Morphocarabus and Eucarabus in Serbia
by Ivo Tosevski
Populations of subgenus Morphocarabus and Eucarabus in Serbia, their morphology, diversity,
distribution and speciation level.
The genital apparatus, of such importance in taxonomy and systematics of other insect groups, is
poorly researched in genus Carabus Lineus, 1758. If we except studies of Ishikawa (1973, 1978, 1979,
1984), contributions in this sense are almost symbolical, especially when European authors are
concerned. Thus the monograph published by Breuning (1932-1937) is still the most significant work
dealing with taxonomy and systematics of Palearctic Carabus species.
The genital apparatus of the male, i.e. its internal structure endophallus with all its specialised
structures represents the significant morphological characteristic for determination of the taxonomic
status of species. Phylogenetically, endophallus as morphological characteristic clearly groups similars
species within a single taxon. In literature, such an group of similar species is treated as subgenus,
although similarity and stability of endophallus morphology suggest that these taxons could be treated
as separate genera. It seems that the separation of Carabus into several genera would introduce more
order in systematics of these insects. On the other hand, Carabus species as distinctly terricolous insects
have diminished possibility of dispersion due to unfunctional wings, most often present in these insects in
rudimentary form. The consequence is great species diversity which in some species results in formation
of a great number of subspecies with clear geographical distribution. Diversity is not necessarily present
in all populations of a species, but is more often distinct in areas labeled as refugial. This seems to be
one of the reasons why diversity of certain species of Central and Northern Europe is less present when
compared with the same species in the Balkans region. Therefore, the study of endophallus morphology
as an objective morphological characteristic of these insects would surely advance taxonomy and
systematics of Carabus species. In literature, the taxonomic status of a species, relationship between
species and the status of their populations, is often the fruit of author's subjective attitude, while the
methodological way of reaching the conclusions can not be followed.
This site aims to use World Wide Web to present the problem of the taxonomic status of some
Carabus species populations in the Balkans region, speciation degree of populations within
Morphocarabus subgenus (type species Carabus monilis), and pronounced diversity of Carabus
(Eucarabus) ullrichi.
The basis of this presentation is its author's 1990 study entitled "Betraig zur Kenntnis der
serbischen Population der Gattung Morphocarabus Gehin, 1885 (Coleoptera, Carabidae). Acta Mus.
Maced. Sci. nat., 19, 4/157:85-110". In this paper Morphocarabus populations on the territory of the
former Yugoslavia were treated as a subspecies of monilis species and classified - on the basis of the
collected and reviewed material - as 5 subspecies and several nations.
Morphocarabus monilis ssp. simulator Kraatz, 1876
Morphocarabus monilis ssp. praecellens Palliardi, 1825
Morphocarabus monilis ssp. illigeri Dejean, 1826
Morphocarabus monilis ssp. bjelasnicensis Apfelbeck, 1902
Morphocarabus monilis ssp. semetrica Kraatz, 1878
This classification was, in a way, the result of trends present in literature from 70's and 80's mainly
based on contributions of Mandl 1965, Machard 1979 and Ivanovs 1981.
Later additional research of morphological characteristics of endophallus in these populations
showed that differentiation and the speciation degree of above quoted subspecies are developed up to
the level of bona species. Thus the concept of C. (Morphocarabus) monilis as a singular species with
important number of subspecies in this region is untenable. A disproportion in divergence of external
morphological characteristics and endophallus morphology is striking. It is evident that monilis species
populations originating in France have pronounced diversity of external morphology of elytra which is not
followed by the morphological differentiation on endophallus level. On the contrary, the external
morphology diversity in East and South-East Europe is often symbolical, but the differentiation in
endophallus morphology is exceptionally pronounced. In the material concerning Morphocarabus
population in Serbia, the morphological differentiation of endophallus in neighbouring populations was
observed (stable and constant for each of populations studied). In geographical sense, these populations
occupied clearly defined distribution areals.
