















Pipistrellus cinnamomeus Miller 1902
Referred to the Genus Myotis

BY

E. RAYMOND HALL and WALTER W. DALQUEST

University of Kansas Publications
Museum of Natural History

Volume 1, No. 25, pp. 581-590, 5 figures in text
January 20, 1950

University of Kansas
LAWRENCE
1950


UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY

Editors: E. Raymond Hall, Chairman, Edward H. Taylor,
A. Byron Leonard, Robert W. Wilson

Volume 1, No. 25, pp. 581-590, 5 figures in text
January 20, 1950

UNIVERSITY OF KANSAS
Lawrence, Kansas

PRINTED BY
FERD VOILAND. JR., STATE PRINTER
TOPEKA, KANSAS
1950

23-1545

[Transcriber's Note: Words surrounded by tildes, like ~this~ signifies
words in bold. Words surrounded by underscores, like _this_, signifies
words in italics. Male symbol is shown as [M] and female symbol is
[F].]




Pipistrellus cinnamomeus Miller 1902
Referred to the Genus Myotis

By

E. RAYMOND HALL AND WALTER W. DALQUEST


Miller (Proc. Acad. Nat. Sci. Philadelphia, 1902, p. 390, September
3,1902) based the name _Pipistrellus cinnamomeus_ on a skin and skull of
a vespertilionid bat obtained on May 4, 1900, at Montecristo, Tabasco,
Mexico, by E. W. Nelson and E. A. Goldman. A single specimen was
available to Miller when he proposed the name _P. cinnamomeus_. Dalquest
and Hall (Jour. Mamm., 29:180, May 14, 1948) reported three additional
specimens collected in 1946 by W. W. Dalquest on the Río Blanco, twenty
kilometers west-northwest of Piedras Negras, Veracruz, Mexico. No other
published information concerning this species is known to us, although
the name has, of course, appeared in regional lists, for example in the
"List of North American Recent Mammals, 1923" (Bull. U. S. National
Museum, 128:75, April 29, 1924) by Gerrit S. Miller, Jr.

Additional specimens, nevertheless, are known. Two collected on April 18
and 20, 1903, at Papayo, Guerrero, by Nelson and Goldman, are in the
Biological Surveys Collection in the United States National Museum. A
skin, probably of this species, for which the skull cannot now be found,
was taken on October 27, 1904, at Esquinapa, Sinaloa, by J. H. Batty and
is in the American Museum of Natural History. This is the skin referred
by Miller and Allen (Bull. U. S. Nat. Mus., 144:100, May 25, 1928) to
_Myotis occultus_. Three additional specimens, each a skin with skull,
were collected twenty kilometers east-northeast of Jesús Carranza, at
200 feet elevation, Veracruz, by Walter W. Dalquest, two on April 13,
1949, and one on May 16 of the same year. These are in the Museum of
Natural History of the University of Kansas, as also are the three
previously reported by Dalquest and Hall (_loc. cit._). A total of ten
specimens, from five localities, all in Mexico, thus is accounted for.

On page 392 of the original description--which our study of the holotype
shows to be accurate--Miller wrote: "This bat differs so widely from the
other known American species of _Pipistrellus_ as to need no special
comparisons. Superficially it has much the appearance of an unusually
red _Myotis lucifugus_, and only on examination of the teeth do the
animal's true relationships become apparent." In referring to the teeth
Miller almost certainly was thinking of the premolars of which there are
only two on each side of the upper jaw and on each side of the lower jaw
in _Pipistrellus_, including his _Pipistrellus cinnamomeus_, whereas
_Myotis_ at that time was thought always to have three premolars on each
side of both the upper and lower jaw, except in rare instances where one
premolar might be lacking on one side of one jaw or even more rarely on
both sides of the upper jaw. In his original description of _P.
cinnamomeus_, Miller mentioned also that it had the "Inner upper incisor
distinctly smaller than the outer, not approximately equal to it as is
the case in _P. subflavus_."

