Part 44

_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.