SYSTEMATIC ACCOUNTS OF SPECIES AND SUBSPECIES (continued 8)

Remarks.—When Barber and Cockerell named this subspecies in 1898, they had three specimens. Only two others are known to have been taken before this time. These are a skeleton, without corresponding skin, taken at Lozier, Texas, in 1890 by Wm. Lloyd, and no. 21779/36482, U. S. Nat. Mus., taken on April 6, 1893, by R. D. Lusk at Tombstone, Arizona. On the back of a label recently attached to the last mentioned specimen the name C. K. Worthen appears and probably signifies that the specimen was purchased from this dealer in vertebrate specimens.

M. f. neomexicana has a large geographic range. The old male from Liberal, Seward County, Kansas, extends the known range far to the northeast. Geographically, this occurrence is logical for the southwestern desertlike conditions extend to this part of Kansas. Probably the subspecies occurs in southeastern Colorado and in the panhandle of Oklahoma where conditions are similar. Bailey (1905:198) lists neomexicana as a member of the mammalian fauna of Texas. As stated by him (loc. cit.:198) this inclusion is based on geographic grounds and not on actual specimens. Strecker (1926:13) also includes neomexicana in his list of Texas mammals but writes me, under date of January 9, 1928, that "I included Mustela frenata neomexicana as a Texas mammal on the strength of its being mentioned by Bailey. . . ." On better ground, Bailey (1928:97) lists the subspecies as occurring in southeastern New Mexico at Carlsbad Cavern. However, Bailey (loc. cit.) knew of the existence of weasels just below El Paso and at Langtry, Texas. An unsexed skeleton, no. 167891, in the United States National Museum, from Lozier, Texas, is not certainly identifiable to subspecies. If, as I think, the animal is a female, its skull is intermediate between that of frenata and neomexicana although when all features are considered it is seen to be nearest the latter. The large size (basilar length of 46.5 mm.) may reflect some relationship to texensis. The field notes of the collector furnished me by Dr. H. H. T. Jackson (MS), describe the color as brownish yellow above and sulphur below. The admission of this subspecies to the list of mammals of Texas is made certain by the female (no. 1572, Texas Cooperative Research Collection) taken on July 28, 1940, 1-1/2 mi. NW Kent, Texas, by C. E. Scull.

The skull alone from Durango (City of), extends the known range far to the south. This skull is typical of neomexicana. Skins from the same place would be especially interesting as showing the approach, if any, in color, of neomexicana to M. f. leucoparia.

Mr. D. D. Stone of Casa Grande, Arizona, writes, under date of February 2, 1927, that a weasel was seen by an acquaintance of his in a field near Chandler, Maricopa County, Arizona. Probably this was neomexicana. If so, its range extends much farther west than collected specimens show.

Actual intergradation with M. f. frenata is not shown by the material at hand. The two females from Albuquerque, although typically neomexicana as regards color, have smaller, less prominently ridged skulls than females of neomexicana of the same age from farther south and approach M. f. nevadensis.

Probably the geographic ranges of M. f. neomexicana and M. f. latirostra do not meet; the only evidence of the existence of weasels in all of the large area, comprising western Arizona and the deserts of eastern California, which intervenes between the ranges of the two subspecies is the skull of a young individual, no. 68842, Mus. Vert. Zoöl., from 10 miles northeast of Bard, Imperial County, California. There, on December 29, 1932, Jack C. vonBloeker, Jr., retrieved the weathered skull with some of the vertebrae attached, from the top of a wood rat's nest beneath a mesquite tree near the west bank of the Colorado River.

The idea that the carcass may have been washed down the river from far upstream gains no support from a comparison of the specimen itself for the tympanic bullae are larger than in nevadensis and the skull is larger than the largest males seen of arizonensis, the two upriver races. On the basis of size the skull could be either a male of latirostra or a female of neomexicana. These two subspecies, like arizonensis and the skull in question, have much inflated bullae. However, the immaturity of the specimen conceals any other diagnostic cranial features, and prevents referring it certainly to either neomexicana or latirostra. In any event the specimen provides no evidence of intergradation between the two forms last mentioned. Speculating on its identity, I should say that it might be either an intergrade between arizonensis and nevadensis, from southern Utah or northwestern Arizona, or a member of an unnamed race resident in the lower part of the valley of the Colorado River.

Whereas M. f. panamensis and M. f. aureoventris are the darkest-colored weasels and occur in regions of heavy rainfall, M. f. neomexicana is the lightest-colored American weasel and occurs in an extremely arid region where the rainfall and humidity are slight.

According to Barber and Cockerell (1898:189) "The type specimen was shot in the grass on the shore of Armstrong's Lake. . . ." Bailey (1928:97) found the tracks of one of these animals "in the great pit at the west entrance to" Carlsbad Cavern and supposes they "hunt the cave walls for mice and other small game." Data on the label attached to no. 230973 states that the specimen was taken, two miles west of Willcox, Arizona, in a prairie dog town.

Only two of the 23 skulls show evidence of infestation of the frontal sinuses by parasites.

Specimens examined.—Total number, 28, arranged alphabetically by states and from north to south by counties in each state. Unless otherwise indicated specimens are in the United States National Museum.

Arizona. Graham County: Safford, 1. Cochise County: 2 mi. W Willcox, 1; Willcox, 1; 8000 ft., Chiricahua Mts., 1; 6000 ft., Pinery Canyon, Chiricahua Mts., 1[33]; Tombstone, 1; Sulphur Spring Valley, 1[74].

Durango. "Durango City," 1.

Kansas. Seward County: Liberal, 1[93].

New Mexico. Bernalillo County: 3 mi. NW Albuquerque, 2. Lincoln County: 7800 ft., South Fork Eagle Creek, White Mts., 1. Chaves County: Pecos River, 10 mi. E Roswell, 8[74]; Dexter, 1[74]. Otero County: Cloudcroft, 9000 ft., 1[90]. Dona Ana County: Mesilla Park, 2 (1[75], 1[7]); Berino, 2.

Texas. Culberson County: 1-1/2 mi. NW Kent, 1[90]. Terrel County: Lozier, 1.

Mustela frenata texensis Hall

Long-tailed Weasel

Plates 22, 23 and 24

Mustela frenata texensis Hall, Carnegie Instit. Washington Publ. 473:99, November 20, 1936.

Mustela frenata, Strecker, The Baylor Bull., 27:14, September, 1924.

Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926 (part).

Type.—Male, adult, skull with skin of head, neck and tail; no. 14821, Amer. Mus. Nat. Hist.; Kerr County, Texas; September 17, 1897; obtained by H. P. Attwater.

The skull (plates 22-24) and dentition are complete and unbroken. The preserved parts of the skin are not stuffed.

Range.—Lower Sonoran and possibly Upper Sonoran life-zones of central Texas. See figure 29 on page 221.

Characters for ready recognition.—Differs from Mustela frenata arthuri in possessing white facial markings and postorbital breadth less than distance between posterior borders of P4 and P2; from M. f. frenata in larger size of body and skull, the basilar length of which in adult males is more than 52.5; from M. f. neomexicana in Brussels Brown rather than Buckthorn Brown color of upper parts and basilar length of skull more than 52.5.

