Hydroporus libanus Régimbart, 1901: 101 ZIMMERMANN (1931: 155) ZAITZEV (1953b: 172 1972: 182 WEWALKA (1989: 149) Hydroporus jacobsoni Zaitzev, 1927: 17 ZAITZEV (1933: 335 1946: 88 1953a: 93 Hydroporus longicornis Sharp, 1871 FICHTNER (1974: 189) New species of the Hydroporus longulus-group from Iran, Armenia and Turkey with a synopsis of the group (Coleoptera: Dytiscidae) Fery, Hans Acta Entomologica Musei Nationalis Pragae 2009 2009-12-15 49 2 529 558 6MJFR Regimbart, 1901 Regimbart 1901 [393,809,1377,1402] Insecta Dytiscidae Hydroporus Animalia Coleoptera 9 538 Arthropoda species libanus      Hydroporus libanus Régimbart, 1901: 101(original description). –  ZIMMERMANN (1931: 155)(misidentification). –  ZAITZEV (1953b: 172;  1972: 182) (misidentification). –  WEWALKA (1989: 149)(redescription).    Typelocality. Lebanon, Brummānā (= Broumana), ca. 10 kmE Beyrouth, ca. 33.88N 35.62E{25}.  Type material(not studied). LECTOTYPE: J, ‘Mt. Liban’, ‘Broumana’ {25}, an illegible small label, probably a male sex symbol, ‘Type’ [red, printed], ‘ Hydroporuslibanicus [sic!] Rég. n. sp., (Régim - vid.)’ [probably hw Régimbart], ‘Museum Paris, Coll. M. Pic’ [printed], ‘ Hydroporus libanusdet. Jäch [19]90’ [added by M.A. Jäch in 1990] ( MNHN). Notes: I have not been able to locate the lectotypenor other syntypeswhen visiting MNHN several times during the last years.  Notes on the type specimens: RÉGIMBART (1901: 102)stated that he studied ‘un très petit nombre d’exemplaires’ [= a very small number of specimens].These (former) syntypeshave not been found and, thus, not studied by WEWALKA (1989). Nevertheless, they must be regarded as paralectotypes. WEWALKA (1989: 149)did not explicitly designate a lectotype, but instead wrote ‘Type J: [...]’. According to Article 74.5 of the ICZN (1999), this proceeding has to be treated as a valid designation of the lectotype. I am not absolutely sure about the typelocality because another village with the name Brummānā is located ca. 20 kmdue NE of the first one (ca. 34.00N 35.80E). This locality also fits RÉGIMBART’ s (1901: 102) description: ‘sur le versant Ouest de ce massif montagneux [= on the western slope of this mountain range]’. It seems impossible to find out which of these two villages is meant by Régimbart, however, the first one was certainly easier to access in 1900, and is therefore assumed to be the typelocality.  RÉGIMBART (1901: 101)compared  H. libanuswith  H. obsoletus. ZIMMERMANN (1931: 155), after having studied specimens from Lebanon, stated that  H. libanusis extremely similar to  H. obsoletusand suspected that the former might be a subspecies of the latter. However, already WEWALKA (1989: 149)stated that the similarity of both species ‘is only superficial’. In addition, I have found one specimenof  H. obsoletusstanding under  H. libanusin the collection of the ZSM with the following labels: ‘Samml. A. Zimmermann’ [printed], ‘ Hydroporus libanus?’ [hw?], ‘  Hydroporus obsoletusAubé, det. G. Wewalka [19]89’ [hw Wewalka in part]. This is most probably the specimen that Zimmermann studied before 1931, causing his misidentification of  H. libanus.  Additional material studied. TURKEY:1J, ‘TR 24.5.1987, Amanos Geb. [= Amanus mountains], leg. Jäch (21)’ [printed] {26}, ‘ Hydroporus libanus Reg, det. G. Wewalka [19]87’ ( NMW). WEWALKA (1989: 149)provided the following additional data for this specimen: ‘Soğukoluk, 15 kmS Iskenderun’ (ca. 36.49N 36.17E, ca. 300 kmdue N of the typelocality). The Amanus mountains are also known as Nur Dağları or Gavur Dağları and are situated north of Hatay(= Antakya).  