Hydroporus glasunovi dolini, Fery & Stastny, 2010

Fery, H. & ŠġASTNÝ, J., 2010, On the identity of Hydroporus glasunovi ZAITZEV 1905, with the description of Hydroporus glasunovi dolini nov. ssp. (Coleoptera, Dytiscidae), Linzer biologische Beiträge 42 (2), pp. 1005-1028 : 1012-1028

Hydroporus glasunovi dolini nov.ssp.

? Hydroporus glazunovi [sic!] ZAITZEV; ZAITZEV 1953: 171 (partim). - ZAITZEV 1972: 181

(partim).

H o l o t y p e (): " Kyrgyzstan: Talassky Ala-Tau, Tschatkal [= Chatkal], ca. 3000 m, 30.V.1997, leg. Dolin (K-5/97)", "upper course of Chatkal river" [printed], " Holotypus, Hydroporus glasunovi dolini ssp. nov., Fery & ŠĢastný det. 2010" [red, printed], ca. 42.10N 71.70E {loc. 7} (NMW).

P a r a t y p e s: K y r g y z s t a n: 5, 4 exs. " Kyrgyzstan, Tshatkal Rg. Mt. Tshapshama pass, 2800, 3.06.[19]98, V. Dolin, R. Andreeva ", ca. 41.53N 70.80E {loc. 2} ( CJS, NMPC). 1, " Kyrgyzstan, Talass Rg., Bord of Kasakhstan , env. Majdantal-pass [= Maydantalskiy pereval; see SCHÜTZE & KLEINFELD 1999], 2640 m, 28.05.1997, V. Dolin leg.", ca. 42.30N 70.88E {loc. 3} ( NMB). N o t e s: In the following records the locality "Stab", called also "Shtab", is given. We have not been able to find that locality on any map or elsewhere, but A. Putchkov communicated: "All these [specimens] were collected in the environs of the town Kanysh-Kiya [ca. 41.75N 71.03E], within 5 km from this town. Mostly in the valley of the Chatkal River , upwards along the current. The valley in the environs of Kanysh-Kiya is rather broad, and there are opportunities to climb and to drive around." And R.V. Andreeva (Dolin's widow) added: "Shtab [headquarters] is a ruined concrete construction in the middle reaches of the Chatkal River between its tributaries Kedey-Say and Chanach." 1, " Kyrgyzstan, Mt. Rg. , Tshatkal, 2000, Shtab, 28\2906.[19]98", ca. 41.75N 71.03E {loc. 4} ( CJS). 3, " Kyrgyzstan: W-Tien Shan , Chatkal gorge, Stab. env., 29.VI.1998, 1800 m, leg. Dolin (K-9/98)" {loc. 4} ( NMW). 1, 2, " Kyrgyzstan, Tshatkal valley , Shtab , 2000 m, 28.07.[19]99, V. Dolin leg." {loc. 4} ( NMPC). 25, 5, " 28.7.1999 Kyrgyzstan, Tschatkal mountain range , nr Stab , Bach und Pfützen, Dolin leg." {loc. 4} ( NMW) .

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3, 3, 9 exs., " Kyrgyzstan: Sandalasch G.K., Chatkal riv., ca. 2000 m, 5.VII.1997, leg. Dolin (K-9/97)" {loc. 4} (NMW). 2, 2, " Kyrgyzstan, Talassky Rg., Kara-Bura pass, 3200- 2700 m, 29- 30.05.1997, W. Dolin leg.", 42.20N 71.58E {loc. 6} (NMB). 2, 7, same data as the holotype {loc. 7} (NMW). 1, "Khirgizia: Kichik Alai, Kara-Goy, 2400 m, 21.-22.V.1993, leg. Schawaller", ca. 40.20N 72.45E {loc. 9} (SMNS). 3, 4, "Khirgizia: Kichik Alai, above Kara-Goy, 3000-3200 m, 23.V.1993, leg. Schawaller", ca. 40.20N 72.45E {loc. 9} (SMNS, CHF). 3, 5, " 19.07.1996 Kyrgyzstan, Alaj-Bergkette, Umg. Taldyk-Pass [NW Sary-Tash], 2800 m, Dolin leg. (N 42a)", ca. 39.75N 73.16E {loc. 10} (NMW). 5, 5, " Kyrgyzstan: Ferganskyj-G.K., E-slopes, Urumbasch val., 2000 m, 14.VI.1997, leg. Dolin (K-7/97)", ca. 41.20N 73.40E {loc. 11} (NMW). 1, " Kyrgyzstan, Ferganski range, eastern slopes, nr. Bergut pass, 2015 m, 24.6.1996, leg. Dolin (n 37)", roughly estimated to 41.25N 73.7E {loc. 12} (NMW). N o t e s: This rather immature specimen was incorrectly given under H. goldschmidti in FERY & PETROV (2006: 255). We have not been able to find the exact co-ordinates of the "Bergut pass" on any map, however, A. Putchkov communicated that it is situated on the road from Kazarman to Karaalma. 1, 1, " 13.07.1996 Kyrgyzstan, Suusamyr Plateau, 2250 m, Dolin leg. (N 40)", ca. 42.28N 73.83E "(possibly situated 30 km more SW, at ca. 42.10N 73.57E) {loc. 13} (NMW). 23, 31, " 5.8.1999 Kyrgyzstan, Suusamyr Plateau, nr Tjuz-Aschu pass, 2300 m, Dolin leg." {loc. 13} (NMW). 1, 1, "Kirghizstan Kirghizsky G.-K., Karabalty-Schlucht [= gorge], 2000 m 1- 6.06.[19]94", ca. 42.67N 73.88E {loc. 14} (CJS). 1, 2, "1.- 2.06.1996 Kyrgyzstan, Karabalta- Schlucht, Sosnowka Umg., 1520 m, Dolin leg. (N 23)" {loc. 14} (NMW). 1, " Kyrgyzstan: Sosnowka env., Karabalta gorge, 1520 m, 1.- 2.6.1996, Dolin leg. (n 23)" {loc. 14} (NMW). 1, " Kyrgyzstan: 14.6.1995 Fergansky G.K., 2000 m, Kasarman env. leg. Dolin (23)", ca. 41.41N 74.04E {loc. 15} (CJS). 2, 1, "N Kirghizia Kirghizky Mt., Range Ala-Archa, 1600 m, 29.10.2000, leg.?, Coll. L. Hendrich", ca. 42.6N 74.5E {loc. 16} (CLH). 1, 2, "Kirghizstan, Moldo-Too, 2300, Kurtka valley, 10.07.[19] 95 W. Dolin", roughly estimated to 41.7N 74.7E {loc. 17} (NMB). 11, 8, " Kyrgyzstan, Inner Tien-Shan, Arpa, 2800 m, 14.-18.VII.2000, unknown collector, Coll. Hendrich", ca. 40.78N 74.78E {loc. 18} (CLH). 2, " 27.07.1996 Kyrgyzstan, Moldotoo Bergkette, nördlich vom Naryn Fluss", "ca. 100 km westlich Naryn, unter Karago-Bergpass, 1750 m, Dolin leg. (N 46)", ca. 41.75N 74.83E {loc. 19} (NMW). U z b e k i s t a n: 1, " Uzbekistan, Tashkent Province, Yakka-tut near Burchmulla, 2.7.- 4.7.2005, 41°38'N 70°03'E [= 41.63N 70.05N], Hendrich leg. Loc. 1a/b" {loc. 1} (CLH). N o t e s: All paratypes with the respective red printed paralectotype label.

