Part 1

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OCCASIONAL PAPERS

of the MUSEUM OF NATURAL HISTORY --The University of Kansas Lawrence, Kansas

NUMBER 2 APRIL 29, 1971

NOTEWORTHY RECORDS OF BATS FROM NICARAGUA, WITH A CHECKLIST OF THE CHIROPTERAN FAUNA OF THE COUNTRY

By

J. KNOX JONES, JR.,[1] JAMES DALE SMITH,[2] RONALD W. TURNER[3]

Nicaragua occupies a strategic position in Central America with respect to mammalian distributional patterns, but relatively little has been published concerning the fauna of the country and its zoogeographic relationships. The present paper records information on distribution, variation, and natural history of 40 species of bats from Nicaragua, 14 of which are here recorded for the first time from the country. Appended is a checklist of the chiropteran fauna of Nicaragua in which only primary literature with actual reference to specimens from the republic is cited.

The specimens upon which this report is based are, with few exceptions, in the collections of the Museum of Natural History of The University of Kansas. Some of our material was obtained in 1956 by J. R. and A. A. Alcorn, field representatives of the Museum and sponsored by the Kansas University Endowment Association; most of the specimens, however, were obtained by field parties of which we were members that worked in Nicaragua in 1964, 1966, 1967, and 1968 under the aegis of a contract (DA-49-193-MD-2215) between the U.S. Army Medical Research and Development Command and The University of Kansas. Place-names associated with localities mentioned in the text from which specimens at Kansas were collected are plotted on Fig. 1.

[1] Curator, Division of Mammals, Museum of Natural History, University of Kansas.

[2] Assistant Professor, Department of Biology, California State College, Fullerton, California.

[3] Assistant Professor, Department of Biology, St. Benedicts College, Atchison, Kansas.

In the accounts that follow, departments in Nicaragua are listed alphabetically, but localities within each department are arranged from north to south; elevations are given in meters or feet, depending on which was used on specimen labels. All specimens are in the Museum of Natural History of The University of Kansas unless noted otherwise. We are indebted to Drs. Charles O. Handley, Jr., and Ronald Pine of the U.S. National Museum (USNM) for lending us certain critical specimens.

ACCOUNTS OF SPECIES

Saccopteryx leptura (Schreber, 1774)

Two specimens from El Paraíso, 1 km N Cosigüina, 20 m, Chinandega, on the Cosigüina Peninsula, provide the fourth locality of record for this white-lined bat in Nicaragua. Jones (1964a:506) and Davis _et al._ (1964:375) earlier reported a total of eight specimens from the departments of Managua and Zelaya. The species is known as far north in Middle America as Chiapas (Carter _et al._, 1966:489).

Our two bats, both females, were shot on the evening of 1 March 1968 as they foraged around a yard light. One carried an embryo that measured 8 mm (crown-rump), whereas the other was reproductively inactive.

Peropteryx macrotis macrotis (Wagner, 1843)

Four females (one young and three adult) captured 5 km N and 9 km E Condega, 800 m, in Madriz, on 23 June 1964, provide the first record of this small sac-winged species from Nicaragua. The bats were shot from daytime roosts in small, well-lighted, cave-like spaces formed among immense blocks of granite in a small patch of tropical deciduous forest surrounded by extensive pine-oak woodland. None of the adult females was reproductively active. _Glossophaga soricina_, _Diphylla ecaudata_, and a large nursery colony of _Desmodus rotundus_ were found in association with the _Peropteryx_. Measurements of our specimens agree closely with those reported for material from El Salvador (Felten, 1955:284) and Costa Rica (Starrett and Casebeer, 1968:3-4).

Noctilio labialis labialis (Kerr, 1792)

_Specimens._--_Boaco_: 4 km W Teustepe, 140 m, 9. _Chontales_: Hato Grande, 13 km S, 8 km W Juigalpa, 60 m, 49. _Rivas_: 4 km S, 1.5 km E Alta Gracia, 40 m, Isla de Ometepe, 1; Finca Amayo, 13 km S, 14 km E Rivas, 40 m, 4. _Zelaya_: S side Río Mico, El Recreo, 25 m, 1; Cara de Mono, 50 m, 2.

