Myotis alcathoe
Encyclopedia
Myotis alcathoe is a European bat in the genus Myotis. Known only from Greece and Hungary when it was first described in 2001, its known distribution has since expanded to Spain, England, Sweden, and Azerbaijan, among other countries. It is similar to the whiskered bat
Whiskered bat
The whiskered bats Myotis mystacinus and related species, are small European bats with long fur. Although uncommon, M. mystacinus is often found around human habitation and around water; it is similar to Brandt's bat Myotis brandtii, from which it was distinguished as a separate species only in...

 (Myotis mystacinus) and other species and is difficult to distinguish from them. However, its brown fur is distinctive and it is clearly different in characters of its karyotype
Karyotype
A karyotype is the number and appearance of chromosomes in the nucleus of an eukaryotic cell. The term is also used for the complete set of chromosomes in a species, or an individual organism.p28...

 and DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

s. Although some genetic data suggest that it is related to Geoffroy's bat
Geoffroy's Bat
Geoffroy's Bat is a species of vesper bat in the Vespertilionidae family.It can be found in the following countries: Afghanistan, Algeria, Armenia, Austria, Azerbaijan, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, France, Georgia, Germany, Greece, Hungary, Iran, Israel,...

 (Myotis emarginatus), other analyses do not support a close relationship between M. alcathoe and any other species.

With a forearm length of 30.8 to 34.6 mm (1.2 to 1.4 in) and body mass of 3.5 to 5.5 g (0.1 to 0.2 oz), Myotis alcathoe is a small bat. The fur is usually reddish-brown on the upperparts and brown below, but more grayish in juveniles. The tragus (a projection on the inner side of the ear) is short, as is the ear itself, and the inner side of the ear is pale at the base. The wings are brown and the baculum
Baculum
The baculum is a bone found in the penis of most mammals. It is absent in humans, but present in other primates, such as the gorilla and chimpanzee.The bone aids in sexual intercourse.-Purpose:...

 (penis bone) is short and broad. M. alcathoe has a very high-pitched echolocation
Animal echolocation
Echolocation, also called biosonar, is the biological sonar used by several kinds of animals.Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects...

 call, with a frequency that falls from 120 kHz at the beginning of the call to about 43 kHz at the end.

Usually found in old-growth deciduous forest near water, Myotis alcathoe forages high in the canopy and above water and mostly eats flies. The animal roosts in cavities high in trees. Although there are some winter records from caves, it may also spend the winter in tree cavities. Several parasites have been recorded on M. alcathoe. The IUCN Red List
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...

 assesses Myotis alcathoe as "Data Deficient
Data Deficient
Data Deficient is a category applied by the IUCN, other agencies, and individuals to a species when the available information is not sufficient for a proper assessment of conservation status to be made...

", but it is considered threatened in several areas because of its rarity and vulnerability to habitat loss.

Taxonomy

The whiskered bat
Whiskered bat
The whiskered bats Myotis mystacinus and related species, are small European bats with long fur. Although uncommon, M. mystacinus is often found around human habitation and around water; it is similar to Brandt's bat Myotis brandtii, from which it was distinguished as a separate species only in...

 (Myotis mystacinus) and similar species in Eurasia (collectively known as "whiskered bats") are difficult to distinguish from each other; for example, the distantly related Brandt's bat
Brandt's Bat
Brandt's Bat is a species of vesper bat in the Vespertilionidae family.It is found throughout most of Europe and parts of Asia.It is named for the German zoologist Johann Friedrich von Brandt.-Echolocation:...

 (Myotis brandtii) was not recognized as distinct from M. mystacinus until the 1970s. Small, unusual M. mystacinus-like bats were first recorded in Greece in the 1970s, but it was not until the advent of genetic studies that these bats could be confirmed as representing a distinct species, named Myotis alcathoe. In 2001, the species was described by German zoologists Otto von Helversen and Klaus-Gerhard Heller on the basis of specimens from Greece and Hungary. Although it also differs from other whiskered bats by morphological characters, Myotis alcathoe is most clearly distinct in its genetics, including DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

s and the location of the nucleolus organizer region
Nucleolus organizer region
The nucleolus organizer region or nucleolar organizer is a chromosomal region around which the nucleolus forms. This region is the particular part of a chromosome that is associated with a nucleolus after the nucleus divides. The region contains several tandem copies of ribosomal RNA genes...

s.

