existence of one or more additional subspecies since animals from each
of the three localities now represented differ from those from the
other two and some of these differences are correlated with geographic
position. However, specimens from all three localities agree in
several features. For example all of them have the dorsal outline of
the skull highly convex, transversely, and, more especially,
longitudinally. In this respect they are sharply differentiated from
any other American weasel. Nevertheless, _M. f. panamensis_ is clearly
a link between the North and South American subspecies and _panamensis_
intergrades with the adjacent subspecies. The large size of the skull
and teeth and the slightly more ventrally projected tympanic bullae of
no. 10112 from Boquete approach features seen in _M. f. costaricensis_.
The smaller size of skull and teeth of no. 178970 from Mt. Pirre are
points of resemblance to _M. f. meridana_.
The type specimen was selected from a region where _M. f. panamensis_
is thought to have its distinctive characters well developed. The
specimen is not adult and, therefore, does not show as many
differential characters as does a nontypical adult from Boquete.
Nevertheless, the majority of the above mentioned differential
characters are shown by the type specimen and an adult from the same
place would, it is judged, show all the differential characters better
than would an adult from Boquete.
Of the 11 skulls examined, 6 show no signs of having had the frontal
sinuses infested with parasites.
_Specimens examined._--Total number, 19, listed by localities from
north to south and unless otherwise indicated in the United States
National Museum.
=Panamá=: Boquete, 10 (3[75], 1[8], 1[2], 3[4], 1[7]); Río Gariche
[é], 5300 ft., 1[1]; Siola, 1[1]; Chiriquí, 1[7]; Río Indio, near
Gatún, 1; Mt. Pirre, 3 (2[1]); Calovébora, 1[7] (locality not
found, possibly misspelling of Calovébora); no locality more
definite than Panamá, 1[4].
=Mustela frenata meridana= Hollister
Long-tailed Weasel
Plates 25, 26, 27, 37, 38 and 39
_Mustela meridana_ Hollister, Proc. Biol. Soc. Washington, 28:143,
July 10, 1914.
_Putorius affinis_, Robinson and Lyon, Proc. U. S. Nat. Mus.,
24:147, October, 1901; Allen, Bull. Amer. Mus. Nat. Hist.,
30:256, December 2, 1911.
_Mustela affinis_, Osgood, Field Mus. Nat. Hist. Publ. 155, zoöl.
ser. 10:61, January 10, 1912.
_Putorius macrurus_, Allen, Bull. Amer. Mus. Nat. Hist., 31:92,
April 19, 1912.
_Mustela affinis affinis_, Allen, Bull. Amer. Mus. Nat. Hist.,
35:100, April 28, 1916 (part).
_Mustela affinis costaricensis_, Allen, Bull. Amer. Mus. Nat.
Hist., 35:101, April 28, 1916 (part).
_Mustela frenata meridana_, Hall, Carnegie Instit. Washington Publ.
473:110, November 20, 1936; Hall, Physis, 16:175, 1939.
_Type._--Male, subadult, skull and skin; no. 123341, U. S. Nat.
Mus., 1630 meters elevation, Montes de Mérida, near Mérida,
Venezuela; August 14, 1903; obtained by S. Briceno.
The skull (plates 25 and 26) lacks the right exoccipital condyle
and posterior half of the right zygomatic arch. The teeth all are
present, unworn and entire. The skin is well made and complete.
_Range._--Near sea level (San Julián) to 8500 feet (Montes de
Culata, Mérida, Venezuela), and 9000 feet (Santa Elena, Colombia).
Temperate to Subtropical life-zones of Venezuela and northern and
western Colombia. 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 _M. frenata panamensis_ in
lighter color of upper parts (tone 3 rather than tone 4, pl. 344,
Reddish Black, Oberthür and Dauthenay), flat rather than convex
dorsal outline of skull immediately behind postorbital processes
(see pl. 27); from _M. f. affinis_, in males, by lesser average
breadth and length of skull and greater actual and relative size
(see measurements) of facial part of skull; from _M. f.
aureoventris_, in males, by lighter upper parts (tone 3 rather
than tone 4, pl. 344, Reddish Black, Oberthür and Dauthenay) and
by smaller skull and teeth (basilar length less than 45, length of
m1 less than 6.3, width of M1 less than 4.8, outside length of P4
less than 5.7).