T y p e l o c a l i t y: Kyrgyzstan: Talassky Ala-Tau, upper reaches of Chatkal River, altitude ca. 3000 m, ca. 42.10N 71.70E {loc. 7}.

M a t e r i a l n o t c o n s i d e r e d p a r a t y p e s The following specimens from K y r g y z s t a n are not considered paratypes because they cannot be assigned undoubtedly to one of the two subspecies. Their shape of the body and that of the median lobe as well as the reticulation of the venter is as in H. g. dolini nov.ssp., the measurements also (see e.g. column "Maydan" in tab. 1) give values clearly closer to the new subspecies than to the nominotypical one. However, the lighter colouration of the upper side and that of the antennae and legs make these specimens similar to H. g. glasunovi (see also section Populations of unclear identity): 4, " Kyrgyzstan: Alaj-G.K., Majdan [= Maydan] gorge, ca. 2000 m, 12.VII.1997, leg. Dolin (K- 11/97)", ca. 39.70N 72.15E' {loc. 8} (NMW). 4, 5, 1 ex. without genitalia, " Kyrgyzstan: Alaj-G.K., Majdan (right) gorge, 2270 m, 13.VII.1997, leg. Dolin (K-12/97)", ca. 39.70N 72.15E' {loc. 8} (NMW). 7, 12, " Kirgisistan, Sumsar, 1600 m, 15.6.1993, M. Danilevsky leg. ", ca. 41.28N 71.32E {loc. 5} (CLH, CHF). The following specimen is extremely immature and, thus, safe identification is impossible. The few better recognisable characters, however, allow us to assume that the specimen belongs to the new subspecies: 1, "Turkestan, Utsch Tjube [= Uch- Tyube], Alai Geb. [= Alai mountains], 2370 m. 10.10.[18]90, Conradt S." [light brown labels, printed], " Hydroporus glazunovi [sic!] Zaitz, det. G. Wewalka [19]88", ca. 39.90N 73.44E, ca. 40 km NE Sary-Tash (somewhat NE of loc. 10) (MNB).

The following specimens from K a z a k h s t a n (localities 20, 21 and 22) are not considered paratypes because some of them look indeed like typical H. g. dolini nov.ssp., but others show strong tendencies to the nominotypical subspecies. Considering in addition the distance of about 500 km between these localities and the closest ones of H. g. dolini nov.ssp., we prefer to treat these specimens not as paratypes of the new subspecies. 1, 1, "Distr. Panfilov [= Sharkent], Umg. Koktal, 26.VI.1989 ", "USSR, O-Kasachtan (sic!), V. Dolin", the male with additional " Hydroporus glasunovi Zaitz , det. A. Nilsson", ca. 44.13N 79.80E {loc. 20} (NMB). 1, "USSR, Kasakhstan, Dzhungar A. [= Dzungarskiy Alatau] Tyshkan, 11.V.1985, leg. Shilenkov",

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roughly estimated to 44.5N 80.0E {more or less same as loc. 21, see below} (HNHM). 1, "SE Kazakhstan, S Dzhungarskìy Alatau, Tyschkan Valley N Sarybel, 1700-1800 m, 5.-7.VII.2001, leg. W. Schawaller", ca. 44.51N 80.09E {loc. 21} (SMNS). 2, "Tyschkan Tau, 10 km E Sarybeh [= Sarybel, ca. 44.25N 80.02E]", " 2250-2600 m, 12.VII.1990 ", "USSR, O Kasachstan, V. Dolin", ca. 44.39N 80.16E {loc. 22} (NMB). Another specimen has very inexact locality data: 1, " 3.06.1996 Süd-Kasachstan [= southern Kazakhstan], Talass Bergkette [= mountain range], Vorgebirge [= foothills], 900 m, Dolin leg. (N24)" (NMW). N o t e s: The Talas mountain range extends south of the city Talas and the river of the same name and (in its eastern part) north of the river Naryn from ca. 42.33- 42.50N 70.50- 73.00E (SCHÜTZE & KLEINFELDT 1999: 169), with its largest parts in Kyrgyzstan, a small western part in southern Kazakhstan and eastern Uzbekistan. Dolin collected on June 1-2, 1996, in the Karabalta gorge {loc. 13}, possibly he subsequently left that region and already one day later collected in the Kazakh part of the Talas mountains, somewhat west of {loc. 3}.

