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© RUSSIAN ENTOMOLOGICAL JOURNAL, 2006
Russian Entomol. J. 15(4): 401–407
Further investigation of Dorcadionini (Coleoptera: Cerambycidae)
endophallus with a revision of taxonomical position of the genus
Trichodorcadion Breuning, 1942
Äàëüíåéøåå èçó÷åíèå ñòðîåíèÿ ýíäîôàëëþñà Dorcadionini
(Coleoptera: Cerambycidae) ñ ïåðåñìîòðîì òàêñîíîìè÷åñêîãî
ïîëîæåíèÿ ðîäà Trichodorcadion Breuning, 1942
M.L. Danilevsky1 & D.G. Kasatkin2
Ì.Ë. Äàíèëåâñêèé1, Ä.Ã. Êàñàòêèí2
1 A.N. Severtzov Institute of Ecology and Evolution, Russian Academy of Sciences, Leninsky prospect 33, Moscow 119071,
Russia; e-mail: dorcadion@telecont.ru
2 Rostov branch of FGU “All Russian Centre of Plant Quarantine”, 20th line str. 43\16, Rostov-on-Don, 344037, Russia; e-mail:
kassatkind@mail.ru
1 Институт проблем экологии и эволюции РАН им. А.Н.Северцова, Ленинский проспект 33, Москва 119071, Россия.
2 Ростовский филиал ФГУ "Всероссийский центр карантина растений", ул. 20-я линия 43/16, Ростов-на-Дону 344037,
Россия
KEY VWORDS: Cerambycidae, Dorcadionini, Dorcadion, Trichodorcadion, endophallus, morphology, tax-
onomy, new synonyms.
КЛЮЧЕВЫЕ СЛОВА: Cerambycidae, Dorcadionini, Dorcadion, Trichodorcadion, эндофаллюс, морфология,
таксономия, новые синонимы.
though of general D. (Cribridorcadion) type. So D.
(Cribridorcadion Pic, 1901) = Dorcadion ( Bergerianum
Pesarini et Sabbadini, 2004), syn.n.
РЕЗЮМЕ: Строение эндофаллюса Tricho-
dorcadion gardneri Breuning, 1942 очень близко эн-
дофаллюсу Iberodorcadion Breuning, 1943 (а не
Eodorcadion Breuning, 1947, как можно было бы
ожидать из географических соображений), так как
центральный ствол слит с преапикальной камерой
без явной перетяжки, а преапикальная камера отде-
лена от апикальной камеры глубокой перетяжкой
(отсутствующей у Eodorcadion), внутренняя мемб-
рана в основании апикальной камеры хорошо раз-
вита (отсутствует у Eodorcadion), но хорошо отли-
чается очень крупными дистальными структурами.
Род Trichodorcadion Breuning, 1942 перенесён в
Dorcadionini. Описаны и изображены эндофаллюсы
5 видов, выделенных ранее в 2 подрода Dorcadion —
Megalodorcadion Pesarini et Sabbadini, 1999: D. (M.)
ledereri Thomson, 1865 ( типовой вид), D. (M.) walteri
Holzschuh, 1991 и Bergerianum Pesarini et Sabbadini,
2004: D. (B.) chrysochroum Breuning, 1943 (типовой
вид), D. (B.) hampei Mulsant et Rey, 1863, D. (B.) sonjae
Peks, 1993. D. (Megalodorcadion) признан естествен-
ной группой родственных видов, однако, строение
эндофаллюса не позволяет рассматривать её в ранге
подрода, так как вполне типично для D. (Cribri-
dorcadion Pic, 1901). Таким образом D. (Cribri-
ABSTRACT: Endophallus morphology of Tri-
chodorcadion gardneri Breuning, 1942 is very close to
Iberodorcadion Breuning, 1943 (but not to Eodorcadi-
on Breuning, 1947, as could be more expectable be-
cause of geographical reasons), because of central trunk
fused with preapical bulb without constriction, and
preapical bulb delimited from apical bulb by deep con-
striction (absent in Eodorcadion), internal membrane at
the base of apical bulb well developed (absent in Eodor-
cadion), but clearly differs because of very big distal
structures. Genus Trichodorcadion Breuning, 1942 is
transferred to Dorcadionini. Endophallic structures of 5
species being composed before in 2 subgenera of Dor-
cadion: Megalodorcadion Pesarini et Sabbadini, 1999:
D. (M.) ledereri Thomson, 1865 (type species), D. (M.)
walteri Holzschuh, 1991 and Bergerianum Pesarini et
Sabbadini, 2004: D. (B.) chrysochroum Breuning, 1943
(type species), D. (B.) hampei Mulsant et Rey, 1863, D.
