ANNALES HISTORICO-NATURALES MUSEI NATIONAL1S HUNG A R I Cl Tomus 63. PARS ZOOLOGICA 1971.

Taxonomic and Zoogeographie Investigations on the Subspecies of Leistus spinibarbis Fabr.

Ry S. HORVATOVICH, Budapest

During the examination of the entire Leistus material of the Hungarian Natu­ral History Museum, Budapest, i t was found that the species Leistus spinibarbis F A B R . , L . magnicollis MÖTSCH., L . montanus STEPH., L . apfelbecki GANGLB. , L . glaciális F I O R I , L . gracilis Fuss, heretofore considered distinct, represent in fact one species or a specific Formenkreis. After a study of the relevant literature i t became clear that there are also other forms, regarded unti l now as distinct species, belonging to L . spinibarbis FABR. , namely L . sutomorensis R E I T T . , L . austriacus SCHAUBERGER, L . imitator R R E I T , L . punctatissimus R R E I T , L. pgrenaeus K R A A T Z , L . noesskei BÄNNINGER, and L . ucrainicus LAZORKO. The oldest available name of all is L . spinibarbis FABR. , 1775, hence, according to the rules of priority, this name enjoys precedence over the other ones, and is thus the valid name of the species. Of the names listed in the Coleopterorum Catalogus, BÄNNINGER drew in the name L . munganasti R E I T T . , as a junior synonym. The holotype of L . munga-nasti R E I T T . could be included in the material I studied, with the result that its inclusion in the species L . spinibarbis F A B R . appears to be justified. I t could happen, however, that we are dealing here wi th a subspecies, a question which can be resolved only by further investigations founded on a more extensive mate­rial. The holotype is a female, and REITTER'S Collection contains merely this single specimen under this name.

The major part of my research material consisted of the R E I T T E R Collection. In spite of the comparatively rich material, the presence of the subspecies may in many cases be only an assumption. The reason of this situation appears to lie in the followings. We are dealing wi th a species comprising a great number of sub­species. There is often a wide zone of intermixture between the several subspecies, considerably obstructing or hindering a safe identification. There are also some hardly accessible areas in which only a meagre material has hitherto been collected. Owing to the reasons listed above, there are many synonymous names, and the systematic place of the forms varies per author. The aim of the present paper is the separation of L . spinibarbis F A B R . from the other species and the relegation of forms, heretofore considered distinct species, to this species. Detailed drawings by the aid of a drawing apparatus have been made from the characteristic animals and series (Figures 1—5). Outlines have been made of the penis and prothorax of the males, but only of the prothorax of the females. The detailed drawings, showing identical morphologic parts, were all made by the same scale. For the males, in ­variably the drawing of the penis is given first, followed by that of the prothorax. There are also animals whose drawing of the penis is missing, because, owing to the weak chitinization, the penis was unsuitable for study.

I n the following, I propose to introduce, by the shape of the penis and the pro thorax and on the basis of the available material, the various forms; at the end of the paper, the unification into one species of the diverse forms wi l l be discussed.

Concerning external morphology, penis structure, and distribution the sub­species of L . spinibarbis F A B R . can be relegated into three groups: subspecies living in alpine, montane, and lower regions, that is, the "oreobius"', the "montanus", and the "spinibarbis" groups.

Except for L . spinibarbis bjelasnicensis APFELBECK, the oreobius-group contain forms which, together wi th the stil l valid species L . ouipennis CHATJD., formerly constituted the subgenus Oreobius. They are characterized by the following common morphologic characters: 1. decrease in body size, 2 . degeneration of the membranous wings and roundig of the elytral shoulders. The penis in a lateral view (as figured throughout in the illustrations) is thin, apically attenuating to acute. The following subspecies are assignable here: apfelbecki GANGLB. , austriacus SCHAUBERGER, bjelasnicensis APFELBECK, glaciális F I O R I , gracilis Fuss, imitator B R E I T , noesskei BÄNNINGER, punctatissimus B R E I T , pyrenaeus K R A A T Z , ucrainicus LAZORKO. Among these subspecies some are known by one or merely a few speci­mens only: austriacus SCHAUBERGER: 1 0* ; punctatissimus B R E I T : 1 $ ; ucrainicus LAZORKO: 1 <f 1 $ ; pyranaeus K R A A T Z : a few specimens. I n my research material only four subspecies were found to occur:

Leistus spinibarbis ÍJ lacialis F I O R I

Distribution: I taly: Gran Sasso dTtalia, Majella. Examined material: 1 $ (Fig. 1. A) .

