Natalus primus

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 : 9-22

Natalus primus Anthony, 1919

Figures 1–3

Natalus major primus Goodwin, 1959 .

Natalus stramineus primus Linares, 1971 .

HOLOTYPE: AMNH 41009 , a fossil right dentary, collected by H. E. Anthony in 1917, Cueva de Los Indios (locality 29 in the appendix), Daiquirı´, Santiago de Cuba, Cuba. The holotype is missing the coronoid process plus the incisors, canine, and first premolar, and is stained dark brown. (A second right dentary, designated by Anthony as a topo­ type, is in the vial with the holotype. It is complete, but is missing all teeth except the third premolar and the first molar).

DISTRIBUTION: Cuba, Isle of Pines, and the Bahamas (Abaco, Andros, New Providence). There is only one known extant locality (Cueva La Barca, Cuba; Tejedor et al., 2004) and 14 fossil localities (fig. 5).

DIAGNOSIS: Largest extant species in the genus Natalus (forearm length 46.1–51.2 mm, greatest skull length 18.1–19.9 mm); ear tip rounded, notch on outer margin of ear almost imperceptible; rostrum longer than in any other Greater Antillean Natalus , point of flexion between rostrum and braincase posterior to anterior edge of orbit; braincase not rising abruptly from rostrum, forming an angle smaller than 558 with dorsal plane of rostrum; braincase oval­shaped in dorsal profile,

breadth smaller than length; postorbital constriction relatively wide, its sides markedly diverging anteriorly; postorbital sphenoid fenestra present; maxillary convex above molars; two pairs of relatively large fenestra on palate, between molars; basisphenoid pits deeply concave; posterior edge of ascending ramus of mandible forming an angle smaller than 708 with basal plane of dentary, and often with a notch below condyloid process ( table 1 ).

DESCRIPTION: Natalus primus is the largest of the Greater Antillean Natalus in mean forearm length, greatest skull length, and breadth across molars (P, 0.001; table 2 , fig. 6). Its mean braincase breadth, however, is relatively small compared to those of N. jamaicensis and N. major (P, 0.001; fig. 6). The ears of N. primus are very large (20.2–21.2 mm). Males of N. primus are significantly, although only slightly, larger than females in body weight, length of tibia, greatest skull length, and zygomatic breadth (P, 0.01; table 2 ; fig. 6).

The hair is long and lax. Dorsal and ventral hairs are 8 mm long. The dorsal hairs are bicolored, with bases lighter than tips, and the ventral hair is monocolored. There are two pelage­color morphs, one bright yellowish and another grayish, with some individuals showing intermediate coloration. In the bright­yellowish phase the bases of the dorsal hairs are buff and the tips are sepia, and the ventral hair is entirely cream buff. In the grayish phase the dorsal hairs are drab to smoke gray with fuscous tips and the ventral hairs are pale smoke gray. There is a fringe of hairs along the free border of uropatagium. Natalus primus has mustachelike facial hair tufts that are sepia to fuscous in col­ or, and are not connected across the dorsum of the muzzle by a 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 long and dorsoventrally flattened. The small, oval nostrils open ventrolaterally. The tip of the muzzle, between nostrils, projects slightly beyond upper lip, a feature that is most noticeable in ventral view. The lower lip is thickened and has a shallow central groove. The ears are rather squareshaped, with a straight anterior margin and lacking a notch on the posterior margin, giving the tip a relatively round appearance. The ears of Natalus primus show rudimentary folds, varying in number from five to seven.

The braincase of Natalus primus is moderately inflated, and does not rise abruptly from the rostrum, forming an angle smaller than 558 with the dorsal plane of the nasals. The sagittal crest is pronounced and its interparietal region is slightly higher than its supraoccipital region. The postparietal region is relatively narrow in dorsal view and the ridge formed between this region and the occipital is relatively low. The postorbital constriction is relatively wide, with sides markedly diverging anteriorly. The rostrum is very long and moderately broad. The sulcus between the nasals is long and deep. The premaxillary region projects anteriorly, so that in lateral view the diastema between canines and incisors is conspicuous. The maxillary is convex dorsal to the molars. The rostrum tips slightly downward. The infraorbital foramen opens laterally, close to the alveolar margin. The anterior palatal foramina are very small and near the palatine emargination, at the level of the anterior edge of the canines. There are two relatively large palatal fenestra at the level of the last two molars. The bony palate extends for less than half the distance between posterior edge of last molars and tip of pterygoids. The pterygoid processes are convergent posteriorly, and vary from broadly triangular to acutely pointed in lateral view. The inner edge of each pterygoid has a prominent right­angled point visible in ventral view. The basisphenoid pits are deep, forming a large, double concavity that is conspicuously deeper than the basisphenoid furrows between the auditory bullae.

The dentary bone is thin and not markedly curved. A rudimentary mandibular angle is sometimes present. The angular process is long and thin, rather straight, and with a hooked end. The base of the angular process is at the level of the alveolar plane. The coronoid process is at about the same height or slightly higher than the condyle above the alveolar plane. The ascending ramus is not markedly upturned. The anterior border of the ascending ramus is slightly concave, rising from the alveolar plane in an angle of about 808. The posterior border of the ascending ramus rises from the base of the angular process in an angle smaller than 708. Usually the posterior edge of the ascending ramus is not straight, having a shallow notch immediately above the base of the angular process. The mental foramen is between the canine and the first premolar.

