A current generic classification of sap beetles (Coleoptera, Nitidulidae)

2008·0.11 MB·

by  A. G. Kirejtshuk

© Zoological Institute, St. Petersburg, 2008
A current generic classifi cation of sap beetles
(Coleoptera, Nitidulidae)
A.G. Kirejtshuk
Kirejtshuk, A.G. 2008. A current generic classifi cation of sap beetles (Coleoptera, Nitidu-
lidae). Zoosystematica Rossica, 17(1): 107-122.
The list of generic and subgeneric taxa arranged into subfamilies and tribes proposed for
sap beetles is given, which includes also new genera [Parapocadius gen. nov. (Nitidulinae:
Nitidulini) and Interfaxia gen. nov. (Nitidulinae: Cyllodini)] and subgenera [Semocarpolus
subgen. nov. Gaplocarpolus subgen. nov. Askocarpolus subgen. nov. (Carpophilinae: Car-
pophilus Stephens, 1829)]. The Meoncerus Sharp, 1891; Apsectochilus Reitter, 1874 and
Lordyrops Reitter, 1875 are considered as quite distinct each from other above mentioned
as from all other generic taxa. In the list there are given the complete synonymy, including
new synonymy of generic and subgeneric names [Crepuraea Kirejtshuk, 1990 and Nyujwa
Perkovsky, 1990 syn. nov.; Haptoncus Murray, 1864 and Haptoncurina Jelínek, 1977, syn.
nov.; Ecnomaeus Erichson, 1843 and Somaphorus Murray, 1864, syn. nov.; Ecnomorphus
Motschulsky, 1858; Tribrachys LeConte, 1861 syn. nov.; Stauroglossicus Murray, 1864, syn.
nov. and Microxanthus Murray, 1864, syn. nov.; Pria Stephens, 1829 and Allopria Kirejtshuk,
1980, syn. nov.; Megauchenia Macleay, 1825 and Orvoenia Dajoz, 1980, syn. nov.; Tetrisus
Murray, 1864 and Pseudoischaena Grouvelle, 1897, syn. nov.; Neopocadius Grouvelle, 1906
and Pseudostelidota Grouvelle, 1906, syn. nov.; Cychramus Kugelann, 1794 and Aethinopsis
Grouvelle, 1908, syn. nov.; Mystrops Erichson, 1843, and Cryptoraea Retter, 1873, syn. nov.;
Cyllodes Erichson, 1843 and Mecyllodes Sharp, 1891, syn. nov.; Grammorus Murray, 1868
and Colopteroides Watrous, 1982, syn. nov.; Cryptarcha Shuckard, 1839 and Priatelus Broun,
1881, syn. nov.]. For some taxa the rank is changed, namely, Lordyra Gemminger & Harold,
1868, stat. nov. is regarded as a subgenus of Lasiodactylus Perty, 1830-1834; Brounthina
Kirejtshuk, 1997, stat. nov. as a subgenus Neopocadius Grouvelle, 1906 and Teichostethus
Sharp, 1891, stat. nov. as a subgenus of Hebascus Erichson, 1843, while the taxa Coxollodes
Kirejtshuk, 1987 (stat. nov.) and Onicotis Murray, 1864 (stat. nov.) are regarded as separate
genera. The new taxa and new taxonomical proposals are supplied with corresponding data
in the notes below the list. In these notes there are also proposed the new synonymy for the
following species names: Carpophilus (Ecnomorphus) acutangulus Reitter, 1884 and C. (E.)
cingulatus Reitter, 1884, syn. nov.; C. (E.) bakeweli Murray, 1864; C. (E.) planatus Murray,
1864, syn. nov. and C. (E.) aterrimus Macleay, 1864, syn. nov.; C. (E.) debilis Grouvelle,
1897 and C. (E.) opaculus Grouvelle, 1897, syn. nov.; C. (E.) luridipennis Macleay, 1873
and C. (E.) loriai Grouvelle, 1906, syn. nov.; C. (E.) murrayi Grouvelle, 1892 and C. (E.)
hebetatus Grouvelle, 1908, syn. nov.; C. (E.) plagiatipennis (Motschulsky, 1858) and C.
