Part 48

_Color._--Rarely a few white hairs anterior to each ear; posterior fifth of each upper lip white; top of head, posteriorly to slightly behind ears, black, grading into color of upper parts of body; dark spots at angles of mouth absent; tip of tail black; remainder of upper parts near (_n_) Argus Brown and Carbon Brown, tone 3 (pl. 342, Oberthür and Dauthenay); chin whitish; remainder of underparts Warm Buff; color of underparts extends distally on posterior sides of forelegs to wrists but not reaching foot-soles and on hind legs to slightly below knees. Least width of color of underparts 24 per cent of greatest width of color of upper parts in each of two males and 19 to 30 per cent in three females. Black tip of tail longer than hind foot and averaging 40 (39-42) per cent of length of tail-vertebrae.

_Skull and teeth._--Male (based on type specimen and adult no. 24132): See measurements and plates 27-29. As described in _Mustela frenata macrura_ except that: Weight, 4.5 (4.2 and 4.8); basilar length, 44.6 (44.0-45.3); zygomatic breadth more than distance between condylar foramen and M1 or than between anterior palatine foramen and anterior margin of tympanic bulla; breadth of rostrum more or less than (approximately equal to) length of tympanic bulla; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of rostrum; anterior margin of masseteric fossa posterior to m2 by length of that tooth.

Female (based on nos. 24134 to 24136): See measurements. As described in _Mustela frenata macrura_ except that: Weight, 1.7 (1.5-1.9) grams; basilar length, 36.5 (35.3-38.1); zygomatic breadth less than distance between anterior palatine foramen and anterior margin of tympanic bulla; least width of palate more or less than (approximately equal to) outside length of P4; length of tympanic bulla less than length of rostrum.

The skull of the female averages 62 per cent lighter than that of the male.

The skull of the male is generally large and heavy as are the teeth. Comparison with _macrura_ is made in the account of that subspecies. From males of _affinis_ those of _helleri_ differ in: skull shorter; breadth of rostrum and interorbital breadth actually and relatively greater.

_Remarks._--The five specimens examined of this subspecies were taken by Edmund Heller for the Field Museum of Natural History in 1922 and 1923. It is to honor his contributions to mammalogy that the subspecies is named _helleri_. No. 24135 is the specimen carried as a pet for some time by Mr. and Mrs. Heller and of which Mrs. Heller (1924:481) has given an account.

This subspecies is insufficiently known, especially as to geographic range; probably it occupies a considerable range in the Tropical Life-zone along the eastern base of the Andes. The three females, two from Ambo and one from Huanuco, come from a much higher altitude than do the two males and the climate is said to be arid at Ambo and Huanuco. The skulls of the females are 62 per cent lighter and correspondingly smaller in measurements, than those of males. This difference is more than that found in any other South American weasel and it may be that the females are of a subspecies other than _helleri_.

The type specimen has a broad skull with major proportions strikingly like those of _Mustela stolzmanni_. Possibly the similar climatic conditions under which the two live have left their impress in similar fashion in this part of each of the two species. The teeth, tympanic bullae, and certain other parts of the skull are, however, so differently proportioned as to show that the skulls represent two species. The referred male has a much longer skull than the type specimen and the relative proportions of breadth and depth of the two skulls differ widely. Judging from large series of weasels examined from localities outside the range of _M. f. helleri_, the two skulls probably represent almost the maximum of individual variation occurring in one subspecies.

The dark color is as might be expected since _helleri_ inhabits the humid Tropical Zone.

None of the five skulls shows signs of having had the frontal sinuses infested by parasites.

_Specimens examined._--Total number, 5, all in the Field Museum of Natural History.

=Perú=: 3500 ft., Hacienda Buena Vista, Río Chinchao, 1; 3000 ft., Hacienda San Antonio, Río Chinchao, 1; Huanuco, 1; Ambo, 2.

=Mustela frenata agilis= Tschudi

Long-tailed Weasel

Plates 27, 28, 29, 39 and 40

_Mustela agilis_ Tschudi, Fauna Peruana, p. 110, 1844; Gray, Proc. Zoöl. Soc. London, 1865:113, 1865; Taczanowski, Proc. Zoöl. Soc. London, 1874:311, 1874; Taczanowski, Proc. Zoöl. Soc. London, 1881:648, 1881; Allen, Bull. Amer. Mus. Nat. Hist., 35:104; April 28, 1916; Thomas, Proc. U. S. Nat. Mus., 58-224, 1920.

