Transcriber's Notes
The text presented in this file is that contained in the original
printed version. Only one typographical error was noted in the
conversion of the printed document to digital format.
Typographical Error
Page 598: P.h. veracrucis => P. s. veracrucis
[Pg 591]
A Synopsis of the American Bats of the Genus Pipistrellus
BY
E. RAYMOND HALL and WALTER W. DALQUEST
University of Kansas Publications
Museum of Natural History
Volume 1, No. 26, pp. 591-602, 1 figure in text
January 20, 1950
University of Kansas
LAWRENCE
1950
[Pg 592]
University of Kansas Publications, Museum of Natural HistoryEditors: E. Raymond Hall, Chairman, Edward H. Taylor,
A. Byron Leonard, Robert W. Wilson
Volume 1, No. 26, pp. 591-602, 1 figure in text
January 20, 1950
University of KansasLawrence, Kansas
PRINTED BY
FERD VOILAND, JR., STATE PRINTER
TOPEKA, KANSAS
1950

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[Pg 593]
A Synopsis of the American Bats
of the Genus Pipistrellus
By
E. RAYMOND HALL AND WALTER W. DALQUEST
Four nominal species of the genus Pipistrellus are currently recognized
in North America. They are Pipistrellus subflavus (F. Cuvier)
of eastern North America, Pipistrellus hesperus (H. Allen) of western
North America, Pipistrellus veracrucis (Ward) from Veracruz,
Mexico, and Pipistrellus cinnamomeus Miller from Tabasco, Mexico.
In the past three years, specimens have been obtained in Veracruz
(by Dalquest) of each of the southern species. One of these, P.
cinnamomeus, previously was known from a single specimen; the
other, P. veracrucis, was known only from six specimens which now
are lost or misplaced. The results of our study of these recently
acquired Mexican specimens constitute our principal contribution in
this paper; we have done little more with the material from the
United States and Canada than to codify the findings of other mammalogists
with respect to the systematic status and geographic distribution.
Study of the available specimens reveals that there are only two
species, Pipistrellus hesperus and Pipistrellus subflavus; Pipistrellus
veracrucis proves to be only a subspecies (geographic race) of P.
subflavus, and Pipistrellus cinnamomeus proves to be a species of
another genus, Myotis (see Hall and Dalquest, page 583 of this
volume).
Genus Pipistrellus Kaup
1829. Pipistrellus Kaup, Skizzirte Entw.-Gesch. u. natürl. Syst. europ.
Thierw., Vol. 1, p. 98, Type, Vespertilio pipistrellus Schreber (not seen
by us, after Miller, N. Amer. Fauna, 13:87, 1897).
Remarks.—There are two species in North America. Their geographic
ranges, as now known, meet, but do not overlap. Certain
[Pg 594]
differences between the two species are listed in the parallel columns
below. Most of these differences in the skull and teeth are illustrated
in figures 22 and 23 on page 92 of Miller's "Revision of the North
American bats of the family Vespertilionidae (N. Amer. Fauna, 13,
1897)."
Structure | P. hesperus | P. subflavus |
|
Color | Predominately gray | Predominately brown |
Foot | Less than half as long as tibia | More than half as long as tibia |
Thumb, length of | Less than 4.9 mm. | More than 4.9 mm. |
Tragus | Blunt, terminal part bent forward | Narrow, straight |
Skull (dorsal profile) | Nearly straight | Dish-faced |
Braincase (viewed from above) | Small | Large |
Palate | Extending far behind molars; spine short, narrow at base | Extending short distance long, wide at base |
I2 | Unicuspidate | Bicuspidate |
I3 | Accessory cusp present on anterointernal face | Accessory cusp absent on anterointernal face |
P1 (occlusal view) | Less than a seventh as large as canine | More than a seventh as large as canine |
P1 (labial view) | Concealed by C1 and P4 | Not concealed |
P4 | Touching canine | Not touching canine |
i3 | Touching i2 and c1 | Separated by space from i2 and c1 |
p3 | Lower than anterior cusp of canine | As high as anterior cusp of canine |
Distance from c1 to m1 | Less than length of m2; premolars crowded | More than length of m2; premolars less crowded |
|
[Pg 595]
Pipistrellus hesperus
(Synonomy under subspecies)
Range.—Arid Sonoran life-zones of western North America from Washington
southward to Jalisco.
