Systematic Accounts of Species and Subspecies (part 4)

The name Mustela aureoventris Gray has been regarded by most authors as preoccupied by Mustela auriventer Hodgson (1841:909). However, the writer is not of this opinion and agrees with Thomas (1920:224) that "The name aureoventris is not invalidated by the auriventer of Hodgson, as, apart from 'one-letterist' differences, its first half comes from the adjective aureus, while Hodgson's name is based on the substantive aurum, so that not only the spellings but the derivations are different." The spelling of Gray's name should be aureoventris for this is the spelling in the original description which in pagination precedes the colored plate of the animal that is labeled Mustela aureoventris. Putorius brasiliensis var. aequatorialis Coues (1877:142) is the only name known to the writer that has been proposed as a substitute for Mustela aureoventris Gray.

Thomas (1920:224) treats Mustela macrura Taczanowski as a synonym of Mustela aureoventris Gray. Allen (1916:101) also treats the two names as applying to the same kind of weasel but regards aureoventris as preoccupied and therefore uses the name macrura. Taczanowski's original description (1874:311) and plate of Mustela macrura indicate an animal that is lighter colored than M. f. affinis. Gray's original description (1864:55) and plate of aureoventris indicate an animal that is darker colored than M. f. affinis. Indeed Gray (1865:115) in speaking of the type of aureoventris as compared with an adult from New Granada [= Colombia] that probably later became the type specimen of Mustela affinis, states: "The young from Quito is much darker than the adult;. . . ." Comparison of the plates accompanying the original descriptions of aureoventris and macrura well illustrate the difference stated in the written descriptions. My examination of the type specimens of M. macrura and M. f. aureoventris shows them to have been fairly accurately portrayed in the plates accompanying the original descriptions. Accordingly the two names are used for the two kinds of animals which appear, however, to be only subspecifically distinct.

Comparison of Gray's plate (1864, pl. 8) with the available specimens from South America indicates that the name aureoventris is based on an individual that is lighter colored than no. 34677 Amer. Mus. Nat. Hist., from Gualea, Ecuador, but on one which resembles no. 34677 more than it does the lighter-colored specimens from the Temperate Zone of Ecuador and northern Perú. Because Quito, Ecuador, is in the Temperate Life-zone and because the available specimens from this zone in Ecuador and northern Perú are distinctly lighter colored than Gray's plate representing the type of aureoventris shows this specimen to be, it is judged to have come from an altitude lower than that of Quito (9350 feet, according to Chapman, 1926:717); probably it came from the Subtropical Life-zone of Ecuador. Indeed Gray (1864:55) did not say that the specimen was collected or obtained at Quito but that it was ". . . received from Quito. . . ." Chapman (1926:717) has pointed out that Quito, since 1846 has been the distributing point for bird skins which specimens ". . . come from the vicinity of the city, from the 'Napo' region on the Amazonian slopes of the Andes, and from Nanegal, Gualea, and other localities on the Pacific side rarely below the Subtropical Zone." It is also pointed out that only some of the specimens are labeled with their approximate place of capture and that even then these localities cannot be accepted as definite; they indicate mainly whether the specimen is from the eastern or western side of the Andes.

The above mentioned considerations and information gained by study of the specimens cause me to think that the type is an intergrade tending toward the lighter-colored Mustela f. macrura of the Temperate Zone although sufficiently dark to be referred to the dark subspecies represented by no. 34677 Amer. Mus. Nat. Hist., from Gualea, Ecuador.

The skull of no. 34677 shows no infestation of the frontal sinuses by parasites.

Specimens examined.—Total number, 3, as follows:

Ecuador: Gualea, 1, Amer. Mus. Nat. Hist.

Colombia: 8325 ft., Munchique, 1, Amer. Mus. Nat. Hist. In the British Museum of Natural History, the type, (1).

Mustela frenata helleri Hall

Long-tailed Weasel

Plates 27, 28 and 29

Mustela frenata helleri Hall, Proc. Biol. Soc. Washington, 48:143, August 22, 1935; Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:175, 1939.

Type.—Male, adult, skull and skin; no. 24133, Field Mus. Nat. Hist.; 3000 feet, Hacienda San Antonio, Río Chinchao, Perú; August 22, 1922. Obtained by Edmund Heller. Original no. 6589.

The skull (plates -) is complete and unbroken. The teeth all are present, entire and but slightly worn. The skin is well made, unfaded, and in good condition.

Range.—Three thousand feet (type locality) to 6700 feet (Ambo), Tropical and Subtropical life-zones of eastern Perú. See figure on page .

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 macrura by darker color (Carbon Brown, tone 3, pl. 342 rather than Chocolate, tone 3, pl. 343, Oberthür and Dauthenay) of upper parts.

Description.—Size.—Male: Measurements of the type specimen and topotype, no. 24132, are, respectively, as follows: Total length, 382, 418; length of tail, 152, 164; length of hind foot, 52, 48. Tail 66 and 65 per cent as long as head and body. Hind foot more than basal length.

Female: Measurements of two referred females, no. 24134 from Ambo and no. 24136 from Huanuco, are, respectively, as follows: Total length, 328 and 303; length of tail, 118 and 103; length of hind foot, 39 and 38.5. Tail 56 and 51 per cent as long as head and body. Hind foot shorter than basal length.

The average differences in external measurements of the two sexes are: Total length, 85; length of tail, 49; length of hind foot, 11.

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

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 -. 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 , , , and

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 on page .

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 .

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 -. 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 and . 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 p, 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 , , , , , , , and

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 -, ), 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 on page .

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 .

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.