At this point it is well to make clear that each of the genera
_Pipistrellus_ and _Myotis_ contains a large number of species and that
the differences between the two genera are few. Our examination of
American specimens reveals only one differential character: In _Myotis_
the outer upper incisor is distinctly larger than the inner, whereas the
two incisors are of approximately equal size in _Pipistrellus_. It may
be noted that the outer upper incisor of several, but not all, species
of _Myotis_ has a well-developed concave surface directed toward the
canine whereas this surface is flat or convex in _Pipistrellus_. In both
features, the type of _Pipistrellus cinnamomeus_ Miller agrees with
_Myotis_ and differs from _Pipistrellus_.

Five years after naming and describing _Pipistrellus cinnamomeus_,
Miller published his monumental work entitled "The families and genera
of bats" (Bull. U. S. Nat. Mus., 57, June 29, 1907) wherein he points
out the differences in the upper incisors between _Pipistrellus_ and
_Myotis_ (by a _lapsus plumae_ ascribes subequal incisors to _Myotis_
and unequal incisors to _Pipistrellus_) but seemingly failed to
reëxamine _P. cinnamomeus_ in the light of this better understanding of
the two genera, or if he did examine _P. cinnamomeus_ he possibly was
misled still by the absence of the third premolar on each side of both
the upper and lower jaw.

In 1928 when Miller and Allen published their account of "The American
bats of the Genera _Myotis_ and _Pizonyx_" (Bull. U. S. Nat. Mus., 144,
May 25, 1928) they examined specimens of _Myotis occultus_ which they
implied (_op. cit._: 99-100) had only two instead of three premolars on
each side of both the upper and lower jaws. In preparing this taxonomic
account of bats of the genus _Myotis_, the specimens (type and two from
Papayo) of _Pipistrellus cinnamomeus_ seem not to have been examined.
Indeed, it is almost certain that they were not examined for the species
was renamed; the new name, _Myotis lucifugus fortidens_ Miller and
Allen (Bull. U. S. Nat. Mus., 144:54, May 25, 1928), was based on a
skull with the corresponding body in alcohol. The characters of this
specimen are almost exactly those of _Pipistrellus cinnamomeus_, named
and described by Miller 26 years earlier. The type locality (Teapa) of
_M. l. fortidens_ is 80 miles westerly from the type locality of _P.
cinnamomeus_; both are in the state of Tabasco, and in the same
life-zone, at equivalent elevations (neither higher than 50 meters).
Since there are no characters of taxonomic worth to distinguish the two
named specimens, _Myotis lucifugus fortidens_ Miller and Allen 1928
falls as a synonym of _Pipistrellus cinnamomeus_ Miller 1902. But,
according to Miller and Allen (Bull. U. S. Nat. Mus., 144:19, 197),
_Vespertilio cinnamomeus_ Wagner 1855 is a name based on _Myotis ruber_
(E. Geoffroy, 1806) from Paraguay and hence _Myotis cinnamomeus_
(Miller) 1902 is a homonym of _Myotis cinnamomeus_ (Wagner) 1855 and is
unavailable for the animal from Montecristo when it is transferred to
the genus _Myotis_; the species of animal concerned will take the next
available name, which seems to be _Myotis lucifugus fortidens_ Miller
and Allen 1928.

It may reasonably be asked if _Myotis_ and _Pipistrellus_ should be
retained as separate genera if the only constant difference between the
two is subequal versus unequal upper incisors. In our opinion it would
be worth-while for someone who had access to adequate material from both
the Old World and the New World to investigate this question. We lack
adequate material from the Old World.

When Miller and Allen named _M. l. fortidens_ they had only two
specimens, the holotype from Teapa, Tabasco, and a referred specimen
from Fort Hancock, El Paso County, Texas, approximately 1,200 miles
north-northwest of Teapa. We have examined this specimen from Texas (U.
S. Nat. Mus., 21083/36121, skin and skull) and regard it as _Myotis
lucifugus carissima_ Thomas. Furthermore, we regard the holotype of
_Myotis lucifugus fortidens_ Miller and Allen 1928 as specifically
distinct from _Myotis lucifugus_ of Miller and Allen 1928. The Cinnamon
Myotis, described below, therefore may stand as:


~Myotis fortidens~ Miller and Allen

CINNAMON MYOTIS

     _Pipistrellus cinnamomeus_ Miller, Proc. Acad. Nat. Sci.
     Philadelphia, p. 390, September 3, 1902, type from
     Montecristo, Tabasco (preoccupied by _Vespertilio
     cinnamomeus_ Wagner, Schreber's Säugethiere, suppl., 5:755,
     1855, a renaming of _Vespertilio ruber_ E. Geoffroy
     Saint-Hilaire).