Description.—Size.—Male: Measurements taken from the dried skins of a young male, no. 15476, Mus. Comp. Zoöl., from Kerr County, Texas, and a subadult male, no. 2017, Baylor Univ. Mus., from 5 mi. N Waco, Texas, are, respectively, as follows: Total length, 600 and more than 510; length of tail, 200 and 225; length of hind foot, 52 and 52.

Female: Skins unknown.

Externals.—As described in Mustela frenata frenata.

Color.—As described in Mustela frenata frenata.

Skull and teeth.—Adult male: See measurements and plates 22-24. As described in Mustela frenata frenata except that: Weight, 8 grams; basilar length 54; least width of palate less than length of P4; anterior margin of masseteric fossa anterior to middle of m2.

Female: Skull unknown.

Remarks.—The type specimen, taken by the veteran collector of Texan mammals, H. P. Attwater, appears to have been the first one of these animals to find its way into the collection of any museum or naturalist. The second specimen from Kerr County was secured by, or through, the well-known commercial collector, F. B. Armstrong. Two trade skins, from Kerr County, taken on December 10, 1938, are in the Texas Cooperative Research Collection, as is also the skeleton of a young animal from Fredericksburg. The two other specimens from McLennan County (both males contrary to the statement of Strecker, 1924:14), owe their preservation to the alertness of John K. Strecker, Curator of the Baylor University Museum, who has given a complete account of their history.

The range of this subspecies is thought to include much of central Texas.

The preserved parts of the skin of the type specimen show it to have been generally large. The part of the tail preserved measures 226 millimeters and the skin of the head and neck is correspondingly large. The skin alone, no. 427, from near Waco, Texas, has, as now stuffed, a body 365 millimeters long. Individuals of this race attain larger size than those of any other American member of the subgenus Mustela with the possible exception of Mustela frenata macrophonius from Veracruz, México. In addition to large size, texensis and macrophonius are analogous in that each has a small geographic range at the northern end of an extensive range of its similarly colored southern relative from which it differs mainly in size. Each of the two groups, goldmani and macrophonius on the one hand and perotae, frenata and texensis on the other, has relatively uniform color, color pattern and body proportions over a large region but at its northern extremity develops a "giant" population, M. f. macrophonius and M. f. texensis, respectively. The skull of the type specimen of M. f. texensis is the largest one seen of any American weasel. The type specimen of M. f. macrophonius has a basilar length that is greater by one-tenth of a millimeter but in every other measurement taken the skull of M. f. texensis is the larger. Its weight, 8 grams, also shows it to be larger.

The broad, white bands in front of the ears are confluent with the white patch between the eyes on both sides in two specimens and on one side only in one other specimen. A white patch between the ears is present in four specimens. The dark spot at each angle of the mouth is absent on both sides in four specimens and on one side only in one other specimen. Thus out of a possible twelve cases, the broad bands in front of the ears are confluent with the spot between the eyes in five cases. Four of the six specimens have a white spot between the ears. The dark spot at each angle of the mouth is present three out of a possible twelve times.

The skull of no. 2017, from five miles north of Waco, is smaller than either of the two skulls seen from Kerr County and in this respect approaches M. f. frenata. There is no actual evidence of intergradation with any other subspecies but intergradation probably does take place with M. f. neomexicana and possibly with M. f. arthuri and M. f. primulina.

Strecker (1924:14) remarks that of the two specimens obtained near Waco, one was taken in a trap baited for mink and the other was shot in a hen house. None of the four skulls had the frontal sinuses infested with parasites.

Specimens examined.—Total number, 7, arranged by counties from north to south.

Texas. McLennan County: 5 mi. N Waco, 1[3]; Erath, 1[3]. Gillespie County: Fredericksburg, 1[90]. Kerr County: 4[75]; 1[2]; and 2[90] trade skins.

Mustela frenata frenata Lichtenstein

Long-tailed Weasel

Plates 1, 22, 23, 24, 36, 37, 38 and 40

Mustela frenata Lichtenstein, Darstellung neuer oder wenig bekannter Säugethier, 1832, pl. 42, and corresponding text, unpaged; Seton, Lives of game animals, 2:576, 1929.

Mustela brasiliensis Sevastianoff, Mem. de l'Acad. Imp. Sci. St. Petersburg, 4:356-363, tab. 4, 1813, name on plate only, the description being in the text (not of Gmelin, 1788); Gray, Proc. Zoöl. Soc. London, 1865:114.

Putorius frenatus, Baird Mamms. N. Amer., p. 173, 1858; Merriam, N. Amer. Fauna, 11:26, pl. 3, figs. 1, 1a, 1b, June 30, 1896; Bailey, N. Amer. Fauna, 25:198, October 24, 1905.

Putorius (Gale) brasiliensis aequatorialis Coues, Fur-bearing animals, p. 142, 1877, part? ("merely as a substitute for Gray's [supposedly] preoccupied name" that is, aureoventris).

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Putorius mexicanus Coues, Fur-bearing animals, p. 142, 1877, [nomen nudum, cited by Coues in synonymy as "Putorius mexicanus, Berlandier, MSS. ic. ined. 4 (Tamaulipas and Matamoros)"].

Putorius brasiliensis frenata, Allen, Bull. Amer. Mus. Nat. Hist., 3:219, April 17, 1891.

Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 6:197, May 31, 1894; Bangs, Proc. Biol. Soc. Washington, 10:9, February 25, 1896; Allen, Bull. Amer. Mus. Nat. Hist., 8:74, April 22, 1896.

Mustela frenata frenata, Strecker, The Baylor Bull., 27:12, August, 1926; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.

Type.—Female, adult, skull and skin; no. 991, Berlin Zool. Mus., México City, México; June, 1829; obtained by F. Deppe.

The specimen once mounted, now is remade into a study skin and lacks the distal part of the tail. The skull (plates 36-38, 40) lacks the basicranial region.

Range.—Altitudinally, sea level (Brownsville, Texas) to 7600 feet (Tlalpam, México); from southern Texas as far south as México City; Lower Sonoran to at least Transition life-zone. See figure 29 on page 221.

Characters for ready recognition.—Differs from M. f. perotae in nonextension of blackish over anterior fourth of neck, least width of color of underparts more than 37 per cent of greatest width of color of upper parts; height of tympanic bulla more than distance from its anterior margin to foramen ovale; from M. f. leucoparia by restricted white facial markings that cover less than half surface of head in front of ears, by nonextension of black of head onto anterior half of neck and by wider (more than 7.8) tympanic bullae; from M. f. neomexicana by Brussels Brown rather than Buckthorn Brown color of upper parts and mastoid breadth less than postpalatal length; from M. f. texensis by smaller size of body and skull (basilar length in adult males less than 52.5); from M. f. arthuri by white facial markings and postorbital breadth less than distance between posterior borders of P4 and P2; from M. f. tropicalis by nonextension of blackish over anterior fourth of neck, least width of underparts more than 37 per cent of greatest width of upper parts, postorbital breadth of adult males less than distance between posterior borders of P4 and P2.

