Cabassous Centralis (Cingulata: Dasypodidae)

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45(898):12–17 Cabassous centralis (Cingulata: Dasypodidae)

VIRGINIA HAYSSEN,JORGE ORTEGA,ALBERTO MORALES-LEYVA, AND NORBERTO MARTI´NEZ-MENDEZ

Department of Biology, Smith College, Northampton, MA 01063, USA; [email protected] (VH) Laboratorio de IctiologıáyLimnologıá, Posgrado en Ciencias Quimicobiolo´gicas, Departamento de Zoologıá, Escuela Nacional de Ciencias Biolo´gicas, Instituto Politećnico Nacional, Prolongacioń de Carpio y Plan de Ayala s/n, Col. Sto. Tomas, 11340, Me´xico, Distrito Federal, Me´xico; [email protected] (JO), [email protected] (AM-L), [email protected]. mx (NM-M)

Abstract: Cabassous centralis (northern naked-tailed armadillo) is a small armadillo with a slender tail. The distinctive tail has widely spaced, thin plates that are gray-pink in color. This fossorial armadillo occurs in diverse tropical habitats and has a distributional range from southern Mexico, through Central America, and into northern South America (Colombia, Venezuela, and Ecuador). C. centralis is listed as ‘‘Least Concern’’ by the International Union for Conservation of Nature and Natural Resources, and is rare everywhere.

Key words: Edentata, edentate, Neotropical, northern naked-tailed armadillo, Xenarthra

Ó 30 May 2013 American Society of Mammalogists Synonymy completed 11 May 2012 DOI: 10.1644/898.1 www.mammalogy.org

Cabassous centralis (Miller, 1899) Cabassous hispidus: Neveu-Lemaire and Grandidier, Northern Naked-tailed Armadillo 1911:106, footnote. Not Dasypus hispidus Burmeister, 1854. Dasypus gymnurus: Frantzius, 1869:309. Not Tatus gymnu- [Cabassous] lugubris: Yepes, 1928:467 [page 7 of reprint rus Olfers, 1818. separate]. Part; not Ziphila lugubris Gray, 1873. X[enurus]. hispidus: True, 1896:345. Not Dasypus hispidus Cabassous lugubris: Moeller, 1968:420. Part; not Ziphila Burmeister, 1854. lugubris Gray, 1873. Dasypus [Xenurus] hispidus True, 1896:346. Not Dasypus hispidus Burmeister, 1854. CONTEXT AND CONTENT. Order Cingulata, family Dasypo- Dasypus gymnurus: Alfaro, 1897:46. Not Tatus gymnurus didae, subfamily Tolypeutinae, tribe Priodontini. Synonymy Olfers, 1818. Xenurus gymnurus: J. A. Allen, 1897:43. Not Tatus gymnurus Olfers, 1818. [Lysiurus (Lysiurus)] unicinctus: Trouessart, 1898:1146. Part; not Dasypus unicinctus Linnaeus, 1758. [Lysiurus (Lysiurus)] hispidus: Trouessart, 1898:1147. Part; not Dasypus hispidus Burmeister, 1854. Tatoua (Ziphila) centralis Miller, 1899:4. Type locality ‘‘Chamelecon, [Corte´s,] Honduras.’’ Tatoua (Ziphila) lugubris: Miller, 1899:6. Not Ziphila lugubris Gray, 1873. C[abassous]. (Ziphila) centralis: Palmer, 1899:72. Name combination. Cabassous hispidus: Bangs, 1900:89. Not Dasypus hispidus Burmeister, 1854. Cabassous (Ziphila) lugubris: J. A. Allen, 1904:421. Not Ziphila lugubris Gray, 1873. Fig. 1.—A male Cabassous centralis from Centro de Recepciońy [Cabassous (Ziphila)] lugubris: Trouessart, 1905:821. Part; Rehabilitacioń de Fauna Silvestre in Colombia. Used with not Ziphila lugubris Gray, 1873. permission of the photographer Alba Lucıá Morales. 45(898)—Cabassous centralis MAMMALIAN SPECIES 13 modified from Gardner (2005) and Wetzel et al. (2008). C. centralis is monotypic (Wetzel et al. 2008).

NOMENCLATURAL NOTES. The generic name, Cabassous,is from the native language Galibi (French Guiana), derived from capocou meaning armadillo. The species name, centralis, refers to the Central American (Honduras) origin of the holotype (Tirira 2004). Other common names are armadillo hediondo, armadillo rabo de carne de occidente, armadillo rabo de puerco, armadillo zopilote, armado de zopilote, cabasu´, cachicamo morrocoy, cucuso, cusuco venenoso, cuspa montan˜era zuliana, cuspita, douro´, morrocoy, No¨rdliches Nacktschwanz-Gu¨rteltier, pitero de un˜a, rabo de carne, rabo de molle, ta´kan ta´kan, tatu´-aı´, tatu´ de rabo molle, timba, tumbo, tumbo armado, and wai-wech (True 1896; Wetzel 1982; Emmons and Feer 1990; Superina and Aguiar 2006).