The constriction of the prothorax is of the smallest rate among the four alpine subspecies. B I N A G H I ( 1 9 5 9 ) described the form ssp. relictus from the Gran Sasso d'Italia, but since this range is immediately adjoining the Majella, the validi ty of the subspecies needs revision.

A B C D E F G H I J K

L M N O P Q R S T U

Fig . 1 : A : Outline of pro thorax of Leistus spinibarbis glaciális FIORI, Majella, $ ; B — K : outline of male genitalia of L . spinibarbis bjelasnicensis APFELBECK; B — J : BjelaSnica; K : '"Bosnien"; L — U : outl ine of prothorax of L . spinibarbis bjelasnicensis APFELBECK; L — Q ;

BjelaSnica, çfçf; R — U : BjelaSnica, OÇ

r

A B C D E F G H I )

Fig . 2: A — G : Outline of male genitalia of Leistus spinibarbis apfelbecki GAJÎGLB., V o l u j a k ; D — F : out l ine of pro thorax of L . spinibarbis apfelbecki GANGLB., Vo lu jak , $ $ ; G : narrower paramere of L . spinibarbis gracilis Fuss; H — P: outl ine of male genitalia of L . spinibarbis gracilis Fuss; H : R e t y e z á t ; I : Fogaras Alps ; J : Rosenauer Gebirge; K — P : Bucsecs; Q — T : out l ine of

pro thorax of L . spinibarbis gracilis Fuss, Bucsecs -tftf

Leistus spinibarbis bjelasnicensis APFELBECK

Distribution: Yugoslavia: Bjelasnica-planina, Veles-planina. Material examined: Bjelasnica-planina: 1 4 tf (Fig. 1 , B—J, L —Q) 1 6 $ (Fig. 1 ,

R—U); Veles-planina: 1 tf; "Herzegovina" (Coll. LICHTNECKERT): 1 tf ; "Halma" (South Bosnia; Coll. REITTER) : 1 tf ; "Bosnia" (Coll. LICHTNECKERT): 1 tf (Fig. 1 , K ) . * Among the specimens bearing identical locality data, morphological vari­

ations of the prothorax can be observed, both in the males and the females. Mainly the constriction as well as the proportion of width to length of the prothorax display great variety. A similar variation is discernible also in the shape of the penis end. There are specimens in which the two minute apices of the penis are hardly visible, indeed, they may completely disappear. The end of the penis varies from an acute (70°) to a right angle. Nor do the exemplars deriving from the other localities show any greater differences, except for the one from Rosnia whose penis end is even more acute than the extreme value of that from Rjelasnica, although the genital organ is wholly chitinized. A t the same time, the prothorax of this specimen lies between the extreme values from Rjelasnica.

Leistus spinibarbis apfelbecki GANGLB.

Distribution: Yugoslavia: Volujak, Vlasulja-planina, Cvristnica-planina, Trescavica-planina.