Description.—Size.—Male: Fifteen adults and subadults from Brownsville, Texas, yield average and extreme measurements as follows: Total length, 485 (430-556); length of tail, 202 (165-250); length of hind foot, 48 (40-55). Averages believed to be reliable but extremes probably are not. Tail averages 71 per cent as long as head and body. Length of hind foot less than basal length. Corresponding measurements of an adult male (topotype, no. 50826) from Tlalpam, México, are: 505, 203, 53. Another adult male, from Miquihuana, Tamaulipas, México, measures: 520, 215, 52.

Female: Six adults, subadults and young from Brownsville, Texas, yield average and extreme measurements as follows: Total length, 420 (362-456); length of tail, 173 (126-200); length of hind foot, 41 (40-46). Tail averages 70 per cent as long as head and body. Length of hind foot more (with possible exception of no. 36362/48732 U. S. Nat. Mus.) than basal length.

The average differences in external measurements of the two sexes are: Total length, 65; length of tail, 29; length of hind foot, 7.

Externals.—Longest facial vibrissae black and reaching beyond ear; carpal vibrissae same color as underparts and extending to apical pad of fifth digit; hairiness of foot-soles as shown in figure 20.

Color.—Spot between eyes, broad band, confluent with color of underparts, on each side of head extending anterodorsally anterior to each ear, and posterior two-thirds to one-half of each upper lip, white; remainder of sides and top of head, posteriorly to line connecting posterior margins of ears, blackish; dark spot posterior to angle of mouth present on both sides in about half the specimens; tip of tail black; remainder of upper parts Brussels Brown; chin white; remainder of underparts near (16´a) Ochraceous-Buff (same color in juveniles and young), which color extends distally on posterior sides of forelegs over forefeet and on medial sides of hind legs to feet and sometimes onto upper sides of toes. Least width of color of underparts averaging, in a series of seventeen males from Brownsville, Texas, 47 (extremes 38-53) per cent of greatest width of color of upper parts. Black tip of tail, in same series, averaging 49 (extremes 40-55) mm. long, thus about equal to length of hind foot and averaging 24 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on ten adults from Brownsville): See measurements and plates 22-24; weight (three adults, one topotype and two from Brownsville, Texas), 6.2 (5.3-7.2) grams; basilar length, 49.8 (48.2-51.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; mastoidal breadth less than postpalatal length; postorbital breadth less than length of upper premolars (less than distance between posterior borders of P4 and P2) and not greater (usually less) than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth not greater 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 3 or 4 (including I3) upper incisors; height of tympanic bulla more 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 or shorter (usually longer) than rostrum; anterior margin of masseteric fossa just behind m2.

Female (based on two adults from Brownsville, Texas): See measurements and plates 36-38, 40; weight, 3.4 (3.3-3.5) grams; basilar length (six, adult to young) 43.3 (41.3-47.3); zygomatic breadth more or less than distance between condylar foramen and M1 and more than distance between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth less than length of upper premolars and more or less than (about equal to) width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate less than outside length of P4; tympanic bulla as far posterior to foramen ovale as width of 2 to 3-1/2 (including I3) upper incisors; height of tympanic bulla more 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 or shorter than rostrum.

The skull of the female averages 45 per cent lighter than that of the average male.

Comparison of the skull with those of M. f. arthuri, tropicalis, perotae, leucoparia and neomexicana has been made in accounts of those subspecies. As compared with M. f. texensis (known only from males), the only difference detected is smaller size.

Remarks.—As Merriam (1896:27) has said: "In 1813 a Russian naturalist, Sevastianoff, gave the name 'Mustela brasiliensis' to a weasel brought to St. Petersburg by Capt. A. J. Krusenstern on his return from a voyage around the world. The animal was said to have come from Brazil, but no definite locality was given." This name was long applied by many European naturalists to American weasels which had white facial markings, and several American naturalists adopted the name. However, Lichtenstein in 1832 applied the name Mustela frenata to the weasels of the vicinity of México City and that name was used for bridled weasels from México and the southwestern United States by most subsequent German writers and by several Americans. In 1896 Merriam (1896:27) showed that Sevastianoff's Mustela brasiliensis, 1813, although probably the same as Mustela frenata, was preoccupied by Gmelin's Mustela brasiliensis, 1788, applied to an otter and that Lichtenstein's name must be used as the next available one. Since that time, 1896, frenata has been the name applied to the large bridled-weasels of Texas and the high table land of México south to México City. It may be added that in 1937 search by the writer among the specimens and records at the Russian Academy of Sciences, in Leningrad, failed to reveal any trace of the type specimen of Sevastianoff's Mustela brasiliensis.

The geographic range of this subspecies is relatively large and, as might therefore be expected, specimens show geographic variation. The specimens from Tlalpam, which Merriam (op. cit.:27) regards as topotypes, differ in certain respects from specimens from Texas. The skull of the adult male "topotype," no. 50826, differs from any other adult male seen in that the basilar length, the length of the upper tooth-rows, the orbitonasal length, the ratio of the same to the basilar length, the mastoidal breadth, the zygomatic breadth, the depth of the skull at the posterior margins of the upper molars, and the length and breadth of M1, are greater. The height of the tympanic bullae is less than the average height for these structures in more northern specimens. The specimens from Tlalpam have also larger external measurements than the average of more northern specimens. All of these features show an approach to the subspecies of more southern distribution. On the other hand, the blackish of the head is not more intense or more extended posteriorly onto the neck than in specimens from Brownsville, Texas. The skin, with skull crushed, no. 767, in the Paris Museum, from 3200 meters elevation near Toluca, does have the black color of the head extended 30 millimeters posteriorly to the ears. In this feature, and also in the extensively white face on which the white bar in front of each ear connects with the frontal spot, as well as with the color of the underparts, the specimen resembles leucoparia. Better material from the western part of the state of México may show the range of leucoparia to extend eastward almost or quite to Toluca.

An adult male, taken on July 15 at Miquihuana, Tamaulipas, is unique in several respects. The top of its head is black, rather than blackish, and this color extends posteriorly on the top and sides of the neck almost halfway to the shoulders. All of the upper parts are much more darkly colored than in other specimens of this race. The least width of the color of the underparts is 63 per cent of the greatest width of the color of the upper parts; thus the color of the underparts is considerably more extensive than in any other specimen seen. The underparts are more intensely colored than in the average specimen. The mastoidal breadth is greater than in any other adult male and amounts to more than the postpalatal length. On available maps the elevation of Miquihuana is given as 1892 meters (about 6200 feet). Thus the dark colors can hardly be ascribed to more tropical conditions than those under which animals from Brownsville, Texas, live. Brownsville is only a few feet above sea level and only 235 miles farther north. The difference noted, therefore, seems to be of geographic significance. However, there is from Alvarez, San Luis Potosí, approximately 115 miles south of Miquihuana, a young (nearly subadult) female, no. 21968, which is as light colored as specimens from Brownsville, Texas, or Tlalpam, México. The only distinctive feature of this specimen is the much greater extent of its white facial markings; they are more extensive even than in the specimen from Miquihuana.

