resembles _M. f. frenata_ of northern México and the high mountain
forms of southern México more than it does the lowland tropical forms.
This is true as regards size of entire animal, proportions of its
parts, and size, general angularity and major proportions of its skull.
The marked postorbital constriction, convex supralacrymal face of
rostrum, width of tympanic bullae and angularity of the braincase place
it nearest _M. f. frenata_ as does also the color and color pattern.
The ventrally convex squamosal anterior to each tympanic bulla and the
slight degree of projection from the cranium of the anterior margin of
each tympanic bulla are intermediate in degree between the condition in
_M. f. macrophonius_ and that in _M. f. frenata_. Thus _M. f. perotae_
combines several characters of _M. f. frenata_ on the one hand with
some of _M. f. macrophonius_ on the other and in some features, for
instance in the size, shape and degree of inflation of the tympanic
bullae, presents intermediate stages of development.
On the eastern plain below the high mountain, Cofre de Perote, there
ranges the similarly colored, smaller, tropical weasel, _Mustela
frenata tropicalis_. Between _M. f. perotae_ and _M. f. tropicalis_
there is marked differentiation in the skulls with much less
differentiation in coloration. The differences in typical skulls of the
two subspecies are so pronounced that one would, at first glance,
hardly believe it possible for direct intergradation to occur between
them on the sides of this mountain. Merriam (1896:30) thought that it
did not. The two skulls figured by him (_op. cit._:31) are a topotype
of _M. f. tropicalis_ from Jico and the one which now is the type
specimen of _M. f. perotae_. They show the great difference in size and
proportions and are females of comparable ages, not of different ages
as I suspected before examining the skulls. However, despite this
marked difference in the skulls, there is some, although not
conclusive, evidence of intergradation furnished by a young female from
Xuchil, Veracruz. This specimen is described in connection with _M. f.
tropicalis_ (see p. 366).
None of the seven skulls shows marked deformity of the interorbital
region, but two of the three adults appear to have had these parts
infested with nematodes.
_Specimens examined._--Total number, 7, all from México, listed by
localities from north to south. Specimens from Veracruz and Oaxaca
in the United States National Museum; remainder in Texas
Cooperative Research Collection.
=México=: Monte Río Frío, 45 Km. ESE México City, 1; 55 Km. ESE
México City, 1; N slope Mt. Popocatépetl, 13,555 ft., 1.
=Puebla.= Río Otlati, 8700 ft., 1.
=Veracruz.= Cofre de Perote, 12,500 ft., 1; Perote, 1.
=Oaxaca.= Cerro San Felipe, 10,000 ft., 1.
=Mustela frenata goldmani= (Merriam)
Long-tailed Weasel
Plates 1, 24, 25, 26 and 30
_Putorius frenatus goldmani_ Merriam, N. Amer. Fauna, 11:28, June
30, 1896; Elliot, Proc. Biol. Soc. Washington, 18:236, December
9, 1905.
_Mustela frenata goldmani_, Miller, U. S. Nat. Mus. Bull., 79:100,
December 31, 1912; Hall, Carnegie Instit. Washington Publ.
473:109, November 20, 1936.
_Type._--Male, adult, skull and skin; no. 77519, U. S. Nat. Mus.,
Biol. Surv. Coll.; Pinabete, Chiapas, México; February 10, 1896;
obtained by E. A. Goldman (on attached label collectors recorded
as Nelson and Goldman); original no. 9279.
The skull (plates 24 and 30) has the rostrum badly injured. All
the right, and part of the left nasal, the upper part of the right
maxilla, the postorbital process and intervening area of frontals
are missing. Each zygomatic arch is broken but the parts are
present and attached to the skull. The frontal and interorbital
regions are greatly malformed owing to parasites that infested the
sinuses. Right I2 and I3, right and left i3, and the medial parts
of the paraconid and protoconid of right m1 are missing. The light
facial markings are less extensive than in any of the referred
specimens. These markings consist of a separate spot between the
eyes and a white line, confluent with the color of the underparts,
on each side of the head, that extends from the base of the ear to
above the eye. The dark color of the underparts is represented at
the angles of the mouth by a spot on the left side and a similar
dark area, confluent with the dark color of the face, on the right
side. The large size, characters of the skull, and scrotal pouch
on the skin prove the specimen to be a male as stated on the
label.