D i f f e r e n t i a l d i a g n o s i s H a b i t u s in dorsal view elongate oval ( fig. 2 ), but sides somewhat more rounded than in H. g. glasunovi, not subparallel; discontinuity between pronotum and elytra more distinct, because sides of elytra behind shoulders more curved. Maximum width near middle of total length, between first and second third of elytral length; sides more rounded and less pointed at apex. In lateral view outline of dorsum more or less evenly curved from anterior margin of pronotum to apex of elytra; body distinctly more vaulted and in posterior third more descending towards apex than in nominotypical subspecies. Pronotum black, elytra lighter, of dirty dark brown; contrast between pronotum and elytra present, but not conspicuous.

H e a d darker than in H. g. glasunovi, only indistinctly lighter than pronotum. Interocular distance smaller than half of pronotal width at posterior angles. Polygonal meshes on surface somewhat smaller. Punctures on clypeus near anterior margin sparser; behind of about same diameter as meshes, on frons denser and coarser.

P r o n o t u m almostentirelyblack,onlylateralbeadinposteriorhalfbrownish;anterior and posterior margins narrowly transparent brownish, but very indistinctly so. Lateral bead near anterior angle somewhat broader. Sides of pronotum more converging from posterior towards anterior angles. Near lateral margins punctures slightly longitudinally impressed, posterolaterally still deeper and coarser. Surface smooth only near central puncture and in small transversal area between this puncture and posterior margin.

E l y t r a dirty dark brown, not reddish; only very indistinctly transparent in anterior third. In dorsal view sides in anterior two thirds evenly rounded, in posterior third more rounded, apically less pointed, conjointly rounded. In lateral view elytral margin less ascending towards humeral angle than in nominotypical subspecies. Discal puncture line indistinct near elytral base, behind base formed by coarse punctures, in posterior third these becoming progressively weaker; other puncture lines less recognisable because difference in diameter of normal surface punctures and those of lines smaller. Elytral reticulation at apex less impressed than in H. g. glasunovi.

V e n t r a l surface with colouration similar to that of nominotypical subspecies, but brownish parts darker. Prosternal process in cross-section almost tectiform. Large parts of ventral surface without reticulation. Genae reticulated, gula smooth except in anterior third. Metacoxal plates with traces of irregular transverse lines (or strongly reduced transverse meshes). Last three abdominal ventrites with transverse meshes, weakly impressed on fourth and fifth, strongly impressed on apical half of sixth ventrite. Last abdominal ventrite, with sculpture similar to that of H. g. glasunovi, but surface here more roughly sculptured. Antennae, palpi, mouthparts and legs lighter brownish, but not as light as in nominotypical subspecies and contrast to black venter less prominent. Fifth to

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eleventh antennomeres and last palpomere darkened distally. Trochanters and mesocoxae darker brownish, procoxae in distal third dark brownish, elsewhere almost black. Femora slightly darkened centrally. Fifth to tenth antennomeres not much longer than 1.5 of their greatest width.

: Protarsal claws slightly prolonged, medially more or less straight, before apex more curved; differences to female claws more distinct than in H. g. glasunovi. Median lobe of aedeagus in dorsal and lateral view as in fig. 4 ; left paramere as in fig. 8 .

: Externally similar to males, except simple protarsal claws. Gonocoxae as in fig. 5 , gonocoxosternum as in fig. 6.

M e a s u r e m e n t s: The holotype has the following values: TL = 3.6 mm, TL-H = 3.25 mm, MW = 1.75 mm, TL/MW = 2.06, TL-H/MW = 1.86, WPr/IOD = 2.23, MW/WPr = 1.21. Further data can be found in tab. 1.

D e r i v a t i o n o m i n i s The new subspecies is named in honour of the collector of the holotype and most parts of the other material, Vladimir Gdalich Dolin from Kiev, Ukraine, specialist in Elateridae , who left us far too early in 2004 (obituary by CATE 2005).

N o t e s o n r e l a t e d s p e c i e s Hydroporus g. dolini nov.ssp. is somewhat similar to Hydroporus transgrediens GSCHWENDTNER 1923 from south-eastern Europe, Asia Minor and the Middle East. This species, however, can be easily distinguished from the new subspecies by the rather large non-reticulated area before the base of the pronotum and by the shape of the male genitalia (see FERY & PETROV 2006). Hydroporus goldschmidti is another similar species, which – beside other differentiating characters – is on the average larger (3.5-4.2 mm) and has different shape of the male median lobe (see FERY & PETROV 2006). In the latter publication the range of the TL of H. goldschmidti is incorrectly given as " 3.3-4.2 mm ", the lower value being due to a single female from "Bergut pass" cited there with the remark "specimen with pronotum totally reticulated, but elytra smooth". This specimen was now proved to belong to H. g. dolini nov.ssp. (see above).

D i s t r i b u t i o n: Kazakhstan, Kyrgyzstan, Uzbekistan (see fig. 9).