Finally, the series from Brownsville, Texas, indicates that the animal there is smaller than frenata from the vicinity of México (city). The skull is similarly proportioned except that relative to the basilar length the orbitonasal length is more. Several other measurements of the skull of the adult male from Tlalpam, as pointed out above, are actually, although not relatively, greater than in any specimen from Brownsville. The similarities between specimens from the two localities, Tlalpam and Brownsville, are striking; since the two localities lie at opposite, extreme ends of the range more geographic variation would be expected. All that is known of the characters of populations from intermediate localities is that the one specimen from Alvarez shows no peculiarities whereas the one from Miquihuana suggests the existence there of a geographic variant.

None of the specimens seen shows actual intergradation with M. f. neomexicana or with M. f. arthuri but it is supposed that frenata intergrades with each of these subspecies. The difference between frenata and arthuri is greater than between frenata and neomexicana. Bailey (1905:198) records tracks of a weasel seen just below El Paso which he supposed had been made by a weasel of the neomexicana type. He also cited the taking of a weasel at Langtry which suggested to him (op. cit.) ". . . a continuous range from the country of frenatus up the Rio Grande to the type locality of neomexicanus at Mesilla Valley," New Mexico. Other records of occurrence in Texas cited by Bailey, in addition to those provided by specimens examined by the writer, are San Diego, Beeville, and Port Lavaca. The Port Lavaca record is the easternmost one assigned to the subspecies frenata; possibly specimens from there would be referable to arthuri.

The series of thirty-four specimens from Brownsville, Texas, permits measuring the amount of individual and age variation in several features. For instance, the material is sufficient to show that external measurements of subadults and those that fall in the upper part of the category designated as "young" may be included with the measurements of adults, because the mentioned measurements are not appreciably greater in adults. The series of skulls, although not providing more than six of any one age, shows the range of variation in size and proportion of certain parts and enables the student the better to evaluate cranial characters of nearby races known from only a few specimens. For example, not one of the twenty skulls of males from Brownsville and immediate vicinity is as large as either of the two specimens of texensis from Kerr County.

The white facial markings vary much in size and shape. In the series of thirty-four skins from Brownsville the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in three specimens and on one side only in six other specimens. These bands are confluent with the color of the underparts in all but two specimens. In one specimen the connection is lacking on both sides and in the other on one side only. A white patch between the ears is present in two specimens. The dark spot at each angle of the mouth is absent on both sides in eleven specimens and absent on one side only in ten others.

In six other specimens from parts of Texas north of Brownsville, the broad white bands in front of the ears are confluent with the white patch between the eyes on both sides in one specimen. A white spot between the ears is present in one specimen. The dark spot at each angle of the mouth is absent on both sides in six specimens and on one side only in three other specimens.

In eleven specimens from México, the broad white bands in front of the ears are confluent with the white spot between the eyes on both sides in two specimens and on one side only in one other specimen. The white spot between the ears is present in one specimen. The dark spot at each angle of the mouth is absent on both sides, in six specimens, and on one side only in one other specimen.

Thus, in 51 specimens the broad bands (one in front of each ear) are confluent with the white patch between the eyes in nineteen out of 100 instances, and not with the color of the underparts in three instances. A white spot between the ears is present in four specimens. The dark spot at each angle of the mouth is present 47 out of a possible 98 times.

Four juvenal specimens from Brownsville, Texas, with their dates of capture and probable age, are as follows: no. 58574, ♀, three weeks old, taken on February 15; no. 17318/24239, ♂, four weeks old, taken on March 16; no. 45899, ♀, forty days old, taken on May 21; no. 21778/36481, ♂, thirty days old, taken on October 20. In the order given, the dates of birth of these four juveniles would be approximately as follows: January 25, February 15, April 1, and September 20. The dates of birth of other specimens less than three months old as judged by the stage of development of the skull, and reckoning backward from the dates of capture, are as follows: April 1, April 30, May 25, October 12, and December 21. Thus, young appear to be brought forth at Brownsville, Texas, in the fall, winter and spring, that is to say from the latter part of September until the latter part of May.

Mustela frenata frenata is either free of the parasites that infest the frontal sinuses of most weasels, or withstands their presence remarkably well, for only one skull shows a definite pathological condition of the frontal sinuses.

Allen (1896:74) quotes H. P. Attwater, with respect to this species in Bexar County, Texas, as follows: "Not common, but occasionally met within the chaparral and cactus lands, where Wood Rats, Rabbits and Quail abound. They were frequently met with around San Antonio during the great 'Tramp Rat' [= Sigmodon hispidus texianus, see Bailey (1905:116)] invasion of 1889-90."

Specimens examined.—Total number, 63, arranged by counties, and in México by states, from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.

Texas. Bexar County: San Antonio, 2 (1[2]). Goliad County: Charco, 1. Nueces County: Corpus Christi, 1[2]. San Diego County (not found), 1. Hidalgo County: La Hacienda, 1. Duval County: San Diego, 2[7]. County in question: Lower Rio Grande, 1. Cameron County: Brownsville, 34 (3[2], 4[1], 3[93], 2[75], 1[59], 1[60], 1[4]); no locality more definite than county, 2.

Nuevo León. Río Ramis, 20 mi. NW Montemorelos, 1[90].

Tamaulipas. Matamoros, 6; Miquihuana (now in Nuevo León), 1[75].

San Luis Potosí. Alvarez, 1[75].

México: Region montagneuse des environs de Toluca, Nevada Toluca, 3200 M., 1[84]

Distrito Federal. City of México, 2 (1[4]); Tlalpam, 2. No locality more definite than México, 4 (1[4], 3[7]).

Mustela frenata leucoparia (Merriam)

Long-tailed Weasel

Plates 1, 24, 25, 26, 29, 30, 36, 37 and 38

Putorius frenatus leucoparia Merriam, N. Amer. Fauna, 11:28, June 30, 1896.

Putorius brasiliensis frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 2:165, October 21, 1889.

Putorius frenatus frenatus, Allen, Bull. Amer. Mus. Nat. Hist., 22:259, July 25, 1906.

Mustela frenata leucoparia, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Hall, Carnegie Instit. Washington Publ. 473:108, November 20, 1936.

Type.—Male, adult, skull and skin; no. 34914/47179, U. S. Nat. Mus., Biol. Surv. Coll.; Pátzcuaro, Michoacán, México; July 27, 1892; obtained by E. W. Nelson; original no. 2960.

The skull (plates 29 and 30) lacks most of the braincase; a fragment, consisting of the supraoccipital and the coalesced frontals and parietals remains. The rostrum, left zygomatic arch, palate, left pterygoid, left glenoid fossa and right postorbital process are intact. The teeth all are present and entire. The lower jaw lacks the right coronoid process and the lateral part of the articular condyle. The skin is well made and in good condition. It differs from an adult male topotype (36855, U. S. Nat. Mus.) and other referred specimens in having: the black of the head extended farther posteriorly on the neck, the maximum amount of white on the head, and a white stripe 50 mm. long extending down the middle of the nape from a point between the ears more than half way to the shoulders.

Range.—Sonoran and Transition life-zones of mountains west of México (city) in Michoacán and Nayarit. See figure 29 on page 221.

