SYSTEMATICS OF MILLER S LONG-TONGUED BAT, GLOSSOPHAGA LONGIROSTRIS, WITH DESCRIPTION OF TWO NEW SUBSPECIES

OCCASIONAL PAPERS THE MUSEUM TEXAS TECH UNIVERSITY NUMBER 100 18 MARCH 1986 SYSTEMATICS OF MILLER S LONG-TONGUED BAT, GLOSSOPHAGA LONGIROSTRIS, WITH DESCRIPTION OF TWO NEW SUBSPECIES Wm. David Webster and Charles O. Handley, Jr. Glossophaga longirostris is a common nectar-feeding bat that occurs in northern South America and on some Caribbean islands as well. Despite its relatively wide distribution, few scien¬ tific collections contain a sufficient number of specimens from throughout its geographic range to permit an adequate evaluation of the variability in this species. Miller (19136) examined 204 specimens of G. longirostris in his revision of the genus; he divided what he recognized as the “longirostris-group” into G. /. longirostris from coastal Colombia and Venezuela, G. I, rostrata from Grenada and the Grenadines, and G. elongata from Curasao. More recently, Goodwin (1958) named and described G. major from Trinidad, but Koopman (1958) subsequently recog¬ nized it as a subspecies of G. longirostris and also arranged indi¬ viduals from the Netherlands Antilles as G. longirostris elongata. Specimens from Boqueron de San Francisco (ca. Bogota), Colom¬ bia, originally described by J. A. Allen (1916) as Glossophaga apolinan and listed by Cabrera (1958) as Glossophaga longirostris apolinari, actually represent Anoura geoffroyi according to San¬ born (1933) and our own observations, and they are not discussed beyond. Four subspecies of Glossophaga longirostris currently are recognized (Jones and Carter, 1976): G. 1. longirostris from north¬ ern Colombia and Venezuela; G. /. elongata from Aruba, Curasao, and Bonaire; G. /. major from Trinidad and Tobago; and G. 1. 2OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY rostrata in the southern Lesser Antilles, from Grenada northward to St. Vincent. A significant number of specimens of G. longirostns have been collected in the years since Miller’s revision, the most noteworthy additions examined by us being those of the Smithsonian Vene¬ zuelan Project (SVP) (Handley, 1976) and the small but critical collection of O. P. Pearson from Huila, Colombia (deposited in the Museum of Vertebrate Zoology). These specimens allow us to assess the extent of geographic and nongeographic variation in Glossophaga longirostris, and to define better certain aspects of the biology that are poorly known in the species. This revision is based on the examination of 1188 specimens of G. longirostris, most of which are conventional skins accompanied by skulls; however, some consisted of skulls and postcranial skeletons, skulls without accompanying skins, or specimens preserved in fluid (from some of which the skulls had been extracted). All data on labels associated with specimens were recorded. Molt patterns were mapped for each individual found to have been in the pro¬ cess of molt. Measurements used in the morphometric analyses were taken from adults (completely fused phalangeal epiphyses and well-ossified cranial sutures) with dial calipers in millimeters; for definitions of cranial measurements see Handley (1959). Pelage coloration was assessed from individuals of Glossophaga longiros¬ tris deposited in the National Museum of Natural History under and Exam-O-Lite; color terminology from Ridgway (1912). Glossophaga longirostris Miller’s Long-tongued Bat Glossophaga longirostris is known from northern South Amer¬ ica (Colombia eastward to Guyana) and several of the Caribbean islands immediately adjacent to the mainland, including the Netherlands Antilles (Aruba, Curasao, and Bonaire), the continen¬ tal islands of Margarita, Trinidad, and Tobago, and the southern Lesser Antilles (Grenada, the Grenadines, and St. Vincent). Hummelinck (1940) reported G. soricina from the Testugos and several of the islands mentioned above. Some or all of