PHYLLOSTOMIDAE Leaf-nosed Bats FAUNA PARAGUAY

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See below for a key to the Paraguayan species.

PHYLLOSTOMIDAE: LEAF-NOSED BATS
General characteristics: Twenty-one Paraguayan species in this large and varied family confined to the New World. The most notable characteristic of the family is the leaf-like growth on the nose - an adaptation for echo-location. The uropatagium, where present completely encloses the short tail and the foot is unmodified. Tragus is well-developed. Bats of this family are broad-winged with slow, lazy flight designed for flying through dense vegetation. Typically they feed on stationary objects such as fruit or insects gleaned from foliage. As a result of the dense habitat in which they live they use a mixture of sight, olfaction, sound and echolocation when searching for prey. Echolocation calls of phyllostomid bats are relatively uniform in structure, consisting primarily of faint, broadband, and steep frequency modulated calls (Weinbeer & Kalko 2007). Six subfamilies occur in Paraguay and the characteristics of each are given below.
Cranial characteristics: Lacks a postorbital process. Premaxillaries are complete and fused with each other and the maxillary. Click here to view the Phyllostomidae Comparative Skull Gallery
Skeletal characteristics: Digit II with well-developed metacarpal and small phalanx. Digit III has three bony phalanges. Humerus with well-developed trochiter and double scapula articulation. Seventh cervical vertebrae is free from first thoracic. Fibula cartilaginous proximally.
Taxonomy: According to Gardner (2007) the Phyllostomidae can be split into six subfamilies, five of which are represented in Paraguay. The Carolliinae are sometimes included within the Stenodermatinae. A key to the subfamilies is offered below.

Key to Subfamilies of Paraguayan Leaf-nosed Bats

1a Nose leaf present. Two upper incisors present on each side.............................................................................2
1b Nose leaf absent/poorly-developed. Tail absent, uropatagium much reduced so that the legs appear free. Agile and rapid when walking on the ground. One extremely sharp and pointed upper incisor on each side. .....................................................................................................................................................Desmodontinae

2a Ears long, extending beyond snout when laid forward. Tail and uropatagium well-developed. Tip of tail free of dorsal surface of uropatagium. Nose leaf large with horseshoe shape encompassing nostrils...Phyllostominae
2b Ears do not extend beyond snout when laid forward. Tail extremely short or absent.......................................3

3a Snout long and narrow. Incisors reduced or absent. Tongue extremely long with brush tip....Glossophaginae
3b Snout short and broad. Incisors present and well-formed. Tongue of normal length, lacking brush tip............4

4a Uropatagium much reduced or absent. Tail extremely short or absent. Many species with conspicuous spots or stripes on pelage.....................................................................................................................Stenodermatinae
4b W-shaped uropatagium well-developed. Tail extremely short. Pelage without spots or stripes........Carolliinae

Subfamily Phyllostominae Gray, 1825
General characteristics: Seven Paraguayan species in five genera. Large pointed nose-leaf and a free horseshoe-shape below encompassing the nostrils. The uropatagium is long and completely encloses the tail, the tip of which departs from the dorsal side of the membrane. The forehead is steep and sloping angled straight down to the long snout. The ears are large.
Dental characteristics: Molars characteristically W-shaped.
Skeletal characteristics: Humerus with definite secondary articulation with scapula. Epitrochlea large with slightly developed spinous process.
Ecology: Phyllostomids are carnivorous or insectivorous bats, snatching prey from a substrate or water with the mouth, and not habitually taking prey from the air.