Although phenotypically very close, all above mentioned populations manifested difference in form
of endophallus. This difference was significant even in very similar populations where differences in
sculpture of elytrae - as the most significant element in comparison of external morphology - was not so
pronounced. Endophallus, as a morphological characteristic clearly systematized the populations studied
into separate taxons. Great diversity of Carabus (Morphocarabus) monilis species is well known in
literature, while the species nomenclature comprises 186 names (Deuve, 1994), whose taxonomic status
is still not clear. Findings of stable characteristics in endophallus morphology in species of this subgenus
suggest the differentiation of these species on the level of bona species for the Balkans populations.
2
Similar situation can be observed in Romanian Morphocarabus populations. It is certain that the
intensive spetiation process occured in near past, so that they cling, in geographical sense, to their strict
distribution areals; up to now the sympatricity of neighbouring species was not confirmed. Over long
enough time period geographical barriers divided these populations so that the stability of endophallus
morphology - representing potential mechanical isolation during mating - is evidently genotypically
determined. Also, each population in its own distribution areal manifests different degrees of diversity in
regard to the populations of the neighbouring taxon. The case of Balkans Morphocarabus populations
thus represents a example of geographical (allopatric) speciation. Therefore, we maintain it is justified to
treat thus differentiated taxons as bona species:
Carabus (Morphocarabus) bjelasnicensis Apfelbek, 1902
Carabus (Morphocarabus) praecellens Palliardi, 1825
Carabus (Morphocarabus) kollari ssp. semetricus Kraatz, 1878
Carabus (Morphocarabus) simulator Kraatz, 1876
In spite of pronounced differences in endophallus morphology of Balkans Morphocarabus
populations, phenotypically, differences between these species are not especially significant. On the
contrary, in Eucarabus subgenus the species are, phenotypically clearly differentiated, while endophallus
morphology seems sigificant only on the level of species. Although populations of these species manifest
considerable diversity (Battony & Breuning, 1970; Pavicevic & Tosevski, 1988), clear changes of
endophallus morphology were not observed even in the populations which are geographically distant and,
phenotypically, significantly different. In the Serbian fauna, from the species of this subgenus only is
Carabus (Eucarabus) ullrichi Germar, 1824 present. This species is widely distributed and occupies
very different types of habitats, while Morphocarabus populations are local and occupy very narrowly
defined eco-systems.
Basic differences observable in Serbian populations of ullrichi species are diffrences in elytra
sculpture and colour. These differences are sometimes very pronounced in neighbouring populations, but
can also be very discrete; therefore it is, methodologically, difficult to establish an objective model that
could be clearly applicable in determination of their taxonomical status. Even in highly homogeneous
populations, according to observed taxonomic characteristics (e.g. arrogans, gornjakensis, jaroslawi), can
be found specimens (most frequently < 10%) which differ from the general habitus of the population
observed. In this sense it can be said that each such population on the areal border with the neighbouring
population represents a very heterogenous whole wherein predominate specimens possessing common
taxonomic characteristics of both populations. Such transitory populations are most frequent, and they
inhabit the greatest part of areal. Populations homogenously differentiated in regard to habitus are
commonly encountered in a very limited geographical area.
If we accept Mayer's definition of subspecies as "an aggregate of phenotypically similar
populations inhabiting a geographical subdivision of the range of that species and differing
taxonomically from other populations of that species", then, generally speaking, colour of population
is the only characteristic which determines the main phenotypical characteristics of populations so that
they can be divided into monochromatic and polychromatic. Completely blue coloured specimens are
genotypic characteristic of individua and probably are dominantly-recessively inherited. Blue colour is
characteristic of some populations in the eastern part of distribution areal of ullrichi species. In Serbia,
such specimens are present only in n. jarosawi and n. transdiernae.