these bats may represent G. longirostris (see Koopman, 1958), but Handley examined specimens of G. soricina from Isla Margarita in the SVP collections (Handley, 1976), Grenada (BMNH 67.5.4.5), and Bequia, Grenadines (BMNH 91.5.15.9). WEBSTER AND HANDLEY—GLOSSOPHAGA LONGIROSTRIS 3 Diagnosis Largest member of the genus in most external and cranial mea¬ surements; upper incisors noticeably and equally procumbent, 12 equal to II in bulk (occlusal view); P4 with reduced lingual cingular shelf; Ml narrow; parastyle of Ml usually absent or, if pres¬ ent, minute and directed posterolabially from paracone; mesostyle of Ml reduced, continuous with labial outline of tooth; fourth commissure of Ml long, well developed, and always longer than third; M2 similar to Ml except parastyle better developed, directed labially; lower incisors large and usually in contact, subtriangular in occlusal view, equal in bulk; p4 narrow, similar to p2 and p3 in bulk; premaxillae elongate anteriorly; pterygoid "wings” absent; presphenoid ridge usually high and complete throughout; mandibular symphyseal ridge absent, chin of mandible receding at a 45° angle; pelage bicolored, the tips of the individual hairs darker than the paler bases, Wood Brown to Fuscous dorsally, Avellaneous to Clove Brown ventrally; weight averaging 13.25 (10.3-16.0) grams in males, 12.81 (9.8-14.3) in nonpregnant females from throughout the range of the species. Comparisons The morphology of the anterior region of the cranium and mandible in five species of Glossophaga is shown in Fig. 1. Glossophaga longirostris can be distinguished from G. commissarisi by its larger size in most external and cranial measurements. In addition, in G. longirostris the upper incisors are noticeably pro¬ cumbent; the fourth upper premolar has a reduced lingual cingular shelf; the parastyle of Ml is absent or greatly reduced; the lower incisors are large, subtriangular in occlusal view, and usu¬ ally in contact; the presphenoid ridge is high and complete throughout; the slope of the rostrum is not abrupt; and the man¬ dibular symphyseal ridge is absent. In G. commissarisi, the upper incisors are not noticeably procumbent; P4 has a distinct posterolingual cingular shelf; parastyle of Ml is well developed; lower incisors are reduced in size and subcircular in occlusal view, with distinct gaps between the teeth; presphenoid ridge is flattened subterminally; slope of the rostrum is more abrupt; and mandibular symphyseal ridge is well developed. Compared with G. longirostris, G. leachii is smaller in most external and cranial measurements. In addition, upper incisors are noticeably procumbent in G. longirostris (not noticeably procum¬ bent in G. leachii), P4 has a reduced lingual cingular shelf (well 4OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY Fig. I.—Incisor morphology in five species of Glossophaga. Oblique and dorsal view of the anterior rostral region (from top to bottom), and a dorsal view of the anterior mandibular region, in a typical specimen of, from left to right, G. commissarisi, G. leachii, G. longirostris, G. mexicana, and G. soncina. WEBSTER AND HANDLEY—GLOSSOPHAGA LONGIROSTRIS 5 by the larger dimensions of most external and cranial measure¬ ments. Pterygoid “wings” and mandibular symphyseal ridge are absent in G. longirostris (well developed in G. soricina), upper incisors are similar in bulk in G. longirostris (12 smaller than II in G. soricina), lingual cingular shelf of P4 is reduced in G. lon¬ girostris (prominent in G. soricina), parastyle of Ml is absent in G. longirostris (well developed and directed anterolabially in G. soricina), mesostyles of Ml and M2 are poorly developed in G. longirostris (well developed in G. soricina), and p4 is narrow and similar to p2 and p3 in bulk in G. longirostris (p4 wider than p2 and p3 in G. soricina). Tamsitt and Valdivieso (1963) reported the noseleaf of G. longirostris to be smaller