Евразиатский энтомол. журнал 9(4): 631–640
© EUROASIAN ENTOMOLOGICAL JOURNAL, 2010
Genital structures of some East Palaearctic species of the subtribe Staphylinina Latreille, 1802 (Coleoptera, Staphylinidae) Ãåíèòàëüíûå ñòðóêòóðû íåêîòîðûõ âîñòî÷íîïàëåàðêòè÷åñêèõ âèäîâ ïîäòðèáû Staphylinina Latreille, 1802 (Coleoptera, Staphylinidae) E.A. Khachikov*, A.V. Shavrin** Ý.À. Õà÷èêîâ*, À.Â. Øàâðèí** * Rostov Branch of Russian Entomological Society, Alexandrovsky spusk 59, Rostov-on-Don 344030 Russia. * Ростовское отделение Русского энтомологического общества, Александровский спуск 59, Ростов-на-Дону 344030 Россия. ** Institute of Systematic Biology, Daugavpils University, Vienibas 13, Daugavpils LV-5401 Latvia. E-mail:
[email protected]. ** Институт биологической систематики, Даугавпилсский университет, Виенибас 13, Даугавпилс LV-5401 Латвия.
Key words: Staphylinidae, Staphylininae, Staphylinina, Hadropinus, Dinothenarus, Liusus, Platydracus, genitalia, endophallus, terminalia. Ключевые слова: Staphylinidae, Staphylininae, Staphylinina, Hadropinus, Dinothenarus, Liusus, Platydracus, гениталии, эндофаллус, терминалии. Abstract. The morphology of genitalia and terminalia is described for five East Palaearctic species of the subtribe Staphylinina Latreille, 1802: Platydracus dauricus (Mannerheim, 1830), P. sp. pr. mortuorum (Bernhauer, 1912), Dinothenarus chrysocomus (Mannerheim, 1830), Hadropinus fossor Sharp, 1889 and Liusus hilleri (Weise, 1877). Резюме. Описывается морфология структур гениталий и терминалий пяти восточнопалеарктических видов подтрибы Staphylinina Latreille, 1802: Platydracus dauricus (Mannerheim, 1830), P. sp. pr. mortuorum (Bernhauer, 1912), Dinothenarus chrysocomus (Mannerheim, 1830), Hadropinus fossor Sharp, 1889 и Liusus hilleri (Weise, 1877).
Introduction The paper presents preliminary results of the first author’s study of the genital structures of species from the subtribe Staphylinina [Khachikov, 2005, 2006]. It deals with the detailed descriptions of genitalia and terminalia of Platydracus dauricus (Mannerheim, 1830), P. sp. pr. mortuorum (Bernhauer, 1912), Dinothenarus chrysocomus (Mannerheim, 1830), Hadropinus fossor Sharp, 1889 and Liusus hilleri (Weise, 1877), all distributed in East Palaearctic region. Many structures of the internal sac of the aedeagus are described and illustrated for the first time, and their new nomenclature is proposed.