Skull and teeth.—Male (based on type specimen and no. 562): See measurements and plates -; weight, not known; basilar length, 43.2 (40.8 and 45.5); zygomatic breadth more or less than distance between condylar foramen and M1 and more than that between anterior palatine foramen and anterior margin of tympanic bulla; mastoid breadth more or less than postpalatal length; postorbital breadth more than length of upper premolars and greater than width of basioccipital, measured from medial margin of one foramen lacerum posterior to its opposite; interorbital breadth more 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 than inside length of P4; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 4 (including I3) 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 and longer or shorter than rostrum; anterior margin of masseteric fossa below or behind m2.

Female (based on no. 564, from Cutervo, Perú, type specimen of Mustela jelskii Taczanowski): See measurements and plates -; weight, not known; basilar length, 38±; zygomatic breadth less than distance between condylar foramen and M1 and not greater than that between anterior palatine foramen and anterior margin of tympanic bulla; postorbital breadth more than alveolar length of upper premolars and (probably) more than width of basioccipital measured from medial margin of one foramen lacerum posterior to its opposite; least width of palate more than inside length of P4; tympanic bulla as far posterior to foramen ovale as width of at least 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.

As compared with that of helleri, the skull of the male of macrura from Junín southward has a lesser mastoid breadth, notably smaller teeth, and a flatter skull which averages lighter throughout. The skulls of females available indicate that the skull and teeth are larger than in agilis.

Remarks.—Seven years after Taczanowski named this subspecies, he applied the name jelskii to a female taken farther north than the original examples of macrura. As indicated in synonymy, various other names have been applied to animals included by the present author in this subspecies.

Mustela frenata macrura intergrades with M. f. affinis as shown by practically all the referred specimens from north of Junín. As one proceeds northward the color of the weasels becomes progressively darker and the teeth become larger until the conditions found in affinis are met with near the northern border of Ecuador. From the material available it appears that the light-colored upper parts found in macrura characterize weasels of, at least, the Temperate Zone, from Marcapata, Perú, to near Quito, Ecuador. West of the range of macrura there exists the still lighter-colored subspecies, M. f. agilis. Immediately adjacent on the north, east, and south, darker-colored weasels occur. So far as color is concerned, the geographic range of the subspecies M. f. macrura is not difficult to define. However, the small size of the teeth characterizes only that part of this light-colored subspecies from Junín southward including the subspecies boliviensis at the southern extremity of the range of the species. From Cutervo northward the light-colored weasels of the Temperate Zone have teeth similar in size to those of the darker, more northern affinis. To designate the slightly larger-toothed, light-colored animals from Ecuador as a subspecies distinct from affinis and macrura is one solution but at present it seems best to refer all of these light-colored animals to macrura.

The type specimen and topotype no. 562 differ more in the amount of inflation of the tympanic bullae than adult males of comparable ages from a given locality usually do. In other respects, the differences between the two skulls are not greater than those ordinarily found in specimens from the same locality. No. 562 has the tympanic bullae greatly, relative to the other South American weasels, inflated posteriorly. Otherwise, the bullae agree with those of the type specimen.

Specimens from southwestern Ecuador, average large, and include the largest specimens of the species Mustela frenata seen from South America. A subadult male, no. 61406, in the American Museum of Natural History, is the largest. Its external measurements are 482, 191, 56. The basilar length of the skull is 48.2 and the zygomatic breadth is 30.3. Although not so large as this specimen, the corresponding measurements of specimens from Alamor, El Chiral, and even from as far away as Sigsig also are distinctly large.

The skull of the female from Ollantaytambo and that of the male from Marcapata have teeth equally as small as do the specimens from Lake Junín.

The skin alone, no. 194328, from Ollantaytambo has the color of the underparts extended over the entire upper sides of the forefeet. The male from Marcapata has less of this color on the forefeet and is in this respect intermediate between the specimens from Lake Junín and the one from Ollantaytambo.

In size of teeth the female, type specimen of M. jelskii, from Cutervo, shows an approach to the larger-toothed weasels of the northern part of the range of macrura.

The specimens in the Riksmuseum from the vicinity of Quito, Ecuador, have been rather fully described by Lönnberg (1921:11-17) and need little comment here, except to say that they show, as he suggested, that the weasel of the Temperate Zone of Ecuador is an intermediate link between M. f. macrura and M. f. affinis.

The adult female and juvenal male labeled as from Ambato have little left of the skulls except some of the teeth and the assignment of the specimens to the subspecies macrura is made mainly on geographic grounds. These two specimens probably are part of the shipment of birds and mammals of which Chapman (1926:703) speaks as follows: "A small collection of native-made skins purchased by the American Museum from a commission merchant in New York City as from 'Ambato' proved to be from the eastern slope of the Andes." Another skin in the same Museum, labeled by a native collector as from "Baeza arriba" [= above Baeza] is so dark colored and has the color of the underparts so much restricted, as to suggest that it belongs to the race aureoventris. Possibly, therefore, it was taken not at Baeza, Ecuador, which I find to the eastward of Quito at 77° 55' W and O° 25' S, but at some place of the same name on the Pacific Slope, unless the locality has been altogether wrongly recorded on the label. If the specimen was taken near the Baeza above referred to, then it gives evidence of an unnamed race of Mustela on the eastern slope of the Andes, characterized by its dark color. Unfortunately the specimen is young and its skull therefore offers insufficient basis for the judging of its subspecific relationships.

Other specimens, in the British Museum of Natural History, recorded as taken "near Quito" and here tentatively listed under macrura, mostly, include specimens so dark colored as to lead me to think they came from country, lower than Quito, adjacent to the range of aureoventris.

