Descriptions of new bats from Panama, CO Handley Jr

Tags: USNM, AMNH, Brazil, Darien, zygomatic breadth, Panama, interfemoral membrane, specimens, Tacarcuna Village, type locality, subspecies, braincase, Charles O. Handley, Jr., row length, NATIONAL MUSEUM, Venezuela, Florida, Panama Canal Zone, Mexico, Body size, Lasiurus borealis, Mimon crenulatum, eastern Panama, Handley, Gorgas Memorial Laboratory of Panama, Gorgas Memorial Laboratory, Sagittal crest, Harvard University Museum of Comparative Zoology, Museum of Natural History, Ecuador, Malaria Control and Survey Branch, Pedro Galindo, Robert Altman, Fort Gulick, Color Standards, Lasiurus intermedins H. Allen, SMITHSONIAN INSTITUTION, Chicago Natural History Museum, Lasiurus floridanus Miller, Lasiurus ega Juscatus Thomas, Lasiurus ega panamensis Thomas, hairs, CNHM, Lasiurus borealis teliotis H. Allen, median band, Lasiurus borealis frantzii Peters, Lasiurus egregius Peters, Lasiurus ega argentinus Thomas, Chiroderma villosum villosum Peters, Rio Amazonas, Coronoid process, floridanus, dorsum, Osceola County, Florida, Auditory bullae, Rio Pucro, upper incisor, Baja California, Texas
Content: Proceedings of
the United States National Museum
Smithsonian Institution · WASHINGTON, D.C.
Volume 112
Number 3442
By Charles O. Handley, Jr.
In its studies of tropical diseases, the Gorgas Memorial Laboratory of Panama is conducting a mammal surve3r of the Republic. The work in 1959 centered on the headwaters of the Rio Pucro, near Cerro Tacarcuna, Province of Darien, eastern Panama. Among the mammals collected on the Rio Pucro were 43 species of bats, all caught in mist nets. Many of these species had not been taken previously in Central America. Three species and one subspecies herein described are new. Either there is a surprising amount of endemism in this region, which is continuous with the Choco of Colombia, or else there are a surprising number of undescribed widespread species of bats in the Neotropical fauna. The Malaria Control and Survey Branch of the Office of the Chief Surgeon, U.S. Army Caribbean, has also been collecting bats in Pan- ama. Included in its collections are several species not previously taken in the Republic and one of the new subspecies here described. I am indebted to the personnel of the Gorgas Memorial Laboratory, particularly Carl Johnson, Pedro Galindo, and Rudolpho Hinds for their support and cooperation. Likewise, I thank Robert Altman, Marvin Keenan, and Vernon Tipton of the Malaria Control and Survey Branch for their assistance and for the opportunity to study their collections. I am also grateful to the authorities of the American 459
vol. 112
Museum of Natural History (AMNH), Academy of natural sciences of Philadelphia (ANSP), Chicago Natural History Museum (CNHM), and Harvard University Museum of Comparative Zoology (MCZ) for allowing me to study specimens under their care. Capitalized color terms in the following descriptions are from Ridg- way (Color Standards and Color Nomenclature, 1912). All measurements are in millimeters and are explained in Handley (1959, pp. 98-99).
Mimon crenulatum keenani, new subspecies
USNM Holotype:
311951, adult male, skin and skull, collected
August 25, 1959, by C. M. Keenan, Fort Gulick, Panama Canal Zone,
original number 4127.
Distribution: Central Panama to northwestern Venezuela and
western Ecuador. The Panamanian specimen was found in the day-
time hanging on the outside of a decayed hollow stump in a sparsely
wooded area. The Ecuadorean specimens likewise were taken in a
hollow rotted tree stump in a wooded area (Tate, 1931, p. 250).
Description: Dorsum bright mahogany brown (between Carob
Brown and Black); individual hairs monocolored from base to tip;
T j
tail; large prominent yellow-orange spots at posterior bases of ears.
Underparts orangish, individual hairs fuscous at base. Membranes
and ears blackish. Noseleaf hairy, slightly crenulated at base; wing
membranes attached to metatarsus; calcar long. Rostrum relatively
long, shallow, and dorsally flattened; sagittal crest relatively low;
posterior extension of palate relatively long and broad; molariform
teeth broad and massive.
Measurements: See table 1.