Material, methods and terminology The material from the collection of the second author as well as the material obtained from colleagues (see the Acknowledgements) was used in the present
study. The specimens are deposited in the private collections of both authors (CS — private collection of A. Shavrin, Daugavpils; CK — private collection of E. Khachikov, Rostov-on-Don) and the dissected genital structures are deposited in the collection of the first author (CK). Dissected genital structures were stained by fuchsine due to obtain a better visibility of transparent tissues. For permanent mount, the structures were placed on transparent glass and imbedded into transparent nail polish (dry preparation) or placed into closed small containers with 30 % solution of glycerin and water (wet preparation). The preparation methods protocol and most anatomical terms are adopted from various papers [Blackwelder, 1936; Coiffait, 1953; Kasatkin, 2006; Khachikov, 2005, 2006; Khachikov, Popov, 2006; Shvanvich, 1949]. The new terms, and definitions proposed in the paper are listed below. Phallomer, a general term, used for topological designation of various parts of endophallus. Some parts of phallomer are specifically named on the basis of their permanent presence in the internal sac of the aedeagus, such as dorsal titillator-brush. Operculum is a separate sclerite, located in posterior dorsal part of penis. The penis tube, which is not entirely closed along full length of dorsal surface and bearing operculum, is named semiclosed. Titillator is a spine in a form of an elongated hollowed structure, consisting of two or more tittilatorvirgae; distal piece of ejaculatory duct is situated inside the tittilator. Agonoporial triada consists of three titillators permanently present in apical part of endophallus, the
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Table 1. List of terms Òàáëèöà 1. Ñïèñîê òåðìèíîâ Term
Abbreviation
Usage*
Synonyms and references (if present)
Operculum
op
original
"opercule du penis" [Coiffait, 1953]; "dorsal plate" [Tikhomirova, 1968]
Agonoporial triada
-
original
"agonoporial sclerites" [Khachikov, 2005]; "agonoporial titillators" [Khachikov, 2006]
-
-
used
basal-ventral
tbv
original
titillator-brush
tb
used
"needle sclerite" [Khachikov, Popov, 2006; Khachikov, 2006]
titillator-lamina
tl
used
[Khachikov, 2006]
bititillator
bt
original
"agonoporial sclerite" [Khachikov, 2005]
monotitillator
mt
original
"apical sclerite" [Khachikov, 2005]
titillator-spine
tsp
original
"virga" [sensu Khachikov, 2006; non Shvanvich, 1949]; flagellum [Blackwelder, 1936]
Virga
vi
used
[Shvanvich, 1949]
Apical spiculae
sa
used
[sensu Kasatkin, 2006]
-
-
used
[sensu Kasatkin, 2006; Khachikov, 2006; Khachikov, Popov, 2006]
lateral
sl
original
-
p
used
Titillator
Swelling
Projection
dorsal
pd
original
basal-dorsal
pbd
original
basal-ventral
pbv
original
-
pr
used
lateral
prl
original
dorsal-lateral
prdl
original
dorsal
prd
original
-
-
generally accepted
medial-lateral
chml
original
medial-ventral
chmv
original
basal-ventral
chbv
original
-
-
used
proximal
php
original
medial
phm
used
distal
phd
original
Appendix of endophallus
ap
used
Ductus ejaculatorius
de
generally accepted
Apodema
apd
used
Prominence
Chamber
Phallomer
Secondary agonopor
ag
generally accepted
-
vag
generally accepted
provagina
pvag
original
metavagina
mvag
original
vaginolamina
vl
original
sex gland
sg
original
spermatheca
sp
original
enter to spermatheca
es
original
proctovaginus
prv
original
proctiger
prc
original
Contour topological scheme
sc
original
Vagina
[Shvanvich, 1949; Khachikov, 2006] "basal tittillator" [Khachikov, 2006]
[Khachikov, 2006]
-
[Khachikov, 2006]
-
[sensu Shvanvich, 1949; Kasatkin, 2006] "basal phallomer" [Kasatkin, 2006] [Kasatkin, 2006] "apical phallomer" [Kasatkin, 2006] [sensu Kasatkin, 2006] [sensu Shvanvich, 1949]
-
"ventral spicula" [Khachikov, 2005, 2006]
-
"proctiger" [Khachikov, 2005, 2006] -
* «Usage» means limited usage of the term in some papers in the field of morphology and anatomy of beetle genitalia. Accepted abbreviations are used in the text and in figures here. * «Óïîòðåáëåíèå» îçíà÷àåò îãðàíè÷åííîå èñïîëüçîâàíèå òåðìèíà â íåêîòîðûõ ïóáëèêàöèÿõ ïî ìîðôîëîãèè è àíàòîìèè ãåíèòàëèé æåñòêîêðûëûõ. Ïðèíÿòûå ñîêðàùåíèÿ èñïîëüçóþòñÿ çäåñü, à òàêæå â òåêñòå è ðèñóíêàõ.