(E.) nigricans Grouvelle, 1897, syn. nov.; C. (E.) terminalis Murray, 1864 and C. (E.)
gentilis Murray, 1864, syn. nov.; Lasiodactylus brunneus Perty, 1830; L. centralis Cline et
Carlton, 2004, syn. nov.; L. falini Cline et Carlton, 2004, syn. nov. and L. kelleri Cline et
Carlton, 2004, syn. nov.); Pallodes opacus Grouvelle, 1906 and P. loriai Grouvelle, 1906,
syn. nov.; Cyllodes fauveli Grouvelle, 1903 and Pallodes vagepunctus Grouvelle, 1903, syn.
nov.; Pallodes jucundus Reitter, 1873 and Mecyllodes nigropictus Sharp, 1891, syn. nov.;
Pallodes birmanicus Grouvelle, 1892 and P. kalingus Kirejtshuk, 1987, syn. nov.; P. gestroi
Grouvelle, 1906 and P. misellus Grouvelle, 1906, syn. nov.; P. rufi collis Reitter, 1873 and
P. cyanescens Grouvelle, 1898, syn. nov.; Grammophorus caelatus Gerstäcker, 1864 and
Colopterus striaticollis Murray, 1864, syn. nov. In connection with a preliminary revision
of many type series of the family and the mentioned taxonomical changes for some species
names are established new taxonomical interpretation, namely: Pleoronia nitida (Grouvelle,
1898), comb. nov. (Axyra : Axyrodes); Parapocadius immerizi (Grouvelle, 1899), comb. nov.
(Pallodes); Camptodes rufi cornis (Grouvelle, 1898), comb. nov. (Pallodes); Neopallodes
aestimabilis (Grouvelle, 1906), comb. nov. (Pallodes); N. alluaudi (Grouvelle, 1899), comb.
nov. (Pallodes); N. aterrimus (Grouvelle, 1906), comb. nov. (Pallodes); N. dorsalis (Grou-
velle, 1896), comb. nov. (Pallodes); N. fairmairei (Grouvelle, 1906), comb. nov. (Pallodes);
N. incertus (Grouvelle, 1906), comb. nov. (Pallodes); N. klugi (Grouvelle, 1896), comb. nov.
(Pallodes); N. limbicollis (Reitter, 1880), comb. nov. (Pallodes); N. militaris (Grouvelle,
1906), comb. nov. (Pallodes); N. niger (Grouvelle, 1906), comb. nov. (Pallodes); N. nigro-
cyaneus (Grouvelle, 1906), comb. nov. (Pallodes); N. nitidus (Grouvelle, 1906), comb. nov.
(Pallodes); N. orthogonus (Grouvelle, 1906), comb. nov. (Pallodes); N. perrieri (Grouvelle,
1906), comb. nov. (Pallodes); N. scutellaris (Grouvelle, 1906), comb. nov. (Pallodes); N.
108 A.G. Kirejtshuk: Generic classifi cation of sap beetles • ZOOSYST. ROSSICA Vol. 17
sicardi (Grouvelle, 1906), comb. nov. (Pallodes); N. sikordi (Grouvelle, 1896), comb.
nov. (Pallodes); N. variabilis (Grouvelle, 1896) , comb. nov. (Pallodes); Coxollodes
cyrtusoides (Reitter, 1884), comb. nov. (Pallodes); C. amamiensis (Hisamatsu, 1956),
comb. nov. (Pallodes); Coxollodes opacus (Grouvelle, 1906), comb. nov. (Pallodes); C.
parvulus (Grouvelle, 1908), comb. nov. (Pallodes); C. reitteri Kirejtshuk, 1987, comb.
nov. (Pallodes); Pallodes fauveli (Grouvelle, 1903), comb. nov. (Cyllodes); Cyllodes
jucundus (Reitter, 1873), comb. nov. (Pallodes); Interfaxia fasciata (Sharp, 1891), comb.
nov.; Onicotis auritus Murray, 1864 comb. nov.; Platyarcha biguttata (Motschulsky,
1858), comb. nov. (Carphophilus: Ecnomorphus); Cryptarcha optanda (Broun, 1881),
comb. nov. (Priateles). Besides, because of the new interpretation for Pallodes laetus
Grouvelle, 1898, which should be transfered to the genus Camptodes, C. grouvellei nom.
nov. (non Camptodes laetus Kirsch, 1873) is proposed. For the generic names Perilopa
Erichson, 1843 and Meoncerus Sharp, 1891 the type species are designated as well as
for Pallodes laetus Grouvelle, 1898 (Camptodes grouvellei nom. nov.) the lectotype
designation is made.
