Part 45

Comparison of the skull of the male with that of _M. f. nicaraguae_ has been made in the account of that subspecies. The skull of the male as compared with that of _M. f. tropicalis_ has shorter tympanic bullae, deeper braincase at anterior margin of basioccipital, lesser zygomatic and palatal breadth and smaller P4 and m1. The skull of the female is larger in every measurement taken except those reflecting width of the preorbital portion. This part is actually narrower but probably mainly because the females of _perda_ are younger than those of _tropicalis_. Features in which three skulls of subadults of _M. f. perda_ differ from the five adults of _M. f. goldmani_ and show no overlap are: lesser basilar length, lesser weight, greater relative length of upper tooth-rows, greater relative width of rostrum, greater relative length of rostrum, lesser mastoid and zygomatic breadths, lesser width, length and height of tympanic bullae; lesser outside length of P4 and greater relative depth of braincase at anterior margin of basioccipital and at posterior margin of M1. Features in which _perda_ averages less are: length of tooth-rows, interorbital breadth, orbitonasal length, relative zygomatic breadth, length of m1, outside and inside lengths of P4, width and length of M1, and depth of skull at posterior margin of M1. Features in which _perda_ averages more than _goldmani_ are: relative interorbital breadth, relative mastoid breadth and depth of skull at anterior margin of basioccipital. The length of the inner half of M1 averages the same. As compared with _goldmani_, the skull of the male of _perda_ is shorter, otherwise generally smaller, but relatively broader except across the zygomatic arches, and relatively deeper. The anterior margins of the tympanic bullae project slightly less from the braincase and the squamosals immediately in front of these bullae are slightly more convex ventrally.

_Remarks._--Described by Merriam in 1902 as a subspecies of _Putorius tropicalis_, the form _perda_ was regarded by Allen (1916:99) as not subspecifically distinct from _P. t. tropicalis_.

This is the eastern, lowland subspecies of the Tropical Life-zone, corresponding to _M. f. goldmani_ of the higher mountains just as _M. f. tropicalis_ corresponds to _M. f. frenata_ and _perotae_ of the high mountains and table land. The difference in size between _perda_ and _nicaraguae_ and between _perda_ and _tropicalis_ is slight. _M. f. perda_ is slightly less richly colored than _M. f. nicaraguae_ but has the color of the underparts more restricted and has a longer black tip on the tail. In these respects it is second only to _M. f. panamensis_ among Central American weasels. Evidence of intergradation with _goldmani_ is furnished by the specimens from Cobán, Guatemala, and the nearby locality San Cristóbal in Verapaz, Guatemala. Reduced size as compared with _goldmani_ suggests affinity with _perda_ but the greater width of the light-colored underparts, which averages 24 (extremes 18-32) per cent of the greatest width of the color of the upper parts, shows approach to _goldmani_. Farther north, in Chiapas, however, specimens of _perda_ from San Cristóbal and San Vicente are readily distinguishable from those of _goldmani_ taken a few miles away at Pinabete and near Teopisca. The latter two localities are, however, several thousand feet higher than San Cristóbal (Chiapas) and San Vicente.

Two of the nine skulls (only 3 adult) examined for malformation of the frontal sinuses reveal infestation by parasites.

_Specimens examined._--Total number, 18, listed by localities from north to south, and unless otherwise indicated in the United States National Museum.

=Veracruz.= Catemaco, 1.

=Tabasco.= Teapa, 2 (1[7]).

=Chiapas.= San Cristóbal, 1; San Vicente, 1; no locality more definite than state, 1.

=Yucatán.= Chichén-Itzá, 1[76].

=Guatemala=: Cobán, 2 (1[7], 1[4]); Finca la Providenci, S. Cristóbal, Verapaz, 3[76]; central Guatemala, 1; no locality more definite than Guatemala, 5 (2[7]).

=Mustela frenata nicaraguae= Allen

Long-tailed Weasel

Plates 1, 25, 26, 27 and 30

_Mustela tropicalis nicaraguae_ Allen, Bull. Amer. Mus. Nat. Hist., 35:100, April 28, 1916.

_Putorius tropicalis_, Allen, Bull. Amer. Mus. Nat. Hist., 24:661, 1908.

