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INTRODUCTION
Until recently, the representatives of the genus
Zophohelops Reitter, 1901 were known only from the
mountain systems of the Northern and Western Tien-
Shan (Reitter 1922, Medvedev 1987). In the course of
examination of the material on Helopini it was found,
that one species of this genus, Z. humeridens (Reitter,
1901), is distributed in Armenia. The problem is that the
generic placement of this species was not hitherto
revealed. This species was originally described by Retter
(1901) as Xanthomus humeridens. Subsequently it was
assigned to the subgenus Omaleis Allard, 1877 of the
genus Cylindronotus Faldermann, 1837 (Reitter 1922).
It is noteworthy, that the characteristics of this species
were given by Reitter not in the key, but in the footnote.
Probably, Reitters doubts about the generic placement
of this species were caused by the large geographical
gap between the distribution of the species from Tien-
Shan, on the one hand, and Armenia, on the other. How-
ever, the problematic species from Armenia undoubtedly
belongs to the genus Zophohelops. Such assignment is
supported by the structure of male genitalia, epipleura
and pronotum. A material from south-western Tadzhik-
istan was studied. As a result a new Zophohelops Z.
michajlovi sp. nov. was discovered. Thus, a disjunct
range of the genus Zophohelops includes Tien-Shan,
Hissaro-Darvaz, and south of Armenia plateau. It seems
possible, that in the future this genus will be found also
in northern Iran. There are also other examples of widen
disjunct generic ranges in the family Tenebrionidae in
the Palearctic region. In particular, the genus Seidlitzel-
lus Reitter, 1919 (tribe Apolitini). includes two species,
S. tristis (Faldermann, 1837) and S. turcestanicus Reit-
ter, 1919. The former species is distributed in Transcau-
casus and Balkan Peninsula, whereas the latter was
described from Tien-Shan.
ACKNOWLEDGEMENTS AND DEPOSITORIES
OF THE MATERIAL EXAMINED
This study is based on the examination of material
from the following institutions:
HNHM Hungarian Natural History Museum, Buda-
pest, Hungary (O. Merkl);
IZA Institute of Zoology of Armenia, Erevan,
Armenia (M. Kalashian);
KES Kharkov Entomological Society, Kharkov,
Ukraine (V. Michajlov);
MLLK Museum of local lore of Kharkov, Kharkov,
Ukraine (A. Drogvalenko);
ZIN Zoological Institute, Russian Academy of
Sciences, St.-Petersburg, Russia;
ZMRSU Zoological Museum of the Rostov State
University, Rostov-on-Don, Russia.
TAXONOMIC NOTES ON THE GENUS ZZOOPPHHOOHHEELLOOPPSS
REITTER, 1901 WITH DESCRIPTION OF A NEW SPECIES
FROM TADZHIKISTAN AND A NEW GENUS
PPSSEEUUDDOOPPRROOBBAATTIICCUUSSGEN. NOV.
(COLEOPTERA: TENEBRIONIDAE: HELOPINI)
MAXIM V. N ABOZHENKO
Rostov State University, Biological Faculty, Department of Zoology; 344711, Bolshaya
Sadovaya street, 105, Rostov -on-Don, Russia, e-mail: nalassus@mail.ru
Abstract. A new species Zophohelops michajlovi sp. nov. is described from Tadzhikistan.
Generic position of Zophohelops humeridens (Rtt.), comb. nov. is specified. Genus Zophohelops
has found to be distributed not only in Tien-Shan, but also in Hissaro-Darvaz and Southern Arme-
nia. A new genus Pseudoprobaticus gen. nov. (type species Helops granipennis Allard, 1876) is
described. Relationships and position of the new genus in tribe Helopini is discussed.
Ë
Key words. Coleoptera, Tenebrionidae, Helopini, Zophohelops, new species, Tadzhikistan,
Pseudoprobaticus, new genus, taxonomy, distribution.
