sign in sign up
<<
Home , Belisarius (scorpion), Buthus occitanus, Calchas (scorpion), Euscorpius, Euscorpius flavicaudis, Isometrus maculatus

ACTA ZOOLOGICA BULGARICA Acta zool. bulg., 62 (1), 2010: 3-12

Scorpions of

Victor FET

Marshall University, Huntington, West Virginia, USA; email: [email protected]

Abstract: This brief review summarizes the studies in systematics and zoogeography of European . The current “splitting” trend in is only a reasonable response to the former “lumping.” Our better understanding of scorpion systematics became possible due to the availability of new morphologi- cal characters and molecular techniques, as well as of new material. Many taxa and local are still under revision. The total number of native scorpion in Europe could easily be over 35 (, 8; , 22-24; , 1; , 3) belonging to four families and six genera. The northern limit of natural (non-anthropochoric) scorpion distribution in Europe is in Saratov Province, Russia, at 50°40’54”N, for Mesobuthus eupeus (Buthidae).

Keywords: Buthidae, Euscorpiidae, Iuridae, Chactidae, , Mesobuthus, , , ,

One thinks of scorpions primarily as inhabiting Already Aristotle distinguished between toxic deserts, and indeed the rich Palearctic scorpiofaunas European Buthidae and non-dangerous Euscorpius of North , and Central are well (Fe t et al. 2009). Small but interesting scorpiofauna known—albeit not always well understood. However, of Europe received a lot of attention starting with it could be a surprise to many zoologists that many en- Linnaeus himself who in 1767 described Scorpio demic Palearctic scorpion taxa (especially Euscorpiidae carpathicus (Fe t , So l e g l a d 2002). A substantial re- and Iuridae) do not in fact live in arid habitats at all but view on the Aegean published by Ki n z e l b a c h are found in quite temperate and even humid and cold (1975). Our latest reviews (Fe t et al. 2004, Ka l t s a s environments, including mountains (, , et al. 2008) address details on taxonomy and dis- Taurus, ) up to 2500 m. tribution of most European taxa; see also Vi g n o l i , The latitudinal boundary of scorpions in Europe Sa l o m o n e (2008). is not easy to determine since many populations are BUTHIDAE introduced by humans, e.g. Two most prominent in Europe buthids are in , E. tergestinus in Austria, and E. italicus in Buthus occitanus (Am o r e u x , 1789) in and many areas. The northern boundary of natural (post- France, and Mesobuthus gibbosus (Br u l l é , 1832) glacial dispersal) scorpion distribution in the west- in the Balkans. These two species, which occupy ern Europe is probably marked by the northernmost xeric habitats, are well-known as the only relatively record of Euscorpius germanus in Austria (North toxic scorpions in Europe. These two taxa exem- Tyrol, 47°39’N; Ko m p o s c h et al. 2001), but in the plify two routes used by Buthidae (as well as by (lower Volga in Russia, Saratov numerous other xeric elements) in their dispersal Province) Mesobuthus eupeus reaches almost 51°N to Europe: in Buthus, from Africa (via the Iberian (see below).

3 Fet V.

Peninsula), and in Mesobuthus, from Asia (via the Biology and ecology of M. gibbosus in Greece Balkan ). has been intensively studied (Cr u c i t t i , Ma r i n i 1987; The genus Buthus Le a c h , 1815 is an African Ka l t s a s et al. 2006, 2008; Ka l t s a s , My l o n a s 2007). element, with its center of diversity in . An impressive phylogeographic study (based on Buthus occitanus was described from southern mtDNA markers) was published by Pa r m a k e l i s et al. France; it is widespread in Spain, and was also re- (2006a). The origin of Rhodes population was stud- corded from Portugal (a record from Greece in the ied by Ga n t e n b e i n , La r g i a d è r (2002). Ga n t e n b e i n , Europaea database is clearly erroneous). The Ke i g h t l e y (2004) used both M. gibbosus and M. same species (with numerous local subspecies) was cyprius to assess rates of molecular evolution in also traditionally reported for the large part of Africa scorpions. north of the equator and from the Middle East. In addition, two predominantly Asian species Recently, Lo u r e n ç o (2002, 2003) reconsidered many (or most likely complexes of species), Mesobuthus of these populations, elevated subspecies to species caucasicus (No r d m a n n , 1840) and Mesobuthus eu- rank, described new species, and reduced Buthus oc­ peus (C.L. Ko c h , 1839), are found in the fringes of citanus to European populations. Lo u r e n ç o (2002) geographic Europe. also suggested that the colonization of Europe by B. Mesobuthus caucasicus was reported from occitanus was rather recent (). This spe- southern Ukraine (Odessa and Kherson Provinces), cies has been an object of detailed genetic studies highly disjunct from its main range; specimens from to evaluate gene flow across the Strait of Gibraltar the Odessa Province exist in collections (Fe t 1989). (Ga n t e n b e i n , La r g i a d è r 2003; Ga n t e n b e i n 2004). The same species is found in the Northern Caucasus Cr u c i t t i et al. (1994) outlined the ecological factors areas of Russia (Dagestan, Chechnya) that geo- influencing the distribution of Buthus occitanus in graphically belong to Europe. Europe. Another widely ranging Asian species, M. eu­ An unexpected discovery of two endemic peus, is found in the lower Volga, south of European Buthus species in Spain, Buthus ibericus Lo u r e n ç o Russia, where it was reported for Astrakhan Province et Va c h o n , 2004 and Buthus montanus Lo u r e n ç o (Fe t 1989). These populations were described both et Va c h o n , 2004, brings more attention to this genus as M. e. bogdoensis (Bi r u l a , 1896) and M. e. vol­ (Тe r u e l , Pé r e z -Bo t e 2005). A new, endemic spe- gensis (Bi r u l a , 1925); they likely belong to the cies of Buthus was also found on Cyprus (Lo u r e n ç o Central Asian “thersites” complex. Disjunct repro- et al. in press); we include Cyprus in the account ducing populations were recently documented even of European fauna due to its coverage in Fauna further north at the right bank of Volga, such as one Europaea (Cr u c i t t i 2004). in the Volgograd Province (Sa g a l a e v 1997). The The second buthid genus in Europe, Mesobuthus Saratov Province record in Krasnoarmeisky District Vachon, 1950, is an arid Asian element, with its (misidentified as Buthus occitanus; An i k i n , Ki r e e v highest species diversity in Iran. The taxonomy of 1998; V. An i k i n , pers. comm. 2010), at 50°40’54”N Mesobuthus is in flux; a molecular survey (Ga n t e n b e i n 45°38’36’’E, is the northern boundary of natural et al. 2003) confirms a separate position of Balkan- scorpion distribution in Europe. Anatolian M. gibbosus. In Europe, it is found in Incidentally, M. eupeus has been recently also Albania, SW Bulgaria, Macedonia (Doiran), Greece, documented in the Orenburg Province of Russia Montenegro, and European Turkey. In Bulgaria, it (Da v y g o r a , Ru s a k o v 2001), which constitutes the was first reported byT e r u e l et al. (2004) from Pirin; northernmost limit of natural scorpion distribution it is unclear if resident populations exist. An endem- in Asia and the Palearctic. This record in Aituar ic species Mesobuthus cyprius Ga n t e n b e i n Steppe of the Orenburg Natural Reserve lies at et Kr o p f , 2000, has been recently described from 51°13’N 57°75’E (S. Es y u n i n , pers. comm. 2010); Cyprus (previously treated as M. gibbosus). compare with Gr o m o v (2001). Only some records