Chrotopterus W.Peters, 1865: Woolly False Vampire
A monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Vampyrus W.Peters 1865:305. Not Vampyrus Leach (1821).
Chrotopterus W.Peters 1865:505. Type species Vampyrus auritus W.Peters (1865) by original designation. Proposed as a subgenus of Vampyrus Leach (1821).
Chrotoptems Dobson 1878:478. Incorrect spelling.
Chrotoperus RJ Baker, RM Fonseca, DA Parish, CJ Phillips & FG Hoffman 2004:4. Incorrect spelling.
General characteristics: One of the largest of South American bats. Ears extremely large, well-separated and simple. Pelage is long and woolly and tail is reduced to the point of sometimes appearing absent. Calcar is longer than the foot. Lips and chin almost smooth.
Cranial characteristics: See species account.
Skeletal characteristics: Wide rib cage. Manubrium of sternum with ventral projection providing additional attachment for pectoralis muscle. Large, elongated and pointed scapulae.
Dental characteristics: See species account.
Genetic characteristics: See species account.
Evolution: Fossils are known from the Pleistocene of Brazil.
Paraguayan Species:
Chrotopterus auritus - Woolly False Vampire

Lophostoma d´Orbigny, 1836: Round-eared Bats
Six species, two present in Paraguay. Synonyms adapted from Gardner (2007).
Synonyms:
Lophostoma d´Orbigny 1836:pl.6. Type species Lophostoma silvicolum d´Orbigny (1836) by monotypy.
Phyllostoma JA Wagner 1843:365. In part. Not Phyllostoma G.Cuvier (1800).
Vampyrus Pelzeln 1883:110. In part. Not Vampyrus Leach (1821).
Tonatia Palmer 1898:110. In part. Not Tonatia Gray in Griffith, Hamilton-Smith & Pidgeon (1827).
Chrotopterus JA Allen 1910:147. In part. Not Chrotopterus W.Peters 1865.
Tonatia O.Thomas 1910:184. In part. Not Tonatia Gray in Griffith, Hamilton-Smith & Pidgeon (1827).
Tonatia Goodwin 1942:205. In part. Not Tonatia Gray in Griffith, Hamilton-Smith & Pidgeon (1827).
Tonatia WB Davis & Carter 1978:6. In part. Not Tonatia Gray in Griffith, Hamilton-Smith & Pidgeon (1827).
Tonatia Genoways & Williams 1980:205. In part. Not Tonatia Gray in Griffith, Hamilton-Smith & Pidgeon (1827).
Tonakia Ascorra, Gorchov & Cornejo 1994:537. Incorrect spelling.
General characteristics: Small to medium-sized bats with short fur and calcar longer than foot. Similar in general appearance to Tonatia, but these bats curl the ears when handled which Tonatia does not and the facial fur is much shorter so that the snout may appear naked.
Cranial characteristics: Postorbital constriction narrow, less than 90% of the breadth across the cingula of the canines.
Dental characteristics: Dental Formula I2/1 C 1/1 P2/3 M3/3 = 32.
Taxonomy: Formerly considered part of Tonatia.
Evolution: Lophostoma silvicolum is known from the late Pleistocene of northwestern Peru (Czaplewski 1990).
Paraguayan species:
Lophostoma brasiliense - Pygmy Round-eared Bat
Lophostoma silvicolum - d'Orbigny's Round-eared Bat

Macrophyllum Gray, 1838: Long-legged Bat
Monotypic genus. Synonyms adapted from Gardner (2007).
Synonyms:
Phyllostoma G.Cuvier 1800: Table 1. In part. Unjustified emendation of Phyllostomus Lacépède (1799).
Macrophyllum Gray 1838:489. Type species Macrophyllum nieuwiedii Gray (1838) by monotypy.
Dolichophyllum Lydekker 1891:673. Replacement name for Macrophyllum which was presumed to be preoccupied by Macrophylla Hope (1837) a Coleopteran.
Dolychophyllum Trouessart 1904:110. Incorrect spelling.
Mesophyllum COC Vieira 1942:311. Incorrect spelling.
Maerophyllum George, Marques, Vivo, Branch, Gomes & Rodrigues 1988:39. Incorrect spelling.
General characteristics: One of the smallest members of the family. Tail is long and extends to the posterior margin of the uropatagium. Ventral surface of uropatagium with rows of dermal denticles. Feet greatly elongated.
Cranial characteristics: See species account.
Dental characteristics: See species account.
Genetic characteristics: See species account.
Paraguayan species:
Macrophyllum macrophyllum - Long-legged Bat

