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line:xlsx:hash://sha256/181a039844a33e66a35a457b7ece741051086608e425a040051b79581d606b97!/Sheet1!/L170	application/vnd.openxmlformats-officedocument.spreadsheetml.sheet	Eptesicus nilssoni	Eptesicus nilssoni	Eptesicus nilssonii	Eptesicus nilssoni	Eptesicus nilssoni	Eptesicus nilssonii	Eptesicus nilssonii	Eptesicus nilssonii	Eptesicus nilssonii	Eptesicus nilssonii	Eptesicus nilssonii	Eptesicus nilssonii	Cnephaeus nilssonii	Cnephaeus nilssonii	Cnephaeus nilssonii		[MSW2] Subgenus Eptesicus. Includes propinquus; see W. B. Davis (1965:230). Includes japonensis; see Corbet (1978c:57), but also see Yoshiyuki (1989). Includes gobiensis; see Corbet (1978c); but see also Strelkov (1986), who discussed status, relations, and distribution, and Pavlinov and Rossolimo (1987). Revised by Wallin (1969).; [MSW3] Subgenus Eptesicus. Includes propinquus; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis; see Yoshiyuki (1989). Does not include gobiensis; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbet and Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus and possibly paraphyletic with respect to that species; see Mayer and von Helversen (2001a). Specific epithet has often been spelled nilssoni, but the correct spelling is nilssonii. Placed in the subgenus Amblyotus by Horácek et al. (2000).; [HMW] Vesperus nilssonii Keyserling & Blasius, 1839 , Sweden . Eptesicus milssonii appears to be sister to E. serotinus , and they are often paraphyletic in genetic studies using mitochondrial genes due to extensive ancient hybridization. Nevertheless, when using nuclear genes, LE. nmilssonu forms a monophyletic clade not closely related to E. serotinus . Eplesicus nilssonii 1s sometimes considered to include E. japonensis , but they are generally recognized as distinct species based on morphological data. Eptesicus gobiensis has often been included under E. nilssonii , but it is usually considered a distinct species now based on genetic and morphological distinctions. There is also considerably uncertainty as to whether the north Indian endemic E. tate: represents a distinct species or a subspecies of E. nilssonii , although it is geographically well separated from E. nilssonii and is considered a distinct species here. Distribution of E. nilssonii has been reported differently in a number of sources, making actual distribution somewhat uncertain, and distribution here should be considered as tentative. Two subspecies ( nilssonii and parvus) are usually recognized, but genetic data do not support this view because there is low genetic variability between these two taxa. Monotypic.; [batnames2022] Subgenus Cnephaeus .  Includes propinquus ; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis ; see Yoshiyuki (1989). Does not include gobiensis ; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbetand Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus andpossibly paraphyletic with respect to that species; see Mayer and von Helversen (2001 a ). Specific epithet has often been spelled nilssoni , but the correct spelling is nilssonii . Placed in the subgenus Amblyotus by Horácek et al. (2000).; [IUCN] Some authors consider Eptesicus japonensis to be a synonym of E. nilssonii , but here we consider it to be a separate species (see Simmons 2005 and references therein).; [batnames2023] Subgenus Cnephaeus .  Includes propinquus ; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis ; see Yoshiyuki (1989). Does not include gobiensis ; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbetand Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus andpossibly paraphyletic with respect to that species; see Mayer and von Helversen (2001 a ). Specific epithet has often been spelled nilssoni , but the correct spelling is nilssonii . Placed in the subgenus Amblyotus by Horácek et al. (2000).; [MDD2025_2.0] moved from Eptesicus to Cnephaeus; [batnames2025_1.7] Includes propinquus; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis; see Yoshiyuki (1989). Does not include gobiensis; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbetand Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus andpossibly paraphyletic with respect to that species; see Mayer and von Helversen (2001a). Specific epithet has often been spelled nilssoni, but the correct spelling is nilssonii. Placed in the subgenus Amblyotus by Horácek et al. (2000).; [MDD2025_2.2] moved from Eptesicus to Cnephaeus				propinquus, japonensis		atratus, centralasiaticus, gobiensis, japonensis, kashgaricus, parvus, propinquus.	nilssoni, gobiensis, kashgaricus, centrasiaticus, parvus, japonensis	nilssonii, parvus	atratus, borealis, kuhli, propinquus			nilssonii, parvus	nilssonii - atratus, borealis, kuhli, propinquus	borealis, kuhli, nilssonii, atratus, propinquus, parvus	Some authors consider Eptesicus japonensis to be a synonym of E. nilssonii , but here we consider it to be a separate species (see Simmons 2005 and references therein).	nilssonii, parvus	nilssonii - atratus, borealis, kuhli, propinquus	borealis, kuhli, nilssonii, atratus, propinquus, parvus	borealis, kuhlii, nilssonii, nilsonii, atratus, propinquus, nilssoni, parvus, kuhli, varangus	nilssonii, parvus	nilssonii - atratus, borealis, kuhli, propinquus 	borealis (Nilsson, 1838) [preoccupied]|kuhlii (Nilsson, 1838) [preoccupied]|nilssonii (von Keyserling & J. H. Blasius, 1839)|nilsonii (Lesson, 1842) [incorrect subsequent spelling]|atratus (F. A. Kolenati, 1858)|propinquus (W. C. H. Peters, 1872)|nilssoni (G. S. Miller, 1897) [incorrect subsequent spelling]|parvus (Kishida, 1932)|kuhli (Simmons, 2005) [incorrect subsequent spelling | not used as valid]|varangus (Lopatin, 2023)		Corbet, G.B. and Hill, J.E. 1980. A World List of Mammalian Species. British Museum (Natural History), London, 226 pp.	