This species has been reported previously from Nicaragua by several authors. All our specimens were netted over small streams or shot as they foraged; parts of scarabids and lepidopterans were found in the mouths of several individuals shot at Finca Amayo. Twenty-six of 31 autopsied females taken in April were pregnant, each containing a single embryo--average crown-rump length 16.7 (5-26) mm. Testes of 15 males collected in April had an average length of 4.6 (2-7) mm, those of four taken in June, 5.2 (4-6) mm.

We follow Cabrera (1958:55), Husson (1962:63), and Handley (1966b:758) in use of the subspecific name _labialis,_ the type locality of which is the "Mosquito shore" of Nicaragua, rather than Perú as suggested by Hershkovitz (1949:433-434).

Noctilio leporinus mexicanus Goldman, 1915

_Specimens._--_Chinandega_: Potosí, 5 m, 2. _Chontales_: Hato Grande, 13 km S, 8 km W Juigalpa, 60 m, 4. _Rivas_: 4 km S, 1.5 km E Alta Gracia, 40 m, Isla de Ometepe, 4; Mérida, 40 m, Isla de Ometepe, 2; Finca Amayo, 13 km S, 14 km E Rivas, 40 m, 1.

This fish-eating species, first reported from Nicaragua by Davis _et al._ (1964:376), apparently occurs throughout Middle America, although known from the region by comparatively few records. We have 13 additional Nicaraguan specimens as listed above.

The two individuals (both females, one pregnant with an embryo that measured 20 mm) from Potosí were caught on 6 March in a mist net set over a large pool in a shallow estuarine stream; the mouth of the stream opened into the Gulf of Fonseca approximately 200 yards below our netting site. Other individuals of this species were observed as they foraged over large pools formed at high tide near the mouth of the stream. Our other specimens were caught in mist nets set over fresh water streams near Lake Nicaragua or along the shores of the lake. A female from near Alta Gracia, caught on 27 March, carried a single embryo that measured 41 mm, whereas one from Finca Amayo was lactating on 25 June. Four males taken on Isla de Ometepe in late March and early April had a mean testicular length of 9.5 (8-10) mm.

Pteronotus davyi fulvus (Thomas, 1892)

_Specimens._--_Chontales_: Cuapa, 4. _Matagalpa_: 3 mi E San Ramón, 126.

This small naked-backed bat has not been reported previously from Nicaragua. Autopsy of seven females collected on 9 May near San Ramón revealed that four were pregnant, each with a single embryo--average crown-rump length 25.0 (21-29) mm.

The distribution of _P. davyi_ is poorly known in much of Central America. The species was reported only recently from Costa Rica (Starrett and Casebeer, 1968:8) and is unknown from Panamá. We assign our specimens tentatively to the subspecies _fulvus_ on geographic grounds.

Pteronotus parnellii fuscus (J. A. Allen, 1911)

_Specimens._--_Boaco_: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 1; Los Cocos, 14 km S Boaco, 220 m, 1. _Chinandega_: 6.5 km N, 1 km E Cosigüina, 10 m, 1. _Zelaya_: Bonanza, 850 ft, 1; 2 mi SW Bonanza, 600 ft, 1; S side Río Mico, El Recreo, 25 m, 1; Cara de Mono, 50 m, 4.

Although this species is widespread in Middle America, it has been known previously from Nicaragua only by a specimen from "Chontales" (Miller, 1902:402). All of our specimens were captured in mist nets. Two adult females (one taken on 28 February and the other on 8 March) each carried a single embryo (13 and 18 mm in crown-rump length, respectively); a female taken in April was lactating and had a flaccid uterus suggesting relatively recent parturition. Adult females captured in the months of June, July, and August evidenced no sign of reproductive activity. Two of these were in dark, fresh pelage, but one captured on 24 June and another on 28 July were molting. In both individuals, active molt evidently had slowed or subsided and remnants of the old pelage (rich ochraceous orange) were confined to a narrow strip at the lateral edges of the body.