Nematodes taken from the right frontal sinus of no. 562 from Junín proved to belong to the superfamily Oxyuriodea according to Professor W. B. Herms and Mr. O. L. Williams, who have independently identified them. Because these worms had been dried fifty-five years in the mounted specimen and were later boiled in cleaning the skull, a more accurate determination was impossible and whether or not they pertain to the same species found in North American weasels cannot be said. Of 18 adult skulls examined for this type of infestation, 13 were found affected as judged by the evident malformation of the frontal region.

Specimens examined.—Total number, 74, arranged by localities from north to south and unless otherwise indicated in the American Museum of Natural History.

Ecuador: Ibarra, 6600 ft., and 7500 ft., 2; San Antonio, 8000-8500 ft., 5 mi. N Quito, 4 (2, 2); Nono, 10000 ft., 1; Mindo, 1; Zambiza, 8000-8100 ft., NE Quito, 4 (2, 2); Carapungo, 8500 ft., NE Quito, 1; Panecillo, 10000 ft., near Quito, 2; Guapulo, 8800 ft., 3 mi. E of Quito, 1; Pichincha, 10500 ft., and 12000 ft., 2 (1, 1); San Ignacio, 11500 ft., Pichincha, 1; Santa Rosa, 9600 ft., Río Pita, 2; near Santa Rosa, 9000 ft., 1; Río San Rafel, 9000 ft., 1; N side Quito, 9000 ft., 1; Quito, 1; near Quito, 5; Nára Papallacta, 11000 ft., 1; below Papallacta, 9000 ft., 1; Chillo Valley, 1; "Hacienda Hda," 10000 ft., Pintag, Valencia, 1; Baeza arriba, 1; Ambato, 2; San Francisco, 8000 ft., E of Ambato, 1; Chunchí, Pagma Forest, 6400 ft., 1; Canar, 2600 M., 1; Malletura, 7600 ft., 1; Contrayerbas, 11000 ft., 1; Sisig, 8500 ft., 3; El Chiral, 1; Almor, 1; Guainche, 3200 ft., 1; no locality more definite than Ecuador, 4; "Received from Quito," 1; Quisaya, 6000 ft. (locality not found), 1; La Carolina (locality not found), 1.

Perú: La Lejía, 1; Huancabamba, 4 (2); Cutervo, 9000 ft., 1; Condechacha, 7000 ft., Río Utcubamba, 1; San Pedro, 8600-9400 ft., S of Chachapoyas, 1; Celendín, 1; Junín, 2; Yana Mayo, Río Tarma, 1; Ollantaytambo, 9000 ft., 3 (1, 2); Ocabamba, 1; Anta Cuzco, 3400 and 3500 M., 2; Marcapata, 1.

Mustela frenata boliviensis Hall

Long-tailed Weasel

Plates , and

Mustela frenata boliviensis Hall, Proc. Biol. Soc. Washington, 51:67, March 18, 1938.

Mustela frenata macrura, Hall, Carnegie Instit. Washington Publ. 473:110, November 20, 1936; Hall, Physis, 16:176, 1939 (part).

Type.—Male, adult, skull and skin; no. 72587, Amer. Mus. Nat. Hist.; Nequejahuira, 8000 feet, Bolivia; May 19, 1926; obtained by G. H. H. Tate; original no. 4135 (see plates -).

Range.—As now known 8000 to 9500 feet in the Andes from Limbaní, Perú, south to Nequejahuira, Bolivia; upper Subtropical and Temperate life-zones. See figure on page .

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 macrura by darker color of upper parts (tone 4 or darker of Carbon Brown, pl. 342 rather than tone 3 of Chocolate, pl. 343, Oberthür and Dauthenay) and lesser size (in males hind foot less than 45 and m1 less than 5.6); from Mustela frenata agilis by darker color of upper parts (as given above rather than tone 2 of Chocolate, pl. 343, of Oberthür and Dauthenay).

Description.—Size.—Male: The type and two young specimens from Limbaní, Perú, measure respectively, as follows: Total length, 383, 368, 304; length of tail, 140, 132, 115; length of hind foot, 43, 44, 41. Tail 55 per cent as long as head and body. Length of hind foot approximately equal to basal length.

Female: Unknown.

Externals.—As described in Mustela frenata macrura.

Color.—Top of head blackish posteriorly to behind ears; upper lips same color as upper parts of head; dark area at angle of mouth not separated from upper parts as a distinct spot; tip of tail black; remainder of upper parts near (n) Mars Brown of Ridgway and tone 4 or darker of Carbon Brown (pl. 342, Oberthür and Dauthenay); underparts Cream-Colored with strong wash of Ochraceous-Buff; whitish on insides of forelegs to just below elbow; color of underparts extended distally on forelegs over ankles onto antipalmar faces of inner toes, and on hind legs to knees. Least width of color of underparts averages 15 (11-19) per cent of greatest width of color of upper parts. Black tip of tail in type longer than hind foot and amounting to 36 per cent of length of tail-vertebrae.

Skull and teeth.—Male (based on the type): See measurements and plates -. As described in Mustela frenata macrura except that: Weight, 2.8 grams; basilar length, 41.6; zygomatic breadth less than distance between anterior palatine foramen and anterior margin of tympanic bulla; anterior margin of tympanic bulla as far posterior to foramen ovale as width of 5 upper incisors.

Female: Skull unknown.

Remarks.—Apparently the first specimens of this race to find their way into a zoölogical collection were the two young males taken on February 17, 1904, at Limbaní, by Geo. Ockenden [sic.].