Phyllostomus Lacépède, 1799: Spear-nosed Bats
Four species in this genus, two of which occur in Paraguay. Synonyms adapted from Gardner (2007).
Synonyms:
Vespertilio Pallas 1767: 7. In part. Not Vespertilio Linnaeus 1758.
Pteropus Erxleben 1777:136. Not Pteropus Brisson 1762.
Phyllostomus Lacépède, 1799:16. Type species Vespertilio hastatus Pallas 1767 by monotypy.
Phyllostoma G.Cuvier 1800: Table 1. Unjustified emendation.
Phylostoma G.Fischer 1817:373. Incorrect spelling.
Alectops Gray 1866:114. Type species Alectops ater by monotypy.
Phyllostomum Thenius 1926:257. Incorrect spelling.
General characteristics: Medium to large, robust bats with moderately-small nose leaf. Tail reaches middle of uropatagium. Ears small and separated. Calcar length when compared to foot length varies between species. Males with well-developed gular gland.
Cranial characteristics: Skull heavy with broad, low and flattish rostrum. Sagittal crest broadly expanded. Paroccipital processes expanded and shelf-like. Zygomata heavy and slightly expanded anteriorly and posteriorly. Basisphenoid pits shallow. Floor of braincase forms an angle wth the roof of the posterior nasals. Auditory bullae small and flat covering approximately half of the cochlea and much wider than they are high.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/2 M3/3 = 32. Two incisors and two premolars in lower toothrow. Hypocone low in lower first molar and obsolete in second. Premolars with thick, short shafts and slightly developed cingula. Middle upper incisor simple and slightly projecting, concave on posterior edge and with anterior surface almost as long as it is high. Lateral incisor small and blunt barely rising above the cingulum of the canine. Upper canine with a broad, flattish groove on anterior face of shaft. Lower incisors form almost straight row between the cingula of the canines, though the outer teeth are slightly smaller. Upper surface of crown slightly longer than the height of the anterior surface. Cutting edge slightly trifid and middle lobe smallest.
Physiology: Valdivieso & Tamsitt (1964) describe the histology of the chest gland of P.discolor. Hoffman et al (2008) describe the auditory cortex of P.discolor.
Evolution: Remains of P.hastatus are known from late Pleistocene caves in Minas Gerais, Brazil (Santos et al 2003). Van den Bussche & Baker (1993) concluded that Phyllostomus jointly with Phylloderma W.Peters, 1865 formed a natural assemblage and that P.hastatus forms a sister group with P.elongatus (Geoffroy St. Hilaire, 1810). The four species of Phyllostomus and Phylloderma are estimated to have diverged at approximately the same time.
Paraguayan species:
Phyllostomus discolor - Pale Spear-nosed Bat
Phyllostomus hastatus - Greater Spear-nosed Bat

Tonatia Gray, 1827: Round-eared Bats
General characteristics: Medium-sized bats with short fur and calcar longer than the foot. Similar in general appearance to Lophostoma, but these bats do not curl the ears when handled as does Lophostoma and the snout is well-furred. Tail extends to the middle of the uropatagium.
Cranial characteristics: Postorbital constriction broad, greater than 90% of the breadth across the cingula of the canines.
Dental characteristics: Dental Formula I2/1 C 1/1 P2/3 M3/3 = 32.
Tonatia bidens - Spix's Round-eared Bat

Key to Adult Spear-nosed Bats of Paraguay
1a Forearm >70mm...............................................................................................................................................6
1b Forearm <65mm...............................................................................................................................................2

2a Very small, forearm <36mm. Tail long, enclosed in uropatagium and reaching almost to posterior margin. Ventral surface of uropatagium with rows of dermal denticles. Feet elongated.......................................................................................................Macrophyllum macrophyllum
2b Forearm in range 36-65mm. Tail extends less than half the length of the uropatagium. No dermal denticles on ventral surface of uropatagium. Feet of normal size.........................................................................................3

3a Ears long and rounded. Tail approximately half length of uropatagium with tip poking free on dorsal surface. Forearm typically <56mm. Pelage long, soft and not dense..................................................................................4
3b Ears broad and triangular. Tail approximately one third the length of the uropatagium and totally enclosed within it. Forearm typically >56mm. Pelage smooth, dense and grizzled, not extending to membranes................................................................................................................Phyllostomus discolor