Characters for ready recognition.—Differs from M. f. goldmani in least width of color of underparts more than 47 per cent of greatest width of color of upper parts, hind feet colored like underparts rather than like upper parts; postorbital constriction less than, rather than more than, combined length of upper premolars; from M. f. macrophonius by same details of coloration as from goldmani and by ventrally concave rather than ventrally convex pretympanic part of squamosal; from M. f. perotae by least width of color of underparts more than 40 per cent of greatest width of color up upper parts; height of tympanic bulla more than three-fifths distance from its anterior margin to foramen ovale; from M. f. frenata by white facial markings that cover half of surface of head in front of ears, by extension of black of head onto neck halfway to shoulders and by narrower (less than 7.8) tympanic bullae; from M. f. neomexicana by Argus Brown rather than Buckthorn Brown color of upper parts and distance from anterior margin of tympanic bulla to foramen ovale more, rather than less, than four-fifths of height of tympanic bulla.

Description.—Size.—Male: Two adults and one young from Los Reyes and Pátzcuaro, Michoacán, yield average and extreme measurements as follows: Total length, 514 (510-523); length of tail, 206 (196-215); length of hind foot, 55 (52-58). Tail averages 67 per cent as long as head and body. Length of hind foot more than basal length.

Female: One adult from Artenkiki, Jalisco, and one subadult from Pátzcuaro, Michoacán, measure, respectively, as follows: Total length, 412, 400; length of tail, 159, 159; length of hind foot, 41, 42. Tail averages 64 per cent as long as head and body. Length of hind foot equal to or greater than basal length.

The average differences in external measurements of the two sexes are: Total length, 108; length of tail, 47; length of hind foot, 13.

Mustela frenata leucoparia has a greater total length and length of tail than either M. f. frenata or goldmani. The hind foot is longer than that of frenata and approximately the same as in goldmani. Relative to the body length, the tail averages longer than that of goldmani and shorter than that of frenata.

Externals.—As described in Mustela frenata frenata.

Color.—Broad white bands on sides of head, extending anterodorsally anterior to each ear, confluent with white spot between eyes and with color of underparts; posterior third of each upper lip white; remainder of sides and top of head, and neck posteriorly to point halfway to shoulders from ears, black; no dark spots at angles of mouth; tip of tail black; remainder of upper parts Argus Brown; chin white and sometimes also chest, neck and medial sides of hind legs; remainder of underparts near (16´) Ochraceous-Buff (near (a) Ochraceous-Buff in juvenal female), which color extends distally over all of each foreleg (except its lateral face proximally from about middle of forearm) and on medial side of hind leg and over most of upper side of each foot. Least width of color of underparts averaging, in eight specimens, 54 (extremes 44-61) per cent of greatest width of color of upper parts; black tip of tail averaging, in four males, 52 (extremes 38-78) mm. long, thus averaging 25 per cent of length of tail-vertebrae.

As compared with M. f. frenata and goldmani: white facial markings more extensive; color of underparts less restricted and more extended on legs; black tip of tail relatively of about same extent as in frenata and thus much less than in goldmani; black color of head extending farther posteriorly than in frenata but not so far as in goldmani.

Skull and teeth.—Male (adult): See measurements and plates 24-26, 29, 30. As described in Mustela frenata frenata except that: Weight (no. 128972) 6.3 grams; basilar length, 51.2; interorbital breadth 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 4 or 5 upper incisors; height of tympanic bulla more or less than (about equal to) distance from its anterior margin to foramen ovale; anterior margin of masseteric fossa anywhere from slightly anterior, to slightly posterior, to m2.

Female (based on no. 26153): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 44.5; zygomatic breadth less than distance between condylar foramen and M1, or than between anterior palatine foramen and anterior margin of tympanic bulla; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla not more than distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row or than length of rostrum.

The skull of the female is 43 per cent lighter than that of the male.

Comparison of the skull with those of M. f. perotae, goldmani and neomexicana has been made in the accounts of those subspecies. As compared with that of frenata the main difference is the less inflated tympanic bulla, the height of which is approximately equal to, rather than decidedly more than, distance from its anterior margin to foramen ovale.

Remarks.—The first specimen known to have been preserved is the alcoholic in the British Museum of Natural History, taken in September, 1891, on the Río Santiago in Jalisco, by D. A. C. Buller. The other known specimens of this white-faced weasel are divided between the American Museum and the United States National Museum. The two referred specimens from Jalisco were the last of several helpful ones collected in México and Central America by J. H. Batty, and these two were taken less than three months before Batty's tragic death in Chiapas (see Allen, J. A., 1906:191). The five specimens from Michoacán were taken by Nelson or Nelson and Goldman together. Merriam had only three of these when he named the subspecies and remarked (1896:29) that "This form is the poorest subspecies described in the present paper." Although the form is not strongly marked, the two additional specimens from Michoacán and better comparative material than Merriam had confirm several of the differential characters ascribed to it by him and indicate the existence of still other characters.

M. f. leucoparia occurs in the Sonoran and Transition life-zones. No. 27258 from Los Masos, and no. 26153 from Artenkiki (see specimens examined for other spellings) approach true frenata in coloration. Each of these specimens has a few white hairs between the ears and the white patch between the eyes is confluent on one side only with the lateral white bands on the side of the head. No. 27258 from Los Masos has a dark spot at each angle of the mouth. The 7 other specimens are relatively uniform in coloration. Each has the white spot between the eyes confluent on both sides with the extensive white areas on each side of the face. None has a dark spot at either angle of the mouth. Of these 7 specimens, the type specimen and three others have white hairs forming a median line between the ears and a fifth specimen has a white spot behind each ear.

M. f. leucoparia is most like M. f. frenata. Unlike frenata, leucoparia has tympanic bullae that are less inflated, narrower and less projected, at their anterior margins, from the cranium. In these characters leucoparia is intermediate between M. f. frenata and M. f. goldmani. The latter subspecies has the least inflated, narrowest and least projecting tympanic bullae of the three. The black color of the head extends, on the average, farther posteriorly than in M. f. frenata but not so far as in M. f. goldmani. The general color, too, is intermediate between that of M. f. frenata and that of the much darker M. f. goldmani. The white facial markings are more extensive than in either M. f. frenata or M. f. goldmani. This applies to both the white area between the eyes and the one on each side of the head between the ear and eye. M. f. neomexicana, whose range possibly meets that of M. f. leucoparia, also has more extensive white facial markings than M. f. frenata but less extensive markings than M. f. leucoparia.

On the basis of skulls alone, specimens of frenata from Tlalpam and those of leucoparia from Los Reyes can hardly be distinguished. This fact, and the circumstance that the specimens from the northern part of the range of leucoparia closely resemble frenata in color, constitute sufficient evidence for regarding the two as only subspecifically distinct. The female, no. 26153 from Artenkiki, as mentioned above, approaches true frenata in coloration. On this account it is not to be regarded as typical and it was because no other skulls of adult females were available that this one was used for comparison with females of allied races.

M. f. leucoparia is, then, a subspecies of the large, temperate-zone group and is unique in possessing the maximum extent of white facial markings.

None of the seven skulls shows signs of having had the frontal sinuses infested with parasites.