_Range._--Two thousand five hundred feet (El Cipres, Guatemala) to
9500 feet (near Tecpám, Guatemala), Upper Tropical Life-zone of
mountains and western coasts of southern México, Guatemala and
Salvador. See figure 29 on page 221.
_Characters for ready recognition_ (characters based on
males).--Differs from _M. f. nicaraguae_ and _M. f. perda_ by
larger size (total length of adult males more than 489), least
width of color of underparts not less than 26 per cent of greatest
width of color of upper parts, weight of skull of adult male more
than 5 grams; from _M. f. macrophonius_ by smaller size (total
length of adult males less than 540), skull of male with basilar
length less than 52.5 and weight less than 6 grams; from _M. f.
perotae_ (typical specimens of same sex not available) by darker
color of upper parts which are Argus Brown or darker rather than
Brussels Brown; nonextension of color of underparts onto hind
feet; from _M. f. leucoparia_ in least width of color of
underparts not more than 37 per cent of greatest width of color of
upper parts; color of underparts not extended onto hind feet;
black tip of tail two-fifths rather than one-fourth as long as
tail-vertebrae; height of tympanic bulla less than four-fifths
distance from its anterior margin to foramen ovale.
_Description._--_Size._--Male: Four adults yield average and
extreme measurements as follows: Total length, 508 (500-512);
length of tail, 196 (185-207); length of hind foot, 55.5 (54-58).
Tail averages 63 (59-67) per cent as long as head and body. Length
of hind foot more than basal length.
Female: Typical specimen unknown.
_Externals._--Longest facial vibrissae black and reaching beyond
ear; carpal vibrissae wholly or in part of same color as upper
parts and reaching as far as hypothenar pad; hairiness of
foot-soles distinctly less than that shown in figure 20 on page
60.
_Color._--Spot between eyes, band, confluent with color of
underparts, on each side of head extending anterodorsally anterior
to each ear and posterior third of each upper lip, white;
remainder of sides and top of head and neck posteriorly to or
slightly behind shoulders, black; dark spots at angles of mouth
usually absent; tip of tail black; remainder of upper parts Argus
Brown or near (_n_) Argus Brown; chin, throat and breast white;
remainder of underparts near (16' _c_) Ochraceous-Buff; color of
underparts extending distally on posterior sides of forelegs onto
medial toes and on hind legs to points between knees and heels.
Least width of color of underparts, in five adult males, averaging
28 (extremes 26-33) per cent of greatest width of color of upper
parts; black tip of tail, in four adult males, averaging 40 per
cent of length of tail-vertebrae.
_Skull and teeth._--Male (based on five adults): See measurements
and plates 24-26, 30; weight, 5.4 (5.3-5.5) grams; basilar length,
49.9 (49.6-51.3); zygomatic breadth (except in no. 12523 from
Salvador) more than or equal to distance between condylar foramen
and M1 or between anterior palatine foramen and anterior margin of
tympanic bulla. Mastoid breadth less than postpalatal length;
postorbital breadth more or less than length of upper premolars
and greater than width of basioccipital measured from median
margin of one foramen lacerum posterior to its opposite;
interorbital breadth less than distance between foramen opticum
and anterior margin of tympanic bulla; breadth of rostrum less
than length of tympanic bulla; least width of palate more or less
than length of P4; anterior margin of tympanic bulla as far
posterior to foramen ovale as width of five 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 shorter than or equal to length
of rostrum; anterior margin of masseteric fossa immediately behind
m2.
Female: Typical skull unknown.
Comparison of male skull with that of _M. f. perda_ made in discussion
of that form. Comparison with that of _M. f. nicaraguae_ shows similar
differences, some of which are more pronounced. For example, squamosals
anterior to tympanic bullae more convex ventrally and these bullae
project less from braincase than in _M. f. perda_; thus the difference
in these features is greater between _goldmani_ and _nicaraguae_ than
between _goldmani_ and _perda_.
As compared with the skull of the male of _M. f. macrophonius_, each
one of the skulls of the adult males of _M. f. goldmani_ is smaller in
every measurement taken, with two exceptions. The width of the tympanic
bullae was more in three specimens of _M. f. goldmani_ as was also the
depth of the same in three specimens. Relative to the basilar length
all but two of these measurements average less in _goldmani_; the
exceptions are the zygomatic breadth and depth of the skull at the
anterior margin of the tympanic bullae which average more. Relative to
the basilar length, the orbitonasal length and depth of the skull at
the posterior margin of M1 are less in each skull of _goldmani_. Thus,
excepting the width and height of the tympanic bullae and the relative
zygomatic breadth and relative depth of the braincase posteriorly, the
skull of _goldmani_ is shorter and relatively as well as actually
narrower and lighter throughout.
