The Project Gutenberg eBook of 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

Pipistrellus cinnamomeus Miller 1902
Referred to the Genus Myotis

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."[Pg 584] 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[Pg 585] 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:[Pg 586]

Myotis fortidens Miller and Allen

Cinnamon Myotis

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.

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[Pg 587] (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.

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.[Pg 588]

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.

University of Kansas Museum of Natural History, Lawrence, Kansas. Transmitted October 31, 1949.[Pg 589]

Cranial Measurements of Myotis fortidens

No.	Sex Age	Locality	Greatest length	Condylobasal length	Zygomatic breadth	Interorbital constriction	Breadth of braincase	Mandible	Maxillary tooth-row	Maxillary breadth at M3	Mandibular tooth-row	Wear of teeth
25030	♂	Esquinapa	....	....	...	...	...	....	...	...	...	?
126650	♀	Papayo	15.0	14.2	9.7	3.9	7.1	11.5	5.5	5.6	6.0	0
126651	♀	Do.	15.1	13.8	9.4	3.8	6.8	10.6	5.6	5.9	6.0	0
17834	♂	P. Negras[1]	....			4.1		10.6	5.6	5.7	6.0	0
17835	♀	Do.	15.5	14.9	9.6	4.2	7.2	11.0	5.7	6.0	6.1	2
17836	♀	Do.	15.5	14.5	9.7	4.2	7.3	10.9	5.4	5.9	5.7	3
32112	♂	J. Carranza[2]	15.3	14.4	9.7	4.1	7.3	11.5	5.7	5.9	6.3	1
32113	♂	Do.	15.0	14.0	9.5	4.2	7.2	10.9	5.5	5.9	5.9	1
32114	♂	Do.	15.0	13.9	9.7	4.1	7.2	10.8	5.4	6.0	5.9	1
88.8.8.18	♀[3]	Teapa	15.0	13.8	9.6	3.8	7.4	....	5.4	5.8	5.8	1
100231	♀[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 3: Type of Myotis lucifugus fortidens; measurements after Miller and Allen, Bull. U. S. Nat. Mus., 144:100; 101, May 25, 1928.]

External Measurements of Myotis fortidens

No.	Sex Age	Locality	Total length	Head and body	Tail	Tibia	Foot	Forearm	Thumb	Third metacarpal	Fifth metacarpal	Ear from notch
25030	♂	Esquinapa	..	..	..	14.2	8.1[5]	35.6	5.5	33.3	30.8	..
126650	♀	Papayo	..	..	..	14.7	8.2[5]	38.3	5.4	35.1	32.4
126651	♀	Do.	..	..	..	14.8	7.9[5]	35.6	5.7	32.7	31.1
17834	♂	P. Negras[6]	95	55	40	14.7	9.0[5]	37.0	5.7	33.8	32.0	15
17835	♀	Do.	93	55	38	15.6	9.4[5]	37.5	6.0	35.4	32.2	15
17836	♀	Do.	94	55	39	14.3	8.4[5]	37.6	6.0	34.5	32.7	15
32112	♂	J. Carranza[7]	94	53	41	14.5	8.9[5]	38.2	5.0	35.1	33.8	16
32113	♂	Do.	94	57	37	14.2	8.0[5]	36.5	5.3	34.9	32.7	16
32114	♂	Do.	90	53	37	....	...	37.0	5.1	34.2	33.0	16
88.8.8.18	♀[8]	Teapa	..	46	39	15.6	8.0	38.6	6.2	34.8	33.0
100231	♀[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 8: Type of Myotis lucifugus fortidens; measurements after Miller and Allen, Bull. U. S. Nat. Mus., 144:100, 101, May 25, 1928.]

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