A.G. Kirejtshuk, Zoological Institute of Russian Academy of Sciences, Sainct Petersburg,
199034, Russia. E-mail: [email protected], [email protected], alexander_kirejtshuk@
yahoo.com
The system of the family Nitidulidae has been publication (the principle of priority). Besides, if a
greatly changed by efforts of many students and genus can be divided into recognizable subgenera,
essentially improved during the last 20-30 years. the latter were listed as well. To prepare such a
However there are no comprehensive publications complete arrangement of the taxa above the spe-
devoted a general view on the system of this family. cies level the author was obliged to include some
It is partly connected with the fact that some im- new data (new synonymy or other taxonomic
portant aspects of the system of this group are still propositions) which are explained bellow the list
needed to be considered. Recently the chapter on of taxa and these explanations in the necessary
Nitidulidae in the Palaearctic “catalogue” appeared cases are provided with appropriate comments
(Jelínek & Audisio, 2007). It contains some prin- (diagnoses, composition and so on). In all cases
cipal defi ciencies stimulated a preparation of this new synonymy is proposed on base of re-testing
paper. These defi ciencies can be explained that not the type species of genera and type series of all
all necessary taxonomic data achieved were taken synonymysed species names. Nevertheless, some
into account by the author of the catalogue and a new taxa of the tribe and genus rank are still in
part of published information was dropped from preparation by the author and his collaborators
the author’s consideration. The author of this pa- and not included in the list and notes to it. In some
per is aware that such a situation became possible important cases the type species are designated,
because he having known nearly all groups of this although in other cases, when they have already
family obtained from the recent and past faunas designated in previous publications, the mention
could publish only a small portion of these data and of them are omitted to shorten the volume of this
he has paid not enough attention for preparation of paper.
detailed generalizations on the family system. This The systematic construction of the family here
paper aims partly to compensate the mentioned represented should be regarded as a preliminary
defects, although a more thorough analysis of the version of a system based on the phylogeny and
systematic blocks of this family will be discussed history. At the moment, not-convergent (mono-
in the coming monographs (Kirejtshuk, in prepa- phyletic) origin of Cillaeinae and Cryptarchinae
ration). These monographs will cover a detailed is accepted only because they have the formal
review of bibliography, and therefore the writer structural characters uniting these subfamilies.
restricted the references only by the list of most It can be expected that a more detailed study of
appropriate sources supplementary to the notes historic development of these groups and more
here included and the publications with authors and detailed comparison of their structures will dem-
years of taxa available in the Zoological Records onstrate their polyphyletic state. While the origin
and different catalogues (Grouvelle, 1913; Jelínek of the family in general remains rather obscure,
& Audisio, 2007; Ponomarenko & Kirejtshuk, the relationship of Nitidulina ecliva Martynov,
2008 etc.) are mostly omitted to make this paper 1927 is not clear and this name is omitted in the
as compact as possible. list below. To clarify the systematic position of
The core of the paper includes the list of generic this and some other fossil species needs a further
names arranged into subfamilies, tribes (when careful research of many Cucujoiformian groups
they can be outlined) and the synonyms of each from the Jurassic and Cretaceous. The recent