_Description._--_Size._--Male: Average and extreme measurements of
topotypes (as recorded by collectors on labels, and so uniform as
to show them not to be accurate to within more than 5 mm.) are as
follows: Total length, 434 (410-460); length of tail, 164
(150-180); length of hind foot, 50 (no variation in collectors'
measurements). Tail averages 61 per cent as long as head and body.
Length of hind foot more than basal length. Corresponding
measurements of no. 22191, a young male from Mérida, measured by
Osgood or Conover, are 439, 165, 54. The adult male no. 18703,
from Páramo de Tama (eastern boundary of Venezuela) has the
following measurements written on the label by Osgood: 404, 150,
47.
Female: Average and extreme measurements of topotypes (as recorded
by collectors on labels and so uniform as to show them not to be
accurate to within more than 5 mm.) are as follows: Total length,
347 (320-370); length of tail, 128 (120-130); length of hind foot,
40 (no variation in collectors' measurements). Tail averages 57
per cent as long as head and body. Length of hind foot more than
basal length. Two females, adult no. 11034 and young no. 11033
from Cincinnati, Santa Marta, Colombia, measured by M. A.
Carriker, Jr., measure, respectively, as follows: 371, 330; 140,
140; 38, 36. No. 14463, adult, from Río Zapata, Colombia, measured
(by J. H. Batty), 315, 138, 39. No. 32182, adult, from Mira
Flores, Cauca, Colombia, measured (by W. B. Richardson), 375, 150,
43.
The average differences in external measurements of the two sexes,
at Mérida, are: Total length, 87; length of tail, 36; length of
hind foot, 10.
_Externals._--Longest facial vibrissae black (few rarely white)
and extending beyond ear; carpal vibrissae colored like underparts
or upper parts, and not extending beyond apical pad of fifth
digit; hairiness of foot soles slightly greater than shown in
figure 21.
_Color._--As described in _Mustela frenata panamensis_ except
that: Posterior fourth of each upper lip rarely, and small spot in
front of ear usually, white; black of head proper not extending
back of ears and grading insensibly into color of upper parts;
anterior half of upper parts of adults "frosted" with numerous
white hairs (tick bites?), upper parts near (_n_) Bay or tone 2 of
Reddish Black (pl. 344, Oberthür and Dauthenay) or tone 3 in
freshest, unfaded pelage. Least width of color of underparts (in
ten males from Mérida) 20 (17-23) per cent of greatest width of
color of upper parts. Black tip of tail, in same series, 60 to 75
mm. long, thus longer than hind foot and 41 (40-44) per cent as
long as tail-vertebrae.
_Skull and teeth._--Male (based on type specimen and seven
topotypes, five adults and three subadults): See measurements and
plates 25-27; weight, 4.1 (3.8-4.3) grams; basilar length, 43.6
(42.3-44.3); zygomatic breadth more than distance between condylar
foramen and M1 or than between anterior palatine foramen and
anterior margin of tympanic bulla; mastoid breadth less than
postpalatal length; postorbital breadth greater than length of
upper premolars or than width of basioccipital measured from
medial margin of one foramen lacerum posterior to its opposite;
interorbital breadth not less than distance between foramen
opticum and anterior margin of tympanic bulla; breadth of rostrum
greater than length of tympanic bulla; least width of palate
greater than length of P4; anterior margin of tympanic bulla as
far posterior to foramen ovale as width of 4 to 5 upper incisors;
height of tympanic bulla less than distance from its anterior
margin to foramen ovale; length of tympanic bulla more or less
than (approximately equal to) alveolar length of lower molar and
premolar tooth-row and shorter than rostrum; anterior margin of
masseteric fossa posterior to m2 and confined to posterior third
(34 per cent average, 32 minimum, 37 maximum) of mandible.