Specimens examined.—Total number, 8, all from México. Localities are listed by states from north to south. Specimens from Michoacán are in the United States National Museum; one from Río Santiago is in the British Museum of Natural History; all others are in the American Museum of Natural History.

Nayarit. Tepic, 1.

Jalisco. Río Santiago, 1; Los Masos, 1; "Artenkiki" (J. A. Allen, 1906, p. 238, writes "Artenkikil" and, on p. 259, "Artenkiki."), 1.

Michoacán. Zamora, 1; Los Reyes, 1; Pátzcuaro, 3.

Mustela frenata perotae Hall

Long-tailed Weasel

Plates 36, 37 and 38

Mustela frenata perotae Hall, Carnegie Instit. Washington Publ. 473:100, November 20, 1936.

Putorius frenatus, Merriam, N. Amer. Fauna, 11: pl. 3, fig. 2, June 30, 1896.

Type.—Female, adult, skull and skin; no. 54278, U. S. Nat. Mus., Biol. Surv. Coll.; 12,500 feet, Cofre de Perote, Veracruz, México; May 26, 1893; obtained by E. W. Nelson; original no. 4864.

The skull (plates, 37-39) lacks the right zygomatic arch. Left p2 is missing. The skin is fairly well made and in good condition except that the extreme tip of the tail has been broken off and there are two holes in the right hind leg. The underparts show the beginning of a spring molt.

Range.—From 7500 (?) feet (Perote) to 13,500 feet (Popocatépetl), Upper Sonoran, Transition and Boreal life-zones of mountains along Puebla-México boundary, eastward to western central Veracruz and south into Oaxaca. See figure 29 on page 221.

Characters for ready recognition.—Differs from M. f. frenata, its nearest relative, in extension from head of blackish onto anterior fourth of neck; restriction of color of underparts (least width of same less than 37 per cent of greatest width of color of upper parts), height of tympanic bulla less than distance from its anterior margin to foramen ovale; from M. f. macrophonius and M. f. goldmani in presence of, rather than absence of, color of underparts on hind feet; upper parts (black) Brussels Brown rather than Argus Brown or darker; from M. f. tropicalis in larger size (adult female with total length more than 400, basilar length more than 40, weight of skull more than 3 grams); postorbital breadth less than combined length of upper premolars; m1 more than 5.4 long; from M. f. leucoparia in white facial markings so restricted that spot between eyes is not confluent with white stripe in front of ear, or, if so, narrowly (less than 4 wide) confluent; color of upper parts extending onto antipalmar face of forefoot, least width of color of underparts not more than 40 per cent of greatest width of color of upper parts; height of tympanic bulla not more than three-fifths distance from its anterior margin to foramen ovale.

Description.—Size.—Male: A nontypical specimen from Cerro San Felipe, Oaxaca, measures: Total length, 500; length of tail, 205; length of hind foot, 52.

Female: The type specimen, measures: Total length, 418; length of tail, 160; length of hind foot, 45.

In this male the tail is 70, and in the female, 62 per cent as long as the head and body. In each the hind foot is longer than the basal length.

The differences in external measurements between these two specimens, representing the two sexes, are: Total length, 82; length of tail, 45; length of hind foot, 7.

Externals.—As described in Mustela frenata frenata.

Color (based on type specimen).—Color and color pattern as described in Mustela frenata frenata except that: blackish of sides and top of head extends one-fourth of way back to shoulders from ears; throat and breast as well as chin white; remainder of underparts near (16´ c) Ochraceous-Buff; least width of color of underparts equals 36 per cent of greatest width of color of upper parts; black tip of tail equal to 28 per cent of length of tail-vertebrae.

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 page 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

Putorius macrophonius Elliot, Proc. Biol. Soc. Washington, 18:235, December 9, 1905.

Mustela macrophonius, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912.

Mustela frenata macrophonius, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, adult, skull and skin; no. 14063, Field Mus. Nat. Hist.; Achotal, Veracruz, México; January 15, 1904; obtained by Edmund Heller and Charles M. Barber; original no. 3424.

The skull (plates 24-26, 30) is complete and unbroken. Excepting right P2, which has been aborted or broken away, all the teeth are present. The skin is well made and in good condition. As shown by the scrotal pouch, the specimen is a male.

Range.—Tropical Life-zone, probably into Boreal life-zones, of mountains along eastern border of southern Veracruz. See figure 29 on page 221.

Characters for ready recognition.—Differs from M. frenata frenata and M. f. perotae and M. f. leucoparia in lacking color of underparts on hind feet and in larger skull (skulls of adult males with basilar length more than 52.5); from M. f. goldmani by larger size of skull (see above) and entire animal and wider tympanic bullae; from M. f. tropicalis and M. f. perda by larger size (total length of adult males more than 510), postorbital breadth amounting to less than combined length of upper premolars.

Description.—Size.—Male: External measurements of the type specimen, an adult, are: Total length, 598; length of tail, 246; length of hind foot, 59. Tail 70 per cent as long as body; length of hind foot more than basal length.

Female: The skin, without field collector's measurements, of an adult female from Pérez, Veracruz, shows this sex to be correspondingly large. Because the skin is understuffed and because the hind feet are skinned out, reliable measurements can not be obtained from the dried skin.

Externals.—As described in Mustela frenata goldmani except that all carpal vibrissae are of same color as upper parts and that hairiness of foot-soles is halfway between that shown in figures 20 and 21.

Color.—As in darkest individuals of M. f. goldmani, thus, color of upper parts on posterior part of back near (n) Argus Brown. Color of underparts near (12) Mikado Orange in a juvenile, extending distally on posterior sides of forelegs onto inner toes and on hind legs to points between knees and heels. Least width of color of underparts 28 per cent of greatest width of color of upper parts. Black tip of tail 34 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on type specimen): See measurements and plates 24-26, 30. As described in Mustela frenata frenata except that: Weight, 6.9 grams; basilar length, 54.1; zygomatic breadth less than distance between condylar foramen and M1 or that between anterior palatine foramen and anterior margin of tympanic bulla; interorbital breadth 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 4 to 6 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; anterior margin of masseteric fossa below posterior half of m2.

Female (based on no. 132528): See measurements and plates 37-39. As described in Mustela frenata frenata except that: Weight, 3.6 grams; basilar length, 43.5; zygomatic breadth less than distance between condylar foramen and M1 and more or less than (in the single specimen, equal to) that between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more or less than (about equal to) outside length of P4; tympanic bulla as far posterior to foramen ovale as width of 4 or 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale.

The skull of the female is 48 per cent lighter than that of the male.

Comparison of the skull with that of M. f. goldmani is made in the account of that subspecies. Similar differences probably exist between males of perotae and macrophonius. As compared with skulls of males of M. f. tropicalis and perda, the skull of the male of macrophonius is larger in every measurement taken. The postorbital constriction is less, rather than more, than the combined length of the upper premolars. Relative to the basilar length, the following measurements are less than in any specimen of tropicalis or perda: length of tooth-rows; orbitonasal length; depth of skull at posterior border of upper molars; and depth of skull at anterior margin of basioccipital.