DIAGNOSIS

Cabassous centralis (Fig. 1) is similar to C. unicinctus (southern naked-tailed armadillo) but their ranges do not overlap: C. centralis occurs west of the Andes and C. unicinctus occurs east of the Andes. C. centralis differs from C. unicinctus in having the external surface of pinna without scales. The elongate, oval teeth are similar to those of C. tatouay (greater naked-tail armadillo), but in C. centralis the long axis of midposterior teeth may be diagonal to long axis of the skull (Wetzel 1985). Posterior surface of pinna is naked in C. centralis, whereas in other species the pinna is partially or completely scaled. Interorbital (24.3 mm) and zygomatic (40.8 mm) widths are smaller in C. centralis than in C. tatouay (33.8 and 56.3, respectively) and C. unicinctus (26.6 and 44.7, respectively—Wetzel 1980).

GENERAL CHARACTERS

A small armadillo with a slender skull (Fig. 2), naked cheeks and pinnae, mediolaterally compressed teeth, and Fig. 2.—Dorsal, ventral, and lateral views of skull and lateral view scutes on the first 2 complete rows of scapular shield that are of mandible of an adult Cabassous centralis (26621, Instituto de much wider than long (Wetzel 1980). Counts of scutes (SD, Biologıá, Universidad Nacional Autońoma de Me´xico) from Selva n) are: cephalic shield, 35.3 (4.3, 23); 1st complete band of Lacandona, Chiapas, Mexico. Greatest length of skull is 77.6 mm. scapular shield, 18.1 (1.3, 16); last band of scapular shield, Photographs provided by Linda Saldanã, Daniela Labastida, and Fernanda Caciano. 27.2 (1.6, 19); 3rd movable band, 28.3 (1.9, 19); 4th movable band, 28.6 (1.5, 21); 1st band of pelvic shield, 25.9 (0.9, 18); last band of pelvic shield, 8.3 (0.9, 18); total number of movable bands, 12.1 (0.6, 22—Wetzel 1980). length of ear, 31, 33, 32, 32, 37, —. Ranges (mm) of external External measurements (mm) for 6 adult Cabassous measurements from the Neotropics were: length of head and centralis (2 females, 4 males, respectively) in the National body, 305–417; length of tail, 106–183; length of hind foot, Museum of Natural History (formerly the United States National Museum [USNM]) were: length of head and body, 60–76; length of ear, 27–62; mass, 2–3.5 kg (True 1896; 336, 337, 335, 363, 378, 400; length of tail, 164, 145, 134, Bangs 1900; Allen 1904; Meritt 1985; Wetzel 1985; Emmons 160, 183, 173; length of hind foot, 69, 62, 61, 60, 74, 79; and Feer 1990; Carrillo and Wong 1992). 14 MAMMALIAN SPECIES 45(898)—Cabassous centralis