We tentatively apply the subspecies name _fuscus_ to our Nicaraguan bats on geographic grounds; one of us (Smith) currently is investigating geographic variation in the genus _Pteronotus_.

Pteronotus suapurensis (J. A. Allen, 1904)

_Specimens._--_Chontales_: Cuapa, 1. _Matagalpa_: 3 mi E San Ramón, 24. _Zelaya_: Cara de Mono, 25 m, 4.

This relatively poorly known monotypic species occurs from southern Veracruz to the Amazon Basin. It has been reported from several localities in Central America including one in Nicaragua (Goodwin, 1942a:88). Three pregnant females from near San Ramón (9 May) carried embryos that had crown-rump lengths of 27, 27, and 28 mm, and two netted on 24 April at Cara de Mono each had an embryo that measured 22 mm. All of our specimens are in bright pelage that is fulvous red in color.

Micronycteris megalotis mexicana Miller, 1898

_Specimens._--_Carazo_: 3 km N, 4 km W Diriamba, 600 m, 5. _Granada_: Hda. Mecatepe [2 km N, 11.5 km E Nandaime], 5. _Managua:_ 5 mi NW Managua, 1. _Matagalpa_: Río Viejo, 7 mi WNW Darío, 4; 11 mi SE Darío, 1. _Zelaya_: S end Isla del Maíz Grande, 9.

The individuals from Isla de Maíz Grande were shot on 30 June as they flew from small caves and crevices in rocky outcrops on the south end of the island. Of the three adult females obtained at that time, two were lactating; each was accompanied by a young, the forearms of which measured 19.7 (male) and 21.3 mm (female). Two adult males from Isla del Maíz Grande had testes 2 mm in length.

TABLE 1.--Selected measurements of adults of two subspecies of _Micronycteris megalotis_ from Nicaragua. Superscript numbers indicate fewer specimens averaged than indicated in left-hand column.

Table Legend:

Col. A: Number of specimens averaged or catalogue number, and sex Col. B: Length of forearm Col. C: Greatest length of skull Col. D: Zygomatic breadth Col. E: Breadth of braincase Col. F: Length of maxillary toothrow

=================================================================== A B C D E F ------------------------------------------------------------------- _Micronycteris megalotis mexicana_, west-central Nicaragua[a]

Average 8 35.4 19.5[6] 9.5 8.0[7] 7.5 (3[Male], 5 [Female]) Minimum 33.0 18.8 9.1 7.7 7.3 Maximum 38.1 20.3 9.8 8.2 7.7

Isla del Maíz Grande, Nicaragua

Average 6 35.6 19.4 9.2 7.8 7.4 (3 [Male], 3 [Female]) Minimum 34.5 19.1 9.0 7.7 7.3 Maximum 37.0 19.8 9.3 7.9 7.5

_Micronycteris megalotis microtis_, Greytown, Nicaragua (holotype)

USNM 16366/23364, [Male] 31.5 18.2 8.8 7.6 6.9

Río Coco, Nicaragua (AMNH)

Average 6 33.6 18.9[3] -- 7.5[3] 7.1[3] (3 [Male], 3 [Female]) Minimum 32.7 18.8 -- 7.4 7.0 Maximum 34.2 19.0 -- 7.6 7.2

Bonanza, Nicaragua

KU 96251, [Male] 32.3 18.4 8.8 7.6 7.0 -------------------------------------------------------------------

[a] Specimens labeled with reference to Darío, Diriamba, and Managua.