Von Helversen and Heller argued that none of the old names now considered synonyms
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...

 of M. mystacinus could apply to M. alcathoe, because these names all have their type localities in western or central Europe. However, the more recent discovery of M. alcathoe further to the west renders it possible that an older name may be discovered. In addition, Russian researcher Suren Gazaryan has suggested that the name caucasicus Tsytsulina, 2000 (originally proposed for a subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...

 of M. mystacinus from the Caucasus) may prove to be applicable to M. alcathoe; in that case, the species would be renamed Myotis caucasicus. The species may have remained undetected in Germany for so long because bat researchers did not sample its preferred habitats and would dismiss unusual-looking whiskered bats as being abnormal M. mystacinus or M. brandtii.

On the basis of mitochondrial DNA
Mitochondrial DNA
Mitochondrial DNA is the DNA located in organelles called mitochondria, structures within eukaryotic cells that convert the chemical energy from food into a form that cells can use, adenosine triphosphate...

 sequence analysis, Myotis alcathoe was at first thought to be most closely related to Geoffroy's bat
Geoffroy's Bat
Geoffroy's Bat is a species of vesper bat in the Vespertilionidae family.It can be found in the following countries: Afghanistan, Algeria, Armenia, Austria, Azerbaijan, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Czech Republic, France, Georgia, Germany, Greece, Hungary, Iran, Israel,...

 (Myotis emarginatus) of southern Europe, North Africa, and southwestern Asia. However, a study of the mitochondrial cytochrome b
Cytochrome b
Cytochrome b/b6 is the main subunit of transmembrane cytochrome bc1 and b6f complexes. In addition, it commonly refers to a region of mtDNA used for population genetics and phylogenetics.- Function :...

 gene incorporating many Myotis species did not support this relationship, and could not place M. alcathoe securely at a specific position among Eurasian Myotis. Two groups with slightly divergent mitochondrial DNA sequences (separated by 1.3 to 1.4% sequence divergence) are distinguishable within the species, which probably correspond to different glacial refugia where M. alcathoe populations survived the last glacial period. One, known as the "Hungarian" group, has been recorded from Spain, France, Austria, Hungary, and Slovakia, and probably corresponds to a refugium in Iberia; the other, the "Greek" group, is known only from Greece and Slovakia.

The specific name, alcathoe, refers to Alcathoe, a figure from Greek mythology
Greek mythology
Greek mythology is the body of myths and legends belonging to the ancient Greeks, concerning their gods and heroes, the nature of the world, and the origins and significance of their own cult and ritual practices. They were a part of religion in ancient Greece...

 who was turned into a bat when she refused the advances of the god Dionysus
Dionysus
Dionysus was the god of the grape harvest, winemaking and wine, of ritual madness and ecstasy in Greek mythology. His name in Linear B tablets shows he was worshipped from c. 1500—1100 BC by Mycenean Greeks: other traces of Dionysian-type cult have been found in ancient Minoan Crete...

. She was associated with gorges and small streams, the preferred habitat of Myotis alcathoe in Greece. In their original description, von Helversen and colleagues described her as a nymph
Nymph
A nymph in Greek mythology is a female minor nature deity typically associated with a particular location or landform. Different from gods, nymphs are generally regarded as divine spirits who animate nature, and are usually depicted as beautiful, young nubile maidens who love to dance and sing;...

, and the common name
Common name
A common name of a taxon or organism is a name in general use within a community; it is often contrasted with the scientific name for the same organism...

 "nymph bat" has therefore been used for this species. However, none of the classical sources speak of Alcathoe as a nymph; instead, she was a princess, the daughter of King Minyas
Minyas (mythology)
In Greek mythology, Minyas was the founder of Orchomenus, Boetia. As the ancestor of the Minyans, a number of Boeotian genealogies lead back to him, according to the classicist H.J. Rose...

 of Orchomenos. Therefore, Petr Benda recommended in 2008 that the common name "Alcathoe bat" or "Alcathoe myotis" be used instead. Other common names include "Alcathoe's bat" and "Alcathoe whiskered bat".