     _Myotis lucifugus fortidens_ Miller and Allen, Bull. U. S.
     Nat. Mus., 144:54, May 25, 1928.

_Type._--"Adult female (in alcohol) No. 88.8.8.18, British Museum
(Natural History). Collected at Teapa, Tabasco, Mexico, by H. H. Smith,
January 5, 1888. Presented by Messrs. Salvin and Godman [after Miller
and Allen, Bull. U. S. Nat. Mus., 144:54, May 25, 1928]."

_Range._--Known only from the lower part of the Tropical Life-zone of
the region of the Isthmus of Tehuantepec and east and west coasts of
Mexico.

[Illustration: FIG. 1. Map showing localities from which _Myotis
fortidens_ has been recorded.]

_Diagnosis._--Among American species of the genus, over-all size medium
(total length 94 mm); body long (54); tail short (39); forearm of medium
length (37); tibia short (14.5); foot long (58 per cent of length of
tibia); wing membrane arising from side of foot at distal end of
metatarsal; calcar simple (not keeled) and 7 mm long; ears 15 to 16 mm
long measured in the flesh from the notch (posteroventral border of the
meatus); tragus, measured from same place, 7 to 8 mm high with
posterobasal lobe; third metacarpal longest and second metacarpal
shortest; fifth shorter than fourth; ears brownish; membranes of wing
and tail blackish; uropatagium almost hairless, the few hairs that are
present being almost invisible; pelage of back 5 mm long with some
overhairs 8 to 9 mm long; basal 3 mm of fur black, remainder
Cinnamon-Brown (capitalized color terms, after Ridgway, Color Standards
and Color Nomenclature, Washington, D. C., 1912); outline of skull
viewed dorsally similar to that of _Myotis lucifugus_; sagittal crest
well developed; distance across upper canines equal to or slightly
exceeding interorbital constriction; braincase low; two premolars on
each side in upper jaw and also in lower jaw, the one remaining small
premolar in contact with both the canine and the fourth premolar.

[Illustration: FIGS. 2-5. Four views of the skull of _Myotis fortidens_.
No. 32112, University of Kansas Museum of Natural History, [M], obtained
20 kilometers east-northeast Jesús Carranza, 200 feet elevation,
Veracruz, Mexico, on May 16, 1949, by Walter W. Dalquest; original no.
12869. ×2.]

_Remarks._--_Myotis fortidens_ is known only from the Tropical
Life-zone. The skin, without a skull, from Esquinapa, Sinaloa, agrees in
color with the undoubted specimens of _M. fortidens_ from Papayo,
Guerrero, but can be matched also by selected skins of _Myotis occultus_
from Blythe, Riverside County, California. Without the skull the
reference of this specimen to _M. fortidens_ is provisional. Reason for
referring it to _fortidens_ rather than to _M. occultus_ is provided,
however, by a series of eleven specimens of _M. occultus_ from Álamos,
Sonora. These are Saccardo's Umber rather than Cinnamon-Brown and they
are geographically intermediate between the reddish _M. occultus_ of
California and the reddish _M. fortidens_ of Mexico. Furthermore, these
specimens from Álamos have large skulls of slightly different
proportions than those of _M. fortidens_ or than those of _M. occultus_
from California; possibly the animals from Álamos are representative of
the larger, duller-colored variation for which Hollister proposed the
name _Myotis baileyi_ (Proc. Biol. Soc. Washington, 22:44, March 10,
1909). This duller-colored type of animal intervenes between the
geographic ranges of undoubted _M. occultus_ and undoubted _M.
fortidens_. The specimen from Esquinapa, in the geographic sense, is on
the _fortidens_ side rather than on the _occultus_ side of the _baileyi_
population. This geographic position is the basis on which the specimen
from Esquinapa is referred to _M. fortidens_. The third premolar is
lacking from each side of both the upper and the lower jaws of each
individual of this series from Álamos.

