Budapest 1971 - Magyar Természettudományi Múzeum

Budapest 1971 - Magyar Természettudományi Múzeum


Tomus 63. PARS ZOOLOGICA 1971.

Taxonomic and Zoogeographie Investigations

on the Subspecies of Leistus spinibarbis Fabr.


During the examination of the entire Leistus material of the Hungarian Natural

History Museum, Budapest, it was found that the species Leistus spinibarbis

FABR., L. magnicollis MÖTSCH., L. montanus STEPH., L. apfelbecki GANGLB.,

L. glaciális FIORI, L. gracilis Fuss, heretofore considered distinct, represent in

fact one species or a specific Formenkreis. After a study of the relevant literature

it became clear that there are also other forms, regarded until now as distinct

species, belonging to L. spinibarbis FABR., namely L. sutomorensis REITT., L.

austriacus SCHAUBERGER, L. imitator RREIT, L. punctatissimus RREIT, L. pgrenaeus

KRAATZ, 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 REITT., as a junior synonym. The holotype of L. munganasti

REITT. could be included in the material I studied, with the result that its

inclusion in the species L. spinibarbis FABR. appears to be justified. It could

happen, however, that we are dealing here with 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 REITTER 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 with 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 FABR. 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, invariably

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.

In 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 will be


Concerning external morphology, penis structure, and distribution the subspecies

of L. spinibarbis FABR. 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 with the still 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 FIORI, gracilis Fuss, imitator

BREIT, noesskei BÄNNINGER, punctatissimus BREIT, pyrenaeus KRAATZ, ucrainicus

LAZORKO. Among these subspecies some are known by one or merely a few specimens

only: austriacus SCHAUBERGER: 1 0* ; punctatissimus BREIT: 1 $ ; ucrainicus


A B C D E F G H I )

Fig. 2: A — G: Outline of male genitalia of Leistus spinibarbis apfelbecki GAJÎGLB., Volujak;

D —F: outline of prothorax of L. spinibarbis apfelbecki GANGLB., Volujak, $$; G: narrower

paramere of L. spinibarbis gracilis Fuss; H— P: outline of male genitalia of L. spinibarbis gracilis

Fuss; H: Retyezát; I: Fogaras Alps; J: Rosenauer Gebirge; K—P: Bucsecs; Q —T: outline of

prothorax of L. spinibarbis gracilis Fuss, Bucsecs -tftf

Leistus spinibarbis bjelasnicensis APFELBECK

Distribution: Yugoslavia: Bjelasnica-planina, Veles-planina.

Material examined: Bjelasnica-planina: 14 tf (Fig. 1, B—J, L —Q) 16 $ (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 variations

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. At 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, Trescavicaplanina.

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. In a

* In the enumeration of the examined material, wherever no nearer locality is given or if

Ihe locality of the label could not be identified with recourse to our maps, the locality names will

be listed in quotation-marks followed by the owner's name of the collection in 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): 19 tf (Fig. 2, K-P, Q-T), 13 9.

Of the four alpine species discussed here, this one occurred in the greatest

individual numbers in the research material. However, the distribution per locality

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: it 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 FABR.,

namely the montan«s-group, has an enormous range, occurring from Scotland to

Libanon. Aside from some exceptions, they are identical with the earlier L. montanus

STEPH. races. The subspecies bjelasnicensis APF. was formerly considered

a race of L. montanus STEPH. MARAN (1941) relegated among the races of L.

montanus STEPH. also the form L. spinibarbis abdominalis REICHE, 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, it in

fact corresponds with 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 HEER.

Distribution: Alps: Switzerland, Austria, Yugoslavia; Czechoslovakia and Panold:

Krkonose, Babia Gora.

