1"Rt. 98" and "15 mi. S Waukenna" both in Jefferson Co.
2Only nine specimens.
3N Island, Grand Lake, Iberville Parish."
4Some in Amer. Mus. Nat. History.
5Females, 8; males, 3; unsexed, 1.
6Only 11 specimens.
710 mi. W, 2 mi. S Piedra, Tamaulipas.
816 mi. W, 3 mi. S Piedra, Tamaulipas.
Lasiurus ega
Southern Yellow Bat
Diagnosis.—Upper parts yellowish-brown (much as inLasiurus intermedius floridanusfrom Louisiana) having overlay of grayish or blackish anterior to shoulders; hair on basal half of interfemoral membrane more yellowish than elsewhere; size medium (forearm 42.7-52.2; condylocanine length 14.6-16.3).
Diagnosis.—Upper parts yellowish-brown (much as inLasiurus intermedius floridanusfrom Louisiana) having overlay of grayish or blackish anterior to shoulders; hair on basal half of interfemoral membrane more yellowish than elsewhere; size medium (forearm 42.7-52.2; condylocanine length 14.6-16.3).
This species occurs from the southwestern United States (Palm Springs, California, and Tucson, Arizona) southward into Uruguay and northeastern Argentina. Of the six currently (see Handley, 1960) recognized subspecies ofL. ega, four occur only in South America, and two occur only in North America.
Cabrera (1958:115) regardedDasypterus ega fuscatusThomas (1901:246), based on three specimens from Río Cauquete, Río Cauca, Colombia, as a synonym ofDasypterus ega panamensisThomas (loc. cit.) that was based on a specimen from Bogava, 250 meters elevation, Chiriquí, Panamá. The latter name has line priority overfuscatus. Cabrera (1958:116) remarked that: "Las diferencias que Thomas señaló entre elDasypterusde Panamá y el de Colombia (fuscatus) nos parecen estar dentro de los límites de la variación individual, siendo además muy raro que una especie de quiróptero este representada en Colombia y en Panamá por razas diferentes."
On July 16, 1958, at the British Museum of Natural History, one of us (Hall) examined the holotypes ofpanamensisandfuscatus, as well as other materials used by Thomas, and readily perceived the differences that he pointed out. Thomas' description, although terse, is accurate.L. e. fuscatusis much more blackish thanpanamensis. We are inclined to retain the two names as applicable to two subspecies. Whether or notfuscatusis synonymized underpanamensis, the holotype ofpanamensisis an intergrade between the almost black Colombian animal (fuscatus) and the paler individuals in Central America and territory north thereof. Even so, the holotype ofpanamensismore closely resembles the blackish Colombian population than the paler populations to the north and the namepanamensis, therefore, is correctly applicable to the bat from Panamá, but not to bats of the speciesLasiurus egafrom farther north as most authors (see, for example, Hall and Kelson, 1959:194, map 143; and Handley, 1960:474) suggested was the case. For the populations north of Panamá the nameLasiurus ega xanthinus(Thomas) (1897:544) needs to be used.
