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Chapter II: Part 2

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Bats of the genus _Molossus_ are in need of thorough systematic study. We think most, if not all, mainland populations of small _Molossus_ with pale-based hairs pertain to the species _Molossus molossus_, originally described from the Lesser Antilles. Although there is some variation in size and overall color among our Nicaraguan samples, all seem to represent that species. Because specimens from Nicaragua closely resemble examples of _M. m. aztecus_ from Jalisco, Mexico (Table 3), in external and cranial proportions, we tentatively refer them to _aztecus_.

In our field experience in both Middle and South America, colonies of _M. molossus_ tend to be extremely localized, with individual populations or demes frequently separated by many miles of territory in which the species seemingly does not regularly occur.

TABLE 3.--Selected measurements of adult _Molossus molossus_.

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: Condylobasal length Col. E: Zygomatic breadth Col. F: Mastoid breadth Col. G: Breadth of braincase Col. H: Postorbital constriction Col. I: Length of maxillary toothrow Col. J: Breadth across upper molars

====================================================================== A B C D E F G H I J ---------------------------------------------------------------------- Western Jalisco, Mexico

KU 109075, [Male] 38.2 18.0 15.3 11.0 10.5 9.1 3.8 6.2 7.9 KU 109076, [Male] 37.3 17.8 15.3 11.0 10.5 9.2 3.6 6.3 8.1 KU 120540, [Male] 37.7 17.8 15.3 11.0 10.8 8.9 3.9 6.3 8.1 Average 15 ([Female]) 37.1 17.2 14.8 10.7 10.3 8.9 3.8 6.1 7.9 Minimum 36.0 16.5 14.2 10.4 10.1 8.5 3.7 6.0 7.7 Maximum 38.1 17.7 15.2 11.0 10.5 9.2 3.9 6.2 8.2

Departamento de Chinandega, Nicaragua

USNM 337552, [Male] 39.0+- 18.2 15.5 11.0 10.7 9.0 3.7 6.0 8.0 KU 114140, [Female] 37.3 16.8 14.7 10.7 9.9 8.7 3.4 5.9 7.4

3 mi SW Managua, Nicaragua

KU 71009, [Male] 39.8 18.7 16.1 11.1 10.9 8.9 3.8 6.3 8.0 KU 71011, [Male] 39.4 18.5 16.5 -- 11.0 9.2 3.7 6.7 8.0 Average 5 ([Female]) 38.9 17.5 15.4 10.7 10.3 9.0 3.6 6.1 7.8 Minimum 38.1 17.2 15.1 10.5 10.1 8.9 3.5 5.9 7.6 Maximum 39.7 18.0 15.9 10.9 10.6 9.1 3.7 6.3 8.0

Santa Rosa, Boaco, Nicaragua

Average 6 ([Female]) 36.0 17.5 14.7 10.8 10.6 9.1 3.7 6.0 8.0 Minimum 35.5 17.3 14.4 10.6 10.4 9.0 3.6 5.7 7.8 Maximum 36.7 17.8 15.1 11.0 10.8 9.2 3.8 6.2 8.2

Rivas, Nicaragua

KU 106291, [Male] 38.2 18.8 16.1 11.5 10.9 9.4 3.8 6.6 8.3 KU 106290, [Female] 39.6 17.4 15.0 11.0 10.5 9.1 3.6 6.0 7.7 KU 106293, [Female] 37.2 17.3 14.8 10.7 10.2 9.0 3.5 5.7 7.8 ----------------------------------------------------------------------

The localized and presumably highly inbred populations may have diverged morphologically, in some cases at least, to a degree that mensural differences can be demonstrated even between samples from the same general geographic area. Localized variation and relatively marked secondary sexual variation (unrecognized by some earlier workers), superimposed on geographic variation, have resulted in application of a relatively large number of names to these small _Molossus_. Felten (1957:13-14), for example, apparently used different specific names for males and females from El Salvador, and Gardner (1966) employed three different specific names for North American specimens. Only when material is available for a detailed study of variation throughout the Neotropics can the perplexing mosaic of characters in these small _Molossus_ be assessed adequately.

We have seen no specimens from Nicaragua that are identifiable as _Molossus bondae_, another relatively small species that has been reported from Greytown (Miller, 1913a:89) and from elsewhere in Central America by other authors (Goodwin, 1942c:145; Handley, 1966b:772; Gardner _et al._, 1970:727). Our examination of the female holotype of _M. bondae_ reveals that it is larger than _M. molossus_, corresponding in size to females recently reported from Costa Rica by Gardner _et al._ (_loc. cit._), and that _bondae_ has dark-based hairs. The two males reported by Goodwin (_loc. cit._) from Honduras as _bondae_, would seem to be too small for that species, based on the measurements listed; also, these specimens allegedly have white-based hairs and probably represent _M. molossus_ as here defined.