than that of sympatric G. soricina in central Colombia. Specimens of both species that we examined from the savannas of the Rupununi in Guyana and the llanos of Venezuela show the opposite relationship (specimens from each locality were collected on the same day and preserved in fluids in identical manner)—the noseleaf of G. longirostris averages 6.06 (5.7-6.4), whereas that of G. soricina averages 5.28 (4.6-5.7). Taxonomic Conclusions Patterns of geographic variation (Table 1) and a fragmented geographic distribution indicate that there are six subspecies of Glossophaga longirostris. Bats from the llanos of Venezuela and from low elevation grasslands around the Kanuku Mountains in Guyana represent an undescribed race that is relatively small in external and cranial dimensions, and has moderately large postor¬ bital swellings. Specimens from Aruba, Curasao, and Bonaire, to which the trinomial Glossophaga longirostris elongata Miller applies, are characterized by small measurements of the wing and a moderately long cranium that is narrow throughout. Bats from northern Colombia and northwestern Venezuela constitute another distinct race, Glossophaga longirostris longirostris Miller, that is moderately large in external and cranial dimensions, has reduced postorbital swellings, and a slightly dished facial profile. Another subspecies, Glossophaga longirostris major Goodwin, occurs along the xeric coast of Venezuela southward through the High Llanos of Colombia and Venezuela, and eastward to Trin¬ idad. This race has a short and high braincase, dished facial pro¬ file, and inflated postorbital swellings. The disjunct population of bats in the upper Magdalena River Valley represents an unde¬ scribed subspecies that is distinguished by its massive size in OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY £ £ d o +i -r d +1 9 2 c? °0 d od +1 9 00 tc d d +1=V o 9cm O o +1 9 9 — ri 2° cm : CO o O d CO 5T ? * s: o S : d 6 -H t £ ®> 00 © 2? q d «, -Hcm +1cm £9 d 00 oc°o CO 9 CM ~ CM _ 5 « s 2 (ooo id® — r. +j ^ ■ CM CM Si 2CM 2 O CM o TT HH lO 0 cmor, d ^ ■H w g _■ 05 ■ oo ■ oo ■ CO Sn g n S W ffi CMco ^ in 2 ^ CM CM - ^GO -H e? Q on +1<N CO CO • CM 94 S3 24 CMST d d +1CM ? 9 S3 =4 d d +1CM S3 9 a o is £ q |g o 52. CMLO 2 5 3'S w Sf 94 ^ ■ CM ^ ^ M3 ^00 CM O ao O * -H ^ -He 00 rL 40 c ■— Tf 1 ■ cm c OT cm ' ^5 X* UO 40 40 ooo -H *» S 9 S3 S ° 2 S3 9 ° 7 SR O So q SjrSi i! TT d 2 CTl CM 2 ~ ° q ~<0 q rd 2 d cr> mC 54 d ~ 2:! o Kg? 2 MO O . +1 9 00 °0 -24 o q +1 q S300 2 40 o X d -H 9 9 5 d 9 -H 9 d 9. iTi 00 o cm +1 9 CM +1 9 ,9 ^ d 9 +|oo S10 00 2 2S 2SS 22 os 00 O CO o T -H , ^ as •05 ao oo 'f rl on 3*0 lo -H ^ S3 9 od * 2S o 54 +1 9 m 9 d « CM 00 d 2. : _Si 92 o 9 +1 9 Sm 00 2 -H 9 CO(yj d 2 OO c.® - 2 O o—; +1 9 +1 d oo00 WEBSTER AND HANDLEY— GLOSSOPHAGA LONGIROSTRIS t©’ 00 *3 I d <n \i Si IS 11 d 00 d ^ «i iig I! d g _ OO IS - ^ S-- b'S £b dos JrS tC & g*4n § IS 3si ; o • * !S IS °T ~ 2 si § b '<j> • so li ob h ss s< 5s sI si £ si I b i II II Is II fi II cS »S = 14 S - S S IS 5? l| M S CJ C4 -i S in o5> dos -H T" SS 11 ll Is IS Is 2 £ gS £3 83 a a is Ssi O O : c. w! 3 04 g m ?S is 5 o d ^ b +j n ^ s§ n d s. oijfc IS IK £ - S cn £ 53 IS d on £ d 5 d d 32. d b OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY external and cranial measurements and well-developed postpalatal processes. The insular populations that are known from Tobago and the Windward Islands from Grenada northward to St. Vin¬ cent represent another distinct race, Glossophaga longirostris rostrata Miller, that has a narrow rostrum, moderately reduced post¬ orbital swellings, a low and narrow braincase, and zygomata that converge anteriorly. Glossophaga longirostris campestris, new subspecies Holotype. —Adult female, skin and skull, no. 389033, U.S. National Museum of Natural History; from Hato San Jose, 20 km. W. Paragua (=146 km. S, 7 km. E Ciudad Bolivar), Bolivar Venezuela, 300 m.; obtained on 8 April 