Scorpions of Europe Victor FET ACTA ZOOLOGICA BULGARICA ‘Acta rool. bul, 62 (1), 2010: 3-12 “Marshall Univesity, Huntington, West Virginia, USA email: fev@marshal.edy Abstract: ‘This brief review summarizes the studies in systematics and zoogeography of European scorpions, The current “splitting” tend in scorpion taxonomy is only a reasonable response tothe former “lumping,” Our better understanding of scorpion systematies became possible due to the availability of new morphologi- cal characters and molecular techniques, as well as of new material, Many taxa and local faunas are still, under revision. The total number of native scorpion species in Europe could easly be over 35 (Buthidae, 8; Euscorpiidae, 22-24; Chactidae, 1; Iuridae, 3) belonging to four families and six genera. The northern limit of natural (non-anthropochorie) scorpion distribution in Europe is in Saratov Province, Russia, at 50°40°S4"N, for Mesobuthus eupeus (Buthidac), juthidae, Euscorpiidae, Turdae, Chactidae, Buthus, Mesobuthus, Fuscorpius, lurus, Calchas, Belisarius One thinks of scorpions primarily as inhabiting deserts, and indeed the rich Palearctic scorpiofaunas of North AMfica, Middle East and Central Asia are well known—albeit not always well understood, However, it could be a surprise to many zoologists that many en demic Paleareticscorpion taxa (especially Euscorpiidae and luridae) do not infact live in arid habitats at all but are found in quite temperate and even humid and cold environments, including mountains (Alps, Balkans, Taurus, Caucasus) up to 2500 m ‘The latitudinal boundary of scorpions in Europe is not easy to determine since many populations are introduced by humans, e.g. Euscorpius flavicaudis in France, E. tergestinus in Austria, and E. italicus in many areas. The northern boundary of natural (post- glacial dispersal) scorpion distribution in the west- em Europe is probably marked by the northernmost record of Euscorpius germanus in Austria (North, Tyrol, 47°39°N; Koposca et al. 2001), but in the easter Europe (lower Volga in Russia, Saratov Province) Mesobuthus eupeus reaches almost 51°N (see below), Already Aristotle distinguished between toxic European Buthidae and non-dangerous Euscorpius (Fer et al. 2009), Small but interesting scorpiofauna of Europe received a lot of attention starting with Linnaeus himself who in 1767 described Scorpio carpathicus (Fer, SoLeaLaD 2002). A substantial re- view on the Aegean region published by Kiszenpactt (1975). Our latest reviews (Fer et al. 2004, Kautsas et al, 2008) address details on taxonomy and dis- tribution of most European taxa; see also Vicxots, ‘Satomone (2008). Family BUTHIDAE, ‘Two most prominent in Europe buthids are Buthus oceitanus (Amorevx, 1789) in Spain and France, and Mesobuthus gibbosus (Bruit, 1832) in the Balkans. These two species, which occupy xeric habitats, are well-known as the only relatively toxie scorpions in Europe. These two taxa exem- plify two routes used by Buthidae (as well as by numerous other xeric elements) in their dispersal to Europe: in Buthus, from Africa (via the IberianFet V. Peninsula), and in Mesobuthus, from Asia (via the Balkan Peninsula). The genus Buthus Leact, 1815 is an African element, with its center of diversity in North Africa Buthus occitanus was described from southem Franee; it is widespread in Spain, and was also re- corded from Portugal (a record from Greece in the Fauna Europaea database is clearly erroneous). The same species (with numerous local subspecies) was also traditionally reported for the large part of Attica north of the equator and from the Middle East Recently, Loumco (2002, 2003) reconsidered many of these populations, elevated subspecies to species rank, described new species, and reduced Buthus oc- citanus to European populations. Lourenco (2002) also suggested that the colonization of Europe by B. occitanus was rather recent (Quaternary). This spe- cies has been an object of detailed genetic studies to evaluate gene fiow across the Strait of Gibraltar (Ganrenseix, Larciapie 2003; Gaxtenssin 2004) Cavern et al (1994) outlined the ecological factors influencing the distribution of Buthus occitanus in Europe ‘An unexpected discovery of two endemic Buthus species in Spain, Buthus ibericus Lourexco et Vacnon, 2004 and