Molossus pretiosus pretiosus Miller, 1902

_Specimens._--_Boaco_: Los Cocos, 14 km S Boaco, 220 m, 28; San
Francisco, 19 km S, 2 km E Boaco, 200 m, 3. _Carazo_: 3 km N, 4 km
W Diriamba, 600 m, 25. _Managua_: 6 mi WSW Managua, 3.

This relatively large mastiff bat has not been reported previously from Nicaragua. Specimens from several localities in Boaco were captured in mist nets over streams; most of those from northwest of Diriamba were shot in the early evening as they foraged high around large trees in a coffee finca, but several were netted over a water-filled concrete tank or found in the water in the tank as detailed in the account of _Eptesicus furinalis_. At Los Cocos, bats that we netted seemed to be emerging from a hollow located high in a tree over the stream.

Selected measurements of _M. p. pretiosus_ from Nicaragua, which compare favorably with those of topotypes from Venezuela, are listed in Table 4 along with measurements of _M. ater_ and _M. sinaloae_. The taxonomic relationships of _M. pretiosus_ and _M. ater_ are less than clear, and some authors (Handley, 1966b:773, for instance) have suggested that the two may be conspecific. Whatever their ultimate relationships may prove to be, two distinctive taxa seem to be present in Nicaragua; the larger is assignable to _ater_ and the smaller to _pretiosus_, as currently understood. Furthermore, the presumed presence of two large _Molossus_ with dark-based hairs elsewhere in Central America (Dilford C. Carter, personal communication) and in southeastern Mexico (Goodwin, 1956:4; Goodwin and Greenhall, 1964:20) argues for specific recognition of _pretiosus_.

The species _ater_ and _pretiosus_ differ mainly in size (Table 4), some measurements clearly separating the two when sexual dimorphism is considered. Also, the average weights of 18 nonpregnant females and four males of _pretiosus_ (all adults) collected on 20 February 1968 at Los Cocos, were 20.9 (14.6-23.8) and 27.0 (24.6-31.7) gms, respectively, significantly smaller than corresponding figures for 11 nonpregnant females and nine males of _M. a. nigricans_ taken two weeks later on the Cosigueina Peninsula--29.1 (26.1-33.0) and 32.9 (29.3-35.1) gms. It is of note that we have not collected these two large species at the same localities in Nicaragua, and it is possible that one competitively excludes the other in local situations.

TABLE 4.--Selected measurements of adults of three species of _Molossus_ from Nicaragua.

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: Condylobasal length Col. E: Zygomatic breadth Col. F: Breadth of braincase Col. G: Length of maxillary toothrow Col. H: Breadth across upper canines

==================================================================== A B C D E F G H --------------------------------------------------------------------

_Molossus ater nigricans_, Departamento de Chinandega, Nicaragua

Average 10 ([Male]) 49.5 23.2 20.2 14.3 11.0 8.2 6.1 Minimum 48.2 22.8 19.9 14.1 10.5 8.0 6.0 Maximum 50.7 23.5 20.5 14.9 11.2 8.3 6.3 Average 10 ([Female]) 49.0 22.0 19.2 13.6 10.7 7.8 5.8 Minimum 47.1 21.5 18.8 13.3 10.5 7.6 5.6 Maximum 51.2 22.5 20.0 14.1 11.0 8.2 6.1

_Molossus pretiosus pretiosus_, Departamento de Boaco, Nicaragua

Average 8 ([Male]) 45.1 21.7 18.8 13.4 10.6 7.6 5.9 Minimum 44.1 21.0 18.3 13.1 10.2 7.4 5.6 Maximum 46.1 22.2 19.1 13.7 10.9 7.9 6.1 Average 23 ([Female]) 43.6 20.2 17.7 12.5 10.1 7.3 5.2 Minimum 41.0 19.5 17.2 12.2 9.5 7.1 5.0 Maximum 44.8 20.7 18.2 13.1 10.4 7.6 5.6

6 mi SW Managua, Nicaragua

KU 70135, [Male] 44.8 21.7 18.6 13.0 10.2 7.7 5.8 KU 70137, [Male] 45.1 21.6 19.0 13.2 10.7 7.7 5.8 KU 70136, [Female] 43.5 19.9 17.8 12.7 10.3 7.5 5.5