_Mustela frenata nicaraguae_, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

_Type._--Male, subadult, skull and skin; no. 30754, Amer. Mus. Nat. Hist., Matagalpa, Nicaragua; April 16, 1910; obtained by W. B. Richardson; original no., 712.

The skull (plates 25-27, 30) of the type specimen lacks the entire right zygomatic arch. Otherwise it is complete. The teeth all are present and unbroken. The skin is complete and unfaded but only partly stuffed.

_Range._--Honduras and Nicaragua. Altitudinal and zonal limits unknown. See figure 29 on page 221.

_Characters for ready recognition._--Differs from _M. f. costaricensis_ and _M. f. goldmani_ in shorter black tip of tail (not more than 35 per cent of length of tail) and lesser width (usually not more than 7 mm.) of tympanic bulla; from _M. f. perda_ in greater extent of color of underparts (22 or more per cent of greatest width of color of upper parts), shorter black tip on tail (not more than 35 per cent as long as tail) and narrower skull, the mastoid breadth in adult males being less than 23.9 and the zygomatic breadth less than 27.

_Description._--_Size._--Male: Average and extreme measurements of five subadults and one young (four from Matagalpa and one from San Rafel del Norte) are: Total length, 450 (420-480); length of tail, 178 (150-190); length of hind foot, 48 (46-50). Tail averages 65 (extremes 56-69) per cent as long as head and body. Length of hind foot (measurements from dried skins) more than basal length.

Female: Measurements unrecorded.

_Externals._--As described in _Mustela frenata goldmani_, except that hairiness of foot soles (between that shown in figures 20 and 21) is less, slightly less even than in _M. f. perda_.

_Color._--As described in _Mustela frenata goldmani_ except that: Back near (_n_) Argus Brown, or Carbon Brown, tone 4 of Oberthür and Dauthenay, pl. 342. Underparts Ochraceous-Buff. Least width of color of underparts, in four males, young, subadult and adult, 24 (extremes 22-26) per cent of greatest width of color of upper parts; the corresponding per cent in one female is 32; black tip of tail, in two subadult males, averaging 29 (extremes, 28-30) per cent of length of tail-vertebrae; corresponding per cent in one female, 36.

_Skull and teeth._--Male (based on type specimen, one adult topotype [?] and one subadult from San Rafel del Norte): See measurements and plates 25-27, 30. As described in _Mustela frenata perda_ except that: Weight, 4.2 grams (estimated for adults); basilar length 45.0 (44.8-45.5); interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of four to five upper incisors; length of tympanic bulla not less than length of lower molar and premolar tooth-row; anterior margin of masseteric fossa below anterior margin of m2 or posterior to that tooth.

Female: Skull unknown.

Comparison of the skull of the male with that of _M. f. costaricensis_ is made in the account of that subspecies. As compared with that of _M. f. perda_, which it most closely resembles, the skull of the male has a narrower, shorter rostrum, lesser interorbital breadth, lesser mastoid and zygomatic breadths and slightly shallower braincase, measured at anterior margin of basioccipital. The tympanic bullae are slightly less projected, at their anterior margins, from the braincase and the squamosal, directly anterior to each, is a little more convex ventrally. The skull of _M. f. nicaraguae_ is, then, slightly shorter than that of _M. f. perda_ and relatively narrower.

_Remarks._--When naming this form, Allen (1916:100) characterized it as "Similar to _M. tropicalis tropicalis_ but general coloration much darker and the white face markings somewhat reduced in area." In the sentence preceding the one quoted, _Putorius tropicalis perdus_ was placed as a synonym of _Putorius tropicalis tropicalis_. _M. f. nicaraguae_ and _M. f. perda_ are nearly alike in color and color pattern but differ in cranial characters. _M. f. perda_ and _M. f. tropicalis_ are widely different in color and more especially in color pattern but differ only slightly in cranial characters. The aggregate difference between _perda_ and _nicaraguae_ is less than that between _perda_ and _tropicalis_. All three are lowland forms and each is smaller than the adjacent highland forms, namely, _M. f. goldmani_, _macrophonius_, _perotae_ and _frenata_.