The specimens of _M. fortidens_ are all distinguishable by their color
from other kinds of _Myotis_ found in the same area. Occasional
individuals of _Myotis velifer_, as for example three from Las Vigas,
Veracruz, also are reddish but they are of brighter tone. In addition,
the larger size and cranial features of these specimens of _M. velifer_
permit ready differentiation of them from specimens of _M. fortidens_.
One specimen (No. 32113) of _M. fortidens_ from twenty kilometers
east-northeast of Jesús Carranza is lighter than the others, being near
(_j_) Cinnamon-Brown above and is lighter on the under-parts than on the
upper parts. Another individual (No. 32112) is duller colored than the
others, being Snuff Brown both above and below. Otherwise the specimens
of _M. fortidens_ agree in color.

Among named kinds of _Myotis_, _M. fortidens_ resembles _Myotis
lucifugus_ and _Myotis occultus_. From the former, _M. fortidens_
differs in possessing a strong sagittal crest and in lacking the third
premolar in both the upper jaw and the lower jaw. _M. fortidens_ lacks
the glossy sheen found on the pelage of many individuals of _M.
lucifugus_. From _M. occultus_, _M. fortidens_ differs in having the
rostrum (viewed from above) smaller in relation to the braincase. This
is true of specimens with the teeth showing much wear as well as in
specimens with the teeth unworn or only moderately worn. Also, _M.
fortidens_ is longer bodied as may be seen by comparing the measurements
given here with those recorded for _M. occultus_ by Miller and Allen
(Bull. U. S. Nat. Mus., 144:100, May 25, 1928). We are agreed that _M.
fortidens_ is as closely related to _M. occultus_ as to any other named
kind of _Myotis_, and that it is more closely related to it than to most
other species of the genus, but one of us (Dalquest) thinks that _M.
fortidens_ is specifically distinct from _M. occultus_, whereas the
other author (Hall) inclines to the view that additional specimens from
localities intermediate between the known geographic ranges of _M.
occultus_ and _M. fortidens_ will reveal intergradation between the two
kinds. However that may be, there is no proof at present of such
intergradation and the binomial is therefore used for the Cinnamon
Myotis.

     _Specimens examined._--Total number, 10, all from Mexico,
     each a skin with skull except the skin-only from Sinaloa.
     _Sinaloa_: Esquinapa, 1 (Amer. Mus. Nat. Hist.). _Guerrero_:
     Papayo, 2 (U. S. Biological Surveys Collection). _Veracruz_:
     20 km. WNW Piedras Negras, 3 (Mus. Nat. Hist., Univ.
     Kansas); 20 km. ENE Jesús Carranza, 200 ft. elevation, 3
     (Mus. Nat. Hist., Univ. Kansas). _Tabasco_: Montecristo, 1
     (U. S. Biological Surveys Collection).

     _Additional record._--Tabasco: Teapa, the holotype of
     _Myotis lucifugus fortidens_ Miller and Allen 1928.

_University of Kansas Museum of Natural History, Lawrence, Kansas.
Transmitted October 31, 1949._

CRANIAL MEASUREMENTS OF _Myotis fortidens_

Column headings:

A: Greatest length
B: Condylobasal length
C: Zygomatic breadth
D: Interorbital constriction
E: Breadth of braincase
F: Mandible
G: Maxillary tooth-row
H: Maxillary breadth at M3
I: Mandibular tooth-row
J: Wear of teeth

