iowa are mentioned in the account of _spadix_.
The specimen from Swartz, Louisiana, suggests intergradation with
_arthuri_ in that the anteromedial part of the tympanic bulla is less
inflated than in typical _primulina_.
Intergrades with _longicauda_ are available from Riley and Pratt
counties, Kansas. No. 7182, Univ. Kans., subadult male in winter
pelage, from near Winkler, has a skull of larger size as in
_longicauda_ with which race it seems to agree in large size of body,
tail and hind foot, although the collector's measurements are lacking.
Color pattern and relative proportions of the skull throughout are as
in _primulina_. The young male, no. 3495, Univ. Kans., from Pratt,
Kansas, agrees in external measurements and large size of skull with
_longicauda_, but has the color and color pattern precisely as in
_primulina_. The teeth are smaller as in _primulina_. Immaturity
prevents judging of its relationships on the basis of relative
proportions of the skull.
The two specimens, skins only, available from Oklahoma, are
provisionally referred to _primulina_. These are remarkable for the
restriction of the color of the underparts and for the intensity of the
yellow coloration of the underparts. The specimen from Norman has the
color of the underparts entirely absent from the hind legs and not
extending posteriorly to the penis. On the chest and lower throat,
large spots of color of the upper parts are present and the yellow area
of the underparts on the belly is narrower than in any other specimen
of _primulina_ examined. The specimen from 8 miles northwest of
Stillwater has the color of the underparts only a little less
restricted although this color does extend over the inguinal region
almost to the knees. The skin of the posterior part of the body of a
weasel is available from 10 miles south of Sulphur Springs, Texas. It,
likewise, is only provisionally referred to _primulina_. The coloration
is about as in the specimens from Oklahoma but the distribution of the
color of the underparts cannot be made out.
The dark color of the upper parts occurs far westward in animals which
otherwise display characters of _longicauda_. Among these intergrades,
the larger size of _longicauda_ generally is combined with this dark
color. This geographic behavior of the dark color of the upper parts is
analogous to the condition described in _M. f. spadix_. Stated in
another way, the dark color of the upper parts is the character, of the
eastern animal, last to disappear as one goes westward across the
Mississippi Valley toward the range of _longicauda_ which is a
subspecies of markedly different size, shape of skull, and coloration.
Only two of 29 specimens from Kansas show infestation of the frontal
sinuses. All four of the specimens from Missouri have the frontal
sinuses malformed as do 9 of the 14 from Arkansas examined in this
respect.
An adult female from Boone County, Iowa, bears the date May 9, 1938,
and the annotation by T. G. Scott, "Fox-killed."