Comparisons: Brighter and more ornate than the contiguous races
crenulatum and longijolium. Rostrum longer and slightly deeper,
posterior palatal elongation longer and broader, teeth larger, and
calcar longer than in crenulatum. Rostrum shallower and dorsally
flatter than in longijolium. Remarks: The nominal genera Anihorhina and Mimon are not distinguishable even as subgenera. The bats of Anihorhina differ from those of Mimon principally in having a smaller anterior upper
premolar (P1 ); shorter lower incisor; larger auditory bulla; stouter z}^gomatic arch; shorter, less woolly fur; smaller ears; and hairy
noseleaf. It is doubtful that either of the nominal genera contains more than a single species. Dalquest (1957, p. 45) cleared up some of the con- fusion surrounding the supposed species M. bennettii and cozumelae, presently known only by a few specimens from Brazil and Mexico, respectively. Dalquest failed, however, to note that his only speci- men of bennettii was a juvenile, and that he distinguished it from his
NEW BATS FROM PANAMA HANDLEY aiqirea pujqaq qipcajq pujsoa
vol. 112
-- adult cozumelae on the basis of its juvenile characters smaller size; darker, less ornate coloration; and bulging braincase. Apparently bennettii and cozumelae are very similar, but until adult specimens of each are compared, their true relationship cannot be determined. Four species have been named in Anthorhina, but all seem to have been based on distortions of preservation or individual, seasonal, or geographic variations of a single species. Crenulation and hairiness of the noseleaf are individually variable throughout. The wings apparently are always inserted on the distal portion of the metatarsus, but this feature may be obscured by labels tied to the ankle. The calcar varies individually and geographically, and often is difficult to measure accurately in preserved specimens. Coloration varies with age and season, as well as geographically. Coloration of the fresh adult pelage approximates black and white, but with age these colors became progressively obscured with yellow, orange, and reddish. Juveniles are duller, blacker, and have less prominent markings than adults. Crowding of the maxillary tooth row is individually variable. Size of the ears in dried specimens depends to some extent on the conditions of drying. The genus Mimon includes the following: Mimon bennettii Gray: Type locality Ipanema, Sao Paulo, Brazil; and Mimon cozumelae Goldman: Type locality Cozumel Island, Quintana Roo, Mexico, diagnosis: Body size large (forearm aver- ages 55 mm.) ; coloration pale brownish, unmarked except for whitish postauricular patches; fur long and woolly; ears very large; noseleaf plain, naked; anterior upper premolar (P1 ) about equal to inner upper incisor (I1 ) in size; lower incisor longer than wide; auditory bullae small; zygomata fragile, specimens examined: M. bennettii: Brazil: Ipanema, Sao Paulo, 1, USNM. M. cozumelae: Mexico: Cozumel Island, Quintana Roo, 1, MCZ, 2 including type of cozumelae, USNM; Izamel, Yucatan, 2, USNM. Mimon crenulatum crenulatum E. Geoffroy St. Hilaire: Type locality, Brazil. 1 diagnosis: body size small (forearm averages 50 mm.) ; dorsal coloration blackish-brown, marked with whitish postauricular patch and median dorsal stripe, both varying from fairly prominent to obscure; fur medium-long, lax; underparts whitish to rusty, bases of hairs grayish ; cars large ; noseleaf more or less crenulated on margins toward the base, fringed with long hairs; P1 about equal to I2 in size; lower incisor as wide as long; auditory bullae large; zygomata stout; rostrum
1 Cabrera (1958, p. 66) further restricted the type locality to Bahia, hut in view of the uncertainty of the distribution of variation in the species in eastern Brazil, the restriction appears premature and perhaps detrimental. Present indications are that the subspecies crenulatum occurs at least from eastern Venezuela and Trinidad to the lower Amazon.
relatively short, shallow, and posteriorly narrowed; posterior palatal extension short and narrow; sagittal crest relatively low; molariform teeth relatively small, specimens examined: Brazil: Santarem, Rio Tapajoz, 4, MCZ. AMNH. Trinidad: Port-of-Spain, 1, Mimon crenulatum keenani Handley: Type locality, Fort Gulick, Panama Canal Zone, diagnosis: see description and comparisons above. Specimens from Rio Tocuyo, Venezuela, show intergradation with crenulatum. These are as ornate as typical keenani, but are duller and have gray ventral hair bases. In shape of rostrum and shortness of calcar, they approach crenulatum. Specimens from Bahia de Caraquez, Ecuador, seem to show intergradation with longifolium. Colorwise they are close to keenani, but are slightly duller and have gray based ventral hairs. Skull is most like that of longifolium. specimens examined: Ecuador: Bahia de Caraquez, 5, AMNH. Panama Canal Zone: Fort Gulick, 1, USNM. Venezuela: Rio Tocuyo, 500 meters, 16, AMNH. Mimon crenulatum longifolium Wagner: Type locality, Villa Maria, Mato Grosso, Brazil (peruanum Thomas, type locality Rfo Pachitea, Huanuco, Peru, is a synonym), diagnosis: This is the least ornate subspecies. Dorsal coloration dull blackish brown; ventral hairs gray based; postauricular patches and median dorsal stripe usually reduced, often obscure, and occasionally absent; rostrum relatively long, deep, and not dorsally flattened; sagittal crest high; calcar long. specimens examined: Brazil: Cacao Pereira Igarape, Rio Negro, 3, AMNH; Santo Antonio da Uayara, Rio Madeira, 4, AMNH. Colombia: Tahuapunto, Rio Vaupes, 8, AMNH. Ecuador: Boca de Rio Curaray, 1, AMNH. Peru: Montealegre ?, 2, AMNH. Venezuela: Mount Duida, 350 meters, 1, AMNH. No exact locality, 1, USNM. Mimon crenulatum picatum Thomas: Type locality, Lamarao, 300 meters, Bahia, Brazil, diagnosis: Apparently similar to crenulatum in length of calcar and rostrum, but may be brighter and more ornate in coloration. M. c. picatum may be a synonym of crenulatum, or it may represent a bright-colored southeastern population. It is known only by Thomas' type specimen.
Anoura cultrata, new species
USNM Holotype:
309396, adult female, skin and skull, collected
February 7, 1959, by Charles O. Handley, Jr., and B. R. Feinstein,
Tacarcuna Village, 3,200 ft., Rio Pucro, Darien, Panama, original
number 4747.
