Escalera's Bat
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Escalera's bat (''Myotis escalerai'') is a European bat in the genus '' Myotis'', found in Spain (including the
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), Portugal, and far
southern France Southern France, also known as the South of France or colloquially in French language, French as , is a defined geographical area consisting of the regions of France that border the Atlantic Ocean south of the Marais Poitevin,Louis Papy, ''Le midi ...
. Although the species was first named in 1904, it was included in Natterer's bat (''Myotis nattereri'') until molecular studies, first published in 2006, demonstrated that the two are distinct species. ''M. escalerai'' is most closely related to an unnamed species from Morocco. Unlike ''M. nattereri'', which lives in small groups in tree holes, ''M. escalerai'' forms large colonies in caves. Females start to aggregate in late spring in maternity colonies, and their young are born in summer. The species spends each winter in hibernation colonies, usually in caves or basements. ''M. escalerai'' is a medium-sized, mostly gray bat, with lighter underparts. It has a pointed muzzle, a pink face, and long ears. The wings are broad and the species is an agile flyer. Wingspan is and body mass is . Though very similar to ''M. nattereri'', it differs from that species in some features of the tail membrane. The conservation status of ''M. escalerai'' is assessed as " Vulnerable" or "
Data Deficient A data deficient (DD) species is one which has been categorized by the International Union for Conservation of Nature (IUCN) as offering insufficient information for a proper assessment of conservation status to be made. This does not necessaril ...
" in various parts of its range.


Taxonomy

''Myotis escaleraii'' was named by Angel Cabrera in 1904, on the basis of four specimens from two localities in eastern Spain. He named the new species after the Spanish entomologist Manuel Martínez de la Escalera, who collected two specimens of the species in Bellver, Catalonia. Cabrera did not designate either of the two localities (Bellver and Foios, Valencia) as the type locality, and later authors have listed both. Currently, Foios, which was listed first by Cabrera, is accepted as the type locality. Cabrera commented that ''M. escalerai'' was close to Natterer's bat (''Myotis nattereri''), and in 1912, Gerrit S. Miller listed ''escalerai'' as a
synonym A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
of that species. He argued that one of the features Cabrera had listed as distinguishing the two was an artefact of the preservation of the specimens of ''M. escalerai'' in alcohol. Miller's classification was followed for almost a century,Simmons, 2005, p. 513 and indeed, Cabrera himself accepted in 1914 that ''M. escalerai'' was not a valid species.Salicini et al., in press, p. 8 However, a 2006 study by Carlos Ibáñez and colleagues found that ''M. nattereri'' in fact included several
cryptic species In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
with highly distinguished
DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
s characteristics, even though morphological differences were small or nonexistent. One, which they recorded in the southern Iberian Peninsula, was identified as ''M. escalerai''. Populations in the mountains of northern Spain represent another species (now known to be ''
Myotis crypticus The cryptic myotis (''Myotis crypticus'') is a European vespertilionid bat. It is a member of the Natterer's bat (''M. nattereri'') species complex, and is also the closest living relative of ''M. nattereri''. It is mostly distributed across Eur ...
''), which is now also known from the Alps.García-Mudarra et al., 2009, p. 441 A 2009 study using data from the
mitochondrial A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is use ...
genes cytochrome ''b'' and ND1 found that ''M. escalerai'' is most closely related to an unnamed species from Morocco previously included in ''M. nattereri'' (now known to be ''Myotis'' ''zenatius''), and more distantly to other members of the ''Myotis nattereri'' group.García-Mudarra et al., 2009, fig. 2 ''M. escalerai'' and ''M. zenatius'' are estimated to have diverged about 2 million years ago.García-Mudarra et al., 2009, p. 442 Later in 2009, ''M. escalerai'' was also recorded for the first time from France. One 2011 study found a fifth putative species in the complex ("''Myotis'' sp. C"), occurring in the Italian peninsula and most closely related to ''M. crypticus'', but another study published in the same year included these populations in ''M. crypticus''. The latter study, by I. Salicini and colleagues, used sequences from six
nuclear Nuclear may refer to: Physics Relating to the nucleus of the atom: * Nuclear engineering *Nuclear physics *Nuclear power *Nuclear reactor *Nuclear weapon *Nuclear medicine *Radiation therapy *Nuclear warfare Mathematics *Nuclear space *Nuclear ...
genes to confirm the distinctiveness of ''M. escalerai'' and its close relationship with ''M.'' ''zenatius''.