The superior incisors are relatively long and slightly hooked in lateral view. The first incisor is situated more anteriorly than the second, so that in lateral view the first incisor is easily visible. In occlusal view, the base of the canine is slender, its width comprising about two thirds of its length. The cingulum of the canine is well developed. The premolars are successively larger, in occlusal view, and relatively long and not crowded. The 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 inferior incisors are small, tricuspid, and subequal. The canine is relatively small and straight. In occlusal view, the canine and premolars are longitudinally elongate, with the first premolar longer than the canine and about as long as the other two premolars. The molars are approximately equal in size and shape, with the first more projected labially than the other two. The last premolar and the first molar are not markedly crowded (fig. 3).

COMPARISONS: Natalus primus is the largest of all extant Natalus , and overlaps in range of forearm length only with Natalus jamaicensis (mean forearm length, however, is significantly different between the two species; P, 0.001; table 2 ). Natalus primus , therefore, can be distinguished by size alone from most other species of the genus.

Externally N. primus can be identified by its rather rounded ear tip that lacks a notch or emargination along the posterior margin of the ear, whereas in all other species of Natalus the ear tip is markedly pointed and the lateral ear margin has a notch below the ear tip. Similarly, N. primus lacks the band of dark hairs that in all other species of Natalus connects the labial mustachelike hair tufts characteristic of the genus.

Cranially, Natalus primus is unlike any other species of Natalus in that its basisphenoid pits are very deep and steep­sided, whereas in the remaining species of the genus the basisphenoid pits are shallow. Also, in N. primus , the rostrum appears proportionately longer, relative to skull length, than in all other species of Natalus . This greater length of the rostrum in N. primus is concomitant with (1) the elongation of the premaxillary region, resulting in an anterior projection of the incisors, and (2) the position of the dorsal point of flexion of the skull, which, in lateral view, lies posterior to the anterior edge of the orbit. In all other species of Natalus , the premaxilla is not markedly elongated, so that the incisors are at or near the level of the canines, and the dorsal point of flexion of the skull, in lateral view, lies dorsal to the anterior edge of the orbit. In N. primus , the posterior edge of the ascending ramus of the mandible is not perpendicular to the basal plane of the dentary, and usually shows a distinctive notch, absent in all other species of Natalus , that is dorsal to the angular process.

REMARKS: Fossil Natalus collected in the western Bahamas were referred to N. major primus by Morgan (1989), but were reported by him to be noticeably larger than those of Cuba. Although it is possible that the Bahamian fossils represent a species distinct from N. primus , we refrain from excluding them from the Cuban taxon until specimens from both island groups are compared in greater detail.

Natalus primus

THE BAHAMAS

Abaco

15. Hole in the Wall Cave, (fossils), Morgan (2001).

Andros

16. Coleby Bay Cave, Morgans Bluff (258109N, 788029W). FLMNH 79324 (fossil), Morgan (1989).

17. King Cave, Morgans Bluff (258109N, 788029W). FLMNH 79110 (fossil), Morgan (1989).

New Providence

18. Banana Hole (258019N, 778339W). FLMNH 79986–79990 (fossils), Morgan (1989).

19. Hunts Cave (258029N, 778229W). FLMNH 27694, 79751–79759, 79761–79766, 79843– 79848 (fossils), Morgan (1989).

CUBA

Camagüey

20. Cueva de los Portales de Pinto, Jaronú (218489N, 778579W), Esmeralda. MCZ (fossil), Koopman and Ruibal (1955).

21. Cueva del Círculo, Cairije (218369N, 778409W), Tomás Betancourt. IES (fossil), Silva­Taboada (1979).

Cienfuegos

22. Cueva de la Macha [5 Cueva de los Machos, Cueva de Vilches], Cantera de San Antón, Soledad (228079N, 808199W). AMNH (fossil), Goodwin (1959).

Isla de la Juventud

23. Cueva del Abuelo, Sierra de Caballos (218529N, 828469W). IES (fossil), Silva­Taboada (1979).

Matanzas

24. Cueva de la Eloísa, Bellamar (238019N, 818339W), Guanábana. IES (fossil), Silva­Taboada (1979).

Pinar del Rio

25. Cueva La Barca, 20 km E of Cabo de San Antonio (218509N, 848569W), Guanahacabibes Peninsula, Manuel Lazo. MNHN 1–26 G. Sil­ va­Taboada field numbers (male), 27–51 G. Silva­Taboada field numbers (female), Tejedor et al. (2004). FLMNH 26810 (female).

Sancti Spiritus

26. Lomas de Judas (22869N, 788279W), Yaguajay. IES (fossil), Silva­Taboada (1974). ISEZ (fossil), Wolozyn and Silva­Taboada (1977).

27. Sistema Cavernario Masones­Jagüey, Trinidad (218489N, 798599W). IES (225 fossil specimens), Silva­Taboada (1974); ROM 59133– 59135 (fossils).

Santiago de Cuba

28. Cueva de la Cantera, Siboney (198579N, 758429W), Damayajabo. IES (fossil), Silva­Taboada (1979).

29. Cueva de Los Indios, Daiquirí (198559N, 758399W), AMNH 41009 (fossil, holotype), Anthony (1919).

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.