Part 43

_Skull and teeth._--Male (based on a referred specimen from Cerro San Felipe which certainly is nontypical): See measurements. As described in _Mustela frenata frenata_ except that: Weight, 4.9 grams; basilar length, 49.2; postorbital breadth more than distance between posterior borders of P4 and P2; 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; zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla.

Female (based on type specimen, an adult): See measurements and plates 37-39. As described in _Mustela frenata frenata_ except that: Weight 3.4 grams; basilar length, 43.5; 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 width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla one-half to three-fifths 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 is 33 per cent lighter than that of the nontypical (and smaller than average) male from Cerro San Felipe.

Comparison of the skull with that of _M. f. tropicalis_ is made in the account of that subspecies. Compared with the skull of _M. f._ _macrophonius_, that of the female of _perotae_ is more flattened, has the longitudinal dorsal outline distinctly concave rather than flat just behind the postorbital processes, and much wider tympanic bullae. Accordingly, the basioccipital is slightly narrower in _perotae_. The more marked postorbital constriction of the type specimen of _perotae_ possibly is due to its relatively greater age. As compared with the skull of _M. f. leucoparia_, that of the female of _perotae_ has less inflated tympanic bullae, the height of each being half as great as distance from its anterior margin to foramen ovale, whereas, in _leucoparia_ (as represented by no. 26153) the two distances are equal. As compared with that of _M. f. frenata_, the skull of the female of _perotae_ differs mainly in the lesser inflation of the tympanic bullae and their relative position. The height of each bulla is in _perotae_ only half as much as, but in _frenata_ more than, the distance from its anterior margin to foramen ovale. The anterior margin of the bulla is much less projected from the floor of the braincase in _perotae_. The squamosal anterior to each bulla is convex ventrally in _perotae_ but flat or concave ventrally in _frenata_.

_Remarks._--The type specimen and a juvenal female from the town of Perote were taken in the spring of 1893 by E. W. Nelson. Of these two, the type specimen was mentioned and figured by Merriam (1898:30, fig. 16 [= fig. 15], pl. 3, fig. 2) as _Putorius frenatus_. The referred nontypical specimen from Cerro San Felipe, Oaxaca, was referred by Merriam (op. cit.:29) to _Putorius frenatus goldmani_ with the comment that it was intermediate ". . . both in coloration and cranial characters, between typical _frenatus_ and _goldmani_;. . . ." No other published references to this subspecies, or specimens of it, have been seen. In 1941 and 1942, W. B. Davis and associates took four specimens along the boundary between the states of Puebla and México.

Although the specimen from Cerro San Felipe, Oaxaca, is referred to _Mustela frenata perotae_, to the description of which it answers best, that specimen, on account of its structural characters and geographic position relative to adjacent races, is in reality an intergrade between several of the adjacent races. Some of its intermediate characters are pointed out in the discussion of _M. f. goldmani_. In the specimens from 45 and 55 kilometers ESE of México (city) the black color of the top of the head does not extend so far behind the ears as in the holotype of _M. f. perotae_ and in this feature the two specimens show intergradation between the two subspecies, _perotae_ and _frenata_.

The type specimen taken on May 26, is acquiring new hair on the belly and lower sides which appears to be the result of a normal molt.

As would be expected from its geographic position, _M. f. perotae_ resembles _M. f. frenata_ of northern México and the high mountain forms of southern México more than it does the lowland tropical forms. This is true as regards size of entire animal, proportions of its parts, and size, general angularity and major proportions of its skull. The marked postorbital constriction, convex supralacrymal face of rostrum, width of tympanic bullae and angularity of the braincase place it nearest _M. f. frenata_ as does also the color and color pattern. The ventrally convex squamosal anterior to each tympanic bulla and the slight degree of projection from the cranium of the anterior margin of each tympanic bulla are intermediate in degree between the condition in _M. f. macrophonius_ and that in _M. f. frenata_. Thus _M. f. perotae_ combines several characters of _M. f. frenata_ on the one hand with some of _M. f. macrophonius_ on the other and in some features, for instance in the size, shape and degree of inflation of the tympanic bullae, presents intermediate stages of development.