This division seems most acceptable as the whole corpus of ullrichi species could be divided into
two subspecies: nominal (monochromatic) ssp. ullrichi Germar, 1824 and polychromatic ssp.
fastuosus Palliardi, 1825. Numerous populations within these subspecies with all their differences,
commonly observable in the quality of elytrae sculpture, nevertheless possess only infraspecific
characteristic and the most acceptable procedure is to treat them as nations, although from the
standpoint of valid ICZN they are of no taxonomic significance. Pronounced diversity within populations,
particularly visible in East Serbia is significant and objectively present, but insufficient to treat some of
these populations as a separate taxon on the level of subspecies. This is the most sensitive of the
decision a taxonomist makes when faced with a similar case: should such populations be proclaimed a
subspecie s, thereby meeting ICZN criter ia or should the present diversity be shown in a manner that
objectively reflects the field situation. In that context, a nation represents a population or a group of
populations (which can be dispersed within the areal) with their own taxonomic characteristics but
belongs to an agregate of phenotypically similar populations marked as subspecies, possessing a clear
geographical distribution. In a geographical sense, there is a distinct border of areal inhabited by
monochromatic and polychromatic populations. Such a clear geographical demarcation is drastically
manifest in Gornjak Gorge where two nations, two different subspecies are sharply separated by a small
mountain river.
Literature:
Battoni, S. & Breuning, S., 1970. Una nuova natio di Carabus ullrichi Germ. della Serbia Orientale. Bullettino della
Societa Entomologica Italiana. Volume 102: 9-10. Genova.
3
Breuning, S. 1932-1937. Monographie der Gattung Carabus L.- Bestimmungstabellen der europäischen
Coleopteren, 104-110, ed. Troppau.
Deuve T., 1994. Une classification de Genre Carabus. Vol. 5, pp. 296. ed. Sciences Nat.
Ishikawa, R., 1973. Notes on some basic problems in the taxonomy and the phylogeny of the subtribe Carabina
(Coleoptera, Carabidae). Bull. Natn. Sci. Mus. Tokyo. 191-215.
Ishikawa, R., 1978. A revision of the higher taxa of the subtribe Carabina (Coleoptera, Carabidae). Bull. Natn. Sci.
Mus. Ser. A (Zool.), (1):45-68.
Ishikawa, R., 1979. A preliminary revision of the Carabogenici of the subtribe Carabina (Coleoptera, Carabidae).
Bull. Natn. Sci. Mus. Serr. A (Zool.), 5(2):95-114.
Ishikawa, R., 1984. Phylogeny and Subgeneric classification of the Genus Chaetocarabus (Coleoptera Carabidae).
Kontyu. Tokio (1):94-109.
Pavicevic D., & Tosevski I., 1988. Einige neue nationes von Carabus (Eucarabus) ullrichi fastuosus Palliardi aus
Nordost-Serbien (Coleoptera, Carabidae). Fragmenta Balcanica, Vol.13:113-118.
Tosevski I., 1990. Betraig zur Kenntnis der serbischen Population der Gattung Morphocarabus Gehin, 1885
(Coleoptera, Carabidae). Acta Mus. Maced. Sci. nat., 19, 4/157:85-110.
Dr. Ivo Tosevski:
tosevski@eunet.yu
Endophallus morphology in diferent Carabus species
Lateral view of endophallus in Carabus species belonging to different subgenera (with exeption of
croaticus and violaceus which belong to subgenus Megodontus).
Eucarabus - adults and endophallus morphology
Evident similarity of endophallus morphology of different species within a subgenus can sugest that the
differentiation of morphological characteristicts is more important than subgenus level.
4
Adult morphology
Caudal view of endophallus
5
Lateral view of endophallus
Distribution of the Balkans Morhocarabus populations
The map shows the areas where the presence of Morphocarabus populations was reliably
ascertained. The data encountered in literature (primarily data quoted by Breuning, 1932-1937) were not
included, due to the impossibility of their taxonomic verification and possible errors in locality identification
(e.g. the mention of comptus from locality Golubinje, East Serbia).
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Carabus (Morphocarabus) bjalasnicensis Apfelbek, 1902 bona sp.
Distribution: West of the Velika Morava river (Central, North-West, West and South-West Serbia
and accross the Drina river up to Central and North-West Bosnia). Northern border of areal is the Sava
river.