Buthus montanus Lourenco et VAcHON, 2004, brings more attention to this genus (Crnuet, Pérez-Bore 2005). A new, endemic spe- cies of Buthus was also found on Cyprus (Lourenco ef al. in press); we include Cyprus in the account of European fauna due to its coverage in Fauna Europaea (Caucrrni 2004), ‘The second buthid genus in Europe, Mesobuthus Vachon, 1950, is an arid Asian element, with its highest species diversity in Iran. The taxonomy of _Mesobuthus sin flux: amotecular survey (GANTENBEIN et al, 2003) confirms a separate position of Balkan- Anatolian M. gibbosus. In Europe, it is found in Albania, SW Bulgaria, Macedonia (Doiran), Greece, Montenegro, and European Turkey. In Bulgaria, it was first reported by Turi etal, (2004) from Pirin; it is unclear if resident populations exist. An endem- ic island species Mesobuthus eyprius Gawrevasin et Krorr, 2000, has been recently described from Cyprus (previously treated as M. gibbosus) 4 Biology and ecology of M. gibbosus in Greece has been intensively studied (Crucrrm, Marist 1987; Karrsas et al. 2006, 2008; Katrsas, Mvtovas 2007), ‘An impressive phylogeographic study (based on mtDNA markers) was published by PaRMAkEU (2006a), The origin of Rhodes population was stud- ied by Gaxrenpen, Larotanin (2002). Ganrinnein, Keioirtey (2004) used both M. gibbosus and M. cyprius to assess rates of molecular evolution in scorpions, In addition, two predominantly Asian sp (or most likely complexes of species), Mesobuthus caucasicus (Norway, 1840) and Mesobuthus eu- peus (C.L. Kock, 1839), are found in the fringes of geographic Europe. Mesobuthus caucasicus was reported from southem Ukraine (Odessa and Kherson Provinces), highly disjunct from its main range; specimens from the Odessa Province exist in collections (Fer 1989) ‘The same species is found in the Northem Caucasus areas of Russia (Dagestan, Chechnya) that geo- graphically belong to Europe Another widely ranging Asian species, ‘reus, is found in the lower Volga, south of European Russia, where it was reported for Astrakhan Province (Fer 1989). These populations were described both a8 M. ¢. bogdoensis (But, 1896) and M, e, vol- gensis (Brut, 1925); they likely belong to the Central Asian “thersites” complex. Disjunct repro- ducing populations were recently documented even further north at the right bank of Volga, such as one in the Volgograd Province (Saatary 1997). The Saratov Province record in Krasnoarmeisky District (misidentified as Buthus occitanus; Axixix, Kineey 1998; V. Anixin, pers. comm. 2010), at 50°40°S4"N 45°38°36"E, is the northem boundary of natural scorpion distribution in Europe. Incidentally, M. eupeus has been recently also documented in the Orenburg Province of Russia (Davycora, Rusakov 2001), which constitutes the northernmost limit of natural scorpion distribution in Asia and the Palearctic, This record in Aituar Steppe of the Orenburg Natural Reserve lies at 51°13°N 57°75°E (S. Esvunmy, pers. comm. 2010); compare with Gromiov (2001). Only some records etal.Review Scorpions of Europe of Paruroctonus boreus (Gieaxd, 1854) in south- em Alberta, Canada, lie slightly further north (D, on~sson, pers. comm. 2010). In total, eight native buthid species are docu- ‘mented in Europe. Further increase in their number is not expected (but so unexpected were new species, discoveries in Spain and Cyprus). Family EUSCORPIDAE This family is represented in Europe by its ‘unique genus Euscorpius Thorell, 1876, a European- Mediterranean endemic, the core and diverse element of European scorpiofauna, found in diverse habitats from sea coast to the high Alps and Balkans. In many areas, two or more congeners (or even “‘consubgen- ers”) are found sympatrically. Currently, 17 species are valid (see below) but their accepted ranges do not include all known populations of Euscorpius. 