Description

Myotis alcathoe is the smallest European Myotis species. The fur is brownish on the upperparts, with a reddish tone in old specimens, and a slightly paler gray-brown below. Younger animals may be completely gray-brown. The brown fur distinguishes adult M. alcathoe from other whiskered bats, but juveniles cannot be unambiguously identified on the basis of morphology. M. alcathoe is similar to Daubenton's bat
Daubenton's bat
Daubenton's Bat, Myotis daubentonii, is a Eurasian bat with quite short ears. It ranges from Britain to Japan and is considered to be increasing its numbers in many areas.The name commemorates the French naturalist Louis-Jean-Marie Daubenton....

 (Myotis daubentonii) and M. emarginatus in color. On the upper side of the body, the hairs are 6 to 8 mm long and have dark bases and brown tips. The hairs on the lower side of the body are only slightly paler at the tip than at the base.

The face and the upper lips are reddish to pink. Although most of the face is hairy, the area around the eyes is bare. The nostrils are heart-shaped, and their back end is broad, as in M. brandtii, not narrow as in M. mystacinus. Several gland
Gland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...

s are present on the muzzle, most prominently in reproductively active males. The ears are brown and are lighter on the inside than the outside. There is a notch at the edge of the ear, and the pointed tragus (a projection inside the ear that is present in some bats) extends up to this notch. The base of the inner side of the ear is white; it is much darker in M. mystacinus. The feet and the thumbs are very small. The small size of the ear, tragus, feet, and thumb distinguishes M. alcathoe from the slightly larger M. mystacinus and M. brandtii, but the feet are relatively larger than in M. mystacinus.

The wings are brown, but lighter than those of M. mystacinus. The plagiopatagium (the portion of the wing between the last digit and the hindlegs) is attached to the fifth toe. The tail extends only about 1 mm beyond the back margin of the uropatagium (the portion of the wing membrane between the hindlegs). The calcar
Calcar
The calcar, also known as the calcaneum , is the name given to a spur of cartilage arising from inner side of ankle and running along part of outer interfemoral membrane in bats , this is to help spread the interfemoral membrane , which is part of the wing membrane between the tail and the hind...

, a cartilaginous spur supporting the uropatagium, is slender. With a width around 1.3 mm, the penis is narrow, and it lacks a broadened tip. The baculum
Baculum
The baculum is a bone found in the penis of most mammals. It is absent in humans, but present in other primates, such as the gorilla and chimpanzee.The bone aids in sexual intercourse.-Purpose:...

 (penis bone) is about 0.5 mm long. The short and broad shape of this bone distinguishes M. alcathoe from M. brandtii as well as M. ikonnikovi.

The skull is similar in shape to that of M. mystacinus and M. brandtii, but the front part of the braincase is higher. The second and third upper premolars (P2 and P3) are tiny and pressed against the upper canine (C1) and fourth premolar (P4). The canine is less well-developed than in M. mystacinus. There is a clear cusp
Cusp (dentistry)
A cusp is an occlusal or incisal eminence on a tooth.Canine teeth, otherwise known as cuspids, each possess a single cusp, while premolars, otherwise known as bicuspids, possess two each. Molars normally possess either four or five cusps...

 present on the side of the P4. The accessory cusp known as the protoconule is present on each of the upper molars when they are unworn. M. mystacinus lacks the P4 cusp and the protoconules on the molars.

As usual in Myotis species, Myotis alcathoe has a karyotype
Karyotype
A karyotype is the number and appearance of chromosomes in the nucleus of an eukaryotic cell. The term is also used for the complete set of chromosomes in a species, or an individual organism.p28...

 consisting of 44 chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...

s, with the fundamental number of chromosomal arms equal to 52. However, a 1987 study already found that M. alcathoe (then called "Myotis sp. B") differs from both M. mystacinus and M. brandtii in the pattern of active nucleolus organizer regions on the chromosomes. M. alcathoe also differs from other Myotis species in the sequences of the mitochondrial genes 12S rRNA and NADH dehydrogenase
NADH dehydrogenase
NADH dehydrogenase is an enzyme located in the inner mitochondrial membrane that catalyzes the transfer of electrons from NADH to coenzyme Q...

 subunit 1 by at least 5% and 13%, respectively.