Remarks.—This large weasel appears to have escaped the notice of naturalists until the spring of 1903 when J. Friesser obtained an adult female and juvenal male at Pérez for the collection of the United States Bureau of Biological Survey. These specimens were tentatively referred to Mustela tropicalis. In the following January, Edmund Heller and Charles M. Barber obtained the adult male that was made the type specimen by Elliot who did not see, or if he did, did not mention, the specimens from Pérez. He did, however, refer a young female from Xuchil, Veracruz, to his Putorius macrophonius. This young female is here referred to Mustela frenata tropicalis.

The extent of the geographic range of this subspecies is not well known.

Mustela frenata macrophonius and M. f. texensis are the largest American weasels. The basilar length in the type specimen is greater by one-tenth of a millimeter than in the type specimen of M. f. texensis. The other cranial measurements taken are greater in M. f. texensis. The skull of the female from Pérez is one of the largest skulls examined of that sex. The juvenal male has teeth as large as those of the type specimen and the skull is the largest for its age of any seen. Although the skin of the female is understuffed and hence does not provide reliable measurements, it shows that the female is also large.

The white bands in front of the ears are confluent with the white patch between the eyes on one side only in one specimen. It is the juvenal male. These bands are not confluent with the color of the underparts on either side in the female and on one side only in the adult male. None of the specimens has a white patch between the ears. The dark spot at each angle of the mouth is present only in the juvenile where it occurs on each side. Of the three specimens, the juvenile is the darkest and the adult male the lightest. The white facial markings are most extensive in the juvenal male and the least extensive in the adult female.

M. f. macrophonius most closely resembles M. f. goldmani but in the relatively flattened braincase, deep constriction of the postorbital region and general angularity of the skull approaches M. f. perotae and M. f. frenata.

Only one of the three skulls, that of the female, shows evidence of infestation of the frontal sinuses by parasites, and this did not result in malformation of the interorbital region.

Specimens examined.—Total number, 3, all from México, listed by localities from north to south.

Veracruz. Achotal, 1 (Field Mus. Nat. Hist.); Pérez, 2 (U. S. Nat. Mus.).

Mustela frenata tropicalis (Merriam)

Long-tailed Weasel

Plates 25, 26, 27, 30, 37, 38 and 39

Putorius tropicalis Merriam, N. Amer. Fauna, 11:30, pl. 3, figs. 5, 5a, 6, 6a, text fig. 16, June 30, 1896; Merriam, Proc. Biol. Soc. Washington, 15:68, March 22, 1902.

Putorius frenatus, Merriam, N. Amer. Fauna, 11:27, June 30, 1896.

Mustela tropicalis tropicalis, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912; Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916.

Mustela frenata tropicalis, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, adult, skull and skin; no. 54994, U. S. Nat. Mus., Biol. Surv. Coll.; Jico, Veracruz, México; July 9, 1893; obtained by E. W. Nelson; original no. 5195.

The skull (plates 25-27, 30) is complete. All the upper incisors, except the second and third on the left side, are missing. The right upper canine is broken. The skin is well made and in good condition.

Range.—Up to 5000 feet (as now known) in Tropical Life-zone of Veracruz, México. See figure 29 on page 221.

Characters for ready recognition.—Differs from M. f. frenata and M. f. perotae in least width of color of underparts not exceeding 36 per cent of greatest width of color of upper parts and in postorbital breadth exceeding length of upper molar and premolar tooth-rows; from M. f. macrophonius and M. f. perda in least width of color of underparts averaging more than 29 per cent of greatest width of color of upper parts; and from M. f. perda by longer tympanic bullae which in males are more than 14.9; and from M. f. macrophonius by lesser basilar length (not more than 48) and in postorbital breadth exceeding length of upper molar and premolar tooth-row.

Description.—Size.—Male: The type specimen and no. 12764/11058, a subadult, from Jalapa, Veracruz, measure, respectively, as follows: Total length, 444, 442; length of tail, 175, 160; length of hind foot, 50, 47. The tail is 65 and 57 per cent as long as the head and body. The hind foot is more or less than (approximately equal to) the basilar length.

Female: Merriam (1896:31) gives the measurements of a female topotype (probably no. 54993, U. S. Nat. Mus., which has no measurements written on the attached label) as: Total length, 333; length of tail, 121; length of hind foot, 37. The length of the tail amounts to 57 per cent of the length of the body. The length of the hind foot of no. 54993, U. S. Nat. Mus. is the same as the basal length.

The differences in external measurements between the male and the female topotypes are: Total length, 111; length of tail, 54; length of hind foot, 13.

Externals.—As described in Mustela frenata frenata except that carpal vibrissae do not reach apical pad of fifth digit and hairiness of foot soles is less.

Color.—As described in M. f. frenata except that: Blackish of head extends half way or more from ears to shoulders; upper parts near (14) Brussels Brown or slightly faded tone 2 of Maroon of Oberthür and Dauthenay, pl. 341; underparts of juvenal pelage near (a) Ochraceous-Buff. Least width of color of underparts averaging (in three specimens from Jico and one from Jalapa) 34 (extremes 30-37) per cent of greatest width of color of underparts. Black tip of tail, in two male topotypes, 57.5 (55 and 60) mm. long; thus longer than hind foot and in each individual comprising 34 per cent of length of tail-vertebrae.

As compared with M. f. frenata: White facial markings slightly less extensive; blackish (not black) of head extending onto neck; upper parts slightly darker; ventral side of tail noticeably darker; color of underparts more restricted, averaging approximately one-third rather than nearly one-half width color of upper parts; black tip of tail one-third rather than one-fourth length of tail and much longer than hind foot. Similar differences of lesser amount exist between perotae and tropicalis. M. f. perda, macrophonius and goldmani bear the opposite relation to tropicalis. That is to say, in the latter: White facial markings slightly more extensive; blackish of head less extended over neck; upper parts markedly lighter; color of underparts less restricted and black tip of tail shorter.

Skull and teeth.—Male (based on type specimen and a subadult, no. 11058, from Jalapa): See measurements and plates 25-27, 30. As described in Mustela frenata perda except that: Weight 4.7 (4.6 and 4.7) grams; basilar length 46.7 (45.5 and 47.8); zygomatic breadth more or less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more than length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 (including I3) upper incisors; anterior margin of masseteric fossa below middle of m2 or posterior to that tooth.

Female (based on no. 54993 and no. 1060): See measurements and plates 37-39. As described in Mustela frenata perda except that: Weight (of 54993) 2.2 grams; basilar length, 37.5 (36.0-39.0); zygomatic breadth more or less than distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more than greatest length of P4; height of tympanic bulla equal to one-third to three-fourths of distance from its anterior margin to foramen ovale.

The skull of the adult female is 53 per cent lighter than that of the type specimen, a male.

Comparison of the skulls of males and females with those of M. f. perda, the nearest relative, has been made in the discussion of that subspecies. Some of the features that readily distinguish skulls of M. f. tropicalis from those of M. f. frenata, perotae and macrophonius are as follows: Weight less than 4.8 grams; basilar length less than 48; postorbital breadth more than length of upper M-Pm tooth-row. The skulls of male frenata, perotae and macrophonius are much larger, heavier, and are decidedly more angular with more constricted postorbital region the least width of which is less than the length of the upper premolars. In frenata the anterior margins of the tympanic bullae are protruded much farther from the braincase. The skull of the female of M. f. tropicalis is smaller, weighing less than 3 grams; basilar length less than 41; postorbital breadth more than length of upper molar and premolar tooth-row.