M. f. boliviensis is smaller than any other South American weasel except possibly agilis. Better material of the two races probably will show even agilis to be larger.

Early in my study of Mustela after examination of the one young specimen, from Limbaní, in the United States National Museum, an account of this race was drawn up, but the account was discarded for want of satisfactory material and the animal was referred to macrura. Then, in 1937, when the two other specimens were studied, the race was formally characterized as different from previously recognized kinds.

The collector has noted on the labels of the two young from Limbaní that they were shot in the afternoon when running together beneath bushes. The frontal sinuses of the type are malformed as a result of infestation by parasites.

Specimens examined.—Total number, 3, as follows:

Perú: Carabaya, Limbaní, 2 (one in U. S. Nat. Mus. and one in Berlin Zool. Mus.).

Bolivia: Nequejahuira, 1 Amer. Mus. Nat. Hist.

Mustela frenata (?) gracilis (Brown)

Plates and

Putorius gracilis Brown, Mem. Amer. Mus. Nat. Hist., 9(pt. 4):182, pl. 17, 1908.

Mustela gracilis, Hay, Iowa Geol. Surv. Bull., 23:32, 1914; Hay, Carnegie Instit. Washington Publ. 322A:252, October 15, 1924; Hay, Carnegie Instit. Washington Publ. 390(vol. 2):528, 1930.

Mustela frenata gracilis, Hall, Carnegie Instit. Washington Publ. 473:112, November 20, 1936.

Type.—Adult skull without lower jaws, probably of a female, no. 12431, Amer. Mus. Nat. Hist.; from Conard Fissure, four miles west of Willcockson, Newton County, Arkansas; obtained sometime in the period 1903 to 1905 inclusive. (See plates and .)

Range.—Known only from the Pleistocene deposit in Conard Fissure, at the type locality in northern Arkansas.

Description.—Skull. Probably female (based on the type): See measurements and plates and ; weight unknown; basilar length, 38.1; least width of palate less than greatest length of P4; tympanic bulla as far posterior to foramen ovale as width of 3 to 5 upper incisors; height of tympanic bulla less than distance from its anterior margin to foramen ovale; length of tympanic bulla less than length of rostrum.

Comparison and remarks.—The type specimen was the only individual referred by Brown (1908) to this species. The remaining material of weasels from this deposit was referred by Brown to his Putorius cicognanii angustidens. Examination of the original materials convinces the writer, too, that the specimens, except no. 12431, are of the species erminea [= cicognanii of Brown]. No. 12431 itself may possibly be erminea but is far more probably of the species frenata. The uncertainty is due to the fact that an occasional skull alone of a subadult male erminea is extremely difficult certainly to distinguish from a skull alone of an adult female frenata. This is true among Recent specimens in the northern Mississippi Valley today; more exactly in Iowa and southern Minnesota the females of frenata, oftentimes intergrades between the subspecies Mustela frenata longicauda, M. f. noveboracensis and M. f. primulina, by only the skulls are next to indistinguishable from certain, unusually slender skulls of male erminea. At other places where the ranges of the two species meet, this difficulty is not so often encountered. Also, the type of gracilis has the skull broken in such a way that the postglenoid length in relation to the length of the skull as a whole could not be accurately determined in this particular skull.

The type specimen of gracilis surely is an adult and because of its small size is thought to be a female. Of known long-tailed weasels of the species frenata, gracilis is structurally nearest to M. f. primulina which occurs in the same region today and to M. f. noveboracensis, the long-tailed weasel of the eastern United States. M. gracilis differs from noveboracensis and agrees with primulina in possessing well-marked temporal ridges which fuse to form a low sagittal crest, in having the mastoid processes projecting farther, laterally, beyond the braincase, in having the anterior ends of the tympanic bullae produced below the squamosal rather than on the same plane with the squamosal, and in having the bullae more inflated anteromedially. M. gracilis differs from both noveboracensis (97 ♂ and 56 ♀ with skulls of comparable age) and primulina (64 ♂ and 24 ♀ with skulls of comparable age) in that the zygomatic breadth amounts to less than 58 per cent of the basilar length. Another difference from any one of the skulls of females of primulina is the longer rostrum, which, when measured from the posterior base of the postorbital process of the frontal to the anterior end of the nasal on the same side, amounts to more than 35 per cent of the basilar length. As pointed out by Brown (1908:182) this specimen represents the extreme of slender skull among known kinds of American weasels.

Selected measurements of no. 12431, the type specimen of Mustela gracilis, are as follows: Basilar length of Hensel, 38.1 mm.; length of upper tooth-rows, 14.3 to 14.4; breadth of rostrum, 11.0; interorbital breadth, 8.5; orbitonasal length, 13.6; mastoid breadth, 18.2; length of tympanic bulla, 13.0; breadth of tympanic bulla, 6.3; depth of tympanic bulla, 3.25; outside length of P4, 4.5; inside length of P4, 4.7; breadth of M1, 3.4; length of inner moiety of M1, 1.8; depth of skull at anterior margin of basioccipital, 12.2; depth of skull at posterior borders of last upper molars, 11.3; distance from foramen ovale to tympanic bulla, 3.6 mm.

Mustela Africana Desmarest

Tropical Weasel

(Synonymy under subspecies)

Type.—Mustela africana Desmarest, Nouv. Dict. d'Hist. Nat., vol. 19, p. 376. 1818.

Range.—Known from the headwaters of the Amazon in eastern Perú and from near the mouth of the same river, on its southern side in Brazil, all within the Tropical Life-zone. See figure on page .