Characters.—Smoke Gray to Buff Brown (Capitalized color terms after
Ridgway, Color Standards and Color Nomenclature, Washington, D. C., 1912)
dorsally; total length, 60 to 86; foot less than half as long as tibia; tragus
blunt with terminal part bent forward; skull nearly straight in dorsal profile;
inner upper incisor unicuspidate; outer upper incisor with accessory cusp on
anterointernal face; P1, viewed from occlusal face, less than a seventh of area
of canine, and from labial aspect concealed by canine and fourth premolar;
lower, third premolar lower than anterior cusp of canine; lower premolars
crowded, distance between canine and first molar less than length of second
lower molar.
Remarks.—In the United States and in the northern part of Mexico,
P. hesperus is the smallest bat found. Little is known about its
habits. It emerges earlier in the evening than other species of bats.
The frequency with which it is seen near cliffs suggests that it finds
concealment under rocks. In winter, in Nevada (Hall, Mammals of
Nevada, p. 150, 1946), P. hesperus has been found singly in crevices
in the roofs of mine tunnels.
In the United States National Museum in July, 1949, the specimen
providing the easternmost record station of occurrence was examined
by us. This is No. 23591, in alcohol, taken on August 24, 1890, by
William Lloyd, original No. 88, at the mouth of the Pecos River in
Texas. In the same collection there is a specimen of Pipistrellus
subflavus providing the westernmost record of occurrence of that
species. This specimen, a skin with skull, is No. 126729, , taken
on May 3, 1903, by Jas. H. Gaut, original No. 1271, at Comstock,
Texas. The two localities concerned are in the Valley of the Rio
Grande, and are only about five miles apart. Nevertheless, the two
specimens are clearly referable to their respective species and show
no tendency toward intergradation. Consequently, confidence is felt
in treating Pipistrellus hesperus and Pipistrellus subflavus as two
distinct species.
The most recent report upon geographic variation throughout the
entire species, Pipistrellus hesperus, was that by Hatfield (Jour.
Mamm., 17:257-262, August 14, 1936). Later, as explained below
in the account of P. h. australis, Burt (Miscl. Publ., Mus. Zool.,
Univ. Michigan, 39:25, February 15, 1938) examined specimens from
Sonora, Mexico, and for them and for specimens from southern Arizona
proposed a different nomenclatural arrangement.
[Pg 596]

Fig. 1. Map showing the geographic ranges of species and subspecies of Pipistrellus. |
1. Pipistrellus h. hesperus | | 5. Pipistrellus h. santarosae | 2. Pipistrellus h. merriami | | 6. Pipistrellus s. subflavus | 3. Pipistrellus h. australis | | 7. Pipistrellus s. obscurus | 4. Pipistrellus h. maximus | | 8. Pipistrellus s. veracrucis |
|
Pipistrellus hesperus hesperus (H. Allen)
Scotophilus hesperus H. Allen, Smithsonian, Miscl. Coll., No. 165,
Vol. 7 (art. 1): p. 43, June, 1864.
Vesperugo hesperus True, Proc. U. S. Nat. Mus., 7:602, 1885.
Pipistrellus hesperus Miller, N. Amer. Fauna, 13:88, October 16, 1897.
Type locality.—Old Fort Yuma, Imperial County, California, on right bank
of Colorado River, opposite present town of Yuma, Arizona.
Range.—Intermontane region of the United States from south-central Washington
south to Cataviñá, Baja California, and from southeastern California
eastward to southeastern Utah. Marginal occurrences (unless otherwise indicated,
after Hatfield, Jour. Mamm., 17:258, 1936) are: Washington (Dalquest,
Univ. Kansas Publ., Mus. Nat. Hist., 2:165, 1948): Maryhill; Vantage; Almota.
Oregon: Watson. Idaho: 8 mi. W Rogerson (Davis, Mamms. Idaho,
p. 120, 1939). Nevada: Middle Stormy Spring (Hall, Mamms. Nevada, p.
[Pg 597]
151, 1946). Utah: Goodridge. Arizona: 11 mi. NW Kayenta; Tinajas Altas.