Lasiurus ega xanthinus(Thomas)
1897.Dasypterus ega xanthinusThomas, Ann. Mag. Nat. Hist., ser. 6, 20:544, December, type from Sierra Laguna, Baja California.1953.Lasiurus ega xanthinus, Dalquest, Louisiana State Univ. Studies, Biol. Ser., 1:61, December 28.Geographic distribution.—Southern California, southern Arizona, and northern Coahuila southward through México to southern Costa Rica.Diagnosis.—Yellowish-brown with an overlay of grayish anterior to the shoulders; forearm, 42.7-47.2.Remarks.—Specimens from Baja California and the adjacent western part of the mainland of México average paler than specimens from Veracruz and some places in Central America but the differences are slight.Records of occurrence.—Specimens examined, 21, as follows:Baja California.—Comondú, 1 (USNM); Sierra Laguna, 4 (1 USNM, 3 British Mus.).Coahuila.—4 mi. W Hacienda La Mariposa, 2300 ft., 2 (KU).Zacatecas.—Concepción del Oro, 7680 ft., 4 (KU).Tamaulipas.—Sierra de Tamaulipas, 1200 ft., 10 mi. W, 2 mi. S Piedra, 5 (KU); 16 mi. W, 3 mi. S Piedra, 1 (KU).Sinaloa.—1 mi. S Pericos, 1 (KU).Veracruz.—Achotal, 1 (Chicago Mus.).Yucatan.—Yaxcach, 1 (USNM).Costa Rica.—Lajas, Villa Quesada, 1 (AMNH); San José, 1 (AMNH).Additional records:California: Palm Springs (Constantine, 1946:107).Arizona: Tucson (Cockrum, 1961:97).Baja California(Handley, 1960:474): Santa Ana; Miraflores.Sinaloa: Escuinapa (Handley, 1960:475).Durango: Aguajequiroz, 12 mi. SSW Mapimí, 5000 ft. (Greer, 1960:511).San Luis Potosi(Dalquest, 1953:62): 1½ mi. E Río Verde; 19 km. SW Ebano; 4 mi. SSW Ajinche.Quintana Roo: 7 mi. N, 37 mi. E Puerto de Morelos (Ingles, 1959:384).Honduras: Tegucigalpa (Handley, 1960:474).
Geographic distribution.—Southern California, southern Arizona, and northern Coahuila southward through México to southern Costa Rica.
Diagnosis.—Yellowish-brown with an overlay of grayish anterior to the shoulders; forearm, 42.7-47.2.
Remarks.—Specimens from Baja California and the adjacent western part of the mainland of México average paler than specimens from Veracruz and some places in Central America but the differences are slight.
Records of occurrence.—Specimens examined, 21, as follows:Baja California.—Comondú, 1 (USNM); Sierra Laguna, 4 (1 USNM, 3 British Mus.).Coahuila.—4 mi. W Hacienda La Mariposa, 2300 ft., 2 (KU).Zacatecas.—Concepción del Oro, 7680 ft., 4 (KU).Tamaulipas.—Sierra de Tamaulipas, 1200 ft., 10 mi. W, 2 mi. S Piedra, 5 (KU); 16 mi. W, 3 mi. S Piedra, 1 (KU).Sinaloa.—1 mi. S Pericos, 1 (KU).Veracruz.—Achotal, 1 (Chicago Mus.).Yucatan.—Yaxcach, 1 (USNM).Costa Rica.—Lajas, Villa Quesada, 1 (AMNH); San José, 1 (AMNH).
Additional records:California: Palm Springs (Constantine, 1946:107).Arizona: Tucson (Cockrum, 1961:97).Baja California(Handley, 1960:474): Santa Ana; Miraflores.Sinaloa: Escuinapa (Handley, 1960:475).Durango: Aguajequiroz, 12 mi. SSW Mapimí, 5000 ft. (Greer, 1960:511).San Luis Potosi(Dalquest, 1953:62): 1½ mi. E Río Verde; 19 km. SW Ebano; 4 mi. SSW Ajinche.Quintana Roo: 7 mi. N, 37 mi. E Puerto de Morelos (Ingles, 1959:384).Honduras: Tegucigalpa (Handley, 1960:474).
Lasiurus ega panamensis(Thomas)
1901.Dasypterus ega panamensisThomas, Ann. Mag. Nat. Hist., ser. 7, 8:246, September, type from Bogava [= Bugaba], Chiriquí, 250 meters, Panamá.1960.Lasiurus ega panamensis, Handley, Proc. U.S. Nat. Mus., 112:474, October 6.Geographic distribution.—Panamá; also recorded by Handley (1960:474) from Venezuela.Diagnosis.—"General colour dark brownish clay-color, something between Ridgway's 'raw-umber' and 'clay-color'. Fur black basally, then dull brownish buffy, the extreme tips black. Center of face similar to back, cheeks from eyes to lips contrasting black. Rump and hairy part of interfemoral verging toward brownish fulvous. Under surface similar to upper." (Thomas, 1901:246.) Forearm of holotype, 46.5.Remarks.—Notes taken down by one of us (Hall) on July 16, 1958, at the British Museum, Natural History, contain the following: "Color accurately described by Thomas. The blackish stands out. The difference between the types ofD. e. panamensisandD. e. xanthinusis tremendous."Record of occurrence.—Specimen examined, one, the type (British Mus.).