3 km N, 4 km W Diriamba, Nicaragua

Average 7 ([Male]) 44.8 21.2 18.5 13.0 10.5 7.4 5.5 Minimum 43.8 20.7 18.3 12.7 10.2 7.2 5.4 Maximum 45.7 21.3 18.8 13.3 10.9 7.6 5.5 Average 9 ([Female]) 44.0 20.7 18.0 12.6 10.2 7.3 5.3 Minimum 42.0 20.0 17.4 12.0 10.0 7.0 5.0 Maximum 45.5 21.2 18.6 13.0 10.6 7.5 5.5

_Molossus sinaloae sinaloae_, El Recreo, Nicaragua

Average 4 ([Male]) 47.6 21.2 18.6 12.3 9.8 7.7 5.4 Minimum 46.7 20.7 18.3 12.0 9.7 7.6 5.3 Maximum 48.6 21.5 18.9 12.6 10.0 7.9 5.5 Average 10 ([Female]) 47.5 20.2 17.8 11.8 9.7 7.3 5.1 Minimum 46.2 20.0 17.6 11.5 9.5 7.1 5.0 Maximum 48.3 20.4 18.0 12.0 9.8 7.5 5.5 --------------------------------------------------------------------

Among females from Departamento de Boaco, none of 18 collected on 20 February was pregnant, whereas three of four obtained on 4 April were gravid (embryos 13, 15, and 17 mm in crown-rump length) as was one (embryo 21 mm) taken on 18 July. Among those from northwest of Diriamba, two of three were pregnant on 30-31 March (embryos 7 and 13 mm in length), whereas only three of eight carried embryos (29, 31, and 34 mm) in mid-August at a time when flying young also were collected. Additionally, a female captured 6 mi WSW Managua on 27 March had an embryo that measured 11 mm.

Four adult males collected on 20 February had an average testicular length of 5.7 (4-7) mm, whereas those of seven obtained in late March averaged 5.0 (4-6) mm; testes of two adults taken on 18 July were 6 and 6.5 mm, and those of three taken in the period 12-15 August were 5 mm in length.

Molossus sinaloae sinaloae J. A. Allen, 1906

_Specimens._--_Boaco_: Santa Rosa, 17 km N, 15 km E Boaco, 300 m,
1. _Managua_: 1 km N Sabana Grande, 1; 3 mi SW Managua, 1; 8 km SW
Managua, 400 m, 3. _Rivas_: San Juan del Sur, 20 m, 1. _Zelaya_: S
side Rio Mico, El Recreo, 25 m, 73.

This mastiff bat can be distinguished easily from the other two large species of _Molossus_ in Nicaragua in that the hairs of the dorsum are relatively long and are pale, frequently white, basally. The species has been reported twice previously from Nicaragua--from Greytown by Miller (1913a:89) and by Goodwin and Greenhall (1964:13), who listed four individuals from Rio Grande.

Most of our specimens were taken from buildings. A large series obtained in a house at El Recreo in late June 1966 consisted only of females and young, many of which were nearly full grown; some females still were lactating and at least two were pregnant (on 23 June--crown-rump length of embryos 12 and 14 mm). A group taken from another house at El Recreo (from cracks in the stone wall and from between rafters and the corrugated metal roof) in late July 1966 consisted of adult males, lactating females, and young. Similarly, an adult male, a pregnant female (embryo 14 mm in crown-rump length), and a hairless juvenile were shot, on 16 February 1964, from a small colony in an attic of a large house southwest of Managua; _Glossophaga soricina_ also was taken in this attic. The testes of an adult male captured in June were 6 mm in length; those of four taken in July were 5, 5, 6, and 7 mm, and one obtained in August had testes that measured 3 mm.

Selected measurements of _M. s. sinaloae_ from El Recreo are given in Table 4.

CHECKLIST OF NICARAGUAN BATS

Following is a checklist of the 68 species of bats thus far reported from Nicaragua. Citations are given only to publications that are primary sources of information on Nicaraguan specimens. In some instances, specimens were reported in the literature under a name different than the one here employed; we have mentioned earlier-used names where it seemed appropriate to do so. Taxa preceded by an asterisk are included in the present paper.

Family EMBALLONURIDAE

_Rhynchonycteris naso_ (Wied-Neuwied, 1820).--J. A. Allen (1908:669, 1910:110), Sanborn (1937:327), Davis _et al._ (1964:375, 379).

_Saccopteryx bilineata_ (Temminck, 1838-1839).--J. A. Allen (1910:110), Sanborn (1937:331), Jones (1964a:506).

*_Saccopteryx leptura_ (Schreber, 1774).--Davis _et al._ (1964:374), Jones (1964a:506).

