Part 28

_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."

_Specimens examined._--Total number, 131, arranged alphabetically by states and from north to south by counties in each state. Except as otherwise indicated, specimens are in the University of Kansas, Museum of Natural History.

=Arkansas.= _Boone County_: 3 mi. E Bergman, 4[74]; 3 mi. SE Bergman, 1[74]; 3 mi. S Bergman, 1[74]; 3 mi. SW Bergman, 1[74]; 4 mi. SE Bergman, 2[74]; 5 mi. SE Bergman, 1[74]; 4-1/2 mi. SE Bergman, 3[74]; 5 mi. SE Bergman, 1[74]; 5 mi. S Bergman, 2[74]; 5 mi. SW Bergman, 2[74]. _Washington County_: Fayetteville, 1[96]. _Crawford County_: 10 mi. S Winslow, 1. _Sebastian County_: Fort Smith, 1[91].

=Iowa.= _Fayette County_: Fayette, 3[12]. _Dubuque County_: Dubuque, 1[12]; Green's Island, 1[65]. _Butler County_: New Hartford, 1[12]. _Hardin County_: Union, 1[65]. _Benton County_: Vinton, 5[12]. _Linn County_: Cedar Rapids, 1[12]. _Boone County_: Worth Township, Sec. 21, 1[65]; 2 mi. E Ledges St. Park, 1[65]. _Story County_: Kelley, 1[65]. _Cedar County_: Tipton, 3[12]. _Scott County_: Davenport, 2[12]. _Muscatine County_: Muscatine, 4[12]. _Henry County_: Hillsboro, 5[91]. _Van Buren County_: Keosaqua, 1[65]; no locality more definite than county, 1[50]. _Taylor County_, 1.

=Kansas.= _Riley County_: near Winkler, 1. _Pottawatomie County_: Onaga, 1[83]. _Atchison County_: Doniphan Lake, 1; 5 mi. NE Muscotah, 1; no locality more definite than county, 1. _Douglas County_: Lawrence, 8; 6 mi. NW Lawrence, 1; 1-1/2 mi. W Lawrence, 1; 6 mi. S Lawrence, 1; 7 to 7-1/2 mi. SW Lawrence, 14; 10 mi. W Lawrence, 1; Clinton, 4; Baldwin, 1; no locality more than county, 29 (2[74]). _Woodson County_: 1-1/2 mi. S Neosho Falls, 1[59]. _Greenwood County_: 8 mi. SW Toronto, 2. _Pratt County_: Pratt, 1.

=Louisiana.= _Quachita Parish_: Swartz, 1[71].

=Missouri.= _Carroll County_: Bridge Creek, 1[91]. _Lincoln County_: Silex, 1[74]; 1 mi. E Silex, 1[74]. _Reynolds County_: 4 mi. S Lesterville, 1[74]. _Jasper County_: 5 mi. NE Avilla, 1[91]; 7-1/2 mi. SE Carthage, 1[74].

=Oklahoma.= _Payne County_: 8 mi. NW Stillwater, 1[82]. _Cleveland County_: Norman, 1[100].

=Texas.= _Hopkins County_: 10 mi. S Sulphur Springs, 1[43].

=Mustela frenata arthuri= Hall

Long-tailed Weasel

Plates 16, 17 and 18

_Mustela noveboracensis arthuri_ Hall, Proc. Biol. Soc. Washington, 40:193, December 2, 1927.

_Mustela frenata arthuri_ Hall, Carnegie Instit. Washington Publ. 473:105, November 20, 1936.

_Type._--Male, subadult, skull and skin; no. 37515, Mus. Vert. Zoöl.; Remy, St. James Parish, Louisiana; December 15, 1926; obtained by Stanley C. Arthur.

The skin is stuffed and well preserved. The skull (plates 16-18) is unbroken. The teeth all are present and entire. The presence of a well-developed scrotal pouch shows the specimen to be a male. Contrary to what was stated in the original description the specimen was taken in 1926 and not in 1925.

_Range._--Lower Austral Life-zone of southeastern Texas, Louisiana, and into Mississippi. See figure 29 on page 221.

_Characters for ready recognition (of males)._--Differs from _M. f. olivacea_ in smaller size (adult males with hind foot and basilar length less than 45), depth of skull at anterior margin of basioccipital, ignoring sagittal crest, amounting to more than 63 per cent of mastoid breadth, and greater convexity of dorsal outline of skull in longitudinal axis (see pls. 16-18); from _M. f. noveboracensis_, in males, by zygomatic breadth not less than distance between anterior palatine foramen and anterior margin of tympanic bulla and by convex dorsal outline of skull in longitudinal axis; from _M. f. primulina_ by evenly spreading zygomatic arches, lesser inflation of tympanic bullae anteromedially than posteromedially, and convex dorsal outline of skull in longitudinal axis; from _M. f. texensis_ and _M. f. frenata_ by absence of white facial markings and postorbital breadth more than distance between posterior borders of P4 and P2.

