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 28-30. 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 39 and 40
_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 39 and 40.)
_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 39 and 40; 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
[M] and 56 [F] with skulls of comparable age) and _primulina_ (64 [M]
and 24 [F] 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 29 on page
221.
_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
-, -, ---, -; teeth heavy; medial lobe of M1 but slightly larger than
3 1 2 2
lateral lobe. See plates 28, 29, 30, 39 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
---, is also intermediate between the numbers
2
3 2
- and - of the North American _Mustela_ and the Patagonian
3 2
_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 28, 29, 30 and 41
_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 29 on page 221.
_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 22.
_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 28-30). 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."