Female (based on four adult topotypes): See measurements and
plates 37-39; weight (no. 143665), 2.3 grams; basilar length 37.2
(36.3-38.2); zygomatic breadth more or less than distance between
condylar foramen and M1 or than distance between anterior palatine
foramen and anterior margin of tympanic bulla; postorbital breadth
(sinuses badly infested with parasites) more than length of upper
premolars or width of basioccipital measured from medial margin of
one foramen lacerum posterior to its opposite; least width of
palate more than length of P4; tympanic bulla as far posterior to
foramen ovale as combined width of 4 to 5 upper incisors; height
of tympanic bulla less than (one half to three fourths of)
distance from its anterior margin to foramen ovale; length of
tympanic bulla more than length of lower molar and premolar
tooth-row.
The skull of the female is 44 per cent lighter than that of the
average male.
Comparisons of the skull with those of _M. f. panamensis_ and _affinis_
have been made in the accounts of those subspecies. As compared with
the skull of the male of _M. f. aureoventris_, that of _meridana_
averages smaller in every measurement taken. Indeed, none of the skulls
of _meridana_ equals that of _aureoventris_ in basilar length, length
of tooth-rows, length of tympanic bulla, depth of skull at anterior
margin of basioccipital or at posterior margin of upper molars, or
measurements of teeth. Relative to the basilar length, most of the
measurements are greater in _meridana_. Exceptions are the relative
length of the tooth-rows, and the two measurements of depth of the
skull which average less.
_Remarks._--In 1914 when Hollister named this weasel he compared it
with _M. f. affinis_ and most of the differential characters which he
ascribed to _meridana_ were merely "more than" or "less than" in
_affinis_. In _affinis_, Hollister included specimens from Chiriquí,
Panamá, and the coast of Venezuela. The specimens from these three
places were referred by Allen (1916:101) to _Mustela affinis
costaricensis_, and he restricted (_op. cit._:100) the type locality of
_Mustela affinis affinis_ to Bogotá, Colombia, and synonymized _Mustela
meridana_ with _M. a. affinis_. Hollister probably would not have named
_meridana_ had he had specimens from Bogotá for comparison and had he
regarded them as topotypes of _affinis_ for the difference is slight.
Nevertheless, within the large geographic range of _M. f. meridana_
there is some geographic variation. There is more of such variation in
the color of the pelage than in shape and size of the skull. The
specimen from San Julián is darker than the average and in this respect
approaches true _panamensis_. San Julián is situated at a relatively
low elevation on the coast of Venezuela.
_M. f. meridana_ so closely resembles _M. f. affinis_ that the writer
has no quarrel with anyone who would synonymize _meridana_. However, as
represented by topotypes, the two races unquestionably are, _on the
average_, different, and specimens from the southeastern part of the
range of _affinis_ probably are individually distinguishable from
topotypes of _meridana_.
Variation in the skulls of the series from Mérida is relatively small.
This applies to both males and females. The external measurements
recorded by native collectors are not accurate to within more than five
millimeters but, considering this, variation in external measurements
also seems to be slight. The difference in size of the two sexes
appears to be uniformly greater than in weasels from Central America.
The twenty-six topotypes show that the color and color pattern are
relatively uniform. All are of nearly the same tone except juveniles or
young which are, as in the case of _panamensis_, much brighter colored
on the underparts. Also, the young have darker-colored upper parts. The
adults, without exception, have numerous white hairs scattered over the
back of the head, neck and between the shoulders. I have no
trustworthy evidence to support the suggestion that these white hairs
are the results of tick bites or that they are caused by other
parasites which damage the hair follicles. The white facial markings
vary relatively little in the 45 specimens carefully examined in this
regard. Also, the variation in color pattern of the two sides of the
head is small. Indeed, within rather narrow limits, the color of the
two sides of the head is the same in every specimen except two. In
these two the white spots anterior to the ears are confluent with the
color of the underparts. Only one specimen, no. 21342, has a white spot
between the eyes and this spot is small. Ten of the twenty-six
specimens have a definite white spot or band in front of each ear. Two
specimens have such a spot on one side only. The dark spots at the
angles of the mouth are present on two sides in three specimens and on
one side only in three others. The mentioned spots are, then, present
nine out of a possible fifty-two times. When the spots are absent, dark
color usually is present in the required area but is confluent with the
color of the upper parts.