As compared with the skull of the male of _M. f. leucoparia_, that of
_M. f. goldmani_ averages a trifle shorter and no skull of _goldmani_
equals that of _leucoparia_ in actual or relative zygomatic and mastoid
breadths or length or height of tympanic bullae. In depth, the skull of
_goldmani_ averages actually and relatively greater. Its teeth are
smaller. The squamosal anterior to each tympanic bulla is convex
ventrally whereas it is concave ventrally in _leucoparia_ as in
_frenata_.
_Remarks._--When Merriam (1896:28) named this subspecies, he had only
one specimen but he called attention to the more important diagnostic
characters, which additional specimens show pertain to the race as a
whole.
_M. f. goldmani_ in typical form occurs in high mountains of the Upper
Tropical Life-zone and is most closely related to _M. f. frenata_ and
_M. f. macrophonius_. The altitude at which the two specimens were
taken, twenty miles southeast of Teopisca in Chiapas, is not known.
Merriam (1896:28) states that the type specimen was obtained at "about
8200 feet." The specimen taken by Stirton in Salvador comes from 8000
feet and the one obtained by Barber in Guatemala from 9500 feet. The
specimen from Dueñas, the skin alone of a young animal, is not
instructive.
As regards size, _goldmani_ is larger than the immediately adjacent
subspecies from the Lower Tropical Life-zone but is smaller than _M. f.
leucoparia_ or _macrophonius_. As compared with _M. f. frenata_,
_goldmani_ is longer, has an actually as well as relatively shorter
tail, and a much longer hind foot.
The most outstanding difference in externals from _frenata_ is the
naked foot soles.
Molting probably takes place twice each year although actual proof of
this is lacking. In number 133254 from twenty miles southeast of
Teopisca, taken on May 12, the molt is well advanced. Another specimen
from the same place still retains the winter coat.
In color, _goldmani_ is much darker than _frenata_, has less extensive
white facial markings, longer black tip on tail, more restricted color
of underparts, and lacks the extension of color of the underparts onto
the hind feet.
Of the adult males from the high mountains, the type specimen from
Chiapas is lightest, and the one from Salvador is darkest. This
progressively darker color to the southward probably is geographic
variation.
In total length and relative and actual length of tail, the specimen
from Salvador is the smallest of the five adult males from the higher
mountains. In addition to its darker color and smaller size, no. 12523
from Salvador shows certain distinctive cranial characters. The
zygomatic breadth is less than, rather than more than, or equal to, the
distance between the condylar foramen and M1 or than that between the
anterior palatine foramen and the anterior margin of the tympanic
bulla. This difference appears to be correlated with geographic
position, since no. 15953 from Guatemala has the three distances about
equal and therefore is intermediate in this respect between the
specimen from Salvador and those from Chiapas, in which the zygomatic
breadth is greater than the other two measurements. Also in the greater
depth of the skull and smaller size of the teeth, this specimen from
Salvador approaches the subspecies of the Lower Tropical Life-zone. It
has, however, the longest, highest and widest tympanic bullae of any of
the five specimens. The amount of ventral convexity of the squamosal in
front of each tympanic bulla appears not to be greater than in the
other specimens.
As indicative of intergradation with _perotae_, _leucoparia_ and
possibly _frenata_, there is the specimen from Cerro San Felipe,
Oaxaca. The degree of restriction of the color of the underparts is
intermediate between that of _goldmani_ and _leucoparia_. The same is
true as regards the amount of projection from the braincase of the
anterior margins of the tympanic bullae. The squamosal immediately
anterior to each tympanic bulla is flat instead of ventrally convex as
in _goldmani_ or ventrally concave as in _leucoparia_ and _frenata_. In
accordance with the custom adopted in this paper of referring every
specimen to some one subspecies, this specimen from Cerro San Felipe is
referred to _Mustela frenata perotae_, to the description of which it
most nearly answers.
Possibly _goldmani_, as here constituted, is a composite form. The
specimens from the high mountains closely resemble one another.