Baja California: Cataviñá; San José; Laguna Hanson. California: Dos
Palmos Spring; Banning; Victorville; 12 mi. below (down river) Bodfish;
Little Lake; 2 mi. S Benton Station. Nevada: 2 mi. NW Morgans Ranch;
Deephole. Oregon: Princeton.
Diagnosis.—Size medium for the species; total length, 71.8(66-74); tibia,
12.0(10.7-13.5); forearm, 29.4(27.8-31.8); greatest length of skull, 11.9(11.5-12.3);
breadth of braincase, 6.3(6.1-6.4). Color between Drab Gray and Smoke Gray,
dorsally; between Smoke Gray and Pale Smoke Gray, ventrally (after Hatfield,
Jour. Mamm., 17:257, 1936).
Pipistrellus hesperus merriami (Dobson)
Vesperugo merriami Dobson, Ann. and Mag. Nat. Hist., 18(ser. 5):
124, August, 1896.
Pipistrellus hesperus merriami Grinnell, Proc. California Acad. Sci.,
3(ser. 4):279, August 28, 1913.
Type locality.—Red Bluff, Tehama County, California.
Range.—California west of the Sierra Nevada; the Sacramento Valley, the
San Joaquin Valley, and the Coast Range from San Francisco Bay south to
San Diego County. Marginal occurrences (after Hatfield, Jour. Mamm., 17:260,
1936, unless otherwise noted) are: California: Dales on Paines Creek; Fyffe;
Yosemite Valley; Shaver Ranger Station; Springville; Fort Tejon; Painted
Gorge (P. H. Krutzsch, MS); Carrizo Creek; thence northward up the coast
probably to San Francisco Bay; in the Sacramento Valley west to Rumsey.
Diagnosis.—Size medium for the species; total length, 71.3(66-78); tibia,
11.2(10.6-11.7); forearm, 28.9(27.5-30.8); greatest length of skull, 11.8(11.3-12.2);
breadth of braincase, 6.4(6.0-6.6). Color Buffy Brown to Army Brown, dorsally;
Wood Brown to Buffy Brown, ventrally (after Hatfield, op. cit.: 258, 260).
Pipistrellus hesperus australis Miller
Pipistrellus hesperus australis Miller, N. Amer. Fauna, 13:90, October
16, 1897.
Pipistrellus hesperus apus Elliot, Field Columb. Mus., pub. 90, zool.
ser., 3:269, March 8, 1904. Type from Providencia Mines, Sonora,
Mexico.
Type locality.—Barranca Ibarra, Jalisco, Mexico.
Range.—Central Arizona south to Jalisco and including the southern half
of Baja California. Marginal occurrences (after Hatfield, op. cit.: 261, unless
otherwise indicated) are: Arizona: Camp Verde; Fort Bowie. Sonora:
Pilares (Burt, Miscl. Publ., Mus. Zool., Univ. Michigan, 39:24, 1938). Jalisco:
Barranca Ibarra (Miller, orig. descr.). Baja California: Miraflores; San
Ignacio. Arizona: Bates Well.
Diagnosis.—Size small for the species; total length, 67.1(60-72); tibia,
11.3(10.1-12.3); forearm, 28.4(26.3-30.0); greatest length of skull, 11.7(11.3-12.0);
breadth of braincase, 6.1(5.9-6.3). Color: between Cinnamon Drab and Drab,
dorsally; Wood Brown to Light Drab, ventrally (after Hatfield, op. cit.:260).
Remarks.—Hatfield (op. cit.) examined no specimens from Mexico
(Baja California excepted) and Burt (op. cit.) who did examine
[Pg 598]
some specimens (from Sonora), referred one from northwestern
Sonora to P. h. hesperus and those from northeastern Sonora to P. h.
merriami. Since our treatment of subspecies of Pipistrellus (P. s.
veracrucis excepted) aims merely to reflect the latest systematic
treatment accorded the animals, we would follow Burt (op. cit.)
were it not for the fact that he shows the geographic range of P. h.
merriami separated by the range of P. h. hesperus into two parts.
This is inconsistent with the ordinarily accepted concept of subspecies.
Consequently, we have followed Hatfield (op. cit.). Clearly,
a critical study is needed of adequate material of Pipistrellus hesperus
of Mexico.
