Artibeus lituratus (Olfers, 1818)

Don E. Wilson & Russell A. Mittermeier, 2019, Phyllostomidae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 444-583 : 575

196.

Great Fruit-eating Bat

Artibeus lituratus

French: Artibée rayée / German: GroRer Fruchtvampir / Spanish: Artibeo grande

Other common names: Great Artibeus

Taxonomy. Phyllostomus lturatus Olfers, 1818 ,

“ Paraguay.” Restricted by A. Cabrera in 1958 to Asuncion, Paraguay.

Artibeus lituratus is in subgenus Artibeus . Its name is based on the “chauve-souris obscure et rayée” described by EF d’Azara in 1801. The bat came from “Pueblo mi amigo Don Pedro Blas Noseda,” which would be closer to San Ignacio but not Asuncion, which raises some doubt about this type locality restriction. Delimitation of subspecies is uncertain at best because diverse morphometric and molecular analyses failed to unambiguously define independent evolutionary lineages. Some authors treated A. intermedius as a valid species, but here the most common taxonomy is maintained, and it considered a subspecies. Four subspecies recognized.

Subspecies and Distribution.

A.l.lituratusOlfers,1818—fromSoftheOrinocoBasininVenezuela,EthroughtheGuianas,andSthroughSEColombia,Ecuador,Brazil,Peru,andBoliviaintoParaguayandNEArgentina.

A.l. intermediusJ.A.A.l.,1897—fromC&SMexico(includingYucatanPeninsulaandCozumelI)throughCentralAmericatoNWColombia.

A.l.koopmaniWilson,1991—TresMariasIs,offNayarit,Mexico.

A. l. palmarumJ. A. A.l. & F. M. Chapman, 1897 — from N & E Colombia, Venezuela N of the Orinoco Basin, Trinidad and Tobago Is, and the S Lesser Antilles ( St. Vincent, the Grenadines, Grenada, and Margarita Is).

Descriptive notes. Head-body 72-95 mm (tailless), ear 19-26 mm, hindfoot 12-21 mm, forearm 59-68 mm; weight 43-59 g for subspecies intermedius. Head—body 80-101 (tailless), ear 20-26 mm, hindfoot 15-21 mm, forearm 69-77 mm; weight 55-90 g for subspecies palmarum. Greatest lengths of skulls are 27-4-30-2 mm (intermedius) and 30-6-32-9 mm (palmarum). The Great Fruit-eating Bat is among the largest species of Artibeus , showing extensive morphological variation throughout its distribution. This variation has been used as criteria for distinguishing taxonomic subdivisions, with subspecies palmarum being the largest and intermedius the smallest. Dorsal fur is brownish, ranging from pale or grayish brown to reddish brown (or grayish brown with reddish tinge) or even brown-chocolate; ventral fur is slightly paler, being brownish to dark brown but never frosted with white-tipped hairs. Dorsal fur is long (7-8 mm) and tricolored, with pale bases. Faint facial stripes are present; upper pair is always conspicuous, and lower ones can be obscure to faint (in intermedius). Horseshoe of noseleafis not attached to upper lip; chin has central wart surrounded by smaller ones. Body fur extends to contiguous areas of wings, which is more conspicuous on ventral view. Dorsal one-half of forearm is well furred, with long hairs. Dorsally, uropatagium is densely covered with thin hairs at least to knees, and usually legs are also covered with long brownish hairs. Skull is long and robust; preorbital and postorbital processes are well developed (rostral shield is present). Dental formulais12/2,C1/1,P2/2,M 2/3 (x2) = 30. I' has distinctly bilobed edges; M,is rarely missing. Chromosomal complement has 2n = 30-31 and FN = 56, with Y, Y, sex chromosome system.

Habitat. Wide variety of habitats from sea level to elevations of ¢. 2620 m. Subspecies lituratus and palmarum are typically found in tropical rainforests but also tropical deciduous forests. Subspecies intermedius and koopmani are found in subtropical deciduousforests, tropical thorn forests, and pine-oak vegetation. Great Fruit-eating Bats also occur in tropical semideciduous forests, cloud forests, savannas, seasonally dry forests, and human-modified habitats, such as agricultural land, especially in fruit trees and even urban areas.

Food and Feeding. The Great Fruit-eating Bat is a fruit generalist but prefers figs. It can consume flowers, leaves, pollen, and insects. Figs ( Ficus , Moraceae ) comprise most of the diet, being available year-round and thus securing food resources when other fruits are scarce. The role of the Great Fruit-eating Bat as a seed disperseris well studied; it is an important disperser for at least ten plant species in rainforests but also disperses at least 42 plant species in 17 families (e.g. Cecropia spp., Urticaceae ; Anacardium sp., Spondias sp., and Mangifera sp., all Anacardiaceae ; Musa sp., Musaceae ; Persea sp., Lauraceae ; Cyclanthus sp., Cyclanthaceae ; Cavendishia sp., Ericaceae ; Vismia sp., Hypericaceae ; Psidia sp., Myrtaceae ; Guazuma sp., Malvaceae ; Piper spp-, Piperaceae ; Prunus sp., Rosaceae ; and Solanum spp., Solanaceae ). Leaves of at least six species of plants have been also identified in diets (leaves are chewed, fluids ingested, and fibrous material not swallowed but dropped as pellets), probably to supplement protein requirements during certain reproductive periods. Insects can be a frequent food item as some specific habitats. Broad feeding niche of the Great Fruit-eating Bat suggests great ecological flexibility throughout the various habitats is occupies and allowing it to respond to changing seasonal and annual availability of resources.

