Part 46

_Externals._--Longest facial vibrissae black and extending beyond posterior border of ear; carpal vibrissae wholly, or in part, black and extending as far as hypothenar pad; hairiness of foot-soles as shown in figure 21.

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

_Skull and teeth._--Male (based on three adults from Boquete): See measurements and plates 25-30; weight, 5 (4.5-5.4) grams; basilar length, 45.2 (42.8-48.3); zygomatic breadth more or less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth more than length of upper premolars and more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum approximately same (more or less than) length of tympanic bulla; least width of palate less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4-1/2 to 5-1/2 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or slightly less than (approximately equal to) length of lower molar and premolar tooth-row or length of rostrum; anterior margin of masseteric fossa directly below posterior fourth of m2.

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

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

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

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

_Remarks._--This subspecies had not been recognized by previous workers because specimens from Panamá were supposed to be _Mustela affinis_ Gray up until 1916, when Allen (1916:100) restricted the type locality of _M. affinis_ to Bogotá, Colombia. At that time Allen referred specimens from Panamá to _Mustela affinis costaricensis_, and Goldman (1920:161) followed Allen.

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

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

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

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

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

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

=Panamá=: Boquete, 10 (3[75], 1[8], 1[2], 3[4], 1[7]); Río Gariche [é], 5300 ft., 1[1]; Siola, 1[1]; Chiriquí, 1[7]; Río Indio, near Gatún, 1; Mt. Pirre, 3 (2[1]); Calovébora, 1[7] (locality not found, possibly misspelling of Calovébora); no locality more definite than Panamá, 1[4].

=Mustela frenata meridana= Hollister

Long-tailed Weasel

Plates 25, 26, 27, 37, 38 and 39

_Mustela meridana_ Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.

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

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

_Putorius macrurus_, Allen, Bull. Amer. Mus. Nat. Hist., 31:92, April 19, 1912.

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

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

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

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

The skull (plates 25 and 26) lacks the right exoccipital condyle and posterior half of the right zygomatic arch. The teeth all are present, unworn and entire. The skin is well made and complete.

_Range._--Near sea level (San Julián) to 8500 feet (Montes de Culata, Mérida, Venezuela), and 9000 feet (Santa Elena, Colombia). Temperate to Subtropical life-zones of Venezuela and northern and western Colombia. See figure 29 on page 221.

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

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

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

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

_Externals._--Longest facial vibrissae black (few rarely white) and extending beyond ear; carpal vibrissae colored like underparts or upper parts, and not extending beyond apical pad of fifth digit; hairiness of foot soles slightly greater than shown in figure 21.

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

_Skull and teeth._--Male (based on type specimen and seven topotypes, five adults and three subadults): See measurements and plates 25-27; weight, 4.1 (3.8-4.3) grams; basilar length, 43.6 (42.3-44.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; postorbital breadth greater than length of upper premolars or than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum greater than length of tympanic bulla; least width of palate greater than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 to 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla more or less than (approximately equal to) alveolar length of lower molar and premolar tooth-row and shorter than rostrum; anterior margin of masseteric fossa posterior to m2 and confined to posterior third (34 per cent average, 32 minimum, 37 maximum) of mandible.

Female (based on four adult topotypes): See measurements and plates 37-39; weight (no. 143665), 2.3 grams; basilar length 37.2 (36.3-38.2); zygomatic breadth more or less than distance between condylar foramen and M1 or than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth (sinuses badly infested with parasites) more than length of upper premolars or width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than length of P4; tympanic bulla as far posterior to foramen ovale as combined width of 4 to 5 upper incisors; height of tympanic bulla less than (one half to three fourths of) distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row.

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

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

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

_M. f. meridana_ so closely resembles _M. f. affinis_ that the writer has no quarrel with anyone who would synonymize _meridana_. However, as represented by topotypes, the two races unquestionably are, _on the average_, different, and specimens from the southeastern part of the range of _affinis_ probably are individually distinguishable from topotypes of _meridana_.

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

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

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

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

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

=Venezuela=: San Julián, 1[91]; Carácas, 2; Galipare, Cerro del Avila, 6500 feet, 1; San Esteban, 1[2]; Mérida, 45 (10[91], 14[2], 10[4], 2[60], 2[14], 1[78]); Páramo de Tama, 1[60].