The weasels from Honduras definitely are not typical of _nicaraguae_ as it is known from the specimens from Nicaragua itself. The specimens from the state of Tegucigalpa, Honduras, are larger. Some are darker than topotypical _nicaraguae_. The dorsal outline of the skull is more nearly flat (less convex) in some. In these and several other differential features studied, the average of specimens from Tegucigalpa is intermediate toward _goldmani_, but everything considered the animals seem best placed with _nicaraguae_ rather than with _goldmani_ or _perda_, to which latter also, they show some resemblance. With better material from Nicaragua and additional specimens from Salvador (here referred to _goldmani_) a restudy of all the material now referred to the three races named would be profitable. Aims of this restudy might be to determine if a highland race additional to _goldmani_ should be recognized and if the lowland races _perda_ and _nicaraguae_ differ from one another in the way that the existing specimens indicate.

In the five males from Matagalpa, the narrow white band in front of each ear is confluent with the color of the underparts on one side only in one specimen and on both sides in two specimens. None of these bands is confluent with the white patch between the eyes. A dark spot at the angle of the mouth is present on one side in one specimen. The corresponding area is dark colored in all other specimens but not separated from the color of the upper parts. In the specimen from San Rafel del Norte the white bands are not confluent with the color of the underparts. The female from Mambacho has the mentioned bands confluent with the color of the underparts. This female approaches _M. f. costaricensis_ in the dark color of the upper parts but has more extensive white facial markings than some specimens from much farther north. Like a female seen of _M. f. costaricensis_, this one has a "frosted" nape but the white hairs on the back of the neck are less numerous than in the female of _M. f. costaricensis_.

_M. f. nicaraguae_ in typical form, then, is thought of as a small, lowland, tropical subspecies only slightly differentiated from _M. f. perda_. By reason of its intermediate characters, it constitutes a link between the lowland forms, and the larger animals called _M. f. goldmani_ and _M. f. costaricensis_.

None of the four skulls from Nicaragua shows signs of infestation of the frontal sinuses by parasites.

_Specimens examined._--Total number, 16, listed by localities from north to south. Specimens are in the American Museum of Natural History, unless otherwise indicated.

=Honduras=: Alto Cantoral, 2; Cerro Grande La Paz, 1. La Flor Archaga, 1[75]; Comayagüela, 1[75]; vicinity of Tegucigalpa, 2; no locality more definite than Honduras, 1[4].

=Nicaragua=: San Rafel del Norte, 1; Matagalpa, 6; Ma[o]mbacho, 1.

=Mustela frenata costaricensis= Goldman

Long-tailed Weasel

Plates 25, 26, 27, 28, 29 and 30

_Mustela costaricensis_ Goldman, Proc. Biol. Soc. Washington, 25:9, January 23, 1912.

_Mustela brasiliensis_, Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):374, 1874.

_Putorius (Gale) brasiliensis frenatus_, Coues, Fur-bearing animals, p. 142, 1877 (part).

_Putorius affinis_, Merriam, N. Amer. Fauna, 11:31, June 30, 1896 (part).

_Mustela affinis costaricensis_, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Lönnberg, Arkiv för Zool., 14(no. 4):16, 1921.

_Mustela frenata costaricensis_, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

_Type._--Male, young, skull and skin; no. 13770/37149, U. S. Nat. Mus.; San José, Costa Rica; obtained by C. H. Van Patten.

The skull (plates 28-30) is complete and unbroken. All teeth are present and unworn. The skin apparently has been remade. It lacks the distal two-thirds of the tail. The head is somewhat shrunken. The color is possibly faded but if so only to a slight degree. Otherwise, the skin is in good condition. The orange color of the underparts is so intense as to suggest that the full, adult pelage has not been acquired. No white markings are present on the face. There is no sex mark on the label attached to the skin but the size and proportions of the skull and the scrotal pouch on the skin prove that the specimen is a male. The presence of sutures on the dorsal face of the rostrum and the short, wide, and low sagittal crest show the specimen to be young.

_Range._--Costa Rica. Altitudinal and zonal range unknown. See figure 29 on page 221.

_Characters for ready recognition._--Differs from _M. f. panamensis_ in lighter color of upper parts (tone 2 rather than tone 4 of Reddish Black of Oberthür and Dauthenay, pl. 344) and longitudinally flat interorbital region of skull; from _M. f. nicaraguae_ in darker color of upper parts (of Oberthür and Dauthenay, tone 2 of pl. 344 rather than tone 4 of pl. 342) and greater width (more than 7) of tympanic bulla.