1967 by N. Peterson, D. Peacock, and R. Peacock, Smithsonian Venezuelan Project, origi¬ nal no. SVP 12679. Selected external and cranial measurements of the holotype are: total length, 72; length of tail, 8; length of hind foot (dry), 12; length of ear from notch, 15; length of forearm (dry), 38.3; length of tibia (dry), 14.3; length of calcar (dry), 4.5; greatest length of skull, 22.2; zygomatic breadth, 9.6; breadth of braincase, 8.4; depth of braincase, 6.9; postorbital breadth, 4.6; length of maxillary toothrow, 7.6; maxillary breadth, 5.9; breadth across canines, 4.0; weight, 11.7 grams. Distribution. —Llanos and Rio Ventuari Basin of central Vene¬ zuela and savanna-clad lowlands around the Kanuku Mountains in Guyana and adjacent Brazil (Fig. 2); known altitudinal distri¬ bution from 50 to 300 meters. Diagnosis. —For a Glossophaga longirostris, this subspecies is small both externally and cranially (Table 1). G. /. campestris is variable in cranial proportions, but on average the rostrum is short and narrow, braincase shallow and moderately low, facial profile moderately dished, and postorbital swelling moderately large. Comparisons. —G. /. campestris can be distinguished from other subspecies by its small size and short, low braincase. Intergrada¬ tion between G. 1. major and G. /. campestris is evident in speci¬ mens from southern Guarico and northern Apure, Venezuela. The braincase of these bats is deeper and more bulbous (similar to G. 1. major) than in typical G. 1. campestris; however, they resemble the latter in having a short, narrow rostrum and in small overall size. WEBSTER AND HANDLEY—GLOSSOPHAGA LONGIROSTRIS 9 Remarks .—The isolated populations around the Kanuku Mountains suggests that G. /. campestris may be distributed dis¬ continuous^ in suitable habitat throughout the Gran Sabana, especially along river valleys. However, it was not found at Icaburu at the southern edge of the Gran Sabana in Venezuela. Spec¬ imens of G. 1. campestris have been collected from roosts in rock crevices, hollows of trees, and buildings. Pregnant females are known from December, and reproductively inactive females have been collected in April, June, and December. The Latin epithet campestris, of or pertaining to fields or grass¬ lands, refers to the preferred habitat of this race of G. longirostris. Specimens examined (292).— Brazil. Roraima: Lucetania Ranch, 8 (ROM). Guyana. Rupunum: Achimeriwau Mouth, 1 (ROM); “Bush Island,’’ Dadanawa, 2 (AMNH); Chipirari Wau Mouth, 15 mi. E Dadanawa, 1 (ROM); Contanrib Island, 5 mi. above Dadanawa, 10 (ROM); 15 mi. ENE Dadanawa, 1 (ROM); 0-20 mi. E Dadanawa, 18 (2 AMNH, 16 ROM); Illia Wau River, 1 (ROM); Karanambo, 1 (AMNH); Kuitaro River, 5 (ROM); Mt. Tawatawun, ca. 8 km. E Dadanawa, 3 (ROM); Ruawau River, 30 mi. SW Dadanawa, 3 (ROM); Rock Cave, ca. Sand Creek Reservation, 5 (ROM); Sand Creek Village, 8 (ROM); “southern savannas,’’ 17 (USNM); upper Sawariwau River, 7 (ROM); Wee Wee Tan, Makow Tan, 5 (ROM); Weri More, 4 (ROM). Venezuela. Apure: Hato Cariben, La Villa, 60 km. (=32 mi.) NE Puerto Paez, 76 m., 97 (USNM); Hato “La Guanota,” 6 km. W San Fernando de Apure, 100 m., 1 (TCWC); Rio Cinaruco, 41 km. NW Puerto Paez, 24 (USNM); ca. 4 km. W San Fernando de Apure, 66 m., 2 (TCWC); San Rafael de Atamaica, 45 km. S, 6 km. E San Fernando de Apure, 100 m., 7 (USNM). Bolivar: Hato La Florida, 14 km. S, 45 km. E Caicara, 50 m., 10 (USNM); Hato San Jose, 20 km. W La Paragua (=146 km. S, 7 km. E Ciudad Bolivar), 300 m., 10 (USNM); Isla de Cuba (Playa del Medio), Rio Orinoco, 3 (UCV); Paso Caruachi, Rio Caroni, 3 (UCV). T. F. Amazonas: Chaparito, 9 km. SE Puerto Ayacucho, 119 m., 6 (USNM); Coromoto, 25 km. SSE Puerto Ayacucho, 126 m., 1 (USNM); Guayabal, 28 km. S Puerto Ayacucho, 135 m., 1 (USNM); Las Queseras, 12 km. SSE Puerto Ayacucho, 135 m., 4 (USNM); Morocoy, 65 km. SSW Puerto Ayacucho, 161 m., 3 (USNM); Paria, 25 km. SSE Puerto Ayacucho, 114 m., 2 (USNM); Puerto Ayacucho, 2 (UCV); 0.75 km. E San Juan, Rio Manapiare, 1 (USNM); ca. 4 km. San Juan, Rio Manapiare, ca. 163 km. ESE Puerto Ayacucho, ca. 155 m., 13 (USNM); Tamanaco, 4 km. NE San Juan, Rio Manapiare, 155 m., 2 (USNM). Glossophaga longirostris elongata Miller 1900. Glossophaga elongata Miller, Proc. Biol. Soc. Washington, 13:124, 6 April. 