Northern bat	France, Norway – E Siberia, Japan, Iraq, Tibet	Honacki, J.H., Kinman, K.E. and Koeppl, J.W. 1982. Mammal Species of the World: A Taxonomic and Geographic Reference. Allen Press, Lawrence, 694 pp.	Eptesicus nilssoni	Sweden.	Keyserling and Blasius	1839	Arch. Naturgesch., 5(1):315.	Distribution: Ranging across the northern Palearctic from western Europe to Japan, south to northern Iran, Nepal, and western China.		Corbet, G.B. and Hill, J.E. 1991. A World List of Mammalian Species. Third edition. Oxford University Press, London, 243 pp. ISBN 0-19-854017-5	Northern bat	France, Norway – E. Siberia – Japan, N India	Koopman, K.F. 1993. Order Chiroptera. Pp. 137–242 in Wilson, D.E. and Reeder, D.M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference. Second edition. Smithsonian Institution Press, Washington, 1206 pp.	Keyserling and Blasius	1839	Arch. Naturgesch., 5(1):315.	Subgenus Eptesicus. Includes propinquus; see W. B. Davis (1965:230). Includes japonensis; see Corbet (1978c:57), but also see Yoshiyuki (1989). Includes gobiensis; see Corbet (1978c); but see also Strelkov (1986), who discussed status, relations, and distribution, and Pavlinov and Rossolimo (1987). Revised by Wallin (1969).	W and E Europe to E Siberia and NW China; north beyond Arctic Circle in Scandinavia, south to Bulgaria, Iraq, the Elburz Mtns (N Iran), The Pamirs and W China (not Tibet); Nepal; Honshu, Hokkaido (Japan); Sakhalin Isl (Russia).	Sweden.		KEYSERLING & BLASIUS	1839	Rostrum of medium length but fairly broad; not flattened dorsally. Inner upper incisor bicuspid and outer upper incisor well developed. Last upper molar well developed. Braincase fairly low. Basicranial pits well developed. Size medium (forearm length, 37-43 mm).	Distribution: Ranging across the northern Palearctic from western Europe to Japan, south to northern Iran, Nepal, and western China.	Six subspecies are currently recognized:	E. n. nilssoni (western Europe at least to central and perhaps to eastern Siberia), E. n. gobiensis (Iran and Mongolia, perhaps to southeastern Siberia), E. n. kashgaricus (Afghanistan to northwestern China), E. n. centrasiaticus (west-central China), E. n. parvus (Korea and perhaps Sakhalin and Hokkaido in Japan), E. n. japonensis (Honshu in Japan). Subspecies boundaries, however, are far from certain.	119	species	E. nilssoni	KEYSERLING & BLASIUS	1839	Eptesicus	subgenus	Eptesicus nilssoni				Rostrum of medium length but fairly broad; not flattened dorsally. Inner upper incisor bicuspid and outer upper incisor well developed. Last upper molar well developed. Braincase fairly low. Basicranial pits well developed. Size medium (forearm length, 37-43 mm).	Six subspecies are currently recognized:		1. E. nilssoni (KEYSERLING & BLASIUS 1839) [nilssoni group].	1	_C. n. nilssonii_ (Keyserling & Blasius, 1839) (synonyms: _atratus_ (Kolenati, 1858), _borealis_ (Nilsson, 1838), _kuhlii_ (Nilsson, 1838), _propinquus_ (Peters, 1872)); _C. n. parvus_ (Kishida, 1932); _C. n. varangus_ (Лопатин, 2023) (fossil)			Don E. Wilson & DeeAnn M. Reeder (editors). 2005. Mammal Species of the World. A Taxonomic and Geographic Reference (3rd ed), Johns Hopkins University Press, 2,142 pp. (Available from Johns Hopkins University Press, 1-800-537-5487 or (410) 516-6900, or at http://www.press.jhu.edu).	CHIROPTERA	Vespertilionidae	Vespertilioninae	Eptesicini	Eptesicus nilssonii	Eptesicus	Eptesicus	nilssonii	Keyserling and Blasius	y	1839		Arch. Naturgesch.	5	1	315		Northern Bat	Sweden.	W and E Europe to E Siberia and NW China; north beyond Arctic Circle in Scandinavia, south to Bulgaria, Iraq, the Elburz Mtns (N Iran), The Pamirs and W China (not Tibet); Korea; Hokkaido (Japan); Sakhalin Isl (Russia).	IUCN 2003 and IUCN/SSC Action Plan (2001) – Lower Risk (lc).	atratus Kolenati, 1858; borealis Nilsson 1838 [not Müller, 1776]; kuhli Nilsson 1836 [not Kuhl, 1819]; propinquus Peters, 1872; parvus Kishida, 1932.	Subgenus Eptesicus. Includes propinquus; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis; see Yoshiyuki (1989). Does not include gobiensis; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbet and Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus and possibly paraphyletic with respect to that species; see Mayer and von Helversen (2001a). Specific epithet has often been spelled nilssoni, but the correct spelling is nilssonii. Placed in the subgenus Amblyotus by Horácek et al. (2000).	4C3D87E8FFA66A1AFA84921F16BCBAC9	Handbook of the Mammals of the World – Volume 9 Bats, Barcelona: Lynx Edicions	978-84-16728-19-0	hbmw_9_Vespertilionidae_716.pdf.imf	hash://md5/b004ff90fffb6a44fffc96591e00bb32	851	zip:hash://sha256/ec5fd314a06aba1a7b0b72f23e54ac625ae272bd98f82f1d01f4c09627d9e8e0!/treatments-xml-main/data/4C/3D/87/4C3D87E8FFA66A1AFA84921F16BCBAC9.xml	Eptesicus nilssonii	Vespertilionidae	Eptesicus	nilssonii		1839	Sérotine de Nilsson @fr | Nordfledermaus @de | Murciélagohortelano nortefo @es | Northern Bat @en	Vesperus nilssonii Keyserling & Blasius, 1839 , Sweden . Eptesicus milssonii appears to be sister to E. serotinus , and they are often paraphyletic in genetic studies using mitochondrial genes due to extensive ancient hybridization. Nevertheless, when using nuclear genes, LE. nmilssonu forms a monophyletic clade not closely related to E. serotinus . Eplesicus nilssonii 1s sometimes considered to include E. japonensis , but they are generally recognized as distinct species based on morphological data. Eptesicus gobiensis has often been included under E. nilssonii , but it is usually considered a distinct species now based on genetic and morphological distinctions. There is also considerably uncertainty as to whether the north Indian endemic E. tate: represents a distinct species or a subspecies of E. nilssonii , although it is geographically well separated from E. nilssonii and is considered a distinct species here. Distribution of E. nilssonii has been reported differently in a number of sources, making actual distribution somewhat uncertain, and distribution here should be considered as tentative. Two subspecies ( nilssonii and parvus) are usually recognized, but genetic data do not support this view because there is low genetic variability between these two taxa. Monotypic.	