taxon are put together according to dates of their subfamilies Calonecrinae and Maynipeplinae
ZOOSYST. ROSSICA Vol. 17 • A.G. Kirejtshuk: Generic classifi cation of sap beetles 109
are certainly closely related to other sap beetles choropsis Grouvelle, 1912; Haptoncognathus
are provisionally regarded in the composition of Gillogly, 1962)
the Nitidulidae, although these small groups are Subgenera
so distinct that they could be excluded from this Amystrops Grouvelle, 1906
family as two taxa with the family rank. Finally, Mandipetes Kirejtshuk, 1997
the palaeoendemic genera with unclear familiar Parepuraea Jelínek, 1977
attribution described among the Nitidulidae which Ceratomedia Kirejtshuk, 1990
should be regarded rather far from this family are Trimenus Murray, 1864
also omitted in the list [f.i., Sinonitidulina Hong, Ecnomaeus Erichson, 1843 (=Somaphorus
1983; Sinosoronia Zhang, 1992 etc.] Murray, 1864, syn. nov.) (3)
Platychorina Grouvelle, 1905
I. Subfamily CALONECRINAE Kirejtshuk, Baloghmena Kirejtshuk, 1987
1982 Stauromenus Kirejtshuk et Kvamme, 2001
Calonecrus Thomson, 1857 Tribe Taenioncini Kirejtshuk, 1998
Taenioncus Kirejtshuk, 1984
II. Subfamily MAYNIPEPLINAE Kirejtshuk,
Raspinotus Kirejtshuk, 1990
1998
Taeniolinus Kirejtshuk, 1998
Carpocryraea Kirejtshuk, 1998
Maynipeplus Kirejtshuk, 1998
Csiromenus Kirejtshuk et Kvamme, 2001
Eutaenioncus Kirejtshuk et Kvamme, 2001
Carpophilinae-lineage
IV. Subfamily CARPOPHILINAE Erichson, 1943
III. Subfamily EPURAEINAE Kirejtshuk, 1986
Procarpophilus de Jong, 1953
Tribe Epuraeini Kirejtshuk, 1986
Carpophilus Stephens, 1829
Subgenera
Crepuraea Kirejtshuk in Kirejtshuk et Pono- Carpophilus Stephens, 1829
marenko, 1990 (Nyujwa Perkovsky, 1990, syn. Megacarpolus Reitter, 1919
nov.) (1) Semocarpolus Kirejtshuk, subgen. nov. (4)
Epanuraea Scudder, 1892 Gaplocarpolus Kirejtshuk, subgen. nov. (5)
Epuraea Erichson, 1843 Askocarpolus Kirejtshuk, subgen. nov. (6)
Subgenera Plapennipolus Kirejtshuk, 1997
Epuraea Erichson, 1843 Ecnomorphus Motschulsky, 1858 (=Tri-
Ceroncura Kirejtshuk, 1994 brachys LeConte, 1861, syn. nov.; Eidocolas-
Dadopora Thomson, 1859 tus Murray, 1864, syn. nov.; Stauroglossicus
Strophoraea Kirejtshuk et Kvamme, 2001 Murray, 1864, syn. nov. ; Microxanthus Mur-
Epuraeanella Crotch, 1874 (=Omosiphora
ray, 1864, syn. nov.; Idocolastus Gemminger
Reitter, 1875)
et Harold, 1868) (7)
Ommoraea Kirejtshuk, 1998
Caplothorax Kirejtshuk, 1997
Micruria Reitter, 1874 (=Micrurula Reit-
Myothorax Murray, 1864
ter, 1884)
Nitops Murray, 1864 (=Endomerus Murray, 1864)
Aphenolia Reitter, 1884
Subgenera
Africaraea Kirejtshuk, 1989
Nitops Murray, 1864
Apria Grouvelle, 1919
Urocarpolus Kirejtshuk, 1997
Haptoncus Murray, 1864 (=Haptoncura
Ctilodes Murray, 1864
Reitter, 1875;
Loriarulus Kirejtshuk, 1987
Haptoncurina Jelínek, 1977, syn. nov.) (2)
Vulpixenus Kirejtshuk, 1990
Blackburnaea Kirejtshuk et Kvamme, 2001
Urophorus Murray, 1864 (?=Heterodontus Mur-
Marinexa Kirejtshuk, 1989
ray, 1864, nom. nudum)
Polinexa Kirejtshuk, 1989
Subgenera
Horniraea Kirejtshuk et Pakaluk, 1996
Urophorus Murray, 1864
Orthopeplus Horn, 1879
Anophorus Kirejtshuk, 1990
Grouvellia Kirejtshuk, 1984
Mystronoma Kirejtshuk, 1990
V. Subfamily AMPHICROSSINAE Kirejtshuk,
Amedanyraea Kirejtshuk et Pakaluk, 1996
1986
Amystrops Grouvelle, 1906 (=Propetes Reitter,
1875, non Walker 1851; Amystrops Grouvelle, Amphicrossus Erichson, 1843 (=Cametis Mo-
1906; Platychorinus Grouvelle, 1906; Platy- tschul sky, 1863; Lobostoma Fairmaire, 1892;
110 A.G. Kirejtshuk: Generic classifi cation of sap beetles • ZOOSYST. ROSSICA Vol. 17
Rhacostoma Berg, 1898; Nitidopecten Reichens- Omosiphila Kirejtshuk, 1990
perger, 1913) Temnoracta Kirejtshuk, 1988
Hisparonia Kirejtshuk, 2003
Ornosia Grouvelle, 1899
Nitidulinae-lineage
Pleoronia Kirejtshuk, 2003 (=? Axyrodes Mur-
ray, 1867, nom. nudum) (9)
VI. Subfamily MELIGETHINAE Thomson,
Amphotis Erichson, 1843
1859 (Meligethina)
Macleayania Kirejtshuk, 2003
Sebastianiella Kirejtshuk, 1995