ANNALES ZOOLOGICI (Warszawa), 2001, 51(4): 511-515
I thank the curators and their institutions mentioned
above for the loan of the material. I am grateful to G. S.
Medvedev, B. A. Korotyaev and A. Yu. Solodovnikov for
critically and carefully reviewing this paper and making
useful comments.
ZZoopphhoohheellooppss hhuummeerriiddeennss (Reitter, 1901) comb. nov.
(Figs 14)
Xanthomus humeridens Reitter, 1901: 223;
Cylindronotus (Omaleis)humeridens: Reitter, 1922: 139.
TTyyppeess (examined)..Holotype (male) and paratypes
(1 male and 2 females) with the labels: Caucasus. Arax-
esthal. Leder. Reitter, coll. Reitter, Xanthomus
humeridens m. 1900 (HNHM).
OOtthheerr mmaatteerriiaall eexxaammiinneedd.. Armenia: Vaykskiy Distr.,
Gogi Mts. (3000 m), 1 VII 1935, 1 female (IZA); Kafanskiy
Distr., Atkiz (2500 m), 6 VI 1954 (Akramovskiy), 1 male
(IZA); Megrinskiy Range, 25 V 1987, 2500 m (G. Davidi-
an), 1 male (ZIN); Vaykskiy Distr., Dzhermuk, 6 VI 1988
(M. Savitskiy), 1 female (ZIN); Megrinskiy Pass, 6 VI
1993 (M. Kalashian), 1 male (ZMRSU).
ZZoopphhoohheellooppss mmiicchhaajjlloovvii sp. nov.
(Figs 58)
DDiiaaggnnoossiiss. Similar to Zophohelops humeridens
Reitter, 1901 but differs in the narrow and long antennal
segments, longitudinal segments of the fore tarsus, not
confluent punctures of striae and the structure of aedea-
gus and abdominal sternite VIII.
DDeessccrriippttiioonn. Body pale
fuscous with strong sheen,
legs and antennae paler.
Anterior margin of clypeus
straight. Head widest at the
level of eyes. Eyes small, con-
vex; ratio of head width in its
widest part to distance
between eyes 1.52. Lateral
margin of head at junction of
gena and clypeus without
emargination; genae very
weakly rounded. Head with-
out transverse depression at
junction of clypeus and frons.
Punctation of head very fine
and scarce (distance between
punctures 34 times diameter
of puncture). Antennae long,
extending behind base of
pronotum by their three last
segments; width to length
ratio in antennal segments
2
nd
11
th
, respectively, as fol-
lows: 1, 2.5, 1.6, 1.6, 2, 2, 2, 1.4,
1.4, 1.7. Pronotum weakly
transverse (1.18 times as
wide as long), widest in the
middle; anterior margin
weakly widely emarginate;
base straight; lateral margins
rounded in the middle, but
straight behind and before
the middle; anterior and pos-
terior angles obtuse, rounded;
lateral margins and base nar-
rowly bordered; border of
anterior margin widely inter-
rupted in the middle. Disc reg-
ularly convex, not flattened at
512 M. V. NABOZHENKO
Figures 18. Zophohelops Reitter. 14. Zophohelops humeridens. 58. Zophohelops michajlovi sp. nov.
(1, 5) Aedeagus, ventral view; (2, 6) aedeagus, lateral view; (3, 7) abdominal sternit VIII; (4, 8) antenna.
1 mm
6
7
5
8
4
3
21
sides. Pronotal punctation fine and scarce, distances
between punctures 35 times diameter of puncture. Pro-
pleura smooth, weakly rugose near coxae. Apterous. Ely-
tra oblong-oval, strongly convex; elytral intervals slight-
ly convex, moderately coarsely punctate (two, rarely
three punctures across interval). Lateral reflexed mar-
gin of elytra very narrow, visible only at base (viewed
from above). Visible abdominal sternites smooth, finely
and scarcely punctate. Fore tibia straight, strongly and
gradually widened to apex, its outer margin thorn-
shaped, oblong on the apex; middle and hind tibiae
straight; fore tarsus weakly widened, tarsomeres elon-
gated; middle and hind tarsi not widened, with dense
golden pubescence ventrally.