Geographic distribution.—Panamá; also recorded by Handley (1960:474) from Venezuela.
Diagnosis.—"General colour dark brownish clay-color, something between Ridgway's 'raw-umber' and 'clay-color'. Fur black basally, then dull brownish buffy, the extreme tips black. Center of face similar to back, cheeks from eyes to lips contrasting black. Rump and hairy part of interfemoral verging toward brownish fulvous. Under surface similar to upper." (Thomas, 1901:246.) Forearm of holotype, 46.5.
Remarks.—Notes taken down by one of us (Hall) on July 16, 1958, at the British Museum, Natural History, contain the following: "Color accurately described by Thomas. The blackish stands out. The difference between the types ofD. e. panamensisandD. e. xanthinusis tremendous."
Record of occurrence.—Specimen examined, one, the type (British Mus.).
RELATIONS BETWEEN THE SPECIES OF LASIURUS
As suggested by Dalquest in 1953 (p. 62) and by Handley in 1959 (p. 119) and 1960 (p. 473), the yellow bats,Lasiurus ega(Gervais) andLasiurus intermediusH. Allen, so closely resemble the hoary bat,Lasiurus cinereus(Palisot de Beauvois), and the red bats,Lasiurus borealis(Müller) and the seven related species listed below, that all are properly included in a single genus. Many of the common characteristics are enumerated above in the diagnosis of the genus (see also Handley, 1960:473).
Diagram of bones
Fig. 3.Diagram of bones of right arm and third finger (middle digit) including cartilage on distal end of terminal (3rd) phalanx. Percentages are in terms of the over-all length of the arm and third finger.
A listing of the differences between the species is less impressive than a listing of the resemblances. The yellow bats differ less from the red bats than does the hoary bat,L. cinereus, which differs from all of the others as follows: talonid on m3 larger; p4 single-rooted instead of double-rooted; hypocone on M1 and M2 smaller; coronoid process lower; ossified part of tympanic ring, which shields the petrosal, larger; humerus relatively shorter; forearm relatively longer; first phalanx of middle finger relatively shorter; presternum including keel longer than wide instead ofvice versa. The differences in the sternum and proportions of the forelimb reflect the more rapid flight of the hoary bat. The yellow bats differ from the red bats and hoary bat in long rostrum, pronounced sagittal crest, high coronoid process, absence of the first upper premolar, long first phalanx of the third digit and short terminal (3rd) phalanx of the same digit. Features in which the red bats are extreme in the genus are short rostrum, short forearm, and relatively longer second phalanx of the third finger. The red bats differ only slightly one from another.
Next to nothing is known of extinct Tertiary ancestors of species of the genusLasiurus. Also relatively little is known aboutLasiurusin the Pleistocene. Consequently, evolution of the living species has to be inferred almost entirely from what is known about their structure, habits, and geographic distribution. Figure 4 presents some ideas concerning relationships.
GraphFig. 4.Postulated relationships of species of the genusLasiurus.
LIST OF NAMED KINDS OF THE GENUS LASIURUS
The words "type from" indicate that a specimen or specimens served as basis for the name. The words "type locality" signify lack of knowledge as to whether a specimen was preserved.
Red Bats
Lasiurus borealis borealis(Müller), 1776, type from New York.
[Vespertilio]noveboracensisErxleben, 1777, based, in part, on "Der Neujorker" of Müller (ante).
Vespertilio lasiurusSchreber, 1781, type locality, North America.
Vespertilio rubellusPalisot de Beauvois, 1796, type locality unknown.
Vespertilio rubraOrd, 1815, based on the red bat of Wilson, Amer. Ornith., 6:60.