Meligethinus Grouvelle, 1906 (=Prianella
Annachramus Kirejtshuk, 1995
Reit ter, 1919)
Stenoronia Kirejtshuk, 2003
Pria Stephens, 1829 (=Laria Scopoli, 1763,
Cormyphora Laporte de Castelneau, 1840; pars; Prometopia-complex
Allopria Kirejtshuk, 1980, syn. nov.) (8)
Microporodes Endrödy-Younga, 1978 Prometopia Erichson, 1843
Anthystrix Kirejtshuk, 1984 Subgenera
Micropria Grouvelle, 1899 (=Metapria Grou- Prometopia Erichson, 1843
velle, 1908/1909)) Parametopia Reitter, 1884
Cyclogethes Kirejtshuk, 1979 + Palaeometopia Kirejtshuk et Poinar,
Cryptarchopria Jelínek, 1975 2007
Kabakovia Kirejtshuk, 1979 Cacconia Sharp, 1890
Horakia Jelínek, 2000
Microporum C. Waterhouse, 1876 (=Probaenus Megauchenia-complex
C. Waterhouse, 1876; Microporellus Endrödy-
Younga, 1978) Axyra Erichson, 1843 (=Galaor Thomson, 1858)
Palmopria Endrödy-Younga, 1978 Megauchenia Macleay, 1825 (=Ischaena Erich-
Cornutopria Endrödy-Younga, 1978 son, 1843; Orvoenia Dojoz, 1980, syn. nov.) (10)
Lechanteuria Endrody-Younga, 1978 (=Pria- Tetrisus Murray, 1864 (=Pseudoischaena Grou-
nella Lechanteur, 1955, non Reitter, 1919) velle, 1897, syn. nov.) (11)
Meligethes Stephens, 1829 Megaucheniodes Audisio et Jelínek, 1993
Subgenera Taraphia Audisio et Jelínek, 1993
Chromogethes Kirejtshuk, 1989 Pseudoplatychora Grouvelle, 1890
Lariopsis Kirejtshuk, 1989
Ipidia-complex
Clypeogethes Scholtz, 1932 (=Idiogethes
Kirejtshuk, 1977)
Ipidia Erichson, 1843
Meligethes Stephens, 1829 (=Odontogethes
Subgenera
Reitter, 1871)
Ipidia Erichson, 1843
Astylogethes Kirejtshuk, 1992
Hemipidia Kirejtshuk, 1992
Acanthogethes Reitter, 1871
Platychora Erichson, 1843 (=Pherocopis
Thomson, 1858) (12)
VII. Subfamily NITIDULINAE Latreille, 1802
Taracta Murray, 1867
Psilotus Fischer, 1829 (=Cerophorus Laporte
A. Tribe Nitidulini Erichson, 1843 (=Thalycrina de Castelneau, 1840)
Thomson, 1859; Pocadiini Seidlitz, 1872; Orvoe- Perilopa Erichson, 1843
nini Dajoz, 1980, syn. nov.) (10)
Phenolia-complex
Nitidula-complex
Stelidota Erichson, 1843
Nitidula Fabricius 1775 Phenolia Erichson, 1843 (Lordites auctorum,
Omosita Erichson, 1843 (=Saprobia Gangl- non Erichson, 1843; Lasiodactylus auctorum, non
bauer, 1899) Perty, 1830-1834)
Subgenera
Soronia-complex Aethinodes Blackburn, 1891
Lasiodites Jelínek, 1999
+Microsoronia Kirejtshuk et Kurochkin, 2008 Phenolia Erichson, 1843
Soronia Erichson, 1843 (=Platipidia Broun, Plesiothina Kirejtshuk, 1990
1893) Gaulodes Erichson, 1843
Lobiopa Erichson, 1843 (=? Axyrodes Murray, Ussuriphia Kirejtshuk, 1992
1867, nom. nudum) (9) Ostomarcha Kirejtshuk, 2006
ZOOSYST. ROSSICA Vol. 17 • A.G. Kirejtshuk: Generic classifi cation of sap beetles 111
Perilopsis-complex Pocadites Reitter, 1884
Pocadius Erichson, 1843
Tagmolycra Kirejtshuk et Leschen, 1998
Epuraeopsis Reitter, 1875
Parapocadius gen. nov. (16)
Perilopsis Reitter, 1875
Pleuroneces Olliff, 1891
Subgenera
Thalycra Erichson, 1843 (=Perthalycra Horn,
Perilopsis Reitter, 1875
1879)
Testudoraea Kirejtshuk, 1986
Pseudothalycra Howden, 1962
Cratonura Reitter, 1875
Quadrifrons Blatchley, 1916
Neothalycra Grouvelle, 1899
Aethina-complex
Thalycrinella Kirejtshuk in Kirejtshuk et Le-
schen, 1998
Psilonitidula Heller, 1916 Pocadionta Lucas, 1920 (=Pocadiopsis Grou-
Lasiodactylus Perty, 1830-1834 (=Nitidulingen velle, 1898, non Fairmaire, 1896)
Gillogly, 1965) (13) Thalycrodes Blackburn, 1891
Subgenera Rixerodes Kirejtshuk et Lawrence, 1992
Lasiodactylus Perty, 1830-1834 Australycra Kirejtshuk et Lawrence, 1992
Lordyra Gemminger & Harold, 1868, stat. Pocadiolycra Kirejtshuk et Leschen, 1998
nov. (13)
Neopocadius Grouvelle, 1906 (13) B. Tribe Cychramini Lacordaire, 1854 (1855)
Subgenera (Cychramides)
Neopocadius Grouvelle, 1906 (=Pseudosteli-
dota Grouvelle, 1906, syn. nov.) (13) +Cychramites Wickham, 1913
Brounthina Kirejtshuk, 1997, stat. nov. (13) Cychramus Kugelann, 1794 (=Campta Ste-
Australaethina Kirejtshuk et Lawrence, 1999 phens, 1830; Aethinopsis Grouvelle, 1908, syn.