Notes on variability

If a "normal" H. g. glasunovi and a "normal" H. g. dolini nov. ssp. are directly compared, one would by no means have the idea that both are closely related. The body outline of H. g. dolini nov.ssp. is more rounded in dorsal view, more vaulted in lateral and frontal view; the general colouration of H. g. glasunovi is lighter with a distinct tendency to reddish, while it is darker and more dirty brownish in H. g. dolini nov.ssp.; the contrast between the lighter elytra and head on the one hand and the darker pronotum on the other hand is more prominent in the nominotypical subspecies. Some other characters which can be helpful are the following:

x colouration of sides of pronotum, prothoracic epipleuron and proepisternum: brownish in H. g. glasunovi, dark brownish to black in H. g. dolini nov.ssp. (except parts of the bead in most specimens);

x colouration of legs: reddish brown in H. g. glasunovi, brownish and darkened in part in H. g. dolini nov.ssp.;

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x colouration of antennomeres: unicolourous and lighter reddish brown in H. g. glasunovi, darker brownish and darkened distally in H. g. dolini nov.ssp.;

x length of fifth to tenth antennomeres: about 2 times as long as wide in H. g. glasunovi, about 1.5 times as long as wide in H. g. dolini nov.ssp.;

x extent of smooth area on pronotum: rather large in H. g. glasunovi, smaller in H. g. dolini nov.ssp.;

x punctation on last abdominal ventrite: denser and coarser and apex roughly sculptured in H. g. dolini nov.ssp., less so in the nominotypical subspecies;

x shape of median lobe: in lateral view more curved near base in the nominotypical subspecies, wider curved in H. g. dolini nov.ssp. (compare figs 3 and 4 ).

However, all these characters (and also others not listed here) are varying considerably in part and any particular specimen can by no means be identified on the basis of only one of them. For instance, we have studied several specimens of H. g. dolini nov. ssp. with unicolourous antennae (possibly due to immaturity) and several specimens of H. g. glasunovi with distally slightly darkened antennomeres; some H. g. glasunovi have the last abdominal ventrite more strongly punctured and sculptured than several H. g. dolini nov.ssp. According to our experience, however, identification should be relatively safe if any particular specimen displays the majority of the characters of one subspecies.

Populations of unclear identity

Originally, we wanted to describe the new taxon as a species and not as a subspecies of H. glasunovi . This treatment would have been supported by the fact that – to our present knowledge – H. g. glasunovi is restricted to the south-west and H. g. dolini nov.ssp. to the north and north-east of the discussed area.

The relatively high variability of several characters of both subspecies, however, and the possible overlap between both areas of distribution south of the Fergana plain, where specimens have been found which might be interpreted as representing an intermediate form, induced us to treat both populations as subspecies (localities J and 8; situated in zoogeographical zone I/3, but not far from the south-eastern part of zone I/5). We have no information on the mechanisms which once might have led to an isolation of part of the originally single uniform population, and which later allowed an eventual contact between both. Anyway, the chance to study more material from that transition region in the south-east of the Fergana plain would be very welcome. The specimens from locality 5 also show intermediate characters, but are especially problematic, because this locality is situated near the border between zoogeographical zones I/4 and I/5 (i.e. north of the Fergana plain), and not in zone I/3. At present we have no explanation for this observation except to assume a mix-up of the label data with those from another locality.

In addition to the unclear identity of the populations from localities J, 8 and 5, we have studied further material for which we have found no satisfactory solution. On the one hand, these are the specimens from Kangxiwar {loc. K}, China, which so far have been treated as belonging to H. g. glasunovi. On the other hand, all the specimens from the Issyk-Kul region {locs X and Y} cannot be assigned to one of the known species or subspecies. Below we list the respective material and give some comments.

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S p e c i m e n s f r o m C h i n a: The paralectotypes of H. glasunovi from China listed above under H. g. glasunovi must be included here. The fact that they are paralectotypes of this taxon does not affect in any way the following considerations.

1, 1, "4.X.[18]90, Gromb. [= Grombtschewski, collector], King Shiver [= Kangxiwar; almost illegible]" [hw Grombtschewski?], " Syntypus?, Hydroporus glazunovi Z." [red, hw R.E. Roughley?], ca. 36.20N 78.77E {loc. K} ( ZISP). N o t e s: Both specimens mounted on one pin, but separated by us and provided with photocopies of the original labels. These two specimens, most probably, belong also to the series of specimens which Grombtschewski collected in 1890 at King Shiver (Kangxiwar). We assume, however, that they have not been studied by Zaitzev when he described his new species, because (1) they have not been stored in the ZISP together with those listed above under Paralectotypes, and because (2) according to the original description ( ZAITZEV 1905: 26) there should exist only three syntypes from King Shiver. 4, 2, "Chin. Turkestan, King Schewer [= Kangxiwar], Kuschgar [= Kashgar], 2000 m. 3.-4.10.[18]90, Conradt S." [light brown labels, printed], four specimens with additional " Hydroporus glazunovi [sic!] Zaitz, det. G. Wewalka [19]88", ca. 36.20N 78.77E {loc. K} ( MNB). N o t e s: Kashgar (or Kashi ) is the administrative centre of the Kashgar prefecture in the western part of the Xinjiang Uyghur Autonomous Region of China. The name on the label has most probably the meaning of "Kashgaria" which is the ancient name of the western part of Xinjiang (SCHÜTZE & KLEINFELD 1997) .

These specimens are on a first glance similar to H. g. glasunovi, but are still lighter, in particular, the pronotal disc is not blackish, but reddish brown to dark reddish brown; the contrast between the light elytra and the darker pronotum is not very prominent; the shape of the body is even more elongate parallel (compare the values for TL/MW, TL- H/MW, MW/WPr in tab. 1); the venter is not reticulated except the last two abdominal ventrites. On the other hand, it is not quite clear whether the lighter appearance of the specimens is not due to immaturity or any aging process. Considering these observations and the large distance (ca. 700 km) between Kangxiwar and the known distribution area of H. g. glasunovi, it seems not to be unlikely that in future the study of freshly collected material – eventually supported by molecular methods – may lead to the result that the Chinese population is a distinct species.