_Cormura brevirostris_ (Wagner, 1843).--J. A. Allen (1910:110--specimens listed as _Peropteryx canina_ from Pena Blanca, see Sanborn, 1937:348), Miller (1924:38), Sanborn (1937:349), Goodwin (1942c:119).

_Peropteryx kappleri kappleri_ Peters, 1867.--J. A. Allen (1908:669, 1910:110--as _P. canina_), Sanborn (1937:344).

*_Peropteryx macrotis macrotis_ (Wagner, 1843).

_Balantiopteryx plicata plicata_ Peters, 1867.--Jones _et al._ (1962:152), Davis _et al._ (1964:375), Jones (1964a:506).

_Diclidurus virgo_ Thomas, 1903.--Alston (1879-82:30).

Family NOCTILIONIDAE

*_Noctilio labialis labialis_ (Kerr, 1792).--Goodwin (1942c:121, 1946:297), Davis _et al._ (1964:376), Kohls _et al._ (1965:344), Wenzel _et al._ (1966:578).

*_Noctilio leporinus mexicanus_ Goldman, 1915.--Davis _et al._ (1964:376).

Family PHYLLOSTOMATIDAE

*_Pteronotus davyi fulvus_ (Thomas, 1892).

*_Pteronotus parnellii fuscus_ J. A. Allen, 1911.--Miller (1902:402), Rehn (1904:203).

*_Pteronotus suapurensis_ (J. A. Allen, 1904).--Goodwin (1942a:88, 1946:299, 1953:246).

_Micronycteris brachyotis_ (Dobson, 1879).--Goodwin (1946:302--as _M. sylvestris_), Sanborn (1949:224--as _M. platyceps_), Goodwin and Greenhall (1961:231).

_Micronycteris hirsuta_ (Peters, 1869).--Valdez and LaVal (1971:247).

*_Micronycteris megalotis mexicana_ Miller, 1898.--G. M. Allen (1929:130).

*_Micronycteris megalotis microtis_ Miller, 1898.--Miller (1898:328, 1900:154), Lyon and Osgood (1909:263), J. A. Allen (1910:110), Poole and Schantz (1942:133).

_Micronycteris minuta_ (Gervais, 1856).--Valdez and LaVal (1971:248).

_Micronycteris schmidtorum_ Sanborn, 1935.--Davis _et al._ (1964:378).

_Macrophyllum macrophyllum_ (Schinz, 1821).--Davis _et al._ (1964:378), Jones (1964a:506).

*_Tonatia nicaraguae_ Goodwin, 1942.--Goodwin (1942b:205, 1946:305, 1953:246), Davis and Carter (1962:67), Valdez and LaVal (1971:248).

_Phyllostomus discolor verrucosus_ Elliot, 1905.--Jones (1964a:507), Wenzel _et al._ (1966:598).

*_Phyllostomus hastatus panamensis_ J. A. Allen, 1904.--Goodwin (1942c:126), Wenzel _et al._ (1966:597).

*_Trachops cirrhosus coffini_ Goldman, 1925.--Carter _et al._ (1966:491).

*_Chrotopterus auritus auritus_ (Peters, 1856).

_Vampyrum spectrum nelsoni_ (Goldman, 1917).--Gray (1844:18), Dobson (1878:471), Alston (1879-82:39), J. A. Allen (1910:111), Goldman (1917:116), Goodwin (1946:310).

_Glossophaga commissarisi_ Gardner, 1962.--Davis _et al._ (1964:380), Jones (1964a:507).

_Glossophaga soricina leachii_ (Gray, 1844).--Gray (1844:18), Dobson (1878:501), Alston (1879-82:44), J. A. Allen (1910:111), Miller (1913b:419), Alvarez (1963:400), Davis _et al._ (1964:380), Starrett and de la Torre (1964:57).

*_Anoura geoffroyi lasiopyga_ (Peters, 1868).

*_Choeroniscus godmani_ (Thomas, 1903).--Handley (1966a:86).

*_Lichonycteris obscura_ Thomas, 1895.--Thomas (1895:57), Davis _et al._ (1964:380).

*_Carollia castanea_ H. Allen, 1890.---Davis _et al._ (1964:379).

_Carollia perspicillata azteca_ Saussure, 1860.--Hahn (1907:112), J. A. Allen (1908:669, 1910:111), Davis _et al._ (1964:379), Starrett and de la Torre (1964:58).

_Carollia subrufa_ (Hahn, 1905).--Davis and Carter (1962:71), Davis _et al._ (1964:379).

_Sturnira lilium parvidens_ Goldman, 1917.--Davis _et al._ (1964:379), Starrett and de la Torre (1964:59), Jones (1964a:507).

*_Sturnira ludovici ludovici_ Anthony, 1924.