(=Idaethina Reitter, 1875, non Gemminger et nov.) (17)
Harold, 1868. et non Olliff, 1884) Xenostrongylus Wollston, 1854 (=Strongyllodes
Aethina Erichson, 1843 (=Aethinopa Reitter, Kirejtshuk, 1992)
1875; Pseudomystrops Grouvelle, 1912/1913; Subgenera
Meligethopsis Rebmann, 1944) Xenostrongylus Wollaston, 1854
Subgenera Strongylolasius Reitter, 1911
Aethina Erichson, 1843 Oxystrongylus Reitter, 1911
Cleidorura Kirejtshuk et Lawrence, 1999 Kirejtshukostrongylus Audisio et Jelínek in
Idaethina Gemminger et Harold, 1868, Audisio, Mariotti, Jelínek & DeBiase 2001
(=Macroura Reitter, 1873, non Meuschen, Axychramus Kirejtshuk, 1996
1778, non Loew, 1845; Idaethina Olliff, Ceratochramus Kirejtshuk, 1986
1884, non Reitter, 1875; Olliffura Jelínek
et Kirejtshuk, 1986) C. Tribe Mystropini Murray, 1864 (Mystropidae)
Circopes Reitter, 1873
Ithyra Reitter, 1873 Mystrops Erichson, 1943 (=Priops Reitter,
Anister Grouvelle, 1901 (=Oturovana Reitter, 1873; Cryptoraea Retter, 1873, syn. nov.; Eumys-
1915) trops Sharp, 1889) (18)
Anthepurops Kirejtshuk, 1996
Pocadius-complex (14) Anthocorcina Kirejtshuk, 1996
Platychorodes Reitter, 1884
+Omositoidea Schaufuss, 1891 Cychrocephalus Reitter, 1873 (=Cychropiestus
Atarphia Reitter, 1884 Reitter, 1875)
Hebasculinus Kirejtshuk, 1992 Nitidulora Reitter, 1873
Hebascus Erichson, 1843 Palmostrops Kirejtshuk et Jelínek, 2000
Subgenera
Hebascus Erichson, 1843 D. Tribe Cyllodini Everts, 1898 (=Strongylini
Teichostethus Sharp, 1891, stat. nov. Sturm, 1844 (Strongylinae), non Müller, 1780;
(Trichostethus) (15) Arborotubini Leschen et Carlton, 2004)
Hyleopocadius Jelinek, 1977
Kryzhanovskiella Kirejtshuk, 2006 +Cyllolithus Kirejtshuk in Kirejtshuk et Pono-
Niliodes Murray, 1868 marenko, 1990
Physoronia Reitter, 1884 (=Lordyrodes Reitter, Somatoxus Sharp, 1891 (=Somatorus Grou-
1884; Pocadioides Ganglbauer, 1899; Osotima velle, 1891, err.)
Rebmann, 1944) Cyclocaccus Sharp, 1891
112 A.G. Kirejtshuk: Generic classifi cation of sap beetles • ZOOSYST. ROSSICA Vol. 17
Cyllodes-complex Eupetinus Sharp, 1908 (=Apetinus Scott, 1908)
Prosopeus Murray, 1864 (=Prosopius Gem-
Camptodes Erichson, 1843 (=Eucamptodes minger et Harold, 1868; Nesopeplus Sharp, 1908;
Sharp, 1890) Nesopetinus Sharp, 1908)
Meoncerus Sharp, 1891 (19) Cillaeopeplus Sharp, 1908 (=Notopeplus Sharp,
Apsectochilus Reitter, 1874, distinct genus (19) 1908)
Lordyrops Reitter, 1875, distinct genus (19) Colopterus Erichson, 1842 (=Colastus Erich-
Carinocyllodes Leschen, 1999 son, 1843)
Cyllodes Erichson, 1843 (=Strongylus Herbst, Colopterus Erichson, 1843
1792, non Muller, 1780; Volvoxis Kugelann, 1794; Cyllopodes Murray, 1864
Mecyllodes Sharp, 1891, syn. nov.; Pseudocamp- Grammorus Murray, 1868 (=Grammophorus
todes Grouvelle, 1896) (20) Gerstäcker, 1864, nec Solier, 1851; Colopteroides
Eusphaerius Sharp, 1891 Watrous, 1982, syn. nov.) (23)
Viettherchnus Kirejtshuk, 1985 Grouvellepeplus Kirejtshuk, 2001
Ceramphosia Kirejtshuk et Kirk-Spriggs, 1996 Brachypeplus Erichson, 1842 (=Nitidulopsis
Camptomorphus Grouvelle, 1908 Walker, 1858; Tasmus Murray, 1864)
Pallodes Erichson, 1843 (21) Subgenera
Coxollodes Kirejtshuk, 1987, stat. nov. (21) Brachypeplus Erichson, 1842
Neopallodes Reitter, 1884 Selis Murray, 1864
Cyllodesus Reitter, 1877 (=Strongylomorphus Leiopeplus Murray, 1864
Reitter, 1875, non Motschulsky, 1853) Idosoronia Schaufuss, 1891
Teloconus Grouvelle, 1916
Oxycnemus-complex Tokocillaeus Kirejtshuk, 2001
Onicotis Murray, 1864, stat. nov. (24)
Oxycnemus Erichson, 1843 (22)
Campsopyga Murray, 1864 (=Hypodetus Mur-
Eugoniopus Reitter, 1884 (22)
ray, 1964).