On the eastern plain below the high mountain, Cofre de Perote, there ranges the similarly colored, smaller, tropical weasel, _Mustela frenata tropicalis_. Between _M. f. perotae_ and _M. f. tropicalis_ there is marked differentiation in the skulls with much less differentiation in coloration. The differences in typical skulls of the two subspecies are so pronounced that one would, at first glance, hardly believe it possible for direct intergradation to occur between them on the sides of this mountain. Merriam (1896:30) thought that it did not. The two skulls figured by him (_op. cit._:31) are a topotype of _M. f. tropicalis_ from Jico and the one which now is the type specimen of _M. f. perotae_. They show the great difference in size and proportions and are females of comparable ages, not of different ages as I suspected before examining the skulls. However, despite this marked difference in the skulls, there is some, although not conclusive, evidence of intergradation furnished by a young female from Xuchil, Veracruz. This specimen is described in connection with _M. f. tropicalis_ (see p. 366).

None of the seven skulls shows marked deformity of the interorbital region, but two of the three adults appear to have had these parts infested with nematodes.

_Specimens examined._--Total number, 7, all from México, listed by localities from north to south. Specimens from Veracruz and Oaxaca in the United States National Museum; remainder in Texas Cooperative Research Collection.

=México=: Monte Río Frío, 45 Km. ESE México City, 1; 55 Km. ESE México City, 1; N slope Mt. Popocatépetl, 13,555 ft., 1.

=Puebla.= Río Otlati, 8700 ft., 1.

=Veracruz.= Cofre de Perote, 12,500 ft., 1; Perote, 1.

=Oaxaca.= Cerro San Felipe, 10,000 ft., 1.

=Mustela frenata goldmani= (Merriam)

Long-tailed Weasel

Plates 1, 24, 25, 26 and 30

_Putorius frenatus goldmani_ Merriam, N. Amer. Fauna, 11:28, June 30, 1896; Elliot, Proc. Biol. Soc. Washington, 18:236, December 9, 1905.

_Mustela frenata goldmani_, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

_Type._--Male, adult, skull and skin; no. 77519, U. S. Nat. Mus., Biol. Surv. Coll.; Pinabete, Chiapas, México; February 10, 1896; obtained by E. A. Goldman (on attached label collectors recorded as Nelson and Goldman); original no. 9279.

The skull (plates 24 and 30) has the rostrum badly injured. All the right, and part of the left nasal, the upper part of the right maxilla, the postorbital process and intervening area of frontals are missing. Each zygomatic arch is broken but the parts are present and attached to the skull. The frontal and interorbital regions are greatly malformed owing to parasites that infested the sinuses. Right I2 and I3, right and left i3, and the medial parts of the paraconid and protoconid of right m1 are missing. The light facial markings are less extensive than in any of the referred specimens. These markings consist of a separate spot between the eyes and a white line, confluent with the color of the underparts, on each side of the head, that extends from the base of the ear to above the eye. The dark color of the underparts is represented at the angles of the mouth by a spot on the left side and a similar dark area, confluent with the dark color of the face, on the right side. The large size, characters of the skull, and scrotal pouch on the skin prove the specimen to be a male as stated on the label.

_Range._--Two thousand five hundred feet (El Cipres, Guatemala) to 9500 feet (near Tecpám, Guatemala), Upper Tropical Life-zone of mountains and western coasts of southern México, Guatemala and Salvador. See figure 29 on page 221.