4 Review Scorpions of Europe of Paruroctonus boreus (Gi r a r d , 1854) in south- Anthropochory is documented for many populations ern Alberta, , lie slightly further north (D. of E. flavicaudis in France (Va c h o n 1969); a similar Jo h n s o n , pers. comm. 2010). trend is observed for other Euscorpius. In total, eight native buthid species are docu- Subgenus Polytrichobothrius Bi r u l a , 1917 mented in Europe. Further increase in their number Euscorpius italicus (He r b s t , 1800), the larg- is not expected (but so unexpected were new species est species in the genus, is found from France to discoveries in Spain and Cyprus). Caucasus (including the Black coast of Turkey, Family EUSCORPIIDAE Georgia, and Russia), and also in Northern Africa This family is represented in Europe by its (Algeria, Morocco, Tunisia). For its distribution in unique genus Euscorpius Thorell, 1876, a European- Slovenia, see Fe t et al. (2001); for biology in , Mediterranean endemic, the core and diverse element see Co l o m b o (2006) and Cr u c i t t i et al. (2007). of European scorpiofauna, found in diverse habitats For distribution and biology in Switzerland, see from sea coast to the high Alps and Balkans. In many Br a u n w a l d e r (2001, 2005). A very low genetic di- areas, two or more congeners (or even “consubgen- vergence was detected by Fe t et al. (2006) between ers”) are found sympatrically. Currently, 17 species European and Anatolian populations of E. italicus are valid (see below) but their accepted ranges do (as compared to its sister species E. naupliensis). not include all known populations of Euscorpius. I This suggests a recent dispersal from a glacial re- estimate the total number of Euscorpius species in fugium, possibly even in historical times—which Europe as at least 22-24. Most populations from could explain a puzzling absence of E. italicus from Greece and the Balkans still require a detailed study. all Aegean and Mediterranean . The spe- Although family Euscorpiidae is represented cies is clearly anthropophilic and anthropochoric; in mountainous Asia (subfamily Scorpiopinae), Br a u n w a l d e r (2001, 2005) notes its predominant the closest taxa to Euscorspius (also in subfamily occurrence in human habitations. Many of its dis- Euscorpiinae) are Mexican genera Megacormus and junct populations are clearly introduced (e.g. in Iraq Plesiochactas (So l e g l a d , Si s s o m 2001). Some spe- and Yemen), and some probably became extinct, e.g. cies of Euscorpius are very clearly defined, but in from the Don River in , discovered many other cases traditional morphological criteria in the 1920s but never reported again (Fe t 1989). have not been sufficient. For recent reviews, see: For decades, E. italicus was considered the only spe- Fe t et al. (2004); Ka l t s a s et al. (2008), and Vi g n o l i , cies of its distinct subgenus. However, Ga n t e n b e i n Sa l o m o n e (2008; they also give a good review of et al. (2002), based on both molecular and morpho- known ecology). logical data, restored Euscorpius naupliensis (C.L. Ko c h , 1837), an endemic of Greece ( Subgenus Tetratrichobothrius Bi r u l a , 1917 and Zakynthos Island), and a senior synonym of E. Euscorpius flavicaudis (DeGe e r , 1778), the italicus zakynthi Ca p o r i a c c o , 1950 from Zakynthos. only species of this subgenus, is limited to the Western Its ecology was studied in the Peloponnese (Cr u c i t t i , Mediterranean: Spain, Baleares, southern France, Bu b b i c o , 2001; as E. italicus) and on Zakynthos , Italy, and . It is also found in North (Co l o m b o 2006). Based on the genetic divergence Africa, and has introduced populations in England between E. naupliensis and E. italicus, Ga n t e n b e i n and even Brazil and Uruguay. Its biology and ecology et al. (2002) suggested that their split predates the was studied by Be n t o n (1991, 1992, 1993) in the in- Messinian salin­ity crisis (5.2 Myrs ago). troduced population in England. High genetic homo- geneity (Ga n t e n b e i n et al. 2002) between its popu- Subgenus Alpiscorpius Ga n t e n b e i n et al., 1999 lations in mainland France and Corsica suggests that This subgenus was established to accommodate E. flavicaudis could be a relict that passed through the topology of a pilot 16S mtDNA phylogeny that a bottleneck and has been dispersing with humans. demonstrated a separate position of a group of spe-