4a Forearm in range 50-56mm ..............................................................................................................................5
4b Forearm <45mm long. Muzzle appears naked or sparsely-furred.....................Lophostoma brasilense

5a Ears >30mm long connected by a piece of skin, passing tip of nose when laid forwards. Forearm almost naked. Muzzle appears naked or sparsely-furred. May have white throat patch of variable size, but lacks median white head stripe....................................................................................................Lophostoma silvicolum
5b Ears <30mm long and separated, not reaching the nose when laid forwards. Forearm furred. Muzzle well-furred. Variably conspicuous pale median head stripe present, but lacks white gular patch.................................................................................................................................Tonatia bidens

6a Ears long (>45mm) and well-separated. Pelage long and woolly, extending over basal half of forearm. Wing membranes dark, tipped white...................................................................................Chrotopterus auritus
6b Ears <30mm and well-separated. Pelage short, dense and velvety extending onto uropatagium and along basal third of forearm. Wing membranes lack white...............................................Phyllostomus hastatus

Subfamily Glossophaginae Bonaparte, 1845
General characteristics: Two Paraguayan species of nectarivorous bat with small ears, slender heads and long snouts furred to at least around the eye. These are small bats with an extremely short tail (absent in Anoura). The nasal leaf is reduced and lacks the horseshoe of other Phyllostomids and the tongue is very long with a tip covered in papillae - an adaptation for nectar-feeding. Tongue lacks a groove along the lateral margins. The pelage is paler at the base, especially noticeable around the neck and shoulders. The lower lip is V-shaped and surrounded by warty growths.
Cranial characteristics: Braincase large, rounded with a large foramen magnum. Zygomatic arches weak or incomplete. Auditory bullae small.
Dental characteristics: Dental Formula I2/0-2 C 1/1 P2-3/3 M2-3/2-3 = 26-34. Dentition is comparatively weak. Molars elongated antero-posteriorly and upper molars lack a hypocone. Premolars with recuced cusps and styles.
Skeletal characteristics: Atlas twice as wide as other cervical vertebrae, which are wider than postcervical vertebrae. Sternum is T-shaped with low ventral keel. Scapula with reduced spinous processes, large acromion process and large coracoid process.

Anoura Gray, 1838: Tailless Bats
General characteristics: One species present in Paraguay. Medium-sized Glossophagines that lack a tail and have an extremely narrow interfemoral membrane.
Cranial characteristics: Zygomatic arch frequently not fully ossified and appears incomplete.
Dental characteristics: Dental Formula I2/0 C 1/1 P3/3 M3/3 = 32. Upper incisors in two pairs separated by a gap. Outer incisors blade-like, inner incisors peg-like. Lower incisors absent. Canines show sexual dimorphism, uppers being larger in males. Hypocone absent but all other major cusps present on molars, though reduced in size.
Anoura caudifer - Lesser Tailless Bat

Glossophaga E. Geoffroy-St-Hilaire, 1818: Long-tongued Bats
General characteristics: One species present in Paraguay. Small Glossophagines with a short tail 50% as long as the long interfemoral membrane.
Cranial characteristics: Rostrum shorter than the braincase. Zygomatic arches complete.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/3 M3/3 = 34. Outer upper incisors smaller and more pointed than inners. Upper and lower molars lack hypocone, other cusps and lophs are well-developed.
Glossophaga soricina - Pallas's Long-tongued Bat

Key to Adult Nectarivorous Bats of Paraguay
1a Tail absent and uropatagium greatly reduced. .............................................................................Anoura caudifer
1b Tail present but extremely short in conspicuous W-shaped uropatagium...............................Glossophaga soricina