1 estimate the total number of Euscorpius species in Europe as at least 22-24, Most populations from Greece and the Balkans still require a detailed study. Although family Euscorpiidae is represented in mountainous Asia (subfamily Scorpiopinae), the closest taxa to Buscorspius (also in subfamily Euscorpiinae) are Mexican genera Megacormus and Plesiochactas (So.eGLAD, Sissom 2001). Some spe cies of Euscorpius are very clearly defined, but in many other cases traditional morphological criteria have not been sufficient, For recent reviews, see: Ferret al, (2004); Karrsas et al, (2008), and Viewors, SaLomons: (2008; they also give a good review of known ecology). Subgenus Tetratrichobothrius Buew1., 1917 Euscorpius flavicaudis (DeGeex, 1778), the only species of this subgenus, is limited to the Western Mediterranean: Spain, Baleares, southern France, Corsica, Italy, and Sardinia. It is also found in North Africa, and has introduced populations in England and even Brazil and Uruguay. Its biology and ecology was studied by Bextos (1991, 1992, 1993) in the in- troduced population in England. Iligh genetic homo- geneity (Gantrnaran ef al. 2002) between its popu- lations in mainland France and Corsica suggests that E, flavicaudis could be a relict that passed through. a bottleneck and has been dispersing with humans. Anthropochory is documented for many populations of E. flavicaudis in France (VaCHoN 1969); a similar trend is observed for other Buscorpius. Subgenus Polytrichobothrius BiRULs, 1917 Euscorpius italicus (Heeast, 1800), the larg- est species in the genus, is found from France to Caucasus (including the Black Sea coast of Turkey, Georgia, and Russia), and also in Northern A\tica (Algeria, Morocco, Tunisia). For its distribution in Slovenia, sce Fer et al. (2001); for biology in Italy, see CoLowso (2006) and Crucirtt ef al. (2007) For distribution and biology in Switzerland, see BauNwaLper (2001, 2005). A very low genetic di- vergence was detected by Fer et al. (2006) between European and Anatolian populations of F, italicus (as compared to its sister species E. naupliensis) This suggests a recent dispersal from @ glacial re- fugium, possibly even in historical times—which could explain a puzzling absence of E. italicus from all Aegean and Mediterranean islands. The spe- cies is clearly anthropophilic and anthropochorie; Braunwaner (2001, 2005) notes its predominant occurrence in human habitations. Many of its dis- junct populations are clearly introduced (e.g, in Iraq and Yemen), and some probably became extinct, e.g from the Don River in southern Russia, discovered in the 1920s but never reported again (Fer 1989) For decades, E.italicus was considered the only spe- cies of its distinct subgenus. However, GaNTENBEIN etal. (2002), based on both molecular and morpho- logical data, restored Euscorpius naupliensis (C.L Kocu, 1837), an endemic of Greece (Peloponnese and Zakynthos Island), and a senior synonym of E: italicus zakynthi Caroixcco, 1950 from Zakynthos, Its ecology was studied in the Peloponnese (Cructr, Bussico, 2001; as E, italicus) and on Zakynthos (Covoymo 2006). Based on the genetic divergence between £. naupliensis and E. italicus, GaNTENBEIN ef al, (2002) suggested that their split predates the Messinian salinity evisis (5.2 Myrs ago). Subgenus Aipiscorpius Gavennewy et al., 1999 ‘This subgenus was established to accommodate the topology of a pilot 16S mtDNA phylogeny that demonstrated a separate position of a group of spe-Fet V. cies previously included in the subgenus Euscorpius s. str, Morphology clearly confirms this compact group; a revision is in progress (Fer et al.). The sub- gomus ranges from Austria to Caucasus. It includes some high-mountain forms, up to 2170 m in East Tirol (Austria) and 2250 m in Swiss Alps (B. ger- manus), and 2200-2400 m in the Balkans (E. bero- ni). Currently five species are valid; the speci el composition of Aipiscorpius is still unclear in the ‘main part of its range, i. the Balkans and Anatolia Euscorpius alpha Caporiacco, 1950 was el- evated to species level from a subspecies of E. ger- manus by GaxvenneIn et al. (2000a). It is found in NW Italy (Piedmont, Valle d’Aosta and Lombardy) and Switzerland. F, germanus and E, alpha are al- lopatric, sibling Alpine species (Frt et al. 2004) The degree of molecular divergence suggests that these two species have evolved in the area before the Pleistocene glaciations, maybe since the orogenesis of the Alps (GanTENBEIN et al. 2000a). E. germanus also has isolated populations in the Apennines; it is not known whether they are relict or introduced (Fer ef al. 2004), For distribution and biology in Switzerland, see BRaunwatneR (2001, 2005). Euscorpius beroni Ft, 2000, was described from high