Examined material: Volujak: 7 tf (Fig. 2, A—C), 3 $ (Fig. 2, D - F ) ; "Bosnia" (Coll. REITTER) : 1 tf ; "Herzegovina": 1 tf 1 $ ,

Standing very near to the preceding subspecies, but slightly smaller. The relatively unexplored state of the high mountains of Yugoslavia renders very difficult the clarification of the taxonomy and range of the alpine subspecies. I n a

* I n the enumeration of the examined mater ia l , wherever no nearer loca l i ty is given or i f Ihe loca l i ty of the label could not be identified w i t h recourse to our maps, the local i ty names w i l l be l isted in quotat ion-marks followed by the owner's name of the collection i n brackets.

lateral view, the penis is somewhat wider than that of the preceding subspecies. The variability of its apex is also similar, but none subtending a right angle was found to occur in the examined material. The prothoracic shape of the three females originating from Volujak is strongly varying. Those of the female exemplars bearing Herzegovinian and Bosnian locality data fell between the extreme values dispayed by the variations from Volujak.

Leistus spinibarbis gracilis Fuss

Distribution : Rumania : South Carpathians : the Szörény Alps, Retyezát, the Fogaras Alps, Bucsecs, Keresztényhavas.

Examined material : Mehadia : 1 tf 1 9 ; Retyezát : 1 tf (Fig. 2, H) , 2 9 '•> Fogaras Alps : 1 tf (Fig. 2, I) ; Rosenauer Gebirge ( = Keresztényhavas = Cristian Mare) : 3 tf (Fig. 2, J), 2 Ç; Bucsecs ( = Bucegi): 1 9 tf (Fig. 2, K - P , Q - T ) , 1 3 9 .

Of the four alpine species discussed here, this one occurred in the greatest individual numbers in the research material. However, the distribution per local­i ty is very uneven. Aside of the series from the Bucsecs, our collection contains but few specimens from the other localities. The specimen from Mehadia had a wholly unchitinized penis, entirely unsuitable for examination. The penis of the single male specimen from the Retyezát differs from that of the other exemplars originating from the other localities: i t is narrower, nearly parallel-sided, the apex almost rectangular. On the other hand, the genital organ of the male exemplar from the Fogaras Alps does not differ from that of the individuals deriving from the Keresztényhavas and the Rucsecs. In the specimens from this latter locality, the variation of the prothoracic and penis forms also occurs.

The subspecies group living in montane habitats of L . spinibarbis F A B R . , namely the montan«s-group, has an enormous range, occurring from Scotland to Libanon. Aside from some exceptions, they are identical wi th the earlier L . mon­tanus STEPH. races. The subspecies bjelasnicensis A P F . was formerly considered a race of L . montanus STEPH. M A R A N ( 1 9 4 1 ) relegated among the races of L . montanus STEPH. also the form L . spinibarbis abdominalis R E I C H E , described from Jerusalem. The single specimen serving for the basis of his statement originates from the mountainous region of Libanon, and, according to the description, i t in fact corresponds wi th a L . montanus STEPH. race in the earlier sense, but the exemplars of the Museum collection do not belong to the montanus-gr oup.

Leistus spinibarbis rhaeticus H E E R .

Distribution: Alps: Switzerland, Austria, Yugoslavia; Czechoslovakia and Panold: Krkonose, Babia Gora.

Examined material: "Helvetia" (Coll. I . FRIVALDSZKY) : 2 tf (Fig. 3, A ) ; "Schweiz" (Coll. R E I T T E R ) : 1 9 ; Kraina: Greifenburg: 2 ? (Fig. 3 / B - C ) .

The subspecies should possibly be split into several forms but not enough material was available to decide the question. The female specimens from Greifen­burg differ from each other in body size, as shown also by the dimensions of the prothorax. The genital organ of the single male, suitable for study, from Switzer­land resembles to a great extent that of the oreobius-group.

