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.
_Skull and teeth._--Male (based on type specimen and no. 562): See
measurements and plates 27-30; 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 37-40;
weight, not known; basilar length, 38±; zygomatic breadth less
than distance between condylar foramen and Ml 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[7]; San Antonio,
8000-8500 ft., 5 mi. N Quito, 4 (2[7], 2[78]); Nono, 10000 ft., 1;
Mindo, 1[78]; Zambiza, 8000-8100 ft., NE Quito, 4 (2[78], 2[95]);
Carapungo, 8500 ft, NE Quito, 1[78]; Panecillo, 10000 ft., near
Quito, 2[78]; Guapulo, 8800 ft., 3 mi. E of Quito, 1[78];
Pichincha, 10500 ft., and 12000 ft., 2 (1[78], 1[95]); 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[78]; Quito, 1[4]; near Quito, 5[7]; Nára
Papallacta, 11000 ft., 1[78]; below Papallacta, 9000 ft., 1[78];
Chillo Valley, 1[78]; "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[1]; Canar, 2600 M.,
1[7]; Malletura, 7600 ft., 1; Contrayerbas, 11000 ft., 1; Sisig,
8500 ft., 3[7]; El Chiral, 1; Almor, 1; Guainche, 3200 ft., 1; no
locality more definite than Ecuador, 4[95]; "Received from Quito,"
1[7]; Quisaya, 6000 ft. (locality not found), 1[7]; La Carolina
(locality not found), 1[78].
=Perú=: La Lejía, 1; Huancabamba, 4 (2[75]); Cutervo, 9000 ft.,
1[73]; Condechacha, 7000 ft., Río Utcubamba, 1[7]; San Pedro,
8600-9400 ft., S of Chachapoyas, 1; Celendín, 1[7]; Junín, 2[73];
Yana Mayo, Río Tarma, 1[7]; Ollantaytambo, 9000 ft., 3 (1[7],
2[91]); Ocabamba, 1[7]; Anta Cuzco, 3400 and 3500 M., 2[4];
Marcapata, 1[91].
=Mustela frenata boliviensis= Hall
Long-tailed Weasel
Plates 28, 29 and 30
_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 28-30).
_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 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 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_.