Description

A medium-sized gray bat, ''Myotis escalerai'' is similar to ''Myotis nattereri''. The fur is long and soft; with a brown tone on the back, and the brighter underparts approaching white. The feet are dark gray. Much of the face is pink, and the muzzle is pointed, with long hairs on the upper lip resembling a moustache. The long ears are brown to gray. The tragus, a projection on the inner side of the outer ear, is long and reaches to the middle of the ear and colored gray to yellow, becoming darker from the base towards the tip. According to several authors, it differs from ''M. nattereri'' in showing a distinct fringe of hairs on the tail membrane, but bat specialist A.M. Hutson writes that this feature does not distinguish the two species. In addition, the presence of an S-shaped spur on the uropatagium (membrane between the hind legs), which approaches the middle of the membrane, is a distinctive feature of this species. With its broad wings, low flight, and rapid wingbeats, the species is capable of precise, agile flight.Serra-Cobo et al., 2008a, p. 167 The head body length is , tail length is , forearm length is , ear length is , wingspan is , and body mass is .Serra-Cobo et al., 2008a, p. 166


Distribution and ecology

The range of ''Myotis escalerai'' remains poorly constrained and may turn out to be larger than currently known.Evin et al., 2009, p. 143 ''M. escalerai'' is widespread in Spain and Portugal. For example, it occurs widely, though localized, in Aragón, where ''Myotis'' sp. A (the only other species in the ''M. nattereri'' complex to occur there) is known from a single locality only. Similarly, in Catalonia, ''M. escalerai'' is widespread and occurs from sea level up to an altitude of .Flaquer et al., 2010, p. 42 The species also occurs on the
Balearic Islands The Balearic Islands ( es, Islas Baleares ; or ca, Illes Balears ) are an archipelago in the Balearic Sea, near the eastern coast of the Iberian Peninsula. The archipelago is an autonomous community and a province of Spain; its capital is ...
of
Mallorca Mallorca, or Majorca, is the largest island in the Balearic Islands, which are part of Spain and located in the Mediterranean. The capital of the island, Palma, is also the capital of the autonomous community of the Balearic Islands. The Bal ...
, Menorca, and
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. The sole French record is from a cave in
Valmanya Valmanya (; ca, Vallmanya) is a commune in the Pyrénées-Orientales department in southern France. Geography Valmanya is located in the canton of Le Canigou and in the arrondissement of Prades. Population See also *Communes of the Pyr ...
,
Pyrénées-Orientales Pyrénées-Orientales (; ca, Pirineus Orientals ; oc, Pirenèus Orientals ; ), also known as Northern Catalonia, is a department of the region of Occitania, Southern France, adjacent to the northern Spanish frontier and the Mediterranean Sea. ...
.Evin et al., 2009, p. 142 Relatively little is known of the biology of ''M. escalerai''. Females begin to form reproductive colonies in April and May, either small ones or larger aggregations that may also contain males. However, most males remain solitary in this period, although some also form colonies. The single young is born in June or July and becomes independent after some six weeks. Mating usually takes place in fall, but sometimes in winter. The formation of large reproductive colonies in caves, which may consist of several hundreds of individuals, distinguishes ''M. escalerai'' from ''M. nattereri'' as well as ''M.'' sp. A, which roost in smaller groups in tree holes.Ibáñez et al., 2006, p. 286 In Aragón, colonies contain 50 to 880 individuals, and Catalan colonies are known to contain over a hundred bats. Reproductive colonies may be formed in a variety of structures, including caves, mines, tree holes, and human-made structures such as bridges and houses. However,
hibernation Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
colonies need constant temperatures between , and are usually located in caves or basements. ''M. escalerai'' is considered a sedentary species, and does not usually migrate over long distances, although it does move between reproduction and hibernation colonies. Rabies has been identified in a Spanish specimen of ''M. escalerai''.


Conservation status

The IUCN Red List classifies ''Myotis escalerai'' as " least concern". ''M. nattereri'' and ''M. escalerai'' are also listed separately on the Annex to the Agreement on the Conservation of Populations of European Bats. Portugal lists ''M. escalerai'' as " vulnerable", though noting that populations may be increasing.Ministério do Ambiente e do Ordenamento do Território, 2010, table 1 Because of its restriction to caves, it is considered vulnerable in Aragón. In Catalonia, the species appears tolerant of different habitats and of human disturbance, but it is listed as "
data deficient A data deficient (DD) species is one which has been categorized by the International Union for Conservation of Nature (IUCN) as offering insufficient information for a proper assessment of conservation status to be made. This does not necessaril ...
". In France, where the species was only discovered in 2009, it is also listed as "data deficient".Gauthier, 2009, p. 2