A B C D E F G H 1 3

K L M N O P Q R S T

Fig . 3: Outl ine of male genitalia of Leistus spinibarbis rhaeticus H E E K . , " H e l v e t i a " ; B — G : outline of prothorax of L . spinibarbis rhaeticus H E E R . , Greifenburg, 9 9 ? D — F : ouline of male genitalia and prothorax of L . spinibarbis montanus S T E P H . ; D , E : Nimes, t f ; F : "Aragonien" , Ç ; G — K : out l ine of male genital ia and pro thorax of L . spinibarbis parvicollis C H A U D . ; G, I : Ost r i -Medvedjak; H , J : Maranai , tf ; K : A t t i ca , tf ; L : outline of pro thorax of L . spinibarbis cyprius M A R A X , Larnaka , 0 ; M — P : outline of male genitalia and p ro thorax of L . spinibarbis cretensis M A Ê A X ; M , N , P: " K r é t a " ; O: Canea; Q : outl ine of pro thorax of L . spinibarbis cephallonicus M A R A X , "Kepha l lon i a " , Ç ; R — T : outline of male genitalia of L . spinibarbis maynicollis M Ö T S C H . ,

Ipek.

Leistus spinibarbis montanus STEPH.

Distribution: England, Scotland, Ireland, France, Western Switzerland, Mountainous ranges of Northern Spain.

Examined material: Nîmes: 2 tf (Fig. 3, D —E); "Aragonien" (Coll. R E I T T E R ) : 1 9 (Fig. 3, F).

Similarly to the situation in the preceding subspecies, here, too, examinations of series are needed. The prothorax of the female specimen from Aragónia ( = the holotype of L . munganasti REITTER) is much more constricted than that of the males from Nîmes whose penis agrees with the shape characteristic of the sub­group.

Leistus spinibarbis parvicollis CHAUD.

Distribution: Mountainous areas from Istria through South Yugoslavia and Albania to South Greece.

Examined material: Yugoslavia: Ostri Medvedjak (Fig. 3, G, I ) ; 1 tf; Crkvenica: 1 9 ; Velebit : 1 tf ; 2 9 ; Cemerno : 2 9 ; Albania : Maranai : 1 tf (Fig. 3, H , J) ; Greece: "Attica" (Coll. REITTER) : 1 tf (Fig. 3, K ) ; 1 9 ; "Turciä" (Coll. L FRIVALDSZKY) : 1 9 •

There is no series from an identical locality in the material to show the var­iability of the penis apex. Of the four male individuals, the one from Ostri Med­vedjak seems to be transitional in the shape of both the prothorax and the male genitalia to the subspecies magnicollis MÖTSCH. I t is only the prothoracic shape of the animal from the Albanian Maranai which resembles that of magnicollis MÖTSCH. On the other hand, the form of the prothorax of the specimen from Attica stands

near that of the subspecies rufipes G A N G L B . These characteristics also corroborate the assumption that both magnicollis and montanus represent taxonomically but a subspecies form each.

Leistus spinibarbis cyprius M A R A N

Distribution: Cyprus. Examined material: Larnaka: 1 0 (Fig. 3, L) .

The prothorax of the examined female occupies an intermediate place between that of the subspecies parvicollis C H A U D , and the subspecies rufipes G A N G L B . The description, too, was based on a female exemplar; a knowledge of the male genitalia is still needed.

Leistus spinibarbis cretensis M A R A N

Distribution: Crete. Examined material: "Krete" (Coll. PAGANETTI) : 3 tf (Fig. 3, M, N , P), 1 Ç ; Canea:

1 tf (Fig. 3, O).

The shape of the prothorax almost wholly agrees wi th that of rufipes GANGLB. However, the form of the penis is widely different, corresponding to that of the forms belonging to the montanus-group. The genital organ of the males also varies.

Leistus spinibarbis cephallonicus M A R A N

Distribution : Cephalonia. Examined material: "Kephallonia" (Coll. STREDA): 1 $ (Fig. 3, Q).

The subspecies rufipes also inhabits this island; i t differs widely by the shape of its prothorax. The distribution of the two subspecies within the island is un­known to me. The prothoracic shape of the examined female stands very near to that of the subspecies parvicollis C H A U D .

The subspecies constituting the thi rd group, the spinibarbis-group, inhabit within the range of the species the geographically southernmost and physiogeogra-phically lowest regions. Knowledge concerning the area of the several subspecies is still highly incomplete. A special difficulty in identification lies in the fact that the differences between the taxa are the least conspicuous in this group, rendering further obstructions also in recognizing the zones of intermixture. Their common morphological features are: the most (angularly) developed elytral shoulder and apically concave penis, the apex and the sides subtending (more or less) a rectangle.