Distribution: Known only from the type locality, where in Febru-
ary and March 1959 eight specimens were caught in mist nets set over
a stream in a mountain forest.
vol. 112
Description: Coloration shiny blackish (between Blackish Brown-3 and Black) throughout; individual hairs of dorsum pale grayish on basal two-thirds, those of underparts black to base. Pelage short and crisp ; interfemoral membrane reduced to a narrow, densely furred band; tail present. Skull most like that of Anoura geoffroyi, but differing as follows: larger; braincase more tapering anteriorly; rostrum thickened; zj^gomata complete; posterior margin of palate more deeply incised beside posterior palatal extension; pterygoids inflated pos- teriorly so as to narrow the elongated mesopterygoid fossa. Symphysial region of mandibles shortened and trough between canines deepened and broadened (anterior end of mandible correspondingly depressed) ; coronoid process reduced in height ; ventral edge of man- dible with a low process just anterior to the angular process. Outer upper incisor (I2) enlarged, bladelike; upper canine enlarged, roughly triangular in cross-section at base, with distinct anterointernal, anteroexternal, and posterior basal cusps; internal face anteroposteriorly concave, and anterior face flat, with prominent longitudinal sulcus from base of crown to near tip , P3 and P4 reduced in height and thick- M M ness and in prominence of cusps; 2 and 3 reduced in size; lower canine reduced in height, with distinct cingulum on anterior and internal faces, and with a posterointernal cingular cusp ; anterior lower premolar (Px) bladelikc and enormously enlarged (lengthened and thickened), with its highest point in the posterior half of the tooth; P3 and P4 reduced in height and thickness and in prominence of cusps. Measurements (holotype): Total length 94, tail vertebrae 6, hind foot 14, ear from notch 16, forearm 43.2, tibia 15.9, calcar 3.8. Greatest length of skull 26.3, zygomatic breadth 10.7, interorbital breadth 4.7, braincase breadth 10.3, braincase depth 8.0, maxillary tooth row M length 9.0, postpalatal length 9.4, palate breadth outside of 3 5.7, rostral breadth at base of canines 4.7. Comparisons: Probably the closest relative of cultrata is Anoura geqffroyi, but as indicated in the description above it is strongly dif- ferentiated from this species in many characters. Specimens examined: Panama: Tacarcuna Village, 3,200 ft., Rio Pucro, Darien, 8, USNM.
Chiroderma gorgasi, new species
USNM 6x3 Holotype:
309903, adult male (testis
mm.), skin
and skull, collected March 6, 1959, by Charles O. Handley, Jr., and
B. R. Fcinstein, Tacarcuna Village, 3,200 ft., Rio Pucro, Darien,
Panama, original number 5436.
Distribution: Known only from the type locality in eastern Pan-
ama, where five individuals were caught in February and March 1959
in mist nets set over a stream in a mountain forest.
Description: Body size small (forearm 37.5-38.5 mm., greatest
length of skull 20.2-20.9 mm.). Dorsal coloration yellowish brown,
brown (between Sudan Brown and Prout's Brown in holotype), or
grayish, paler anteriorly; a white median stripe extends from upper
back to base of tail; prominent white stripes above and below eye,
extending from noseleaf to crown at inner base of ear and from pos-
terior part of upper lip to outer base of ear; area about eye a little
darker than remainder of body; individual hairs of dorsum tricolored,
dusky at base, bufl'y medially, and brown or gray at tip; underparts
uniformly grayish or brownish gray, very slightly washed with whitish.
Membranes blackish; tragus and basal portion and margins of ear
yellow; tip of ear yellowish gray. Eye large; noseleaf broad, with
simple tip; interfemoral membrane hairy at base but naked on pos-
terior margin. Nasal aperture short, extending only to level of
anterior edge of orbits; supraorbital region scarcely ridged, but lachry-
mal region sharply ridged; sagittal and lambdoidal crests poorly de- veloped; inner upper incisors slender; canines and P4 low, but anterior
lower premolar (Pi) large and anterior cusp half or two-thirds the height of P4 . Measurements: Male holotype of gorgasi, followed by female para-
type (USNM 309902), and in parentheses the female holotype of
trinitatum: total length 56, 57 (-) ; hind foot 10, 11 (12); ear from
notch 17, 18 (15 in alcohol) ; forearm 38.5, 37.6 (41.0); tibia 12.4, 12.4
(15.2) ;calcar 4.9, 4.5 (4.5). Greatest length of skull 20.9, 20.7 (22.2);
zygomatic breadth 12.8, 13.1 (13.9) ; interorbital breadth 5.4, 5.6 (5.6)
braincase breadth 9.4, 9.6 (9.8); braincase depth 7.8, 7.9 (7.8); maxil-
lary tooth row length 7.3, 7.3 (7.7); postpalatal length 5.4, 6.0 (5.8); M palatal breadth outside of 3 9.5, 9.4 (9.7); rostral breadth behind
canines 4.8, 4.9 (5.1).
Comparisons: The closest relative of gorgasi appears to be trinita-
tum Goodwin, which is known only by the type specimen from Trinidad. Together these species stand well apart from all other known
Chiroderma, and additional collecting may show them to be conspecific.