Diversity: on the territory of Serbia Carabus (Morphocarabus) bjelasnicensis populations are
represented by three subspecies: ssp. koshanini, ssp. tarensis i ssp. cerensis. Morphologically, by the
form of endophallus they clearly belong to bjelasnicensis but they also clearly differ among themselves,
i.e. they manifest their pecularities. In external morphology these subspecies differ in organisation of
primary, secondary and tertiary elytrae sculpture, dominant population colour, and the type of habitat.
Subspeciae koshanini and tarensis are polychrome, with roughly expresed sculpture, while the cerensis
subspecies is practically monochromatic (blue coloured specimens are dominant) and have considerably
finer sculpture.
Habitat: the koshanini is clearly platicolous species, inhabits alpine and subalpine meadows,
tarensis is silvicolous, while cerensis inhabits the edges of light deciduous forests and clearings with
sparse vegetation.
Endophallus morphology: tarensis and koshanini evidently have the greatest similarity with the
nominal subspecies bjelasnicensis. In these subspecies caudal part of distal lobe seen from dorsal side in
region of pygmenteum field is almost square, while in cerensis this part of lobe is longer and rectangular.
Nomenclature: It is not quite clear which name has priority for the populations designated
bjelasnicensis. The reason is unclear or unknown type locality of current names. On the other hand, all
populations in this region are phenotypically similar, and their objective status in regard to other
populations can be determined only on the basis of endophallus morphology. The case of bjelasnicensis
Apfelbek, 1902, name is very indicative. According to the nomenclature quoted by Breuning (1932-1937)
at least 5 names have priority.
illigeri Dejean, 1826 (type locality "Croatia"?!)
laticollis Sturm, 1826 (type locality "Croatia"?!)
stentzi Villa, 1833 (type locality not known)
parallelus Kraatz, 1878 (Mt. Pljesevica, Bosnia)
curtulus Gnglb., 1892 (type locality: hills surrounding Sarajevo)
bjelasnicensis Apfelbek 1902 (Mt. Bjelasnica, near Sarajevo, Bosnia)
Due to the lack of material for genitalisation and analysis of endophallus morphology as the only
reliable sign in proving of phylogenetic affiliation of a population, it was not possible to confirm that
curtulus Gnglb., 1892 represents senior synonym of bjelasnicensis Apfelbek, 1902, although it seems
only possibility. Other quoted names also were potential candidates as it can be expected that parallelus
Kraatz, 1878 (locus typicus Pljesevica, Bosnia) belongs, due to the type of endophallus morphology, to
the populations from the vicinity of Sarajevo. The material studied for this presentation belonged to the
population from Mt. Bjelasnica (locus typicus od bjelasnicensis); there fore this name is used as valid in
this presentation.
The illigeri Dejean, 1826 name probably can not be considered current where central Bosnian
populations are concerned because of insufficiently clearly defined locus typicus. On the basis of the
existing material no conclusions can be drawn, although the population from the vicinity of Karlovac
(village Bosanci) would be the most acceptable solution. It should be pointed out that the populations
from this region (Karlovac, Bosanci) differ from central Bosnian populations (=bjelasnicensis) in
endophallus morphology. The serious study of these populations could suggest that in this case we also
deal with a clearly differentiated species.
C. (Morphocarabus) bjelasnicensis ssp. tarensis Tosevski, 1990
Mt. Tara, W Serbia, 1250m., June1988, lgt. & coll.Tosevski.
7
C. (Morphocarabus) bjelasnicensis ssp. cerensis Tosevski, 1990
Mt. Cer, NW Serbia, 400m., June 1987, lgt. & coll.Tosevski,
C. (Morphocarabus) bjelasnicensis ssp. koshanini, Chiki, 1904
Mt. Cer, NW Serbia, 400m., July 1988, lgt. & coll.Tosevski,
Carabus (Morphocarabus) praecellens Palliardi, 1825
Distribution: The populations of this species are common in Srem (Serbia) and Slavonia (Croatia),
East to the Danube, while Southern and Western areal border is the Sava river.
Diversity: Phenotypically and by endophallus morphology populations from Mt. Fruska Gora
(Srem, North Serbia) and those from Mt. Papuk (Croatia, Slavonia, Pakrac) are practically identical.