Vespertilio tesselatusRafinesque, 1818, type locality unknown.
Vespertilio monachusRafinesque, 1818, type locality unknown.
Vespertilio rufusWarden, 1820, based on the red bat of Wilson,ibid.
Lasiurus funebrisFitzinger, 1870, type locality, Tennessee.
Myotis quebecensisYourans, 1930, type from Anse-à-Wolfe, Quebec.
Lasiurus borealis frantzii(Peters), 1871, type from Costa Rica.
Lasiurus borealis teliotis(H. Allen), 1891, type probably from California.
Lasiurus borealis ornatusHall, 1951, type from Penuela, Veracruz.
Lasiurus borealis varius(Poeppig), 1835, type from Antuco, Provincia de Bió-Bió, Chile.
Nycticeus poepingiiLesson, 1836, type from Chile.
Lasiurus borealis salinaeThomas, 1902, type from Cruz del Eje, Cordoba, Argentina.
Lasiurus borealis blossevilliiLesson and Garnot, 1826, type from Montevideo, Uruguay.
Vespertilio bonariensisLesson, 1827, type from Buenos Aires, Argentina.
Lasiurus ensleniiLima, 1926, type from São Lourenço, Rio Grande do Sul, Brazil.
Lasiurus pfeifferi(Gundlach), 1861, type from Cuba.
Lasiurus degelidusMiller, 1931, type from Sutton's, District of Vere, Jamaica.
Lasiurus minorMiller, 1931, type from "Voute l'Eglise," 1350 ft., a cave near the Jacmel road a few kilometers N Trouin, Haiti.
Lasiurus seminolus(Rhoads), 1895, type from Tarpon Springs, Pinellas Co., Florida.
Lasiurus castaneusHandley, 1960, type from Tacarcuna Village, 3200 ft., Río Pucro, Darién, Panamá.
Lasiurus egregius(Peters), 1871, type from Santa Catarina, Brazil.
Lasiurus brachyotis(J. A. Allen), 1892, type from San Cristóbal Island, Galapagos Islands.
Yellow Bats
Lasiurus golliheri(Hibbard and Taylor), Contributions Mus. Paleo., Univ. Michigan, 16:162, fig. 10F, July 1, 1960 [an extinct species], type from [a stratum of Late Pleistocene Age] "Below the caliche bed in the Kingsdown formation; Cragin Quarry local fauna, locality 1 (Sangamon age); Big Springs Ranch, SW ¼ sec. 17, T. 32 S., R. 28 W. (Kansas University Locality 6), Meade County, Kansas."
Lasiurus ega xanthinus(Thomas), 1897, type from Sierra Laguna, Baja California.
Lasiurus ega panamensis(Thomas), 1901, type from Bugaba, Chiriquí, Panamá.
Lasiurus ega fuscatus(Thomas), 1901, type from Río Cauquete, Colombia.
Dasypterus ega punensisJ. A. Allen, 1914, type from Isla de Puná, Ecuador.
Lasiurus ega ega(Gervais), 1856, type from Ega, Estado de Amazonas, Brazil.
class="indent2"Lasiurus caudatusTomes, 1857, type from Pernambuco, Brazil.
Lasiurus ega argentinus(Thomas), 1901, type from Goya, Province of Corrientes, Argentina.
Lasiurus intermedius intermediusH. Allen, 1862, type from Matamoros, Tamaulipas, México.
Lasiurus intermedius floridanus(Miller), 1902, type from Lake Kissimmee, Osceola Co., Florida.
Lasiurus intermedius insularisHall and Jones, 1961, type from Cienfuegos, Las Villas Province, Cuba.
Hoary Bats
Lasiurus fossilisHibbard, Contributions Mus. Paleo., Univ. Michigan, 8(No.6): 134, fig. 5, June 20, 1950 [an extinct species], type from [an early Pleistocene or a late Pliocene deposit] "Rexroad formation, Rexroad fauna. Locality UM-K1-47, Fox Canyon, XI Ranch, Meade County, Kansas."