Psilopyga LeConte, 1853 (22)
Cillaeus Laporte de Castelneau, 1835
Interfaxia gen. nov. (22)
Subgenera
Triacanus Erichson, 1843 (=Tricanus auctorum)
Cillaeus Laporte de Castelneau, 1835
Monafricus Kirejtshuk, 1995
Xanthopeplus Fairmaire, 1880
Gymnocychramus Lea, 1922
Paracillaeopsis Kirejtshuk, 2001
Pycnocnemus Sharp, 1891
Cillaeopsis Grouvelle, 1899
Halepopeplus Murray, 1864 (=Chalepopeplus
Arborotubus-complex
Gemminger et Harold, 1868)
Liparopeplus Murray, 1864 (=Carpophilops
Arborotubus Leschen et Carlton, 2004
Grouvelle, 1898)
E. Tribe Cychramptodini Kirejtshuk et Lawrence, Halepopeplus Murray, 1864
1992 Adocinus Murray, 1864
Platynema Ritsema, 1885 (=Orthogramma
Cychramptodes Reitter, 1874 Murray, 1864, not Guenée 1852 et nec R. L.
Miskoramus Kirejtshuk et Lawrence, 1992 (Reichenbach, Leipzig), 1817)
Cylindroramus Kirejtshuk et Lawrence, 1992 Ithyphenes Murray, 1864
Macrostola Murray, 1864
F. Tribe Lawrencerosini Kirejtshuk, 1990 Macrostolops Grouvelle, 1916
Conotelus Erichson, 1843
Lawrencerosus Kirejtshuk, 1990
Krakingus Kirejtshuk, 1990 X. Subfamily CRYPTARCHINAE Thomson,
Koryaga Kirejtshuk, 1990 1859 (=Ipinae Erichson, 1843; Ipsomorpha
Koryaginus Kirejtshuk, 1990 Reitter, 1873)
VIII. Subfamily CILLAEINAE Kirejtshuk et A. Tribe Cryptarchini (=Pityophagini Fauconnet,
Audisio, 1986 1896; Glischrochilini Iablokoff-Khnzoryan, 1966)
Gonioryctus Sharp, 1878 (=Goniothorax Sharp, Cnips Philippi, 1864
1908; Nesapterus Sharp, 1908; Eunitidula Sharp, Cnipsarcha Jelínek, 1982
1908) Cryptarcha Shuckard, 1839 (=Cryptarchus
Orthostolus Sharp, 1908 Heer, 1843; Priatelus Broun, 1882 (pro Priateles
Apetasimus Sharp, 1908 (=Cyrtostolus Sharp, Broun, 1881), syn. nov.; Lepiarcha Sharp, 1891;
1908) Liarcha Sharp, 1891; Cryptarchula Ganglbauer,
ZOOSYST. ROSSICA Vol. 17 • A.G. Kirejtshuk: Generic classifi cation of sap beetles 113
1899; Africanips Lechanteur, 1959; Cryptarchina (type species: Crepuraea archaica Kirejtshuk
Iablokoff-Khnzoryan, 1966) (25) in Kirejtshuk et Ponomarenko, 1990, by origi-
Homepura Broun, 1893 (=Inopria Broun, 1921) nal designation), while Nyujwa syn. nov. (type
Kaszabena Kirejtshuk, 1987 species: Nyujwa zherichini Perkowsky, 1990,
Paromia Westwood, 1850 (=Lioschema Fair- by monotypy) appeared in the 4th one of the
maire, 1861; Aparomia Redtenbbacher, 1867) same year. The latter completely corresponds the
Paromidia Reitter, 1873 representatives of the former originated from the
Pityophagus Shuckard, 1839 same site (Baissa), they should be regarded as
Glischrochilus Reitter, 1873 (Ips Fabricius, synonyms. Another problem is that among the
1777, non DeGeer, 1775) congeners described in composition of Crepuraea
Subgenera one species named as C. zherichini and in this
Glischrochilus Reitter, 1873 case N. zherichini should be also recognized as a
Librodor Reitter, 1884 (=Cryptarchips junior homonym of C. zherichini. At the present,
Reitter, 1911; Cephalips Arrow, 1931) the relation of the holotype of N. zherichini with
Gymnoparomius Kirejtshuk, 1987 other members of the genus is still unclear and a
further comparison of the specimens collected in
B. Tribe Platyarchini Kirejtshuk, 1998
Baissa is needed.