Remarks.—This subspecies was originally described by Merriam as a full species. Later he described Putorius tropicalis perdus as another subspecies. Allen (1916) placed P. t. perdus in synonymy but named Mustela tropicalis nicaraguae as new. In the present paper all three forms are recognized but are regarded as only subspecifically distinct from the other bridled weasels of México and Central America.

The limits of the geographic range of tropicalis are fairly well known on the south and west but the only specimen available from the tropical coastal region north of Jico, is a young female from a point 50 miles south of Victoria. Thus, how far north along the coast it ranges toward Matamoros, where M. f. frenata occurs, is not known. The three specimens from Jico, a young female from Jalapa and another adult collected by J. Potts and labeled as coming from México City, are assumed to be typical. The latter specimen certainly came from an elevation lower than that of México City because M. f. frenata occurs there. Although the female from Jalapa, agrees well with specimens from Jico, a male, no. 12764/11058, from Jalapa, has a relatively broader skull, as in perda, although the tympanic bullae are short as in tropicalis. The resemblances to perda in features of coloration are: slightly darker upper parts, and the termination just below the knees of the color of the underparts. There are three specimens labeled as from Orizaba that indicate intergradation with perotae as does also the coloration of the juvenal female from 5 kilometers north of Jalapa. The specimens labeled as from Orizaba are old, poorly-prepared skins, only two of which have partial skulls. The size and coloration of the skins suggest perotae as do also the partial skulls in some respects although the skulls show greater resemblance to those of tropicalis.

The topotype, female, no. 54993, was figured by Merriam (1896, fig. 16, p. 31) along with that of what now is the type specimen of M. f. perotae. Merriam called attention to the great difference in size between the skulls of the two sexes of M. f. tropicalis and compared the condition to that found in noveboracensis. Although the skull of the female from Jico is fully adult, it probably is exceptionally small.

The young female from Xuchil is indistinguishable in coloration from the juvenal female of M. f. perotae from Perote, but in size of skull and size of teeth is intermediate between the female of tropicalis from Jalapa and the females from Cofre de Perote.

There is then, indication of intergradation with M. f. perotae as well as with M. f. perda. M. f. tropicalis differs from M. f. perotae and M. f. frenata in about the same way that M. f. perda differs from M. f. goldmani and M. f. macrophonius. M. f. tropicalis and perda each is smaller and more intensely colored than goldmani and macrophonius, and inhabits the lowland to the east of their highland relative.

At least five of the nine skulls have the frontal sinuses infested by parasites.

Specimens examined.—Total number, 13, all from México, listed by localities from north to south. Unless otherwise indicated specimens are in the collection of the United States National Museum.

Tamaulipas. 50 mi. S Victoria, 1[71]. Near? México City, 1.

Veracruz. Jico, 3; 5 km. N Jalapa, 1[90]; Jalapa, 2 (1[2], 1[75]); Xuchil, 1[60]; Orizaba, 4 (2[75], 1[4]).

Mustela frenata perda (Merriam)

Long-tailed Weasel

Plates 25, 26, 27, 30, 37, 38 and 39

Putorius tropicalis perdus Merriam, Proc. Biol. Soc. Washington, 15:67, March 22, 1902.

Putorius (Gale) brasiliensis frenatus, Coues, Fur-bearing animals, p. 142, 1877 (part).

Mustela tropicalis perda, Miller, U. S. Nat. Mus. Bull., 79:100, December 31, 1912.

Mustela tropicalis tropicalis, Allen, Bull. Amer. Mus. Nat. Hist., 35:99, April 28, 1916.

Mustela frenata perda, Hall, Carnegie Instit. Washington Publ. 473:109, November 20, 1936.

Type.—Male, subadult, skull and skin; no. 100041, U. S. Nat. Mus., Biol. Surv. Coll.; Teapa, Tabasco, México; March 31, 1900; obtained by E. W. Nelson and E. A. Goldman; original no., 14074.

The skull (plates 25-27, 30) is unbroken and all the teeth are present and entire. The skin is well made and in good condition.

Range.—Fifty feet (Chichén Itzá) to 4000 feet (San Vicente) in Lower Tropical Life-zone south from southern Veracruz through southern México into Guatemala. See figure 29 on page 221.

Characters for ready recognition.—Differs from M. f. nicaraguae in lesser extent of color of underparts (not more than 22 per cent of greatest extent of color of upper parts), black tip of tail more than 38 per cent of length of tail, and broader skull (in adult males, mastoid breadth more than 23.9 and zygomatic breadth more than 27.4); from M. f. tropicalis in more restricted color of underparts (least width of color of underparts less than 28 per cent of greatest width of color of upper parts) and shorter tympanic bullae, which in males are less than 15; from M. f. goldmani by total length not exceeding 489, least width of color of underparts not exceeding 24 per cent of greatest width of color of upper parts, weight of adult skull less than 5 grams and basilar length less than 48.5.

Description.—Size.—Male: The type specimen and another subadult from San Vicente, Chiapas, measure, respectively, as follows: Total length, 473 and 443; length of tail, 184 and 169; length of hind foot, 51 and 51.5. The tail is 62 and 64 per cent as long as the head and body. The length of the hind foot is greater than the basal length.

Female: Estimates made from the dried skin of no. 218036 are: Total length, 375; length of tail, 140; length of hind foot, 40. The hind foot of no. 65422 from Catemaco also measures 40.

The average differences in external measurements of the two sexes are: Total length, 83; length of tail, 37; length of hind foot, 11.

Externals.—As described in Mustela frenata goldmani except that hairiness of foot soles is slightly less.

Color.—As described in Mustela frenata goldmani except that: back near (n) Argus Brown or Carbon Brown, tone 3, of Oberthür and Dauthenay, pl. 342; underparts Ochraceous-Buff. Least width of color of underparts, in four specimens, averaging 20 (extremes 18-22) per cent of greatest width of color of upper parts; black tip of tail, in two subadult males, averaging 48 (extremes 46-49) per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on type specimen and subadult no. 132997 from San Vicente): See measurements and plates 25-27, 30; weight 4.4 grams (same for each); basilar length 45.7 (45.3 and 46.1); zygomatic breadth 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 or less than (approximately equal to) length of upper premolars and greater than width of basioccipital measured from medial 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 not greater 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 than length of lower molar and premolar tooth-row and longer or shorter than rostrum; anterior margin of masseteric fossa below middle of m2.

Female (based on two subadults, nos. 65422 and 218036): See measurements and plates 36-39; weight, 2.4 (2.3-2.5) grams; basilar length, 40.5 (40.4-40.6); zygomatic breadth less than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than 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 outside length of P4 and less than inside length of same; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 or 6 upper incisors; height of tympanic bulla equal to one-third to one-half distance from its anterior margin to foramen ovale; length of tympanic bulla more than length of lower molar and premolar tooth-row and more or less than (about equal to) length of rostrum.

The skull of the female averages 48 per cent lighter than that of the male.

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.