ANNALES HISTORICO-NATURALES MUSEI NATIONAL1S HUNG ARI 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, 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
K L M N O P Q R S T
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
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. 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
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
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
K L M N O P Q R S
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
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),
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
A B C D E F G H
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
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Balkanhalbinsel. Berlin: 48—51. — BÄNNINGER, M. (1931): Über Carabinae,
Ergänzungen und Berichtigungen (Col.) 17. Beitrag. Deutsche Ent. Zeitschrift,
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