_Description._--_Size._--Male: The type, a subadult male, measures (inches and quarter fractions thereof, transposed into millimeters) as follows: Total length, 390; length of tail, 113; length of hind foot, 44. Tail is 41 per cent as long as head and body. Length of hind foot less than basal length.

Typical female unknown.

_Externals._--Longest facial vibrissae black, or dark brown (both colors in the type) and extending beyond ear; carpal vibrissae same color as underparts and extending to within 3.5 millimeters of apical pad of fifth digit. Hairiness of foot-soles in type slightly less than shown in figure 20.

_Color._--Upper parts in summer tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304; underparts as described in _M. f. olivacea_. In winter, upper parts (based on type) near (1) Brussels Brown or grayer than tone 4 of Burnt Umber of Oberthür and Dauthenay, pl. 304, darker on top of head from nose to a line connecting posterior margins of ears. Chin and posterior third of each upper lip white. Remainder of underparts white with wash of Warm Buff. Tip of tail black. Color of underparts extends distally on posterior sides of forelegs over toes but represented on antipalmar faces of feet by only a few scattered hairs. Color of underparts extends distally on medial sides of hind limbs only to knees. Least width of color of underparts amounting to 15 per cent of greatest width of color of upper parts. Black tip of tail 50 mm. long; thus longer than hind foot and 44 per cent as long as tail-vertebrae.

_Skull and teeth._--Male (based on type and 2 subadults): See measurements and plates 16-18. As described in _M. f. noveboracensis_ except that: Weight, 4.0 (3.7-4.3) grams; basilar length, 43.5 (43.3-43.6); zygomatic breadth not less than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than length of upper premolars; interorbital breadth more than distance between foramen opticum and anterior margin of tympanic bulla; least width of palate more or less than length of P4; tympanic bulla longer than rostrum.

Female: Typical skull unknown. The skull from 12 miles east of Eagle Lake, Texas, lacks the convexity in the dorsal longitudinal axis and the skull agrees with those of larger individuals of _primulina_ except that the anteromedial faces of the tympanic bullae are less inflated, and the mastoid and zygomatic breadths are greater than in any female seen of _primulina_. Probably this greater breadth is the result of intergradation with _M. f. frenata_ to the westward.

Compared with the skull of _M. f. olivacea_ that of _arthuri_ differs as follows: Averaging smaller in every part measured; basilar length 5 mm. less; by weight a fourth lighter; relative to basilar length, interorbital breadth greater and zygomatic and especially mastoid breadth less; dorsal outline of skull more convex in longitudinal axis; tympanic bullae narrower and less inflated especially on anteromedial faces. Compared with the skull of _noveboracensis_ that of _arthuri_ has the zygomatic breadth equal to or exceeding the distance from the anterior palatine foramen to the anterior margin of the tympanic bulla, whereas the zygomatic breadth is less than this distance in _noveboracensis_. Also, in _arthuri_, the rostrum is relatively shorter, the braincase is more inflated anteriorly, the zygomatic arches are more uniformly spreading, and the dorsal outline of the skull is distinctly convex, both transversely and longitudinally, whereas it is transversely more nearly flat in _noveboracensis_ and longitudinally is concave in the interorbital region.

Compared with _M. f. primulina_, _arthuri_ has narrower bullae, which are much less inflated on their anteromedial faces, a less marked postorbital constriction, a braincase which is narrower across the mastoid region and broader anteriorly, and a skull, which, in longitudinal axis, has the dorsal outline markedly more convex.

Compared with the skull of _M. f. texensis_ that of _arthuri_ is smaller in every part measured; length one-fifth less; one-half as heavy; postorbital constriction less marked; braincase relatively narrower posteriorly and tympanic bullae less inflated especially anteromedially. Compared with the skull of _M. f. frenata_ that of _arthuri_ is smaller in every part measured; basilar length 6 mm. less; a third lighter; postorbital constriction less marked; relative to the basilar length the rostrum is broader, longer and deeper; the zygomatic expanse and breadth of the braincase across the mastoids is less; the dorsal profile of the skull is more convex in longitudinal axis; zygomata evenly spreading rather than abruptly protruding from skull posteriorly; tympanic bullae less inflated anteromedially.

_Remarks._--In 1926, Stanley C. Arthur, then Director of the Division of Wild Life, for the Louisiana State Department of Conservation, obtained specimens of this weasel. Some of them were mounted and the remainder were placed in the collections of the United States National Museum and the Museum of Vertebrate Zoölogy. In 1938 to 1940 George H. Lowery saved specimens from Baton Rouge, which showed the color of the summer pelage and revealed that the size of males was more than was indicated by the original materials. In 1940 and 1941 Rollin H. Baker obtained specimens from eastern Texas which greatly extended the known geographic range.