Myotis alcathoe has the highest-frequency echolocation
Animal echolocation
Echolocation, also called biosonar, is the biological sonar used by several kinds of animals.Echolocating animals emit calls out to the environment and listen to the echoes of those calls that return from various objects near them. They use these echoes to locate and identify the objects...

 call of any European Myotis. In open terrain, the call has an average duration of 2.5 ms
Millisecond
A millisecond is a thousandth of a second.10 milliseconds are called a centisecond....

, but it may be up to 4 ms long. At the beginning, its frequency is around 120 kHz, but it then falls fast, subsequently falls slightly slower, and at the end falls faster again. The call reaches its highest amplitude at around 53 kHz. It terminates at around 43 to 46 kHz; this characteristic is especially distinctive. In different experiments, the time between calls was found to be around 85 and 66 ms, respectively. The high-pitched call may be an adaptation to the animal's occurrence in dense vegetation.

Head and body length is about 4 cm (2 in) and wingspan is around 20 cm (8 in). Forearm length is 30.8 to 34.6 mm (1.2 to 1.4 in), tibia
Tibia
The tibia , shinbone, or shankbone is the larger and stronger of the two bones in the leg below the knee in vertebrates , and connects the knee with the ankle bones....

 length is 13.5 to 15.9 mm (0.5 to 0.6 in), hindfoot length is 5.1 to 5.8 mm (0.2 to 0.2 in), and body mass is 3.5 to 5.5 g (0.1 to 0.2 oz).

Distribution and habitat

Although Myotis alcathoe was initially known only from Greece and Hungary and was thought to be restricted to southeastern Europe, records since then have greatly expanded its range, and it is now known from Spain and England to Sweden and European Turkey. In several European countries, focused searches were conducted to detect its occurrence. Its habitat generally consists of moist, deciduous, mature forest near streams, for example in ravines or in alluvial forest (forest near a river), where there are many decaying trees that the bat can use as roosting sites. In Germany, its preferred habitat consists of mixed deciduous forest. In the south of the continent, it usually occurs in mountain ranges, but the factors affecting its distribution in the north are less well known. Its range appears to be similar in shape to those of the greater
Greater Horseshoe Bat
The Greater Horseshoe Bat is a European bat of the Rhinolophus genus. Its distribution covers Europe, Africa, South Asia and Australia. It is the largest of the European Horseshoe Bats and is thus easily distinguished from other species...

 and lesser horseshoe bat
Lesser horseshoe bat
The Lesser Horseshoe Bat , is a type of European bat related to but smaller than its cousin, the Greater Horseshoe Bat...

s (Rhinolophus ferrumequinum and R. hipposideros) and Myotis emarginatus. It may yet be found in other European countries, such as the Benelux
Benelux
The Benelux is an economic union in Western Europe comprising three neighbouring countries, Belgium, the Netherlands, and Luxembourg. These countries are located in northwestern Europe between France and Germany...

 countries, Ireland, and Moldova. Although there are abundant records from some areas, such as France and Hungary, the species appears to be rare in most of its range.

Known records are as follows:
Albania: A single specimen was caught in 2006 in a forest of planes (Platanus orientalis
Platanus orientalis
Platanus orientalis, or the Oriental plane, is a large, deciduous tree of the Platanaceae family, known for its longevity and spreading crown. The species name derives from its historical distribution eastward from the Balkans, where it was recognized in ancient Greek history and literature....

) and poplars (Populus spp.) next to a small stream. M. mystacinus was recorded at the same place.
Austria: Three specimens of Myotis alcathoe were caught in Burgenland
Burgenland
Burgenland is the easternmost and least populous state or Land of Austria. It consists of two Statutarstädte and seven districts with in total 171 municipalities. It is 166 km long from north to south but much narrower from west to east...

, southeastern Austria, in 2006. They were caught near fishponds in a region dominated by oak (Quercus petraea) and hornbeam (Carpinus betulus).
Azerbaijan: The species was recorded around 2009.
Belgium: The species is know from the 21 of July 2011 in two place around the city of Rochefort
Rochefort, Belgium
Rochefort is a Walloon municipality located in the Belgian province of Namur close to the Ardennes. On January 1, 2006 Rochefort had a total population of 12,038. The total area is 165.27 km² which gives a population density of 73 inhabitants per km²...