_Description._--_Size._--Male: No collector's measurements available of fully grown animals. Estimated measurements of adult males: Total length, 470; length of tail, 165; length of hind foot (taken from dried skins of 3 adults), 52 (50-52). Tail estimated to average 55 per cent as long as head and body. Length of hind foot more or less than (about equal to) basal length.

_Female_: A subadult or adult, from the Candelaria Mountains, and a subadult from Irazú, measure, respectively: Total length, 370, 385; length of tail, 130, 150; length of hind foot, 40, 31. Tail 59 per cent as long as head and body. Length of hind foot probably about equal to basal length.

The estimated differences in external measurements of the two sexes are: Total length, 92; length of tail, 25; length of hind foot, 16 (probably average difference is less).

_Externals._--As described in _M. f. panamensis_ (figure 21) except that foot soles are slightly more hairy.

_Color._--As described in _Mustela frenata panamensis_ except that: back is near Reddish Black, tone 2 of Oberthür and Dauthenay, pl. 344; chin, lips, and throat white or whitish; remainder of underparts near (_c_) Ochraceous-Buff; color of underparts rarely extending distally onto toes of forefeet. Least width of color of underparts, in eleven specimens, averaging 23 (10-36) per cent of greatest width of color of upper parts; black tip of tail, in six specimens, averaging 36 (31-38) per cent of length of tail-vertebrae.

_Skull and teeth._--Male (based on 2 adults, no. 3.2.1.6. from vicinity of San José and no. 11408, U. S. Nat. Mus., from "Costa Rica"): See measurements and plates 25-30; weight, 5.9 grams; basilar length 49 +; 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 in undiseased skulls less than length of upper premolars (less than distance between posterior borders of P2 and P4) and less than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more or less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum more or less (about equal to) 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 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; tympanic bulla longer or shorter than (about equal to) lower molar and premolar tooth-row and longer or shorter than (about equal to) rostrum; anterior margin of masseteric fossa directly below posterior border of m2.

Female: Skull of adult unknown.

Comparison of the skull of the male with that of _M. f. panamensis_ has been made in the account of that subspecies. As compared with that of _M. f. nicaraguae_ the skull of _M. f. costaricensis_ is heavier and in every measurement taken is larger. The skull is generally more massive and it follows that most measurements of depth and width are greater in relation to the basilar length as well as actually greater. The individual teeth are larger and the tympanic bullae wider and at their anterior ends are more projected from the braincase. Indeed the skull is more like that of _M. f. goldmani_ than like that of _M. f. nicaraguae_.

_Remarks._--The half dozen ill-prepared skins, with partial skulls inside, of this form in the United States National Museum long were referred either to _Mustela brasiliensis_ or _Mustela affinis_. It was not until 1912 when Goldman studied these specimens that the distinctive characters of the Costa Rican weasel were recognized and made the basis of the name _costaricensis_.

_M. f. costaricensis_ is well differentiated from _M. f. nicaraguae_ and _M. f. goldmani_ which occur to the northward and from _M. f. panamensis_ which occurs to the southward and is a large, heavy-skulled, dark-colored animal with white facial markings restricted or absent. In the type specimen and the female from the Candelaria Mountains the white facial markings are only narrow facial bars or a few white hairs, but in the young male from Cervantes there is a well developed bar 6 millimeters wide on each side of the face and a separate nasofrontal spot, 10 x 12 mm. The young female from Cachí has a V-shaped frontonasal spot, on the right side of the face a white bar 5 mm. wide and 17 mm. long connected with the color of the underparts, and on the left side a white spot in front of the ear and another between the ear and eye. White facial markings were not recorded in the other specimens. The color of the upper parts is only a little less dark than those of _M. f. panamensis_. Owing to the numerous white hairs on the dorsal side of the neck, the nape of the female from the Candelaria Mountains has a frosted appearance not present in other specimens.