E France to N Europe well above the Arctic Circle in Norway , Sweden , and Finland , Central Europe, Caucasus, NW Iran , and Central Asia (including N Kazakhstan ) through Russia and N Mongolia to Russian Far East including Sakhalin I and Kamchatka Peninsula and S through Amur region into NC & NE China ( Inner Mongolia [= Nei Mongol ], Heilongjiang , Jilin , and Shandong ) and Korean Peninsula, including Hokkaido and Okushirito inJapan and Kuril Is (Kunashir and Iturup); there are occasionally vagrant records throughout Europe as far S as Bulgaria and Italy and W into S Britain.	Head-body 54-68 mm, tail 35-50 mm, ear 13-17-5 mm, hindfoot 10-12 mm, forearm 37-1-44-2 mm; weight 9-13 g. Fur of the Northern Serotine is long, silky, and shaggy. Dorsal pelage is dark brown to blackish and frosted with goldyellow tips, giving glossy yellowish appearance. Ventral pelage is lighter pale brown to beige. Bare face, ears, and membranes are distinctly dark brown. Ears are short, rounded, and fleshy, with five transverse folds; tragus is short and rounded, curving inward. Wings are broad and long (wingspans of 240-280 mm) and are attached to base of toes. Tail extends a little past uropatagium, and calcaris about halfway to tail; there is no postcalcarial lobe. Baculum is short (1-3-1-5 mm), relatively long for Eptesicus , and Y-shaped, and it has deep basal bifurcation, with either side going outward from shaft. Skull is relatively delicate and tapered; rostrum is laterally rounded; condylobasal lengths are 14-2—-15-6 mm; zygomatic breadths are 9-5-10-5 mm; braincase is low and subspherical; sagittal crest is very low or absent; lambdoidal crests curve slightly toward point of contact at middle; interorbital region is hourglass-shaped; and lacrimal swelling is present. Dentition is robust; I? is bicuspid; I is unicuspid and more than one-half the height of I’; and lower molars are myotodont. Chromosomal complement has 2n = 50 and FNa = 48 (throughout Europe).	Boreal habitats and high mountain ecosystems, mostly forests, but also desert regions, lowland, and foothills from sea level up to elevations of ¢. 2920 m . The Northern Serotine is the most northern distributed bat species. Due to its fast flight, it generally needs open areas to hunt such as open woodlands or forest edges, parks and gardens in urban areas,lakes, wetlands, meadows or swamps in high mountains.	The Northern Serotine forages along vegetation edges, forest edges or hedgerows in natural landscapes and around streetlamps in urbanized areas. It is adapted to hunt insects at just above freezing temperatures. It has been reported feeding primarily on small Diptera (specifically Nematocera), but it also captures Coleoptera , Lepidoptera , and Heteroptera in smaller quantities. It generally hunts by aerial-hawking, capturing prey in flight at heights of 2-20 m rather than capturing them on vegetation or the ground. Nevertheless, because it is not clutter specialist,it is capable of visually detecting and recognizing large (greater than 50 mm ) and brightly colored insects on the ground and capturing them using visual and acoustic senses. In someareas, it tends to select areas with streetlamps. In localities close to the ocean, Northern Serotines and Eurasian Serotines ( E. serotinus ) have been observed foraging over the ocean where insects are abundant.	Pregnant Northern Serotines give birth to 1-2 young from mid-June to end ofJuly, occurring earlier at lower latitudes. In Germany , births usually occur in mid-June, but in Scandinavia and further north, births occur in late June to early July, or even in midand late July in some regions. This later birth date might be attributed to females entering torpor more often during the day in cooler weather to conserve energy while pregnant, which would delay fetal development. Young startto fly atjust two ( Sweden ) or three ( Germany ) weeks of age. Females become sexually active during their first year but do nottypically breed during theirfirst three years. Males disperse more widely than females during theirfirst year, and females do not change maternity colonies very often. The Northern Serotine is one of the few bat species that has been found breeding above the Arctic Circle. Maximum longevity in the wild has been recorded at c.20 years.	