.
Bulgaria: The species is known from six localities in the south and west of the country; the first record dates from 2003. Habitats include river and mountain forests.
Caucasus: Bats collected in the Russian part of the Caucasus
Caucasus
The Caucasus, also Caucas or Caucasia , is a geopolitical region at the border of Europe and Asia, and situated between the Black and the Caspian sea...

 and in Abkhazia
Abkhazia
Abkhazia is a disputed political entity on the eastern coast of the Black Sea and the south-western flank of the Caucasus.Abkhazia considers itself an independent state, called the Republic of Abkhazia or Apsny...

 (a breakaway part of Georgia) from 2003 to 2009 may represent Myotis alcathoe. They are small and morphometrically
Morphometrics
Morphometrics refers to the quantitative analysis of form, a concept that encompasses size and shape. Morphometric analyses are commonly performed on organisms, and are useful in analyzing their fossil record, the impact of mutations on shape, developmental changes in form, covariances between...

 distinct from other local Myotis mystacinus-like bats. However, the identity of these bats has not been confirmed molecularly.
Croatia: In 2003, Myotis alcathoe was recorded here on the basis of two specimens.
Czech Republic: Here, the species was recorded at nine sites clustered in three regions, with the first record dating from 2001, in addition to records from roadkilled specimens at three further sites. The typical habitat was mature oak-hornbeam forest near water with dead, decaying trees, at altitudes ranging from 170 to 390 m (558 to 1280 ft). Both M. mystacinus and M. brandtii occur in some of the same places in this country. M. alcathoe has a limited, patchy distribution within the country, but reaches a high abundance in suitable habitat.
France: Myotis alcathoe was informally recognized in France in 2000 as a small Myotis similar to Myotis mystacinus, the "Murin cantalou"; in 2002, it was realized that this bat represents Myotis alcathoe. A large number of sites are known, mostly in the north of the country. The species reaches altitudes of up to 2000 m (6562 ft). It is usually found close to water, but it has been found in a variety of habitats, including farmlands, swamps, forests, and wooden grounds. In late summer and autumn, it occurs in caves.
Germany: The species is known from two different areas in the country. In 2005 and 2006, specimens were caught in an old moist forest near the Rhine in western Baden-Württemberg
Baden-Württemberg
Baden-Württemberg is one of the 16 states of Germany. Baden-Württemberg is in the southwestern part of the country to the east of the Upper Rhine, and is the third largest in both area and population of Germany's sixteen states, with an area of and 10.7 million inhabitants...

. Two other bats were found in highway tunnels close to this site. The species is also known from the Kyffhäuser
Kyffhäuser
The Kyffhäuser is a range of hills located on the border of the German state of Thuringia with Saxony-Anhalt. It stands on the southern edge of the Harz. The range has a length of and a width of . It reaches its highest point at the Kulpenberg , situated in Thuringia...

 hill range of Thuringia
Thuringia
The Free State of Thuringia is a state of Germany, located in the central part of the country.It has an area of and 2.29 million inhabitants, making it the sixth smallest by area and the fifth smallest by population of Germany's sixteen states....

 in central Germany, an island of relatively warm habitat with some unusual wildlife. There, bats were caught near a spring in a karst
KARST
Kilometer-square Area Radio Synthesis Telescope is a Chinese telescope project to which FAST is a forerunner. KARST is a set of large spherical reflectors on karst landforms, which are bowlshaped limestone sinkholes named after the Kras region in Slovenia and Northern Italy. It will consist of...

 landscape amid oak-dominated deciduous forest. The species was also recorded in deciduous forest at a former Soviet military training site in eastern Thuringia. Myotis alcathoe has also been recorded in the nearby states of Saxony-Anhalt
Saxony-Anhalt
Saxony-Anhalt is a landlocked state of Germany. Its capital is Magdeburg and it is surrounded by the German states of Lower Saxony, Brandenburg, Saxony, and Thuringia.Saxony-Anhalt covers an area of...

 and Saxony
Saxony
The Free State of Saxony is a landlocked state of Germany, contingent with Brandenburg, Saxony Anhalt, Thuringia, Bavaria, the Czech Republic and Poland. It is the tenth-largest German state in area, with of Germany's sixteen states....