5 Fet V. cies previously included in the subgenus Euscorpius Euscorpius gamma Ca p o r i a c c o , 1950, de- s. str. Morphology clearly confirms this compact scribed from Slovenia as a subspecies of E. ger­ group; a revision is in progress (Fe t et al.). The sub- manus, was elevated to species rank by Sc h e r a b o n et genus ranges from Austria to Caucasus. It includes al. (2000). E. gamma belongs to the “E. mingrelicus some high-mountain forms, up to 2170 m in East complex.” It is confirmed for SE Austria (Carinthia), Tirol (Austria) and 2250 m in Swiss Alps (E. ger­ Croatia, NE Italy (Friuli), and Slovenia. Details of manus), and 2200-2400 m in the Balkans (E. bero­ distribution are available for Slovenia (Fe t et al. ni). Currently, fivespecies are valid; the species-lev- 2001) and Austria (Ko m p o s c h et al. 2001; Ko m p o s c h el composition of Alpiscorpius is still unclear in the 2004, 2009). The species is included in the Red Data main part of its range, i.e. the Balkans and . Lists of Austria (Ko m p o s c h 2009). Euscorpius alpha Caporiacco, 1950 was el- In addition to E. beroni and E. gamma, “E. evated to species level from a subspecies of E. ger­ mingrelicus complex” includes a number of Balkan manus by Ga n t e n b e i n et al. (2000a). It is found in populations, for which species boundaries are not NW Italy (, Valle d’Aosta and Lombardy) yet clear. Their range includes Croatia, Bosnia and Switzerland. E. germanus and E. alpha are al- and Herzegovina, Montenegro, Kosovo, Serbia, lopatric, sibling Alpine species (Fe t et al. 2004). Macedonia, SW Bulgaria (Pirin), and NW Greece The degree of molecular divergence suggests that (Epiros). Several names are available for these pop- these two species have evolved in the area before the ulations, technically still under E. mingrelicus sub- Pleistocene glaciations, maybe since the orogenesis species (Ka l t s a s et al. 2008). of the Alps (Ga n t e n b e i n et al. 2000a). E. germanus Subgenus Euscorpius Th o r e l l , 1876 also has isolated populations in the Apennines; it The nominotypic subgenus, after separation of is not known whether they are relict or introduced Alpiscorpius (Ga n t e n b e i n et al. 1999), corresponds (Fe t et al. 2004). For distribution and biology in to what is often vaguely named “E. carpathicus com- Switzerland, see Br a u n w a l d e r (2001, 2005). plex.” For decades, most of its species were classi- Euscorpius beroni Fe t , 2000, was described from high Prokletije Mountains of Albania. It be- fied as subspecies of a (very polytypic) “E. carpathi­ longs to the Balkan portion of the “E. mingrelicus cus.” Currently, nine species are valid. The subgenus complex.” ranges from the Baleares in the west (E. balearicus) to the in the east (E. tauricus) and from the Euscorpius germanus (C. L. Ko c h , 1837). This traditional species was restricted by Ga n t e n b e i n et Southeast Alps in the north (E. tergestinus) to North al. (2000) only to some populations in Austria, NE Arica in the south (E. sicanus). Italy, Slovenia (east), and Switzerland. In Slovenia, Euscorpius balearicus Ca p o r i a c c o , 1950, an subspecies E. g. marcuzzii Ca p o r i a c c o , 1950 is found endemic of the Balearic Islands (Spain), was elevated sympatrically with E. gamma. Details of distribution to species rank, using morphological and molecular are available for Slovenia (Fe t et al. 2001), Austria data, by Ga n t e n b e i n et al. (2001), and redescribed (Ko m p o s c h et al. 2001; Ko f l e r 2002; Ko m p o s c h by Fe t , So l e g l a d (2002). 2004, 2009), and Switzerland (Br a u n w a l d e r 2001, Euscorpius carpathicus (Li n n a e u s , 1767). 2005). The species is included in the Red Data Lists Fe t , So l e g l a d (2002) studied the existing type of of Austria (Ko m p o s c h 2009). Linnaeus, and limited this traditional species to (Ke s s l e r , 1874) was Romania (Transylvanian Alps). As a result, all taxa described from Georgia. In Europe it is found, in my formerly classified as E. carpathicus or its subspe- opinion, only in the Northern Caucasus of Russia cies, belong to other species (of which some are yet (Krasnodar Province). The Balkan populations, still not described). The first mtDNA marker data for the technically included in E. mingrelicus as subspecies, Romanian population were published by Fe t et al. are under revision (see below). (2002).