Nightly activity of the Northern Serotine is generally bimodal, with two activity peaks after sunset and around dawn. Pregnant females usually have unimodal or bimodal activity pattern, with peaks always after dusk and sometimes before dusk; non-pregnant females forage intermittently throughout the night. Reproducing females at 57° N left the roost 47 minutes after sunset and spent an average of three hours outside the maternity roost each night. During pregnancy, however, ¢.20% of their time outside the maternity colony was spent roosting in trees, foraging for an average of 150 minutes each night. Lactating females foraged an average of 120-130 minutes/night just after giving birth, 220-260 minutes/night in mid-lactation, and 320 minutes/night at peak lactation, reflecting energetic costs of lactation. Female Northern Serotines tend to emerge increasingly later as pregnancy advances. Due to the cold climate in its distribution, swarming period occurs from mid-July into late autumn. Hibernation tends to occur from November or December until March or April. Northern Serotines typically choose hibernacula that have ambient temperatures of 0-2°C but will occasionally use roosts that get as cold as =5°C, roosting in houses, cellars, and sometimes caves and mines. Maternity colonies in summer are commonly located in buildings but sometimes in hollow trees. In most ofits distribution, nursery colonies are found in roofs of houses, walls, or empty large cavities below metal sheets. Search calls of the Northern Serotine are not well differentiated from the Eurasian Serotine and have broad overlap with some species of Nyctalus . Pulses are up to 20 milliseconds long and typically shaped as an FM/QCF sweep. Peak frequencies are usually 26-32 kHz. On Hokkaido , Japan , start frequencies were 51-8-64-7 kHz, end frequencies were 24-3-27.7 kHz, peak frequencies were 28-3-31-6 kHz, durations were 4-4-8 milliseconds, and interpulse intervals were 95-185 milliseconds. Swedish recordings had an average peak frequency of 32-4 kHz, end frequency of 30-4 kHz, duration of 7 milliseconds, and interpulse interval of 100 milliseconds. Social calls seem to be quite distinct, consisting of long QCF sweep, with duration of up to 60 milliseconds and powerful second and third harmonics.	In winter, the Northern Serotine tends to move toward its hibernation sites. During hibernation, only single individuals have been found. Although it shifts roosts quite often between seasons, these movements are rather short, and maximum migratory distance is ¢. 450 km in Europe. A few banded and recaptured individuals suggest that it might be a short-distance migratory species, only occasionally flying long distances. During the breeding season, females form maternity colonies. Maternity colonies can contain tens to hundreds of individuals (very large colonies have been reported in Japan ). Single individuals, males, or non-reproductive females can be found roosting in a wide variety of different roosts. Males commonly roost alone year-round, rarely in small groups. Northern Serotines can fly up to 70 km each night to reach foraging areas, but they typically commute only 1-10 km, using up to eight different sites. Foraging areas range from 0-2 km? in summer to 66 km ? during swarming and are strongly affected by length of the night (0-91-17-2 km?). Females often establish small feeding territories (c. 100 m ?) with abundant insects, where they return every night, sometimes for subsequent years. Feeding territories of males and females are vigorously defended against intruders, involving aerial chases and audible vocalizations with maximum amplitudes of 14 kHz. These same territorial behaviors have been observed outside of hibernacula in autumn, suggesting that roosts are also defended. Foraging localities and activity peaks might be partially affected by competition with other bat species that forage in open spaces but do not forage at the same time, e.g. Eurasian Particolored Bats ( Vespertilio murinus ), Common Noctules ( Nyctalus noctula ), and Eurasian Serotines. Species that used narrow spaces foraged simultaneously with Northern Serotines (e.g. Daubenton’s Myotis , Myotis daubentonii ). Population density decreases at higherlatitudes, with densities at 57° N being five times higher than in similar landscapes at 65° N .	Classified as Least Concern on The IUCN Red List. The Northern Serotine is quite common, occurs in high densities in several places in its distribution, and is widespread, sometimes considered the most abundant bat species in northern countries. Nevertheless, in some areas, they are very rare and locally endangered (e.g. Germany ). Climate change is the most acknowledged threat to the Northern Serotine because much its current distribution is within northern habitats that will be heavily altered by increasing temperatures.	Abe et al. (2005) | Ahlén et al. (2009) | Artyushin, Bannikova et al. (2009) | Artyushin, Kruskop et al. (2018) | Artyushin, Lebedev, Bannikova & Kruskop (2012) | Benda et al. (2012) | Corbet (1978) | Coroiu (2016b) | Dietz & Kiefer (2016) | Duvergé et al. (2000) | EkIof et al. (2002) | Frafjord (2013) | Fukui et al. (2004) | Gerell & Rydell (2001) | Haupt & Schmidt (2007) | Haupt et al. (2006) | Hutterer et al. (2005) | Jensen et al. (2001) | Jére et al. (2018) | Jo Yeong-Seok et al. (2018) | de Jong (1994) | Juste, Benda et al. (2013) | Juste, Bilgin et al. (2009) | Kawai & Helgen (2011) | Keyserling & Blasius, (1839) | Lu¢an (2007) | Ohdachi et al. (2009) | Pacifici et al. (2013) | Pavlini¢ & Tvrtkovié (2003) | Rydell (1992a, 1993a, 1993b, 1993c, 1998, 1999) | Rydell & EkI6f (2003) | Rydell et al. (1994) | Siivonen & Wermundsen (2008) | Simmons (2005) | Smith & Xie Yan (2008) | Spitzenberger (2002) | Tress (1994) | Vintulis & Pétersons (2014) | Volleth et al. (2001) | Yoshiyuki (1989)	https://zenodo.org/record/6398226/files/figure.png	197. Northern Serotine Eptesicus nilssonii French: Sérotine de Nilsson / German: Nordfledermaus / Spanish: Murciélago hortelano nortefo Other common names: Northern Bat Taxonomy. Vesperus nilssonii Keyserling & Blasius, 1839 , Sweden . Eptesicus milssonii appears to be sister to E. serotinus , and they are often paraphyletic in genetic studies using mitochondrial genes due to extensive ancient hybridization. Nevertheless, when using nuclear genes, LE. nmilssonu forms a monophyletic clade not