(B.) sonjae Peks, 1993 — are described and figured. D.
(Megalodorcadion) is recognized as a natural group of
relative species, but endophallic morphology does not
allow to regard it as a subgenus, being very typical for
D. (Cribridorcadion Pic, 1901). So D. (Cribridorcadi-
on Pic, 1901) = D. (Megalodorcadion Pesarini et Sabba-
dini, 1999), syn.n. D. (Bergerianum) is totally artificial
complex consisting of three parts without any relative
connections. Each of three is more or less similar to
different groups of D. (Cribridorcadion) with the ex-
ception of D. crysochroum, which is more or less unique,
402 M.L. Danilevsky & D.G. Kasatkin
dorcadion Pic, 1901) = D. (Megalodorcadion Pesarini
et Sabbadini, 1999), syn.n. D. (Bergerianum) представ-
ляет собой искусственный комплекс не родственных
видов, состоящий из трёх очень разных групп. Стро-
ение эндофаллюса в каждой из них напоминает эн-
дофаллюс тех или иных групп внутри D. (Cribri-
dorcadion) за исключением D. crysochroum, эндо-
фаллюс которого несколько своеобразен, хотя впол-
не укладывается в план строения D. (Cribri-
dorcadion). Таким образом, D. (Cribridorcadion Pic,
1901) = Dorcadion (Bergerianum Pesarini et
Sabbadini, 2004), syn.n.
Introduction
In our previous publication on Dorcadionini en-
dophallus [Danilevsky et al., 2005] we supported the
separation of Dorcadion subgen. Megalodorcadion
Pesarini et Sabbadini, 1999 on the base of endophallic
structure of D. glabrofasciatum Daniel, 1901, which
was included in Megalodorcadion by C. Pesarini and
A. Sabbadini in the original description of the subge-
nus. The endophallus of the type species of Mega-
lodorcadion: D. ledereri Thomson, 1865 (original des-
ignation) was not investigated by us.
Recently Dorcadion subgenus Bergerianum Pe-
sarini and Sabbadini, 2004 (type species: D. chrysoch-
roum Breuning, 1943 — original designation) was de-
scribed. Originally it includes four species: D. chrysoch-
roum Breuning, 1943, D. hampei Mulsant et Rey, 1863,
D.sonjae Peks, 1993 and D. glabrofasciatum Daniel,
1901. Five species (D. ledereri Thomson, 1865, D. es-
cherichi Ganglbauer, 1897, D. angorense Ganglbauer,
1897, D. parallelum Kuster, 1847 and D. walteri Holzs-
chuh, 1991) are still included in D. (Megalodorcadion).
So, the endophallic structure of D. glabrofasciatum
can not be used for characterization of D. (Megalodorca-
dion). For better understanding of natural separation of
two subgenera we have studied their type species and
more representatives of each subgenus. Now we are able
to describe endophallus of D. ledereri and D. walteri, as
well as D. chrysochroum, D. hampei and D. sonjae.
We have also studied the endophallus of Trichodor-
cadion gardneri Breuning, 1942 (known from Ganges
Figs 1–2. Trichodorcadion gardneri: 1 — male, 2 — female.
Ðèñ. 1–2. Trichodorcadion gardneri: 1 — ñàìåö, 2 — ñàìêà.
1
2
403
Further investigation of Dorcadionini endophallus
Figs 3–4. Trichodorcadion gardneri: 3 — endophallus, lateral view; 4 — basal half of endophallus, ventral view.
Ðèñ. 3–4. Trichodorcadion gardneri: 3 — ýíäîôàëëþñ, ñáîêó; 4 — áàçàëüíàÿ ÷àñòü ýíäîôàëëþñà, ñíèçó.
Details: ab — apical bulb; ae — aedeagus; af — apical furrow; ap — apical phallomer,
bb — apical bubble; bd — central bend; bt — basal tube; cb — central bladder; ct — central
trunk; ds — dorsal swelling of central bladder; lt — lateral tubercles of central bladder;
mt — medial tube; pb — preapical bulb; vp — ventral plate; vs — ventral swelling of central
bladder.