Four of the five specimens taken northwest of Diriamba were shot from a daytime retreat in a culvert; the fifth was caught by hand as it attempted to fly out of a hollow, fallen tree. Two adult females captured on 31 March were pregnant, each carrying a single embryo (13 and 14 mm in crown-rump length), whereas two obtained on 14 August showed no sign of reproductive activity. An adult male, also taken on 14 August, had testes 2 mm in length. Of three additional adult females, one captured on 3 June was pregnant (embryo 21 mm in crown-rump length), whereas two obtained on 14 April evinced no gross reproductive activity.

The subspecies _mexicana_ has not been reported previously from Nicaragua, although Gardner _et al._ (1970:715) recently extended its known distribution southward from Honduras (Goodwin, 1942c:124) and El Salvador (Felten, 1956:180) to west-central Costa Rica. In Nicaragua, as apparently in Costa Rica, _mexicana_ occupies the Pacific versant, the Caribbean lowlands being inhabited by the smaller race, _Micronycteris megalotis microtis_ Miller, 1898 (type locality, Greytown, Nicaragua--reported also from "Río Coco," Nicaragua, by J. A. Allen, 1910:110). Nicaraguan examples of _M. m. mexicana_ are, on the average, considerably larger in cranial dimensions and length of forearm than are specimens of _M. m. microtis_ (see Table 1). The series of _microtis_ from "Río Coco," probably from the headwaters of that stream in the vicinity of San Juan de Río Coco and thus near the divide between Caribbean and Pacific drainages, is intermediate in size between the two subspecies and suggests intergradation between them. As noted also by Gardner _et al._ (1970:715), we find no differences in length of ear between _microtis_ and _mexicana_ (contrary to Miller's claim, in the original description, that _microtis_ had noticeably smaller ears), nor do we note any consistent differences in color between the two races on the basis of the specimens at hand.

Our specimens from Isla del Maíz Grande inexplicably agree rather closely in size (average but slightly smaller) with specimens of _M. m. mexicana_ from western Nicaragua and elsewhere within the range of the subspecies, and are tentatively, therefore, referred to _mexicana_. Four specimens earlier reported (G. M. Allen, 1929:130) from the same island also are relatively large (forearm 35 mm). Further commentary on this insular population must await a time when additional material is available from Central America.

Tonatia nicaraguae Goodwin, 1942

_Specimens._--_Boaco_: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 1. _Chontales_: Hato Grande, 13 km S, 8 km W Juigalpa, 60 m, 1.

Only four specimens of this seemingly rare species have been recorded previously from Nicaragua--one (the holotype) from Kanawa Creek, near Cukra, north of Bluefields, 100 ft, Zelaya (Goodwin, 1942b:205), another from 12.5 mi. S and 13 mi. E Rivas, 125 ft, Rivas (Davis and Carter, 1962:67-68), and two from 6 km N Tuma, 500 m, Matagalpa (Valdez and LaVal, 1971:248). Our specimens were caught in mist nets placed over small, quiet streams that were fringed with tall, gallery forest. The surrounding areas were savanna-like with small stands of secondary forest. An adult male taken on 21 April had testes that measured 5 mm in length, whereas those of another (9 August) were 3 mm.

Handley (1966b:761) synonymized _T. nicaraguae_ with _Tonatia minuta_ Goodwin, 1942, a conclusion with which we agree. However, the name _nicaraguae_ (Goodwin, 1942b:205) has page priority over _minuta_ Goodwin (op. cit.:206) and is the valid name for the species rather than _minuta_ as used by Handley (see also LaVal, 1969:820; Gardner _et al._, 1970:716; Valdez and LaVal, 1971:248).