_Mustela macrura_, Allen, Bull. Amer. Mus. Nat. Hist., 35:103, April 28, 1916.

_Mustela frenata agilis_, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939.

_Type._--No type specimen, or type locality more restricted than cold, barren highlands of the Cordillera [referring to Perú] designated.

_Range._--High, barren Cordillera of Perú (see Tschudi, orig. descr.); as here restricted, Temperate Life-zone and higher in western Andes and intermountain valleys of Perú. See figure 29 on page 221.

_Characters for ready recognition._--Differs from _Mustela frenata macrura_ by lighter color (Chocolate, tone 2 rather than 3, pl. 343, Oberthür and Dauthenay) of upper parts; length of upper tooth-rows, in females, less than 13; inside length of P4 more than 4.6; from _M. f. aureoventris_ by smaller teeth (maximum size just given for _agilis_); from _M. f. boliviensis_ by lighter color, upper parts being Chocolate, tone 2, pl. 343, rather than tone 4 or darker of Carbon Brown, pl. 342 (Oberthür and Dauthenay).

_Description._--_Size._--Male: The stuffed skin of an adult, from Lima, measures: Total length, 460; length of tail, 125; length of hind foot, 45.7. A skin alone from Huarochirí has a body, as now stuffed, 277 mm. long. The tail is missing and the bones of the hind feet have been removed.

Female: The mounted specimen, no. 565, Mus. Polonais d'Hist. Nat., yields measurements, taken by me, as follows: Total length, 250; length of tail, 75; length of hind foot, 32.5. The female, no. 21147, from Macate, measures, 300, 102, 34.

_Externals._--Longest facial vibrissae, either dark-or light-colored and extending beyond ear; carpal vibrissae either dark-or light-colored and extending to apical pad of fifth digit; hairiness of foot soles as shown in figure 20.

_Color._--Tschudi's description of the color is, in substance, as follows: Head, back and tail reddish gray; base of hair gray, followed by broader grayish-yellow ring and then reddish-brown tip; nose simply dark brown or upper lips edged with white; throat, breast, belly and higher parts of inner sides of extremities whitish gray, at times wholly gray, bases of hairs always gray; feet darker than body, almost chestnut brown; tail darker on tip than at base; ears externally dark brown, internally whitish.

No. 565 possibly somewhat faded from exposure to light, has all the upper parts near (14´ _j_) Ochraceous-Tawny or Cinnamon, and tone 4 of Oberthür and Dauthenay, plate 323; posterior half of each upper lip white; no other white facial markings present; dark spot at each angle of mouth (one spot confluent with color of upper parts); tip of tail probably black (tip missing); underparts white, belly probably originally with slight tinge of yellow or allied color; color of underparts extending distally on forelegs to feet and onto upper sides of toes and on hind legs to just above heels. Least width of color of underparts equal to about one-fourth of greatest width of color of upper parts.

No. 21147, subadult, from Macate, has a white band confluent with the underparts extending anterodorsally anterior to each ear and the posterior third of each upper lip white. Top of head near (_n_) Mars Brown, and Carbon Brown, tone 3 (pl. 342, Ober. and Dauth.); tip of tail black; remainder of upper parts near (16" _j_) Tawny-Olive, and Chocolate (tone 2, of pl. 343 of Ober. and Dauth.) or Raw Umber (tone 3 of pl. 301 of Ober. and Dauth.); anterior half of underparts, including posterior sides of forelegs and antipalmar faces of forefeet, white; remainder of underparts tinged with Warm Buff and extended on posterior legs almost to ankles.

No. 8.1.10.1., male adult, from Lima, is also light colored, and as described in no. 21147, except that left side of head has a white spot rather than bar; posterior eighth of each upper lip white; white frontonasal spot present, 11 x 11 mm.; antipalmar faces of forefeet spotted with brown color of upper parts; color of underparts extending distally on hind legs along medial side of foot to point halfway between heel and tip of inner toe.

No. 13257 from Huarochirí in color and color pattern closely resembles no. 21147. It differs from no. 21147 in slightly lighter color of upper parts, entirely white underparts, less extension of color of underparts onto forefeet, few white hairs instead of white band in front of each ear; color of underparts more restricted.