Endophallus morphology of praecellens manifests phylogenetic similarity with Morphocarabus
populations of Hungary and Slovakia, although, phenotypically, they have the greatest similarity with
bjelasnicensis populations.
Carabus (Morphocarabus) praecellens Palliardi, 1825
North Serbia, Srem, Mt. Fruska Gora, June 1987, lgt. & coll. Tosevski
8
Carabus (Morphocarabus) kollari ssp. semetricus Kraatz, 1878 bona sp.
Distribution: East part of Serbian Banat (foothills of West Carpathian range). The areal border with nominal
ssp. kollari Palliardi, 1825 is not clear in the Romanian part of Banat.
Habitat: light groves with plenty of bushy vegetation, southern exposure.
Diversity: Phenotypically, semetricus populations and nominal ssp. kollari (loc. typicus Mehadia) are very
similar. The differences occur in specimen size, while the differences in elytrae sculpture although present are less
pronounced. However, the endophallus morphology differentiates between semetrica and kollari. On the contrary, in
the populations designated as comptus hampei and comptus incompsus there is a significant difference in elytrae
sculpture in regard to semetricus and kollari. However, phylogenetically, the representatives of comptus species (=
rothi according to Deuve, 1994), studied from the standpoint of endophalis morphology, by their type nevertheless
belong to kollari species.
Carabus (Morphocarabus) kollari ssp. semetrica Kraatz, 1878
Banat, Vrsac, Vrsacki Breg, June 1987, lgt. & coll. Tosevski
Carabus (Morphocarabus) kollari ssp. kollari Palliardi, 1825
Romania, Sasca Montana, June 1987, lgt. Budka, coll. Tosevski
Carabus (Morphocarabus) rothi ssp. hampei , Küster, 1846
Romania, Albesti, Sigbisoara, July 1986, lgt. A. Yual, coll. Tosevski.
9
Carabus (Morphocarabus) simulator Kraatz, 1876 bona sp.
Distribution: East Serbia, East from the Velika Morava river.
Diversity: From the morphological and phylogenetic standpoint simulator populations are
exceptionally interesting. Phenotypically, they are almost perfect copies of French monilis species
populations, especially of some its subspecies (e.g. ssp. monilis consitus). However, in the endophallus
morphology simulator is closest to the bjelasnicensis populations. Simulator populations are rather
homogeneous morphologically in their own geographical areal. The species is polychrome and with
relatively few variations in the size or sculpture of elytrae ( smaller, blue coloured specimens were
described as var. serbicus Hopffgarten, 1878, while red and green coloured specimens were described
as var. polychloros Beuthin, 1885). In colour variation simulator completely repeats colour variation
present in French monilis. Subspecies tekijensis stat. nov. described from the vicinity of Tekija, a typical
refugial habitat at the exit of Iron Gate, shows a rougher elytra sculpture and smaller size of specimens.
In this subspecies lateral lobes of distal lobe of endophallus are less pronounced than in nominal
simulator.
Remark: It is not very probable that the name of versicolor Frivaldszky, 1835 should be tied to
simulatror populations from East Serbia. Taking into acount that versicolor was described from vicinity of
Sliven (East Bulgaria) it is not probable that simulator and versicolor are so close to represent one taxon,
bearing in mind the speciation degree among other Balkans populations. In any case, verification through
endophallus analysis would solve the existing nomenclature problem of priority of versicolor or simulator
names.