S p e c i m e n s f r o m K y r g y z s t a n I s s y k - K u l r e g i o n The lectotype of H. macrocephalus (locality X) listed above under H. g. glasunovi must be considered here. We can provide the following details about its external characters: The specimen is somewhat immature, which may be the reason for the rather light appearance of the specimen and the rather weak contrast between the colour of the pronotum and elytra. The head and pronotum are light reddish brown, the pronotum only slightly darker. The elytra are a little lighter than the head and largely transparent. The head appears rather large, but this may be due to the fact that it is slightly protruding. The head and pronotum – except for a very small area near the larger discal puncture – are completely microreticulated. The punctation of the pronotum and elytra is conspicuously weaker than in both subspecies of H. glasunovi , the puncture lines are almost absent. The reticulation of the last two abdominal ventrites is distinct, on the other ventrites and on

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the metacoxal plates as well as on the metasternum, however, only very minute traces of reticulation are perceptible, and this only at high magnification (x160). The punctation of the last abdominal ventrite is distinct, but not coarse and roughly sculptured. The median lobe and the parameres are well sclerotised, and show no distinct differences to that of both subspecies of H. glasunovi . The measurements are given in tab. 1. When comparing the values with those of both subspecies of H. glasunovi , it can be seen that these are closer to those of the nominotypical subspecies.

We have strong doubts that H. macrocephalus is a synonym of H. glasunovi , the external characters make us suspect that it probably should be treated as a distinct species. Lack of more material, however, induces us at present not to change the status of this taxon. See also the notes about the type locality of the taxon under H. g. glasunovi.

Unfortunately, the following two specimens from the same region are not really helpful for solving the problem with H. macrocephalus: 1, "Przhevalsk, Semirech. [= Semirechye (Russian) = "seven rivers land" = "Siebenstromland" (in German)], Pedashen[k]o, 7.IV.[19]08" [translated/transliterated from Russian; the collector being probably Dmitry Pedashenko 1868-1927, zoologist from St. Petersburg] {loc. Y} ( ZISP). N o t e s: "Przhevalsk" has certainly the meaning of "environs Przhevalsk". 1, "Tien-schan., Przewalsk., Karakolthal [= valley of Karakol]", " Hydroporus , glazunovi [sic!] Zaitz., det. A. Nilsson [19]87" [hw Nilsson], "compared with types, 1/4-[19]89 AN" [red, hw Nilsson] {loc. Y} ( CAN). N o t e s: The river Karakol is formed by the confluence of the rivers Kel'-Ter and Uyun-Ter at 42.30N 78.48E, environs of the town of Karakol [formerly named Przhevalsk], and flows into Lake Issyk-Kul at 42.58N 78.28E (H. Schütze, personal communication). The specimen labelled " Przhevalsk, Semirech." is immature, the punctation of the surface is distinctly coarser than in the lectotype of H. macrocephalus, the elytral puncture lines are well visible. A small smooth area between the centre of the pronotal disc and its base is present. The specimen labelled "Karakol valley" is mature, the punctation of the surface is fine, but the puncture lines are distinct. The contrast between the lighter head and elytra and the dark brown pronotum is rather distinct. A small smooth area between the centre of the pronotal disc and its base is also present. The pronotum is rather broad, the base distinctly broader than that of the elytra. The last seven antennomeres are distally darkened. The protarsal claws are unusually short. This specimen is more similar to the "normal" H. g. glasunovi than the one labelled "Przhevalsk, Semirech.", which matches rather H. g. dolini nov.ssp. The colouration of both specimens is different from that of the lectotype of H. macrocephalus. On the other hand, Lake Issyk-Kul lies more or less exactly between the more south-western and the (slightly uncertain) north-eastern limits of the distribution of H. g. dolini nov. ssp. and is far away from those of H. g. glasunovi, a fact that creates additional problems, for which at present we are unable to offer a solution .

As can be seen, neither the lectotype of H. macrocephalus nor the two other specimens from the Issyk-Kul region {loc. Y} can be assigned with certainty to one of both subspecies of H. glasunovi and, additionally, all the three Issyk-Kul specimens themselves show not so many corresponding characters to be sure that they all belong to one and the same taxon. Thus, for the time being, we prefer to leave the status of H. macrocephalus as a junior subjective synonym of H. glasunovi unchanged and to treat the other two specimens as "close to H. glasunovi ".

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Notes on biology

Aquatic habitats in the Fan Mountains area are characterised by very cold water, even in summer months (Kulikalon lakes, max. 10° C), and by fluctuations of the water level due to uneven melting of the glaciers. Suitable conditions for development of water insects are found up to about 3,200 m a.s.l. (based on observations of immature stages). Lakes at altitudes between 2,600 and 3,200 m a.s.l. are inhabited by five species of predaceous diving beetles ( Dytiscidae ): Hydroporus g. glasunovi, Nebrioporus airumlus (KOLENATI 1845) , Agabus winkleri ( GSCHWENDTNER 1923) , Agabus dichrous SHARP 1878 , and Hydronebrius cordaticollis (REITTER 1896) . The junior author collected H. g. glasunovi only at altitudes between 2,700 and 2,900 m a.s.l. {loc. D} ( figs 10-12 ). It was found in abundance in sparsely vegetated littoral zones of lakes and places with stony bottom, where the beetles look for patches of accumulated detritus. Here the subspecies was found in association with N. airumlus and A. winkleri .

In Yakka-tut, near Burchmulla, Uzbekistan {loc. 1} ( fig. 13 ), H. g. dolini nov. ssp. was found in a slowly flowing irrigation channel with sandy and muddy bottom, with green algae and Chara sp. Other Hydradephaga found in the same locality are Agabus basalis (GEBLER 1829) , Agabus bipustulatus (LINNAEUS 1767) , Deronectes abnormicollis SEMENOW 1900 , Deronectes vestitus (GEBLER 1848) , Hydroglyphus geminus (FABRICIUS 1792) , Hydroporus goldschmidti , N. airumlus, Gyrinus distinctus AUBÉ 1838 and Haliplus sibiricus MOTSCHULSKY 1860 (see HENDRICH & HENDRICH 2005: 427).