Subfamily Carolliinae Miller, 1924
General characteristics: A single Paraguayan species of fruit-eating bat with a short broad snout and prominent nose-leaf. Behind the snout the forehead is steep, it does not slope like in the Phyllostominae. The lower lip is V-shaped and surrounded by warts. Pelage is long and soft, banded and basally darker. Tail is very short, enclosed within a W-shaped uropatagium and with the tip extending slightly out of the dorsal side. Ears are medium-sized with pointed tips, shorter than the Phyllostominae.
Cranial characteristics: Generalised and intermediate skull when compared to the extremes shown in the family. Bony palate extends behind the maxillary toothrow in a tube that encloses the naso-pharyngeal cavity. Lateral emarginations on the outer palate separate the tube from the molars.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/2 M3/3 = 32. Cusps of upper and lower molars reduced and located towards lingual margin. Upper molars modified and W-pattern not noticeable.
Skeletal characteristics: Digit III exceptionally long. Digit IV is short as a result of a fourth metacarpal reduced in length. Trochanter impinges on the scapula.

Carollia Gray, 1838: Short-tailed Fruit Bats
General characteristics: Tail short, about 50% of interfemoral membrane. Primarily frugivorous though some insect material may be taken.
Dental characteristics: M1 and M2 both with protocone. Lower molars with well-developed protoconid and hypoconid.
Carollia perspicillata - Seba´s Short-tailed Bat
NB: A second species, Carollia brevicauda may be present in the extreme north or the Oriental region of Paraguay though its presence has yet to be conclusively proved. The two species are extremely difficult to distinguish and due to considerable variability specimens cannot always be assigned conclusively to species. As a general rule C.perspicillata has a forearm >40mm and tail approximately 10mm long whilst C.brevicauda has a forearm in the range 37-42mm and tail <7mm, but there is a zone of overlap in forearm measurements.

Subfamily Stenodermatinae P.Gervais, 1856
General characteristics: Eight Paraguayan species of fruit-eating bats with a short, broad snout, large eyes and a flattish head. The ears are medium-sized and triangular. The tail is greatly reduced or absent, though the uropatagium is present. Nose leaf is present and well-developed. Fruit is plucked from a tree with the mouth in flight and the bat then retires to a concealed perch to eat it, chewing the fruit and dropping the seed and masticated material. Many species show pale longitudinal bands on the pelage.
Dental characteristics: Upper molars with reduced cusps so that W-shape pattern is partly altered. Crowns of lower molars typically reduced with well-developed cusps.
Skeletal characteristics: Trochiter impinges on the scapula.
Taxonomy: Gardner (2007) splits the subfamily into three tribal groupings, the Stenodermatini (represented in Paraguay only by Pygoderma), Sturnirini (represented by Sturnira) and Ectophyllini (containing the remaining Paraguayan genera). Tribal groupings are not recognised here pending further study to clarify relationships.

Artibeus Leach, 1821: Fruit-eating Bats
General characteristics: Medium to large fruit bats, generally with facial stripes but lacking dorsal stripes. Fur is soft and short on the lower back. Triangular ears of medium-size.
Cranial characteristics: Skull short and broad.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/2 M2-3/2-3 = 28-32. Upper incisors larger than lowers and crowded between canines. Inner upper incisors bilobed and larger than outers. Canines robust and lack secondary cusps. Upper premolars with triangular outer cusp and low, slightly concave inner basin. Lower premolars similar but broader and lack posteromedial basin. Molars broad with low cusp. Third molar reduced or absent.
Taxonomy: All the Paraguayan species belong to the subgenus Artibeus, Leach, 1821 as defined by Gardner (2007).
Artibeus fimbriatus - Fringed Fruit-eating Bat
Artibeus lituratus - Great Fruit-eating Bat
Artibeus planirostris - Flat-faced Fruit-eating Bat

Chiroderma W.Peters, 1860: Big-eyed Bats
General characteristics: A single species present in Paraguay. A medium-sized bat with prominent white facial lines and a white dorsal line that arises between the shoulders. Forearms and legs are well-furred and fur extends to the dorsal surface of the uropatagium. Head is short and broad with a deep muzzle and the basal part of the nose leaf is free from the upper lip. External nares dorsally emarginate.
Cranial characteristics: Nasal bones absent.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/2 M2/2 = 28. Upper incisors long and slender, lower incisors smaller and subequal. P1 is small, separated from P2 by a gap but in contact with the canine.
Chiroderma doriae - Brazilian Big-eyed Bat