Original description.After the original description, only a few authors dealt with  H. libanus: ZIMMERMANN (1931: 155)misidentified the species; ZAITZEV (1953b: 172; 1972: 182) repeated Zimmermann’s words, certainly without having studied any specimen. A few other authors listed the species in catalogues or cited it in distributional lists. Only WEWALKA (1989: 149)treated this species in more detail, being seemingly the first author after Régimbart who saw one of the syntypesfrom Lebanonand another specimen from southern Turkey(Amanus mountains). However, even WEWALKA (1989)gave only a few descriptive notes and compared the species shortly with  H. obsoletusand  H. dobrogeanusIeniştea, 1962. This is why I present here a translation of the first part of Régimbart’s Latin description and add some further remarks: ‘Length 3⅔ mm. Oblong-oval, elongate, parallel, depressed, with very fine and obsolete reticulation, entire surface reddish brown, legs and antennae of same colour; head large, rather densely but little strongly punctate, both sides with large and deep grooves anteriorly; pronotum transverse, short, rather strongly and little densely punctate, disc smooth, sides curved and strongly but rather narrowly beaded, posterior angles slightly obtuse, not absent [in the meaning of ‘angles perceptible, not totally rounded’]. Elytra oblong, parallel, posteriorly not narrowed but obtusely rounded, strongly and little densely and almost regularly punctate, punctures with very short hairs [setae]. Ventrally punctation little dense, but larger on coxae [metacoxal plates] and metasternum [metaventrite], more weak on epipleura; legs robust, anterior tibiae large and triangular, tarsi broad, widened, particularly at base.’ ( RÉGIMBART 1901).   Differential diagnosis.The following differential diagnosis is based only on the specimen from the Amanus mountains, because material from Lebanoncould not be studied. I give chiefly the differences from  H. jelineki sp. nov.: Habitus distinctly more elongate, sides less parallel, more evenly rounded over large parts of length. Maximum width of body near middle of total length, between first and second third of elytral length. Discontinuity of body outline in dorsal view between pronotum and elytra weak, because pronotum with sides already converging from posterior angles, maximum width at posterior angles. Punctation on disc of elytra coarser, distance between punctures sometimes less than their diameter, puncture lines rather indistinct. Entire upper surface brownish, clypeus and disc of pronotum slightly darker; head with vertex lighter brownish and parts behind each eye blackish. Venter with meso- and metaepisterna, metaventrite and metacoxal plates blackish brown; genae and gula dark brownish, of same colour; prosternum, epipleura and abdomen brownish, latter with some diffuse weakly darkened areas; appendages brownish; contrast between their colour and that of rest of surface not prominent. JJ. Median lobe of aedeagus ( Fig. 3; see also WEWALKA 1989: 152) in ventral view tapering to apex, far before apex widened and tip very broadly rounded. Paramere as in Fig. 10. Anterior protarsal claws less straight than in  H. jelineki sp. nov.Sucker cups on first tarsomere of pro- and mesotarsomeres present.  ♀♀. Possibly studied by RÉGIMBART (1901), but not described in the literature. I have not seen any female specimens that could be associated with  H. libanus.   Measurements.TL: 3.85 mm, MW: 1.80 mm, TL/MW: 2.23, IO/MP: 0.5.   Distribution.Endemic for Lebanonand southern Turkey, Hatay province( Fig. 26).   Biology.No details are known about the preferred habitats of  H. libanus. It seems to occur like other members of the  H. longulus-group in mountainous regions.    