In addition to the localities represented by specimens examined, Arthur (1928:117) has recorded specimens from Greensburg, St. Helena Parish; Braithwaite, Plaquemines Parish; Geismar, Assumption Parish; Laurel Hill, West Feliciana Parish; French Settlement, Livingston Parish; and Kentwood, Tangipahoa Parish. All these localities lie within the eastern half of southern Louisiana. A skin-only, no. 38902, Mus. Vert. Zoöl., obtained from a fur buyer by Stanley C. Arthur, was taken in Mississippi "south of Jackson." Possibly it is of the subspecies _arthuri_.

Intergradation with _M. f. olivacea_ is indicated by a specimen from Mobile County, Alabama, commented on in the account of _olivacea_. Intergradation with _primulina_ is indicated by the shape of the anteromedial part of the bullae of the specimen from Swartz, Louisiana, that is referred to _primulina_. The lack of specimens from the northern two-thirds of Mississippi and from western Tennessee, prevents any definite statement as to the limits of range of _arthuri_ in those areas. In comparison with animals from the type locality, the slightly larger size of the adult male from Baton Rouge, and the still larger size of the adult male of _primulina_ from Swartz, Louisiana, suggests that the _olivacea_ "influence" may extend farther west in the latitude of northern Louisiana than anywhere else.

None of the skulls examined shows malformation of the frontal sinuses such as results from infestation by parasites in some races. Arthur (1928:115) speaks of the ". . . cut-over swamp land, where the tupelo and cypress have been removed, . . ." as constituting suitable habitat for this animal.

_Specimens examined._--Total number, 13, as follows:

=Texas.= _Colorado County_: 12 mi. N Eagle Lake, 1[43]; 5 mi. W Eagle Lake, 1[43]; 3 mi. S Garwood, 1[43].

=Louisiana.= _East Baton Rouge Parish_: Baton Rouge, 4[71]. _Livingston Parish_: Springville, 1[74]. _Saint James Parish_: Convent, 1[91]; Remy, 2 (1[74], 1[45]). _Assumption Parish_: near Lake Verret, 1[45].

=Mississippi.= _Harrison County_: Saucier, 1[71].

=Mustela frenata olivacea= Howell

Long-tailed Weasel

Plates 16, 17, 18, 31, 32 and 33

_Mustela peninsulae olivacea_ Howell, Proc. Biol. Soc. Washington, 26:139, May 21, 1913.

_Mustela frenata olivacea_, Hall, Carnegie Instit. Washington Publ. 473:104, November 20, 1936.

_Type._--Male, adult, skull and skin; no. 180802, U. S. Nat. Mus., Biol. Surveys Coll.; Autaugaville, Autauga County, Alabama; December 22, 1912; obtained by L. S. Golsan.

The skull (plates 16-18), although cracked at two places in the interorbital region, is in one piece and not warped out of shape. The teeth all are present and entire. The skin is exceptionally well made and in perfect condition except for the extreme tip of the tail which is broken off.

_Range._--Lower and Upper Austral life-zones in eastern Mississippi, Alabama, Georgia, South Carolina, and northern Florida. See figure 29 on page 221.

_Characters for ready recognition._--Differs from _M. f. peninsulae_ in finer, softer pelage and shorter (less than 15.8 in ad. [F]) tympanic bullae; from _M. f. noveboracensis_, in adult males by wider tympanic bulla which is more than rather than less than 8.5, in adult females by total length which is more than rather than less than 345, and by mastoid breadth which is more than rather than less than distance between articular faces of exoccipital condyle and glenoid fossa; from _M. f. arthuri_ in larger size (adult males with hind foot and basilar length each more than 45); depth of skull at anterior margin of basioccipital, ignoring sagittal crest, amounting to less than 63 per cent of mastoid breadth, and lesser convexity of dorsal outline of skull in longitudinal axis (See pls. 16-18).

_Description._--_Size._--Male and female: External measurements of adults are available as follows:

Length Length Catalogue Sex Locality Total of of hind no. length tail foot 47165 [M] Box Springs, 454 160 48 Talbot Co., Georgia

47166 [M] Box Springs, 435 147 47 Talbot Co., Georgia

47167 [M] Box Springs, 422 145 45 Talbot Co., Georgia

41023 [M] Thomas Co., 443 140 47 Georgia

41025 [M] Grady Co., 395 142 47 Georgia

223880 [M] Okefinokee Swamp, 416 145 49 Georgia

198 [M] Okefinokee Swamp, 425 140 48 Georgia

Average 7 [M] 427 146 47

49385 [F] Gainesville, 396 124 45* Alachua Co., Florida

41024 [F] Thomas Co., 380 125 41 Georgia

51527 [F] Talbot Co., 376 128 43 Georgia * [not typical]