6 Review Scorpions of Europe

Euscorpius concinnus (C. L. Ko c h , 1837) was Euscorpius tauricus (C.L. Ko c h , 1837) is restored to species level by Vi g n o l i et al. (2005), and known only from the southern coast of the Crimea reported only from Italy. Sa l o m o n e et al. (2007), in Peninsula in Ukraine; it was restored to species level a large DNA phylogeographic study for Italy, con- by Fe t (2003) based on DNA analysis, morphologi- firmed its separate status as compared to E. sicanus cal investigation pending. For biology observations, and E. tergestinus, and discussed a rather high de- see Ku k u s h k i n (2004). The species is listed as en- gree of divergence between these species. For biol- demic in the Red Data Book of Ukraine (Ve r v e s et ogy in Italy, see Co l o m b o (2006). al. 1999). Euscorpius hadzii Ca p o r i a c c o , 1950 was Euscorpius tergestinus (C.L. Ko c h , 1837) was elevated to species rank and redescribed by Fe t , redescribed by Fe t , So l e g l a d (2002). Vi g n o l i et So l e g l a d (2002) (neotype from Albania). This large, al. (2005, 2007) separated two Italian species from dark Balkan species is found from Albania to SW this taxon, E. concinnus and E. oglasae. E. tergesti­ Bulgaria, often in high mountains. It is easily identi- nus embraces a number of former subspecies from fiable by its additional neobothriotaxic trichobothria Italy, and is confirmed for Albania, Croatia, Italy, on external aspect of pedipalp patella. A population Montenegro, San Marino, and Slovenia. For biology of small-sized E. hadzii (described as E. carpathicus in Italy, see Co l o m b o (2006). In a detailed phylo- lagostae Ca p o r i a c c o , 1950 from Lastovo Island) in- geographic study, Sa l o m o n e et al. (2007) confirmed habits southern Croatian islands in the . separate status of E. tergestinus as compared to E. In Bulgaria, E. hadzii is found only in the southwest sicanus and E. concinnus in Italy. Three disjunct (Fe t , So l e g l a d 2007) as as Zemen, where populations in Austria, likely introduced, were stud- it apparently is closely allopatric with Euscorpius sp. ied using mtDNA markers by Hu b e r et al. (2001); of Stara Planina. they are included in the Red Data Lists of Austria Euscorpius koschewnikowi Bi r u l a , 1900 is (Ko m p o s c h 2004, 2009). known only from Mt Athos in Greece. It was restored In addition to nine species listed above, place- to species rank from a subspecies of E. car­pathicus ment of many European populations of “E. carpathi­ by Fe t , So l e g l a d (2002) who studied the types. cus complex” remains unclear. Vi g n o l i et al. (2007) Euscorpius oglasae Ca p o r i a c c o , 1950 was el- studied populations from Corsica and France listed evated to species rank from a subspecies of E. car­ as E. carpathicus corsicanus Ca p o r i a c c o , 1950 and pathicus by Vi g n o l i et al. (2007). The species is en- E. c. niciensis (C.L. Ko c h , 1841), respectively, and demic to Island (Tuscan Archipelago, indicated their differences from known valid spe- Italy). cies. In a pilot mtDNA phylogeny of Ga n t e n b e i n et Euscorpius sicanus (C. L. Ko c h , 1837), de- al. (2001), French (Alpes-Maritimes) populations scribed from , was restored to species rank from grouped with those of , i.e. E. concinnus. a subspecies of E. carpathicus­ by Fe t et al. (2003) who The Balkan and Aegean populations, which also synonymized with it seven former subspecies, all are currently under revision (Fe t et al. in progress), from Italy. This species absorbed part of the popula­ include Croatia, Bosnia & Herzegovina, Montenegro, tions listed by earlier authors (Ki n z e l b a c h 1975) as Kosovo, Serbia, Macedonia, Bulgaria, and Greece. “E. mesotrichus Hadži” (an invalid name). Sa l o m o n e Several names are available for some of these popu- et al. (2007) confirmed a separate status of E. sica­ lations (i.e. E. carpathicus candiota Bi r u l a , 1903 nus as compared to E. concinnus and E. tergestinus. from Crete), potentially of the species rank; others It is widespread in central and (with likely represent undescribed species (for details, see Sicily and Sardinia) and Greece (Thessaly, Sporades, Ka l t s a s et al. 2008). Especially diverse and compli- Peloponnese), and is also found in Malta, North Africa, cated is Greek fauna, where a pilot DNA phylogeny and Madeira. For biology in Italy, see Co l o m b o (2006), indicates several species that were previously unrec- and in Greece, Cr u c i t t i , Bu b b i c o (2001). ognized (Po u l i k a r a k o u et al. 2009).

7 Fet V.

Family IURIDAE The family placement of Belisarius has been Both genera of Iuridae are recorded in Europe controversial. Early researchers (Ki n z e l b a c h 1975), but Calchas Bi r u l a , 1899 is represented only in largely by geographic association, suggested a close the easternmost islands of Greece: the recently de- relationship to Euscorpius, which is not confirmed. scribed Calchas gruberi Fe t et al., 2009 (formerly Lo u r e n ç o (1998) placed Belisarius in a new fam- reported as C. nordmanni) is found in the Greek is- ily, . Most recently, Belisarius was lands Megisti and Samos as well as in Anatolia (Fe t placed in Chactidae (otherwise limited to the New et al. 2009). World) by Fe t , So l e g l a d (2003). They provided The genus Iurus Th o r e l l , 1876, taxonomy of a justification for its specific placement in a South which for many years was controversial but neglected, American subfamily Brotheinae, with a remark- includes the largest of the European scorpions, Iurus able zoogeographic disjunction. (Even if Belisarius dufoureius (Br u l l é , 1832). Pa r m a k e l i s et al. (2006b) is assigned to Troglotayosicidae, the issue of South studied phylogeography of Iurus using mtDNA mark- American relationship, with genus Troglotayosicus, ers, and demonstrated that populations of Peloponnese, still remains.) Kithyra, and Crete formed a well-supported clade as This review should further complement and cor- opposed to the Eastern Aegean and Anatolian popula- rect the scorpion data available in Fauna Europaea tions. In an extensive recent revision of Iurus, Ko v a ř í k (Cr u c i t t i 2004) database. It reports for Europe two et al. (2010) confirmed this conclusion and provided exotic buthid species, maculatus (DeGe e r , a morphological basis as they limited I. dufoureius 1778) and gracilis (La t r e i l l e , 1804). I. to the “western clade.” The populations of Eastern maculatus is a subcosmopolitan scorpion, introduced Aegean Greek islands (Rhodes, Karpathos, Samos) across the globe; it was once reported for Spain but belong to another, possibly new Iurus species. Based no resident population is confirmed. C. gracilis, on sequence divergence, Pa r m a k e l i s et al. (2006b) which belongs to a large genus, has an suggested that Iurus has been differentiating in the introduced population on Tenerife, Canary Islands southern region of Agaeis at least since the Miocene (also reported as C. nigrescens; Bá e z Fu m e r o (ca. 8 Mya). For details on distribution and morphol- 1984), in the coastal zone of Santa Cruz town. ogy of I. dufoureius, and the genus Iurus in general, An unidentified species ofEuscorpius is also known see Ko v a ř í k et al. (2010). from Tenerife, likely an introduction. Macaronesian fauna is partially covered by the Fauna Europaea Family CHACTIDAE project (except Cape Verde Islands); however, it Probably the most unique element of European does not seem to include any native scorpions. E. i m o n scorpiofauna is a monotypic genus Belisarius S , sicanus record from Madeira (Fe t et al. 2003) could 1879, with its blind (eyeless) species B. xambeui be either introduction from the Mediterranean, or a Si m o n , 1879, from the of France (Pyrénées- local relict. No scorpions have been reported from Orientales) and Spain (Catalunya: Girona and the Azores. Barcelona). Not much study has been done on this It is remarkable that the “hidden diversity” of enigmatic scorpion since its original description. Its European fauna remained unrecognized for over 150 distribution, with a map, has been last summarized by years of intensive arachnological research. The cur- La c r o i x (1992), and it seems to be more common in rent “splitting” trend in scorpion taxonomy is, how- Spanish part of the Pyrenees. Belisarius xambeui has ever, only a response to former “lumping” trend, and been characterized as an endogean, hygrophile lapidi- became possible due to availability of new characters colous species, found in mountain forests (850-1300 and techniques, as well as of new material. Indeed, a m asl) in endogean subterranean habitats. It is the only bothering taxonomic and zoogeographic puzzle for a cave scorpion in Europe, known from three caves in long time was absence of diversity in European scor- France and 15 caves in Spain (La c r o i x 1992). pions; now we see that they behave as any normal

