Natalus major Miller, 1902

TEJEDOR, ADRIAN, TAVARES, VALERIA DA C. & SILVA-TABOADA, GILBERTO, 2005, A Revision of Extant Greater Antillean Bats of the Genus Natalus, American Museum Novitates 3493 (1), pp. 1-23 : 4-22

Natalus major Miller, 1902

Figures 1–4

Natalus major major Goodwin, 1959 .

Natalus stramineus major Linares, 1971 .

HOLOTYPE: USNM 101395 , adult male skull and skin in fluid, collected by W. M. Gabb between 1869 and 1871 ‘‘near Savaneta [locality 10 in the appendix], Dominican Republic’ ’. The skull is complete and the skin is in good condition.

DISTRIBUTION: Hispaniola, including the Dominican Republic and Haiti (fig. 5).

DIAGNOSIS: Forearm length relatively small (42.1–44.8 mm); ear tip pointed, with a shallow notch on the outer margin; rostrum relatively short, with the point of flexion between rostrum and braincase dorsal to the anterior edge of orbit; braincase not rising abruptly from the rostrum, with an angle smaller than 558 between dorsal plane of rostrum and frontal plane of forehead; braincase oval­shaped in dorsal profile, its breadth smaller than its length; postorbital constriction relatively wide, its sides markedly diverging anteriorly; maxillary convex dorsal to molars; one pair of small palatal fenestra sometimes present between last molars; basisphenoid pits shallow, and raised above plane of basisphenoid furrows, in ventral view; posterior edge of ascending ramus of mandible straight and vertical, forming an angle close to 908 with basal plane of dentary, and lacking notch below condyloid process ( table 1 ).

DESCRIPTION: Natalus major has a relatively short forearm (42.0– 44.8 mm) and a relatively short ear (ear length 15.8–17.9 mm). The braincase breadth and greater skull length means of males of N. major appear to average larger than those of females (fig. 6), but these differences were not statistically significant in our sample.

The hair is long (7 mm) and lax, both ventrally and dorsally. Dorsal hairs are bicolored, with bases lighter than tips, and ventral hairs are monocolored. There are two morphs of pelage color, one bright yellowish and another grayish, with a few intermediate individuals. In the bright­yellowish morph, dor­ sal hair bases are buff and the tips are sepia, and ventral hairs are entirely cream buff (fig. 4). In the grayish morph, the dorsal hairs are drab to smoke gray with fuscous tips, and ventral hairs are pale smoke gray. Most individuals have a darker­colored patch on the forehead. There is a fringe of hairs along the free border of the uropatagium. Mustachelike facial hair is sepia to fuscous, connected across dorsum of muzzle by a narrow band of dark hairs. The pelage extends slightly into both faces of the plagiopatagium, dorsally to 6 mm, and ventrally to 10 mm.

The muzzle is relatively short and dorsoventrally flattened. The nostrils are small, oval, and open ventrolaterally. In ventral view, the tip of the muzzle between the nostrils projects slightly beyond the upper lip. The lower lip is thickened and with a shallow central groove. The ears are rather squareshaped. The inner edge of the ear is straight below the tip and has a shallow notch on the outer edge, giving the tip a pointed appearance. The ears of N. major show five to seven rudimentary folds, though they may be entirely absent in some individuals.

The skull is relatively broad and robust. The braincase of Natalus major is not markedly inflated, nor very abruptly elevated above the rostrum, forming an angle smaller than 558 with the dorsal plane of the nasals. The postparietal region is relatively wide in dorsal view. A conspicuous ridge separates the postparietal and occipital regions. The sagittal crest is well developed. The postorbital region is wide, with sides markedly diverging anteriorly, in dorsal view. The rostrum is relatively short and broad. The dorsal point of flexion between the rostrum and the braincase is dorsal to the anterior edge of the orbit. There is a moderately deep sulcus between the nasal bones. The premaxillary region does not project anteriorly, so that in lateral view the small diastema between canines and incisors is not particularly noticeable. The maxillary is convex above molars. The rostrum does not tip downward, or does so only slightly. The anterior palatal foramina are very small and near the palatine emargination, and their anterior margin is posterior to the anterior edge of the canines. Two pairs of small palatal fenestra are present at the level of the last molar in some individuals. The bony palate extends more than half the distance between the posterior edge of the last molar and the tip of pterygoids. The pterygoid processes are slightly convergent and markedly hooked in lateral view. In ventral view, the inner edge of each pterygoid is smoothly convex. In ventral view, the basisphenoid pits, between the maxillary articular facets, are shallow and raised above the level of the basisphenoid furrows.