Hydroporus holzschuhi sp. nov. Typelocality. Turkey, Muş province, near Buğlan geçidi [= Buğlan pass], ca. 13 kmdue WNW Yaygin and ca. 10  km due ESE Solhan (in Bingöl province), ca. 38.93N 41.16E, ca.  1600–1900 ma.s.l {9}. Thecollecting site is in the Muş province, but close to the border with the Bingöl province; the approximate specification on the holotypelabel ‘ 50 kmW Muş’means ‘on road’ ( C. Holzschuh, personal communication).   Type material. HOLOTYPE: J, ‘Asia minor 21.6.[19]72, 50 km W Muş, leg Holzschuh’ [hw Wewalka] {9}, ‘ Hydroporus jacobsoni Zaitz.?, det. G. Wewalka [19]72’ [hw Wewalka in part], ‘ Hydroporus n. sp.?’ [hw Wewalka], ‘ Holotype, Hydroporus holzschuhi sp. n., H. Fery det. 2009’ [red, printed] ( NMW). PARATYPES:  TURKEY:1 J 1 ♀, ‘Turkey, SE. Anat., N of Baskale [ca. 38.2N 44.0E], 2600 m, 21 8.[19]70’ [printed] {8}, ‘Loc. no. 104, Exp. Nat. Mus., Praha’ [printed]. Each paratypeis provided with the respective red printed label ( NMPC). In HOBERLANDT (1974: 20)is given ‘ 30 kmN of Baskale’. The village of Başkale is situated in the SE of the Van province, ca. 75 kmdue SE the province capital Van. It is not clear whether ‘ 30 kmdue N of Baskale’ is meant or ‘ 30 kmon the road’. Most probably, the specimens were collected not far from the road from Başkale to Gürpinar and Tatvan.   Differential diagnosis(including chiefly differences from  H. libanusbut also those from  H. jelineki sp. nov.). Habitus similar to that of  H. libanus; maximum width of body near middle of total length, between first and second third of elytral length. Body outline and shape of pronotum in dorsal view as in  H. libanus. Punctation on disc of elytra similar to that of  H. jelineki sp. nov., less coarse than in  H. libanus; distance between punctures about same as their diameter, puncture lines rather indistinct. Upper surface darker than that of  H. libanus, blackish in large parts, but not as dark as in  H. jelineki sp. nov.; head dark brownish, with vertex lighter, but blackish behind each eye. Pronotum almost black, rim at sides brownish, somewhat darker anteriad. Elytra on disc of same colour as pronotum, brownish stripe along suture distinct, dark colour of disc gradually becoming lighter to sides and lateral third of each elytron thus brownish except near shoulders; these brownish parts in posterior two thirds intersected by a darker narrow longitudinal stripe. Ventral surface predominantly black, colouration similar to that of  H. jelineki sp. nov.; femora somewhat darkened in middle. Antennae and palpi brownish, segments not darkened apically. Contrast between black surface and brownish appendages stronger than in  H. jelineki sp. nov. JJ. Shape of median lobe as in Fig. 4, intermediate between that of  H. libanus( Fig. 3) and  H. erzurumensis Erman & Fery, 2000( Fig. 2; drawing taken from ERMAN & FERY 2000); in ventral view broader than in former, but narrower than in latter; tip broadly rounded, almost as in  H. libanus. Parameres as in Fig. 11. Sucker cups on first tarsomere of pro- and mesotarsomeres present.  ♀♀. The single female studied without conspicuous external differences to males. Gonocoxosternum and gonocoxae as in Figs. 17 and 22.   Measurements.TL: ( holotype/ male paratype/ female paratype): 3.9 / 3.2 / 3.7 mm; MW: 1.9 / 1.6 / 1.9 mm; TL/MW: 1.95–2.05; IO/MP: 0.45–0.50.  Variability.The two paratypesfrom Başkale, in particular the male, are somewhat smaller than the holotypefrom Muş. The female is darker all in all, and the male paratypehas the appendages and the prosternal process lighter than the holotype.   