References


Literature cited

*Alcalde, J.T., Trujillo, D., Artázcoz, A. and Agirre-Mendi, P.T. 2008
Distribución y estado de conservación de los quirópteros en Aragón
Graellsia 64(1):3–16 (in Spanish). *Amengual, B., López-Roig, M., Mas, O., González, J. and Serra-Cobo, J. 2007
Anàlisi d'ADN mitocondrial de cinc espècies de quiròpters de les Illes Balears
Bolletí de la Societat d'Història Natural de les Balears 50:269–277 (in Catalan). *Anonymous. 2010
6th Session of the Meeting of Parties. Prague, Czech Republic, 20–22 September 2010. Resolution 6.2. Amendment of the Annex to the Agreement
EUROBATS.MoP6.Record.Annex5. *Benda, P., Andreas, M., Kock, D., Lučan, R.K., Munclinger, P., Nová, P., Obuch, J., Ochman, K., Reiter, A., Uhrin, M. and Weinfurtová, D. 2006
Bats (Mammalia: Chiroptera) of the eastern Mediterranean. Part 4. Bat fauna of Syria: distribution, systematics, ecology
Acta Societatis Zoologicae Bohemicae 70:1–329. *Cabrera, D.A. 1904
Ensayo monográfico sobre los quirópteros de España
Memorias de la Sociedad Española de Historia Natural 2(5):249–286 (in Spanish). *Evin, A., Lecoq, V., Durand, M., Tillon, L. and Pons, J. 2009
A new species for the French bat list: ''Myotis escalerai'' (Chiroptera: Vespertilionidae)
(subscription required). Mammalia 73(2):142–144. *Flaquer, C., Puig, X., Fàbregas, E., Guixé, D., Torre, I., Ràfols, R.G., Páramo, F., Camprodon, J., Cumplido, J.M., Ruiz-Jarillo, R., Baucells, A.L., Freixas, L. and Arrizabalaga, A. 2010
Revisión y aportación de datos sobre quirópteros de Catalunya: Propuesta de lista roja
Galemys 22(1):29–61 (in Spanish). *García-Mudarra, J.L., Ibáñez, C. and Juste, J. 2009
The Straits of Gibraltar: barrier or bridge to Ibero-Moroccan bat diversity?
(subscription required). Biological Journal of the Linnean Society 96(2):434–450. *Galimberti, A. 2011
DNA barcoding: a link between basic and applied science
PhD thesis, Università degli Studi di Milano-Bicocca. *Gauthier, O. 2009
Implementation of the Agreement on the Conservation of Populations of European Bats. National Report of France. 2006–2009
Inf.EUROBATS.MoP6.19. *Hutson, A.M. 2010
Draft Resolution 6.2: Amendment of the Annex of the Agreement. Review of Species to be listed on the Annex to the Agreement
Doc.EUROBATS.StC4-AC15.12.Rev.2. *Hutson, A.M., Aulagnier, S. and Spitzenberger, F. 2008. . In IUCN. IUCN Red List of Threatened Species. Version 2010.2.
www.iucnredlist.org
. Downloaded on February 12, 2011. *Ibáñez, C., García-Mudarra, J.L., Ruedi, M., Stadelmann, B. and Juste, J. 2006. The Iberian contribution to cryptic diversity in European bats. Acta Chiropterologica 8(2):277–297. *Miller, G.S. 1912
Catalogue of the mammals of western Europe (Europe exclusive of Russia) in the collections of the British Museum
London: British Museum (Natural History), 1019 pp. *Ministério do Ambiente e do Ordenamento do Território. 2010
Agreement on the Conservation of Populations of European Bats. Report on implementation of the Agreement in Portugal – 2010
Inf.EUROBATS.MoP6.35. *Salicini, I., Ibáñez, C. and Juste, J. In press
Multilocus phylogeny and species delimitation within the Natterer's bat species complex in the Western Palearctic
(subscription required). Molecular Phylogenetics and Evolution, in press. *Serra-Cobo, J., Roig, L., Bayer, X., Amengual, B. and Guasch, C. 2008a
Ratpenats. Ciència i mite
Barcelona: Edicions Universitat Barcelona, 267 pp. (in Catalan). *Serra-Cobo, J., Bourhy, H., López-Roig, M., Sánchez, L.P., Abellán, C., Borràs, M. and Amengual, B. 2008b
Rabia en quirópteros: Circulación de EBLV-1 (Lyssavirus de murciélagos europeos tipo 1) en murciélagos de España
Boletín Epidemiológico Semanal 16(15):169–180 (in Spanish). *Simmons, N.B. 2005. Order Chiroptera. Pp. 312–529 in Wilson, D.E. and Reeder, D.M. (eds.)
Mammal Species of the World: A Taxonomic and Geographic Reference
3rd ed. Baltimore: The Johns Hopkins University Press, 2 vols., 2142 pp. {{Taxonbar, from=Q773518 Mammals described in 1904 Mammals of Europe Mouse-eared bats