Leistus spinibarbis magnicollis MÖTSCH.

Distribution: Inner areas of the Balkan Peninsula: from Croatia and the Domoglcd Range to Northern Greece.

Examined material: Rumania: Herkulesfürdő ( = Baia Herculanea): 2 tf 2 9 î Mts. Domoglcd: 1 9 > "Atrum Imre" (Coll. Mihók): ltf ; Yugoslavia: "Croatia" (Coll. B E I T T E R ) : 1 tf (Fig. 4, A ) ; Mustajbeg: 1 9 ; Delnice: ltf; Sutomore: 1 tf; "Bosnia" (Coll. LICHTNECKERT): 1 9 5 "Montenegro" (Coll. B E I T T E R ) : 1 tf (Fig. 4, B ) ; Albania: Merdita: 2 tf; Ipek: 3 tf (Fig. 3, R - T ; 4, C - D ) ; Greece: Veluchi: 1 9 .

A B C D E F G H I 5

K L M N O P Q R S

F i g . 4: A — D : outl ine of male genitalia and pro thorax of Leistus spinibarbis magnicollis MÖTSCH.; A : "Croa t i a " ; B : "Montenegro" , C — D : Ipek , tftf; E — M : outl ine of male genitalia and pro thorax of L . spinibarbis rufipes GANGLB.; E — F : Corfu; G — I : Kepha l lon ia ; J — K : " A t t i c a " ; L — M : Kephal lonia , 9 9 '•> N — Q : outl ine of male genitalia and pro thorax of L . spini­barbis abdominalis REICHE, Adana ; R — S : outl ine of male genitalia and pro thorax of L . spini­

barbis afer COQUEKEL, A i n Draham.

The penis of the three male specimens from the environs of Ipek (Möns Zljeb) vary. The penis form of the exemplar from "Croatia" and that of two individuals from Ipek is transitional towards that of the subspecies spinibarbis F A B R . and rufipes C H A U D . I n these animals the apex of the penis is considerably wider than the characteristic measure. Variations are observable also in the size and shape of the prothorax.

Leistus spinibarbis rufipes CHAUD.

Distribution: From Istria to Southern Greece, also on the islands. Examined material: Yugoslavia: Lie: 1 tf 1 Q ; Fiume: 1 tf; Istria: 1 9 J "Ulyria"

(Coll. D A H L ) : 2 tf; Bakar: 1 9 ; Velebit: 1 9 ; Mostar: 1 9 ; Albania: Maranai: 1 tf 2 9 ; Greece : Corfu : 2 tf (Fig. 4, E, F) ; Kephallonia : 4 tf (Fig. 4, G, I ) , 5 9 (Fig. 4, L - M ) ; Oxyá: 6 9 ; ' 'Attica" (Coll. BEITTER) : 2 tf (Fig. 4, J - K ) , 2 9 .

The subspecies inhabiting the western part of the Balkan Peninsula exhibited prothoracic and penis variations whenever more than one specimen were present in the research material. This variation fluctuates between strikingly great extreme limits in individuals deriving from Kephallonia, but this is also apparent, though at a lesser rate, in specimens originating from the other localities listed above.

Leistus spinibarbis abdominalis R E I C H E

Distribution: Lower elevations of Asia Minor and Israel. Examined material: Turkey: Adabasar: 1 tf; Adana: 2 cf (Fig. 4, M —Q); "Aegyptus"

(Coll. Kertész); 1 cf ; Jerusalem: 1 0 M $ ; "Palestina" (Coll. Reitter): 1 9 .

Standing very near to rufipes C H A U D . ; the variability of the penis and protho­rax is also apparent. Even by the meagre material, i t can safely be stated that they do not belong to the mon/amis-group.

Leistus spinibarbis afer COQTJEREL

Distribution: Mountainous districts of Tunisia and Algeria. Examined material: Tunisia: Ain Draham: 1 tf (Fig. 4 , R, S).