closely related to E. serotinus . Eplesicus nilssonii 1s sometimes considered to include E. japonensis , but they are generally recognized as distinct species based on morphological data. Eptesicus gobiensis has often been included under E. nilssonii , but it is usually considered a distinct species now based on genetic and morphological distinctions. There is also considerably uncertainty as to whether the north Indian endemic E. tate: represents a distinct species or a subspecies of E. nilssonii , although it is geographically well separated from E. nilssonii and is considered a distinct species here. Distribution of E. nilssonii has been reported differently in a number of sources, making actual distribution somewhat uncertain, and distribution here should be considered as tentative. Two subspecies ( nilssonii and parvus) are usually recognized, but genetic data do not support this view because there is low genetic variability between these two taxa. Monotypic. Distribution. E France to N Europe well above the Arctic Circle in Norway , Sweden , and Finland , Central Europe, Caucasus, NW Iran , and Central Asia (including N Kazakhstan ) through Russia and N Mongolia to Russian Far East including Sakhalin I and Kamchatka Peninsula and S through Amur region into NC & NE China ( Inner Mongolia [= Nei Mongol ], Heilongjiang , Jilin , and Shandong ) and Korean Peninsula, including Hokkaido and Okushirito inJapan and Kuril Is (Kunashir and Iturup); there are occasionally vagrant records throughout Europe as far S as Bulgaria and Italy and W into S Britain. Descriptive notes. Head-body 54-68 mm, tail 35-50 mm, ear 13-17-5 mm, hindfoot 10-12 mm, forearm 37-1-44-2 mm; weight 9-13 g. Fur of the Northern Serotine is long, silky, and shaggy. Dorsal pelage is dark brown to blackish and frosted with goldyellow tips, giving glossy yellowish appearance. Ventral pelage is lighter pale brown to beige. Bare face, ears, and membranes are distinctly dark brown. Ears are short, rounded, and fleshy, with five transverse folds; tragus is short and rounded, curving inward. Wings are broad and long (wingspans of 240-280 mm) and are attached to base of toes. Tail extends a little past uropatagium, and calcaris about halfway to tail; there is no postcalcarial lobe. Baculum is short (1-3-1-5 mm), relatively long for Eptesicus , and Y-shaped, and it has deep basal bifurcation, with either side going outward from shaft. Skull is relatively delicate and tapered; rostrum is laterally rounded; condylobasal lengths are 14-2—-15-6 mm; zygomatic breadths are 9-5-10-5 mm; braincase is low and subspherical; sagittal crest is very low or absent; lambdoidal crests curve slightly toward point of contact at middle; interorbital region is hourglass-shaped; and lacrimal swelling is present. Dentition is robust; I? is bicuspid; I is unicuspid and more than one-half the height of I’; and lower molars are myotodont. Chromosomal complement has 2n = 50 and FNa = 48 (throughout Europe). Habitat. Boreal habitats and high mountain ecosystems, mostly forests, but also desert regions, lowland, and foothills from sea level up to elevations of ¢. 2920 m . The Northern Serotine is the most northern distributed bat species. Due to its fast flight, it generally needs open areas to hunt such as open woodlands or forest edges, parks and gardens in urban areas,lakes, wetlands, meadows or swamps in high mountains. Food and Feeding. The Northern Serotine forages along vegetation edges, forest edges or hedgerows in natural landscapes and around streetlamps in urbanized areas. It is adapted to hunt insects at just above freezing temperatures. It has been reported feeding primarily on small Diptera (specifically Nematocera), but it also captures Coleoptera , Lepidoptera , and Heteroptera in smaller quantities. It generally hunts by aerial-hawking, capturing prey in flight at heights of 2-20 m rather than capturing them on vegetation or the ground. Nevertheless, because it is not clutter specialist,it is capable of visually detecting and recognizing large (greater than 50 mm ) and brightly colored insects on the ground and capturing them using visual and acoustic senses. In someareas, it tends to select areas with streetlamps. In localities close to the ocean, Northern Serotines and Eurasian Serotines ( E. serotinus ) have been observed foraging over the ocean where insects are abundant. Breeding. Pregnant Northern Serotines give birth to 1-2 young from mid-June to end ofJuly, occurring earlier at lower latitudes. In Germany , births usually occur in mid-June, but in Scandinavia and further north, births occur in late June to early July, or even in midand late July in some regions. This later birth date might be attributed to females entering torpor more often during the day in cooler weather to conserve energy while pregnant, which would delay fetal development. Young startto fly atjust two ( Sweden ) or three ( Germany ) weeks of age. Females become sexually active during their first year but do nottypically breed during theirfirst three years. Males disperse more widely than females during theirfirst year, and females do not change maternity colonies very often. The Northern Serotine is one of the few bat species that has been found breeding above the Arctic Circle. Maximum longevity in the wild has been recorded at c.20 years. Activity patterns. Nightly activity of the Northern Serotine is generally bimodal, with two activity peaks after sunset and around dawn. Pregnant females usually have unimodal or bimodal activity pattern, with peaks always after dusk and sometimes before dusk; non-pregnant females forage intermittently throughout the night. Reproducing females at 57° N