A young male from San Julián, Robinson and Lyon (1901:147) state ". . .
was shot . . . as it ran over some bowlders in a ravine. Its eyes shone
with the same greenish light as do the eyes of our common weasel, and
it emitted the same strong odor." No. 14463, Am. Mus. Nat. Hist., from
Río Zapata, Colombia, according to data on the label, was "taken in
timber belt in valley in balk hills" and the native name is Cosonebi.
Two specimens taken on the Páramo de Tama, head of Tachira River,
Venezuela and Colombia are commented on by Osgood (1912:61) as follows:
"One . . . was caught in a steel trap baited with birds and set by the
side of a rushing mountain stream. . . . The other was shot in midday
as it came prowling about our 'house' in the clearing. . . ."
Of the thirty-three skulls before me, twelve have the frontal sinuses
malformed by parasites. These twelve include most of the adults for few
of the subadults and fewer of the young show pathologic conditions in
the frontal region.
_Note on localities._--Several of the localities in Colombia mentioned
in "Specimens examined" are described and located by Chapman
(1917:640-656, pl. 41) in his "Distribution of Bird-life in Colombia."
Place names for Colombia on labels, not found on any map, or duplicated
names of which I can not certainly select one, are Río Barrotow, Río
Oscuro, Río Zapata, Río Japata, Guasca and El Baldro. Sonson may or
may not be the town of that name situated some eighty miles northwest
of Bogotá and on the east flank of the Central Andes west of the
Magdalena River on the drainage of the Cauca River. In Venezuela most
of the specimens from Mérida are labeled 1630 meters, Montes de Mérida.
San Julián is some seven miles east of La Guaira (see Robinson and
Lyon, 1901:136). San Esteban is located a little way back from the
coast between Puerto Cabello and Valencia. Páramo de Tama is on the
Venezuelan-Colombian border near the source of the Tachira River (see
Osgood, 1912:35). Mt. Duida is shown as at 3° 30´ N and 65° 40´ W by
Chapman (1931:13) and Mt. Auyán-tepui as near 5° 15´ N and 62° 50´ W by
Chapman (1937:760).
_Specimens examined._--Total number, 78, arranged by localities
from north to south and unless otherwise indicated in the British
Museum of Natural History.
=Venezuela=: San Julián, 1[91]; Carácas, 2; Galipare, Cerro del
Avila, 6500 feet, 1; San Esteban, 1[2]; Mérida, 45 (10[91], 14[2],
10[4], 2[60], 2[14], 1[78]); Páramo de Tama, 1[60].
=Colombia=: Páramo de Tama, 1[60]; Cincinnati, 3[9]; Valdiva, 3800
ft., 1; Medellín, 2; 7200 ft., Barro Blanco, 1[2]; Santa Elena,
9000 ft., 1[2]; Santa Elena, 1[2]; Sonson, 2 (1[91], 1[2]); Mt.
Auyan-tepui, 1[2]; Pueblo Rico, 5200 ft., 1[91]; Mira Flores,
1[2]; Jerico, near Cauca River, 1; Tornel, 20 mi. NE Quitichao, 1;
Mt. Duida, 1[2]; El Tambo, Cauca, 1[78]; El Baldro, 1[2]; Río
Japata, 2[2]; Río Zapata, 4500 ft., 1; Río Oscuro, 3300 ft., 1;
Río Barrotow, 3300 ft., 1; Guasca, 1[75]; no locality more
definite than Colombia, 1.
=Mustela frenata affinis= Gray
Long-tailed Weasel
Plate 30
_Mustela affinis_ Gray, Ann. and Mag. Nat. Hist., 14(ser. 4):375,
1874.