, where it occurs in mixed deciduous forest. Many Saxony-Anhalt records are from near water. However, the species was also recorded in the center of the city of Chemnitz
Chemnitz
Chemnitz is the third-largest city of the Free State of Saxony, Germany. Chemnitz is an independent city which is not part of any county and seat of the government region Direktionsbezirk Chemnitz. Located in the northern foothills of the Ore Mountains, it is a part of the Saxon triangle...

 in Saxony.
Greece: The species has been recorded in the Pindus and Rhodopi Mountains of central and northern Greece. Here, Myotis alcathoe is usually found in stands of plane
Platanus
Platanus is a small genus of trees native to the Northern Hemisphere. They are the sole living members of the family Platanaceae....

 or alder
Alder
Alder is the common name of a genus of flowering plants belonging to the birch family . The genus comprises about 30 species of monoecious trees and shrubs, few reaching large size, distributed throughout the North Temperate Zone and in the Americas along the Andes southwards to...

 trees next to small streams in ravines. The bat hunts close to the trees, within the stand. It is often found together with the lesser horseshoe bat and with M. mystacinus.
Hungary: Myotis alcathoe is not uncommon in the mountain forests of northeastern Hungary. It has been found at brooks and lakes in oak, beech, alder, and hornbeam
Hornbeam
Hornbeams are relatively small hardwood trees in the genus Carpinus . Though some botanists grouped them with the hazels and hop-hornbeams in a segregate family, Corylaceae, modern botanists place the hornbeams in the birch subfamily Coryloideae...

 (Carpinetum) forests at 230 to 670 m (750 to 2200 ft) altitude. Both M. brandtii and M. mystacinus occur together with M. alcathoe there.
Italy: Myotis alcathoe has been recorded in beech forest in Majella National Park
Majella National Park
The Majella National Park is a national park located in the provinces of Chieti, Pescara and L'Aquila, in the region Abruzzo, Italy....

 in the region of Abruzzo
Abruzzo
Abruzzo is a region in Italy, its western border lying less than due east of Rome. Abruzzo borders the region of Marche to the north, Lazio to the west and south-west, Molise to the south-east, and the Adriatic Sea to the east...

. Additional specimens of Myotis alcathoe have been identified in Italy using molecular methods.
Latvia: A small Myotis was captured at a cave in Latvia between 2007 and 2010; pending genetic testing, it is suspected to be M. alcathoe.
Poland: The species was recorded in four caves in southern Poland in 2005 and 2006, and later at several other sites in the south of the country. It is known from 182 to 1294 m (597 to 4245 ft) above sea level, most often in beech forest (Fagus sylvatica), but also in several other forest types.
Romania: A single Myotis alcathoe was captured in 2007 in a nature reserve in the eastern Carpathians; the reserve contains riverine and conifer forest. The species was additionally recorded in a forested valley containing a small stream in Alba County
Alba County
Alba is a county of Romania, in Transylvania, its capital city being Alba-Iulia with a population of 66,406.- Demographics :In 2002, it had a population of 382,747 and the population density was 61/km².* Romanians - 90.4%* Hungarians - 5.4%...

.
Serbia: The species was reported on the basis of three specimens shortly before 2009, but is probably rare.
Slovakia: Here, Myotis alcathoe is known from a single site, a cave in old deciduous forest at 525 m (1722 ft) altitude.
Slovenia: A single specimen was recorded in Slovenia in 2007.
Spain: In Catalonia
Catalonia
Catalonia is an autonomous community in northeastern Spain, with the official status of a "nationality" of Spain. Catalonia comprises four provinces: Barcelona, Girona, Lleida, and Tarragona. Its capital and largest city is Barcelona. Catalonia covers an area of 32,114 km² and has an...

, the species is known from six sites, ranging from sea level to 1200 m (3900 ft) altitude. It is known in beech and riverine forest and was first recorded in 2006. The species is known from three sites in La Rioja
La Rioja (Spain)
La Rioja is an autonomous community and a province of northern Spain. Its capital is Logroño. Other cities and towns in the province include Calahorra, Arnedo, Alfaro, Haro, Santo Domingo de la Calzada, and Nájera.-History:...