Mean cranial measurements (mm) for 7 adult C. centralis in the USNM and ranges (in parentheses) from additional references (True 1896; Allen 1904; Wetzel 1980) were: skull: greatest length, 79.1 (72.09–83.3); width across zygoma at postorbital process, 41.8 (39.2–45.0); length of upper toothrow, 28.7 (26.6–30.0); posterior margin of last molar to tip of premaxilla, 48.2 (45.0–51.2); condylobasal length, 78.0 (72.3–83.1); rostral length, 36.9 (33.6–39.0); palatal length, 45.1 (41.9–48.1); postrostral length, 41.2 (37.5–44.5); palatal width, 11.4 (10.2–12.3); anterior rostral width, 11.6 (10.4–13.8); interlacrimal width, 33.3 (30.9– 36.4); interorbital width, 24.3 (23.6–25.6); zygomatic width, 40.8 (37.4–43.7); mastoidal width, 37.0 (33.6–41.0); height of cranium, 32.2 (30.1–34.0); mandible: greatest length, 60.2 (55.0–63.4); width from angle to coronoid, 17.8 (16.0–19.7); length of lower toothrow, 26.0 (24.6–27.7); posterior margin of last molar to tip of mandible, 42.0 (38.5–43.8). Means of selected limb measurements (mm) for 2 C. centralis were: humeral length, 54.2; proximal humeral length, 39.4; ulnar length, 57.6; olecranon length, 27.4; functional femoral length, 61.5; proximal femur length, 28.4; leg length, 51.2; and midleg width, 20 (Vizcaıńo and Milne 2002). Fig. 3.—Distribution map of Cabassous centralis modified from Mean tooth lengths and widths (mm), respectively, with Wetzel et al. (2008). parenthetical ranges were: maxillary teeth: 4th, 2.9 (2.4–3.3), 2.3 (2.1–2.6); 5th, 3.0 (2.5–3.4), 2.6 (2.3–3.0); 6th, 2.8 (2.4– moveable bands (Emmons and Feer 1990). The slender tail 3.2), 2.8 (2.4–3.2); 7th, 2.7 (2.2–3.4), 2.7 (2.3–3.2); mandib- is distinctive with widely spaced, thin plates with a pinkish ular teeth: 5th, 3.1 (2.7–3.5), 2.6 (2.2–3.0); 6th, 3.1 (2.4–3.8), gray color (Eisenberg 1989). Upperparts of the back mostly 2.7 (2.4–3.2); 7th, 2.7 (1.9–3.8), 2.5 (1.8–2.9—Wetzel 1980). dark gray-brown but the edge of the carapace is yellowish (Eisenberg 1989; Reid 1997). Tufts of hair on the ventrum are in about 20 regular, transverse rows (True 1896). DISTRIBUTION Cabassous centralis has no incisors or canines and 7–10 upper and 7–9 lower uniform cheek teeth (8 specimens at the Cabassous centralis occurs from sea level up to 2,000 m, USNM—True 1896). The long tongue can be ‘‘extruded to a and with a 3,018-m record from Antioquia Department in great length when feeding’’ (Eisenberg 1989:63). The head of Colombia (Sańchez et al. 2004; Dıáz-N. and Sańchez- the mallei of C. centralis is globose and the anterior process Giraldo 2008). C. centralis ranges from southern Mexico, is noticeably thickened (Patterson et al. 1992). through Central America, into South America where it The diploid chromosome number (2n) is 62 and the occurs ‘‘from northern Colombia (including northern inter- fundamental number (FN) is 74 (Benirschke et al. 1969; Hsu Andean valleys) east of the Choco´ region and north of the and Benirschke 1969). The X is submetacentric and the Y is Andes eastward across Venezuela as far as the state of metacentric (Jorge et al. 1985). Monagas’’ (Fig. 3; Wetzel et al. 2008:149). It also is found Function.—A 3.81-kg animal had a basal rate of metabolism on the coastal lowlands of northwestern Ecuador (Orce´s and 1 1 of 0.213 ml O2 g h with a minimal conductance of 0.033 Albuja 1985; Tirira 2007). No fossils are known. 1 1 1 ml O2 g h 8C (McNab 1980, 1985). Derived indexes calculated from limb measurements are FORM AND FUNCTION as follows: functional forearm length (difference between ulnar length and olecranon length) divided by humeral Form.—The snout is short and broad; the ears are length (brachial index), 54.43; length of the olecranon moderately large and funnel shaped; and the eyes are process divided by the difference between ulnar length and extremely small (Emmons and Feer 1990). The forefeet have olecranon length (index of fossorial ability), 90.74; proximal 5 claws; the middle claw is extremely large and sickle shaped femoral length divided by the functional femur length (hip (Emmons and Feer 1990). The dorsal plates are large, moment index), 46.04; leg length divided by the functional roughly square in shape, and arranged in transversal rows femur length (crural index), 79.38; forelimb length divided for the entire length of the body, with 10–13 inconspicuous by hind-limb length (intermembral index), 72.34; and midleg 45(898)—Cabassous centralis MAMMALIAN SPECIES 15 width divided by leg length (leg robusticity index), 39.06 When restrained, C. centralis growls or squeals, urinates (Vizcaıńo and Milne 2002). copiously, and may defecate while twirling its tail (Emmons and Feer 1990). Other vocalizations include a low buzz or growl and low gurgling squeals (Meritt 1985). C. centralis ONTOGENY AND REPRODUCTION has a pungent, musky odor (Wetzel 