left the roost 47 minutes after sunset and spent an average of three hours outside the maternity roost each night. During pregnancy, however, ¢.20% of their time outside the maternity colony was spent roosting in trees, foraging for an average of 150 minutes each night. Lactating females foraged an average of 120-130 minutes/night just after giving birth, 220-260 minutes/night in mid-lactation, and 320 minutes/night at peak lactation, reflecting energetic costs of lactation. Female Northern Serotines tend to emerge increasingly later as pregnancy advances. Due to the cold climate in its distribution, swarming period occurs from mid-July into late autumn. Hibernation tends to occur from November or December until March or April. Northern Serotines typically choose hibernacula that have ambient temperatures of 0-2°C but will occasionally use roosts that get as cold as =5°C, roosting in houses, cellars, and sometimes caves and mines. Maternity colonies in summer are commonly located in buildings but sometimes in hollow trees. In most ofits distribution, nursery colonies are found in roofs of houses, walls, or empty large cavities below metal sheets. Search calls of the Northern Serotine are not well differentiated from the Eurasian Serotine and have broad overlap with some species of Nyctalus . Pulses are up to 20 milliseconds long and typically shaped as an FM/QCF sweep. Peak frequencies are usually 26-32 kHz. On Hokkaido , Japan , start frequencies were 51-8-64-7 kHz, end frequencies were 24-3-27.7 kHz, peak frequencies were 28-3-31-6 kHz, durations were 4-4-8 milliseconds, and interpulse intervals were 95-185 milliseconds. Swedish recordings had an average peak frequency of 32-4 kHz, end frequency of 30-4 kHz, duration of 7 milliseconds, and interpulse interval of 100 milliseconds. Social calls seem to be quite distinct, consisting of long QCF sweep, with duration of up to 60 milliseconds and powerful second and third harmonics. Movements, Home range and Social organization. In winter, the Northern Serotine tends to move toward its hibernation sites. During hibernation, only single individuals have been found. Although it shifts roosts quite often between seasons, these movements are rather short, and maximum migratory distance is ¢. 450 km in Europe. A few banded and recaptured individuals suggest that it might be a short-distance migratory species, only occasionally flying long distances. During the breeding season, females form maternity colonies. Maternity colonies can contain tens to hundreds of individuals (very large colonies have been reported in Japan ). Single individuals, males, or non-reproductive females can be found roosting in a wide variety of different roosts. Males commonly roost alone year-round, rarely in small groups. Northern Serotines can fly up to 70 km each night to reach foraging areas, but they typically commute only 1-10 km, using up to eight different sites. Foraging areas range from 0-2 km? in summer to 66 km ? during swarming and are strongly affected by length of the night (0-91-17-2 km?). Females often establish small feeding territories (c. 100 m ?) with abundant insects, where they return every night, sometimes for subsequent years. Feeding territories of males and females are vigorously defended against intruders, involving aerial chases and audible vocalizations with maximum amplitudes of 14 kHz. These same territorial behaviors have been observed outside of hibernacula in autumn, suggesting that roosts are also defended. Foraging localities and activity peaks might be partially affected by competition with other bat species that forage in open spaces but do not forage at the same time, e.g. Eurasian Particolored Bats ( Vespertilio murinus ), Common Noctules ( Nyctalus noctula ), and Eurasian Serotines. Species that used narrow spaces foraged simultaneously with Northern Serotines (e.g. Daubenton’s Myotis , Myotis daubentonii ). Population density decreases at higherlatitudes, with densities at 57° N being five times higher than in similar landscapes at 65° N . Status and Conservation. Classified as Least Concern on The IUCN Red List. The Northern Serotine is quite common, occurs in high densities in several places in its distribution, and is widespread, sometimes considered the most abundant bat species in northern countries. Nevertheless, in some areas, they are very rare and locally endangered (e.g. Germany ). Climate change is the most acknowledged threat to the Northern Serotine because much its current distribution is within northern habitats that will be heavily altered by increasing temperatures. Bibliography. Abe et al. (2005), Ahlén et al. (2009), Artyushin, Bannikova et al. (2009), Artyushin, Kruskop et al. (2018), Artyushin, Lebedev, Bannikova & Kruskop (2012), Benda et al. (2012), Corbet (1978), Coroiu (2016b), Dietz & Kiefer (2016), Duvergé et al. (2000), EkIof et al. (2002), Frafjord (2013), Fukui et al. (2004), Gerell & Rydell (2001), Haupt & Schmidt (2007), Haupt et al. (2006), Hutterer et al. (2005), Jensen et al. (2001), Jére et al. (2018), Jo Yeong-Seok et al. (2018), de Jong (1994), Juste, Benda et al. (2013), Juste, Bilgin et al. (2009), Kawai & Helgen (2011), Keyserling & Blasius, (1839), Lu¢an (2007), Ohdachi et al. (2009), Pacifici et al. (2013), Pavlini¢ & Tvrtkovié (2003), Rydell (1992a, 1993a, 1993b, 1993c, 1998, 1999), Rydell & EkI6f (2003), Rydell et al. (1994), Siivonen & Wermundsen (2008), Simmons (2005), Smith & Xie Yan (2008), Spitzenberger (2002), Tress (1994), Vintulis & Pétersons (2014), Volleth et al. (2001), Yoshiyuki (1989).	Simmons, N.B. and A.L. Cirranello. 2022B. Bat Species of the World: A taxonomic and geographic database. Accessed on 10/11/2022.	