_Characters for ready recognition_ (characters based on males).--Differs from _M. f. nicaraguae_ and _M. f. perda_ by larger size (total length of adult males more than 489), least width of color of underparts not less than 26 per cent of greatest width of color of upper parts, weight of skull of adult male more than 5 grams; from _M. f. macrophonius_ by smaller size (total length of adult males less than 540), skull of male with basilar length less than 52.5 and weight less than 6 grams; from _M. f. perotae_ (typical specimens of same sex not available) by darker color of upper parts which are Argus Brown or darker rather than Brussels Brown; nonextension of color of underparts onto hind feet; from _M. f. leucoparia_ in least width of color of underparts not more than 37 per cent of greatest width of color of upper parts; color of underparts not extended onto hind feet; black tip of tail two-fifths rather than one-fourth as long as tail-vertebrae; height of tympanic bulla less than four-fifths distance from its anterior margin to foramen ovale.

_Description._--_Size._--Male: Four adults yield average and extreme measurements as follows: Total length, 508 (500-512); length of tail, 196 (185-207); length of hind foot, 55.5 (54-58). Tail averages 63 (59-67) per cent as long as head and body. Length of hind foot more than basal length.

Female: Typical specimen unknown.

_Externals._--Longest facial vibrissae black and reaching beyond ear; carpal vibrissae wholly or in part of same color as upper parts and reaching as far as hypothenar pad; hairiness of foot-soles distinctly less than that shown in figure 20 on page 60.

_Color._--Spot between eyes, band, confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear and posterior third of each upper lip, white; remainder of sides and top of head and neck posteriorly to or slightly behind shoulders, black; dark spots at angles of mouth usually absent; tip of tail black; remainder of upper parts Argus Brown or near (_n_) Argus Brown; chin, throat and breast white; remainder of underparts near (16' _c_) Ochraceous-Buff; color of underparts extending distally on posterior sides of forelegs onto medial toes and on hind legs to points between knees and heels. Least width of color of underparts, in five adult males, averaging 28 (extremes 26-33) per cent of greatest width of color of upper parts; black tip of tail, in four adult males, averaging 40 per cent of length of tail-vertebrae.

_Skull and teeth._--Male (based on five adults): See measurements and plates 24-26, 30; weight, 5.4 (5.3-5.5) grams; basilar length, 49.9 (49.6-51.3); zygomatic breadth (except in no. 12523 from Salvador) more than or equal to distance between condylar foramen and M1 or between anterior palatine foramen and anterior margin of tympanic bulla. Mastoid breadth less than postpalatal length; postorbital breadth more or less than length of upper premolars and greater than width of basioccipital measured from median margin of one foramen lacerum posterior to its opposite; interorbital breadth less than distance between foramen opticum and anterior margin of tympanic bulla; breadth of rostrum less than length of tympanic bulla; least width of palate more or less than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of five upper incisors; height of tympanic bulla 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 shorter than or equal to length of rostrum; anterior margin of masseteric fossa immediately behind m2.

Female: Typical skull unknown.

Comparison of male skull with that of _M. f. perda_ made in discussion of that form. Comparison with that of _M. f. nicaraguae_ shows similar differences, some of which are more pronounced. For example, squamosals anterior to tympanic bullae more convex ventrally and these bullae project less from braincase than in _M. f. perda_; thus the difference in these features is greater between _goldmani_ and _nicaraguae_ than between _goldmani_ and _perda_.

As compared with the skull of the male of _M. f. macrophonius_, each one of the skulls of the adult males of _M. f. goldmani_ is smaller in every measurement taken, with two exceptions. The width of the tympanic bullae was more in three specimens of _M. f. goldmani_ as was also the depth of the same in three specimens. Relative to the basilar length all but two of these measurements average less in _goldmani_; the exceptions are the zygomatic breadth and depth of the skull at the anterior margin of the tympanic bullae which average more. Relative to the basilar length, the orbitonasal length and depth of the skull at the posterior margin of M1 are less in each skull of _goldmani_. Thus, excepting the width and height of the tympanic bullae and the relative zygomatic breadth and relative depth of the braincase posteriorly, the skull of _goldmani_ is shorter and relatively as well as actually narrower and lighter throughout.