Aedeagus short and wide; parameres 1.2 times as
long as fallobase; lateral margin of parameres weakly
broadly emarginate in apical quarter; aedeagal sclerites
wide, fused in the middle.
Body length 4.8 mm; body maximal width 2.7 mm.
EEttyymmoollooggyy. Named for V. A. Michajlov (KES), collec-
tor of the new species.
TTyyppee mmaatteerraall.. Holotype, male: Tadzhikistan, Pyandzh-
skiy Karatau Mts., 5 VI 1959 (V. A. Michajlov) (ZIN).
PPsseeuuddoopprroobbaattiiccuuss gen. nov.
(Figs 911)
TTyyppee ssppeecciieess..Helops granipennis
Allard, 1876.
GGeennddeerr.. Masculine.
DDiiaaggnnoossiiss. The new genus is most
closely related to Zophohelops and
Xanthomus (structure of male genitalia
and structure of epipleura), from which
it differs in the sculpture of elytra, pro-
pleura, epipleura, pro- and mesosterna
(with dense fine tuberculation) and very
complicated structure of the spermathe-
ca. From the representatives of a genus
Probaticus (including subgenus Pelori-
nus), where originally joined by Reitter
(1922) the new genus differs by a struc-
ture of aedeagus (it concern to nalassus
type, instead of to helops type) not
extended fore and middle tarsus of
male, absence of rigid erected setae on
abdominal sternite, and also tuberculed
by a sculpture of cuticle.
DDeessccrriippttiioonn. Body black, dull, ely-
tra fuscous. Head punctation very
coarse and dense; eyes strongly con-
vex; head without eye furrows. Pronotum transverse,
widest in the middle; sides strongly rounded. Propleura
coarsely and densely tuberculated; prosternum with
coarse and dense tubercles; prosternal process weakly
convex. Elytra convex, with deep irregular striae; elytral
intervals flattened, with very coarse and dense tubercles;
outer margin of epipleura (lateral inflexed margins of ely-
tra) wide, reaching elytral apex where they are connect-
ed with the 1st interval; epipleura not reaching elytral
apex, densely tuberculated. Ventrites coarsely and dense-
TAXONOMIC NOTES ON THE GENUS ZOPHOHELOPS REITTER, 1901 513
1 mm
b
a
c
d
f
910
11
Figures 911. Pseudoprobaticus granipennis. (9) Aedeagus, ventral view; (10) aedeagus,
lateral view; (11) female reproductive system (a canal of spermatheca; b short branches;
c 1
st
reser voir; d 2
nd
reservoir; f gland of spermatheca).
ly punctate, with regular contiguous pubescence; last
ventrite without coarse erect setae, only with contiguous
fine pubescence. Tibiae straight; fore and middle tarsi
not widened, their segments narrow and longitudinal.
Aedeagus weakly sclerotized, wide, keel-oblonged at the
apex; parameres arcuate in lateral view (Figs 910).
Female reproductive system (Fig. 11). Spermatheca
complicated, with two reservoirs, one of them situated at
base, the other, in the middle; basal quarter of sper-
matheca with short lateral branches. Basal duct of sper-
matheca short; duct of spermatecal gland entering in the
place of connection of basal duct of spermatheca with
apical part of vagina.