_Putorius (Gale) brasiliensis frenatus_, Coues, Fur-bearing
animals, p. 142, 1877 (part).
_Putorius affinis_, Merriam, N. Amer. Fauna, 11:31, June 30, 1896.
_Mustela affinis affinis_, Allen, Bull. Amer. Mus. Nat. Hist.,
35:100, April 28, 1916; Allen, Bull. Amer. Mus. Nat. Hist.,
35:220, May 31, 1916.
_Mustela frenata affinis_, Hall, Carnegie Instit. Washington Publ.
473:110, November 20, 1936; Hall, Physis, 16:175, 1939.
_Type._--Male, adult, skull with skin; no. 54.1.11.3 (skull
originally numbered 195d, later 54.6.3.4), Brit. Mus. Nat. Hist.;
Colombia [given as new Granada in original description]; purchased
from Mr. S. Stevens. Type locality restricted by Allen (1916:99)
to Bogotá, Colombia.
The skin is in a good state of preservation and has been made over
into a conventional study specimen from a mount on exhibition.
Exposure to light when mounted probably accounts for the faded
color. The skull (plate 30) lacks the middle 9 mm. of the right
zygomatic arch, occiput, basioccipital and posterior two-thirds
of the left tympanic bulla. The teeth all are present and entire.
_Range._--Four thousand six hundred feet (Quetame) to 9154 feet
(El Carmen), Tropical to Temperate life-zones of eastern Andes of
Colombia. 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, by presence of p2 and by two
roots rather than one root on P2; from _M. frenata meridana_, in
case of males, by, on average, greater breadth and length of skull
and lesser actual and relative size (see measurements) of facial
part of skull; from _M. f. aureoventris_ by lighter-colored upper
parts (tone 2 rather than tone 4, pl. 344, Reddish Black of
Oberthür and Dauthenay); from _M. f. macrura_ by darker color
(Reddish Black, tone 2, pl. 344, Ober. and Dauth., rather than
Chocolate, tone 3, pl. 343, Ober. and Dauth.).
_Description._--_Size._--Male: Measurements in life, estimated
from dried skins, are: Total length, 455; length of tail, 175;
length of hind foot, 52. Proportions of parts supposedly as
described in _Mustela frenata meridana_.
Female: Estimates from two dried skins: Total length, 365; length
of tail, 135; length of hind foot, 43. Proportions of parts
supposedly as described in _Mustela frenata meridana_.
The estimated differences in external measurements of the two
sexes are: Total length, 90; length of tail, 40; length of hind
foot, 9.
_Externals._--As described in _Mustela frenata meridana_.
_Color._--As described in _Mustela frenata panamensis_ except
that: posterior fourth of each upper lip and spot in front of each
ear white in approximately half of the specimens; black of head
proper not extending back of ears and grading insensibly into
color of upper parts; upper parts near (_n_) Bay, or tone 2 of
Reddish Black (pl. 344, Oberthür and Dauthenay). Least width of
color of underparts (in five males from vicinity of Bogotá) 24
(15-29) per cent of greatest width of color of upper parts. Black
tip of tail, in same series, 60 to 75 mm. long, thus longer than
hind foot and averaging 38 per cent as long as tail-vertebrae.
_Skull and teeth._--Male (based on three adults and two subadult
topotypes): See measurements and plate 30. As described in
_Mustela frenata meridana_ except that: Weight, 4.5 grams
(estimated); basilar length 45.8±; interorbital breadth not
greater than distance between foramen opticum and anterior margin
of tympanic bulla (type as in _meridana_ where interorbital
breadth is more than distance between foramen opticum and anterior
margin of tympanic bulla); least width of palate not less than
length of P4; masseteric fossa confined to posterior two-fifths
(38 to 40 per cent; average 39 per cent) of mandible and not
extended anteriorly to middle of m2.
Female: No adults examined.