Vespertilionidae	Eptesicus nilssonii	Eptesicus	Cnephaeus	nilssonii	Keyserling & Blasius	1839	1	Arch. Naturgesch.	5(1): 315	Northern Bat	 atratus Kolenati, 1858; borealis Nilsson 1838 [not M&uuml;ller, 1776]; kuhli Nilsson 1836 [not Kuhl, 1819]; propinquus Peters, 1872; <b> parvus </b> Kishida, 1932.	Sweden.	W and E Europe to E Siberia and NW China; north beyond Arctic Circle in Scandinavia, south to Bulgaria, Iraq, the Elburz Mtns (N Iran), The Pamirs and W China (not Tibet); Korea; Hokkaido (Japan); Sakhalin Isl (Russia).	Not listed.	Least Concern	Subgenus Cnephaeus .  Includes propinquus ; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis ; see Yoshiyuki (1989). Does not include gobiensis ; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbetand Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus andpossibly paraphyletic with respect to that species; see Mayer and von Helversen (2001 a ). Specific epithet has often been spelled nilssoni , but the correct spelling is nilssonii . Placed in the subgenus Amblyotus by Horácek et al. (2000).	Mammal Diversity Database. (2023). Mammal Diversity Database (Version 1.11) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.7830771 released 15 April 2023	Eptesicus nilssonii	23	Northern Serotine	Northern Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	VESPERTILIONIFORMES	NA	NA	VESPERTILIONOIDEA	VESPERTILIONIDAE	VESPERTILIONINAE	EPTESICINI	Eptesicus	Cnephaeus	nilssonii	Keyserling & Blasius	1839	1						Sweden.			borealis (Nilsson, 1838) [preoccupied]|kuhli (Nilsson, 1836) [preoccupied]|nilssonii (Keyserling & Blasius, 1839)|atratus (Kolenati, 1858)|propinquus (W. Peters, 1872)|parvus Kishida, 1932	NA	NA	France|Germany|Switzerland|Liechtenstein|Italy|Austria|Czech Republic|Hungary|Slovakia|Slovenia|Romania|Ukraine|Poland|Belarus|Lithuania|Latvia|Estonia|Norway|Sweden|Finland|Russia|Armenia|Azerbaijan|Iran|Kazakhstan|Kyrgyzstan?|Mongolia|China|Japan|North Korea|South Korea	Asia|Europe	Palearctic	LC	0	0	0	Eptesicus_nilssonii	0	sciname match	Eptesicus_nilssonii	0	IUCN. 2022. The IUCN Red List of Threatened Species. Version 2022-1. https://www.iucnredlist.org. Accessed on [28 September, 2022].	7910	Eptesicus nilssonii	ANIMALIA	CHORDATA	MAMMALIA	CHIROPTERA	VESPERTILIONIDAE	Eptesicus	nilssonii	(Keyserling &; Blasius, 1839)	Some authors consider Eptesicus japonensis to be a synonym of E. nilssonii , but here we consider it to be a separate species (see Simmons 2005 and references therein).	20000000	Eptesicus nilssonii	Least Concern		2016	2016-04-25 00:00:00 UTC	3.1	English	Listed as Least Concern because this species is widespread and abundant. No decline in population size has been detected, and there are no known widespread major threats.	A fast flying nocturnal bat species, found in a variety of habitats from mountain taiga to desert. It forages in open areas of diverse habitats, including woodland edge (or above woodland), small-scale farmland, parks and gardens with trees (van der Kooij in litt. 2006), over lakes and rivers and at street lights. It is also found in river valleys where it can remain by a source of freshwater, roosting in tree holes and crevices. In autumn, it forages and displays in high mountains above the tree line (Spitzenberger 2002). Its diet comprises small insects such as Diptera. Summer roosts are located mainly in houses, occasionally in tree holes. It may change roost sites during summer. Winter roosts are found mainly in houses, cellars, and natural and artificial underground habitats. In winter the species roosts singly or in small groups of 2-4 individuals. Long-distance movements of up to 450 km have been recorded by Tress (1994). In Mongolia it hibernates from November to December until March or April, and although it does not migrate, it may shift roosts several times over seasons .	There are no major threats to this widespread species. There are localized threats in some parts of its range but these are not having a significant impact on the species overall.	A widespread and common species over much of its range in Europe, indeed the most abundant bat species in the north (Rydell 1999). Summer maternity colonies usually number 10-100 females. In Mongolia, there have been no population estimates conducted, but the species is believed to be evenly distributed and not rare (M. Stubbe pers. comm.). In Japan, colonies of over 100 individuals can be found.	Stable	This is a widespread Palaearctic species that occurs from France and Norway through northern and central Europe and Asia, east to the Pacific seaboard and northern Japan (found only on Hokkaido (Abe ;et al. ;2005), Rebun, and Rishiri). In Europe, it occurs north to well above the Arctic Circle, but is absent or occasional in the west (the Low Countries, UK, western France, Iberia), and is scarce in the mountains of southern Europe (occurs from southern France across northern Italy and there are scattered occurrences in the Balkans). It has been recorded once from Iran although the population there is marginal to the species' range (M. Sharifi pers. comm. 2005). In Mongolia (subspecies E. n. nilssonii ), it is found throughout the north of the country, in forested areas in north-western Mongol Altai Mountain Range, Hövsgöl, Hangai and Hentii mountain ranges, Mongol Daguur Steppe and Eastern Mongolia. It occurs from sea level up to 2,300 m Asl (van der Kooij in litt. 2006).		Terrestrial	It is protected by national legislation in most range states in Europe. There are also international legal obligations for its protection through the Bonn Convention (Eurobats) and Bern Convention in parts of range where these apply. It is included in Annex IV of EU Habitats and Species Directive, and there is some habitat protection through Natura 2000. It occurs in several protected areas throughout its range.	