Îáîçíà÷åíèÿ: ab — àïèêàëüíàÿ êàìåðà; ae — ýäåàãóñ; af — àïèêàëüíûé øîâ; ap —
àïèêàëüíûé ôàëëîìåð; bb — àïèêàëüíûé ïóçûðü; bd — öåíòðàëüíûé èçãèá; bt —
áàçàëüíàÿ òðóáêà; cb — öåíòðàëüíàÿ êàìåðà; ct — öåíòðàëüíûé ñòâîë; ds —
äîðñàëüíûé âûñòóï öåíòðàëüíîé êàìåðû; lt — áîêîâûå âûñòóïû öåíòðàëüíîé êàìåðû;
mt — ìåäèàëüíàÿ òðóáêà; pb — ïðåàïèêàëüíàÿ êàìåðà; vp — âåíòðàëüíûé ñêëåðèò;
vs — âåíòðàëüíûé âûñòóï öåíòðàëüíîé êàìåðû.
4
3
river valley in India and Nepal) in order to understand
the taxonomic position of the genus Trichodorcadion
Breuning, 1942.
Descriptions
Trichodorcadion gardneri Breuning, 1942
(Figs 1–4)
MATERIAL. #, $: Nepal, Narayani prov., Sauraha SW
Chitwan Nat. Park, 180m, 27°34'80''N, 84°29'49''E, 17.4.2000,
leg. A.Weigel (coll. of M.Danilevsky).
The species was described (on the base of one specimen —
11mm long — most probably male, because of long antennae)
as the type-species of a new genus Trichodorcadion from
India (Uttar Pradesh prov., Dehra Dun). The genus was
placed by its author [Breuning, 1961] in Morimopsini and
never figured.
Due to the curtsey of Mr. Andreas Weigel we have
received a pair of T. gardneri from Nepal (Figs 1–2); length
of male: 11.9mm (including slightly exposed abdomen),
length of female: — 14.8mm. The identification of the
species was made on the base of original description. The
holotype must be preserved in the Museum of Dehra Dun.
A female of the species has just the same elytral design as
male, but relatively wider, anterior tarsi are not narrower
than in male.
Unfortunately the sample of everted and inflated en-
dophallus (Figs 3–4) of our single male was broken during
preparation, but still all main characters can be seen.
af
lt
cb
vp
bt
ae
bb
ab
pb+ct
mt
lt
vp
bt
ae
mt
bd
404 M.L. Danilevsky & D.G. Kasatkin
DESCRIPTION. Endophallus relatively long, much long-
er than elytra (but shorter than elytra and prothorax com-
bined). Basal tube (bt) relatively short, more than two times
shorter than aedeagus, straight, microspicules indistinct,
without transverse rugae, a little swollen distally; ventral
plates (vp) relatively wide and strongly sclerotized. Medial
tube (mt) about as long as aedeagus, straight, microspicules
indistinct, distally swollen forming relatively large central
bladder (cb), with well developed lateral tubercles of central
bladder (lt). Central bend (bd) distinct; central trunk (ct)
turned to the central bladder at the angle of about 30°, long,
wide, relatively cylindrical, not swollen, without tubercles,
covered with dense hardly visible microtrichiae only distal-
ly. Preapical bulb (pb) large, strongly swollen connected
with central trunk without distinct constriction, dorsally,
ventrally and distally with very distinct microspicules; bas-
al and lateral areas of preapical bulb covered with very
small, hardly visible microtrichiae. Apical bulb (ab) large,
spherical, joined to preapical bulb by distinct constriction,
covered with distinct regular spines. Apical furrow (af)
with a well developed internal membrane inside. Apical
bubble (bb) also large hemispherical, without sclerites; long
flagellum seems to be developed, but it is still inside bubble
in our sample. Gonopores situated near apex of apical
bubble.
REMARK. In general endophallus belongs to Iberodor-
cadion Breuning, 1943 type because of preapical bulb con-
nected with central trunk without distinct constriction, inter-
nal membrane present. Same connection is in Neodorcadion
Ganglbauer, 1883 and in Eodorcadion Breuning, 1947, but in
Neodorcadion preapical bulb is not differentiated and in
Eodorcadion apical bulb absent; in Neodorcadion apical mem-
brane present, but in Eodorcadion absent.
Dorcadion (Cribridorcadion) ledereri Thomson, 1865
(Fig. 5)
MATERIAL. #: “TR, vill. Corum, Çerum 10km E, 2–
3.V.1992, lg. Pesarini & Sabbadini” (coll. of M.Danilevsky).