Measurements of the two males (that from Hato Grande listed first) are as follows: total length, 63, 60 mm; length of tail, 5, 6 mm; length of hind foot, 10, 9 mm; length of ear, 23, 23 mm; length of forearm, 34.5, 35.6 mm; weight, 11.4, 8.8 gms; greatest length of skull, 20.2, 20.5 mm; zygomatic breadth, 9.5, 9.5 mm; breadth of braincase, 8.5, 8.2 mm; postorbital breadth, 3.1, 2.9 mm; length of maxillary toothrow, 7.0, 7.1 mm; breadth across upper molars, 6.3, 6.5 mm. We have not compared our specimens directly with others from Middle America. On the basis of available measurements, they resemble material reported from Honduras (LaVal, 1969:820), averaging larger than other specimens for which measurements have been published (see especially Goodwin, 1942b:206; Davis and Carter, 1962:68; Davis _et al._, 1964:379; Gardner _et al._, 1970:716-717).

Phyllostomus hastatus panamensis J. A. Allen, 1904

_Specimens._--_Carazo_: 3 km N, 4 km W Diriamba, 600 m, 1. _Madriz_: Darailí, 5 km N, 14 km E Condega, 940 m, 4. _Managua_: 3 km SW Tipitapa, 40 m, 1; 3 km N Sabana Grande, 50 m, 2; 2 km N Sabana Grande, 40 m, 1; 5 mi S Managua, 2. _Matagalpa_: La Danta, 1 km N, 5 km E Esquipulas, 760 m, 1. _Nueva Segovia_: 4.5 km N, 2 km E Jalapa, 630 m, 2; 1.5 km N, 1 km E Jalapa, 660 m, 1. _Zelaya_: Bonanza, 850 ft, 2; El Recreo, 25 m, 8.

This large spear-nosed species has been reported previously from Nicaragua only from Matagalpa (Goodwin, 1942c:126). _Phyllostomus hastatus_ evidently occurs throughout the country and is relatively common in some places.

Two females taken 5 mi S Managua on 13 March each carried a single embryo (crown-rump length 27 and 32 mm). Lactating females were captured in June, July, and August.

Trachops cirrhosus coffini Goldman, 1925

A male fringe-lipped bat netted over a small stream at Cara de Mono, 50 m, Zelaya, in the Caribbean lowlands, on 24 April, 1968, constitutes the second known occurrence of this species in Nicaragua. Carter _et al._ (1966:491) earlier reported two males from Río Coco, 64 mi NNE Jinotega, 1000 ft, Jinotega. The testes of our specimen were 4 mm long.

Davis and Carter (1962:69), Carter _et al._ (_loc. cit._), and Starrett and Casebeer (1968:11) did not apply a trinomen to bats of this species and noted overlap of measurements between _T. c. cirrhosus_ (Spix) and _T. c. coffini_ Goldman. However, until detailed comparisons can be made, we follow Jones (1966:452) in retaining the subspecific name _coffini_.

Chrotopterus auritus auritus (Peters, 1856)

An adult male captured in a mist net set in a forest clearing at Santa María de Ostuma, 1250 m, Matagalpa, represents the first record of this bat from Nicaragua. The testes of our individual, taken on 1 July 1966, were 6 mm in length. Externally and cranially our Nicaraguan example closely resembles specimens of _C. auritus_ from Veracruz and the Yucatán Peninsula. Handley (1966b:762) and Starrett and Casebeer (1968:12) expressed doubt as to the validity of the currently used subspecific names in this species. Comparisons of cranial and external measurements of the material at hand with those given in various published accounts--Burt and Stirton (1961), Starrett and Casebeer (1968), and Thomas (1905), for example--reveal little variation. Until additional information is available (especially as concerns the South American races), however, we tentatively apply the subspecific name _C. a. auritus_ to Middle American populations.

Anoura geoffroyi lasiopyga (Peters, 1868)

Four specimens of this species, netted on 24 and 25 July 1967 in a banana grove 1.5 km N and 1 km E Jalapa, 660 m, Nueva Segovia, provide the first record of this glossophagine from Nicaragua. Two males each had testes 6 mm in length; one female evidenced no reproduction activity, whereas another carried an embryo 4 mm in crown-rump length. In addition to our material, there is a specimen in the British Museum (BM 8.6.22.4) from Cafetal "Concordia," 4000 ft, Jinotega.