Genital structures of some East Palaearctic species of Staphylinina asymmetrical one is named monotitillator, and the other two are named bititillators. These structures are listed as agonoporial sclerites or titillators in the existing literature. Appendix of endophallus is an interior enlarged appendix made of membranous tissues of different thickness, and it enters into the secondary agonopore. Metavagina is a distal part of vagina (provagina). Proctovaginus is a sclerite situated in a place where vagina and proctiger are joint together; it is named as «proctiger» in the previous papers [Khachikov, 2005, 2006]. Vaginolamina is a sclerite, covering entrance to the vagina, it was previously named as «ventral spicula» [Khachikov, 2005, 2006]. Apodema of secondary agonopore or agonoporial apodema — is a ring-like sclerite, surrounding secondary agonopor. The reasons for proposing these new terms are in more details the complex morphology of genitalia of some genera of the subtribe Staphylinina that will be broadly discussed further. All specific terms used in present paper with their abbreviations, synonyms, etymology and references are presented in the Table 1 below. Under etymology, we explain initial and basic term, from which other morphological terms are derived. Names of those basic terms are in bold.
Results Platydracus Thomson, 1858 Subgenus Platydracus Thomson, 1858 Type species Staphylinus stercorarius Olivier, 1795.
Platydracus dauricus (Mannerheim, 1830) Figs 1, 2.
Material. Irkutskaya oblast, Olkhonskiy raion: without
data, A.S. Pleshanov — 1# (CS); Khuzhir, 20–24.08.2005, A.V. Shavrin — 1$ (CS). Republic of Buryatia, Barguzinskiy nature reserve: Bolshoe Barguzinskoe lake, 08.1997, V.G. Shilenkov — 1$ (CS); Dzhidinskiy raion: Verkhniy Dyrestuy, 6– 9.07.1997, A.V. Shavrin — 1$ (CS). Zabaikalskiy krai: valley of Telenik River, Udunga, 20.07.1992, Kalmykov — 3##; 1$ (CK); Nerchinskiy raion: Pravie Kumaki, Katorginskaya pad’, 3.08.1999, O. Korsun — 2$$ (CK).
Male. Penis tubular of semiclosed type. Operculum enlarged, thickened apically. Endophallus short, weakly curved dorso-ventrally. Titillator-brush (tb) extended almost along the entire length of endophallus, and becoming denser in anterior part. Proximal phallomer (php) consists of the dorso-basal part of dorsal titillator-brush (tb) and the divided baso-ventral titillator (tbv). Distal phallomer (phd) consists of dorso-apical part of dorsal tittilator-brush (tb) and, ventrally of, agonoporial triada consisting of partly sclerotized enlarged monotitillator (mt) and bititillator (bt) forming of two weakly sclerotized swellings. Apex of ejaculatory duct (de) forms on its perimeter an agonoporial apodema (apd), surrounding secondary agonopore in form of broken ring. In this place short membranouse appendix is opening into secondary agonopore (ap). Female. Vagina (vag) large, occupying almost all internal space of genital sclerite, latero-ventrally attached to coxites by coxovaginal mesenteries. In basal part of vagina,
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wide vaginal entrance (ev), immediately under anus, is located. Ventral edge of vaginal entrance is slightly concaved. Vagina consists of wider provagina (pvag) and narrow metavagina (mvag). Provagina is situated immediately under proctiger in dorsal view; both being joined. Surface of provagina is covered by comb-shaped microtrichia. Sex gland (sg) is spherical, laterally attached to vagina by a thin connecting canal, close to vaginal entrance. Oviduct is suited ventrally, with denser area in place of its connection with vagina. Distinctly bordered proctovaginus (prv) is placed inside vagina, its surface covered by microlumps, without microtrichia. Vaginolamina absent.
Platydracus sp. pr. mortuorum (Bernhauer, 1912) Figs 3, 4.