C. gorgasi may be distinguished from trinitatum by smaller size;
relatively broader skull; relatively deeper braincase and more bulging
forehead; shorter rostrum; sharper lachrymal ridge; more rounded
supraorbital region; heavier zygomata; larger outer upper incisors
M (I 2 ); shorter (anterior-posterior)
1 .
Coloration of trinitatum is
P 4,
nasal aperture, and lack of a supraorbital ridge are characters of
gorgasi that distinguish it from the sympatric lyillosum Peters and
salvini Dobson.
vol. 112
Remarks: Chiroderma jesupi J. A. Allen (1900), known only by the type specimen from northern Colombia, was described as a small species. The type is a juvenile with phalangeal epiphyses not ossified. It is the same as the bat subsequently described as C. isthmicum by Miller (1912), and it is conspecific with Chiroderma villosum Peters, type locality Brazil (not Venezuela as stated by Cabrera, 1958, p. 85). The valid species of the genus Chiroderma are thus as follows
Chiroderma doriae Thomas: type locality, Minas Gerais, Brazil {dorsale Lund and villosum Dobson are synonyms). Chiroderma gorgasi Handley: type locality, Tacarcuna Village, Darien, Panama. Chiroderma salvini Dobson: type locality, Costa Rica. Chiroderma trinitatum Goodwin: type locality, Cumaca, Trinidad. Chiroderma villosum villosum Peters: type locality, Brazil. Chiroderma villosum jesupi J. A. Allen: type locality, Cacagualito, Santa Marta, Colombia (isthmicum Miller is a synonym).
Specimens examined: C. gorgasi: Panama: Tacarcuna Village,
Darien, 5, USNM. C. salvini: Costa Rica: Angostura, Cartago, 1, USNM; Cafias Gordas (Agua Buena), Puntarenas, 1, AMNH. Hon-
duras: La Flor Archaga, 32, AMNH; San Marcos, 1, AMNH; De-
partment of Yoro, 2,800 ft., 1, MCZ. Panama: Cana, Darien, 1,
USNM; Cerro Azul, 2, USNM; Tacarcuna Village, Darien, 99,
USNM. C. trinitatum: Trinidad: Cumaca, 1, type of trinitatum,
AMNH. C. villosum villosum: Brazil: Calama, 1,
Diego Martin, 1, AMNH; Maracas Valley, 1, AMNH; Port-of-Spain,
2, MCZ. Venezuela: San Esteban, 1, AMNH. G. v. jesupi: Colom-
AMNH. bia: Cacagualito, 1, type of jesupi, Panama: Barro Colorado
Island, 6, USNM; Cabima, 2, including type of isthmicum, USNM:
Cerro Azul, 1, USNM; Culebra, 1, USNM; Paya Village, Darien, 1,
USNM; Rio Cangandi, San Bias, 1, USNM; Tacarcuna Village,
Darien, 7, USNM. Mexico: Presidio, Veracruz, 1, USNM.
Myotis simus riparius, new subspecies
USNM Holotype:
310255, adult female (with one embryo, 7 mm.
crown-rump), skin and skull, collected February 9, 1959, by Charles O.
Handley, Jr., and B. R. Feinstein, Tacarcuna Village, 3,200 ft., Rio
Pucro, Darien, Panama, original number 4843.
Distribution: Eastern Panama.
Description: Fur short and woolly; dorsum buffy brown (between Warm Sepia and Bister) ; individual hairs slightly burnished at tip,
slightly grayer toward base; underparts yellowish brown, individual
hairs fuscous at base; lips, ears, membranes, and feet blackish. Calcar
keeled; wing membrane attached to foot at base of toes. Body size
large; zygomata heavy and wide spreading; braincase relatively nar-
row; sagittal and lambdoidal crests high and forming a triangular
"helmet" at their juncture in the interparietal region; rostrum long
and shallow; inner cutting edge of outer upper incisor (I2) usually not
crenulated; middle upper premolar (P3) about two-thirds the size of
M M P1 ;
1 and
2 with protoconule, hypocone, metaloph, and cingulum
fairly well developed.
Measurements: Holotype and a female paratype (USNM 310256)
from the t}rpe locality, together with measurements, in parentheses, of two female topotypes of simus (AMNH 76244 and 76246) : Total
length 89, 86 (-, -) ; tail vertebrae 40, 36 (-, -) ; hind foot 8, 8 (9, 10) ear from notch 14, 13 (-, -); forearm 39.1, 35.7 (-, -); tibia 14.3, 13.5
(-, -); calcar 12.8, 13.2 (-, -). Greatest length of skull 13.9, 13.8
(14.0, 13.7); zygomatic breadth 8.8, 8.9 (9.4, -); intcrorbital breadth
3.5, 3.5 (3.9, 3.7) ; braincase breadth 6.4, 6.3 (7.0, 6.7) ; braincase depth
5.0, 5.0 (5.1, 4.9); maxillary tooth row length 5.3, 5.4 (5.1, 5.1); post- M palatal length 4.5, 4.7 (4.7, 4.6) palatal breadth outside of 3 5.5, 5.7 ; (5.7, 5.2); rostral breadth behind canines 3.7, 3.6 (4.0, 3.8).