==============================================================================
         Sex
    No.  Age    Locality        A     B     C    D    E     F    G   H   I   J
------------------------------------------------------------------------------
    25030[M]    Esquinapa      ....  ....  ...  ...  ...  ....  ... ... ...  ?
   126650[F]    Papayo         15.0  14.2  9.7  3.9  7.1  11.5  5.5 5.6 6.0  0
   126651[F]    Do.            15.1  13.8  9.4  3.8  6.8  10.6  5.6 5.9 6.0  0
    17834[M]    P. Negras[1]   ....             4.1       10.6  5.6 5.7 6.0  0
    17835[F]    Do.            15.5  14.9  9.6  4.2  7.2  11.0  5.7 6.0 6.1  2
    17836[F]    Do.            15.5  14.5  9.7  4.2  7.3  10.9  5.4 5.9 5.7  3
    32112[M]    J. Carranza[2] 15.3  14.4  9.7  4.1  7.3  11.5  5.7 5.9 6.3  1
    32113[M]    Do.            15.0  14.0  9.5  4.2  7.2  10.9  5.5 5.9 5.9  1
    32114[M]    Do.            15.0  13.9  9.7  4.1  7.2  10.8  5.4 6.0 5.9  1
88.8.8.18[F][3] Teapa          15.0  13.8  9.6  3.8  7.4  ....  5.4 5.8 5.8  1
   100231[F][4] Montecristo    15.0  14.1  9.0  4.0  7.2  11.4  5.8 ... 6.0  0
  Average                      15.2  14.2  9.5  4.0  7.2  11.0  5.6 5.9 6.0
------------------------------------------------------------------------------

[Note 1: 20 km. WNW Piedras Negras.]

[Note 2: 20 km. ENE Jesús Carranza, 200 ft.]

[Note 3: Type of _Myotis lucifugus fortidens_; measurements after Miller
and Allen, Bull. U. S. Nat. Mus., 144:100; 101, May 25, 1928.]

[Note 4: Type of _Pipistrellus cinnamomeus_ Miller 1902.]


EXTERNAL MEASUREMENTS OF _Myotis fortidens_

Column headings:

A: Total length
B: Head and body
C: Tail
D: Tibia
E: Foot
F: Forearm
G: Thumb
H: Third metacarpal
I: Fifth metacarpal
J: Ear from notch

===============================================================================
         Sex
    No.  Age    Locality        A  B    C     D   E       F   G   H    I    J
-------------------------------------------------------------------------------
    25030[M]    Esquinapa      .. ..   ..   14.2 8.1[5] 35.6 5.5 33.3 30.8 ..
   126650[F]    Papayo         .. ..   ..   14.7 8.2[5] 38.3 5.4 35.1 32.4
   126651[F]    Do.            .. ..   ..   14.8 7.9[5] 35.6 5.7 32.7 31.1
    17834[M]    P. Negras[6]   95 55   40   14.7 9.0[5] 37.0 5.7 33.8 32.0 15
    17835[F]    Do.            93 55   38   15.6 9.4[5] 37.5 6.0 35.4 32.2 15
    17836[F]    Do.            94 55   39   14.3 8.4[5] 37.6 6.0 34.5 32.7 15
    32112[M]    J. Carranza[7] 94 53   41   14.5 8.9[5] 38.2 5.0 35.1 33.8 16
    32113[M]    Do.            94 57   37   14.2 8.0[5] 36.5 5.3 34.9 32.7 16
    32114[M]    Do.            90 53   37   .... ...    37.0 5.1 34.2 33.0 16
88.8.8.18[F][8] Teapa          .. 46   39   15.6 8.0    38.6 6.2 34.8 33.0
   100231[F][9] Montecristo    99 56   44   15.4 9.6    37.0 6.0 .... ....
  Average                      94 53.8 39.4 14.8 8.6    37.2 5.6 34.4 32.4 15.5
-------------------------------------------------------------------------------

[Note 5: Measured on the dried skin.]

[Note 6: 20 km. WNW Piedras Negras.]

[Note 7: 20 km. ENE Jesús Carranza.]

[Note 8: Type of _Myotis lucifugus fortidens_; measurements after Miller
and Allen, Bull. U. S. Nat. Mus., 144:100, 101, May 25, 1928.]

[Note 9: Type of _Pipistrellus cinnamomeus_ Miller 1902.]

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