In each of the four specimens, the least width of the underparts, expressed as a percentage of the upper parts, is as follows: no. 13257, 11 per cent; no. 21147, 29 per cent; no. 565, 31 per cent; no. 8.1.10.1., nineteen per cent.

_Skull and teeth._--Male (based on no. 8.1.10.1.): See measurements and plates 27-29. As described in _Mustela frenata macrura_ except that: Weight 4.1 grams; basilar length, 42.5; zygomatic breadth more than distance between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth less than postpalatal length; tympanic bullae shorter than rostrum.

Female (based on no. 21147): See measurements and plates 39 and 40. As described in _Mustela frenata macrura_ except that: Weight (no. 21147, subadult), 1.5 grams; basilar length, 35.2; least width of palate less than outside length of P4; tympanic bulla as far posterior to foramen ovale as combined width of five upper incisors; no. 565 answers to the same description but differs from no. 21147 in greater basilar length and larger tympanic bullae which are slightly more projected, at their anterior margins, from the braincase.

To judge from the skull of the female from Macate and the skull of the male from Lima, the skull and teeth of _agilis_ are smaller than in any other South American subspecies of _Mustela frenata_, except _M. f. boliviensis_.

_Remarks._--Tschudi almost certainly used the name _Mustela agilis_ in a composite sense. His statement (see quoted matter below) about the marked variation in color of this species, as represented by the skins carried by the Indian women as purses, indicates that the forms here designated as _Mustela macrura_, _M. helleri_ and possibly others additional to the one here called _agilis_ were included by him under the name _Mustela agilis_. Taczanowski took account of _Mustela agilis_ when he described other species from Perú. Allen (1916:104) and Thomas (1920:224) were not convinced that _Mustela agilis_ and _Mustela macrura_ were distinct species or subspecies.

Search on August 28, 1937, in the Musée d'Histoire Naturelle, at Neuchatel, Switzerland, by Mr. Théodore Delachaux, assistant there, and the writer, revealed no trace of weasels from Tschudi's collection, although some other specimens of mammals that he figured in the "Fauna Peruana" are preserved in that Museum. Not only were the collections of specimens examined but the new catalogue and old catalogue of mammals were vainly searched for mention of weasels deposited by Tschudi. Later, at the British Museum of Natural History, on page 105 of a personal notebook, of the late Mr. Oldfield Thomas, record was found of his fruitless search for the same specimens of _Mustela_ in May, 1902, at Neuchatel.

Although Tschudi certainly used the name _Mustela agilis_ in a composite sense, as subspecies are at present understood, his description most nearly applies to the light-colored animals from western Perú--the lightest colored of any South American weasels seen. They are of approximately the same color as North American subspecies inhabiting semiarid regions, for example _Mustela frenata longicauda_ of the Great Plains.

Another, but in my opinion less weighty, justification for applying Tschudi's name _agilis_ to these light-colored weasels of western Perú is that by one line of reasoning, Taczanowski in naming _macrura_ (_jelskii_ is a synonym of it) from farther eastward in Perú, and that Hall in naming _helleri_ from still farther eastward, and _boliviensis_ to the southeastward, geographically restricted the application of the name _agilis_. Hall's action did this because he recognized geographic variation and employed the subspecies concept. Taczanowski, however, proposed his name _macrura_ for a kind of animal which he indicated was specifically (as opposed to subspecifically) distinct from _agilis_ and his account (1881:649) of _jelskii_ indicates that he thought _Mustela agilis_ Tschudi might occur in the same place as the animals which he named as new kinds. Thus, we can not credit Taczanowski with _intent_ to restrict the name _agilis_ geographically, even though later authors may choose to rule that his naming of _macrura_ in effect did so restrict the application of _Mustela agilis_ Tschudi.

The equivalents in millimeters given by Allen (1916:104) for Tschudi's measurements of 9 to 10 inches entire length, and tails of 4 inches to 4 inches and 4 lines, apparently are based on the London scale in use today. If Tschudi employed the Rhine scale also of eight lines to the inch, but one which has the foot longer by an amount of 20 millimeters, or the Leipzig scale in which the foot is 22 millimeters shorter than the London foot, the measurements recorded by Tschudi differ in one direction or the other from those computed by Allen. However, knowledge of which scale Tschudi employed would not help much, if any, in more precise application of the name _agilis_ because he does not indicate whether his measurements are of male or female animals; animals of the two sexes of the same subspecies differ more in external measurements than animals of the same sex of different subspecies of Peruvian weasels.