MMaatteerriiaall eexxaammiinneedd..1 male (MLLK) and 1 female
(ZIN) of Helops granipennis from Turkey
REMARKS
In this part, the following taxa are discussed: Adel-
phinus Fairmaire, 1866; Catomus Allard, 1876; Cylin-
dronotus Faldermann, 1837: C. bellator (Reitter, 1901);
C. batesi Allard, 1876; C. femoratus (Faldermann, 1837);
Ectomopsis Antoine, 1949; Hedyphanes Fisher de
Waldheim, 1822; Nalassus Mulsant, 1854; Caucasonotus
Nabozhenko, 2000; Helopocerodes Reitter, 1922; Nalas-
sus faldermanni (Faldermann, 1837); Helopondrus
Reitter, 1922; Odocnemis Allard, 1876; Heloponotus
Reitter, 1922; Reitterohelops Skopin, 1960; Stenomax
Allard, 1876: Stenomax aeneus (Scopoli, 1763); Tur-
cmenohelops Medvedev, 1987; Xanthomus Mulsant,
1854; Zophohelops Reitter, 1902.
Before elucidating the position of the new genus in
the classification of the tribe, we must briefly outline
some basic morphological features of the darkling bee-
tles of the tribe Helopini.
Evolutionary transformations within the tribe are
those characteristic of the family Tenebrionidae in gen-
eral. It is known, that ancestral forms of Tenebrionidae
were dendrobionts and the family evolved towards adap-
tations to extreme conditions of the open landscapes.
Therefore, the main adaptations within the tribe Helopi-
ni are those to endure deficiency of moisture. First, this
is the structure of elytra, especially their epipleura. The
following character states of elytra seems to be derived:
epipleura reaching the apex of elytra; outer margin of
epipleura (lateral deflexed margin of elytra) reaching
apex of elytra, where it is connected with first interval;
elytral intervals flat; striae of elytra very fine or absent
(in some species of Hedyphanes, Catomus, Reitteroh-
elops, which occur in very dry habitats); junction of ely-
tra very tight; cuticle strongly sclerotized.
The structure of male genitalia is also taxonomically
very important. Their morphological transformations
proceeded in the direction of ensuring reliability of the
copulation. Presumably, as regards the structure of male
genitalia, members of the subtribe Helopina demon-
strate the most derived character state: strongly sclero-
tized parameres, flattened at the apex, without traces of
the middle suture and with the stiff short setae, directed
backwards (or, in some species of Catomus and Adel-
phinus, with dense rasp-shaped puncturation), probably
necessary for fixing parameres inside the vagina, when
penis moving out (Iablokoff-Khnzorian 1964); apical part
of the parameres often curved upwards. In some species
of the subtribe Cylindronotina (Stenomax aeneus) para-
meres are completely curved upwards thus leaving the
apex of penis exposed. In the subtribe Cylindronotina
the structure of parameres is more diverse. In the Cylin-
dronotus group of genera (Cylindronotus, Odocnemis,
Stenomax and Reitterohelops) the parameres are
strongly sclerotized and elongated and, usually, straight
or slightly curved upwards; in apical part, flattened
dorso-ventrally. The structure of genitalia in the Nalas-
sus group of genera (Nalassus, Zophohelops, Xantho-
mus, Ectomopsis, Turcmenohelops and Pseudopro-
baticus) appears more primitive: aedeagus weakly scle-
rotized, parameres short or moderately elongated (Tur-
cmenohelops, many species of Nalassus), oblonged in
apical part in the laterally flattened keel, often with the
noticed track on the apex. Some species of Zophohelops
and also Pseudoprobaticus with rather curved inwards
parameres. Such type of the parameres probably is
ancestral in the subtribe Cylindronotina.