Further localities where H. g. dolini nov. ssp. was found together with other species are the following: Stab {loc. 4}: Agabus conspersus (MARSHAM 1802) , A. bipustulatus , Agabus turcmenus GUIGNOT 1957 , H. goldschmidti (recorded in FERY & PETROV 2006: 255 under the name H. glasunovi ) and N. airumlus ; Talassky Ala-Tau {loc. 7} and Moldoo-To {loc. 19}: A. basalis ; Maydan gorge {loc. 8}: A. conspersus and Agabus sogdianus (JAKOVLEV 1897); Taldyk-Pass {loc. 10}: Hydroglyphus geminus ; Urumbash valley {loc. 11}: A. conspersus and A. winkleri ; Bergut pass {loc. 12}: A. winkleri and H. goldschmidti ; Karabalty gorge {loc. 14}: A. basalis and N. airumlus ; Suusamyr Plateau {loc. 13}: A. dichrous , Hydroporus nigellus MANNERHEIM 1853 , and Hygrotus impressopunctatus (SCHALLER 1783) .

Notes on zoogeography

According to KRYZHANOVSKIJ (1953), the mountain areas of the Central Asian parts of the former USSR can be divided into three separate zoogeographic zones (zones I-III in fig. 9), which are distinguished by different climatic conditions and consequent differences in the structure of the flora and fauna. Each of the areas is inhabited by endemic and apparently unrelated taxa. For our map (fig. 9) we adopted additionally the concept of physical-geographic zones of SOKOLOV & SYROETCHKOVSKIJ (1990) (called "subzones" below, to distinguish them from the larger "zones" of Kryzhanovskij) because of their well defined delimitations according to the botanical and geographical conditions, relief, geological structure, composition of the vegetation and ecological load. Thus, we have the following zones with their sub-zones included:

I. Middle Asian mountains zone (I/1: Southern Tajikistan; I/2: Western Pamir; I/3: Southern Tian-Shan; I/4: Fergana basin; I/5: Western Tian-Shan; I/6: Northern Tian-Shan; I/7: Dzhungaria).

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II. Central Asian mountains zone (II/1: Eastern Pamir; II/2: Inner Tian-Shan).

III. Turanic zone.

The Dzungar-Tian-Shan zone in the concept of SOKOLOV & SYROETCHKOVSKIJ (1990) includes the entire central and eastern Tian-Shan Mountains (corresponding with subzones I/6, I/7, and II/2). It is delimited by the Kyrgyz range in the north, by the Fergana Range in the south-west and by the Barkul and Kurug-Tag ranges and the Dzungarian Alatau Mountains in the east. The environmental conditions in this area are very heterogeneous and influenced mainly by the orientation of the mountain ranges. The climate is continental with marked differences between day and night temperatures. The rainfall increases from the south-west to the north-east; annual precipitation varies between 800 and 930 mm (Suru-Tchelek Nature Reserve). The air humidity in summer is about 45% (Naryn Nature Reserve). The average annual temperature ranges from 2.5 o C (Naryn Nature Reserve) to 6.6 o C (Issyk-Kul Nature Reserve). The vegetation is dominated by steppe associations; trees and shrubs occur only on the moister northern and western slopes. Mountain forests are composed of fir, rowan, birch, currant, willow and berberis (SOKOLOV & SYROETCHKOVSKIJ 1990).

The Western Tian Shan (sub-zone I/5 of SOKOLOV & SYROETCHKOVSKIJ 1990) includes the western slopes of the Fergana Range, the Talas Mountains, the Chatkal, Iskem and Urghan ranges, the Kardzhan Tag and Kara Tag Mountains and their foothills. The climate is typically continental with unevenly distributed rain- and snowfall. The summer rainfall represents only 10-15% of annual precipitation. Winters are very cold, but the snow cover provides good conditions for hibernating organisms. The annual precipitation is about 500-800 mm. The air humidity is about 52% in summer (Besh-Aral Nature Reserve). The flora is very rich and heterogeneous. Large areas are covered by mountain steppes and forests with dominant walnut trees, firs, maples, poplars and hawthorns (SOKOLOV & SYROETCHKOVSKIJ 1990).

The Pamiro-Alai zone in the concept of SOKOLOV & SYROETCHKOVSKIJ (1990) contains the mountains of the Pamiro-Alai, which form prominent mountain ranges in east-west direction (sub-zone I/3: Turkestan, Zeravshan and Gissar ranges), the Nuratau Mountains and the Pamir Mountains (sub-zones I/2 and II/1). Arid climate with marked variations in daily as well as annual temperatures is typical for this area; the variation is higher than in the preceding sub-zones. The annual precipitation is only 250 mm (Fan Mountains). The annual average of air humidity is 30-40% (Gissar Mountains). The vegetation is mainly xerophytic; typical trees in the mountain areas are the ‘archa’ junipers. In the Fan Mountains area (western part of the Pamiro-Alai Mountains), the highest species diversity of plants is found at altitudes from 1,500 to 2,500 m a.s.l. (SOKOLOV & SYROETCHKOVSKIJ 1990).

Sub-zone I/1 is called Southern Tajikistan by SOKOLOV & SYROETCHKOVSKIJ (1990). It contains the low mountain ranges of southern Tajikistan and Uzbekistan and the south Tajik depression (Aktau, Baisuntau, Babatagh, Djilantau, Karatau, Kuhitangtau, Paindjsky Karatau, Sukhaktau and Teriklitau mountain ranges). Sub-zone I/2 is called Western Pamir and contains the Darvazsky, Khazratishoh, Petr Perviy, Rushan, Shakhdara, Shugnan, Yazgulem, Vanch, and Vakhshsky mountain ranges. So far none of the two subspecies has been found in these two zones, however, localities H and I are situated in Afghanistan, not far from the Tajik border, and it is very likely that the character of this region is similar to that of sub-zone I/2.