Comparisons: Compared with Amazonian simus, the Panamanian
specimens have the rostrum longer, shallower, and narrower at the
tip; the braincase narrower, less inflated; the tooth row is longer and
less crowded; the middle upper premolar (P3 ) is larger (two-thirds
P1 );
the protoconule, hypocone, metaloph, and cingulum are better devel-
M M oped in
1 and
2 ;
2 (I )
lated ;
the lambdoidal crests in the interparietal region to form a triangular
helmet not seen in the Amazonian specimens; the fur of the dorsum
is slightly longer and more burnished (thus brighter).
Remarks: Coloration and length of fur are seasonally variable in
simus. In a series of sixteen specimens from Boca de Rio Curaray,
Ecuador, those collected in February and March have short (2-3 mm.
on rump), orange-brown fur, with the individual hairs monocolored
on all parts of the body. The remainder of the series, collected in
October and December, are quite different in appearance. The fur is longer (3-4 mm. on rump), chocolate brown, with slightly burnished
tips, and the individual hairs of the underparts are sharply bicolored.
AMNH Several Brazilian and Peruvian specimens (e.g.
74378, 74380,
74105, 91889, 92702) show molt from short, orange, monocolored fur
to the longer, brown, bicolored pelage.
Several characters ascribed to simus by Thomas (1901, p. 541) and
subsequent authors cannot be substantiated in the specimens of
simus that I have examined. Most important is the supposed inser-
tion of the wing at the ankle. Possibly this feature had been distorted USNM in Thomas' specimen by labels tied to the ankles, as in the two
544232--60 2
vol. 112
specimens relaxed by Miller (1928, p. 206). Actually the wing is attached to the foot at the base of the toes in simus as in most other Myotis. The calcaral keel, described as practically absent, indeed
is absent in a few specimens, but as a rule it is well developed. The
width of the rostrum and crowding of the maxillary tooth row are
geographically variable. Coloration and length of fur are seasonably
Thus, Myotis simus is not so strikingly differentiated from other species of Myotis as Thomas and Miller supposed, and there is little basis for the subgeneric name Hesperomyotis proposed for it by
Cabrera (1958, p. 103). The Panamanian specimens were caught in mist nets set over streams in a lowland semideciduous forest and a mountain forest, and
at the edge of a clearing in a mountain forest.
Specimens examined: M. s. riparius: Panama: Boca de Paya,
Darien, 1, USNM; Cerro Azul, 2,100 ft., 1, USNM; Tacarcuna Village,
3,200 ft., Rio Pucro, Darien, 4, USNM. M. s. simus: Brazil: Auara
AMNH; Igarape, Rio Madeira, 7,
Cacao Pereira Igarape, Rio Negro,
8, AMNH; Igarape Amorin, Rio Tapajoz, 3, AMNH; Rosarinho, Rio
AMNH; Madeira, 4,
Villa Bella Imperatriz, South bank of Rio
Amazonas, 14, AMNH. Ecuador: Boca de Rio Curaray, 16, AMNH,
2, USNM. Peru: Apayacu, Rio Amazonas, 4, AMNH; Orosa, Rio
Amazonas, 3, AMNH; Panya, Boca de Rio Topaya, Ucayali, 1,
AMNH; Rio Pisqui, Ucayali, 1, AMNH; Sarayacu, Rio Ucayali, 12,
topo types of simus, AMNH.
Lasiurus castaneus, new species
USNM Holotype:
310263, adult female (with 2 embryos, 13 mm.
crown rump), skin and skull, collected March 6, 1959, by Charles O.
Handley, Jr., and B. R. Feinstein, Tacarcuna Village, 3,200 ft., Rio
Pucro, Darien, Panama, original number 5445.
Distribution: Known only from the type locality, where a single
specimen was taken in a mist net over a stream in a mountain forest.
Description: Dorsum deep chestnut (between Morocco Red and
Chestnut), shading on rump, interfemoral membrane, and feet to
mahoganjr (between Maroon and Claret Brown); median band of
individual dorsal hairs Cinnamon-Rufous; individual hairs tricolored,
with individual bands (black-amber-chestnut) about equal in extent;
face and muzzle, entirely black; underparts blackish brown with only
scattered buff-tipped hairs except on collar; throat not differentiated
from remainder of underparts in color; hairs surrounding white
humeral spot black tipped; ears, wings, membranes, paid lips entirely
blackish. Distal fourth of interfemoral membrane naked; auricle
M and antitragus relatively large.
