|A drawing by Ángel Cabrera, showing a ventral view of the entire animal, with one wing folded and the other spread; a lateral view of the head; and a view of the inner side of the ear|
|Species complex:||Myotis nattereri|
Ángel Cabrera, 1904
Myotis escalerai is a European bat in the genus Myotis, found in Spain (including the Balearic Islands), Portugal, and far southern France. Although the species was first named in 1904, it was included in Natterer's bat (Myotis nattereri) until molecular studies, first published in 2006, proved that the two are distinct. Myotis escalerai is most closely related to an unnamed species from Morocco. Unlike Myotis 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 the winter in hibernation colonies, usually in caves or basements.
Myotis 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 245 to 300 mm (9.6 to 11.8 in) and body mass is 5 to 9.5 g (0.2 to 0.3 oz). Though very similar to Myotis 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" in various parts of its range.
Myotis escalerai was named by Ángel Cabrera in 1904, on the basis of four specimens from two localities in eastern Spain. He named the new species after one Mr. Martínez de la Escalera, who collected two specimens of the species in Bellver, Catalonia.2 Cabrera did not designate either of the two localities (Bellver and Foyos, Valencia) as the type locality, and later authors have listed both. Currently, Foyos, which was listed first by Cabrera, is accepted as the type locality.3 Cabrera commented that M. escalerai was close to Natterer's bat (Myotis nattereri),4 and in 1912, Gerrit S. Miller listed escalerai as a synonym 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.5 Miller's classification was followed for almost a century,1 and indeed, Cabrera himself accepted in 1914 that M. escalerai was not a valid species.6
|Relationships in the Myotis nattereri complex based on nuclear and mitochondrial DNA sequences7|
However, a 2006 study by Carlos Ibáñez and colleagues found that Myotis nattereri in fact included several cryptic species with highly distinctive DNA sequence characteristics, even though morphological differences were small or nonexistent. One, which they recorded in the southern Iberian Peninsula, was identified as Myotis escalerai.8 Populations in the mountains of northern Spain represent another species ("Myotis sp. A"), which is now also known from the Alps.9 A 2009 study using data from the mitochondrial genes cytochrome b and ND1 found that Myotis escalerai is most closely related to an unnamed species from Morocco previously included in M. nattereri ("Myotis sp. B"), and more distantly to other members of the Myotis nattereri group.10 M. escalerai and the Moroccan species are estimated to have diverged about 2 million years ago.11 Later in 2009, M. escalerai was also recorded for the first time from France.12 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. sp. A,13 but another study published in the same year included these populations in M. sp. A.14 The latter study, by I. Salicini and colleagues, used sequences from six nuclear genes to confirm the distinctiveness of M. escalerai and its close relationship with M. sp. B.15 The common name "Escalera's bat" has been used for M. escalerai.16
A medium-sized, gray bat, Myotis escalerai is similar to M. nattereri. The fur is long and soft with a brown tone on the back and the underparts lighter, 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 forming 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.16 According to several authors, it differs from Myotis nattereri in showing a distinct fringe of hairs on the tail membrane,17 but bat specialist A.M. Hutson writes that this feature does not distinguish the two species.18 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.19
The head body length is 42 to 50 mm (1.7 to 2 in), tail length is 38 to 47 mm (1.5 to 1.9 in), forearm length is 35 to 43 mm (1.4 to 1.7 in), ear length is 14 to 18 mm (0.6 to 0.7 in), wingspan is 245 to 300 mm (9.6 to 11.8 in), and body mass is 5 to 9.5 g (0.2 to 0.3 oz).16
The range of Myotis escalerai remains poorly constrained and may turn out to be larger than currently known.20 Myotis escalerai is widespread in Spain and Portugal.21 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.22 Similarly, in Catalonia, Myotis escalerai is widespread and occurs from sea level up to an altitude of 1500 m (4900 ft).23 The species also occurs on the Balearic Islands of Mallorca, Menorca, and Ibiza.24 The sole French record is from a cave in Valmanya, Pyrénées-Orientales.12
Relatively little is known of the biology of Myotis 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.19 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.8 In Aragón, colonies contain 50 to 880 individuals,25 and Catalonian colonies are known to contain over a hundred bats.23 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 colonies need constant temperatures between 0 and 5 °C (32 and 41 °F), and are usually located in caves or basements. Myotis escalerai is considered a sedentary species, and does not usually migrate over long distances, although it does move between reproduction and hibernation colonies.19 Rabies has been identified in a Spanish specimen of M. escalerai.26
The IUCN Red List does not separate Myotis escalerai from Myotis nattereri, which is listed as "Least Concern",27 but the two species are listed separately on the Annex to the Agreement on the Conservation of Populations of European Bats.28 Portugal lists Myotis escalerai as "Vulnerable", though noting that populations may be increasing.29 Because of its restriction to caves, it is considered vulnerable in Aragón.30 In Catalonia, the species appears tolerant of different habitats and of human disturbance, but it is listed as "Data Deficient".23 In France, where the species was only discovered in 2009, it is also listed as "Data Deficient".31
- Simmons, 2005, p. 513
- Cabrera, 1904, pp. 280–281
- Benda et al., 2006, p. 118, footnote; Ibáñez et al., 2006, p. 286
- Cabrera, 1904, p. 280
- Miller, 1912, p. 174
- Salicini et al., in press, p. 8
- García-Mudarra et al., 2009, fig. 2; Galimberti, 2011, fig. 4.5; Salicini et al., in press, fig. 2
- Ibáñez et al., 2006, p. 286
- García-Mudarra et al., 2009, p. 441
- García-Mudarra et al., 2009, fig. 2
- García-Mudarra et al., 2009, p. 442
- Evin et al., 2009
- Galimberti, 2011, pp. 99–100
- Salicini et al., in press, p. 8
- Salicini et al., in press, p. 5
- Serra-Cobo et al., 2008a, p. 166
- Ibáñez et al., 2006, p. 286; Serra-Cobo et al., 2008a, p. 167; Salicini et al., in press, p. 8
- Hutson, 2010, p. 5
- Serra-Cobo et al., 2008a, p. 167
- Evin et al., 2009, p. 143
- García-Mudarra et al., 2009, fig. 2, pp. 441–442; Ministério do Ambiente e do Ordenamento do Território, 2010; Serra-Cobo et al., 2008a, p. 168
- Alcalde et al., 2008, pp. 8, 15
- Flaquer et al., 2010, p. 42
- Amengual et al., 2007, pp. 274–275; Serra-Cobo et al., 2008a, p. 168
- Alcalde et al., 2008, p. 8
- Serra-Cobo et al., 2008b, p. 171
- Hutson et al., 2008
- Anonymous, 2010
- Ministério do Ambiente e do Ordenamento do Território, 2010, table 1
- Alcalde et al., 2008, p. 15
- Gauthier, 2009, p. 2
- 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. Analisi d' ADN mitocondrial de cinc especies de quiropters 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. Myotis nattereri. 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). ISBN 978-84-475-3344-2
- 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. ISBN 978-0-8018-8221-0