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Sub-zone II/1 (called Eastern Pamir by SOKOLOV & SYROETCHKOVSKIJ 1990) belongs to the Central Asian mountain. It contains the Badahsan-Pamir subdivision of KRYZHANOVSKIJ et al. (1995) and mainly the highlands above 3500 m a.s.l., with mountain xerophytic open woodlands in the west, and high mountain deserts with the prevalence of high-mountain wormwood-winterfat deserts, combined with steppes and cryophytic meadows, in the east (including the high-mountain areas of the Akademiya Nauk, North and South Alichur, Beleuli, Ishkashim, North Tanimas, Sarikul, Vakhan, Zaalay, and Muzkol mountain ranges).

The Dzungar-Tian-Shan zone (I/6, I/7, and II/2) is well separated from the zoogeographic point of view, and shares more features with the fauna of the Altai than with the strongly different Western Tian-Shan (I/5) and Pamiro-Alai sub-zones (I/2, I/3 and II/1). The Western Tian-Shan zone constitutes a transition zone with marked relations to the Pamiro-Alai, and it also shares some common features with the Dzungar-Tian-Shan. According to SOKOLOV & SYROETCHKOVSKIJ (1990), the mountains of the western Talas and Fergana Ranges create a very distinct biogeographic barrier. The Pamiro-Alai is characterised by a considerably high level of endemism (e.g. Hydronebrius cordaticollis and some species of Carabus), and it is virtually unrelated to the fauna of the Tian-Shan Mountains. On the contrary, distinct links of the fauna of the Pamiro-Alai to that of northern Afghanistan, north-western China and Himalaya are known (KRIZHANOVSKIJ 1953). The Pamiro-Alai is separated from the other two zones listed above by the broad valley of the Syrdarya river and by the Fergana basin, which form a distribution barrier for mountain species. A transition zone probably exists in the eastern part of the Alai Mountains (Alai, Trans-Alai, Kichik-Alai ranges) which include more or less the localities J, 8, 9, and 10 (fig. 9).

Two other zones marked on fig. 9 do not contains any localities of the two subspecies. Zone III is named Turanic and encloses the large flat regions in the north-west of the Central Asian Mountains, and sub-zone I/4 – the Fergana basin – is the intermountain depression whose flora is presented by steppes, semi-savannas, and semi-deserts. We have little doubt that the low altitude is the chief reason for the absence of both subspecies in these zones, probably together with some other unsuitable natural conditions.

We can state that – except the somewhat doubtful specimens – the nominotypical subspecies of H. glasunovi is restricted to the zoogeographical sub-zone I/3 (plus a region in northern Afghanistan which is situated very close to sub-zone I/2), while H. g. dolini nov. ssp. occurs in sub-zones I/5 and II/2.

Zusammenfassung

Hydroporus glasunovi ZAITZEV 1905 ist eine nur wenig bekannte Art aus der planus -Gruppe der Gattung Hydroporus CLAIRVILLE 1806 . Sie ist in den Sammlungen der verschiedenen Museen nur spärlich vertreten, wurde nur in wenigen Publikationen behandelt und hier auch meist nur in Auflistungen von Sammlungsmaterial oder in Katalogen, ohne dass der Originalbeschreibung wesentliche neuere Erkenntnisse hinzugefügt wurden. Auch die systematische Stellung der Art wurde von verschiedenen Autoren unterschiedlich gedeutet. Dem Zweitautor gelang es im Jahr 1991 eine grössere Serie von Exemplaren der Art am locus typicus zu sammeln. Weiterhin konnte das Naturhistorische Museum Wien (NMW) Material erwerben, das von V.G. Dolin in Zentralasien gesammelt wurde. Erste Untersuchungen liessen vermuten, dass es sich um H. glasunovi oder zumindest um eine nahe verwandte Art handelt.

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In der vorliegenden Arbeit wird dargelegt, dass das von Dolin gesammelte Material zu einer neuen Unterart gehört, welche zu Ehren des Sammlers den Namen Hydroporus glasunovi dolini nov. ssp. erhält. Die nominotypische Unterart wird neu beschrieben und H. g. dolini nov. ssp. mittels einer Differentialdiagnose charakterisiert. Habitus und Genitale beider Unterarten werden illustriert. Ausserdem werden die Lectotypen von Hydroporus glasunovi ZAITZEV 1905 und Hydroporus macrocephalus GSCHWENDTNER 1923 designiert; der letztere wird trotz einiger Zweifel weiterhin als jüngeres subjektives Synonym von H. g. glasunovi behandelt.

Beide Unterarten sind in deutlich verschiedenen Arealen verbreitet. Die nominotypische Unterart kommt überwiegend im Norden Tadschikistans vor, aber auch im äussersten Südosten Usbekistans, im Südwesten Kirgisiens und im äussersten Norden Afghanistans. Die neue Unterart dagegen tritt im restlichen Kirgisien auf und in einem kleinen Bereich des Nordostens Usbekistans, an der Grenze zu Kirgisien. Allerdings existieren auch Populationen, deren Zuordnung zu einer der beiden Unterarten nicht eindeutig möglich ist. Schliesslich werden die Verbreitung beider Unterarten im Zusammenhang mit zoogeographischen Modellen diskutiert und Angaben zur Biologie und Begleitfauna gemacht.

References

CATE P.C. (2005): In memoriam Vladimir Gdalich Dolin (1932-2004). — Koleopterologische Rundschau 75: 403-416.

FERY H. & P.N. PETROV (2006): Nomenclatural, taxonomic and faunistic notes on selected species of Hydroporus CLAIRVILLE, 1806 (Coleoptera: Dytiscidae). — Russian Entomological Journal 14 (4) (2005): 251-262.