Gastral spicula usually consisting of two lateral rod-
shaped sclerites, common trunk and apical blades, is also
taxonomically important. Strong S-shaped arcuation of
the sclerites of gastral spicula (best visible laterally) may
be considered as derived character state, owing to which
the area of contact of the blades with aedeagus is
increased (reliability of the fixing of aedeagus is ones
turn provided by this) (Medvedev 2001). Significant
degree of fusion of the rod-shaped sclerites and formation
of the common trunk is also a derived character state
(Medvedev 2001). The most derived structure of the gas-
tral spicula in the subtribe Cylindronotina is character-
istic of the species of Cylindronotus [(C. bellator,
C. batesi, C. femoratus and Odocnemis (only subgenus
Heloponotus)], which possess thick rods of the gastral
spicula, strongly S-shaped arcuated and fused on the
apex in short (rarely in long as in C. batesi) common
trunk. The genera Odocnemis and Stenomax are char-
acterized by long common trunk (correlating with the
length of the fallobase) and slight S-shaped arcuation of
the gastral spicula. Species of genera Zophohelops and
Ectomopsis have, in most cases, widely placed and
weakly arcuate branches of the gastral spicula with mod-
erately long common trunk; sclerites of the common
trunk are fused only at the very apex. In the subtribe
Helopina the structure of the gastral spicula is more uni-
form. Its rods are usually thick, long, weakly S-shaped
and fused in long common trunk.
514 M. V. NABOZHENKO
Structure of the female reproductive system in the
subtribe Cylindronotina is rather uniform. Species of the
Nalassus genus group have simple spermatheca without
lateral branches (subgenera Caucasonotus, Helopoce-
rodes and the nominotypical subgenus Nalassus).
Sometimes spermatheca with short tubercles along its
entire length Nalassus (Helopocerodes)falderman-
ni. Species of the Nalassus (Helopondrus) possess
spermatheca with short lateral branches at base. In
species of the Cylindronotus genus group spermatheca
always with short (Cylindronotus, subgenus Helopono-
tus of Odocnemis) or long (many species of Odocnemis)
lateral branches. Spermathecal gland in the Cylin-
dronotus group usually long (length of gland 34 times
length of the beetle itself). The structure of spermatheca
in the subtribe Helopina is more variable and often with
branches and reservoirs.
Judging from the morphology, Pseudoprobaticus
appears a specialized branch within Cylindronotina,
adapted to the semiarid mountainous landscapes of Asia
Minor. Pseudoprobaticus granipennis demonstrates
the most derived character states: strongly sclerotized
cuticle; epipleura (lateral inflexed margin of elytra) with
outer margin reaching the apex of elytra; propleura, epi-
pleura, pro- and mesosterna and elytra with dense fine
tuberculation; very complicated structure of spermathe-
ca. At the some time this species has primitive structure
of aedeagus, resembling that of the genera Zophohelops,
Xanthomus and Ectomopsis. This feature supports
close relationship of the genus Pseudoprobaticus with
the mentioned genera.
REFERENCES
Medvedev, G. S. 1987. [Tenebrionid beetles of the genus Zophoh-
elops Rtt. And allied genera (Coleoptera, Tenebrionidae) from
middle Asia and Kazakhstan]. Trudy Zoologicheskogo Institu-
ta,164: 95129 (in Russian).
Medvedev, G. S. 2001. [Evolution and system of darkling beetles of the
tribe Blaptini (Coleoptera, Tenebrionidae)]. Meetings in memory
of N. A. Cholodkovsky, 53. St.-Petersburg. 332 pp. (in Russian).
Allard, E. 1876. Revision des Helopides vrais de Lacordaire.
lAbeille, 14: 180.
Iablokoff-Khnzorian, S. M. 1964. Bemmerkungen über einige Rei-
itersche Typen aus dem Ungarischen Naturwissenschaftlichen
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293315.
Reitter, E. 1901. Verschiedenes über die Coleopteren der Tenebri-
oniden-Abtheilung Helopinae. Deutsche Entomologische
Zeitschrift, Heft 2: 209224.
Reitter, E. 1922. Bestimmungstabelle der europaischen Coleopteren.
Heft 92. Tenebrionidae. 17. Teil: Unterfamilie Helopina, II. Wien
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TAXONOMIC NOTES ON THE GENUS ZOPHOHELOPS REITTER, 1901 515
Received: July 27, 2001
Accepted: November 10, 2001