Distribution.Endemic in the Muşand Vanprovinces in eastern Turkey( Fig. 26).   Biology: HOBERLANDT (1974: 20)stated that the typelocality was a ‘green grassy valley of a brook, partly swampy, with  Juncus; the bank of the brook narrow with stones.’ Nothing more is known about the biology of this new species.   Etymology.I name this species after the collector of the holotype, Carolus Holzschuh, wellknown specialist in Cerambycidae(Villach, Austria) (noun in apposition in the genitive case).    Hydroporus jacobsoni Zaitzev, 1927      Hydroporus jacobsoni Zaitzev, 1927: 17(original description). –  ZAITZEV (1933: 335;  1946: 88;  1953a: 93; 1953b: 174; 1972: 184). – GSCHWENDTNER (1939: 34). – IENIŞTEA (1978: 297, partim). – SCHAEFLEIN (1983: 16).    Hydroporus longicornisSharp, 1871:  FICHTNER (1974: 189)(misidentification).   Type locality(by present designation of lectotype):  Georgia, Gvileti, ca. 110 kmN Tbilisi, altitude ca.  1500 m( ZAITZEV 1927: 17); according to the label data of the lectotype SW Gvileti, on the way to the Devdorakglacier, NE peak of Mount Kazbek, ca. 42.70N 44.58E{19}.  Type material studied. LECTOTYPE(by present designation): J, ‘oz. u Devdoraksk. budki [= lake (or pond) near Devdorak hut (or ‘booth’ or ‘refuge hut’)], 16.VII.[19]26, Tarnogradsk.’ [in Cyrillic, hw Zaitzev] {19}, ‘ Lectotype, Hydroporus jacobsoni Zaitzev, 1927, des. H. Fery 2009’ [red, printed] ( ZISP). The last four tarsomeres of the right fore leg and the last two of the left hind leg of the lectotypeare disarticulated, but glued on the card. PARALECTOTYPES: 2 ♀♀, ‘Bakuriani, prov. Gori, 24.VI.[19]16’ [ Georgia, ca. 100 kmW Tbilisi, ca. 2600 ma.s.l., ca. 41.74N 43.53E] {17}, one with additional ‘ Syntypus, Hydroporus jacobsoni Zaits. 1927’ [red, hw R. E. Roughley? ( Manitoba, Canada)] ( ZISP). 5 exs., same first label, specimens strongly damaged by dermestids, sex not identifiable; each specimen glued onto its own card, these all mounted on the same pin ( ZISP). Each pin is provided with a respective red printed paralectotypelabel.  Notes on the lectotype.The Devdorak glacier is situated near the peak of Mount Kazbek (northern Georgia, near the Russian border, ca. 42.69N 44.51E). The village of Gvileti (or Gveleti; ca. 42.71N 44.62E) is situated ca. 7 kmNE of Mount Kazbek (a nearby village is Dar’yal’skoye). The hut mentioned on the label must be situated between Gvileti and Mount Kazbek. The following annotations can be found under the URL <http://prielbrusie.narod. ru/library/kazbek/index5.html> (access in January 2009; in Russian, translation by P. Petrov, Moscow, Russia): ‘Climbing Mount Kazbek in 1903–1913. The popularity of the Devdorak way those years is also explained by the fact that at the foot of the glacier there was a hut, in which tourists stopped on their way up to the peak.’ The collector cited on the label should be David Abramovich Tarnogradsky (1891–1974), director of the North-Caucasus Biological Station which published two of Zaitzev’s works, one on Dytiscidaeand one on Gyrinidae. Tarnogradsky was also given as the collector in the original description of  H. jacobsoni, which also cited the collecting date as ‘16.VIII.[19]26’ in contrast to the date on the lectotypelabel. However, I have no doubt that it was erroneously copied by Zaitzev from the label.  Notes on further paralectotypes(not examined): Besides Gvileti and Bakuriani in Georgia, the following further localities in Caucasus were provided in the original description by ZAITZEV (1927: 17):  • ‘near Adaj-choch (  1070 m)’: Adaj-choch[= Adaykhokh or Adaikhokh, {20}] mountains in southern Russia, Northern Ossetia; here is situated the well-known Mamison Pass (= Mamisonskiy Preval, ca. 42.70N 47.63E), ca. 75 kmWSW Vladikavkaz.  • ‘near Mount Bambak(ca.  2500 m), district Majkop’: Majkop[= Maykop, {23}], situated in the Adygeya Republicin southern Russia, NW of north-western Georgia;  Mount Bambak[= gora Bambak] is situated in the Krasnodar Kraynear the border to the Adygeya Republic, ca. 43.91N 40.43E, NNE of the city of Krasnaya Polyana. Except the paralectotypesfrom Bakuriani (see above), I have not been able to locate those from the other two localities. They are not deposited in the ZISP and previous inquiries at the Dzhanashia State Museum of Georgia, Tbilisi, where parts of the Zaitzev collection are stored, have not been successful. Should any former syntypesfrom these localities be found, they will have to be treated as paralectotypes(see Article 74.1.3 of the ICZN 1999).  Additional material studied. GEORGIA:1J, ‘Grusia [= Georgia], 1871’ [hw?], ‘Geor[g]ien’ [hw Wewalka] ( ZISP).  RUSSIA: 1 ♀, ‘Lager [or Camp] Bombang, Kubansk oblast, Go?rulya, 10.VIII.[1]910’; last word meaning most probably the collector, at least one letter illegible; text in Cyrillic; translation by P. Petrov). I have not been able to find ‘Camp Bombang’ on any map; the Kubansk district is situated in north-western Caucasus, Krasnodarskiy Kray. 1 J, ‘ Kabardino-Balkaria, s. [= selo = village] Verkhniye Balkary, 7.VIII.2006, leg. Nabozhenko,Terskov’ [Cyrillic; translation by P. Petrov]; [ca. 43.13N 43.45E; ca. 1100 ma.s.l.] {21} ( CMS). 1 J 1 ♀, ‘Zentr. Kaukasus [= Central Caucasus], 25.6.[19]74, Joost’ [blue ink,hw Joost?] ( CLH, ZSM); the female with additional labels ‘Zentral-Kaukasus, Joost leg., 25.6.1974USSR’ [hw Fichtner (?)], ‘ jacobsoni Zaitz., Fichtnerdet. [19]78’ [hw Fichtner in part]. Both latter specimens belong to a series recorded under the name  H. longicornisby FICHTNER (1974: 189), who specified the collecting locality as ‘near Itkol’. SCHAEFLEIN (1983: 16)studied a specimen from the same series and corrected the determination to  H. jacobsoni. The name ‘Itkol’ denotes a village (chiefly a complex of tourist hotels) and a left tributary of the Baksan river on the south-eastern slopes of Mount Elbrus (north-western Caucasus, Kabardino-Balkar Republic, ca. 43.25N 42.57E, SE of the two peaks of Mount Elbrus; altitude approximately 2000 ma.s.l.) {22}.   Differential diagnosis(based only on the lectotype, paralectotypesnot considered; chiefly differences from  H. jelineki sp. nov.given): Maximum width of body distinctly behind middle of total length, more or less in middle of elytral length. Discontinuity of body outline in dorsal view between pronotum and elytra more distinct, because pronotum with sides in posterior two thirds almost parallel, maximum width near posterior angles. Punctation on disc of elytra slightly less coarse and a little denser than in  H. jelineki sp. nov., but puncture lines more distinct. Head brownish, on vertex lighter brownish, between insertion of antennae and eyes and behind eyes blackish. Pronotum near anterior angles diffusely brownish, rim dark brownish, at posterior angles a little lighter. Elytra very dark brownish, almost black; each elytron in anterior two thirds with brownish stripe parallel to suture. Venter with contrast between brownish appendages and blackish surface more distinct than in  H. jelineki sp. nov.; darkening of middle of femora almost absent.  