Pipistrellus hesperus maximus Hatfield
Pipistrellus hesperus maximus Hatfield, Jour. Mamm., 17:261, August
14, 1936.
Type locality.—Dog Spring, Hidalgo County, New Mexico.
Range.—Southern New Mexico, western Texas and probably the adjoining
parts of Mexico. Marginal occurrences (after Hatfield [op. cit.:261] except
as otherwise indicated) are: New Mexico: Animas Valley; Florida Mountains;
Carlsbad Cave. Texas: Mouth of Pecos River (Bailey, N. Amer. Fauna,
25:210, 1905); Boquillas (Borell and Bryant, Univ. California Publ. Zool.,
48:9, 1942); Glen Spring (Borell and Bryant, loc. cit.).
Diagnosis.—Size large for the species; total length, 80.3(78-83); tibia,
12.3(11.7-13.1); forearm, 32.9(31.8-33.3); greatest length of skull, 12.7(12.3-12.9);
breadth of braincase, 6.6(6.5-6.7). Color between Smoke Gray and Pale Drab
(after Hatfield, op. cit.:261).
Pipistrellus hesperus santarosae Hatfield
Pipistrellus hesperus santarosae Hatfield, Jour. Mamm., 17:261, August
14, 1936.
Type locality.—Santa Rosa, Guadalupe County, New Mexico.
Range.—New Mexico (excepting southern part) and western Colorado.
Marginal occurrences (after Hatfield, op. cit.:262) are: Colorado: Bedrock.
New Mexico: Santa Rosa; Socorro; Laguna.
Diagnosis.—Size large for the species; total length, 82.0(80-86); tibia,
12.4(11.9-13.0); forearm, 32.8(31.7-34.1); greatest length of skull, 12.7(12.3-13.1);
breadth of braincase, 6.6(6.3-6.8). Color between Buffy Brown and Wood
Brown (after Hatfield, op. cit.:261, 262).
Pipistrellus subflavus
(Synonomy under subspecies)
Range.—Canadian to Tropical life-zones of eastern North America from
Quebec southward to Honduras.
Characters.—Sayal Brown to darker than Mummy Brown, dorsally; total
length, 73-89; foot more than half as long as tibia; tragus tapering and straight;
[Pg 599]
dorsal profile of skull convex in interorbital region; inner upper incisor bicuspidate;
outer upper incisor unicuspidate (lacking accessory cusp on anterointernal
face); P1 viewed from occlusal face more than a seventh of area of
canine and visible from labial aspect; lower, third premolar as high as anterior
cusp of canine; lower premolars less crowded than in P. hesperus and
distance between canine and first molar less than length of second lower molar.
Remarks.—In winter this species hibernates in caves in clusters
of fewer than fifty individuals, but in summer fewer of the bats live
there and at this season some have been captured as far as thirty
miles from any such retreat suggesting that the bats inhabit other
types of shelter. The wide range of this species in respect to life-zones
is noteworthy; it occurs in the Canadian Life-zone (Joliet,
Quebec), the Tropical Life-zone (30 km. SSE Jesús Carranza, Veracruz)
and in the intervening life-zones.
The longer thumb of this species, in comparison with that of
Pipistrellus hesperus, was verified by measuring the thumb including
its claw and the pad at the base of the thumb in 12 P. s. veracrucis
and 10 P. h. maximus. In veracrucis the mean was 5.9 millimeters
and the extremes were 5.5 and 6.4. In maximus the
corresponding figures were 3.9, 3.6 and 4.3.
Pipistrellus subflavus subflavus (F. Cuvier)
V[espertilio]. subflavus F. Cuvier, Nouv. Ann. Mus. Hist. Nat. Paris,
1: 17, 1832.
Vespertilio erythrodactylus Temminck, Monogr. de Mamm., II, 13me
monogr., p. 238, 1835-1841 (not seen—after Miller, N. Amer. Fauna,
13:90, October 16, 1897).
Scotophilus georgianus H. Allen, Smithsonian Miscl. Coll., No. 165,
Vol. 7 (art. 1), p. 35, June, 1864.
Vesperugo carolinensis H. Allen, U. S. Nat. Mus. Bull. 43:121, March
14, 1894.
Pipistrellus subflavus Miller, N. Amer. Fauna, 13:90, figs. 22, 23, October
16, 1897.