, where it was recorded in 2004, and occurs amidst beech and riverine forest at 790 to 1390 m (2590 to 4560 ft) altitude. It has also been found at seven localities in Navarre
Navarre
Navarre , officially the Chartered Community of Navarre is an autonomous community in northern Spain, bordering the Basque Country, La Rioja, and Aragon in Spain and Aquitaine in France...

, with the first record dating from 2004. There, it occurs in beech and oak forest at altitudes from 140 to 980 m (460 to 3220 ft). A single locality is known in Galicia.
Sweden: The species was recorded at five sites in the south of the country, starting in 2008, on the basis of echolocation calls.
Switzerland: Myotis alcathoe has been recorded from the Col du Marchairuz
Col du Marchairuz
Col du Marchairuz is a high mountain pass in the Jura Mountains in the canton of Vaud in Switzerland.It connects Le Brassus and Bière. The pass road has a maximum grade of 14 percent.-See also:* List of highest paved roads in Europe...

 in the Jura Mountains
Jura mountains
The Jura Mountains are a small mountain range located north of the Alps, separating the Rhine and Rhone rivers and forming part of the watershed of each...

 (canton of Vaud
Vaud
Vaud is one of the 26 cantons of Switzerland and is located in Romandy, the French-speaking southwestern part of the country. The capital is Lausanne. The name of the Canton in Switzerland's other languages are Vaud in Italian , Waadt in German , and Vad in Romansh.-History:Along the lakes,...

).
Turkey: Eight individuals have been caught at three sites in close vicinity in the European part of the country in 2006.
Ukraine: In 2009, the possible occurrence of Myotis alcathoe in Ukraine was recorded. In 2011, the species was definitively recorded there on the basis of two bats caught in the far southwest of the country in 2009.
United Kingdom: Myotis alcathoe has been recorded in England since 2003, and is known from two swarming sites in the south and a third site in the north of the country. The northern England site, in Ryedale
Ryedale
Ryedale is a non-metropolitan district of the shire county of North Yorkshire in England. Settlements include Helmsley, Kirkbymoorside, Malton, Norton-on-Derwent, Pickering, and Terrington.-Derivation of name:...

, is in a protected area
Protected area
Protected areas are locations which receive protection because of their recognised natural, ecological and/or cultural values. There are several kinds of protected areas, which vary by level of protection depending on the enabling laws of each country or the regulations of the international...

 with many old trees, and the southern sites (in Sussex
Sussex
Sussex , from the Old English Sūþsēaxe , is an historic county in South East England corresponding roughly in area to the ancient Kingdom of Sussex. It is bounded on the north by Surrey, east by Kent, south by the English Channel, and west by Hampshire, and is divided for local government into West...

) are in woodland.
The species is also known from Montenegro and possibly from Bosnia and Herzegovina. Early records of Myotis ikonnikovi—now known to be an eastern Asian species—from Ukraine, Bulgaria, and Romania may also pertain to this species. Because whiskered bats in many cases cannot easily be distinguished from each other without the use of genetic methods, some listings do not differentiate between them; records of Myotis alcathoe and/or M. mystacinus and/or (in some cases) M. brandtii have been reported from Bulgaria, Belgium, and Montenegro.

Ecology and behavior

Myotis alcathoe is a rare species with narrow ecological requirements. According to a study in the Czech Republic, the diet of Myotis alcathoe mostly consists of nematocera
Nematocera
Nematocera , is a suborder of elongated flies with thin, segmented antennae and mostly aquatic larvae, consisting of the mosquitoes, crane flies, gnats, and midges....

n flies, but caddisflies, spider
Spider
Spiders are air-breathing arthropods that have eight legs, and chelicerae with fangs that inject venom. They are the largest order of arachnids and rank seventh in total species diversity among all other groups of organisms...

s, small lepidoptera
Lepidoptera
Lepidoptera is a large order of insects that includes moths and butterflies . It is one of the most widespread and widely recognizable insect orders in the world, encompassing moths and the three superfamilies of butterflies, skipper butterflies, and moth-butterflies...

ns, and neuroptera
Neuroptera
The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order contains some 6,010 species...

ns are also taken. The presence of spiders in the diet suggests that the species gleans prey from foliage. It forages mainly high in the canopy and over water, and is often found in dense vegetation. The parasitic mite Spinturnix mystacina has been found on M. alcathoe, and the mites on M. alcathoe, M. brandtii, and M. mystacinus are genetically closely related. The bat fly Basilia mongolensis nudior has been recorded on M. alcathoe in Thuringia. When caught, individuals of Myotis alcathoe are much calmer than M. mystacinus or M. brandtii.