Palearctic		FALSE	FALSE	Global	Simmons, N. B., & Cirranello, A. L. (2023). Batnames.org Species List Version 1.4 (1.4). Zenodo. https://doi.org/10.5281/zenodo.8136157 	Vespertilionidae	Eptesicus	Cnephaeus	nilssonii	Keyserling & Blasius	1839	1	Arch. Naturgesch.	5(1): 315	Northern Bat	 atratus Kolenati, 1858; borealis Nilsson 1838 [not M&uuml;ller, 1776]; kuhli Nilsson 1836 [not Kuhl, 1819]; propinquus Peters, 1872; <b> parvus </b> Kishida, 1932.	Sweden.	W and E Europe to E Siberia and NW China; north beyond Arctic Circle in Scandinavia, south to Bulgaria, Iraq, the Elburz Mtns (N Iran), The Pamirs and W China (not Tibet); Korea; Hokkaido (Japan); Sakhalin Isl (Russia).	Not listed.	Least Concern	Subgenus Cnephaeus .  Includes propinquus ; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis ; see Yoshiyuki (1989). Does not include gobiensis ; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbetand Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus andpossibly paraphyletic with respect to that species; see Mayer and von Helversen (2001 a ). Specific epithet has often been spelled nilssoni , but the correct spelling is nilssonii . Placed in the subgenus Amblyotus by Horácek et al. (2000).	Eptesicus nilssonii	1005526	23	Northern Serotine	Northern Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	CHIROPTERA	VESPERTILIONIFORMES	NA	NA	VESPERTILIONOIDEA	Vespertilionidae	VESPERTILIONINAE	EPTESICINI	Eptesicus	Cnephaeus	nilssonii	Keyserling & Blasius	1839	1						Sweden.			borealis (Nilsson, 1838) [preoccupied]|kuhli (Nilsson, 1836) [preoccupied]|nilssonii (Keyserling & Blasius, 1839)|atratus (Kolenati, 1858)|propinquus (W. Peters, 1872)|parvus Kishida, 1932	NA	NA				France|Germany|Switzerland|Liechtenstein|Italy|Austria|Czech Republic|Hungary|Slovakia|Slovenia|Romania|Ukraine|Poland|Belarus|Lithuania|Latvia|Estonia|Norway|Sweden|Finland|Russia|Armenia|Azerbaijan|Iran|Kazakhstan|Kyrgyzstan?|Mongolia|China|Japan|North Korea|South Korea	Asia|Europe	Palearctic	LC	0	0	0	Eptesicus_nilssonii	0	sciname match	Eptesicus_nilssonii	0	Burgin, C. J., Zijlstra, J. S., Becker, M. A., Handika, H., Alston, J. M., Widness, J., Liphardt, S., Huckaby, D. G., and Upham, N. S. (2025). How many mammal species are there now? Updates and trends in taxonomic, nomenclatural, and geographic knowledge. Journal of Mammalogy in revision: TBD. https://doi.org/10.1101/2025.02.27.640393	Cnephaeus_nilssonii	1005526	23	Northern Serotine	Northern Bat	Theria	Placentalia	Boreoeutheria	Laurasiatheria	Chiroptera	Yangochiroptera	NA	NA	Vespertilionoidea	Vespertilionidae	Vespertilioninae	Nycticeiini	Cnephaeus	NA	nilssonii	von Keyserling & J. H. Blasius	1	Vesperugo Nilssonii	Keyserling, A. von and Blasius, J.H. 1839. Uebersicht der Gattungs- und Artcharaktere der europäischen Fledermäuse. Archiv für Naturgeschichte 1839(1):293-331.	https://www.biodiversitylibrary.org/page/24904226				Sweden.			moved from Eptesicus to Cnephaeus	Cláudio, V. C., Novaes, R. L., Gardner, A. L., Nogueira, M. R., Wilson, D. E., Maldonado, J. E., ... & Moratelli, R. (2023). Taxonomic re-evaluation of New World Eptesicus and Histiotus (Chiroptera: Vespertilionidae), with the description of a new genus. Zoologia (Curitiba), 40, e22029.				France|Germany|Switzerland|Liechtenstein|Italy|Austria|Czech Republic|Hungary|Slovakia|Slovenia|Romania|Ukraine|Poland|Belarus|Lithuania|Latvia|Estonia|Norway|Sweden|Finland|Russia|Armenia|Azerbaijan|Iran|Kazakhstan|Kyrgyzstan?|Mongolia|China|Japan|North Korea|South Korea	Asia|Europe	Palearctic	LC (as Eptesicus nilssonii)	0	0	0	Eptesicus_nilssonii	0	sciname match	Eptesicus_nilssonii	0	Simmons, N. B., & Cirranello, A. L. (2025). Batnames.org Species List Version 1.7 (1.7). Zenodo. https://doi.org/10.5281/zenodo.14796586	Vespertilionidae	Cnephaeus		nilssonii	Keyserling & Blasius	1839	1	Arch. Naturgesch.	5(1): 315	Northern Bat	atratus Kolenati, 1858; borealis Nilsson 1838 [not M&uuml;ller, 1776]; kuhli Nilsson 1836 [not Kuhl, 1819]; propinquus Peters, 1872; parvus Kishida, 1932.	Sweden.	W and E Europe to E Siberia and NW China; north beyond Arctic Circle in Scandinavia, south to Bulgaria, Iraq, the Elburz Mtns (N Iran), The Pamirs and W China (not Tibet); Korea; Hokkaido (Japan); Sakhalin Isl (Russia).	<a href='https://cites.org/eng/app/appendices.php' target='_blank'>Not Listed</a>	<a href='https://www.iucnredlist.org/species/7910/22116204/' target='_blank'>Least Concern as Eptesicus nilssonii</a>	Includes propinquus; see W. B. Davis (1965). Revised by Wallin (1969). Does not include japonensis; see Yoshiyuki (1989). Does not include gobiensis; see Strelkov (1986), Pavlinov and Rossolimo (1987), and Corbetand Hill (1992). See Rydell (1993), but note that he included japonensis in this species. Closely related to serotinus andpossibly paraphyletic with respect to that species; see Mayer and von Helversen (2001a). Specific epithet has often been spelled nilssoni, but the correct spelling is nilssonii. Placed in the subgenus Amblyotus by Horácek et al. (2000).		Mammal Diversity Database. (2025). Mammal Diversity Database (Version 2.2) [Data set]. Zenodo. https://doi.org/10.5281/zenodo.15007505	NA	Eptesicus nilssonii; Eptesicus nilssonii; Eptesicus nilssonii; Eptesicus nilssonii; Eptesicus nilssonii; Eptesicus nilssonii; nilssonii; parvus; atratus; borealis; kuhli; propinquus; parvus; atratus; borealis; kuhli; propinquus; borealis; kuhli; nilssonii; atratus; propinquus; parvus; Sérotine de Nilsson; Nordfledermaus; Murciélagohortelano nortefo; Northern Bat; Northern Serotine; Northern Bat; Northern Bat; Northern Bat; E. nilssonii