Triaenops afer, Peters, 1876

Don E. Wilson & Russell A. Mittermeier, 2019, Rhinonycteridae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 194-209 : 206-207

4.

African Trident Bat

Triaenops afer

French: Tr aen ps dAlrque German: Atrka-Die zahnblattnase Spanish: Rnonıcterıo africano

Other common names: Chewbacca Bat

Taxonomy. Triamops afar Peters, 1877 ,

“ Mombaca, ” Mombasa County, Kenya.

African populations of Triamops were considered part of TÍ persicus as two subspecies: eastern TÍ p. afar and western TÍ p. majusculus. More recently, '1Î perxiau was demonstrated to be confined to the Middle East and southern Pakistan, and all African populations are now considered to be '1Î afrr. Taxon majusrulus. originally described as larger in body size than East African populations, was shown to be in the dimensional size range of TÍ afersensu stricto and thus not a distinct taxon. Monotvpic.

Distribution. Found in two disjunct regions: largest part in savanna belt of East Africa from Ethiopia, Djibouti, and Somalia S through Kenya, Tanzania (including Pemba and Unguja in Zanzibar Archipelago), extreme SE Zambia, and E Zimbabwe to C Mozambique, with W stretch to E Central African Republic and NE DR Congo; smallest part includes SW Republic of the Congo and extreme NW Angola.

Descriptive notes. Head-body 55-72 mm, tail 25-38 mm, ear 10-16 mm, hindfoot 8-12 mm, forearm 48-60 mm; weight 8-17 g. The African Trident Bat is a large rhinonycterid. Pelage has similar tinge dorsally and ventrally; color is very variable, including beige, pale brown, brownish orange, bright orange, pale yellowish, and dark grayish brown; hairs are darker around eyes. VVng membranes are dark brown, and ears and noseleaf are pale pinkish (unpigmented) and pale gray to dark gravish brown. Noseleaf is large (width 7~7-8 -6 mm) and rounded or pentagonal, with suap-like longitudinal projection that resembles an hourglass in outline, lying across anterior part of anterior leaf, above deep anterior medial emargination. Three tall pointed projections are present on posterior part of posterior noseleaf, and its anterior part in medial position has long lanceolated projection, with sharply pointed tip. Total width of three posterior projections is c.50% the width of posterior leaf. Numerous cells, separated from each other by fleshy septa, scallop lateral parts of posterior leaf, with only one small cell in central position; lateral margins of two of three tall posterior projections are emarginated at their bases. Two supplementary leaflets occur lateral to noseleaf. Ears are short, wide, and pointed, with prominent step-like emargination on anterior margin. Second phalanx of founh wing finger bears unique uansverse bone projection into membrane c. 2~5-3 mm long. Skull has very prominent nasal swellings laterally and posteriorlv. Braincase is almost as high as rostrum and has low but long sagittal crest. Rostrum is relati\ ely short compared with congeners. Wide plates are present on dorsal sides of zygomatic bones. Greatest skull lengths are 17-8-21 -1 mm, condvlo-canine lengths are 15-6-19 -3 mm, zygomatic widths are 8 -2-9-8 mm, and upper tooth row (C -M lengths are 6-1 -7-5 mm. l’ is bilobed; C is slender with slight cingulum and large posterior secondary cusp, extending onehalf C crown height; P’ is small and extruded; C and P‘ are in or nearlv in contact; and M‘ is only little reduced, with metacone. l is bicuspid, I 2 is tricuspid and larger than I, C, is slender, Pl has about twtrthirds the crown area and onehalf the height of P‘, and M is unreduced. Baculum is gracile and c. l -6 mm long, with broad basal epiphvsis and bifurcated distal epiphysis; it has relatively broad diaphvsis (c. l 5% of baculum length), with very long and robust anns at its distal epiphysis (arm length c.0-45 mm; c.28% of baculum length) and robust proximal epiphvsis (basis width 0 -70-0-85 mm; c.45% of baculum length). Chromosomal complement has 2n 36 and FN 60. X-chromosome is medium-sized metacentiic, and Y-chromosome is small subtelocentric.

Habitat. Various habitats across central, eastern, and south-eastern Africa, including rainforests, coastal and riverine forests, dense wooded savannas, mosaics of rainforest and savanna, miombo ( Brachystegia , Fabaceae ) woodlands, bush savannas, dry Sudanian savannas, and fannlands, from sea level to elevations of c. 1700 m.