8 Review Scorpions of Europe fauna, with a realistic combination of local, Asian and Anikin, Janet Beccaloni, Petar Beron, Gergin Blagoev, Alberto Bonacina, Nikos Botsaris, Matt Braunwalder, Michael Brewer, African elements, ancient relicts and recent dispersals, John Cloudsley-Thompson, Marco Colombo, Pierangelo Cru- sympatric congeners and isolated endemics. The total citti, Hieronymos Dastych, Christo Deltschev, Dobrin Dobrev, number of native scorpion species in Europe (exclud- Jason Dunlop, Gerard Dupré, Sergey Esyunin, Galina Fet, Elizabeth Fet, Simon Fet, Benjamin Gantenbein, Matt Graham, ing Macaronesia but including Cyprus) could be now Alexander Gromov, Jürgen Gruber, Dietmar Huber, Dan John- estimated over 35 (Buthidae, 8; Euscorpiidae, 22-24; son, Dimitris Kaltsas, Ivo Karaman, Ragnar Kinzelbach, Marian Komnenov, Christian Komposch, František Kovařík, Mykola Chactidae, 1; Iuridae, 3); the genus Euscorpius is still Kovblyuk, Viktor Krivochatsky, Jean-Bernard Lacroix, Wilson under an intensive revision, especially in the Balkans, Lourenço, Yuri Marusik, Arie van den Mejden, Kirill Mikhailov, Moysis Mylonas, Ivan Pandourski, Aris Parmakelis, Gary Polis, and will clearly yield more species. Valentin Popa, Alexi Popov, Jan Ove Rein, Carles Ribera, Vlado Sakalian, Patrick Schembri, Bernhardt Scherabon, Boris Sket, Michael Soleglad, František Šťahlavský, Iasmi Stathi, Pavel Acknowledgements: I thank dozens of friends and colleagues Stoev, Rolando Teruel, Max Vachon, Marco Valle, Milen Vassi- who, for decades, enjoyably helped me and collaborated in Eu- lev, Valerio Vignoli, and Eric Ythier. I am grateful to Prof. Vassil ropean scorpion studies, including (but not limited to): Vasily Golemanski for inviting me to write this brief review.