1958. Glossophaga longirostris elongata, Koopman, Evolution, 12:437, December. Holotype .—Adult female, skin (from specimen in alcohol) and skull, no. 101871, U.S. National Museum of Natural History; from Willemstad, Curasao; obtained on 4 December 1899 by J. L. Guthrie, no original number. Selected external and cranial measurements of the holotype are: total length, 65; length of tail, 6; length of hind foot (dry), 11; 16 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY Pregnant females have been collected in June and July and a lactating female was taken in June. Two females captured in November evinced no reproductive activity. Specimens in the process of molt have been captured in November. Although the holotype was recorded as not having a tail, the sheathlike portion of the uropatagium that enclosed the caudal vertebrate is clearly visible in that specimen, and the tail length averages 6.67 (range 4-10) in six other specimens from the vicinity of the type locality. The Latin epithet reclusa, separated or removed, refers to the apparently disjunct nature of this population of bats in the upper Magdalena Valley. Specimens examined (25).—Colombia. Cundinimarca: 1 km. NE Giradot, 3 (AMNH). Huila: 5 km. N Villavieja, 1400 ft., 4 (MVZ); 4-7.5 km. E Villavieja, 1400 ft., 15 (MVZ); 17.5 km. SE Villavieja, 1600 ft., 3 (MVZ). Records of G. longnostris from two localities in the upper Magdalena Valley of Cundinimarca (Mesitas del Colegio) and Tolima (Mariquita), as well as another in Meta (Villavicencio) on the eastern Andean slope, by Valdivieso and Tamsitt (1962) were not referred to in subsequent publications by the same authors (Tam¬ sitt and Valdivieso, 1963; Valdivieso, 1964). Glossophaga longirostris rostrata Miller 1913. Glossophaga rostrata Miller, Proc. Biol. Soc. Washington, 26:32, 8 February. 1913. Glossophaga longirostris rostrata Miller, Proc. U.S. Nat. Mus., 46:423, 31 December. Holotype .—Original description based on a specimen assumed to be an adult male (see below), skin and skull, no. 111500, U.S. National Museum of Natural History, from Westerhall Estate, Grenada; date of capture unknown (“1900”), obtained by P. Gellineau, original no. 29. Selected cranial measurements of the holotype are: greatest length of skull, 22.6; zygomatic breadth, 9.8; breadth of braincase, 8.8; depth of braincase, 7.1; postorbital breadth, 4.8; length of maxillary toothrow, 8.0; maxillary breadth, 5.7; breadth across canines, 4.1. Distribution .—Lesser Antilles from St. Vincent southward to Grenada (except Barbados), and Tobago (Fig. 2); known altitudi¬ nal distribution from sea level to approximately 625 meters. Diagnosis .—External and cranial dimensions average in the middle of the range for the species (Table 1); larger overall on Tobago, smaller on St. Vincent. The rostrum is narrow, and the postorbital swellings are moderately reduced. The facial profile is moderately dished and the zygomata converge anteriorly. WEBSTER AND HANDLEY— GLOSSOPHAGA LONGIROSTRIS 17 Comparisons. — G. 1. rostrata is distinguished from other races by its narrow rostrum and reduced postorbital swellings. The braincase is clearly less bulbous than that of G. 1. major from Trinidad and northeastern Venezuela. Remarks .