Platyrrhinus Saussure 1860: Broad-nosed Bats
General characteristics: A single Paraguayan species with a prominent white dorsal stripe and pale facial stripes. Uropatagium narrow and thickly-fringed with hair.
Cranial characteristics: Rostrum as wide as and almost as long as the braincase.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/2 M3/3 = 32. Fairly large upper inner incisors converge at tip. Upper outer incisors bifid, less than 50% of the length of the inner incisors. Lower incisors fill the space between the canines. First two upper molars with almost smooth crushing surfaces.
Platyrrhinus lineatus - White-lined Broad-nosed Bat

Pygoderma Saussure, 1860: Ipanema Bat
General characteristics: Monotypic genus. Dorsal pelgae tricoloured and small white patch of fur on each shoulder. Ventral pelage sparse on the chest in males. Nose leaf prominent. Ears broad and rounded with small tragus. Uropatagium is densely-furred, as are the legs and feet. Facial glands surround the eye, and give a particulalry swollen-eyed look to males.
Cranial characteristics: Rostrum deep and cuboid.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/2 M2/2-3 = 28-30. Third lower molar occasionally present.
Pygoderma bilabiatum - Ipanema Bat

Sturnira Gray, 1842: Little Yellow-shouldered Bat
General characteristics: The single Paraguayan species is a small forest bat with a greatly reduced but thickly-furred uropatagium. Shoulder glands stain the surrounding fur yellow, though it is variable in its conspicuousness and may be related to breeding activity. There is no tail. Ears are short and rounded and nose leaf is well-developed. Diet includes fruit and pollen of many plants, but particularly Solanum sp.
Cranial characteristics: Moderately-developed sagittal crest. Braincase elevated above the rostrum which is more than 50% longer than the braincase.
Dental characteristics: Dental Formula I2/1-2 C 1/1 P2/2 M3/3 = 30-32. Molars with cusps and cuspids located on extreme lingual and labial margins and separated by a U-shaped median groove which continues from the last upper premolar through the upper molars. In the lower molars it is interrupted by the paraconid. Canines sexually dimorphic, being longer in males.
Sturnira lilium - Little Yellow-shouldered Bat

Vampyressa O.Thomas, 1900: Yellow-eared Bats
General characteristics: A single species of small bat with well-defined nose leaf with prominent median rib. Incospicuous facial stripes often present but dorsal stripe lacking. Uropatagium narrow and sparsely-haired.
Cranial characteristics: Skull short and broad with rostrum shorter than braincase. Palate extends beyond the molars. Mesopterygoid fossa long and narrow. Basisphenoid pits deep.
Dental characteristics: Dental Formula I2/2 C 1/1 P2/2 M2/2-3 = 28-30. Inner upper incisors long, separated basally but convergent at the bifid tips. Outer upper incisors small and fill space between inner and canines. Lower incisors small. First upper premolar much smaller than second. Height of inner cusp is reduced in M1 and almost absent in M2 giving irregular profile. First lower premolar is caniniform and smaller than second. Second lower premolar has a single high cusp and is similar in shape to the first lower molar.
Vampyressa pusilla - Southern Yellow-eared Bat

Key to Adult Fruit Bats of Paraguay
1a Uropatagium absent.......................................................................................................................Sturnira lilium
1a Uropatagium present and well-developed, though frequently narrow...............................................................2

2a Two pairs of pale facial lines (sometimes indistinct) extending from nose leaf to the ears. No white spot on shoulder. Upper lip with normal appearance..........................................................................................................3
2b No facial lines. White spot on shoulders. Upper lip grotesquely swollen, giving impression that it is double. Eyes appear swollen. in range 36-65mm...................................................................................Pygoderma bilabiatum

3a Forearm <40mm. Facial lines pronounced. Tragus and edges of ears deep yellow.................Vampyressa pusilla
3b Forearm >40mm..............................................................................................................................................4

4a White dorsal, longitudinal line. Forearm in range 40-56mm.............................................................................5
4b No white dorsal longitudinal line. Forearm >56mm.........................................................................................6