However, a specimen, no. 68541 from "Finca El Cipres," Guatemala, which
place Mr. G. Goodwin tells me is at an elevation of 2500 feet,
approximately 5 miles north of Retalhuleu, has a basilar length of 47.3
and is correspondingly small in other parts. This suggests the
existence of a small, lowland race on the western side of the central
divide corresponding to _perda_ and _tropicalis_ on the eastern side.
From only a few miles away, at San Sebastian, there is available, the
adult skull of a still smaller animal. This skull only, no. 41026, in
the Berlin Zoological Museum, has a basilar length of 46.1, zygomatic
breadth of 27.4, and other cranial measurements notably smaller than
those of specimens from the high mountains. A skin-only, no. 12038,
collection of Donald R. Dickey, from La Cebia, altitude 2150 feet, near
the city of San Salvador, seemingly represents an animal smaller than
typical _goldmani_. This specimen from La Cebia has the light color of
the underparts extended distally on the hind legs to the tips of the
toes as in _M. f. tropicalis_. However, the upper parts are darker and
resemble those of _M. f. goldmani_. A fourth specimen from only 3500
feet elevation, on the south side of Volcano Tajumulco, Guatemala, no.
41768, Field Museum of Natural History, a subadult male, measures only
490 in total length and has the least color of the underparts so
restricted as to amount to only 22 per cent of the greatest width of
the color of the upper parts. Both these features are suggestive of the
lowland races.
These four specimens indicate that the lowland population on the
western side of the divide is smaller than the mountain population. The
juvenile from Carolina and a young male from Finca Cipres, however,
both closely resemble individuals of _goldmani_ from the higher
mountains. All these animals here are referred to _goldmani_. More
specimens may reveal an amount and a pattern of geographic variation in
weasels of this region that will require application of another
subspecific name.
The female, no. 68540, from Puebla agrees remarkably well with the
skull of the female, no. 132528, of _macrophonius_. Differences
displayed by the specimen from Puebla are its slightly narrower
braincase and longer space between the foramen ovale and anterior end
of the tympanic bulla. Considering the far eastern location of Puebla
(just north of Río Motagua, at 89° W, according to a sketch map
provided by Mr. G. G. Goodwin), this specimen might be expected to show
some approach to the small lowland races. Actually, however, it
displays the characters of _goldmani_ better than does the subadult
female from Volcano San Lucas, which is nearer the metropolis of
_goldmani_, and I assume at a higher elevation than Puebla.
Concerning this weasel Merriam (1896:29) says: "Mr. E. W. Nelson writes
me that this fine weasel is found sparingly in the forest about
Pinabete, Chiapas, at an altitude of 7000 to 8000 feet (2100 to 2500
meters). The type specimen was shot in the afternoon while hunting on a
heavily wooded hill slope. It was heard making long, slow leaps over
the dry, crisp leaves. Coming to a log, it stood up and rested its fore
feet on the log, in which position it was shot by Mr. Goldman."
The specimen taken by R. A. Stirton in Salvador comes from an elevation
of 8000 feet in the rain forest of the Upper Tropical Life-zone. Mr.
Stirton tells me that one morning on visiting his traps set for small
rodents, he found in one the partly eaten remains of a _Heteromys_.
Leaving these remains as found he placed a steel trap beside them and
on the following morning found the male weasel in the trap.
At least three of the ten specimens had the frontal sinuses infested
with parasites.
_Specimens examined._--Total number, 15, listed by localities from
north to south, and unless otherwise indicated in the American
Museum of Natural History.
=México=: _Chiapas_: 20 mi. SE Teopisca, 2[91]; Pinabete, 1[91].
=Guatemala=: Puebla, 1; Finca Porvenir, 3500 ft., S slope Volcan
Tajumulco, 1[60]; Sierra [=? Cerro] Santa Elena, 9500 ft. (near
Tecpám), 1[60]; Carolina, 1; Volcano San Lucas, 1; "Finca El
Cipres," 1; "Finca Cipres," 2500 ft., 1; Finca San Isidro, San
Sebastión, Dept. Retalhuleu, 1[4]; Dueñas, 1[7]; no locality more
definite than Guatemala, 1[7].
=El Salvador=: Los Esesmiles, 8000 ft., Chalatenango, 1[59]; La
Cebia, 2150 ft., near San Salvador, 1[59].
=Mustela frenata macrophonius= (Elliot)
Long-tailed Weasel
Plates 24, 25, 26, 30, 37, 38 and 39
_Putorius macrophonius_ Elliot, Proc. Biol. Soc. Washington,
18:235, December 9, 1905.