I n its external morphology very similar to the two preceding subspecies. However, the end of the penis is only very slightly concave and i t subtends an obtuse angle with the dorsal side. This feature displays a relationship wi th the Moroccan subspecies amandatus A N T I O N E , which, again, is closely related to the subspecies expansus PUTZ. , from Spain and Portugal. Hence the examined specimen of afer COQUEREL is a transition between the Eastern and Western Mediterranean branches of the subspecies-group.

Leistus spinibarbis expansus PUTZ.

Distribution: Portugal, Spain. Examined material: Portugal: "Lusitania": 2tf (Fig. 5, A —G); Spain: Durango: 1 $ .

The prothorax of the examined specimens is constricted nearly as much as in the subspecies of the montanus-group. However, the shape of the penis reveals unquestionably that this subspecies belongs to the spinibarbis-group. I n one of the male individuals, the apex of the penis is only weakly concave, and i t subtends an obtuse angle wi th the dorsal side, whereas in the other one no concavity is observable and the obtuseness of the angle is less.

Leistus spinibarbis fiorii L U T S C H N I K

Distribution: Sicily and South Italy. Examined material: "Sicilia" (Coll. LICHTNECKERT): 1 tf (Fig. 5, D): 1 O (Fig. 5, E) ;

Mts. Silla: 1 tf; Vallombrosa: 1 $ (Fig. 5, F).

The prothorax of the female from Vallombrosa (near Florence) is considerably larger than that of the other specimens studied. The form of the prothorax and the penis conforms with that characteristic of the spinibarbis-group.

Leistus spinibarbis spinibarbis F A B R .

Distribution: South England, South Germany, South Poland, West Czechoslovakia, Switzerland, Austria, Northwest Yugoslavia, North Italy, France.

Examined material: "Anglia" (Coll. LICHTNECKERT): 1 tf (Fig. 5, J , O ) ; Germany: Hildesheim: 1 tf; Bünde: 3 tf (Fig. 5, H - I ) , 1 $ ; "Germania" (Coll. R E I T T E R ) : 5 9 (Fig. 5, M, N) ; France: "Pyrenaei" (Coll. I . FRIVALDSZKY): 1 9 ; "Gallia" (Coll. APFELBECK): 1 tf (coll. I . FRIVALDSZKY): 2 9 '•> Yugoslavia: Ludbreg: 1 tf (Fig. 5, G, L), 2 9 ; "Ausztria" (Coll. LICHTNECKERT): 1 tf (Fig. 5, K ) .

The prothoracic shape of the exemplars from the environs of Ludbreg rep­resents a transition towards magnicollis MÖTSCH. A t the same time, the shape of the penis is characteristic of spinibarbis. One of the specimens from Germany has a much smaller prothorax than the average. The extreme values of the penis variations shown by the animals originating from Bünde are shown in Fig. 5. H , L

A B C D E F G H

F i g . 5: A —C: Out l ine of male genitalia and pro thorax of Leistus spinibarbis expansus PUTZ., " L u s i t a n i a " ; D — F : outl ine of male genitalia and prothorax of L . spinibarbis jiorii LUTSCHNIK; D : "S ic i l i a " ; E : "S ic i l i a" , 9 ; F ; Vallombrosa, 9 ; G —O: ouline of male genitalia and prothorax of L . spinibarbis spinibarbis FABR.; G, L : Ludberg , tf;

H - I : B ü n d e ; J , O: " A n g l i a " , M - N : "Germania" , 9 9

To substantiate my statement that the three subspecies groups, discussed in detail on the basis of my research material, constitute a single species, the following arguments can be put forth:

1. The formation of the end of the penis. The penis illustrations submitted in the Figures unequivocally imply that the differences in shape fail to attain the specific level. Transitional forms can namely be found between the subspecific forms whose indentification and relegation cause grave difficulties. I have examined the formation of the male genital organ of all Leistus species represented in the collections of the Museum, and I found that the differences between the species are of a considerably higher order than those exhibited by the subspecies among themselves of L . spinibarbis FABR. , and no transitional forms could be detected between the penis forms of the former species. The difference between the species, related to L . spinibarbis F A B R . on the basis of penis structure ( L . ovipennis CHAUD. , L . nitidus D U F T . , L. rufornarginatus D U F T . , L . sarduus B A U D I ) , and the subspecies of L . spinibarbis F A B R . ist distinct and without transitions.