Breeding. The Great Fruit-eating Bat exhibits a reproductive pattern of bimodal polyestry that varies temporally with latitude and longitude and typically peaks in dry and wet seasons. In Central America, these peaks in pregnancy occur in March and July, with peaks in lactation 2-3 months later. Peaks of pregnancy occur in October-November in Ecuador and Colombia, February in Venezuela and Brazil (Para), and April-May in Trinidad and Tobago. There is a second pregnancy peak in June-July in Venezuela, July in Minas Gerais, Brazil, and August in Amazonas, Brazil. These differences are associated with changing abiotic conditions and local availability of resources. Gestation lasts 3-5-4 months. Females almost always give birth to one young per pregnancy. Based on histological examination of reproductive tracts, populations from south-eastern Brazil exhibited postpartum estrus and bimodal polyestry, with births peak in November and March, coinciding with periods of greater rainfall.

Activity patterns. The Great Fruit-eating Bat becomes active during the first two hours after sunset, flying directly to fruit trees previously identified during commuting flights. These trees are used for almost a week, before moving to a next feeding area (patch of fruiting trees). It mostly forages in higher levels of the forest. It has a larger home range than the Flatfaced Fruit-eating Bat (A. planirostris ) or the Dark Fruit-eating Bat (A. obscurus ), which was suggested to be linked to their ability to flock together to exploit fruit crops of large trees. The Great Fruit-eating Bat usually roosts in dense vegetation, hanging from tree branches under leaves. It takes advantage of broken or crossed palm fronds, in vine-tangled crowns of subcanopy trees, and in dark cave-like recesses on undersides of crowns of canopy trees. It also roosts in large hollow trees ( Ceiba spp.).

Movements, Home range and Social organization. Great Fruit-eating Bats hang singly or in small groups in exposed entrances of caves or well-illuminated openings of large caves, or by hanging from shaded roofs of small niches in cliffs. Groups contain 5-25 individuals. Based on observations of only one adult male in each group,it probably has a harem-based mating system.

Status and Conservation. Classified as Least Concern on The IUCN Red List. The Great Fruit-eating Bat has a wide distribution, occurs in various habitats, and is abundant.

Bibliography. Albuja (1999), Allen (1897), Allen & Chapman (1897), Andersen (1906b, 1908c), Ascorra et al. (1996), d/Azara (1801), Cabrera (1958), Castano et al. (2018), Davis (1970b, 1984), Fleming et al. (1972), Handley (1987), Hershkovitz (1949), Koepcke & Kraft (1984), Larsen et al. (2010b, 2013), Marchan-Rivadeneira et al. (2012), Marques-Aguiar (2008a), Morrison (1980), Olfers (1818), Pacheco et al. (2010), Phillips et al. (1991), Redondo et al. (2008), Simmons (2005), Wilson (1991), Zortéa & Mendes (1993).

Figures

187. Brown Fruit-eating Bat (Artibeus concolor), 188. Honduran Fruit-eating Bat (Artibeus inopinatus), 189. Fraternal Fruit-eating Bat (Artibeus fraterculus), 190. Hairy Fruit-eating Bat (Artibeus hirsutus), 192. Ecuadorian Fruit-eating Bat (Artibeus aequatorialis), 193. Jamaican Fruit-eating Bat (Artibeus jamaicenss), 194. Dark Fruit-eating Bat (Artibeus obscurus), 195. Schwartz’s Fruit-eating Bat (Artibeus schwartzi), 196. Great Fruit-eating Bat (Artibeus lLituratus), 197. Large Fruit-eating Bat (Artibeus amplus), 198. Flat-faced Fruit-eating Bat (Artibeus planirostris), 199. Rosenberg’s Fruit-eating Bat (Artibeus rosenberg), 200. Thomas's Fruit-eating Bat (Artibeus watsoni), 201. Toltec Fruit-eating Bat (Artibeus toltecus), 202. Pygmy Fruit-eating Bat (Artibeus phaeotis), 203. Gervais’s Fruit-eating Bat (Artibeus cinereus), 204. Andersen’s Fruit-eating Bat (Artibeus anderseni), 205. Little Fruit-eating Bat (Artibeus ravus), 206. Aztec Fruit-eating Bat (Artibeus aztecus), 207. Bogota Fruit-eating Bat (Artibeus bogotensis), 208. Silvery Fruit-eating Bat (Artibeus glaucus), 209. Dwarf Fruit-eating Bat (Artibeus gnomus), 210. Jamaican Fig-eating Bat (Ariteus flavescens), 211. Tree Bat (Ardops nichollsi), 212. Red Fruit Bat (Stenoderma rufum), 213. Wrinkle-faced Bat (Centurio senex), 214. Ipanema Broad-nosed Bat (Pygoderma bilabiatum), 215. Visored Bat (Sphaeronycteris toxophyllum), 216. Little White-shouldered Bat (Ametrida centurio), 217. Cuban Fig-eating Bat (Phyllops falcatus)

Subspecies and Distribution. A.l.lituratusOlfers,1818—fromSoftheOrinocoBasininVenezuela,EthroughtheGuianas,andSthroughSEColombia,Ecuador,Brazil,Peru,andBoliviaintoParaguayandNEArgentina. A.l. intermediusJ.A.A.l.,1897—fromC&SMexico(includingYucatanPeninsulaandCozumelI)throughCentralAmericatoNWColombia. A.l.koopmaniWilson,1991—TresMariasIs,offNayarit,Mexico. A. l. palmarumJ. A. A.l. & F. M. Chapman, 1897 — from N & E Colombia , Venezuela N of the Orinoco Basin, Trinidad and Tobago Is, and the S Lesser Antilles ( St. Vincent , the Grenadines , Grenada , and Margarita Is).