3 much reduced, second com-
M M missure shorter than first; hypocone much reduced on 1 and 2 ; P4 double rooted. Rostrum broad and deep; lachrymal ridge not developed; braincase narrow, deep, and tilted up away from plane of palate. Otherwise similar to Lasiurus borealis. Measurements of holotype: Total length 112, tail vertebrae 48, hind foot 11, ear from notch 14, forearm 44.8, tibia 19.5, calcar 15.1. Greatest length of skull 13.0, zygomatic breadth 9.9, interorbital breadth 4.2, braincase breadth 7.6, braincase depth 6.2, maxillary tooth row length 4.7, postpalatal length 5.7, palatal breadth M at 3 6.7, rostral breadth at canine 5.7. Comparisons: Lasiurus castaneus is a member of the borealis group. Its coloration is unique, and extreme tilting of the braincase rlativeto the palatal plane and reduction of parts of the upper molaers like wise set it apart from other species of the borealis group. In overall size and in size of auricle and antitragus it resembles the remote northern borealis and southern varius, but its wings are unusually long. Like seminolus it lacks development of a lachrymal ridge, has a black face, and has the median band of the dorsal hairs reduced. Like varius it has black ears and membranes and darkened face. It scarcely needs comparison with the sympatric Jrantzii, which is much smaller, paler and brighter colored, and plain faced and has a globose braincase and a shorter, narrower rostrum. Remarks : Variation in coloration is remarkably slight in the borealis group. Specimens from California (teliotis), Central America (Jrantzii), and Chile (varius) are scarcely distinguishable colorwise. Differentiation in coloration has occurred in the extreme northeast (borealis) and extreme southeast (blossevillii) along similar lines overlay of the red mass effect with white (sort of a "frosting"). < 'astaneus in Central America, seminolus in the northeast, and perhaps egregius in Brazil, appear to be independent variables. Sexual dimorphism in size and coloration is marked in borealis and seminolus but is slight in western North America, Central America, and South America (see also table 2 on next page). Extension of fur onto the membranes varies with latitude and perhaps with altitude (more at higher latitudes, less at lower latitudes) and is of limited taxonomic value. Bats of the mainland of North and South America that belong to the Lasiurus borealis group are: Lasiurus borealis blossevillii Lesson and Garnot : Type locality, Montevideo, Uruguay (bonariensis Lesson, Buenos Aires, Argentina; enslenii Lima, Sao Lourengo, Rio Grande do Sul, Brazil; and salinae Thomas, Cruz del Eje, Cordoba, Argentina, are probably synonyms). diagnosis: slightly larger than Jrantzii, smaller than varius (larger toward the south) ; dorsum washed with whitish ("frosted") ; reddish
VOL. 112
vol. 112
tips of dorsal hairs shading terminally to blackish (dominantly blackish
in south, more reddish to north) ; buffy median band of dorsal hairs
the dominant band; ears and membranes with reddish markings;
females slightly larger and paler than males; lachrymal ridge devel-
oped, specimens examined: Argentina: Yuto, Jujuy, 2, AMNH.
AMNH; Brazil: Maracaju, Mato Grosso, 1,
Sao Sebastiao, Sao Paulo,
1, USNM. Paraguay: Villa Rica, 3, USNM. Uruguay: Monte-
video, 1, AMNH.
Lasiurus borealis borealis Miiller : Type locality, New York, diag-
nosis: larger than teliotis; dorsum reddish, washed with whitish;
buffy median band of dorsal hairs the dominant band; ears and mem-
branes with reddish markings ; females distinctly larger and paler than males; lachrymal ridge developed, specimens examined: many
from southern Canada, eastern United States, and northeastern
Lasiurus borealis frantzii Peters: Type locality, Costa Rica, diag-
nosis: smaller than contiguous races; dorsal color clear reddish with-
out white wash; buffy median band of dorsal hairs the dominant band;
ears and membranes with reddish markings; females scarcely dif-
ferentiated from males in size and coloration; juveniles similar to adult
blossevillii in coloration; lachrymal ridge developed, specimens
AMNH; examined: Colombia: Pacho, near Bogota, 1,
Meta, 1, AMNH. Costa Rica: San Sebastian, San Jose, 1, AMNH.
Honduras: La Flor Archaga, 1, AMNH. Panama: Boquete, 3,500
USNM; ft., Chiriqui, 1,
Tacarcuna Village, 3,200 ft., Darien, 2,
USNM; USNM. no exact locality, 1,
Peru: Juliaca, 6,000 ft,, 1,
AMNH. Venezuela: Paya, 10 miles north of El Sombrero, Guarico,
1, AMNH; Macuto, 1, USNM; Merida, 1, AMNH.
Lasiurus borealis teliotis H. Allen; Type locality, unknown, probably
California (ornatus Hall, Penuela, near Cordoba, Veracruz, is a syno-
nym), diagnosis: larger than frantzii but similar to it in coloration
(slightly paler northward) and cranial characters, specimens exam- ined: many from the western United States, Baja California, Jalisco,
Michoacan, Oaxaca, and Veracruz.
Lasiurus borealis varius Poeppig: Type locality, Antuco, Bio-Bio,
Chile (poepingii Lesson is a synonym), diagnosis: much larger than
frantzii; similar to frantzii in color, except for darker face; ears and
membranes entirely black; lachrymal ridge developed, specimens examined: Chile: Angol, 3 AMNH; Maquehue, Temuco, 1, AMNH, 2, USNM; Rio Blanco, 4,900 ft., 4, USNM.
Lasiurus castaneus Handley: Type locality, Tacarcuna Village, 3,200 ft., Darien, Panama (see description above).