Characters for ready recognition.—Differs from Mustela frenata, the only geographically adjacent species of the genus, by: presence of thenar pad on forefoot; presence of a longitudinal, median, abdominal stripe of same color as upper parts; upper lips being broadly edged, entirely round, with color of underparts; failure of longest facial vibrissae to reach posterior margin of ear; absence of p2; relative flatness (see pl. 29, fig. i and pl. 39, fig. h) of tympanic bullae.

Characters of the species.—Size large (total length of adults approximately 500 mm.); foot-soles naked; thenar pad present on forefoot; length of claws, measured on concave sides, less than one and one-fourth times depth of claws measured at bases; longest facial vibrissae not reaching posterior margin of ear; tail relatively long haired; tail at all ages terminating in point as is characteristic of only juveniles and very young of Mustela frenata and M. erminea; tip of tail, and muzzle, only slightly darker than remainder of upper parts; upper lips broadly edged with color of underparts; pelage coarse, harsh and sparse; longitudinal, median, abdominal stripe of same color as upper parts present; skull broad and deep; braincase large, rounded, and much inflated anteriorly; palatal region wide; tympanic bullae less inflated than in any other American species of the subgenus; angle of lower jaw reduced; dental formula

3 1 2-3 1
-, -, ---, -;
3 1 2 2

teeth heavy; medial lobe of M1 but slightly larger than lateral lobe. See plates , , , and cranial measurements.

Geographic variation.—The reddish versus chocolate color of the upper parts constitutes the only variation of a geographic nature so far detected.

Remarks.—One of the most noteworthy of the several unique characters of this large, tropical weasel is the longitudinal, median, abdominal band. The species exhibits the minimum degree of development of certain features that become progressively less apparent as one proceeds southward from Central America. The relative uniformity of the coloration of the upper parts (reduction in intensity of black color on the muzzle and tip of the tail) and the reduction of the tympanic bullae are two cases in point. Viewed dorsally the general outline of the skull is most nearly matched by that of the skull of Mustela frenata meridana from Venezuela or that of M. f. helleri from Perú. However, the resemblance is not close. The tympanic bullae, although unique among American weasels, are more like those of M. f. meridana from Venezuela than like those of any other kind. The great postorbital width (relatively less in M. africana than in several South American subspecies of Mustela frenata) and small angular process of the mandible are characters, in varying degrees, also common to all South American weasels. Structurally M. africana clearly is more nearly like other subspecies of M. frenata from South America than it is like any species or subspecies from North America.

Mustela africana is the most primitive of the American weasels. The distinctive cranial and dental characters, excepting the reduction in number of premolars, are of a primitive nature. For example, the relatively wide postorbital region, the large braincase that is inflated anteriorly, and the flattened, tympanic bullae, are points of resemblance to the holarctic Mustela erminea, which species is regarded as nearest the original stem form; also the mentioned characters correspond to ontogenetic stages passed through by other weasels. Mostly on these accounts, one is led to look upon M. africana as a migrant from North America. It may have become isolated from its original stock, by a water barrier in the Central American region, for a length of time sufficient to permit of a degree of differentiation to develop between it and the North American weasels which prevented crossbreeding with the frenata stock when that stock, at a later time, reached South America. This assumption is suggested only by evidence from the Recent specimens. No remains of true weasels (subgenus Mustela) have been recorded from deposits in South America older than the Recent period. The alternate possibility, that M. africana intergrades with some race of M. frenata in western or northern South America, has been considered and regarded as highly improbable.

Cabrera (1940:15) has made the distinctive structural characters of Mustela africana basis of the generic name Grammogale to include the one species africana. I am inclined to accord Grammogale only subgeneric rank.

It is possibly significant that Mustela africana is intermediate in several respects between Lyncodon and typical Mustela. The median, longitudinal, abdominal band of the same color as the upper parts in M. africana and the relative uniformity of the coloration of its upper parts might be considered as an intermediate stage between the dark, bicolored (black muzzle and tail tip and brown body) upper parts and light-colored underparts of the North American weasels on the one hand and the light, unicolored upper parts and dark-colored underparts of the Patagonian weasel (Lyncodon) on the other hand. The number of premolars,

2-3
---,
2

is also intermediate between the numbers

3 2
- and -
3 2

of the North American Mustela and the Patagonian Lyncodon, respectively. The American Mustela and the Patagonian Lyncodon, respectively. The more medially, as opposed to anteriorly, directed medial cusp of P4 (characteristic of approximately half of the specimens examined), and the structure of the skull in general of M. africana also seem to be morphologically intermediate between those parts of Mustela and Lyncodon.

On the chance that Lyncodon is closely enough related to Mustela, to be included in a group with Mustela rather than in a group with Grisonella, it is worth noting that Lyncodon lujanensis Ameghino (1889:324, 325), from the villa of Lujan and at the city of Córdoba, at each place in the Pampean [= Pleistocene] formation of Argentine (see also Cabrera, 1928:263) is the first and only fossil form of this group recorded from the whole of South America. Actually, however, Lyncodon seems to me to be as nearly related to Grisonella, if not more so, than to Mustela. If Lyncodon is more closely related to Grisonella and Grison than to Mustela, then the above remarked intermediacy in characters of M. africana has more of interest as a tendency to parallelism than it has of phylogenetic import. Appraisal of phylogenetic relationships would require appraisal of the ancestral stem forms of the Grison stock and the Mustela stock. None of either is known from deposits of the Pliocene, the period of time immediately preceding the Pleistocene.