Material. Maritime province: Spasskiy raion: 19 km from Vasil’kovka, 15.07.1989, Napolov — 3##; 1$ (CK); Khasanskiy raion: Narvinskiy mountain path, 6.08.1990, Napolov — 2$$ (CK); Bezverkhovskiy mountain path, 25.07.1990, Napolov — 1$ (CK); Lazovskiy raion: Lazo, 20.07.2007, V. Shokhrin — 1# (CS); idem, 1–15.06.2000, L. Sundukova — 1# (CS); Glazkovka, V. Shokhrin, 8–20.08.07 — 1# (CS); Lazovskiy nature reserve, Ezhovaya bay, 15–17.07.2002, Yu. Sundukov — 1# (CS). Remarks. It is possible, that the apical part of endophallus with secondary agonopore was not fully everted, and therefore in reality it may be longer than on present illustrations. Male. Penis tubular of semiclosed type. Operculum enlarged, narrower in basal and medial parts, thickened apically, widely rounded on apical margin. Endophallus short, asymmetrical. Right side of endophallus (in dorsal position) is different than its left side, which has enlarged lateral prominence (prl), dorsal-lateral prominence (prdl) and agonoporial triada. Agonoporial triada consists of triangular monotitillator (mt) and bititillator (bt) and includes two sclerites in form of irregular pyramid. Dorsal side of endophallus includes large, cone-like dorsal projection (pd), lateral swelling (sl), two dorsal asymmetric prominences (right (prd right), left (prd left)), and operculum (op). Female. Vagina (vag) large, attached to coxites by coxovaginal mesenteries latero-ventrally. In basal part of vagina, immediately under proctiger, a wide vaginal entrance (ev) is located. Ventral edge of vaginal entrance slightly concaved. Vagina consists of wider provagina (pvag) and narrow metavagina (mvag). Sex gland (sg) of medium size, oval. Proctovaginus (prv) weakly expressed, without clear borders.
Dinothenarus Thomson, 1858 Type species: Staphylinus pubescens DeGeer, 1774.
Subgenus Parabemus Reitter, 1909 Type species: Staphylinus fossor Scopoli, 1771.
Dinothenarus chrysocomus (Mannerheim, 1830) Figs 5, 6.
Material. Zabaikalskiy krai: Uletovskij raion: Tanga, Shantiyskoe lake, 6.08.07, I. Enustschenko — 1#; 1$ (CS); Chita, 7.06.1992, O. Korsun — 1$ (CK). Maritime province: Spassk-Dal’niy raion: 4 km SE Evseevka, 6–10.08.99, A. Shavrin — 1# (CS); Siniy Mts., 8 km E Evseevka, 7–9.08.1999, A. Shavrin — 2$$ (CS); Barabash-Levada, 7.1991, M. Shestopalov — 2$$ (CK); Sukhanovskiy mountain path, 17.08.1990, Napolov — 1# (CK). Remarks. Endophallus is only partially straight. Therefore description of structures are given for their separate
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fragments. It is also possible that some membranous structures were not revealed. To show the entire picture of the endophallus drawing/scheme (sc) is provided. Male. Penis tubular of semiclosed type. Paramerae with group of stump-like setae apically. Endophallus of medium size, weakly curved in dorso-ventral direction, and topologically divided into proximal phallomer (php), situated between basis of endophallus and dorsal titillator-brush (tb). The endophallus has two symmetrical baso-ventral chambers (chbv), two baso-dorsal projections (pbd) and dorsal titillator-lamina (tl). Medial phallomer (phm) situated between basis of titillator-brush (tb) and agonoporial triada, and also has a titillator-brush and large medial-ventral chamber (chmv). Distal phallomer (phd) includes agonoporial triada consisting of mono- (mt) and bititillator (bt). Monotitillator of triangular beak-like shape, bititillator consisting of two asymmetric flat sclerites, meeting distally in form letter «V»; they are surrounded by secondary agonopore
(ag) and by agonoporial apodema (ap), where appendix of endophallus (ap) is also opened. Female. Vagina wide. Spermatheca (sp) of medium size, and wide. Sex gland (sg) of medium size, oval. Vaginolamina (vl) narrow, curved, covering entrance to spermatheca (es). Proctovaginus (prv) weakly expressed, without clear borders, covered by hair-like microtrichia of different length, denser in distally.