The dentary is curved (upturned) and thick. The angular process is thick, rather straight, and with a blunt spatulate end. The base of the angular process is dorsal to the alveolar plane. The coronoid process is at about the same height than the condyle, above the alveolar plane. The ascending ramus of the mandible is markedly upturned, with its anterior border concave and raising from the alveolar plane at an angle greater than 808. The posterior border of the ascending ramus rises from the base of the angular process at a nearly straight angle with the base of the dentary, giving a rectangular appearance to the ascending ramus. The mental foramen is between the canine and the first premolar.

The upper incisors of Natalus major are small, with a hooked point in lateral view. The first incisor is situated more posteriorly than the second, so that in lateral view the first incisor is not visible. The base of the upper canines is robust, in occlusal view, with a well­developed cingulum that is about as wide as it is long. The upper premolars are successively larger, in occlusal view, and relatively short and crowded. The upper molar row is relatively slender, its width less than half the width of the palate. The first and second molars are approximately equal in size, with the third slightly smaller.

The lower incisors are small, tricuspid, and subequal. The lower canine is large and rather curved, with a large cingulum that is slightly longer than it is wide. The canine and premolars are longitudinally short in occlusal view, and the first premolar is shorter than the canine and the other two premolars. The first two molars are larger and wider than the third; the first projects more labially than the other two. The last premolar and first molar are relatively crowded (fig. 3).

COMPARISONS: Natalus major can be distinguished from most other species of Natalus (with the exception of Natalus jamaicensis ) on the basis of body size and external characters. The forearm length of N. major (42.0–45.0 mm) is smaller than that of Natalus primus (46.1–51.2 mm; P, 0.001; fig. 6), and is larger than those of Natalus stramineus (35.0–41.0 mm; Tejedor et. al., 2004) and Natalus tumidirostris (37.2–41.5 mm; Tejedor, unpublished data). Externally, the ear of N. major has a straight anterior margin and a pointed tip, whereas that of N. primus has a relatively rounded tip and those of N. stramineus and N. tumidirostris have slightly to deeply concave anterior margins. The dorsum of the muzzle of N. major shows a thin band of dark, long hairs connecting the labial mustaches, which is absent in N. primus .

Natalus major is morphometrically distinct from Natalus jamaicensis (figs. 6, 7), but both species overlap in size and are similar in external characters. Nonetheless, N. major can be unambiguously distinguished from N. jamaicensis on the basis of cranial characters ( table 1 , figs. 2–3). In N. major the maxillary bone is convex dorsal to the molars, and in N. jamaicensis it is markedly concave. In N. major , the sulcus between the nasal bones is moderately deep and long, whereas it is short and shallow in N. jamaicensis . In N. major , the upper molar series is narrower than half of the palate width, whereas in N. jamaicensis it is thicker than half of the palate’s width. In N. major , the postorbital constriction is wider than in N. jamaicensis (P, 0.001, fig. 6), with sides markedly diverging anteriorly, in dorsal view, as opposed to sides almost parallel in N. jamaicensis . Also, in N. major , the braincase is much less inflated than in N. jamaicensis , and it is oval­shaped in dorsal profile, instead of almost circular as in N. jamaicensis . Finally, in N. major , the braincase does not rise very abruptly from the rostrum (angle between dorsal plane of rostrum and frontal plane of braincase less than 558), whereas it rises very abruptly from the rostrum in N. jamaicensis (angle between dorsal plane of rostrum and frontal plane of braincase greater than 608).