Lasiurus egregius Peters; type locality, Santa Catarina, Brazil.
diagnosis: Similar to castaneus but larger (forearm 48 mm. in male
type, the only known specimen) ; dorsum bright rufous, darkening almost to chestnut on interfemoral membrane; face noticeably more red tinged than crown and nape; underparts bright red; membranes black (this diagnosis is from notes that G. S. Miller, Jr., made at the Berlin Museum in the early 1900's). Lasiurus seminolus Ilhoads: Type locality, Tarpon Springs, Pinellas County, Florida, diagnosis: similar in size to borealis; dorsum mahogany, washed with whitish; black basal band of dorsal hairs the dominant band; face black; ears and membranes with reddish mark- ings; females larger than males, but similar in coloration; juveniles darker; lachrymal ridge not developed, specimens examined: many from southeastern United States. It should be noted that previous authors have followed Peters (1871) in aligning egregius with the genus Dasypterus, because it lacked the minute upper premolar (P1 ), although in other characters it agreed with the borealis group of the genus Lasiurus. Absence of P1 in a single specimen of Lasiurus {egregius apparently is still known only by the type specimen) is not significant, for although P1 seems always to be absent in Dasypterus, it is also absent in one or both maxillae of about 10 percent of Lasiurus. Table 3, on page 475, lists some of the conspicuous differences between the species of Lasiurus and Dasypterus. More impressive are the following similarities linking these nominal genera and distinguishing them from other vespertilionids and in some cases from all other bats Four mammae and average of two or three young per litter. Spiral effect in scale arrangement on hairs. Reduction of sebaceous glandular tissue and location of the submaxillary salivary gland in the facial area. Bright coloration. Baculum short, J-shaped, with high base and narrow shaft. Distally enlarged and spiny penis. Furry interfemoral membrane. It seems more reasonable to stress the important similarities of these bats and regard them as congeneric, rather than to stress the insignificant differences and regard them as representing distinct genera. I do not believe that Dasypterus is useful even as a subgenus. Recent bats of the ega group may be arranged in the genus Lasiurus as follows: Lasiurus ega argentinus Thomas: Type locality, Goya, Corrientes, Argentina (caudatus Tomes, Pernambuco, Brazil, is a synonym). diagnosis: dorsum pale whitish buff, washed lightly with black; orange hairs of interfemoral membrane contrasting with remainder of dorsum; face blackish; underparts dull whitish buff. Paler than ega. Body size small, specimens examined: Argentina: Tucuman, 1, CNHM, 1, USNM. Bolivia: Caiza, 1, USNM. Brazil: Ipiranga,
vol. 112
Sao Paulo, 1, CNHM; Lago Hyanuary, Pernambuco, 2, MCZ; Salto
Grande, Parand, 1, USNM; Sao Marcello, Bahia, 1, CNHM; Villa
Bella Imperatriz, south bank of Rio Amazonas, 2, AMNH. Para-
CNHM; guay: near Asuncion, 1,
Colonia Nueva Italia, Chaco, 1,
CNHM; Villa Montes, Chaco, 1, CNHM; Villa Rica, 1, USNM.
Uruguay: Salto, 2, CNHM.
Lasiurus ega ega Gervais: Type locality, Ega, Amazonas, Brazil.
diagnosis: dorsum yellowish orange (darker than argentinus), with
inconspicuous blackish wash; hairs of interfemoral membrane not
contrasting with remainder of dorsum; face black; underparts paler.
The Amazonian specimens become progressively darker upstream.
Body size small, specimens examined: Bolivia: Buena Vista, Santa
Cruz, 1, CNHM.
AMNH; Brazil: Baiao, Rio Tocantins, 2,
Rio Madeira, 1, AMNH; Manaos, Rio Negro, 1, AMNH; Rosarinho,
Rio Madeira, 1, AMNH; Santarem, Rio Tapajoz, 1, MCZ; Santo
Antonio da Uayara, Rio Madeira, 1, AMNH. Peru: Pebas, Loreto,
1, ANSP; Sarayacu, Rio Ucayali, 1, AMNH.
Lasiurus ega Juscatus Thomas: Type locality, Rio Cauquete, Rio
Cauca, Colombia (punensis J. A. Allen, Puna Island, Ecuador, is a
synonym), diagnosis: dorsum orange with heavy black wash (com-
pared with panamensis, coloration is much darker and richer, broad
subterminal band of dorsal hair orange rather than buff, and black
tip longer and more dominant in mass effect) ; hairs of interfemoral
membrane not contrasting with remainder of dorsum; entire head
blackish; underparts dusky, becoming buffy posteriorly. Body size small, specimens examined: Colombia: Cali, 1, AMNH. Ecuador: Guayaquil, 1, MCZ; Puna Island, 5, including type of punensis, AMNH.
Lasiurus ega panamensis Thomas: Type locality, Bugaba, Chiriqui,
Panama, diagnosis: dorsum dull sooty yellowish (paler than fuscatus
and duller and more black washed than ega) hairs of interfemoral ; membrane not contrasting with remainder of dorsum; face black;
underparts dull drabby bun", paler posteriorly. Body size small.
AMNH; specimens examined: Costa Rica: San Jose, 1,
Villa Quesada,
Alajuela, 1, AMNH. Honduras: Tegucigalpa, 1, AMNH. Venezuela:
CNHM. Lagunillas, Zulia, 3, Mexico: Achotal, Veracruz, CNHM;
Yaxcach, Yucatan, 1, USNM.