As compared with the skull of the male of _M. f. leucoparia_, that of _M. f. goldmani_ averages a trifle shorter and no skull of _goldmani_ equals that of _leucoparia_ in actual or relative zygomatic and mastoid breadths or length or height of tympanic bullae. In depth, the skull of _goldmani_ averages actually and relatively greater. Its teeth are smaller. The squamosal anterior to each tympanic bulla is convex ventrally whereas it is concave ventrally in _leucoparia_ as in _frenata_.

_Remarks._--When Merriam (1896:28) named this subspecies, he had only one specimen but he called attention to the more important diagnostic characters, which additional specimens show pertain to the race as a whole.

_M. f. goldmani_ in typical form occurs in high mountains of the Upper Tropical Life-zone and is most closely related to _M. f. frenata_ and _M. f. macrophonius_. The altitude at which the two specimens were taken, twenty miles southeast of Teopisca in Chiapas, is not known. Merriam (1896:28) states that the type specimen was obtained at "about 8200 feet." The specimen taken by Stirton in Salvador comes from 8000 feet and the one obtained by Barber in Guatemala from 9500 feet. The specimen from Dueñas, the skin alone of a young animal, is not instructive.

As regards size, _goldmani_ is larger than the immediately adjacent subspecies from the Lower Tropical Life-zone but is smaller than _M. f. leucoparia_ or _macrophonius_. As compared with _M. f. frenata_, _goldmani_ is longer, has an actually as well as relatively shorter tail, and a much longer hind foot.

The most outstanding difference in externals from _frenata_ is the naked foot soles.

Molting probably takes place twice each year although actual proof of this is lacking. In number 133254 from twenty miles southeast of Teopisca, taken on May 12, the molt is well advanced. Another specimen from the same place still retains the winter coat.

In color, _goldmani_ is much darker than _frenata_, has less extensive white facial markings, longer black tip on tail, more restricted color of underparts, and lacks the extension of color of the underparts onto the hind feet.

Of the adult males from the high mountains, the type specimen from Chiapas is lightest, and the one from Salvador is darkest. This progressively darker color to the southward probably is geographic variation.

In total length and relative and actual length of tail, the specimen from Salvador is the smallest of the five adult males from the higher mountains. In addition to its darker color and smaller size, no. 12523 from Salvador shows certain distinctive cranial characters. The zygomatic breadth is less than, rather than more than, or equal to, the distance between the condylar foramen and M1 or than that between the anterior palatine foramen and the anterior margin of the tympanic bulla. This difference appears to be correlated with geographic position, since no. 15953 from Guatemala has the three distances about equal and therefore is intermediate in this respect between the specimen from Salvador and those from Chiapas, in which the zygomatic breadth is greater than the other two measurements. Also in the greater depth of the skull and smaller size of the teeth, this specimen from Salvador approaches the subspecies of the Lower Tropical Life-zone. It has, however, the longest, highest and widest tympanic bullae of any of the five specimens. The amount of ventral convexity of the squamosal in front of each tympanic bulla appears not to be greater than in the other specimens.

As indicative of intergradation with _perotae_, _leucoparia_ and possibly _frenata_, there is the specimen from Cerro San Felipe, Oaxaca. The degree of restriction of the color of the underparts is intermediate between that of _goldmani_ and _leucoparia_. The same is true as regards the amount of projection from the braincase of the anterior margins of the tympanic bullae. The squamosal immediately anterior to each tympanic bulla is flat instead of ventrally convex as in _goldmani_ or ventrally concave as in _leucoparia_ and _frenata_. In accordance with the custom adopted in this paper of referring every specimen to some one subspecies, this specimen from Cerro San Felipe is referred to _Mustela frenata perotae_, to the description of which it most nearly answers.