Specimen no. 565, in the Polish Museum of Natural History, without definite locality, is provisionally referred to this subspecies. The specimen is intermediate in several respects between the female from Macate and the one of _macrura_ from Cutervo.

Tschudi (1844:111-112) has given the following account: "_Lebensweise und geographische Verbreitung_. Das peruanische Wiesel lebt auf den kalten, öden Hochebenen der Cordillera an sonnigen Steinhaufen und Felsen gewöhnlich in Gesellschaft von 8-12 Stücken. Diese Thierchen sind so ausserordentlich behende und scheu, dass bei dem leisesten Geräusche die ganze Schaar mit Blitzesschnelle verschwindet. Es ist uns auch nie gelungen, eines derselben zu erlegen. Die Indianer aber verstehen es, dieselben lebendig einzufangen und zu zähmen. Ein sehr zahmes sahen wir bei einer uns befreundeten Dame in Tarma; gegen alle Fremden biss es mit Wuth und liess sich nicht anfassen, während es sich von seiner Herrin Alles gefallen liess; sie öffnete ihm den Mund und steckte ihm den Finger hinein, ohne dass es eine böse Miene dazu machte, während es bei der geringsten Bewegung, die wir machten, es zu ergreifen, grimmig auf uns lossprang. Wenn es eingeschüchtert wurde, versteckte es sich in den Busen seiner Gebieterin und kroch ihr bald nachher zum Aermel heraus. An den Wänden und Meublen kletterte es mit grosser Behendigkeit und schlüpfte durch so kleine Ritzen und Löcher, dass wir fast an der Möglichkeit dieses Hindurchdringens gezweifelt haben würden, wenn wir es nicht selbst mit angesehen hätten. Wenn es unartig war, wurde es mit einer Schnur an seinem kleinen Halsbande festgebunden; dadurch vermehrte sich sein Zorn, so dass es zuweilen gegen die Dame auffuhr. Mehrmals verschwand es während 8-10 Tagen und kam dann plötzlich wieder zum Vorschein. Seine Nahrung bestand in Gemüse und Fleisch, besonders aber liebte es Zuckerbrod in Milch aufgeweicht; einmal machte es sich an einen Kanarienvogel, den es auch tödtete. Es erhielt seine Strafe und verschwand dann für immer. Die Indianer sollen dieses Wiesel zum Fange der Viscacha abrichten (davon weiter unten). Sie nennen es Comadreja, auch Ardilla. ([footnote] Ardilla ist spanisch und heisst Eichhörnchen. Mit diesem Namen werden sehr verschiedene Thiere bezeichnet; ausser dem Sc. variabilis und der Galictis agilis auch noch mehrere Nager und einige Didelphysarten.) Die Indianerinnen verfertigen sich aus dem kleinen Felle Geldbeutel. Des Sonntags trifft man unter den vielen tausend Punaindianerinnen die nach den grossen Dörfern der Sierra kommen, um ihre Einkäufe zu machen, kaum ein halbes Dutzend, die nicht solche Börsen mit sich führten, und dann kann man auch die verschiedensten Farbennuancen, die bei dieser Species vorkommen, beobachten."

None of the three skulls referred to this subspecies shows infestation of the frontal sinuses by parasites.

_Specimens examined._--Total number, 4.

=Perú=: Macate, 1 (Field Mus. Nat. Hist.); Huarochirí, 1 (Mus. Comp. Zool.); Lima, 1 (British Mus. Nat. Hist.); no locality more definite than Perú, 1 (Mus. Polonais d'Hist. Nat.).

=Mustela frenata macrura= Taczanowski

Long-tailed Weasel

Plates 1, 27, 28, 29, 30, 37, 38, 39 and 40

_Mustela macrura_ Taczanowski, Proc. Zoöl. Soc. London, 1874:311, pl. 48, May 19, 1874; _ibid_., 1881:647, May 17, 1881; _ibid_., 835, November 15, 1881; Lönnberg, Arkiv för Zool., 8 (no. 1):21, 1913 (?); Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914; Allen, Bull. Amer. Mus. Nat. Hist., 35:101, April 28, 1916; Lönnberg, Arkiv för Zool., 14 (no. 4):11, 1921.