DESCRIPTION. Endophallus relatively long, about as
long as elytra. Basal tube (bt) relatively short, about two
times shorter than aedeagus, straight, without microspi-
cules, transversely rugose; ventral plates (vp) relatively
small and poorly sclerotized. Medial tube (mt) much longer
than aedeagus, straight, covered with very small microspi-
cules dorsally and ventrally, distally strongly swollen form-
ing big and longitudinal central bladder (cb), with dorsal (ds)
and ventral (vs) swellings. Central bend (bd) distinct;. cen-
tral trunk (ct) turned to the central bladder at the angle of
about 45°, long, narrow, gradually tapering distally, not
swollen, without tubercles, covered with very dense rela-
tively long brown microtrichiae. Preapical bulb (pb) small,
but wider than distal portion of central trunk, clearly delim-
ited from central trunk by strong constriction, spherical,
glabrous in main portion, but distally with dense brown
microtrichiae similar to those of central trunk. Apical bulb
(ab) joined to preapical bulb by distinct constriction, cov-
ered with distinct regular spines. Apical bubble (bb) with-
out distinct sclerites.
REMARK. In general endophallus is not special. Such
morphology type (with narrow, tapering posteriorly cen-
tral trunk and small elongated distal structures) is known in
several groups of D. (Cribridorcadion), which do not look
to be relatives, for example in D. (C.) apicerufum Breuning,
1943, D. (C.) lugubre Kraatz, 1873 and others [Danilevsky
et al., 2005: Figs 99, 106].
D. (Cr.) walteri Holzschuh, 1991
(Fig. 6)
MATERIAL. # (paratype): “TR (Vill. Bolu), Alput Bey Köyu,
bei Bolu, 800m, 16.4.1990, Heinz leg.” (coll. of M.Danilevsky).
DESCRIPTION. General structure very similar to the pre-
ceding species, but central bladder (cb) hardly pronounced — in
form of gradual dilatation of medial tube (mt), with long and
narrow ventral swelling (vs). Central bend (cb) distinct, cen-
tral trunk (ct) turned to the central bladder at the angle of
about 45°, long, narrow, gradually tapering distally, not swol-
len, without tubercles, covered with very dense relatively
long brown microtrichiae. Preapical bulb (pb) small, narrow,
Fig. 5. Endophallus of Dorcadion (Cr.) ledereri, lateral view.
Explanation see on Figs 3–4.
Ðèñ. 5. Ýíäîôàëëþñ Dorcadion (Cr.) ledereri, ñáîêó.
Îáîçíà÷åíèÿ ñì. íà ðèñ. 3–4.
af
cb
vp
bt
ae
bb
ab
mt
pb
ct
ds
vs
bd
405
Further investigation of Dorcadionini endophallus
elongated, not wider than distal portion of central trunk,
without constriction in between, glabrous in main portion,
but distally with dense brown microtrichiae similar to those
of central trunk. Apical bulb (ab) joined to preapical bulb by
distinct constriction, covered with distinct regular spines.
Apical bubble (bb) without distinct sclerites.
REMARK. According to the endophallus morphology,
D. walteri is rather close to D. ledereri. The presence of
ventral tubercle of central bladder is rather common in such
type of endophallus. Long ventral tubercles are known in
D. (C.) bistriatum Pic, 1898, D. (C.) seminudum Kraatz,
1873, D. (C.) tauricum Waltl, 1838 and others [Danilevsky
et al., 2005: Figs 97, 98, 101].
D. (Cr.) chrysochroum Breuning, 1943
(Fig. 7)
MATERIAL. #: “GRAECIA, Peloponesos b., Ciokos Mts.,
900–1000m, 2km NE Petsaki, 20.5.2004, J.Voøíšek leg.”
(coll. of M.Danilevsky).
DESCRIPTION. Endophallus relatively long, longer than
elytra. Basal tube (bt) relatively short, about two times
shorter than aedeagus, straight, without microspicules, ven-
tral plates (vp) relatively small and poorly sclerotized. Me-
dial tube (mt) much longer than aedeagus, transversely rug-
ose, straight, covered with very small microspicules dorsally
and ventrally, distally strongly swollen forming big and lon-
gitudinal central bladder (cb), with big ventral swelling (vs).
