14.

Greater Noctule

Nyctalus lasiopterus

French: Grande Noctule / German: Riesenabendsegler / Spanish: Noctulo grande

Other common names: Giant Noctule, Greater Noctule Bat

Taxonomy. Vespertilio lasiopterus Schreber, 1780,

?Pisa (uncertain), northern Italy.

Nyctalus lasiopterus is sometimes considered to include N. aviator as a subspecies but it is here treated as distinct, based primarily on their incongruent distributions and morphological distinctions. The two together are sister to N. noctula, all three of which share the same diploid number. Monotypic.

Distribution. Scattered throughout S & E Europe from Iberian Peninsula to W Russia and extreme W Kazakhstan as well as Sicily, Cyprus, W & NE Turkey, the Caucasus, NW Iran, and N Africa, in N Morocco and NE Libya.

Descriptive notes. Head-body 84-104 mm, tail 48-74 mm, ear 18-26 mm, hindfoot 13-16 mm, forearm 59-70 mm; weight 33-74 g. The Greater Noctule is the largest species of the genus. Dorsal pelage is rufous brown (hairs unicolored), while ventral pelage is paler rufous brown and extends onto wings and interfemoral membrane, as in other noctules. Face, ears, and membranes are dark brown, and tail extends a few millimeters past uropatagium. Muzzle is short, with large glands between nostrils and eyes; ears are short and triangular, with 4-5 folds on outer edge; tragus is very short and rounded, mushroom-shaped, asis characteristic of the genus. Wings attach at ankle and calcar reaches halfwayto tail. Postcalcarial lobe is wide with a visible T-shaped piece of cartilage. Skull is large and robust and rostrum is broad; lambdoid crestis moderately developed and there is no sagittal crest. Dentition is massive; P' is small and level with cingulum of P*, being fully displaced lingually; C' and P* are in contact, covering P? lower molars are nyctalodont. Chromosomal complement has 2n = 42 and FNa = 50.

Habitat. Recorded from a variety of forested habitats including mature temperate mixed and deciduous forests throughout Europe, intact Mediterranean forest of cork oak ( Quercus suber) and Algerian oak (Q. canariensis), both Fagaceae, in Morocco, and Mediterranean cypress ( Cupressus sempervirens) and juniper ( Juniperus), both Cupressaceae, forests in Libya. Greater Noctules rely largely on mature and intact forests, as they forage high in the canopy. Although they prefer old-growth forests, they can be found in city parks with older trees in urban areas, at least in the Iberian Peninsula. Greater Noctules can be found at elevations of 0-1900 m.

Food and Feeding. Greater Noctules are primarily insectivorous and occasionally carnivorous. They are fast aerial hawkers that mainly forage high in the canopy, more than 50 m aboveground. In Spain, radar studies show that this species flies up to several hundred meters high, presumably to catch migratory birds. Flying insects make up a large portion of their diet, although some non-flying prey, such as Opiliones, have been recorded from fecal samples, which suggests that the bats must either glean occasionally or catch prey within their day roosts. Although Greater Noctules feed largely on insects, they have also been recorded feeding on nocturnally migrating small birds, e.g. European robins (Erithacus rubecula), Eurasian blue tits (Cyanistes caeruleus). Nocturnally migrating small passerines can make up to 50% oftheir diet in some regions, in autumn and spring.

Breeding. In Europe, the Greater Noctule shows restricted seasonal monoestry, with mating occurring in mid-summer and mid-autumn, before hibernation;this is evident as males develop scrotal testes from early August to October. Females give birth approximately from early May to early June; lactation occurs until early August. Litter size 1s 1-2 young; two young may be more prevalent, as in other noctule species, but no frequency data are available.

Activity patterns. Greater Noctules roost in hollow trees and bat boxes during summer, occasionally also in buildings and dry leaves of tall palms ( Washingtonia filirifera, Arecaceae). In winter, they primarily roost in hollow trees and rock crevices. They are nocturnal, coming out at dusk to forage. They will dive up to 3 m out of the roost to gain speed for flight. Greater Noctules probably hibernate through the winter in northern portions of the distribution, but further research is needed to confirm this. Searchphase call shape at higheraltitudes (more than 50 m aboveground) is a steep/shallow FM call, and only the fundamental harmonic is apparent; start frequency is 25-1 kHz; end frequency is 17-6 kHz; bandwidth is 7-5 kHz; peak frequency is 18-8 kHz; call duration is c¢.12-3 milliseconds; intercall interval is 319-7 milliseconds. At lower altitudes (less than 20 m aboveground), call shape is a steep FM/QCF call and the second harmonic is present. They occasionally use consecutive calls of alternating frequency as in the Common Noctule ( Nyctalus noctula), but only at lower altitudes. Predators include common barn-owls (7yto alba), saker falcons (Falco cherrug), and Beech Marten (Mantes foina).

Movements, Home range and Social organization. Maternity colonies are created during summer with up to 80 females and young, but where males roost is uncertain, probably alone or in small groups. Social calls are high-intensity and each sequence consists of 25 calls with tonal structure, being slightly modulated with end frequencies of 9-15 kHz (this is audible to the human ear unlike most bat calls, sounding metallic). Greater Noctules exhibit a fission-fusion society, where there is a social ranking, but members spread themselves over several roosts every night. Their foraging range may be greater than 30 km from their day roost and is usually 15-40 km, but can reach up to 130 km in a single night. They may also migrate between seasons, although this has not been investigated.

Status and Conservation. Classified as Vulnerable on The IUCN Red List. The primary threat to the Greater Noctule seems to be the logging of mature woodland and loss or disturbance of roosts. Roost destruction can cause the death of many of the individuals living there. Wind turbines are also apparently major sources of mortality to these bats. Although the species has a wide distribution,it is rather patchy, being known from few records in many regions, especially North Africa. Population densities seem to be low throughout the range and populations seem to be generally decreasing as well. The Iberian Peninsula is probably a stronghold for this species, as is northern Hungary, where a large population is present. The species may be commoner throughout its range than is currently known,as it is very hard to survey because it forages high in the canopy. In southern Spain, the introduced rose-ringed parakeet ( Psittacula kramer) is also a threat to the species as it competes for roosting places. The Greater Noctule is protected over much of its distribution, although no conservation action plans have been put forward.

Bibliography. Alcalde et al. (2016), Bec et al. (2008), Benda, Faizolahi et al. (2012), Benda, Hanaket al. (2007), Dondini & Vergari (2000), Esték & Gombkotd (2007), Estdk & Siemers (2009), Fortuna et al. (2009), Hernédndez-Brito, Carrete, Ibanez et al. (2018), Hernandez-Brito, Carrete, Popa-Lisseanu et al. (2014), Hutterer et al. (2005), Ibanez (2013), Ibanez, Guillén-Servent, Agirre-Mendi et al. (2009), Ibanez, Guillén-Servent & Bogdanowicz (2004), Ibanez, Juste et al. (2001), Kovat et al. (2011), Kovalov et al. (2019), Mikula etal. (2016), Paksuz & Ozkan (2011), Popa-Lisseanu et al. (2009), Salgueiro et al. (2007), Smirnov & Vekhnik (2013), Uhrin et al. (2006), Vernier & Vedovato (2008), Vlaschenko, Gashchak et al. (2010), Vlaschenko, Kravchenko et al. (2016), Yigit et al. (2008).