Hadropinus Sharp, 1889 Type species Hadropinus fossor Sharp, 1889.
Hadropinus fossor Sharp, 1889 Figs 7, 8.
Material. Maritime province: Lazovskiy nature reserve, kordon Prosyolochniy (Ta-Chingouza), 1–3.07.2007, A. Shavrin — 8##; 3$$ (CS). Kunashir Island: mouth of Avdeevka river, 14.07.2008, K. Makarov — 1#; 1$ (CK).
Figs 1–2. Platydracus dauricus, genital structures: 1 — male, endophallus; 2 — female. Abbreviations as in the text. Ðèñ. 1–2. Platydracus dauricus, ãåíèòàëüíûå ñòðóêòóðû: 1 — ñàìåö, ýíäîôàëëóñ; 2 — ñàìêà. Ñîêðàùåíèÿ â òåêñòå.
Genital structures of some East Palaearctic species of Staphylinina Male. Penis tubular of semiclosed type. Operculum (op) consists of two sclerites. Endophallus weakly curved in dorso-ventrally, covered ventrally by sparse and short microtrichia. Endophallus includes baso-ventral titillator (tbv), two lateral prominences (prl), two baso-ventral projections (pbv), two medio-lateral chambers (chml), dorsal titillatorbrush (tb), dorsal projection (pd) and agonoporial triada.
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Double tendon, which begins between sclerites of operculum, is situated within endophallus, and extended almost along its entire length. Endophallus includes three phallomers. Proximal phallomer (php) is extended from the basis of endophallus to dorsal titillator-brush (tb), including basalventral titillator (tbv), wide at base and narrower to apex; two baso-ventral projections (pbv), narrow in apical part,
Figs 3–4. Platydracus sp. pr. mortuorum, genital structures: 1 — male, endophallus; 2 — female. Abbreviations see in the text. Ðèñ. 3–4. Platydracus sp. pr. mortuorum, ãåíèòàëüíûå ñòðóêòóðû: 1 — ñàìåö, ýíäîôàëëóñ; 2 — ñàìêà. Ñîêðàùåíèÿ â òåêñòå.
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Figs 5–6. Dinothenarus chrysocomus, genital structures: 1 — male, endophallus; 2 — female. Abbreviations see in the text. Ðèñ. 5–6. Dinothenarus chrysocomus, ãåíèòàëüíûå ñòðóêòóðû: 1 — ñàìåö, ýíäîôàëëóñ; 2 — ñàìêà. Ñîêðàùåíèÿ â òåêñòå.
Genital structures of some East Palaearctic species of Staphylinina
Figs 7–8. Hadropinus fossor, genital structures. 1 — male, endophallus; 2 — female. Abbreviations see in the text. Ðèñ. 7–8. Hadropinus fossor, ãåíèòàëüíûå ñòðóêòóðû: 1 — ñàìåö, ýíäîôàëëóñ; 2 — ñàìêà. Ñîêðàùåíèÿ â òåêñòå.