1980). Cabassous centralis has a single offspring at each birth (Hayssen et al. 1993). At birth C. centralis is naked and CONSERVATION hairless, with closed ear pinnae and closed eyes (Meritt 1985). The claws are hardened; the enlarged foreclaw Cabassous centralis is listed under Appendix III of the resembled that of the adult stage (Meritt 1985). The banding Convention on International Trade in Endangered Species patterns and scutes are present, but the skin is soft, glossy, of Wild Fauna and Flora (2013) and is rare everywhere warm, and pinkish in color (Emmons and Feer 1990). The (Emmons and Feer 1990). This species is considered as nose tip and lower jaw are soft, wrinkled, and slightly ‘‘Least Concern’’ by the International Union for Conserva- distorted (Meritt 1985). tion of Nature and Natural Resources (Superina and Abba 2009). C. centralis represents 2% of the roadkills of vertebrates in Colombia, which is a low percentage ECOLOGY compared with other mammalian groups (Delgado-V. 2007). Automobiles are a major threat in human-dominated Cabassous centralis is one of the most fossorial landscapes. In some local regions of southern Mexico, this armadillos. It prefers dry to mesic forests, but now exists species of armadillo is thought to be poisonous, and it is mainly in patchy, degraded habitat, and it is considered rare killed every time it is encountered. In some regions in the wherever it occurs (Aguiar 2004; Cruz-Rodrı´guez et al. northern Andes of Colombia, C. centralis is sometimes used 2011). The species inhabits dry to moderately moist, as food (Aldana et al. 2006; Castanõ and Corrales 2010). deciduous and semideciduous forests, forest edges in rocky terrain, dry savanna (Genoways and Timm 2003), tropical ACKNOWLEDGMENTS moist montane forests and subparamo (Genoways and Timm 2003, Dıáz-N. and Sańchez-Giraldo 2008), secondary A. L. Morales provided the photograph of the animal. forest, and mixed forest and agricultural land (Superina and The skull plate was created by F. A. Cervantes under Abba 2009). In Chiapas, Mexico, it occurs in tropical consensus of the Coleccioń Nacional de Mamı´feros, evergreen forest, grasslands, and transformed areas of Instituto de Biologıá, Universidad Nacional Autońoma de secondary forest below 500 m (Cuaroń 2005). In Belize, it Me´xico. N. Martıńez-Mendez created the map. Funding was inhabits open savannahs of grasses and sedges with islands from the Blakeslee Grant for Genetics Research at Smith of trees surrounded by pimento palm (McCarthy 1982). In College. Nicaragua, it frequents grasslands with acacias (Genoways and Timm 2003). LITERATURE CITED One C. centralis lived for 5 years and 9 months in captivity and was about 2 years old at time of capture (Weigl AGUIAR, J. M. 2004. The 2004 edentate species assessment workshop: 2005). In the wild, C. centralis feeds primarily on terrestrial species summary and species discussion. Edentata 6:3–26. ants and termites (Cuaroń et al. 1989; Emmons and Feer ALDANA, N. J., M. DIÁZ PORRES,A.FEIJOO, AND M. C. ZUN˜ IGA. 2006. Valoracioń del uso del fauna silvestre en el municipio de Alcala´, 1990; Cuaroń 2005; Wetzel et al. 2007). The fungus Valle de Cauca. Scientia et Technica Anõ XII, 31:291–296. Paracoccidioides brasiliensis was identified from the spleen ALFARO, A. 1897. Mamı´feros de Costa Rica. Tipografıá Nacional, San of C. centralis (Corredor et al. 2005). Jose´, Costa Rica. ALLEN, J. A. 1897. Additional notes on Costa Rican mammals, with descriptions of new species. Bulletin of the American Museum of Natural History 9:31–44. BEHAVIOR ALLEN, J. A. 1904. Mammals from the District of Santa Marta, Colombia, collected by Mr. Herbert H. Smith, with field notes by Mr. Smith. Bulletin of the American Museum of Natural History Cabassous centralis is nocturnal, terrestrial, solitary 20:407–468. (Emmons and Feer 1990; Wetzel et al. 2008), and fossorial, BANGS, O. 1900. List of the mammals collected in the Santa Marta spending most of its time underground in large excavated region of Colombia by W. W. Brown, Jr. Proceedings of New England Zoo¨logy Club 1:87–102. ´ C. centralis tunnels (Cuaron 2005). walks on the tips of its BENIRSCHKE, K., R. J. LOW, AND V. H. FERM. 1969. Cytogenetics studies fore claws with pigeon-toed hind feet (Wetzel 1980). When of some armadillos. Pp. 330–345 in Comparative mammalian burrowing, C. centralis rotates its body, such that the front cytogenetics (K. Benirschke, ed.). Springer-Verlag, New York. BURMEISTER, H. 1854. Systematische Uebersicht der Thiere Brasiliens, portion of its body acts as an auger (Wetzel 1980). C. welche wa¨hrend einer Reise durch die Provinzen von Rio de centralis can swim (Ingles 1953). Janeiro and Minas Gerae¨s gesammelt oder beobachtet wurden 16 MAMMALIAN SPECIES 45(898)—Cabassous centralis

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