None of the skulls of Mustela africana seen or figured has the nasomaxillary sutures entirely obliterated and the specimens would, judged on this character alone and by analogy with North American species, be regarded as young and subadult. However, the sutures close at what seems to be a later age than in M. frenata and M. erminea. The condition of the mammae in the type specimen of M. stolzmanni and in the specimen from Moyobamba, indicate that they have borne young. North American weasels old enough to bear young lack visible traces of the nasomaxillary sutures. I have examined no skulls of africana with greatly worn teeth and hence cannot say if the sutures are obliterated in advanced age.

If available data be correct, this species is unique within the genus in that the two sexes are of approximately the same actual size and of the same relative proportions in the body and in the skull. There was no difference between individuals said to be of different sexes from Pará, described and figured by Goeldi (1904:61-62, pls. 1, 2). The undoubted female, type specimen of Mustela africana stolzmanni, is as large as the undoubted male, no. 37475, of the same species, but of a different subspecies, from Pará, Brazil. All the specimens of M. a. africana that I have handled are labeled male and those of M. a. stolzmanni female. More material may show that the female is smaller than the male, as is the case in all near relatives of M. africana.

Little has been recorded concerning the habits of this species. Tate (1931:254) states that a live individual which he saw in a cage at Pará had been captured "swimming in the salt water of the estuary about half a mile away from the shore." On the label of the specimen from Moyobamba, there appears: "caught in Willow tree."

Subspecies examined.—All described forms, of which there are two.

Mustela africana africana Desmarest

Tropical Weasel

Plates , , and

Mustela africana Desmarest, Nouv. Diction. d'Hist. Nat., 19:376, 1818; Cabrera, Bol. Real Soc. Españ. de Hist. Nat., 13:429, November, 1913; Cabrera, Bol. Real Soc. Españ. de Hist. Nat., 14:175, pl. 1, March, 1914.

Putorius (Mustela) brasiliensis paraensis, Goeldi, Zool. Jahrb. abt. f. systematik, geogr. u., Biol., 10:556, pl. 21, September 15, 1897, type from Pará, Brazil, near Pará, Ward of Marco da Legoa, Brazil; Goeldi, Bol. do Mus. Paraense, 3:195 [translation of orig. descr.], August, 1901.

Putorius paraensis, Goeldi, Bol. do Museu Goeldi, 4:61, pls. 1, 2, 1904.

Mustela affinis paraensis, Hollister, Proc. Biol. Soc. Washington, 28:143, July 10, 1914.

Mustela paraensis, Allen, Bull. Amer. Mus. Nat. Hist., 35:105, April 28, 1916; Tate, Journ. Mamm., 12:253, August 24, 1931.

Mustela stolzmanni paraensis, Hall, Carnegie Instit. Washington Publ. 473:111, November 20, 1936; Hall, Physis, 16:167, pl. 1, figs. 1-4, 1939.

Type.—Male, adult or subadult, mounted; no. 848, Paris Museum; from the "Cabinet de Lisbonne 1808," originally from South America as determined from the characters of the animal; probably came from Brazil, and for the present assumed to be from Pará.

On August 25, 1937, the skull was in the mounted skin and the specimen was in the position shown in the figure published by Cabrera (1914, pl. 1). Except for the loss of the distal part of the tail, and fading because of exposure to light, the specimen was in good condition. See also under remarks.

Range.—Known from the south side of the Amazon River, near its mouth at Pará and Cametá, Río Tocantins, in the Tropical Life-zone of Brazil. See figure on page .

Characters for ready recognition.—Differs from Mustela frenata, the only other geographically adjacent species of the genus, in presence of median, longitudinal, abdominal stripe of same color as upper parts and naked foot-soles, in absence of p2 and in reduced size of tympanic bullae (see pls. 29 and 30) and from Mustela africana stolzmanni by lighter color of upper parts which although near Chestnut-Brown are in adults 10' l (darker in yg. M. C. Z., no. 30802), instead of 11' n as in M. a. stolzmanni.

Description.—Size.—This is a relatively large weasel. Goeldi (1897:559) gives the total length of the type specimen of his P. b. paraensis, a female, as 520 mm. (495 in the flesh) and, by computation from his figures, the length of the tail as 200 (205 in the flesh). These measurements probably include the hairs on the tip of the tail as probably also do the measurements given of two other specimens (see Goeldi, 1904:62). One of these specimens, a female, measured: Total length, 520; length of tail, 200. The other specimen, a male, measured: Total length, 510; length of tail, 200. The skin of no. 37475, Amer. Mus. Nat. Hist., a male, has the following measurements written on the attached label: Total length, 548; length of tail, 234; length of hind foot, 56. The hairs project 20 mm. beyond the tip of the last vertebra of the tail and probably are included in the measurements of total length and length of tail. Collectors' measurements of a young male from Cametá, and a subadult labeled as male, from Pará Murutucu, are respectively as follows: 500, 430; 210, 190; 50 and 54.

Externals.—Foot-soles naked, except for a few scattered hairs on ventral sides of interdigital membranes; length of claws, measured on concave sides, not more than one and one-fifth times depth of claws measured at bases; carpal vibrissae not extending beyond apical pad of first digit (not beyond hypothenar pad except in one young specimen); longest facial vibrissae not extending to posterior margin of ear; superior genal tuft not found, hairiness of foot-soles as shown in figure .