2. The formations of the parameres. Fig. 2, G shows the outlines of the narrower paramere of L . spinibarbis gracilis Fuss; the paramere has a very weakly chitinized and basally widening process, situated on the inner side. Every L . spinibarbis subspecies has this process, but none of the related species.

3. The distribution of the subspecies and subspecies-groups. Figure 6 illustrates the conditions of distribution of the spinibarbis- and oreobius-groups. The pecked line delimits the area of the overlapping, contiguous members of the spinibarbis-group. (The pecked line was plotted also on the sea for the sake of a complete illustration). The distribution of the several alpine subspecies is given by the application of various symbols: the map gives the range of all hitherto known alpine subspecies. As is to be clearly seen, the distribution of the alpine subspecies, except for that of ucrainicus LAZORKO and gracilis Fuss, lies within the area of L . spinibarbis. I have not shown the distribution of the montanus-group which is,

C ssp. opfelbecki Ganglbauer B Ssp. austriacus Schauberger

(D ssp. bjelasnicensis Apfeibeck • ssp. glaciális Fiori

® ssp. gracilis Fuss • ssp. imifaror Breir

ssp. noesskèi Bânningei • ssp. puncrah'ssimus Breir

• ssp. pyrenaeus Kraorz H ssp, ucrainicus Lazorko

F ig . 6: Dis t r ibu t ion of the "spinibarbis" and "oreobius" groups. The broken line embraces the areas of the subspecies of the "spinibarbis" group, whereas the d i s t r ibu t ion symbols explained below indicate the small-range areas of the subspecies of the "oreo­

bius" group

again except for the occurrences in Ireland and Scotland, also within the range of the spinibarbis-group. As pointed out in the discussion, the distribution of the three subspecific groups extends in three ecologically different regions (levels).

The evolvement and spreading of the three subspecies groups may be satis­factorily explained only by taking into account also the climatic and zoogeograph-

ical conditions prevailing during the last (Würm) glacial period. The European core of the Würm glaciation was in the snow cover over Scandinavia. England and Ireland were but partially covered by ice, and had been interconnected both wi th each other and wi th Europe. The presumably much more uniform populations of the species representing the ancestor of the three subspecies-groups were segregated by the ice cover of the W ü r m into three areas of diverse climate. The climatic conditions prevailing in the neighbourhood of the ice cover were uniform for extensive areas and extended, at mountainous regions of a suitable height, also to South Europe and Asia Minor. At the same time, the climate was considerably milder at the lower altitudes above sea level in South Europe. The members of the mont anus-group lived in the former area during the Würm, whereas those of the spinibarbis-group in the letter one. The differences in the climate of the two regions effected the morphological deviations extant between the members of the two subspecies groups. In the south, the areas with a colder climate in the mountainous regions formed a mosaic-like distribution according to the orographic conditions, that is, the several parts were more or less isolated from one another. As a con­sequence, the montanus-group split into considerably more subspecies than in Northern Europe, and the differences among them are also bigger. The differences between the members of the spinibarbis-group are smaller than those between the mon/amzs-group which approximately corresponds to the regional differences in the climatic conditions during the W ü r m .