_Uroderma bilobatum convexum_ Lyon, 1902.--Davis _et al._ (1964:382), Jones (1964a:507), Davis (1968:695).

_Uroderma bilobatum molaris_ Davis, 1968.--Davis _et al._ (1964:382), Davis (1968:697).

*_Uroderma magnirostrum_ Davis, 1968.--Davis (1968:680).

*_Vampyrops helleri_ Peters, 1866.--Davis _et al._ (1964:383), Jones (1964a:507), Valdez and LaVal (1971:248).

*_Vampyrodes major_ G. M. Allen, 1908.--J. A. Allen (1910:112), Goodwin (1946:320).

*_Vampyressa nymphaea_ Thomas, 1909.

*_Vampyressa pusilla thyone_ Thomas, 1909.--Starrett and de la Torre (1964:60).

*_Chiroderma villosum jesupi_ J. A. Allen, 1900.

_Ectophylla alba_ H. Allen, 1892.--H. Allen (1892:441, 1898:267), Lyon and Osgood (1909:266), Poole and Schantz (1942:129), Goodwin (1942c:134, 1946:323), Goodwin and Greenhall (1962:4). Type locality (Rio Segovia) placed in Comarca de El Cabo, Nicaragua, by Miller and Kellogg (1955:77).

_Artibeus inopinatus_ Davis and Carter, 1964.--Davis and Carter (1964:120).

_Artibeus jamaicensis paulus_ Davis, 1970.--Andersen (1908:266), Davis (1970b:121).

_Artibeus jamaicensis richardsoni_ J. A. Allen, 1908.--Andersen (1908:267), J. A. Allen (1908:669), Elliot (1917:168), Goodwin (1953:249), Davis and Carter (1964:120), Davis _et al._ (1964:379), Starrett and de la Torre (1964:60), Davis (1970b:116).

_Artibeus lituratus palmarum_ J. A. Allen and Chapman, 1897.--Andersen (1908:279), Davis and Carter (1964:120), Davis _et al._ (1964:379), Starrett and de la Torre (1964:61).

_Artibeus phaeotis palatinus_ Davis, 1970.--Davis (1970a:401).

_Artibeus phaeotis phaeotis_ (Miller, 1902).--Davis _et al._ (1964:379), Davis (1970a: 399).

*_Artibeus toltecus hesperus_ Davis, 1969.

*_Artibeus toltecus toltecus_ (Saussure, 1860).--Andersen (1908:300), Davis (1969:28).

*_Artibeus watsoni_ Thomas, 1901.--Andersen (1908:290), Davis and Carter (1962:71), Davis (1970a:393).

*_Centurio senex senex_ Gray, 1842.--Goodwin (1946:327, restricted type locality to El Realejo, Nicaragua).

_Desmodus rotundus murinus_ Wagner, 1840.--J. A. Allen (1908:670, 1910:113).

*_Diphylla ecaudata_ Spix, 1823.

Family NATALIDAE

*_Natalus stramineus saturatus_ Dalquest and Hall, 1949.

Family THYROPTERIDAE

_Thyroptera discifera discifera_ (Lichtenstein and Peters, 1854).--Miller (1896:111, 1931:411), Goodwin (1946:330).

Family VESPERTILIONIDAE

*_Myotis albescens_ (E. Geoffroy St.-Hilaire, 1806).--Miller and G. M. Allen (1928:203).

*_Myotis elegans_ Hall, 1962.

*_Myotis nigricans nigricans_ (Schinz, 1821).--Davis _et al._ (1964:379), McDaniel and Coffman (1970:223).

*_Myotis simus riparius_ Handley, 1960.

*_Eptesicus furinalis gaumeri_ (J. A. Allen, 1897).--Miller (1897:100), Davis (1965:234).

_Lasiurus borealis frantzii_ (Peters, 1871).--Davis and Carter (1962:73).

*_Rhogeessa tumida tumida_ H. Allen, 1866.--J. A. Allen (1910:113), Goodwin (1958:3).

Family MOLOSSIDAE

*_Tadarida laticaudata yucatanica_ (Miller, 1902).

*_Eumops auripendulus_ (Shaw, 1800).

*_Molossus ater nigricans_ Miller, 1902.--J. A. Allen (1908:670).

_Molossus bondae_ J. A. Allen, 1904.--Miller (1913a:89).

*_Molossus molossus aztecus_ Saussure, 1860.--Felten (1957:14).

*_Molossus pretiosus pretiosus_ Miller, 1902.

*_Molossus sinaloae sinaloae_ J. A. Allen, 1906.--Miller (1913a:89), Goodwin and Greenhall (1964:13).

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