—Handley and Webster (1985) argued that the holotype, selected by Miller from among the 14 specimens collected by P. Gellineau on Grenada in 1900, consists of a mismatched skin (a male) and skull (a female). They, therefore, restricted the holotype designation to the female skull, the male skin becoming a paratype. Furthermore, they noted that the purported date of col¬ lection, in fact, was the date that Gellineau mailed the specimens to the USNM, not the actual date of capture. Specimens of G. 1. rostrata have been taken from hollow trees in Grenada. They were found roosting with Peropteryx macrotis, Micronycteris megalotis, Phyllostomus hastatus, and Carollia perspicillata on Tobago (Goodwin and Greenhall, 1961). Preg¬ nant females have been captured in August and December, and lactating females are known from March, June, and August. Indi¬ viduals in the process of molt have been collected in August and September. Specimens examined (151).— Lesser Antilles. Grenada: Grand Etang, 2000 ft., 5 (1 AMNH, 4 MCZ); Grenada, 1 (BMNH); Grenville Vale, 5 (AMNH); Pointe Saline, 3 (AMNH); 0-3.5 mi. NE St. George, 16 (6 KU, 9 MCZ, I TTU); St. Pauls, 4 (TTU); True Blue, 3 (AMNH); 0.5 mi. E Vendome, 1000 ft., 7 (1 KU, 6 TTU); Westerhall Estate, 14 (USNM); no locality, 7 (ROM). Grenadines: Carriacou, 23 (17 MCZ, 6 TTU); Mustique, 1 (BMNH); Union Island, 1 (MCZ). St. Vincent: Brighton, 8 (AMNH); Clifton Hill, 400 ft., 22 (KU); Grand Sable Estate, 7 (MCZ); Mesopatamia, 300 ft., 2 (KU); no locality, 4 (2 BMNH, 2 ROM). Tobago. Little Tobago: 2 (AMNH). St. Andrews: Orange Hill Rd., 3 (AMNH); Scarborough Government House, 3 (AMNH). St. David: Grafton, ca. Plymouth, 1 (LSU). St. John: 1 km. N Speyside, 5 (USNM). St. Mary: Pembroke, Gold Borough Estate, 1 (AMNH). St. Patrick: 1 km. SW Crown Point Airport, 3 (USNM). Additional distributional records for G. /. rostrata from Grenada are in G. M. Allen (1911) and from Tobago are in Husson (1954). Biological Observations Ecology. — Glossophaga longirostns is a denizen of numerous lowland tropical and subtropical habitats in northern South America, including arid thorn forests, deciduous and evergreen forests, and savannas. In Venezuela (Handley, 1976), 837 individu¬ als were collected within the following ecological parameters: site moisture—most frequently in dry areas (65 per cent) or near streams and other moist areas (35 per cent); habitat type—thorn forest (49 per cent), savannas and other open areas (43 per cent), 18 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY 50 22 12 6 48 96 38 14 23 14 14 46 Month Fig. 3.—Reproductive activity in Glossophaga longirostns. The histogram represents the per cent of 383 females evincing pregnancy (open) or lactation (stippled) each month; the actual number of pregnant and lactating females is shown in the histogram, and the total number of females examined is below each month. and moist forests (eight per cent); vegetative life zone—tropical dry forest (43 per cent), tropical thorn forest (35 per cent), tropical very dry forest (17 per cent), other dry zones (one per cent), and moist zones (four per cent). Daytime roosts include caves, tunnels, culverts, crevices in rocks, hollow trees, and houses and other buildings. Specimens have been captured from sea level to approximately 650 meters, but most are from less than 500 meters. Other bats known to share daytime roosts with G. longirostris include Peropteryx macrotis, Micronycteris megalotis, Phyllostomus hastatus, Glossophaga soricina, and Carollia perspicillata. Reproduction. —The reproductive condition of 383 females from throughout the geographic distribution of Glossophaga longiros- WEBSTER AND HANDLEY-GLOSSOPHAGA LONGIROSTRIS 19 Table 2.