Cloeotis percivali, Thomas, 1901

Don E. Wilson & Russell A. Mittermeier, 2019, Rhinonycteridae, Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions, pp. 194-209 : 207-208

6.

Percival`s Trident Bat

Cloeotis percivali

French: é he de Per va German: Pei va Kenohr-Drezahnbartna e Spanish: Rn meter dePercrva

Other common names: AtrcanTidenr Bat AfrcanTrdent nosed Bat asrAtrcanTrdent Bat Por iva s Short-cared

Bat Per va Sh fl-GBIOGTIIÖGM Bat Sh n-earedTııdent Bat

Taxonomy. Cloeotis pemivali Thomas, 1901 ,

“Takaungu, N. of Mombasa, British East Africa Kenya].

Subspecies often noted, with nominate perrivalz are smaller in body size a than a subspecies australıs and confined to Kenya (and perhaps Tanzania) vs. southern AF rica. Validity of these two subspecies remains open to question, and no revision based on well representative material is available. Monotypic.

Distribution. SE Kenya, Mafia I ( Tanzania), SE DR Congo, Zambia, Mozambique, Zimbabue, SE Botswana, NE South Africa, and Swaziland.

Descriptive notes. Head-body 30-43 mm, tail 18-36 mm, ear 7-10 mm, hindfoot 5-8 mm, forearm 31-39 mm; weight 3-6 g. Percival's Trident Bat is the smallest species in the family. Pelage usually has slightly darker tinge dorsally than ventrally, 0r venter is sometimes very pale or whitish; color very variable, including bright brownish orange, pale yellowish, pale brown, pale or dark grayish brown, and dull gray; hairs are dark brown, brownish gray, or gray around eyes, noseleaf, and mouth. Wing membranes arc dark grayish brown, and ears and noseleaf are pale pinkish (unpigrnented) to pale grayish brown. Noseleaf is small (width 3 -4-3-9 mm) and rounded, with straplike longitudinal projection that is diamond-shaped in outline and anteriorly forked and lying across anterior part of anterior leaf, above deep anterior medial emargination. Three tall pointed posterior projections are present on posterior pan of posterior leaf, and anterior part in medial position has short and narrow process, with sharply pointed tip. Total width of three posterior projections is c. 50% the width of posterior noseleaf; lateral projections have no emargination at their bases. lateral pans of posterior noseleaf are scalloped by six small cells on each side, plus pair of large cells in central position, separated from each other by fleshy septum. Two supplementary leaflets occur lateral to noseleaf. Ears are very small, rounded, and only bluntly pointed; pinna membrane is thickened. Skull has posteriorly but not laterally prominent nasal swellings. Braincase is much higher than narrow rostrum, and minute sagittal crest is developed only in frontal region. Zygomatic bones are anteriorly strongly convergent and thin, with low dorsal triangular projection on posterior margin. Greatest skull lengths are 13-136 mm, condvlo-canine lengths are 10-9-11 -3 mm, zygomatic widths are 7-7-7 mm, and upper tooth row (C —M‘) lengths are 3 -8-4-4 mm. f is bilobed; C' is slender, with slight cingulum and large posterior secondary cusp, extending about one-third the canine crown height; P‘ is small and extruded; C‘ and P‘ are in or nearly in contact; and M‘ is only little reduced, with metacone. Lower incisors are tricuspid, C, is slender, P, is about one-third to threequarters the crown area and one-half to twothirds the height of P4, and M, is unreduced. Baculum has not been described. Diploid number is 2n 40, but number of autosome arms is unknown.

Habitat. Dry woodland savannas with mopane (Colophospennum mopane, Fabaceae ) and various miombo woodlands, bushvelds, riverine woodlands, coastal forest mosaic, and others habitats from sea level to elevations of c. l 540 m.