Food and Feeding. Moths are considered dominant prey of the African Trident Bat. lt is probably an aerial hawker, preying on flying insects at various heights aboveground in open spaces and among regetation.

Breeding. Reproductive cycle of the African Trident Bat probably is connected with local climatic and yegetation cycles, which differ in Southern and Northern Hemispheres. South of the equator in Africa, spennatogenesis was high in June-October and low or nonexistent in December-January; this was correlated with timing of enlargement and high secretory activity of glandular tissue of medial cell of noseleaf (“ frontal “) in males, which in and lowest in sac was highest june january. Pregnant females with one fetus each were found in early November in Kenya and late December in Tanzania. Copulations probably occur during dry season (lune-October), and births occur in November-January. There no evidence of polyestry, so time span between copulations and births suggests long gestation, with delay in fetal development or spenn storage and postponed fertilization. North of the equator in Ethiopia, pregnant females with one fetus each in late-stage development were found in early May; parturition therefore occurs around mid-May. Thus, the African Trident Bat seems, like other members of the family, to be seasonally monoestrous: females give birth to one young each year, and reproduction is timed with local climatic and vegetation cvcles.

Activity patterns. The African Trident Bat does not enter torpor or hibernate; remains actiye vear-round and forages in all seasons. t is fast and acrobatic flier. It forages at various heights, low to the ground or over water and among or above vegetation. It leaves day roosts shortly before or at sunset and forages for several hours; foraging bats were netted c. 2-3 hours after sunset. Echolocation consists of multihannonic QCF calls where pulse is composed of long CF component, combined with short FM component at end (CF-FM); terminating FM component is steep sweep of c. lo kHz. Sexual dimorphism was found in peak frequencies, with males lower than females. Calls last 5-11 milliseconds (mean 8 -5 milliseconds); interpulse intervals are 35-73 milliseconds (mean 50 -8 milliseconds); peak frequencies are 72-75 kHz (males) and 80-85 kHz (females) in Malawi and 71-75 kHz (males) and 82-85 kHz (females) in Mozambique; and end frequencies are 74-82 kHz (mean 79 kHz).

Movements, Home range and Social organization. The African Trident Bat is colonial and establishes underground colonies of a few to thousands of individuals (reportedly up to c.500,000 individuals), which can be located ven deep in caves and up to 450 m from entrances. Colonies occur in natural caves (sometimes very extensive), manmade structures (mines, cellars, and wells), and perhaps hollow trees. Colonies are mixed sex, regardless of annual reproductive cvcle. When roosting, African Trident Bats hang from ceilings alone or in groups. ln Ethiopia, African Trident Bats roosted in a large karst system with Eloquent Horseshoe Bats ( Rhinolophus eloquens), Iander`s Horseshoe Bats (R. landen), Temminck`s Myotis (Myotis tricolor), African bong-fingered Bats (Minwpmus rrfi-iranus), and Harrison’s Giant Mastiff Bats (Otomops harrisoni).

Status and Conservation. Classified as Least Concern on The IUCN Red List. Some populations of African Trident Bats are considered threatened by disturbance at roost sites and mining activities.

Bibliography. Ae en à Brosset (1968) Benda & Va o (2009) Dur¢ & Mutere (1977) Happod M (2013ab) Hıl (1982 a Kockåı Feten 1980). Lanza era (2015) Ma nova (1979) Marthews (1942) Monadıem, Sdıoeman er a (2010) Monad em Tayor eta (2010) Peters (1877) Pye (1972), Tayor eta (2005)

Figures

1. Orange Diamond-faced Bat (Rhmonícteris aurantia), 2. Persian Trident Bat (Triaenops persicus), 3. Yemeni Trident Bat (Tnaenops paruus), 4. African Trident Bat (Triaenops afer), 5. Rufous Trident Bat (Triaenops menamena), 6. Percival`s Trident Bat (Cloeotis pemivali), 7. Grandidiefs Trident Bat (Paratnaenops auritus), 8. Trouessart’s Trident Bat (Paratriaenops furcula), 9. Paulian ’ Trident Bat s (Paratriaenops pauliani)

Distribution. Found in two disjunct regions: largest part in savanna belt of East Africa from Ethiopia , Djibouti , and Somalia S through Kenya , Tanzania (including Pemba and Unguja in Zanzibar Archipelago), extreme SE Zambia , and E Zimbabwe to C Mozambique , with W stretch to E Central African Republic and NE DR Congo ; smallest part includes SW Republic of the Congo and extreme NW Angola .