— Timing of molt in Glossophaga longirostris based on the examination of 144 adult specimens. Month tris was used to determine the percentage of pregnant and lactating females for each month (Fig. 3). Two periods of pregnancy are evident—one extending from December to April and another from June to October. Lactating females have been collected in every month except February. Thus, the reproductive strategy of G. longirostris appears to be that of monotocous polyestry with a bimodal cycle. Molt. —Molt in adult G. longirostris occurs from June to November (Table 2), and appears to occur annually. The timing of molt is variable among specimens collected from the same place and at the same time and is not highly synchronized. A female from Isla Margarita (KU 118110) was pregnant while in the process of molt; therefore molt may not be delayed in reproductively active females in this tropical species as was found in Eptesicus fuscus in temperate North America (Jones and Genoways, 1967). The sequence of molt in Glossophaga longirostris is similar to that of G. soricina (Jones et al., 1973). Molt begins as a growth of new hair under the old. Then after the new fur has grown to half its normal length, the worn hair drops out in irregular patches rather than in symmetrical fashion. Generally, the hair on the head and shoulders is lost initially, but in some specimens hair on the back is replaced first. Acknowledgments We thank those in charge of the following collections for allowing us access to specimens in their care: American Museum of Natural History (AMNH); British Museum (Natural History) (BMNH); Carnegie Museum of Natural History 20 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY (CMNH); Field Museum of Natural History (FMNH); Museum of Natural His¬ tory, University of Kansas (KU); Los Angeles County Museum of Natural History (LACM); Museum of Zoology, Louisiana State University (LSU); Museum of Comparative Zoology, Harvard University (MCZ); Museum of Vertebrate Zoology, University of California at Berkeley (MVZ); Stovall Museum of Science and His¬ tory, University of Oklahoma (OU); Royal Ontario Museum (ROM); Texas Coop¬ erative Wildlife Collection, Texas A&M University (TCWC); The Museum, Texas Tech University (TTU); Universidad Central de Venezuela (UCV); Museum of Zoology, University of Michigan (UMMZ); and the U.S. National Museum of Nat¬ ural History (USNM). We are grateful to Smithsonian Venezuelan Project (SVP) personnel who collected specimens and natural history data, Jenny Banner and Penny Nelson, who helped measure SVP skulls, and Darylen Handley who com¬ puted the statistics. This research was supported in part by grants to Webster by the Institute of Museum Research, Texas Tech University, and the Theodore Roosevelt Memorial Fund, American Museum of Natural History. Handley’s Smithsonian Venezuelan Project was supported by the Smithsonian Institution and by a contract (DA-49MD-2788) of the Medical Research and Development Command, Office of the Sur¬ geon General, U.S. Army. Literature Cited Albuja, L. 1983. Murcielagos del Ecuador. Escuela Politecnia Nacional, Quito, Ecuador, xii + 285 pp. Allen, G. M. 1911. Mammals of the West Indies. Bull. Mus. Comp. Zool., 54:175-263. Allen, J. A. 1900. List of bats collected by Mr. H. H. Smith in the Santa Marta region of Colombia, with descriptions of new species. Bull. Amer. Mus. Nat. Hist., 13:87-94. -. 1911. Mammals from Venezuela collected by Mr. M. A. Carriker, Jr., 1909-1911. Bull. Amer. Mus. Nat. Hist., 30:239-273. -. 1916. New South American mammals. Bull. Amer. Mus. Nat. Hist., 35:83-87. Cabrera, A. 1958. Catalogo de los mamiferos de America del Sur. Rev. Mus. Argentino Cien. Nat. “Bernardino Rivadavia,” Cien. Zool., 4:iv + 1-307. Genoways, H. H., and S. L. Williams. 1979. Notes on bats (Mammalia: Chiroptera) from Bonaire and Curasao, Dutch West Indies. Ann. Carnegie Mus., 48:311-321. Goodwin, G. G. 1958. Three new bats from Trinidad. Amer. Mus. Novitates, 1877:1-6. Goodwin, G. G., and A. M. Greenhall. 1961. A review of the bats of Trinidad and Tobago. Bull. Amer. Mus. Nat. Hist., 122:187-301. Handley, C. O., Jr. 1959. A revision of American bats of the genera Euderma and Plecotus. Proc. U.S. Nat. Mus., 110:95-246. -. 1976. Mammals of the Smithsonian Venezuelan Project. Brigham Young Univ. Sci. Bull., Biol. Ser., 20(5):l-89. Handley, C. O., Jr., and W. D. Webster. 