Carabus (Morphocarabus) simulator Kraatz, 1876
East Serbia, Majdanpek, Rajkovo, June 1987, lgt&coll. Tosevski
Carabus (Morphocarabus) simulator ssp. tekijensis Tosevski, 1990
East Serbia, Iron Gate, Tekija, June 1987, lgt&coll. Tosevski
10
Carabus (Eucarabus) ullrichi ssp. ullrichi Germar, 1824 and their nations in Serbia
(monochromatic populations)
Carabus (Eucarabus) ullrichi ssp. ullrichi n. jaroslawi Fleisch, 1898
Serbia, Banat, Vrsac,Vrsacki breg, May 20, 1987, lgt. & coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. ullrichi n. nastasi Pavicevic, 1995
Serbia, Beograd, Topcider, 15.05.1987, lgt. & coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. ullrichi n. sokolari Born, 1904
N.Serbia, Srem, Fruska Gora, 15.07.1988, lgt. & coll. Tosevski
11
Carabus (Eucarabus) ullrichi ssp. ullrichi n. werneri Reitter 1899
NW Serbia, Korenita, Loznica, 20.05.1987, lgt. & coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. ullrichi n. gornjakensis Pavicevic & Tosevski, 1988
East Serbia, Gornjacka Klisura, Gornjak, 15.06.1987, lgt. & coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. ullrichi n. kucajensis Pavicevic & Tosevski, 1988
East Serbia, Mt. Kucaj, Babina Glava 500m, 15.06.1986 lgt. Ilic, coll. Tosevski
12
Carabus (Eucarabus) ullrichi ssp. fastuosus Palliardi, 1925 and their nations in
Serbia
(polychromatic populations)
Carabus (Eucarabus) ullrichi ssp. fastuosus n. arrogans Schaum, 1859
East Serbia, Majdanpek, June 15, 1987, lgt&coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. fastuosus n. pseudoarrogans Pavicevic & Tosevski, 1988
East Serbia, Miroc-Donji Milanovac, 15.06.1987, lgt & coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. fastuosus n. ceremosnjensis Pavicevic & Tosevski, 1988
East Serbia, Zdrelo-Krepoljin, 15.06.1987, lgt. & coll. Tosevski
13
Carabus (Eucarabus) ullrichi ssp. fastuosus n. transdiernae Pavicevic & Tosevski, 1988
East Serbia, Tekija, 15.06.1987, lgt. & coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. fastuosus n. serbomacedonicus Kenyery, 1970
South Serbia, Predejane, 15.06.1988, lgt. & coll. Tosevski
Carabus (Eucarabus) ullrichi ssp. fastuosus n. fabrizio Battoni & Breuning, 1970
East Serbia, Negotin, 15.06.1987, lgt. & coll. Tosevski
14
Carabus (Eucarabus) ullrichi Germar, 1824 - distribution in Serbia
Gornjacka Klisura ( Gornjak Gorge)
Site of the sharp demarcation of the populations ssp. ullrichi n. gornjakensis (left) and ssp.
fastuosus n. ceremosnjensis (right). Two populations are divided by the Mlava, a small
mountain river.
15
Titre : Descoperirea coleopterului Carabus Hampei Gutîiensis Takacs et Lie
Titre de la Revue : Acta Musei Porolissensis. [ Acta Mus. Porolissensis. ] , 2002 , vol. 24 , pp. 739 - 740
[ 2 pages. ]
ADALBERT Takacs
Type de document : PERIODIQUE
Cote INIST : 25082
Editeur :
Muzeul Judetean de istorie si arta
Zalau
Roumanie
Langue : roumain
Titre : O valoroasa specie de carabus în nordul tarii: Carabus Seriatissimus Reitter, 1896
Titre de la Revue : Acta Musei Porolissensis. [ Acta Mus. Porolissensis. ] , 2002 , vol. 24 , pp. 735 - 737
[ 3 pages. ]
ADALBERT Takacs
Type de document : PERIODIQUE
Cote INIST : 25082
Editeur :
Muzeul Judetean de istorie si arta
Zalau
Roumanie
Langue : roumain
Titre : Cercetarea biotipurilor unor specii endemice de coleoptere din genul "Carabus Linné" în partea
de nord a Transilvaniei
Titre de la Revue : Acta Musei Porolissensis. [ Acta Mus. Porolissensis. ] , 2002 , vol. 24 , pp. 731 - 734
[ 4 pages. ]
KINGA Bodola , ADALBERT Takacs
Type de document : PERIODIQUE
Cote INIST : 25082
Editeur :
Muzeul Judetean de istorie si arta
Zalau
Roumanie
Langue : roumain