References

An i k i n V. V., E. A. Ki r e e v 1997. Ecological-floristic description Cr u c i t t i P., F. Bu b b i c o 2001. Contributo alla conoscenza del of the habitats of some rare and species genere Euscorpius Thorell, 1876 nel Peloponneso sud- of Lower Povolzhye. – In: Materials of the International occidentale (Scorpiones, Euscorpiidae). – Bolletino del Research-Practical Conference “The Problems of Preserva­ Museo Civico di Storia Naturale di Venezia, 52: 41-57. tion of the in the Arid of Russia”. Vol- Cr u c i t t i P., S. Bu c c e d i , F. Em i l i a n i , M. Ma l o r i , G. Ro t e l l a , and L. gograd, 11-17 September 1998, pp. 106-107 (in Russian). Tr i n g a l i 2007. Ecologia dello scorpione Italiano (Euscorpius Bá e z Fu m e r o M. 1984. Arácnidos. – In J. J. Bacallado Aránega, italicus) in un’area protetta dell’Italia Centrale (Arachnida, J. J. (Еd.): Fauna (Marina y Terrestre) del Archipiélago Scorpiones, Euscorpiidae). – Aldrovrandia, 3: 19-30. Canario, 108-116. Edirca: Las Palmas de Gran Canaria. Cr u c i t t i P., F. Ma r i n i 1987. Ecologia di Mesobuthus gibbosus Be n t o n T. G. 1991. The life history of Euscorpius flavicaudis (Brullé, 1832) nel Pindo (Epiro e Tessaglia, Grecia) (Scor- (Scorpiones, Chactidae). – Journal of Arachnology, 19 piones: Buthidae). – Giornalo italiano Entomologico, 3: (2): 105-110. 291-303. Be n t o n T. G. 1992. The ecology of the scorpion Euscorpius Da v y g o r a A. V., A. V. Ru s a k o v 2001. About northern limits of flavicaudis in England. – Journal of Zoology, London, Mesobuthus eupeus and Galeodes pallasi spreading in the 226 (3): 351-368. south steppes. – In: Biodiversity and bioresources of Be n t o n T.G. 1993. The reproductive ecology of Euscorpius fla­ Urals and adjacent territories. Gaspompechat: Orenburg vicaudis in England. – Memoirs of Museum, (in Russian), 210-211. 33 (2): 455-460. Fe t V. 1989. A catalogue of scorpions (: Scorpiones) of Bo n a c i n a A. 1980. Sistematica specifica e sottospecifica del the USSR. – Rivista del Museo Civico di Scienze Naturali complesso “Euscorpius germanus” (Scorpiones, Chac- “Enrico Caffi” (Bergamo), 13 (1988): 73-171. tidae). – Rivista del Museo Civico di Scienze Naturali Fe t V 2003. The Crimean scorpion, Euscorpius tauricus (C. „Enrico Caffi“ (Bergamo), 2: 47-100. L. Koch, 1837) (Scorpiones: Euscorpiidae): an endemic Br a u n w a l d e r M. E. 2001. Scorpions of Switzerland: summary of species supported by mitochondrial DNA evidence. – Ar­ a faunistic survey. – In: Fe t V., P.A. Se l d e n (Еds.): Scorpi- thropoda Selecta (Moscow), 11 (4): 271-276. ons 2001. In memoriam Gary A. Polis. Burnham Beeches, Fe t V., M. E. Br a u n w a l d e r 2000. The scorpions (Arachnida, Bucks: British Arachnological Society): 279-286. Scorpiones) of the Aegean area: current problems in tax- Br a u n w a l d e r M.E. 2005. Fauna Helvetica 13. Scorpiones onomy and biogeography. – Belgian Journal of Zoology, (Arachnida). Neuchâtel (Centre suisse de cartographie de 130 (Suppl. 1), 17-22. la Faune/Schweizerische Entomologische Gesellschaft), Fe t V., H.El-He n n a w y , J.L.Cl o u d s l e y -Th o m p s o n and M.E. 240 p. (parallel text in German and Italian). Br a u n w a l d e r . 2009. The first observation on scorpion Co l o m b o M. 2006. New data on distribution and ecology of biogeography by Aristotle. – Boletín de la Sociedad Ento­ seven species of Euscorpius Thorell, 1876 (Scorpiones: mológica Aragonesa, 44: 147-150. Euscorpiidae). – Euscorpius, 36: 1-40. Fe t V., B. Ga n t e n b e i n , E.V. Fet and V. Po p a . 2002. Euscorpius Cr u c i t t i P. (ed.) 2004. Fauna Europaea: Scorpiones. Fauna Eu- carpathicus (Linnaeus, 1767) from Roma­nia (Scorpiones: ropaea version 1.1, http://www.faunaeur.org. Euscorpiidae): mitochondrial DNA data. – Biogeographica, Cr u c i t t i P., R. An o t e l l i and S. Fa t t o r i n i 1994. Fattori ecologici 78 (4), 141-147. della distribuzione di Buthus occitanus (Amoreux, 1789) Fe t V., B. Ga n t e n b e i n , A. Ka r a t a ş and Ah. Ka r a t a ş 2006. An in Europa (Scorpiones: Buthidae). Giornalo italiano En­ extremely low genetic divergence across the range of Eu­ tomologico, 7: 57-66. scorpius italicus (Scorpiones: Euscorpiidae). – Journal of Arachnology, 34 (1): 248–253.