Prokletije Mountains of Albania. It be- longs to the Balkan portion of the “E. mingrelicus complex.” Euscorpius germanus (C. L. Koc, 1837). This traditional species was restricted by GanreNBein et al, (2000) only to some populations in Austria, NE Italy, Slovenia (cast), and Switzerland, In Slovenia, subspecies £, g. marcuzzii Caporiacco, 1950 is found sympatrically with E, gamma. Details of distribution are available for Slovenia (Fer et al. 2001), Austria (Komrosc# et al. 2001; Korex 2002; Komrosca 2004, 2009), and Switzerland (BRavnwatpeR 2001, 2005). The species is included in the Red Data Lists of Austria (KowPoscut 2009). Euscorpius mingrelicus (Kessier, 1874) was described from Georgia. In Europe it is found, in my opinion, only in the Northern Caucasus of Russia (Krasnodar Province). The Balkan populations, still technically included in E. mingrelicus as subspecies, are under revision (see below), 6 Euscorpius gamma Caroniacco, 1950, de- scribed from Slovenia as a subspecies of E. ger- ‘manus, was clevated to species rank by ScHERABON et al. (2000). E. gamma belongs to the “E. mingrelicus complex.” It is confirmed for SE Austria (Carinthia), Croatia, NE Italy (Friuli), and Slovenia, Details of distribution are available for Slovenia (Fer et al 2001) and Austria (Komposcut er al. 2001; Komposcit 2004, 2009). The species is included in the Red Data Lists of Austria (Komrosc 2009), In addition to E. beroni and E, gamma, “E. ‘mingrelicus complex” includes a number of Balkan populations, for which species boundaries are not yet clear. Their range includes Croatia, Bosnia and Herzegovina, Montenegro, Kosovo, Serbia, Macedonia, SW Bulgaria (Pirin), and NW Greece (Epiros). Several names are available for these pop- ulations, technically still under 6. mingrelicus sub- species (KALTSAS et al. 2008). Subgenus Euscorpius Tnonet1, 1876 ‘The nominotypic subgenus, after separation of Alpiscorpius (Gaxrexosin et al. 1999), corresponds, to whatis often vaguely named “E. carpathicus com- plex.” For decades, most of its species were classi- fied as subspecies ofa (very polytypic) “E. carpathi- cus.” Currently, nine species are valid. The subgenus ranges from the Baleares in the west (E, balearicus) to the Crimea in the east (E, tauricus) and fiom the Southeast Alps in the north (E. fergestinus) to North ‘Arica in the south (E. sicanus), Euscorpius balearicus Cavowiacco, 1950, an endemic of the Balearic Islands (Spain), was elevated to species rank, using morphological and molecular data, by Gantewnnin et al. (2001), and redescribed by Fer, Soteatan (2002). Euscorpius carpathicus (Lixwavus, 1767), Fer, Sotsotan (2002) studied the existing type of Linnaeus, and limited this traditional species to Romania (Transylvanian Alps). As a result, all taxa formerly classified as E. carpathicus or its subspe- cies, belong to other species (of which some are yet not described). The first mtDNA marker data for the Romanian population were published by Fer et al (2002).Review Scorpions of Europe Euscorpius concinnus (C. L. Koc, 1837) was restored to species level by VieNott et a. (2005), and reported only from Italy, Saomone ef al. (2007), in a large DNA phylogeographic study for Italy, con- firmed its separate status as compared to E. sicanus and E, tergestinus, and discussed a rather high de- gree of divergence between these species. For biol ogy in Italy, see CoLowso (2006) Euscorpius hadzii Cavoriscco, 1950 was elevated to species rank and redescribed by Fer, Souracap (2002) (neotype from Albania). This large, dark Balkan species is found from Albania to SW Bulgaria, often in high mountains. It is easily identi- fiable by its additional neobothriotaxic trichobothria, con external aspect of pedipalp patella. A population of small-sized . hadzii (described as E. carpathicus lagostae Cavoniacco, 1950 from Lastovo Island) in- habits southern Croatian islands in the Adriatic Sea In Bulgaria, £, hadzit is found only in the southwest (Fer, Soteatap 2007) as far north as Zemen, where itapparently is closely allopatric with Euscorpius sp. of Stara Planina, Euscorpius koschewnikowi Buxv.a, 1900 is known only from Mt Athos in Greece, Itwas restored to species rank from a subspecies of E. carpathicus bby Fer, Sousotan (2002) who studied the types. Euscorpius oglasae