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

_Mustela jelskii_ Taczanowski, Proc. Zoöl. Soc. London, 1881:647, May 17, 1881.

_Mustela affinis_, Lönnberg, Arkiv för Zool., 8 (no. 1):21, July 12, 1913.

_Mustela aureoventris_, Thomas, Proc. U. S. Nat. Mus., 58:224, 1920.

_Mustela frenata macrura_, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939.

_Type._--Male, adult, mounted skin, with skull separate; no. 561, Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Lake Junín, central Perú; 1873; obtained by M. Jelski.

The skull (plates 27-29, 30), mounted with the skin but removed by me for study, lacks the right jugal, the basisphenoid, the basioccipital and parts of each exoccipital bearing the exoccipital condyles. The right tympanic bulla, although detached from the skull, is preserved separately. The teeth all are present and entire. The skin is fairly-well mounted, in a good state of preservation, and shows no fading due to exposure to light.

_Range._--Altitudinally, 3200 (Guainche) to at least 12000 feet (Pichincha); Upper Subtropical and Temperate life-zones of central Perú and Ecuador north from the states of Apurimac and Cuzco, Perú, to San Antonio, northern Ecuador. See figure 29 on page 221.

_Characters for ready recognition._--Differs from _Mustela africana stolzmanni_ by absence of median, longitudinal, abdominal stripe of same color as upper parts; presence of p2 and two roots rather than one root on P2; from _Mustela frenata helleri_, _M. f. affinis_ and _M. f. aureoventris_ by lighter color of upper parts which are Chocolate tone 3, pl. 343, Oberthür and Dauthenay, whereas, with reference to the same color standard, the colors are: in _helleri_, Carbon Brown, tone 3, pl. 342; in _affinis_, Reddish Black, tone 2, pl. 344; in _aureoventris_, Reddish Black, tone 4, pl. 344; from _M. f. agilis_ by darker color (Chocolate, tone 3 rather than 2, pl. 343, Oberthür and Dauthenay) of upper parts, length of upper tooth-rows, in females, more than 13, inside length of P4 more than 4.6; from _M. f. boliviensis_ by lighter color of upper parts which are as above rather than tone 4 of Carbon Brown, pl. 342 of Oberthür and Dauthenay, and larger size (in males, hind foot more than 45 and m1 more than 5.6).

_Description._--_Size._--Male (measurements as recorded by Taczanowski in the original description, for two specimens, type and topotype, with correction of the length of tail of his "female" [= male]): Total length, 420, 415; length of tail, 150, 145; length of hind foot, 51, 51. An adult from Yana Mayo, Río Tarma, was measured by Hendee as 394, 134. Hind foot relaxed measures, 47. Tail 55 per cent as long as head and body. Length of hind foot more than basal length.

Female (based on measurements given by Taczanowski (1881:647) of no. 564): Total length, 323; length of tail, 120; length of hind foot, 37. Tail 59 per cent as long as head and body. Length of hind foot approximately equal to basal length.

Differences in external measurements of the two sexes are: Total length, 87; length of tail, 23; length of hind foot, 13.

_Externals._--Longest facial vibrissae extending beyond ear; carpal vibrissae color of either upper parts or underparts; hairiness of foot-soles as shown in figure 20.

_Color._--(Based on specimens from Cutervo and south thereof). Rarely few white hairs between eyes and in front of ears; top of head posteriorly to slightly behind eyes, near (_n_) Chestnut-Brown (Ridgway) and Carbon Brown, tone 2 or darker (pl. 342, Oberthür and Dauthenay); posterior half of upper lip rarely white; dark spots at angles of mouth absent; tip of tail black; remainder of upper parts near (_l_) Russet (Ridgway) and Chocolate, tone 3 (pl. 343, Ober. and Dauth.); underparts white or whitish on medial sides of forelegs, otherwise cream color with tinge of Ochraceous-Buff; color of underparts extended distally on posterior sides of forelegs to just below elbow (in type specimen) or onto forefeet (in specimen from Yana Mayo) and on medial sides of hind legs to points between knees and ankles. Least width of color of underparts averages (in six skins) 17 (14-21) per cent of greatest width of color of upper parts. Black tip of tail longer than hind foot and averaging 36 (32-49) per cent of length of tail-vertebrae.