_M. f. costaricensis_ is a large animal and among its geographic neighbors is approached in size only by a specimen of _panamensis_ from Boquete, Panamá. Also the young male from Cervantes suggests _panamensis_ in the less flattened interorbital region, but even so is more like _costaricensis_. The small size of two young males, one from Navarro and the other from the vicinity of San José, is suggestive of _M. f. nicaraguae_. However, the large size of most of the specimens and the configuration of the skull are more as in _M. f. goldmani_ than in _M. f. nicaraguae_ and thus suggest that the known specimens are of high mountain subspecies. The long black tip of the tail is another point of resemblance to _M. f. goldmani_, the high mountain subspecies to the north. Perhaps in the lowlands of Costa Rica, there are weasels of another subspecies.

Of the eight skulls examined for malformation of the frontal sinuses, each of the two adults and two subadults shows signs of having the frontal sinuses infested with parasites.

_Specimens examined._--Total number, 14, listed by localities from north to south.

=Costa Rica=: Irazú (Frasu or Irasu on label), 3000 M., 1[4]; Cervantes, 1[2]; San José, 1[91]; vicinity of San José, 2[7]; Azahar Cartago, 1[78]; Tucurrique, 1[7]; Cachí, 1[7]; El Muñco [= Muñeco?] (Río Nivarro [= Navarro?]), 4000 ft., 10 mi. S Cartago, Caribbean Slope, 1[76]; Navarro, 1[91]; Candelaria Mts., 1[75]; no locality more definite than Costa Rica, 3[91].

=Mustela frenata panamensis= Hall

Long-tailed Weasel

Plates 1, 25, 26, 27, 28, 29 and 30

_Mustela frenata panamensis_ Hall, Proc. Biol. Soc. Washington, 45:139, September 9, 1932; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

_Mustela brasiliensis_, Alston, Biol. Cent. Amer., Mammalia, p. 78, 1879.

_Putorius affinis_, Bangs, Bull. Mus. Comp. Zoöl., 39:49, April, 1902; Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.

_Mustela affinis_, Goldman, Proc. Biol. Soc. Washington, 25:10, January 23, 1912; Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.

_Mustela affinis costaricensis_, Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Goldman, Smithsonian Miscel. Col., 69 (no. 5): 161, 1920.

_Type._--Female, subadult, skull and skin; no. 170970, U. S. Nat. Mus., Biol. Surv. Coll.; Río Indio, Canal Zone, near Gatún, Panamá; February 17, 1911; obtained by E. A. Goldman; original no. 20897.

The skull is complete and unbroken. The left lower incisor is broken off but all the other teeth are present and entire. The skin is well made and seems to be in faded, worn, first, adult pelage.

_Range._--Sea level (type locality) to 5800 feet (Boquete, see Bangs [1902:49]); Upper Tropical and Lower Tropical life-zones of Panamá. See figure 29 on page 221.

_Characters for ready recognition._--Differs from both _M. f. meridana_ and _M. f. costaricensis_ in darker tone (tone 4 of Oberthür and Dauthenay, pl. 344) of color of upper parts and in convex dorsal outline of skull (Compare figures of mentioned subspecies on plates 25-27).

_Description._--_Size._--Male: Two adults from Boquete in the Museum of Comparative Zoölogy, nos. 10112 and 10113, measure, respectively, as follows: Total length, 480 and 400; length of tail, 170 and 143; length of hind foot, 52 and 43. Hind feet of two other adult males measure 46 on dried skins. Tail, in two specimens mentioned above, is 55 and 56 per cent as long as head and body. Length of hind foot, in each of three adults, slightly longer than basal length. Corresponding measurements of no. 178970 from Mt. Pirre are: 422, 164, 50. Tail 64 per cent (same per cent as in young male, no. 137514 from Boquete) as long as head and body, and hind foot longer than basal length.

Female: An adult and a young from Chiriquí, nos. 18434 and 18435 (Acad. Nat. Sci. Philadelphia), measure, respectively: Total length, 372, 389; length of tail, 138, 144; length of hind foot, 42, 41. The type specimen measures: Total length, 408; length of tail, 159; length of hind foot, 46.5. Tail 64 per cent as long as head and body, and hind foot longer than basal length.

The average differences in external measurements of the two sexes from the vicinity of Boquete are: Total length, 59; length of tail, 15; length of hind foot, 6.