Type locality.—Eastern United States, probably Georgia.
Range.—From approximately 40 degrees North Latitude in Pennsylvania
and Kansas southward to central Florida and at least to extreme southern
Texas; from the Atlantic Coast westward to south-central Kansas and Val
Verde County, Texas. Marginal occurrences are: Kansas (K. U. Collection):
4½ mi. SW Sun City; Ft. Leavenworth. Illinois (Necker and Hatfield, Bull.
Chicago Acad. Sci., 6(3):45, 1941): Quincy; Urbana. Indiana (Lyon, Amer.
Midland Nat., 17:73, 1936): Monroe County; Franklin Co. Ohio (Bole
and Moulthrop, Sci. Publs. Cleveland Mus. Nat. Hist., 5(6):115, 1942: Hamilton
Co.; Smoky Creek. West Virginia (Kellogg, Proc. U. S. Nat. Mus.,
84:449, 1937): Charleston; Smoke Hole Cave. Pennsylvania (Rhoads,
Mamms. Pa. and N. J., p. 211, 1903): Carlisle; Germantown. New Jersey:
Haddonfield (Rhoads, Mamms. Pa. and N. J., p. 211, 1903). Florida: Tarpon
Springs (Sherman., Proc. Florida Acad. Sci., p. 107, 1936). Texas: Brownsville
[Pg 600]
(Bailey, N. Amer. Fauna, 25:211, 1905); Comstock (Bailey, loc. cit.); Kerr
Co. (Taylor and Davis, Game, Fish and Oyster Comm. Bull., 50:17, 1947).
Oklahoma: 10 mi. S and 2 mi. E Sulphur (Blair, Amer. Midland Nat., 22:100,
1939).
Diagnosis.—Size large; eight specimens from Barber and Butler counties,
Kansas, measure in total length, 84(77-89); tibia, 14.8(14.5-15); forearm,
33.5(31.8-35.3); greatest length of skull (exclusive of incisors), 12.8(12.3-13.1);
breadth of braincase immediately above roots of zygomatic arches, 6.5(6.4-6.7).
Color ranging from Snuff Brown to Sayal Brown.
Pipistrellus subflavus obscurus Miller
Pipistrellus subflavus obscurus Miller, N. Amer. Fauna, 13:93, October
16, 1897.
Type locality.—Lake George, Warren County, New York.
Range.—From southern Quebec and southern Ontario south to southern
Ohio and West Virginia; from the Atlantic Coast west into Wisconsin. Marginal
occurrences are: Minnesota: St. Peter (Swanson and Evans, Jour.
Mamm., 17:39, 1936); Marine (Swanson, Tech. Bull. No. 2, Minnesota Dept.
Conservation, p. 60, 1945). Wisconsin: Hurley (Greeley and Beer, Jour.
Mamm., 30:198, 1949). Quebec: Joliet (Anderson, Nat. Mus. Canada, Biol.
ser. No. 31, Bull. 102:30, 1946). Vermont: Brandon (Osgood, Jour. Mamm.,
19:436, 1938). Maine: No locality more precise than the state (Allen, Occ.
Papers Boston Soc. Nat. Hist., 7(3):35, June, 1904). New York: Hastings on
Hudson (Rowley, Abstr. of Proc. Linnean Soc. N. Y., for yr. ending March 11,
1902, p. 57). Pennsylvania: Beaver (Rhoads, Mamms. Pa. and N. J., 1903,
p. 211). West Virginia: Cornwall's Cave (Frum, Jour. Mamm., 25:195, 1944).
Ohio: Cat Run (Bole and Moulthrop, Sci. Publs. Cleveland Mus. Nat. Hist.,
5(6):116, 1942); Symmes Creek (Bole and Moulthrop, loc. cit.); Dry Cave
(Bole and Moulthrop, loc. cit.); "Union County" (Rausch, Jour. Mamm.,
27:275, 1946). Wisconsin: Devils Lake (Jackson, Jour. Mamm., 1:38, 1919).
Diagnosis.—"... color duller and less yellow, and dark tips of shorter
hairs on back more conspicuous" than in P. subflavus subflavus according to
the original description.