Cranially, Natalus major can also be distinguished from Natalus primus , by its small­ er skull size (greatest skull length 17.0– 18.1 mm; P, 0.001), and shallow basisphenoid pits, whereas in the latter the basisphenoid pits are very deep and the skull is larger (greatest skull length 18.1–19.9 mm; P, 0.001; table 2 , figs. 2–3). From Natalus tumidirostris , N. major can be distinguished by its convex yet uninflated maxillary bones, which are markedly inflated in the former, so that the molars are not easily seen in dorsal view. Natalus major differs from Natalus stramineus , as it is currently defined, by its larger skull (greatest skull length: 17.0– 18.1 mm, 15.0– 17.1 mm in N. stramineus ) and by a larger and more voluminous braincase, relative to the overall size of the skull.

REMARKS: Fossil Natalus collected in the eastern Bahamas (Grand Caicos) were referred to N. major major by Morgan (1989). We tentatively include these fossils within N. major until they can be reexamined.

Natalus major

DOMINICAN REPUBLIC

Barahona

3. Los Patos (178589N, 718109W). AMNH 97590 (male).

4. Maniel Viejo (178589N, 718199W, 278 m). AMNH 97589 (female).

Elias Piña

5. Cerro de San Francisco, Bánica (198049N, 718429W). FLMNH (fossil), Morgan (2001).

Independencia

6. Puerto Escondido (188199N, 718349W), 9.9 km S, North slope Sierra de Bahoruco. USNM 542274 (female, mummy).

María Trinidad Sánchez

7. Cueva de Murciélagos, Entrada (198339N, 698549W), Cabrera. AMNH 238148, 238149 (2 females); (AMNH, 4 males, 1 female) AT 44, 45, 49, 51, 52 (field numbers).

Montecristi

8. Pepillo Salcedo (198429N, 718459W), 5 km SE. KU 150713–20 , KU 152361 (7 males, 2 females), Timm and Genoways (2003) .

Sánchez Ramírez

9. Cueva Grande de Julián, Don Miguel, 4 km E Platanal (198079N, 708059W). (AMNH, field numbers) AT 70 (female), AT 77 (male).

Santiago

10. Savaneta (possibly Sabaneta [198409N, 708229W] ‘‘near the north coast’’; see Timm and Genoways 2003). USNM 101395a (male, holotype), 101395b (female, paratype), Miller (1902).

unknown locality FLMNH 5517 (male); USNM 49362, 96496, Goodwin (1959).

HAITI

Departement du L’Ouest

11. Port­au­Prince (188349N, 728179W). BM (1 specimen, unspecified), Sanborn (1941).

Departement du Sud

12. Camp Perrin (188199N, 738529W, 246 m), Les Cayes. KU 150721 (male), Timm and Genoways (2003).

TURKS AND CAICOS

Grand Caicos

13. Conch Bar Cave. FLMNH 246 (fossil), Morgan (1989).

Natalus cf. N. major

CAYMAN ISLANDS

Grand Cayman

14. Bodden Cave, Bodden Town (198169N, 818159W). FLMNH (fossil), Morgan (1994).

Figures

Fig. 1. Lateral view of the skull of three species of Greater Antillean bats of the genus Natalus: A, N. primus (Cuba; MNHN 51, field number); B, N. major (Hispaniola; AMNH AT 51, field number); C, N. jamaicensis (Jamaica; AMNH 246127).

Fig. 2. Dorsal and ventral views of the skull of three species of Greater Antillean Natalus. A, Natalus primus (MNHN 51, field number); B, Natalus major (AMNH AT 51, field number); C, Natalus jamaicensis (AMNH 246127).

Fig. 3. Dorsal view of the right mandibular ramus of three species of Greater Antillean Natalus. A, Natalus primus, B, Natalus major, C, Natalus jamaicensis.

Fig. 4. Photograph of an adult male Natalus major (showing a bright yellowish pelage) captured in Cueva Grande de Julián, Sánchez Ramírez province, Dominican Republic, in January 2002. Photograph by A. Tejedor.