Color.—Upper parts near (10 l) Chestnut-Brown and relatively uniform since tip of tail and muzzle are only slightly darker than remainder; underparts with longitudinal stripe of same color as upper parts extending along median line of belly from throat or breast posteriorly to within 40 to 50 millimeters of anus. Underparts otherwise near (20" a) Olive-Ocher (lips and chin whiter in one young specimen). Color of underparts extends distally on median sides of forelegs to bare foot-soles and on median side of hind legs two-thirds of distance from knee to ankle. Upper lips broadly edged with whitish, which color passes posteriorly below and not touching eye to ventral margin of concha of ear. An inverted, basally broad, V-shaped extension passes upward 4 millimeters, just posterior to the eye.

Skull and teeth.—See measurements (plates -). Male: (based on 3 adult and subadult topotypes and figures and descriptions published by Goeldi, 1897 and 1904.) Weight, 7.0 grams; basilar length, 45.8 (44.6-47.8); skull broad and deep; braincase large, rounded, and much inflated anteriorly; distance from postorbital process to anterior, nasal notch approximately equal to breadth across exoccipital condyles; palatal region wide; tympanic bullae less inflated than in any other species; mastoid bone, laterally, concave; length of upper tooth-rows in adults and subadults less than breadth of palate measured between two outer margins of fourth upper premolars; alveolar distance between C1 and P4 less than length of P4; teeth heavy; medial lobe of M1 only slightly larger than lateral lobe; deuterocone of P4 heavy and often inclined mesially; p2 absent (P2 present above on both sides in only one of seven specimens seen or described); lower jaw heavy; masseteric fossa not extending anteriorly to posterior fourth of talonid of m1; paraconid of m1 low and base of cleft between it and protoconid relatively low on tooth.

Female: No skull examined but from figures published by Goeldi (1904, pl. 2), apparently as described in the male.

Remarks.—Desmarest in 1818 gave a remarkably good description of this animal which he named as a new species, Mustela africana, but mistakenly indicated that the single specimen known to him came originally from Africa. Until 1913 the name was applied, wrongly, to weasels of northern Africa or to those of the Azores Islands and St. Thomas Island. In that year Cabrera (1913:429) identified the species with the one later named Putorius (Mustela) brasiliensis paraensis by Goeldi (1897:556, pl. 21) from Pará, Brazil. Despite Cabrera's clear identification in 1913, and his later mention of the correct application of the name Mustela africana, it was not correctly employed by other authors, including myself who even as late as 1936 (p. 111) instead used Goeldi's name. In 1937 Mr. Cabrera called my attention to his published account of Mustela africana and so permitted me to examine the type specimen in the Paris Museum, whither I was bound when I received Mr. Cabrera's letter. My own examination of the specimen fully confirmed the conclusions published by Cabrera (1913:429).

As a matter of historical interest, however, it is worth noting that Cabrera (1913) originally supposed the type specimen to have been taken as booty of war from Portugal by the French and that Cabrera later, at the request of P. Trouessart, pointed out (1914:176) that the specimen had been acquired in exchange ("a cambio") since according to Dr. Trouessart the Museum register showed that offer had been made to Portugal to return this and other specimens but that Portugal had replied that it had nothing to reclaim. Dr. P. Rode in August, 1937, at the Paris Museum, gave it to me as his opinion that the specimen had been an outright gift from the "Cabinet de Lisbonne" to E. Geoffroy St.-Hilaire on his trip to Portugal in 1808 when he was given also from the same cabinet several primates, all from Brazil. Of the labels attached to the pedestal on which the specimen is mounted, that of most ancient appearance is glued to the bottom of the stand and bears in a hand apparently written before Trouessart's entries on the same label, the information "Du Cabinet de Lisbonne 1808" and "J. H. S. 1809."

The opened mouth of the mounted specimen permits one to determine that P2 is absent on each side above. The stuffed scrotal pouch and hair projecting downward about the preputial opening clearly show the animal to have been a male. The least faded portions of the mounted specimen, its sides, are of the same reddish color as characterizes adults from Pará and not of the darker chocolate color of specimens of M. stolzmanni from Perú. The specimen is indistinguishable from topotypes of P. paraensis of Goeldi and his name will have to fall as a synonym of Mustela africana Desmarest.

Goeldi gave an extended description, with figures of the skull, head, and entire animal, when he named paraensis. As his account shows, he was unaware that Taczanowski had described a similar weasel from the headwaters of the Amazon, or for that matter that any weasel excepting Mustela affinis Gray, had been found in South America. Goeldi's later account of additional specimens (1904:61, pls. 1, 2) gives much useful information about the animal. Photographs of several specimens and photographs and detailed measurements of several skulls are presented by him.

Pará, and Cametá, Brazil, places from which Mustela africana africana is known, are nearly 2000 miles from the localities in eastern Perú and eastern Ecuador from which M. a. stolzmanni is known, and no specimens, from intermediate localities, are available to show actual intergradation of the two. However, the similarity in structure of the two weasels is so great as to indicate close affinity. Furthermore, it is understood that environmental conditions at and between the two localities are similar. These considerations, in the light of our knowledge of actual intergradation of geographic races of weasels in other places, cause me to treat, with a feeling of assurance, M. africana [= P. paraensis Goeldi] and M. stolzmanni Taczanowski as subspecies of a single species. M. Rodolpho Legueira Rodríguez writes me, under date of June 16, 1928, that the type specimen of Putorius (Mustela) brasiliensis paraensis Goeldi is stuffed and preserved in a "vitrine" at the Museum Goeldi (Museum Paraense) De Historia Natural e Ethnographia, Pará, Brazil.

The one young specimen seen, that from Cametá, is darker colored than any of the four older specimens examined. It is almost exactly the Chestnut Brown of Ridgway (1912) and therefore approaches closely in color the adult specimens of M. a. stolzmanni. This same tendency to greater richness of color in young than in adults is seen also in Mustela frenata.