5a White longitudinal line extends along dorsum to interscapular region only (does not reach the head). Large eyes. Uropatagium lightly furred dorsally and broad in the centre. Calcaneum c75% of foot length.............................................................................................................................................Chiroderma doriae
5b White longitudinal line usually extends onto crown. Normal eyes. Uropatagium with deep V-shape, extremely narrow in the centre. Calcaneum <50% of foot length...............................................Platyrrhinus lineatus
NB: Two further species of Platyrrhinus have been reported to occur in Paraguay but the records were considered doubtful by López-González (2005). Platyrrhinus dorsalis and P.helleri. P.helleri is the smaller than the other two species having a forearm <40mm and weight 13-16g - a previous report of the species from Departamento Caaguazú was an incorrectly identified specimen of Pygoderma bilabiatum (López-González 2005). Comparable measurements in P.lineatus are forearm >44mm and weight >18.5g (with a mean of 23g). P.lineatus overlaps in forearm measurements with P.dorsalis but is smaller in all other standard external measurements eg total length <75mm in lineatus, >75mm in dorsalis. Typically dorsalis is darker in pelage colour, being dark grey-brown to black, with lineatus ashy-brown to light brown. P.dorsalis was reported for Paraguay by Redford & Eisenberg (1992) but locality data and whereabouts of a specimen were not provided and the supporting citation (Anderson et al 1982) makes no reference to the species in Paraguay.

6a Facial stripes prominent. Forearm >70mm. Uropatagium furred dorsally with hairs contrasting with membrane colour............................................................................................................................Artibeus lituratus
6b Facial stripes weak or absent. Forearm<70mm. Uropatagium naked or very lightly haired dorsally................7

7a Forearm in range 56-64mm. Pelage typically pale.................................................................Artibeus planirostris
7b Forearm in range 60-70mm. Pelage typically dark..................................................................Artibeus fimbriatus
NB: López-González (2005) states that a fourth species, Artibeus obscurus is of possible occurrence in Paraguay though it has not yet been confirmed. A key for the species is included below but it should be borne in mind that the taxonomy and species limits of the genus Artibeus are not well understood.
7c Forearm in range 50-58mm. Pelage typically dark. Facial stripes weak or absent......................Artibeus obscurus

Subfamily Desmodontinae JA Wagner, 1840
General characteristics: Two Paraguayan species of sanguinophagous (blood-sucking) bats. Tail, calcaneous and uropatagium are reduced and almost absent. Thickened, leathery pads on elbow and muscular legs liberated from uropatagium are adaptations for walking on the ground, assisting in the silent approach to victims. The snout is short and flattened with a reduced nose-leaf (in the form of an M or U), and the lower lip equipped with a “canal” designed to accommodate blood flow from the wound to the mouth. Stomach elasticated and sac-like, specialised for absorption. Members of this family show several unique behavioural traits, including food sharing at the colony amongst unrelated individuals.
Dental characteristics: Incisors are large and triangular, much larger than reduced canines and are designed to form a shearing blade. Premolars and molars are greatly reduced and lack crushing surfaces.
Skeletal characteristics: Long bones of wings and legs are grooved to accommodate muscles. Wing digit III with three bony phalanges. Calcar rudimentary or absent in Paraguayan species.

Desmodus Wied-Neuwied, 1826: Common Vampire Bat
General characteristics: A monotypic genus of medium-sized, short-haired bats. Small, separate and pointed ears. Wings darkly pigmented except along the forearms. Interfemoral membrane is narrow and all membranes are sparsely-haired. Fossils are known from the Pleistocene of South America.
Cranial characteristics: (Fig 1) Large, smoothly-rounded braincase. Short rostrum provides support for enormous dentition.
Dental characteristics: Dental Formula I1/2 C 1/1 P1/2 M1/1 = 20. Large, triangular upper incisors armed with cutting edges completely fill the gap between the canines. Lower incisors small and bilobed- Canines large, long and pointed. Upper and lower cheek teeth are small.
Skeletal characteristics: Calcar reduced to wart-like structure. Thumb greatly elongated and robust with a short metacarpal with a rounded pad at its base and a more elongated pad under its distal half.
Desmodus rotundus - Common Vampire Bat