Central bend (bd) distinct; central trunk (ct) turned to the
central bladder at the angle of about 60°, narrow, but relative-
ly short, about two times longer than wide, cylindrical, not
swollen, without tubercles, covered with very dense, short
microtrichiae. Preapical bulb (pb) big, just a little shorter than
central trunk (unique character!), swollen, elongated, much
wider than distal portion of central trunk, totally covered
with short microtrichiae, which become denser distally. Api-
cal bulb (ab) also big, joined to preapical bulb by distinct
constriction, covered with distinct regular spines, forming
together with hemispherical apical bubble (bb) a large spher-
ical apical phallomer (ap), which is bigger than preapical
bulb; apical bubble without distinct sclerites.
REMARK. In general endophallus is rather special be-
cause of big apical phallomer and short central trunk. Similar
apical phallomers are known in several Dorcadion (Cribri-
dorcadion) groups, which are very far from D. chrysoch-
roum, for example in D. equestre (Laxmann, 1770) or in “D.
semenovi-group” [Danilevsky et al., 2005: Figs 71–76], but
here central trunk longer and more or less swollen, while
preapical bulb strongly transverse. Endophallus of D. chryso-
chroum is not similar the endophallus of any Dorcadion
species, though it is in general of D. (Cribridorcadion) type.
D. (Cr.) hampei aureovittatum Kraatz, 1873
(Fig. 8)
MATERIAL. #: “TR — w. Niksar, w. YOLKONAK, 5.5.1996,
S.Kadlec leg.” (coll. of M.Danilevsky).
DESCRIPTION. Endophallus relatively short, much short-
er than elytra. Basal tube (bt) short, about four times shorter
than aedeagus, straight, with hardly visible microspicules;
ventral plates (vp) distinct and strongly sclerotized. Medial
tube (mt) a little shorter than aedeagus, straight, covered with
very small microspicules dorsally and ventrally, distally a little
dilated, forming hardly pronounced central bladder (cb). Cen-
tral bend (bd) distinct;. central trunk (ct) turned to the median
tube at the angle of about 90°, very wide, but strongly reduced,
short, two times shorter than wide, rather swollen dorsally,
densely covered with distinct microasperities. Preapical bulb
(pb) big, spherical, about as wide as swollen central trunk,
clearly delimited from central trunk by strong constriction,
glabrous basally, in main portion with dense moderately small
microspiculae. Apical bulb (ab) joined to preapical bulb by
distinct constriction, covered with distinct regular spines, cy-
lindrical, tapering distally; internal membrane well developed.
Apical bubble (bb) wide consists of dorsal and ventral por-
tions, without distinct sclerites.
REMARK. General endophallus structure is rather ex-
ceptional, because of reduced, short, transverse central trunk
Fig. 6. Endophallus of Dorcadion (Cr.) walteri, lateral view.
Explanation see on Figs 3–4.
Ðèñ. 6. Ýíäîôàëëþñ of Dorcadion (Cr.) walteri, ñáîêó.
Îáîçíà÷åíèÿ ñì. íà ðèñ. 3–4.
af
vp
bt
ae
bb
ab
mt
pb
ct
vs
bd
406 M.L. Danilevsky & D.G. Kasatkin
swollen dorsally. Such form was known before only in D. (C.)
sturmi Frivaldsky, 1837 (Danilevsky et al., 2005: Fig. 108),
but its close affinity to D. hampei is very doubtful.
D. (Cr.) sonjae Peks, 1993
(Fig. 9)
MATERIAL. # (paratype): “TR (Elazið), Ayvar-Köyü b.,
Elmapinan 850m (23km e Elazið), 15.IV.1992, Heinz leg.”
(coll. of M.Danilevsky).
DESCRIPTION. General structure very similar to the
preceding species, but central bladder (cb) is strongly devel-
oped with very large dorsal (ds) and ventral (vs) swellings;
central trunk (ct) even wider being swollen dorsally and
ventrally.
REMARK. The close natural affinities between D. hampei
and D. sonjae are rather evident from the external characters
of the species. These connections are proved by endophallic
structures.
7
Fig. 7–9. Endophallus of Dorcadion, lateral view: 7 — D. (Cr.) chrysochroum; 8 — D. (Cr.) hampei; 9 — D. (Cr.) sonjae.
Explanation see on Figs 3–4.
Ðèñ. 7–9. Ýíäîôàëëþñ of Dorcadion, ñáîêó: 7 — D. (Cr.) chrysochroum; 8 — D. (Cr.) hampei; 9 — D. (Cr.) sonjae. Îáîçíà÷åíèÿ
ñì. íà ðèñ. 3–4.