Specimens examined.—Total number, 5, all from eastern Brazil, as follows: Pará, 2 (1, 1); Pará Murutucu, 1; Río Tocantins, Cametá, 1; type specimen, 1.

Mustela africana stolzmanni Taczanowski

Tropical Weasel

Plates and

Mustela stolzmanni Taczanowski, Proc. Zoöl. Soc. London, 1881:835, November 15, 1881; Allen, Bull. Amer. Mus. Nat. Hist., 35:105, April 28, 1916.

Mustela stolzmanni stolzmanni, Hall, Carnegie Instit. Washington Publ. 473:111, November 20, 1936; Hall, Physis, 16:167, pl. 1, figs. 5, 6, 1939.

Type.—Female, adult, mounted skin, with skull separate; no. 563, Mus. Polonais d'Hist. Nat. (Warsaw, Poland); Yurimaguas, Perú; 1880; obtained by J. Stolzmann.

The skull (plate ), mounted with the skin but removed by me for study, consists of the premaxillae, maxillae, two halves of the lower jaw and dentition. Of these parts, right m2, left coronoid process, right P4 and M1 and adjacent part of maxilla are lost. The skin is well mounted, in a good state of preservation and shows no fading due to exposure to light. Inguinal mammae are distinctly shown on the skin and prove that the specimen is a female. Except for a few scattered hairs on the lower throat, a spot six by eight millimeters on the medial side of the region of the olecranon of the left foreleg and another of similar size in the left axilla, the underparts are, excepting the ventral longitudinal, abdominal stripe, unmarked by color of the upper parts.

Range.—Known from the Tropical Life-zone of eastern Ecuador and Perú from Jatun Yacu south to Valle del Perené. See figure on page .

Characters for ready recognition.—Differs from Mustela frenata, the only other geographically adjacent species of the genus, in presence of median, longitudinal, abdominal stripe of same color as upper parts and naked foot soles, in absence of p2 and in reduced size of tympanic bullae (see pls. 28, 29, 30, 39 and 40) and from Mustela africana africana by darker color of upper parts which, although near Chestnut Brown, are 11' n instead of 10' l as in M. a. africana.

Description.—Size.—Male: unknown.

Female: Taczanowski (1881:836) gives, among others, the following measurements of the type specimen: Total length, 523, length of body, 260; length of tail without hair, 190 (with hair 224); length of hind foot, 54. Whether or not the measurements were taken from the animal when in the flesh I do not know. Allowing for shrinkage of hind feet and changes due to the posture of the now mounted specimen, I get from it essentially the same measurements. Collectors' measurements of a subadult from Moyobamba and a young female from Valle del Perené, are respectively, as follows: 469, 415; 184, 160; 57, 52. My own measurements of the dry hind feet on the skins are respectively, 48 and 49.

Externals.—As described in M. a. africana except that the length of the concave sides of the claws are approximately one and one-fourth times the depth; thus the claws are relatively longer than in M. a. africana.

Color.—As described in M. a. africana with the following noted exceptions: Upper parts near (11' n) Chestnut-Brown; area of lighter ventral coloration on the throat and sides of head less strongly tinged with yellow; pelage more dense, finer and softer than in M. a. africana.

Skull and teeth.—Male: Skull unknown.

Female: See measurements and plate and . As described in male of Mustela africana africana except that: Weight, 4.7 grams. As contrasted with M. a. africana, the dentition of the lower jaw is lighter; the transverse diameter of m2 is 1.2 mm. in the type and also in the specimen from Moyobamba as against 1.5 to 1.7 in three male topotypes of M. a. africana.

Remarks.—After the Polish naturalist, Stolzmann, in the course of his explorations in Perú, obtained the single specimen which was made the type, no other naturalist, so far as known, visited the type locality until thirty-two years later when Wilfred H. Osgood and M. P. Anderson spent more than a month collecting at Yurimaguas (see Osgood, 1914:147), but secured no topotypes of this little-known weasel. C. O. Schunke took the second specimen in the Valle del Perené in April, 1921; L. Rutter on January 25, 1924, took the third specimen, and W. Clark-MacIntyre took the fourth specimen on the Jatun Yacu. This obscure place name is shown on the map (fig. 4, p. 827) published by Brown (1941) and is the stream flowing from the west to the town of Napo. Napo is situated at approximately 1° 2' S and 77° 49' W.

In the female from Moyobamba there are only 3 pairs of mammae. One pair is inguinal and two pairs are on the posterior part of the abdomen.

Taczanowski (1881:836) relates that this species was taken in the forest to which it appears to be restricted since the inhabitants of the village did not know of the animal. He points out also that the previously known Peruvian species [M. f. macrura and M. f. agilis] live in the treeless territory of eight to eleven thousand feet altitude whereas M. stolzmanni was found in the humid forest of the great plain of the Maynas at an elevation of 500 feet or less above sea level. The frontal sinuses of the specimens seen reveal no malformation as a result of infestation by parasites.

Specimens examined.—Total number, 4, as follows:

Ecuador: R. Tatun [= Jatun] = Yacu, 1, Mus. Comp. Zoöl.

Perú: Yurimaguas, 1 in Mus. Polonais d'Hist. Naturelle, Warsaw; Moyobamba, 2700 ft. [6° S, 77° W], 1 in Brit. Mus. Nat. Hist.; Valle del Perené, 1200 meters, 1 in Amer. Mus. Nat. Hist.

EXPLANATION OF CRANIAL MEASUREMENTS APPEARS ON PAGE 417