It may be noteworthy that the two localities at which this bat is known both are in the highlands of the northern part of Nicaragua, and that we failed to take additional specimens in many hundreds of hours of netting in other places in the country.

Choeroniscus godmani (Thomas, 1903)

Godman's bat was reported from Nicaragua by Handley (1966a:86), who used the locality designation "El Realejo" for the three specimens available to him. Actually, the three are from the nearby Hda. San Isidro, 10 km S Chinandega. We have taken three additional specimens as follows: Santa Rosa, 17 km N and 15 km E Boaco, 300 m, Boaco (an adult female taken on 21 March, which was pregnant with an embryo measuring 16 mm in crown-rump length); 2 km E Yalí, 900 m, Jinotega (an adult male netted on 3 August, testes 4 mm in length); and Santa María de Ostuma, 1250 m, Matagalpa (a nonpregnant, adult female captured on 11 April). Bats of this species have been taken as far north as the western Mexican state of Sinaloa (Jones, 1964b:510).

Lichonycteris obscura Thomas, 1895

Managua is the type locality of this rare long-nosed species and a specimen was reported from 6 mi W Rama, 50 ft, Zelaya, by Davis _et al._ (1964:380). Our collection contains three males, one from Jalapa, 660 m, Nueva Segovia, captured on the night of 27 July 1967 as it flew around a lighted room in a house, and two from the south side of the Río Mico, El Recreo, 25 m, Zelaya, taken on 26 and 27 July 1966 (one was netted and the other caught by hand in the lighted room of a building at night).

Selected measurements of the three males are, respectively: total length, 55, 58, 57 mm; length of tail, 7, 10, 9 mm; length of hind foot, 7, 9, 9 mm; length of ear, 10, 11, 11 mm; length of forearm, 30.9, 31.0, 30.8 mm; weight, 6.8, 6.8, 6.2 gms; length of testes, 1, 2, 2 mm; greatest length of skull, 18.5, 18.0, 17.9 mm; breadth of braincase, 8.1, 8.4, 8.3 mm; length of maxillary toothrow, 5.7, 5.5, 5.7 mm.

Carollia castanea H. Allen, 1890

_Specimens._--_Boaco_: Santa Rosa, 17 km N, 15 km E Boaco, 300 m, 7. _Chontales_: 1 km N, 2.5 km W Villa Somoza, 330 m, 4. _Matagalpa_: 1 km NE Esquipulas, 420 m, 1. _Nueva Segovia_: 7 km N, 4 km E Jalapa, 660 m, 4. _Zelaya_: S side Río Mico, El Recreo, 25 m, 6; Cara de Mono, 50 m, 15.

Bats of this species apparently are common in the Caribbean lowlands of Nicaragua; the highest altitude at which we have taken specimens is 660 meters. The only previous report of _C. castanea_ from Nicaragua evidently is that of Davis _et al._ (1964:379), who mentioned it in passing (from Cacao, Zelaya) in an account of _Macrophyllum macrophyllum_. Two adult females each carried an embryo having crown-rump lengths of 20 (21 March) and 10 mm (27 July). Adult females evincing no gross reproductive activity were taken in February (one), March (one), April (four), June (three), July (two), and August (two). Four adult males netted on 24 April had testes with an average length of 7.0 (5-9) mm, whereas those of four other adult males taken in late June had an average of 4.0 (2-6) mm.

In our collections from Nicaragua, we find at least three kinds of _Carollia_ and we follow Handley (1966b:764-765) in tentatively assigning these to _C. castanea_, _C. subrufa_, and _C. perspicillata_. The systematics of this genus is currently under study by Ronald H. Pine. In Nicaragua, _castanea_ differs from _subrufa_ (with which it has been confused in the past) in being smaller, both externally and cranially, and much darker in color as well as in the additional characters mentioned by Handley (_loc. cit._).

Sturnira ludovici ludovici Anthony, 1924