Possibly _goldmani_, as here constituted, is a composite form. The specimens from the high mountains closely resemble one another. However, a specimen, no. 68541 from "Finca El Cipres," Guatemala, which place Mr. G. Goodwin tells me is at an elevation of 2500 feet, approximately 5 miles north of Retalhuleu, has a basilar length of 47.3 and is correspondingly small in other parts. This suggests the existence of a small, lowland race on the western side of the central divide corresponding to _perda_ and _tropicalis_ on the eastern side. From only a few miles away, at San Sebastian, there is available, the adult skull of a still smaller animal. This skull only, no. 41026, in the Berlin Zoological Museum, has a basilar length of 46.1, zygomatic breadth of 27.4, and other cranial measurements notably smaller than those of specimens from the high mountains. A skin-only, no. 12038, collection of Donald R. Dickey, from La Cebia, altitude 2150 feet, near the city of San Salvador, seemingly represents an animal smaller than typical _goldmani_. This specimen from La Cebia has the light color of the underparts extended distally on the hind legs to the tips of the toes as in _M. f. tropicalis_. However, the upper parts are darker and resemble those of _M. f. goldmani_. A fourth specimen from only 3500 feet elevation, on the south side of Volcano Tajumulco, Guatemala, no. 41768, Field Museum of Natural History, a subadult male, measures only 490 in total length and has the least color of the underparts so restricted as to amount to only 22 per cent of the greatest width of the color of the upper parts. Both these features are suggestive of the lowland races.

These four specimens indicate that the lowland population on the western side of the divide is smaller than the mountain population. The juvenile from Carolina and a young male from Finca Cipres, however, both closely resemble individuals of _goldmani_ from the higher mountains. All these animals here are referred to _goldmani_. More specimens may reveal an amount and a pattern of geographic variation in weasels of this region that will require application of another subspecific name.

The female, no. 68540, from Puebla agrees remarkably well with the skull of the female, no. 132528, of _macrophonius_. Differences displayed by the specimen from Puebla are its slightly narrower braincase and longer space between the foramen ovale and anterior end of the tympanic bulla. Considering the far eastern location of Puebla (just north of Río Motagua, at 89° W, according to a sketch map provided by Mr. G. G. Goodwin), this specimen might be expected to show some approach to the small lowland races. Actually, however, it displays the characters of _goldmani_ better than does the subadult female from Volcano San Lucas, which is nearer the metropolis of _goldmani_, and I assume at a higher elevation than Puebla.

Concerning this weasel Merriam (1896:29) says: "Mr. E. W. Nelson writes me that this fine weasel is found sparingly in the forest about Pinabete, Chiapas, at an altitude of 7000 to 8000 feet (2100 to 2500 meters). The type specimen was shot in the afternoon while hunting on a heavily wooded hill slope. It was heard making long, slow leaps over the dry, crisp leaves. Coming to a log, it stood up and rested its fore feet on the log, in which position it was shot by Mr. Goldman."

The specimen taken by R. A. Stirton in Salvador comes from an elevation of 8000 feet in the rain forest of the Upper Tropical Life-zone. Mr. Stirton tells me that one morning on visiting his traps set for small rodents, he found in one the partly eaten remains of a _Heteromys_. Leaving these remains as found he placed a steel trap beside them and on the following morning found the male weasel in the trap.

At least three of the ten specimens had the frontal sinuses infested with parasites.

_Specimens examined._--Total number, 15, listed by localities from north to south, and unless otherwise indicated in the American Museum of Natural History.

=México=: _Chiapas_: 20 mi. SE Teopisca, 2[91]; Pinabete, 1[91].

=Guatemala=: Puebla, 1; Finca Porvenir, 3500 ft., S slope Volcan Tajumulco, 1[60]; Sierra [=? Cerro] Santa Elena, 9500 ft. (near Tecpám), 1[60]; Carolina, 1; Volcano San Lucas, 1; "Finca El Cipres," 1; "Finca Cipres," 2500 ft., 1; Finca San Isidro, San Sebastión, Dept. Retalhuleu, 1[4]; Dueñas, 1[7]; no locality more definite than Guatemala, 1[7].

=El Salvador=: Los Esesmiles, 8000 ft., Chalatenango, 1[59]; La Cebia, 2150 ft., near San Salvador, 1[59].

=Mustela frenata macrophonius= (Elliot)

Long-tailed Weasel

Plates 24, 25, 26, 30, 37, 38 and 39