Caroxtacco, 1950 was el- evated to species rank from a subspecies of E. car- athicus by Vionou et al, (22007). The species is en- demic to Montecristo Island (Tuscan Archipelago, ttaly) Euscorpius sicanus (C. L. Koc, 1837), de« scribed from Sicily, was restored to species rank from subspecies of, carpathicus by Fer etal. (2003) who also synonymized with it seven former subspecies, all from Italy. This species absorbed part of the popula tions listed by earlier authors (Knwzetnact 1975) as ““E. mesotrichus Had2i" (an invalid name). SaLowowe et al, (2007) confirmed a separate status of E. sica~ ‘nus as compared to E, coneinnus and E. tergestinus, It is widespread in central and southem Italy (with Sicily and Sardinia) and Greece (Thessaly, Sporades, Peloponnese), and i also found in Malta, North Aiea, and Madeira, Forbiology in Italy, see Couowno (2006), and in Greece, Cructrn, Busaico (2001) Euscorpius tauricus (C.L. Koc, 1837) is known only from the southem coast of the Crimea Peninsula in Ukraine; it was restored to species level by Fer (2003) based on DNA analysis, morphologi- cal investigation pending. For biology observations, see Kuxustixny (2004). The species is listed as en- demic in the Red Data Book of Ukraine (Veaves et al. 1999). Euscorpius tergestinus (C.L. Koch, 1837) was redescribed by Fer, Soueaiap (2002). Vicnou et al. (2005, 2007) separated two Italian species from this taxon, E. concinnus and B. oglasae. E. tergesti- nus embraces a number of former subspecies from Italy, and is confirmed for Albania, Croatia, Italy, Montenegro, San Marino, and Slovenia, For biology in Italy, see Coommo (2006). In a detailed phylo- geographic study, SaLomons ef al. (2007) confirmed separate status of F, tergestinus as compared to E. sicanus and E. concinnus in Italy. Three disjunct populations in Austria, likely introduced, were stud- ied using mtDNA markers by Huser et al, (2001); they are included in the Red Data Lists of Austria (Komoscs 2004, 2009), In addition to nine species listed above, place- ment of many European populations of “E. carpathi- cus complex” remains unclear. VioNout et al. (2007) studied populations from Corsica and France listed as E, carpathicus corsicanus Caroxiscco, 1950 and E. c. niciensis (CL. Kocs, 1841), respectively, and indicated their differences from known valid spe- cies. In a pilot mtDNA phylogeny of Gatixpens et al. (2001), French (Alpes-Maritimes) populations grouped with those of Tuscany, ie. E, concinnus. The Balkan and Aegean populations, which are currently under revision (Fer et al, in progress), include Croatia, Bosnia & Herzegovina, Montenegro, Kosovo, Serbia, Macedonia, Bulgaria, and Greece Several names are available for some of these popu- lations (i.e. E. carpathicus candiota Bixv.a, 1903 from Crete), potentially of the species rank; others likely represent undescribed species (for details, see Kaurtsas ef al, 2008). Especially diverse and compli- cated is Greek fauna, where a pilot DNA phylogeny indicates several species that were previously unrec- ognized (PouLiKaakou et al. 2008).Fet V. Family IURIDAE Both genera of luridae are recorded in Europe but Calchas Bruta, 1899 is represented only in the easternmost islands of Greece: the recently de- seribed Calchas gruberi Fer et al., 2009 (formerly reported as C. nordmanni) is found in the Greek ise lands Megisti and Samos as well as in Anatolia (Fer et al. 2009), The genus Jurus THoxr21, 1876, taxonomy of ‘which for many years was controversial but neglected, includes the largest of the European scorpions, Turus dufourcius (BwuLst, 1832). ParMaxstis etal (2006b) studied phylogeography of furus using mtDNA mark- ts, and demonstrated that populations of Peloponnese, Kithyra, and Crete formed a well-supported clade as ‘opposed to the Easter Aegean and Anatotian popula- tions. Inan extensive recent revision of urus, Kovac ef al, (2010) confirmed this conclusion and provided morphological basis as they limited 1. dufoureius to the “westem clade.” The populations of Eastem ‘Acgean Greck islands (Rhodes, Karpathos, Samos) belong to another, possibly new furus species. Based fon sequence divergence, PARMAKELIS ef al. (2006b) suggested that Jurus has been differentiating in the southern region of Agaeis at least since the Miocene (ca. 8 