Lasiurus cinereus cinereus(Palisot de Beauvois), 1796, type from Philadelphia, Pennsylvania. Known from Late Pleistocene time as well as from Recent time (see Hibbard and Taylor, Contributions Mus. Paleo., Univ. Michigan, 16:159, fig. 10A, July 1, 1960, for occurrence in Cragin Quarry local fauna, Sangamon Age, Meade County, Kansas).
Vespertilio pruinosusSay, 1823, type from Engineer Cantonment, Washington Co., Nebraska.
A[talapha]. mexicanaSaussure, 1861, type from an unknown locality, probably from Veracruz, Puebla, or Oaxaca.
Lasiurus cinereus villosissimusÉ. Geoffroy St.-Hilaire, 1806, type locality, Asunción, Paraguay.
Lasiurus grayiTomes, 1857, type from Chile.
Atalapha pallescensPeters, 1871, type from Paramo de la Culata, Andes de Mérida, Venezuela.
Atalapha cinerea brasiliensisPira, 1905, type from Ignape, São Paulo, Brazil.
Lasiurus cinereus semotus(H. Allen), 1890, type from Hawaii.
EXPLANATION AND ACKNOWLEDGMENTS
Hall and Jones are jointly responsible for the accounts of the two species of yellow bats, but Hall alone assumes responsibility for the other parts of the paper. Thanks are extended to the National Science Foundation for financial support (Grant No. 56 G 103) of the study here reported on. We are grateful also to the following persons for the loan of specimens in their care: S. B. Benson, Museum of Vertebrate Zoology, University of California (MVZ); W. F. Blair, Department of Zoology, University of Texas (Univ. Texas); W. B. Davis, Dept. Wildlife Management, Agricultural and Mechanical College of Texas (TAMC or Texas A & M); D. H. Johnson, C. O. Handley, Jr., and W. H. Setzer, U.S. National Museum (USNM); Barbara Lawrence, Museum of Comparative Zoology at Harvard College (MCZ); J. N. Layne, Department of Biology, University of Florida (UF); G. H. Lowery, Jr., Museum of Natural History, Louisiana State University (LSU); P. J. H. van Bree, Department of Mammals, Zoölogisch Museum, Amsterdam (AZM); and R. G. Van Gelder, American Museum of Natural History (AMNH). Thanks are extended also to E. T. Hooper and W. H. Burt, Mus. Zoology, University of Michigan (Univ. Mich.), to Philip Hershkovitz, Chicago Natural History Museum (Chicago Mus.), and to Peter Crowcroft, British Museum, Natural History, for permission to examine specimens there. Mr. Gilberto Silva Taboada arranged the loan of specimens from the Poey Museum, University of Havana and from the Ramsdem Museum, University of Oriente, both in Cuba. Mr. Silva Taboada and Dr. Carlos G. Aguayo of the Poey Museum graciously arranged an exchange of specimens whereby the holotype ofL. i. insularisbecame the property of the Museum of Natural History, University of Kansas. Specimens in the last mentioned institution are identified with the symbol KU.
LITERATURE CITED
Allen, H.
Baker, R. H., and Dickerman, R. W.
Blair, W. F.
Borkhausen, M. B.
Cabrera, A.
Cockrum, E. L.
Coleman, R. H.
Constantine, D. G.
Dalquest, W. W.
Davis, W. B.
de Beaufort, L. F.
Eads, R. B., Menzies, G. C., and Wiseman, J. S.
Frye, O. E., Jr.
Gaumer, G. F.
Gervais, P.
Gray, J. E.
Greer, J. K.
Hall, E. R., and Kelson, K. R.
Hamilton, W. J., Jr.
Handley, C. O., Jr.
Ingles, L. G.
Ivey, R. D.
Lowery, G. H., Jr.
Miller, G. S., Jr.
Moore, J. C.
Peters, W.
Rafinesque, C. S.
Rageot, R. H.
Sanborn, C. C.
Sherman, H. B.
Taylor, W. P., and Davis, W. B.
Thomas, O.
Transmitted June 30, 1961.