_Mustela macrophonius_, Miller, U. S. Nat. Mus. Bull., 79:100,
December 31, 1912.
_Mustela frenata macrophonius_, Hall, Carnegie Instit. Washington
Publ. 473:109, November 20, 1936.
_Type._--Male, adult, skull and skin; no. 14063, Field Mus. Nat.
Hist.; Achotal, Veracruz, México; January 15, 1904; obtained by
Edmund Heller and Charles M. Barber; original no. 3424.
The skull (plates 24-26, 30) is complete and unbroken. Excepting
right P2, which has been aborted or broken away, all the teeth are
present. The skin is well made and in good condition. As shown by
the scrotal pouch, the specimen is a male.
_Range._--Tropical Life-zone, probably into Boreal life-zones, of
mountains along eastern border of southern Veracruz. See figure 29
on page 221.
_Characters for ready recognition._--Differs from _M. frenata
frenata_ and _M. f. perotae_ and _M. f. leucoparia_ in lacking
color of underparts on hind feet and in larger skull (skulls of
adult males with basilar length more than 52.5); from _M. f.
goldmani_ by larger size of skull (see above) and entire animal
and wider tympanic bullae; from _M. f. tropicalis_ and _M. f.
perda_ by larger size (total length of adult males more than 510),
postorbital breadth amounting to less than combined length of
upper premolars.
_Description._--_Size._--Male: External measurements of the type
specimen, an adult, are: Total length, 598; length of tail, 246;
length of hind foot, 59. Tail 70 per cent as long as body; length
of hind foot more than basal length.
Female: The skin, without field collector's measurements, of an
adult female from Pérez, Veracruz, shows this sex to be
correspondingly large. Because the skin is understuffed and
because the hind feet are skinned out, reliable measurements can
not be obtained from the dried skin.
_Externals._--As described in _Mustela frenata goldmani_ except
that all carpal vibrissae are of same color as upper parts and
that hairiness of foot-soles is halfway between that shown in
figures 20 and 21.
_Color._--As in darkest individuals of _M. f. goldmani_, thus,
color of upper parts on posterior part of back near (_n_) Argus
Brown. Color of underparts near (12) Mikado Orange in a juvenile,
extending distally on posterior sides of forelegs onto inner toes
and on hind legs to points between knees and heels. Least width of
color of underparts 28 per cent of greatest width of color of
upper parts. Black tip of tail 34 per cent of length of
tail-vertebrae.
_Skull and teeth._--Male (based on type specimen): See
measurements and plates 24-26, 30. As described in _Mustela
frenata frenata_ except that: Weight, 6.9 grams; basilar length,
54.1; zygomatic breadth less than distance between condylar
foramen and M1 or that between anterior palatine foramen and
anterior margin of tympanic bulla; interorbital breadth less than
distance between foramen opticum and anterior margin of tympanic
bulla; anterior margin of tympanic bulla as far posterior to
foramen ovale as width of 4 to 6 upper incisors; height of
tympanic bulla less than distance from its anterior margin to
foramen ovale; anterior margin of masseteric fossa below posterior
half of m2.
Female (based on no. 132528): See measurements and plates 37-39.
As described in _Mustela frenata frenata_ except that: Weight, 3.6
grams; basilar length, 43.5; zygomatic breadth less than distance
between condylar foramen and M1 and more or less than (in the
single specimen, equal to) that between anterior palatine foramen
and anterior margin of tympanic bulla; least width of palate more
or less than (about equal to) outside length of P4; tympanic bulla
as far posterior to foramen ovale as width of 4 or 5 upper
incisors; height of tympanic bulla less than distance from its
anterior margin to foramen ovale.
The skull of the female is 48 per cent lighter than that of the
male.
Comparison of the skull with that of _M. f. goldmani_ is made in the
account of that subspecies. Similar differences probably exist between
males of _perotae_ and _macrophonius_. As compared with skulls of males
of _M. f. tropicalis_ and _perda_, the skull of the male of
_macrophonius_ is larger in every measurement taken. The postorbital
constriction is less, rather than more, than the combined length of the
upper premolars. Relative to the basilar length, the following
measurements are less than in any specimen of _tropicalis_ or _perda_:
length of tooth-rows; orbitonasal length; depth of skull at posterior
border of upper molars; and depth of skull at anterior margin of
basioccipital.