Mya). For details on distribution and morphol- ogy of L dufoureius, and the genus Jurus in general, see Kovakik et al (2010) Family CHACTIDAE Probably the most unique element of European scorpiofauna is a monotypic genus Belisarius Sivon, 1879, with its blind (eyeless) species B. xambeui Simox, 1879, from the Pyrenees of France (Pyrénées- Orientales) and Spain (Catalunya: Barcelona), Not much study has been done on this enigmatic scorpion since its original description. Its distribution, with a map, has been last summarized by Lackorx (1992), and it seems to be more common in Spanish part of the Pyrenees. Belisarius xambeui has bbeen characterized as an endogean, hygrophile lapidi- colous species, found in mountain forests (850-1300 ‘masl) in endogean subterrancan habitats. Itis the only cave scorpion in Europe, known from th France and 15 caves in Spain (Lacrorx 1992), Girona and 8 The family placement of Belisarius has been controversial. Early researchers (Knvzetsactt 1975), largely by geographic association, suggested a close relationship to Euscorpius, which is not confirmed. Lovrenco (1998) placed Belisarius in @ new fam- ily, Troglotayosicidae, Most recently, Belisarius was placed in Chactidae (otherwise limited to the New World) by Fer, Sotectan (2003). They provided a justification for its specific placement in a South ‘American subfamily Brotheinae, with a remark- able zoogeographic disjunction. (Even if Belisarius is assigned to Troglotayosicidae, the issue of South American relationship, with genus Troglotayosicus, still remains.) ‘This review should further complement and cor- rect the scorpion data available in Fauna Europaea (Cnucrrn: 2004) database. It reports for Europe two exotic buthid species, sometrus maculatus (DeGEER, 1778) and Centruroides gracilis (Lareen.ie, 1804). I. ‘maculatus isa subcosmopolitan scorpion, introduced across the globe; it was once reported for Spain but no resident population is confirmed. C. gracilis, which belongs to a large New World genus, has an introduced population on Tenerife, Canary Islands (also reported as C. nigrescens; B: 1984), in the coastal zone of Santa Cruz. town. ‘An unidentified species of Buscorpius is also known from Tenerife, likely an introduction, Macaronesian fauna is partially covered by the Fauna Europaea project (except Cape Verde Islands); however, it does not seem to include any native scorpions. E sicanus record from Madeira (Fer et al. 2003) could be either introduction from the Mediterranean, or a local relict. No scorpions have been reported from the Azores, It is remarkable that the “hidden diversity” of European fauna remained unrecognized for over 150 years of intensive arachnological research, The cur- rent “splitting” trend in scorpion taxonomy is, how- ever, only a response to former “lumping” trend, and became possible due to availabilty of new characters and techniques, as well as of new material. Indeed, a bothering taxonomic and zoogeographic puzzle for a long time was absence of diversity in European scor- pions; now we see that they behave as any normal z FUMEROReview Scorpions of Europe fauna, with a realistic combination of local, Asian and rican elements, ancient relits and recent dispersals, sympatric congeners and isolated endemics. The total number of native scorpion species in Europe (exelud- ing Macaronesia but including Cyprus) could be now estimated over 35 (Buthidae, 8; Euscorpiidae, 22-24; CChactidae, 1; Turidae, 3); the genus Eusconpius is still ‘under an intensive revision, especially in the Balkans, and will clearly yield more species. Acknowledgements: [thank dozens of friends and colleagues who, for decades, enjoyebly helped me and collaborated in Fu ropean scorpion studies ineluding (but not limited to}: ¥ References ANKIN VV, EA. Kats 1997, Eeologicalorstic description ofthe habitats of some rae insect and arachnid species of Lower Povolzhye.~ In: Materials of the International Research Practical Conference “Dhe Probloms of Prsersa tion ofthe Biodiversity in the rid Regions of Russia Not gograd, 11-17 September 1998, pp. 106-107 (in Russian). ‘Bivz Fonano M. 1984, Asicnidos In J.J, Bacallado Arig, 41. J. (dy: Pauna (Marina y Tereestr) del Archipiclago Canario, 108-116. 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