Figs. 5–8. Median lobe of aedeagus in ventral and lateral view. 5 –  Hydroporus jacobsoni Zaitzev, 1927; 6 –  H. hajeki sp. nov.; 7 –  H. shaverdoae sp. nov.; 8 –  H. kryshtali Bilyashiwski, 1993. JJ. Median lobe of aedeagus ( Fig. 5) with shape in ventral view as described by ZAITZEV (1927: 17; see below), almost parallel in basal two thirds, more or less evenly tapering to tip in apical third; tip very shortly rounded; in lateral view apical third distinctly curved, tip rather broad. Paramere as in Fig. 12. Sucker cups on first tarsomere of pro- and mesotarsomeres present.  ♀♀. Not known from the typelocality. Gonocoxosternum and gonocoxae of the female from Itkol as in Figs. 18 and 23.   Measurements. Lectotype: TL: 3.75 mm, MW: 1.85 mm, TL/MW: 2.03, IO/MP: 0.50. Other specimens: TL: 3.6–3.7 mm, MW: 1.7–1.85 mm, TL/MW: 1.95–2.06, IO/MP: 0.46–0.49.  Figs. 9–15. Left paramere. 9 –  Hydroporus jelineki sp. nov.10 –  H. libanus Régimbart, 1901; 11 –  H. holzschuhi sp. nov.; 12 –  H. jacobsoni Zaitzev, 1927; 13 –  H. hajeki sp. nov.; 14 –  H. shaverdoae sp. nov.; 15 –  H. kryshtali Bilyashiwski, 1993.  Variability.The specimens studied vary to a certain extent in colouration of the dorsal and ventral surface, in punctation and body outline (degree of pronoto-elytral discontinuity, position of maximum width of the body). The shape of the median lobe varies also a little in the four malesstudied, but at present I cannot separate them as distinct populations that could be clearly characterised and possibly described as new taxa (see also the remarks below).   Distribution.  Hydroporus jacobsoniis distributed in the Georgian and Russian Caucasus ( Fig. 26; for several localities see the next section). However, the true distribution of the species is by far not clear, because only females, which cannot be assigned to  H. jacobsoniwith certainty, are known from most localities; such localities are provided with a question mark in Fig. 26. Any future males from other Russian or Georgian localities should be studied to verify whether they belong to  H. jacobsonior to another, possibly still undescribed species. GUÉORGUIEV (1981: 407)recorded  H. jacobsonifrom several localities in Turkey. According to my studies these data are most probably incorrect and must be attributed either to  H. dobrogeanusor to other species, e.g. those described in FERY & ERMAN (2009).    Hydroporus jacobsoniin the literature.  Hydroporus jacobsonihas been rarely treated in the literature and the only essential contributions have been given by Zaitzev. The first seven lines of the original description ( ZAITZEV 1927: 17) are in Latin (see below for the following part in Russian). A shortened translation into German is given by GSCHWENDTNER (1939: 34–35)and a translation into English follows here: ‘J ♀. Similar to  H. longulusMuls.but certainly different: microsculpture of surface less regular, less even, elytra posteriorly more constricted, with punctures coarser and much more approximated, side margin (in lateral view) anteriorly less ascending (but more strongly than in  H. melanarius); pronotum near sides more coarsely (sometimes subrugosely) punctate, sides little curved, lateral beading thicker. Penis until 2/3 of length of rather equal width, then tapering [to tip]. Length 3.4–3.8 mm.’ Notes: In the English translation ( ZAITZEV 1972: 185) of ZAITZEV (1953b: 174)the shape of the median lobe in ventral view is described as ‘then abruptly tapering’ behind the basal two thirds. The adverb ‘abruptly’ seems to be at least somewhat misleading and the phrase should be replaced by ‘from here tapering [until tip]’. All other contributions known to me are in Russian. I give here the translations by P. Petrov of the relevant parts of these works:  ZAITZEV (1927: 17–18)(for the Latin part see above): ‘The new species belongs to the cycle  longulus- nevadensis- cantabricus, but since the latter two Pyrenean species are known to me only by description, so I am comparing our [= my] species only with the former [=  H. longulus], two specimensof which I have before my eyes. It is impossible to consider it the same as  longulusif it were only for the fact that the latter [=  H. longulus] as well as the two others mentioned above are characteristic inhabitants of the western Mediterranean region, one with larger range of distribution, both the others with narrower. However, in APFELBECK [1904: 377]we find a record of the species [=  H. longulus] for Herzegovinaand Attica, but we are still not completely convinced of the correctness of this record. Differences in the shape of the penis do not allow considering our species as a race of  longulus. Judging by the localities where it was found, this species lives in our land only in water bodies of higher mountainous areas. Leder [= SCHNEIDER & LEDER 1877: 86] reports  H. longulusfrom Suram [locality {18}]. No doubt this record refers to our species, the more so because Suram is situated near Bakuriani [locality {17}], where specimens used for this description were taken. Jacobson (Beetles of Russia, p. 426) was absolutely right doubting Leder’s data.’ Suram, better known under the name Surami, is a town in central Georgia(ca. 42.01N 43.25E) NW of the town of Khashuri. A mountain range in the west of Khashuri was formerly called Suramskiy Khrebet and today is named as the Likhskiy Khrebet ( SCHÜTZE & KLEINFELD 2001: 131). In LEDER (1880: 454)it is called ‘Suram-Gebirge’ or ‘Meskisches Gebirge’ [Gebirge = mountains]. This mountain range divides Georgiainto the eastern and western part and connects the Greater Caucasus and the Lesser Caucasus ranges. The distance between Khashuri and Bakuriani is about 30 km.  ZAITZEV (1933: 335): ‘Lake Tabis-kuri, Bakuriani [locality {17}], Suram [locality {18}], then on the main Caucasus range (spring near Mount Adai-khokh [locality {20}], 1070 m, near Vladikavkaz), Gvilety [locality {19}], ca. 1500 m, near Mount Bambak [locality {23}] in environs of Maikop, 2050 m. This species may be considered close to  longulusMuls.and  cantabricusSharp, which both live in mountain water bodies of the Pyrenees, the Alps, and the Balkan Peninsula. There are no closer relatives either of these two species or of  jacobsoniin the north, at present. Probably some species during the postglacial time, when rising higher into the mountains, produced all the three species mentioned above, and maybe also  asturSharp.’ [  Hydroporus astur Sharp, 1882is a junior subjective synonym of  Hygrotus marklini(Gyllenhal, 1813); see BALKE & FERY 1993.]  ZAITZEV (1946: 88): ‘It was found also in the eastern part of the main range (Lagodekhsk nature reserve, river Antsal’-or [ca. 41.76N 46.22E, ca. 120 kmdue E Tbilisi, locality {16}], 7 VIII [19]37 Kakauridze!)’ [the exclamation mark most probably means ‘the collector’].  ZAITZEV (1953a: 93): ‘Considerable material is present from Bakuriani [locality {17}], Gvileti [locality {19}], Lagodekhi (river Antsal’-or) [locality {16}]. In mountain water bodies with flowing water.’ 1750 38.93 The 703 41.16 11 540 1 Bingol C. Holzschuh Mus' 11 540 1 Bingol holotype Georgia 1500 Gvileti 12 541 1 Tbilisi SW Gvileti & Devdorak Georgia 42.7 Mount Kazbek 689 44.58 12 541 1 Tbilisi lectotype Russia 1070 Northern Adaj-choch 13 542 1 North Ossetia Russia 42.7 Vladikavkaz. 689 47.63 13 542 1 North Ossetia Russia Majkop 2500 Mount Bambak Majkop' 13 542 1 Adygeya Russia 43.91 Mount Bambak 684 40.43 Krasnodar Kray 13 542 1 Adygeya Republic