Lasiurus ega xanihinus Thomas: Type locality, Sierra Laguna, Baja
California, Mexico, diagnosis: dorsum pale yellowish, lightly washed
with black (paler, more yellowish, and less mantled with black than
T 3
with remainder of dorsum; face not blackened; underparts orangish,
not becoming significantly paler posteriorly. Body size small, speci-
mens examined: Mexico: Comondu, Baja California, 1, USNM;
AMNH; Miraflores, Baja California, 2,
Santa Anita, Baja California,
3, USNM; Sierra Laguna, Baja California, 1, USNM; Escuinapa, Sinaloa, 1, AMNH. Lasiurus floridanus Miller: Type locality, Lake Kissimmee, Osceola County, Florida, diagnosis: dorsum buffy yellow, lightly washed with black (similar to xantkinus; paler, less orange and with more black than intermedins) ; hairs of interfemoral membrane not contrast- ing with remainder of dorsum; face blackish; undcrparts similar to dorsum in coloration. Body size large, specimens examined: a
Table 3. Differentiating characters of (he species groups of the genus Lasiurus
Red bats (L. borealis, etc.)
Hoary bats (L. cinereus, etc.)
Yellow bats (L. ega, etc.)
Size small (forearm Size large
37-44 mm.)
46-57 mm.)
(forearm Size large (forearm 44-57 mm.)
Lateral wings of presternum equal to body of presternum in width
Lateral wings of presternum equal to body of presternum in width
Lateral wings of presternum considerably broader than body of presternum
Presternum about as Presternum much longer Presternum about as
long as wide
than wide
long as wide
Auditory bullae not Auditory bullae slightly Auditory bullae not
Rostrum short
relatively Rostrum medium
Rostrum long
Sagittal crest very Sagittal crest weak weak
Sagittal crest strong
Coronoid process medi- Coronoid process low um height
P 1 usually present
P 1 usually present
Coronoid process high P 1 always absent
Hypocone slightly re- Hypocone much re- Hypocone slightly re-
M M M M duced on 1 and 2
duced on 1 and 1
M M duced on 1 and 8
M3 very reduced
M3 reduced
M3 slightly reduced
P* double rooted Ma talonid reduced
P* single rooted
P« double rooted
Mi talonid only slightly M| talonid much re-
s 8 a ©
3 cc
vol. 112
total of 47, including the type, from Florida, Georgia, Mississippi, and Louisiana. Lasiurus intermedins H. Allen: Type locality, Matamoros, Tamaulipas, Mexico, diagnosis: dorsum clear yellowish orange with very fine blackish wash (brighter, clearer orange than panamensis, with which it is sympatric) ; hairs of interfemoral membrane not contrasting with remainder of dorsum; face slightly blackened; underparts colored like dorsum. Body size large, specimens examined: Cuba: San German, Oriente Province, 1, USNM. Honduras: Rio Yeguare, between Tegucigalpa and Danli, 1, MCZ. Mexico: San Bartolome, Chiapas, 1, USNM; Tehuantepec, Oaxaca, 1, AMNH; Izamal, Yucatan, 5, USNM; Tekom, Yucatan, 1, CNHM; Matamoros, Tamaulipas, 5, USNM. United States: Brownsville, Texas, 4, AMNH, 1, ANSP, 3, CNHM, 49, USNM; Cameron County, Texas, 6, USNM; Padre Island, Texas, 1, USNM. Although geographic variation in coloration is considerable, geographic variation in body size is not apparent in the small samples of ega that I have examined (see table 4, pp. 476-477). L. intermedins and floridanus must be closely related. Together they stand well apart from ega in larger body size, more massive skull, stronger rostrum, higher crests, and more widely spreading zygomata. Southern Texas and Latin American populations of intermedins average larger in body size than do specimens of floridanus from Louisiana, Georgia, and Florida, but they overlap. Neither this variation nor the variation in coloration appears to be clinal in intermedins and floridanus. However, similarity of the antorbital structure in specimens from Mexico, Texas, and Louisiana, in contrast to this structure in specimens from Georgia and Florida, suggests gene flow between intermedins and floridanus. I have not seen specimens from Texas north of the Rio Grande Valley, where intergradation would be expected to occur if floridanus and intermedins are conspecific.
Literature Cited Cabrera, Angel 1958. Catalogo de los mamiferos de America del Sur. I. Rev. Mus. Argentine* Cienc. Nat. Bernardino Itivadavia. Cienc. Zool., vol. 4, pp. 1-308. Dalquest, Walter W. 1957. American bats of the genus Mimon. Proc. Biol. Soc. Washington, vol. 70, pp. 45-47. Handley, Charles O., Jr. 1959. A revision of American bats of the genera Euderma and Plecotus. Proc. U.S. Nat. Mus., vol. 110, pp. 95-246, 27 figs. Miller, Gerrit S., Jr., and Allen, Glover M. 1928. The American bats of the genera Myotis and Pizonyx. U.S. Nat. Mus. Bull. 144, i-viii, 1-218 pp., 13 maps. Peters, W. 1871. Eine Monographische ubersicht der Chiropterengattungen Nycteris und Alalapha. Monatsb. Preuss. Akad. Wiss. Berlin, 1870, pp. 900-914, 1 pi. Ridgway, Robert + 1912. Color standards and color nomenclature, iv 44 pp., 53 pis. Tate, G. H. II. 1931. Random observations on habits of South American mammals. Journ. Mamm., vol. 12, pp. 248-256. Thomas, Oldfield 1901. New Myotis, Artibeus, Sylvilagus, and Metachirus from Central and South America. Ann. Mag. Nat. Hist., ser. 7, vol. 7, pp. 541-545.

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