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covered by microthorns; and two lateral prominences (prl). Medial phallomer includes two small medio-lateral chambers (chml) and dorsal hook-like curved titillator-brush (tb), situated on membranose dorsal projection (pd). Distal phallomer (phd) forms secondary agonopore and agonoporial triada and consists of flat monotitillator (mt) with pointed apex, and bititillator (bt). The latter consists of two enlarged virgae (vi), with rasp-like surface. All of them form titillator-spine (tsp), bearing in its apical membranous part two small, symmetric apical spiculae (sa) and a secondary agonopore (ag). Appendix of endophallus (ap) of enlarged shape, with thick walls, attached to the base of bititillator. It has interior cavity, which is extended along its entire length. Interior cavity is opened to ejaculatory duct (de) in basal part. Apex of appendix covered by semispherical sclerite with uneven surface. Functionally, the appendix together with titillator-spine form a structure, which is analogous to syringe, where the role of plunger is played by walls of appendix, causing ejection of sperm through canal of titillator-spine into sexual organs of female when contracting. Female. Vagina (vag) wide, oval in apical part. Proctovaginus (prv) large, hardly sclerotized, with middle line wider distally, smooth, without microtrichia, wider than proctiger (prc). Spermatheca (sp) in proximal part wide, and distally thread-like. Sex gland (sg) of medium size, enlarged, pear-shaped. Vaginolamina absent.
Liusus Sharp, 1889 Type species Hadrotes hilleri Weise 1877.
Liusus hilleri (Weise, 1877) Figs 9, 10.
Material. Maritime province: Lazovskiy nature reserve,
Ezhovaya bay, 15–17.07.2002, Yu. Sundukov — 3##; 2$$ (CK, CS).
Male. Penis tubular of closed type. Operculum consists of two oval sclerites. Endophallus short, enlarged, includes baso-ventral titillator (tbv), dorsal titillator-brush (tb) and agonoporial triada. Ejaculatory duct (de) thin, stretching along the entire length of endophallus. Endophallus includes three phallomeres. Proximal phallomer (php) extended from basis of endophallus to dorsal titillator-brush (tb) and has basal-ventral titillator (tbv), wider at base and narrower at apex. Medial phallomer (phm) includes dorsal titillator-brush (tb), situated inside membranose dorsal projection (pd). Distal phallomer (phd) includes secondary agonopore and agonoporial triada, consisting of monotitillator (mt) and bititillator (bt). Monotitillator concaved basally, wide at base and, apically narrowing to a sharp apex; it has laterally two keels, joining together at apex. Bititillator consists of two enlarged virgae (vi), concaved on basal ends and forming together titillator-spine (tsp). Secondary agonopore (ag) and two apical spiculae (sa) are located on the apex of bititillator. Agonoporial appendix enlarged, with thick walls, hollow inside, attached to base of titillatorspine (tsp). Apex of agonoporial appendix with semispherical sclerite of uneven surface. Appendix joined with ejaculatory duct in basal part. Functionally, appendix together with titillator-spine form a structure, analogous to syringe. Walls of appendix act as plunger. Sperm is ejected through ejaculatory duct into sexual organs of female. This interpretation is supported by the fact that the same preparation of appendix (exposed to water or glycerine) was causing some changes in it length (Fig. 9: ap. 1, ap. 2). It shows the ability of contracting and extending the walls of appendix.
Female. Vagina (vag) wide, proctovaginus (prv) weakly expressed, without visible borders, and consists of two symmetric parts. Vaginolamina (vl) narrow, strongly concaved, sickle-shaped. Spermatheca (sp) thick in proximal part, thread-like distally, wider at the end. Sex gland (sg) of middle size, oval.