The subspecifis oreobius-group evolved from those populations which sur­vived the last glacial period in the "massif de refuge" zone of the high mountains; their distribution is insular in character. The differences between the several subspecies are the greatest in this group, since the populaticns of small individual numbers had been completely isolated from both one another and the other sub­species. The number of subspecies may, owing to this small number of individuals constituting the given populations, increase considerably in the wake of future investigations. I t is rather striking when studying the map of distribution, that the subspecies ucrainicus LAZORKO and gracilis Fuss, insular in character, is wholly outside of the range of the spinibarbis-group. Their peripheral position implies that there had been a considerable "massif de refuge" during the W ü r m in the territory of the Eastern and Southern Carpathians and that this area had preserved the species—of a greater distirubtion during the Riss-Würm interglacial—-in the form of new alpine subspecies. The botanical and zoological exploration of the environs of the present ice cover demonstrated that there live a comparatively rich flora and fauna in some smaller areas immediately adjoining the ice cover (LlNDROTH, 1970).

References: ANTOINE, M. (1939): Coléoptères nouveaux du Moyen Atlas. Bul. Soc. Ent. France, 44: 145 — 152. — ANTOINE, M. (1955): Coléoptères Carabiques du Maroc. Paris—Rabat: 38—40. — APFELBECK, V. (1904): Die _Käferfauna der Balkanhalbinsel. Berlin: 48—51. — BÄNNINGER, M. (1931): Über Carabinae, Ergänzungen und Berichtigungen (Col.) 17. Beitrag. Deutsche Ent. Zeitschrift, 57: 177 — 212. — BÄNNINGER, M. (1956): Über Carabinae, Ergänzungen und Be­richtigungen (IV). Entom. Arbeiten aus dem Museum Gg. Frey, 7: 398—411. — B I N A G H I , G . (1959) : Contributo alla conoscenza délia coleotterofauna nivale del Gran Sasso d'Italia. Bol. Soc. Ent. Ital., 89: 6 -12 . — BREIT, J . (1914): Beschrei­bung zwölf neuer palaearktischer Coleopteren-Formen aus der Familie Carabidae. Col. Rundsch., 3: 155 — 170. — Fuss, K . (1860): Leistus gracilis und alpicola zwei neue siebenbürgische Käfer-Arten. Verh. und Mitteil, des siebenbürg. Vereines für Naturwiss., 11: 29—36. — GANGLBAUER, L . (1892): Die Käfer von Mitteleuropa. Wien: 93-97. - HORION, A. (1941): Faunistik der Deutschen Käfer. Band 1:

68—74. — JEANNEL, B . (1941): Coléoptères Carabiques. Paris 1 : 177 — 185. — LAZORKO, W. (1954): Leistus ucrainicus spec. nov. eine neue Carabiden art aus der Ukraine mit einer Übersicht der bisher bekannten Arten der "Oreobius" Gruppe (Col. Carabidae). Entom. Arbeiten aus dem Museum Gg. Frey, 5: 81—91. — LINDROTH, C. (1970): Survival of animals and plants on icefree réfugia during the Peistocene glaciations. Endeavour, 2 9 : 129 — 134. — MAGISTRETTI, M . (1965): Coleoptera, Cicindelidae, Carabidae. Fauna d'Italia: 59—65. — MARAN, J . (1941): O geografickém rozsireni a rassách druhu Leistus montanus Steph. De Leisti montani Steph. distributione et variatione geographica (Coleoptera, Carabidae). Sbornik entom. odd. Zemského Musea v Praze, 19 : 131 — 135. — REITTER, E . (1885): Analytische Übersicht der bekannten europäischen Arten der Coleopterengattung Leistus Fröhl. Wiener Ent. Zeitung, 4 : 213-219. — REITTER, E . (1905): Zur systematischen Gruppeneinteilung des Coleopteren-Genus Leistus Fröhlich und Übersicht der mir bekannten Arten. Wiener Ent. Zeitung, 24 : 209—225. — SCHAUBERGER, E . (1925): Zwei neue paläarktische Carabiciden. Ent. Anzeiger, 5: 19—21. — SCHWEIGER, H . (1970): Neue Leistus-Formen von der Balkanhalbinsel und aus Anatolien (Col., Carabidae). Entomol. Blätter, 6 6 : 57—64.