As compared with _M. f. meridana_ the skull of the male is larger, to
the average amount of 2.2 mm. in basilar length and 1.2 mm. in
zygomatic breadth of adults; length of tooth-rows and mastoid breadth
average greater but relatively less; breadth of rostrum, interorbital
breadth and orbitonasal length average actually and relatively less.
Thus the skull of _affinis_ is longer and broader, but the facial
region is actually, as well as relatively, smaller. As compared with
the skull of the male of _M. f. aureoventris_, that of _M. f. affinis_
is about the same in basilar length. However, in no specimen of
_affinis_ are the measurements of length of tooth-rows or breadth of
rostrum, actually, or relatively, as great as in _aureoventris_. The
same is true of all measurements taken of M1, P4 and m1. The specimens
from the vicinity of Quito and north of there, although referred to
_macrura_, are nearly as dark as typical _affinis_, approach _affinis_
in cranial characters, and indicate intergradation of _affinis_ with
_macrura_.
_Remarks._--_Mustela affinis_ was named by John Edward Gray in 1874 (p.
375) on the basis of a specimen from New Granada. Although usually
synonymized with _Mustela brasiliensis_ by later authors until 1896
when Merriam (1896:31) applied the name to weasels from Costa Rica,
nearly all the South American and several of the Central American
weasels have, at one time or another, had Gray's name, _affinis_,
applied to them. Gray, in 1865 (p. 115) when giving measurements of
_Mustela aureoventris_, probably mentioned the specimen, that later
became the holotype. In 1916 (p. 98) Allen restricted the type locality
to Bogotá, Colombia. Allen's action was a necessary procedure in
clearing up the systematics of South American weasels and was based on
good grounds. As set forth by Allen (_loc. cit._), and more in detail
by Chapman (1917:642), Bogotá has long been the shipping point for
Colombian vertebrate specimens, many of which were obtained in the
mountains to the east. Allen (1916A:220) quotes Thomas as saying that
the type specimen was purchased from Stevens at about the same time
that a number of Colombian birds were purchased from the same dealer.
Also, specimens from Bogotá agree with Gray's description of the type
specimen.
_Mustela frenata affinis_, as here defined, constitutes one of the
several slight geographic variants met with, on the sides of, and
between, the three north and south mountain chains of Colombia. The
others are lumped under the name _Mustela frenata meridana_. _M. f.
affinis_, in common with specimens from the northern part of the range
of _macrura_ has large teeth. Weasels of all of the region from Quito
to Bogotá have large teeth. To the north there is the smaller-toothed
_meridana_ and to the south the smaller-toothed _macrura_ grading into
the still smaller-toothed _agilis_, and _boliviensis_.
Two skins, without corresponding skulls, from Caqueta are lighter
colored than any others of _affinis_; possibly the skins are faded by
exposure to light. Since they probably come from an elevation of less
than 1000 feet in the Amazonian region, they may pertain to another
subspecies.
Complete, unbroken, skulls of _affinis_ are needed to ascertain the
degree to which _affinis_ and _meridana_ differ in cranial features.
The several specimens from the immediate region of Bogotá show well the
color and the color pattern but lack collectors' measurements.
None of the ten skulls examined shows malformation of the frontal
region due to infestation of the frontal sinuses by parasites. Possibly
three of the four adults were infested, although not severely.
_Specimens examined._--Total number, 27, arranged by localities
from north to south and unless otherwise indicated in the United
States National Museum.
=Colombia=: El Carmen, 1[2]; W. Cundinamarca, 1[7]; Muzzo [=
Muzo?], 1[4]; Bogotá, 1; Castillo, near Bogotá, 1[7]; Fambrias,
near Bogotá, 1[75]; Bogotá district, 1[2]; Choachí, 9 (1[75],
2[7], 1[84]); Páramo de Choachí, 2 (1[2], 1[84]); Laguna del
Verjón (= City of Bogotá), 1[75]; Quetame, 2[2]; Fusagasuga, 1;
Caqueta, 2[2]; no locality more definite than Colombia, 3 (1[7]).
=Mustela frenata aureoventris= Gray
Long-tailed Weasel
Plates 27, 28 and 29