Discussion Despite of the fact that our studies were restricted only to five species, we can make some preliminary conclusions on the basis of the examined structure of genitalia. For example, Liusus hilleri, Hadropinus fossor and Creophilus maxillosus Linnaeus, 1758 [see Khachikov, 2006] form one group based on the similarity of their endophallic structures. Thee share the presence of the titillator-spine consisting of bititillator, with the continuation of ejaculatory duct inside; and the titillator-brushes are situated almost in horizontal position along longitudinal axis of endophallus. These species have the appendix with thickened stretchy walls and with semispherical sclerite apically. In general, these similarities support an earlier hypothesis [Khachikov, 2006] about close proximity of these taxa which may be grouped together into a supra-generic taxon (e.g. Creophilinae Kirby, 1837) rank. However, this decision could be made after more comprehensive phylogenetic study is undertaken taking into accounts many more representatives of the subfamily Staphylininae from the world. The morphology of genitalia presented in this paper applies to the other species of the respective genera. For example, Platydracus dauricus is closely related to the Western Palaearctic P. chalcocephalus Fabricius, 1801 [Khachikov, 2006] and has the same type of genitalia. These species share similar structure of endophallus, such as agonoporial triada and basalventral dividing titillator. However, the Eastern Palaearctic Platydracus sp. pr. mortuorum and widespread P. stercorarius Olivier, 1795 probably belong to a separate group within the genus because the structure of their endophallus is different from those of the other studied Palaearctic species with the asymmetrical structure and with the presence of a large dorsal projection in distal part. The structure of agonoporial triada of Dinothenarus chrysocomus is typical for that genus, and for the group of genera, where Dinothenarus belongs.
Acknowledgements We thank the following colleagues for making specimens available for study: K. Makarov (Moscow, Russia), I. Enustshenko (Irkutsk, Russia), O. Korsun (Chita, Russia), Yu. Sundukov (Lazo, Russia), V. Shilenkov (Irkutsk, Russia), A. Pleshanov (Irkutsk, Russia). We are grateful to A. Solodovnikov (Copenhagen, Denmark) and J. Klimaszewski (Quebec, Canada) for help with English and their valuable comments. The research of the second author was conducted within the framework of the project of European Social Fund (No. 2009/0206/1DP/1.1.1.2.0/09/APIA/VIAA/010).
Genital structures of some East Palaearctic species of Staphylinina
Figs 9–10. Liusus hilleri, genital structures: 1 — male, endophallus; 2 — female. Abbreviations see in the text. Ðèñ. 9–10. Liusus hilleri, ãåíèòàëüíûå ñòðóêòóðû: 1 — ñàìåö, ýíäîôàëëóñ; 2 — ñàìêà. Ñîêðàùåíèÿ â òåêñòå.
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References Blackwelder R.E. 1936. Morphology of the coleopterous family Staphylinidae // Smithsonian Miscellaneous Collections. Vol.94. No.13. P.1–102. Coiffait H. 1953. Les Lathrobium (sensu lato) de France et des region voisines (Coleoptera, Staphylinidae) // Annales de la Société Entomologicue de France. T.122. P.85–111. Kasatkin D.G. 2006. [The internal sac of aedeagus of longhorned beetles (Coleoptera: Cerambycidae): morphology, nomenclature of structures, taxonomic significance] // Caucasian Entomological Bulletin. T.2. Vol.1. P.83–104. [In Russian]. Khachikov E.A. 2005. [Internal structures of genitals of some species of genus Ocypus Leach, 1819 and Tasgius Stephens, 1829 (Coleoptera: Staphylinidae) with description of new taxa
from the genus Ocypus] // Caucasian Entomological Bulletin. T.1. Vol.1. P.19–32. [In Russian]. Khachikov E.A. 2006. [New data on the morphology and taxonomy of some species of the subfamily Staphylininae (Coleoptera: Staphylinidae)] // Caucasian Entomological Bulletin. T.2. Vol.2. P.177–182. [In Russian]. Khachikov E.A., Popov D.S. 2006. [New data on morphology and taxonomy of some species of the genus Nicrophorus Fabricius, 1775 (Coleoptera: Silphidae)] // Caucasian Entomological Bulletin. T.2. Vol.1. P.27–40. [In Russian]. Tikhomirova A.L. 1968. [New and little-known Paederinae (Coleoptera, Staphylinidae) of the fauna of USSR] // Sbornik trudov Zoologicheskogo muzeya MGU. T.11. C.195–204. [In Russian]. Shvanvich B.N. 1949. [The course of the general entomology]. Moscow – Leningrad: Sovetskaya nauka. P.1–900. [In Russian].
Поступила в редакцию 9.10.2009