Food and Feeding. Percival`s Trident Bats are aerial hawkers, preying on small flying insects at various heights aboveground and mostly among vegetation. Individuals were documented foraging at various water bodies, in riparian vegetation, and among woodland vegetation. Stomach contents were almost entirely small-sized adult moths (97-1 % by overall volume). All stomachs contained moths, and 86% of them contained only moths. Termites (1 %), beetles 1 -1 %), flies (0-6 %), bugs (04 %). and other items were also consumed. There was little variation in prey type consumed during the night or during annual cycles (moths were 94-7% in summer diets and 983% in winter); small-sized moths were clearly preferred over other similarly sized insects that were equally available.

Breeding. Pregnant Percival's Trident Bats, each containing one fetus, were found in October in Zimbabwe, and pregnant and lactating females with attached young were found in late November in north-eastem South Africa and early December in southern Zambia. This indicated that births occur in November and early December. Examinations of females in other seasons did not reveal any signs of ongoing reproductive activity. Hence, Percival's Trident Bat is, like other species of the family, probably seasonallv monoestrous, with females giving birth to one young each year.

Activity patterns. Percival’s Trident Bat probably does not enter torpor or hibernate. probably an acrobatic flier. leaves day roosts shortly after sunset and forages for several hours or the entire night. Percival's Trident Bat roosts in underground spaces in natural caves and human-made structures (e.g. mine tunnels and dam body corridors). lt roosts in small and narrow crevices and were once found in a cavity on the ground that was thought to be a porcupine den. Echolocation consists of multiharmonic QCF calls where pulse is composed of short CF component, combined with short FM component at end (CF-FM); tenninating FM component is steep sweep up to 29 kHz. Call is unique due to its unusually high peak frequency of first harmonic, well above 200 kHz; however, fundamental frequency often used during foraging. Call has mean durations of 1 -9 milliseconds (fundamental frequency) and 4 -6 milliseconds (first harmonic). Peak frequencies (fundamental frequencies) are 101-6-105 -3 kHz (mean 103 -4 kHz), first hannonics are 204-212 kHz (mean 207 -8 kHz). highest frequency is 212 kHz, and end (lowest) frequency is 183 kHz. Slight sexual dimorphism was detected in CF component of fundamental harmonic frequency: females ha\e, on average, lower frequency (102-9 kHz) than males (103-7 kHz). Extremely high frequency of echolocation calls is speculated to be an adaptation to more efficiently catch moths. Remains of Percivafs Trident Bats were found in small amounts of the diet of the bat hawk (Madmmmphus alnnus).

Movements, Home range and Social organization. Roosting Percival’s Trident Bats hang on cavity ceilings in loose groups, with indiriduals separated from each other by 10-20 cm. Roosting groups have 10-300 individuals, but there are no data on their sexual composition. ln Zambia, a colony of Percival's Trident Bat shared its roost (large karst cave) with colonies of Sundevalfs Leaf-nosed Bats (Hıpposidems mfƒrr), Su-iped Leaf-nosed Bats (Mammycleris vittalus), and Natal bong-fingered Bats (Mmaopterus natalmsis).

Status and Conservation. Classified as beast Ooncem on The IUCN Rad List. Percivafs Trident Bat reportedly highly sensitive to roost disturbance. In South Africa, religious ceremonies by local people are carried out in caves, and these rituals and other forms of disturbance can impact local populations and perhaps cause abandonment of roosts. ln Zambia, Percival's Trident Bat shared a cave with Striped Leaf-nosed Bats that were harvested by locals for food, which seriously disturb roosting Percival's Trident Bats. Due to observed fluctuations in numbers of roosting bats at particular roosts, such activities could cause local extirpations. ln South Africa, a national law protects Percival's Trident Bat.

Bibliography. Baona (2016) Back (1979), Bat* et a (1979) HI (1982a). Jacobs (2013) Monadiam. Res de 8r Lumsden (2007) Monad |em, Tay or era (2010) Reutenbach eta (1993). Sdwoeman G« Jacobs (2008). Seamark (2005), Taylor (1999), Thomas (1901 b), Whitaker 8r B ack (1976)

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. SE Kenya , Mafia I ( Tanzania ), SE DR Congo , Zambia , Mozambique , Zimbabue , SE Botswana , NE South Africa , and Swaziland .