9 Fet V.

Fe t V., M. Ku n t n e r and B. Sk e t . 2001. Scоrpions of Slovenia: occitanus as model organism. – Molecular Phylogenetics a faunistic and biogeographical survey. – In: Fe t V., P. A. and Evolution, 28 (1): 119-130. Se l d e n (Еds.): Scorpions 2001. In memoriam Gary A. Ga n t e n b e i n B., M. E. So l e g l a d and V. F e t 2001. Euscorpius Polis. Burnham Beeches, Bucks: British Arachnological balearicus Caporiacco, 1950, stat. nov. (Scorpiones: Eu- Society), 255-256. scorpiidae): molecular (allozymes and mtDNA) and mor- Fe t V., M. E. So l e g l a d 2002. Morphology analysis supports phological evidence for an endemic Balearic Islands species. presence of more than one species in the Euscorpius – Organisms, Diversity & Evolution, 1 (4): 301-320. carpathicus complex (Scorpiones: Euscorpiidae). – Eu­ Ga n t e n b e i n B., M. E. So l e g l a d , V. Fe t , P. Cr u c i t t i and E. V. Fe t scorpius, 3: 1-51. 2002. Euscorpius naupliensis (C.L. Koch, 1837) (Scorpi- Fe t V., M. E. So l e g l a d 2007. Fauna and zoogeography of scor- ones: Euscorpiidae): elevation to the species level justified pions (Arachnida: Scorpiones) in Bulgaria. – In: Fe t V., by molecular and morphology data. – Revista Inerica de A. Po p o v (Еds.): Biogeography and Ecology of Bulgaria. Aracnologia, 6: 13-43. Springer, 405-422. Gr o m o v A. V. 2001. The northern boundary of scorpions in Cen- Fe t V., M. E. So l e g l a d and B. Ga n t e n b e i n 2004. The Euroscor- tral Asia. – In: Fe t V., P.A. Se l d e n (Еds.): Scorpions 2001. pion: Taxonomy and systematics of the genus Euscorpius In memoriam Gary A. Polis. Burnham Beeches, Bucks: Thorell, 1876 (Scorpiones: Euscorpiidae). – Euscorpius, British Arachnological Society, 301-306. 17: 47-59. Hu b e r D., B. Ga n t e n b e i n , V. Fe t and B. Sc h e r a b o n 2001. Euscor­ Fe t V., M. E. So l e g l a d , B. Ga n t e n b e i n , V. Vi g n o l i , N. Sa l o m o n e , pius carpathicus (L.) from Austria (Scorpiones: Euscorpii- E. V. Fe t and P. J. Sc h e m b r i 2003. New molecular and dae): phylogenetic position clarified by mitochondrial DNA morphological data on the Euscorpius carpathicus species analysis. – In: Fe t V., P. A. Se l d e n (Еds.): Scorpions 2001. complex (Scorpiones: Euscorpiidae) from Italy, Malta, and In memoriam Gary A. Polis. Burnham Beeches, Bucks: Greece justify the elevation of E. c. sicanus (C. L. Koch, British Arachnological Society, 273-278. 1837) to the species level. – Revue suisse de Zoologie, 110 Ka l t s a s D., M. My l o n a s 2006. The population structure of Me­ (2): 355-379. sobuthus gibbosus (Scorpiones: Buthidae) on Koufonisi Fe t V., M. E. So l e g l a d and F. Ko v a ř í k 2009. Etudes on iurids, Island (central Aegean Archipelago, Greece). – Euscorpius, II. Revision of genus Calchas Birula, 1899, with the 55: 1-8. description of two new species (Scorpiones: Iuridae). – Ka l t s a s D., I. St a t h i and V. Fe t 2008. Scorpions of the Eastern Euscorpius, 82: 1−72. Mediterranean. – In: Ma k a r o v S.E., R.N. Di m i t r i j e v i ć Ga n t e n b e i n B. 2004. The genetic population structure of Buthus (Еds.): Advances in Arachnology and Developmental Bi- occitanus (Scorpiones: Buthidae) across the Strait of ology. Papers dedicated to Professor Božidar P.M. Ćurčić. Gibraltar: calibrating a molecular clock using nuclear al- Belgrade-Vienna-Sofia, 209-246. lozyme variation. – Biological Journal of Linnean Society, Ka l t s a s D., I. St a t h i and M. My l o n a s 2006. The effect of insu- 51: 519-534. larity on the seasonal population structure of Mesobuthus Ga n t e n b e i n B., V. Fe t , M. Ba r k e r and A. Sc h o l l 2000. Nuclear gibbosus (Scorpiones: Buthidae). – Euscorpius, 44: 1-8. and mitochondrial markers reveal the existence of two Ka l t s a s D., I. St a t h i and M. My l o n a s 2008. The foraging activity parapatric scorpion species in the Alps: Euscorpius ger­ of Mesobuthus gibbosus (Scorpio­nes: Buthidae) in central manus (C. L. Koch, 1837) and E. alpha Caporiacco, 1950, and south Aegean archipelago. – Journal of Natural His­ stat. nov. (Euscorpiidae). – Revue suisse de Zoologie, 107 tory, 42 (5): 513-527. (4): 843-869. Ki n z e l b a c h R. 1975. Die Skorpione der Ägäis. Beiträge zur Ga n t e n b e i n B., V. Fe t and A.V. Gr o m o v 2003. The first DNA Systematik, Phylogenie und Biogeographie. – Zoologische phylogeny of four species of Mesobuthus Vachon, 1950 Jahrbücher, Abteilung für Systematik, 102: 12-50. (Scorpiones: Buthidae) from . – Journal of Arach­ Ko f l e r A. 2002. Zum Vorkommen des Deutschen Skorpions nology, 31(3): 412-420. Euscorpius germanus (C. L. Koch, 1837) in Osttirol Ga n t e n b e i n B., V. Fe t , C. La r g i a d è r and A. Sc h o l l 1999. First (Arachnida, Scorpiones: Euscorpiidae). – Gredleriana, DNA phylogeny of the genus Euscorpius Thorell 1876 2: 137-145. (Scorpiones, Euscorpiidae) and its bearing on the taxonomy Ko m p o s c h C. 2004. Die Skorpione Osterreichs (Arachnida, Scor- and biogeography of this genus. – Biogeographica, 75 piones). – Denisia, 12, zugleich Kataloge der Oberoster- (2): 49-65. reichischen Landesmuseen Neue Serie 14: 441-458. Ga n t e n b e i n B., P.D. Ke i g h t l e y 2004. Rates of molecular evolu- Ko m p o s c h C. 2009. Rote Liste der Skorpione (Scorpiones) Ös- tion in nuclear genes of east Mediterranean scorpions. – terreichs. – In: Zulka, K.-P. (Ed.): Rote Listen gefährdeter Evolution, 58 (11): 2486-2497. Tiere Österreichs: Flusskrebse, Köcherfliegen, Skorpi- Ga n t e n b e i n B., C. Kr o p f , C.R. La r g i a d è r and A. Sc h o l l 2000. one, Weberknechte, Zikaden. (Grüne Reihe Bd. 14/3). Molecular and morphological evidence for the presence of Böhlau Verlag, Wien: Bundesministerium für Land- und a new Buthid taxon (Scorpiones: Buthidae) on the Island of Forstwirtschaft, Umwelt und Wasserwirtschaft, 359-395. Cyprus. – Revue suisse de Zoologie, 107 (1): 213-232. Ko m p o s c h C., B. Sc h e r a b o n and V. F e t 2001. Scorpions of Ga n t e n b e i n B., C. R. La r g i a d è r 2002. Mesobuthus gibbosus Austria. – In: Fe t V., P. A. Se l d e n (Еds.): Scorpions 2001. (Scorpiones: Buthidae) on the island of Rhodes – Hybrid- In Memoriam Gary A. Polis. Burnham Beeches, Bucks: ization between Ulysses’ stowaways and native scorpi- British Arachnological Society, 267-272. ons? – Molecular Ecology, 11: 925-938. Ko v a ř í k F., M. E. So l e g l a d , V. Fe t and E. A. Ya ğ m u r 2010. Ga n t e n b e i n B., C. R. La r g i a d è r 2003. The phylogeographic im- Etudes on iurids, III. Revision of the genus Iurus Thorell, portance of the Strait of Gibraltar as a gene flow barrier to 1876, with the description of two new species (Scorpiones: terrestrial : a case study with the scorpion Buthus Iuridae). – Euscorpius. (In press).