Myotis alcathoe lives in small groups. In Greece, a maternity colony, containing three females and two juveniles, has been found in a plane tree. Additional roosts were found high in oak trees in Baden-Württemberg and Saxony-Anhalt. Twenty-seven roosting sites have been found in the Czech Republic, all but one in trees (the last was in a concrete pole). Most of the tree roosts were in oaks (Quercus robur); others were in limes (Tilia cordata
Tilia cordata
Tilia cordata is a species of Tilia native to much of Europe and western Asia, north to southern Great Britain , central Scandinavia, east to central Russia, and south to central Spain, Italy, Bulgaria, Turkey, and the Caucasus; in the south of its range it is restricted to...

), birches (Betula pendula), and various other species. Its strong preference for roosting sites in trees is unusual among European bats. Roosts tend to be located high in the canopy, and are often in old trees. In summer, roosts may contain large groups of up to 80 individuals, but autumn roosts in the Czech Republic are occupied by smaller groups.

In Saxony-Anhalt, the species forages deep in valleys when temperatures are above 10 °C (50 °F), but on warmer slopes or rocky areas when it is colder. There, Myotis alcathoe is relatively easy to capture in August, because M. brandtii and M. mystacinus already start swarming in late July. Although there are some records of Myotis alcathoe in caves during the winter, it is also possible that animals spend the winter in tree cavities, and whether swarming behavior occurs in Myotis alcathoe is unclear. An animal found in a cave in Saxony-Anhalt in January was not sleeping deeply. Reproduction may also take place in caves, but pregnant females have been found as late as June. Relatively many juveniles are caught between July and September. In England, one individual of M. alcathoe was captured in 2003 (and identified at the time as M. brandtii) and again in 2009. Three individuals that were telemetrically
Telemetry
Telemetry is a technology that allows measurements to be made at a distance, usually via radio wave transmission and reception of the information. The word is derived from Greek roots: tele = remote, and metron = measure...

 tracked (in eastern France, Thuringia, and Baden-Württemberg, respectively) moved only 800 m (2600 ft), 935 m (3070 ft), and 1440 m (4720 ft) from their night quarters; M. brandtii and M. mystacinus tend to move over longer distances.

Conservation status

Because Myotis alcathoe remains poorly known, it is assessed as "Data Deficient
Data Deficient
Data Deficient is a category applied by the IUCN, other agencies, and individuals to a species when the available information is not sufficient for a proper assessment of conservation status to be made...

" on the IUCN Red List
IUCN Red List
The IUCN Red List of Threatened Species , founded in 1963, is the world's most comprehensive inventory of the global conservation status of biological species. The International Union for Conservation of Nature is the world's main authority on the conservation status of species...

. However, it may be endangered because of its narrow ecological preferences. Reservoir construction may threaten the species' habitat in some places; two Greek sites where it has been recorded have already been destroyed. Forest loss is another possible threat, and the species may be restricted to undisturbed habitats. Because of its patchy distribution and likely small population, it probably does not easily colonize new habitats. The species is protected by national and international measures, but the IUCN Red List recommends further research on various aspects of the species as well as efforts to increase public awareness of the animal. In addition, old forests need to be conserved and the species' cave roosts need to be protected.

In Catalonia, the species is listed as "Endangered" in view of its apparent rarity there. The Red List of Germany's Endangered Vertebrates lists Myotis alcathoe as "Critically Endangered" as of 2009. In Hungary, where the species is probably not uncommon in suitable habitat, it has been protected since 2005. However, the species is declining there and is threatened by habitat loss and disturbance of caves.

Literature cited

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