Examined material: "Helvetia" (Coll. I. FRIVALDSZKY) : 2 tf (Fig. 3, A); "Schweiz"

(Coll. REITTER): 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


Fig. 3: Outline of male genitalia of Leistus spinibarbis rhaeticus HEEK., "Helvetia"; B —G:

outline of prothorax of L. spinibarbis rhaeticus HEER., Greifenburg, 99 ? D —F: ouline of male

genitalia and prothorax of L. spinibarbis montanus STEPH.; D, E: Nimes,tf ; F: "Aragonien", Ç ;

G—K: outline of male genitalia and prothorax of L. spinibarbis parvicollis CHAUD.; G, I : Ostri-

Medvedjak; H, J: Maranai, tf ; K: Attica, tf ; L: outline of prothorax of L. spinibarbis cyprius

MARAX, Larnaka, 0 ; M—P: outline of male genitalia and prothorax of L. spinibarbis cretensis

MAÊAX; M, N, P: "Kréta"; O: Canea; Q: outline of prothorax of L. spinibarbis cephallonicus

MARAX, "Kephallonia", Ç ; R —T: outline of male genitalia of L. spinibarbis maynicollis MÖTSCH.,


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. REITTER):

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


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. It 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 GANGLB. These characteristics also corroborate

the assumption that both magnicollis and montanus represent taxonomically but

a subspecies form each.

Leistus spinibarbis cyprius MARAN

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 CHAUD, and the subspecies rufipes GANGLB. The

description, too, was based on a female exemplar; a knowledge of the male genitalia

is still needed.

Leistus spinibarbis cretensis MARAN

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 with 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 MARAN

Distribution : Cephalonia.

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

The subspecies rufipes also inhabits this island; it 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 CHAUD.

The subspecies constituting the third group, the spinibarbis-group, inhabit

within the range of the species the geographically southernmost and physiogeographically

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. BEITTER): 1 tf (Fig. 4, A); Mustajbeg: 1 9 ; Delnice: ltf; Sutomore: 1 tf;

"Bosnia" (Coll. LICHTNECKERT): 1 9 5 "Montenegro" (Coll. BEITTER): 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


Fig. 4: A—D: outline of male genitalia and prothorax of Leistus spinibarbis magnicollis

MÖTSCH.; A: "Croatia"; B: "Montenegro", C—D: Ipek, tftf; E —M: outline of male genitalia

and prothorax of L. spinibarbis rufipes GANGLB.; E —F: Corfu; G — I: Kephallonia; J —K:

"Attica"; L — M: Kephallonia, 99 '•> N—Q: outline of male genitalia and prothorax of L. spinibarbis

abdominalis REICHE, Adana; R—S: outline of male genitalia and prothorax of L. spinibarbis

afer COQUEKEL, Ain 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 FABR. and

rufipes CHAUD. In 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. DAHL): 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 REICHE

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

Standing very near to rufipes CHAUD.; the variability of the penis and prothorax

is also apparent. Even by the meagre material, it 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).

In its external morphology very similar to the two preceding subspecies.

However, the end of the penis is only very slightly concave and it subtends an

obtuse angle with the dorsal side. This feature displays a relationship with the

Moroccan subspecies amandatus ANTIONE, 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. In one of

the male individuals, the apex of the penis is only weakly concave, and it subtends

an obtuse angle with the dorsal side, whereas in the other one no concavity is

observable and the obtuseness of the angle is less.

Leistus spinibarbis fiorii LUTSCHNIK

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 FABR.

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. REITTER):

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. At 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


Fig. 5: A —C: Outline of male genitalia and prothorax of Leistus spinibarbis expansus

PUTZ., "Lusitania"; D —F: outline of male genitalia and prothorax of L. spinibarbis

jiorii LUTSCHNIK; D: "Sicilia"; E: "Sicilia", 9; F ;

Vallombrosa, 9; G —O: ouline of

male genitalia and prothorax of L. spinibarbis spinibarbis FABR.; G, L: Ludberg, tf;

H-I: Bünde; J, O: "Anglia", M-N: "Germania", 99

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 FABR. on the basis of penis structure (L. ovipennis CHAUD.,

L. nitidus DUFT., L. rufornarginatus DUFT., L. sarduus BAUDI), and the subspecies

of L. spinibarbis FABR. 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 spinibarbisgroup.

(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

Fig. 6: Distribution of the "spinibarbis" and "oreobius" groups. The broken line

embraces the areas of the subspecies of the "spinibarbis" group, whereas the distribution

symbols explained below indicate the small-range areas of the subspecies of the "oreobius"


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 with

each other and with 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 Würm 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 Würm.

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. It 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 Würm 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. —

BINAGHI, G. (1959) : Contributo alla conoscenza délia coleotterofauna nivale del

Gran Sasso d'Italia. Bol. Soc. Ent. Ital., 89: 6-12. — BREIT, J. (1914): Beschreibung

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