8 9
af
vp
bt
bb
ab
mt
pb
ct
ds
vs
af
cb
vp
bt
ae
bb
ab
mt
pb
ct
vs
af
ab
bb
pb
ct
cb
cb
mt
vp
bt
ae
ae
bd bd bd
ab
407
Further investigation of Dorcadionini endophallus
Conclusion
The morphological similarity between Iberodorca-
dion and Trichodorcadion in the general structures of
distal portions seems to be the evidence of mutual
ancient ancestor, younger than morphological type of
Cribridorcadion, because several small features of Tri-
chodorcadion are also same as in Iberodorcadion: the
absence of microtrichiae on the most part of central
trunk, presence of large lateral tubercles of central blad-
der (known only in certain Iberodorcadion), presence
of long flagellum (also known only in Iberodorcadion).
So, the natural position of the genus Trichodorcadion
is inside Dorcadionini. The genus differs from all Ibero-
dorcadion by the spherical shape of preapical bulb (in
Iberodorcadion it is more or less cylindrical) and spher-
ical structure forming by joined apical bulb and apical
bubble (impossible in Iberodorcadion but known in
certain Cribridorcadion).
Imagoes externally is normal Dorcadionini without
wings, with wide membrane between clypeus and labrum
(typical for Eodorcadion), elytra covered with recum-
bent pubescence and numerous stout erect setae.
D. (Megalodorcadion) is a natural group of species,
that it rather clear from the external imaginal morphology.
This point of view can be supported by endophallic
morphology, as endophallus of D. (M.) ledereri is very
close to endophallus of D. (M.) walteri. From the other
side endophallic morphology can not be the base for
separation of this natural group in a subgenus, as it does
not differ from the general D. (Cribridorcadion) morpho-
logical type, so D. (Cribridorcadion Pic, 1901) =
D. (Megalodorcadion Pesarini et Sabbadini, 1999), syn.n.
D. (Bergerianum) is a totally artificial complex con-
sisting of three rather different groups without relative
connections. Exceptional long endophallus of D. gla-
brofasciatum with “S”-shaped central bladder and “S”-
shaped connection between central trunk and preapical
bulb was described by us before (Danilevsky et al.,
2005: Fig. 109). Endophallus of D. chrysochroum (type
species) with its big apical phallomer and short central
trunk is not directly connected with any known Dorca-
dion, but can be easily placed inside D. (Cribridorcadi-
on) morphological type. D. hampei and D. sonjae repre-
sent natural group which has no connections with two
previous species because of reduced, short, transverse
central trunk. Bergerianum was described on the base
of similar shape modification of apical spines of hind
male tibiae. But in reality the longest spines of hind male
tibiae are not dilated in any of species included in the
subgenus. In all four species these spines are not modi-
fied, as it was pictured by the authors of the subgenus.
External morphology of all four species also does not
allow to suppose their relative affinities. So D. (Cribri-
dorcadion Pic, 1901) = Dorcadion (Bergerianum Pe-
sarini et Sabbadini, 2004), syn.n.
ACKNOWLEDGEMENTS. We are very grateful to
Stanislav Kadlec, Walter Heinz, Heinz Peks, Carlo Pe-
sarini, Andrea Sabbadini, Ji i Vo íšek and Andreas
Weigel for providing us with several specimens for
study. Our special thanks to Kirill Makarov for his
invaluable help with arrangement of photographs.
References
Danilevsky M.L., Kasatkin D.G. & Rubenyan A.A. 2005.
Revision of the taxonomic structure of the tribe Dorcadi-
onini (Coleoptera, Cerambycidae) on the base of endophal-
lic morphology // Russian Entomol. J. Vol.13(2004). No.3.
P.127–149.
Breuning S. 1942. Novae species Cerambycedarum XI // Fol.
zool. hydrob. Vol.11. P.113–175.
Breuning S. 1961. Catalogue des Lamiaires du Monde. 5.
Lieferung. Im Verlag des Museums G. Frey. Munchen.
S.287–382.
Pesarini C. & Sabbadini A. 1999. Osservazioni sistematiche su
alcuni Dorcadion della fauna anatolica, con descrizione di
9 nuovi taxa (Coleoptera, Cerambycidae) // Ann. Mus. civ.
St. nat. Ferrara. Vol.1.(1998). P.45–61.
Pesarini C. & Sabbadini A. 2004. Ricerche sui Dorcadiini di
Grecia. I. Le specie del Peloponneso (Coleoptera Ceramby-
cidae) // Atti della Societa Italiana di Scienze Naturali e
del Museo Civico di Storia Naturale in Milano. Vol.145.
No.1. P.133–153.