GSCHWENDTNER L. (1923): Einiges über Ostturkestan und dessen Dytisciden-Fauna. — Archiv für Naturgeschichte 89 (8): 93-111.

GUÉORGUIEV V.B. (1963): Contribution à l'étude des coléoptères hydrocanthares (Haliplidae et Dytiscidae) d'Afghanistan. — Opuscula Entomologica 28: 215-222.

HENDRICH L. & E. HENDRICH (2005): A contribution to the knowledge of the water beetle fauna of Uzbekistan (Coleoptera: Hydradephaga, Hydrophiloidea, Staphylinoidea and Dryopoidea). — Linzer Biologische Beiträge 37 (1): 425-434

KRYZHANOVSKIJ O. L. (1953): Zhuki-zhuzhelitsy roda Carabus Sredney Azii [Ground beetles of the genus Carabus of Central Asia]. — Opredeliteli po faune SSSR, izdavayemye Zoologicheskim institutom AN SSSR [Guides to the Fauna of the USSR. Published by the Zoological Institute of the USSR Academy of Sciences] 52. — Moscow, Leningrad: USSR Academy of Sciences Publisher, 135 pp. [in Russian]

KRYZHANOVSKIJ O.L., BELOUSOV I.A., KABAK I.I., KATAEV B.M., MAKAROV K.V. & V.G. SHILENKOV (1995): A checklist of the ground-beetles of Russia and adjacent lands (Insecta, Coleoptera, Carabidae). — Pensoft Series Faunistica 3: 1-271.

MILLER K.B. & A.N. NILSSON (2003): Homology and terminology: Communicating information about rotated structures in water beetles. — Latissimus 17: 1-4.

NILSSON A. N. (1995): Noteridae and Dytiscidae: Annotated check list of the Noteridae and Dytiscidae of China (Coleoptera), pp. 35-96. — In: JÄCH M. A. & L. JI (eds), Water beetles of China, Vol. 1. — Wien: Zoologisch-Botanische Gesellschaft in Österreich and Wiener Coleopterologenverein, 410 pp.

NILSSON A. N. (2001): World catalogue of insects. Vol. 3. Dytiscidae Coleoptera. — Stenstrup: Apollo Books, 395 pp.

NILSSON A.N. & M. HOLMEN (1995): The aquatic Adephaga (Coleoptera) of Fennoscandia and Denmark. II. Dytiscidae. — Fauna Entomologica Scandinavica 32: 1-192.

SAHLBERG J. (1910): Om Hydroporus semenovi JAKOWL. och närstående arter. — Meddelanden af Socitas Fauna et Flora Fennica 36: 167-176.

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SCHÜTZE H. & F. KLEINFELD (1997): Die Carabenformen Chinas mit dem ausführlichen Verzeichnis ihrer Fundorte. — Schwanfelder Coleopterologische Mitteilungen. Sonderheft 3, 223 pp.

SCHÜTZE H. & F. KLEINFELD (1999): Carabusformen Zentral-Asiens und Sibiriens. — Gleichen, Fürth: privately published, 242 pp.

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ŠġASTNÝ J. (1993): Water beetles in the alpine zone of the Fan Mountains. — Živa 2: 77-78.

[in Czech] WEWALKA G. (1992): Revisional notes on Palearctic species of the Hydroporus planus group

( Coleoptera : Dytiscidae ). — Koleopterologische Rundschau 62: 47-60.

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Entomologicheskoe Obozrenie 5: 25-26.

ZAITZEV F.A. (1953): Nasekomye zhestkokrylye. Plavuntsovye i vertyachki. — Fauna SSSR 4 (N.S. 58): 1-377. [in Russian]

ZAITZEV F.A. (1972): Fauna of the U.S. S.R. Coleoptera . Families: Amphizoidae, Hygrobiidae, Haliplidae, Dytiscidae , Gyrinidae. — Jerusalem: Israel Program for Scientific Translations, 401 pp.

ZIMMERMANN A. (1920): Dytiscidae , Haliplidae, Hygrobiidae, Amphizoidae. — In: SCHENKLING S. (ed.), Coleopterorum Catalogus, Vol. 4, pars 71. — Berlin: W. Junk, 326 pp.

ZIMMERMANN A. (1931): Monographie der paläarktischen Dytisciden, II. Hydroporinae (2. Teil: Die Gattung Hydroporus CLAIRV. ). — Koleopterologische Rundschau 17: 97-159.

Authors’ addresses: Hans FERY

Räuschstr. 73, D-13509 Berlin, Germany

E-mail: [e-mail]

Jaroslav ŠġASTNÝ

KosmonautĤ 359, CZ-45605 Liberec, Czech Republic

E-mail: [e-mail]

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with circles filled

(

.

ssp

.

nov dolini

.

g Hydroporus and

) capitals with squares filled

(

AITZEV Zglasunovi

. g Hydroporus of see symbols text Distribution For) other Fig. 9: numbers

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Figures

Figs 1-2: Habitus of (1) Hydroporus g. glasunovi ZAITZEV; (2) Hydroporus g. dolini nov.ssp.

Figs 3-8: Median lobe of aedeagus of in ventral and lateral view (3) lectotype of Hydroporus g. glasunovi ZAITZEV. (4) holotype of Hydroporus g. dolini nov.ssp. Gonocoxae (5) and gonocoxosternum (6) of Hydroporus g. dolini nov.ssp. Left paramere of (7) lectotype of Hydroporus g. glasunovi ZAITZEV; (8) holotype of Hydroporus g. dolini nov.ssp.

Figs 10-13: Habitats of (10-12) Hydroporus g. glasunovi ZAITZEV, Fan Mountains, Tajikistan {loc. D} (photos by J. ŠĢastný) and (13) Hydroporus g. dolini nov.ssp., Yakka-tut near Burchmulla, Uzbekistan {loc. 1} (photo by L. Hendrich).