Remarks.—No one, as far as we know, has carefully studied the
variation in Pipistrellus subflavus of the United States and Canada
since Miller named P. s. obscurus. With the more abundant material
now available, such an appraisal would be worth-while. The
occurrences cited above for Minnesota and Wisconsin were recorded
in the literature under the specific name without indication of subspecific
affinity. The reference of specimens from these states to
the subspecies P. s. obscurus is an arbitrary assignment on our part;
we have not seen them. However, two specimens in the University
of Kansas Museum of Natural History from Potosi (Snake Cave)
Grant County, Wisconsin, are referable to P. s. obscurus. These
provide the southwesternmost record station of occurrence in Wisconsin
[Pg 601]
but are not shown on the distribution map because the specimens
were received after figure 1 was prepared.
It is noteworthy that the species Pipistrellus subflavus has not
yet, as far as we can ascertain, been recorded from Michigan, northern
Indiana, northern Illinois, or Iowa. Probably the species occurs
in these areas.
Pipistrellus subflavus veracrucis (Ward)
Vesperugo veracrucis Ward, Amer. Nat., 25:745, August, 1891.
Pipistrellus veracrucis Miller, N. Amer. Fauna, 13:93, October 16,
1897.
Type locality.—Las Vigas, 8,500 ft., Veracruz.
Range.—Eastern Mexico, certainly from the type locality southward into
Honduras. Records of occurrence are: Veracruz: Las Vigas (13 specimens
from 4 km. E Las Vigas, 8,500 ft., K. U.); 30 km. SSE Jesús Carranza, 1
(K. U.). Honduras: Jilamo Farm, Tela District, 3 (Univ. Michigan).
Diagnosis.—Size small for the species; measurements of 13 near topotypes
are: total length, 78(73-85); tibia, 12.9(11.8-14.7); forearm, 31.8(29.5-33.1);
greatest length of skull (exclusive of incisors), 12.2(11.8-12.6); breadth of braincase
immediately above roots of zygomatic arches, 6.3(6.0-6.7). Color darker
than Mummy Brown above and below.
Remarks.—The specimen from thirty kilometers south-southeast
of Jesús Carranza, Veracruz, and the three specimens from Honduras
agree in all respects with topotypes. The color of P. s. veracrucis
is much darker than that of P. s. obscurus and is between black and
the darkest brown in Ridgway's (op. cit.) color key. Rinker (Jour.
Mamm., 29:179-180,1948) described the three specimens from Honduras
without assigning a specific name to them because he lacked
topotypes of P. s. veracrucis. We find nothing in his description to
correct, but can add that the upper tooth-rows in many, but not in
all, specimens of P. s. veracrucis are straighter than in P. s. subflavus.
Probably it was this feature to which Rinker referred when
he said that in veracrucis "The tooth rows tend to be more convergent
posteriorly." Rinker did not refer the three specimens from
Honduras to P. veracrucis because Ward's original description states
that veracrucis has evenly spaced lower incisors and a basal cusp
on the lower canine on only its forward edge. Rinker's specimens
from Honduras have the first incisors in contact with each other,
the second incisors in contact with the first incisors and the third
incisor on each side of the lower jaw separated by a space from the
second incisor and from the canine. The specimens from Honduras
have a basal cusp on the hinder edge of the lower canine. In these
two features they agree with the specimens from Veracruz and with
[Pg 602]
specimens of Pipistrellus subflavus from the United States and Canada.
It is clear that Ward (Amer. Nat., 25:747,1891) was mistaken
in stating that the lower incisors of veracrucis were evenly spaced
and that the canine had a basal cusp on only the forward edge.
Ward (loc. cit.) was correct in regarding his Vesperugo veracrucis
as "most closely related to V. georgianus [= Pipistrellus subflavus],"
but for want of actual specimens of P. subflavus to use in comparison
was incorrect in supposing that P. subflavus had only two bands
of color on the fur, more hair on the legs, and a larger area of hair
on the interfemoral membrane. In these respects we perceive no
difference between specimens from Veracruz and the United States.
Vesperugo veracrucis Ward, therefore, proves to be only a subspecies
of Pipistrellus subflavus, but is well characterized by dark
color and small size.
University of Kansas Museum of Natural History, Lawrence,
Kansas.
Transmitted October 31, 1949.

23-1546
Comments on "A Synopsis of the American Bats of the Genus Pipistrellus" :