1986. The supposed occurrence of Glossophaga longirostris Miller on Dominica and problems with the type series of G. rostrata Miller. J. Mamm., in press. Hummelinck, P. W. 1940. Studies on the fauna of Curacao, Aruba, Bonaire, and the Venezuelan Islands, 1:1-130. WEBSTER AND HANDLEY—GLOSSOPHAGA LONGIROSTRIS 21 Husson, A. M. 1954. On Vampyrodes caracciolae (Thomas) and some other bats from the island of Tobago (British West Indies). Zool. Mededelingen, Leiden, 33:63-67. -. 1960. De Zoolgdieren van de Nederlandse Antillen. Natuurwetenschappelijke Werkgroep Nederlandse Antillen, Curasao, 170 pp. Jones, J. K., Jr., and D. C. Carter. 1976. Annotated checklist, with keys to subfamilies and genera. Pp. 7-38, in Biology of bats of the New World family Phyllostomatidae. Part I (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.), Spec. Publ. Mus., Texas Tech Univ., 10:1-218. Jones, J. K., Jr., and H. H. Genoways. 1967. Annotated checklist of bats from South Dakota. Trans, Kansas Acad. Sci., 70:184-196. Jones, J. K., Jr., J. D. Smith, and H. H. Genoways. 1973. Annotated checklist of mammals of the Yucatan Peninsula, Mexico. I. Chiroptera. Occas. Papers Mus., Texas Tech Univ., 13:1-31. Koopman, K. F, 1958. Land bridges and ecology in bat distribution on islands off the northern coast of South America. Evolution, 12:429-439. Linares, O. J. 1968. Quiropteros subfosiles encontrados en las cuevas Venezolanas. Bol. Soc. Espeleol., 1:119-145. Miller, G. S., Jr. 1898. Descriptions of five new phyllostome bats. Proc. Acad. Nat. Set. Philadelphia, 50:326-337. -. 1900fl. Three new bats from the island of Curasao. Proc. Biol. Soc. Washington, 13:123-127, -. 19005. A second collection of bats from the island of Curasao. Proc. Biol. Soc. Washington, 13:159-162. -. 1913a. Five new mammals from tropical America. Proc. Biol. Soc. Washington, 26:31-33. -. 19135. Revision of bats of the genus Glossophaga. Proc. U.S. Nat. Mus., 46:413-429. Morales-Alarcon, A., E. Osorno-Mesa, C. Bernal Cubides, and A. LlerasPizarro. 1968. Aislamiento de virus rabico de murcielagos en Colom¬ bia, S. A. Caldasia, 10:167-172. Pirlot, P. 1964. Nota sobre la ecologia de ciertos quiropteros de la region del Rio Palmar. Rev. Univ. Zulia, Kasmera, 1:289-307. -. 1965. Chiropteres de Test de Venezuela. II. Delta de l’Orenoque. Mammalia, 29:375-389. Ridgway, R. 1912. Color standards and color nomenclature. Washington, D.C., privately printed, 43 pp. Robinson, W., and M. W. Lyon. 1901. An annotated list of mammals collected in the vicinity of La Guaire, Venezuela. Proc. U.S. Nat. Mus. 24:135-162. Sanborn, C. C. 1933. Bats of the genera Anoura and Lonchoglossa. Field Mus. Nat. Hist., Zool. Ser., 20:23-27. Smith, J. D., and H. H. Genoways. 1974. Bats of Margarita Island, Venezuela, with zoogeographic comments. Bull. So. California Acad. Sci., 73:64-79. Tamsitt, J. R., and D. Valdivieso. 1963. Records and observations on Colom¬ bian bats. J. Mamm., 44:168-180. Valdivieso, D. 1964. La fauna quiroptera del Departamento de Cundinimarca, Colombia. Rev. Biol. Trop., 12:19-45. Valdivieso, D., and J. R. Tansmitt. 1962. First records of the pale spear-nosed bat in Colombia. J. Mamm., 43:422-423. 22 OCCASIONAL PAPERS MUSEUM TEXAS TECH UNIVERSITY Webster, W. D., and J. K. Jones, Jr. 1980. Taxonomic and nomenclatorial notes on bats of the genus Glossophaga in North America, with descrip¬ tion of a new species. Occas. Papers Mus., Texas Tech Univ., 71:1-12. Wille, A. 1954. Muscular adaptation of the nectar-feeding bats (subfamily Glossophaginae). Trans. Kansas Acad. Sci., 57:315-325. Addresses of authors: W. D. Webster, Department of Biological Sciences and The Museum, Texas Tech University, Lubbock, Texas 79409; C. O. Handley, Jr., Division of Mammals, Smithsonian Institution, Washington, D.C. 20560; Present address of Webster, Department of Biological Sciences, University of North Caro¬ lina at Wilmington, Wilmington, North Carolina 28403. Received 7 September 1983, accepted 19 November 1984.