10 Review Scorpions of Europe

Ku k u s h k i n O. V. 2004. Data to ecology of the Crimean Scorpion pathicus complex” (Scorpiones: Euscorpiidae) in Italy. – in the South-West Crimea. – Vestnik zoologii, 38 (1): 74. Molecular Phylogenetics and Evolution, 43 (2): 502-514. (In Russian). Sc h e r a b o n B., B. Ga n t e n b e i n , B., V. Fe t , M. Ba r k e r , M. Ku n t - La c r o i x J.-B. 1992. Faune de France (Arachnida : Scorpionida). n e r , C. Kr o p f and D. Hu b e r . 2000. A new species of scor- 7e Note. B – Genre Belisarius Simon, 1879. Arachnides, pions for Austria, Italy, Slovenia and Croatia: Euscorpius 14: 27-38. gamma Caporiacco, 1950, stat. nov. (Scorpiones, Euscor- Lo u r e n ç o W. R. 1998. Panbiogeographie, les distributions disjon- piidae). – Ekologia, (Bratislava), 19: 253-262. tes et le concept de familie relictuelle chez lesScorpions. – So l e g l a d M. E., V. Fe t 2003. High-level systematics and phy- Biogeographica, 74 (3): 133-144. logeny of the extant scorpions (Scorpiones: Orthosterni). – Lo u r e n ç o W. R. 2002. Considérations sur les modèles de dis- Euscorpius, 11: 1-175. tribution et différentiation du genre Buthus Leach, 1815, So l e g l a d , M. E., W. D. Si s s o m 2001. Phylogeny of the family avec la description d’une nouvelle espèce des montagnes Euscorpiidae Laurie, 1896: a major revision. – In: Fe t V., du Tassili des Ajjer, Algérie (Scorpiones, Buthidae) . – P.A. Se l d e n (Eds.): Scorpions 2001. In Memoriam Gary Biogeographica, 78 (3): 109-127. A. Polis. Burnham Beeches, Bucks: British Arachnological Lo u r e n ç o W. R. 2003. Compléments â la faune de scorpions Society, 25-112. (Arachnida) de l’Afrique du Nord, avec des considéra- Тe r u e l R., V. Fe t and L. F. d e A r m a s 2004. A note on the scorpions tions sur le genre Buthus Leach, 1815. – Revue suisse de from the Pirin Mountains, Southwestern Bulgaria (Scorpi- Zoologie, 110 (4): 875-912. ones: Buthidae, Euscorpiidae). – Euscorpius, 14: 1-11. Lo u r e n ç o W. R., M. Va c h o n 2004. Considérations sur le genre Тe r u e l R., J. L. Pé r e z -Bo t e . 2005. Complementos a la descripcion Buthus Leach, 1815 en Espagne, et description de deux de Buthus ibericus Lourenço & Vachon 2004 (Scorpiones: nouvelles espèces (Scorpiones, Buthidae). – Revista Ibérica Buthidae). – Boletín de la Sociedad Entomológica Arago­ de Aracnología, 9: 81–94. nesa, 37: 273-277. Lo u r e n ç o W. R, E. A. Ya ğ m u r & H. Ko c 2010. A new species of Va c h o n M. 1969. Nouvelles remarques sur la répartition en Buthus Leach, 1815 from Cyprus (Scorpiones, Buthidae). – France métropolitaine du Scorpion méditerranéen Euscor­ Turkish Journal of Zoology. (In press). pius flavicaudis (Geer) (famille des Chactidae). – Bulletin Pa r m a k e l i s A., I. St a t h i , M. Ch a t z a k i , L. Sp a n o s , C. Lo u i s and Scientifique de Bourgogne, 26: 189-202. M. My l o n a s 2006a. Evolution of Mesobuthus gibbosus Ve r v e s Y. G., L. A. Kh r o k a l o . R. S. Pa v l y u k and P. G. Ba l a n (Brullé, 1832) (Scorpiones: Buthidae) in the northeastern 1999. Criteria of estimating species of invertebrate Mediterranean region. – Molecular Ecology, 15 (10): for including in the Red Book (in Ukrainian). Zapovidna 2883-2894. sprava v Ukraini, 5 (2): 48-58. Pa r m a k e l i s A., I. St a t h i , L. Sp a n o s , C. Lo u i s and M. My l o n a s Vi g n o l i V., N. Sa l o m o n e 2008. A review of and additions to 2006b. Phylogeography of Iurus dufoureius (Brullé, 1832) the current knowledge of the scorpion genus Euscorpius (Scorpiones, Iuridae). – Journal of Biogeography, 33: Thorell, 1876 (Scorpiones, Euscorpiidae). – Fragmenta 251–260. Entomologica, 40 (2): 189-228. Po u l i k a r a k o u S., I. Mp l a z a k i s , A. Pa r m a k e l i s , I. St a t h i , V. Vi g n o l i V., N. Sa l o m o n e , T. Ca r u s o and F. Be r n i n i 2005. The Fe t and M. My l o n a s 2009. Insights into the molecular Euscorpius tergestinus (C.L. Koch, 1837) complex in Italy: phylogeny of Euscorpius (Scorpiones: Euscorpiidae) in Biometrics of sympatric hidden species (Scorpiones: Eu- Greece. – Abstracts/25th European Congress of Arachnol­ scorpiidae). – Zoologischer Anzeiger, 244: 97-113. ogy, Alexandropouli, Greece, 16-21 August 2009. Vi g n o l i V., N. Sa l o m o n e , F. Ci c c o n a r d i and F. Be r n i n i 2007. The Sa g a l a e v V. A. 1997. Scorpions of Shcherbakovka. – Rodnoy scorpion of Montecristo, Euscorpius oglasae Di Capori- kray, (Volgograd), 3. (In Russian). acco, 1950, stat. nov. (Scorpiones, Euscorpiidae): a paleo- Sa l o m o n e N., V. Vi g n o l i , F. Fr a t i and F. Be r n i n i 2007. Species of the Tuscan Archipelago (northern Tyrrhenian, boundaries and phylogeography of the “Euscorpius car- Italy). – Comptes Rendus Biologies, 330 (2): 113-125.

Accepted: 12.02.2010

11 Fet V. Скорпионите в Европа

В. Фет

(Резюме)

This brief review summarizes the studies in systematics and zoogeography of European scorpions. The current “splitting” trend in scorpion taxonomy is only a reasonable response to the former “lumping.” Our better understanding of scorpion systematics 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 easily 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-anthropochoric) scorpion distribution in Europe is in Saratov Province, Russia, at 50°40’54”N, for Mesobuthus eupeus (Buthidae).

12