2. The synonymy of names Haptoncus (type
Platyarcha Kirejtshuk, 1987
species: Haptoncus tetragonus Murray, 1864, des-
Amlearcha Kirejtshuk, 1987
ignated by Parsons, 1843) and Haptoncurina syn.
nov. (type species: Epuraea angustula Motschul-
C. Tribe Arhinini Kirejtshuk, 1987
sky, 1863, non Epuraea angustula Sturm, 1844,
Ceratarhina Kirejtshuk, 1981 by original designation) should be regarded as
Arhina Murray, 1867 evident, because some Indo-Malayan species of
Arhinella Kirejtshuk, 1981 Haptoncus [E. (H.) concolor Murray, 1864 and E.
(H.) fallax (Grouvelle, 1897)] and the members of
D. Tribe Eucalosphaerini Kirejtshuk, 1987 Haptoncurina show a variability in the subgeneric
disagnostic characters making the discrimination
Eucalosphaera Jelínek, 1978 (=Calosphaera of the subgenera quite problematic.
Jelínek, 1974, non Campbell, 1951) 3. The synonymy of names Ecnomaeus (type
species: Ecnomaeus planus Erichson, 1843, by
X. Subfamily CYBOCEPHALINAE Jaquelin monotypy) and Somaphorus syn. nov. (type spe-
du Val, 1858 (Cybocephalites) cies: Somaphorus ferrugineus Murray, 1864, by
monotypy) should be recognized, as both type
Cybocephalus Erichson, 1844 (=Phantazo- species belong to the same group of clear relatives
merus J. Duval, 1854; Stagnomorpha Wollaston, distriduted in the subequatorial areas of the East
1854; Acribis C.Waterhause, 1877; Dissia Cho- Hemisphere.
baut, 1896) This group was erroneously put into the sub-
Subgenera family Cilaeinae (Kirejtshuk, 1986), because the
Cybocephalus Erichson, 1843 elytra of its species remain two last abdominal
Theticephalus Kirejtshuk, 1988 segments uncovered and somewhat more compact
Pycnocephalus Sharp, 1891 antennal club than in other groups of Epuraeinae.
Hierronius Endrödy-Younga, 1968 Although the structure of the male abdomen in
Pastillodes Endrödy-Younga, 1968 Ecnomaeus species, including shape of the anal
Taxicephomerus Kirejtshuk, 1994 sclerite, ventral plate and genitalia, is certainly
Horadion Endrödy-Younga, 1976 similar to that in other groups of the subfamily
Pastillus Endrödy-Younga, 1962 Epuraeinae, but not to that in Cillaeinae. Besides,
Endrodiellus Endrödy-Younga, 1962 the structure of thorax of Ecnomaeus species,
particularly prosternal process and mesoventrite
INCERTAE SEDIS
is certainly comparable with that in Epuraeinae
rather than other sap beetles.
+Miophenolia Wickham, 1916
4. The subgenus Semocarpolus subgen. nov.
Nodola Brethes 1925 (?=Cybocephalus)
(type species: Carpophilus marginellus Motschul-
+Oligamphotis Theobald, 1937
sky, 1858)
Prioschema Reitter, 1976
Etymology. The name of the subgenus is formed
Notes: from the Greek “sema” (character) and “carpolus”
(Megacarpolus and other generic names with
1. The name Crepuraea was published the the end “polus”) formed in turn from the Greek
in 2d issue of Palaeontological Journal in 1990 “carpus” (fruit, foetus).
114 A.G. Kirejtshuk: Generic classifi cation of sap beetles • ZOOSYST. ROSSICA Vol. 17
Diagnosis: This new subgenus is well character- Brazil) and one species under description from
ized by a raised median carina of the mesoventrite, the Indo-Malayan Region.
almost rectilinear submesocoxal line and undi- 5. The subgenus Gaplocarpolus subgen. nov.
vided fork-sclerite of tegmen. It seems to be close (type-species: Carpophilus (Carpophilus) cunei-
to Carpophilus sensu str. differing from it only in formis Murray, 1864).
the mentioned characters and very shallow and in- Etymology. The name for this taxon is created
distinct punctures on mesoventrite. Species of this from the Greek “gaplos” (meaning simple, soli-
new subgenus have some resemblance to those of tary, lonely) and “carpolus” used for some names
Megacarpolus, but their bodies are reddish brown of the Carpophilinae and formed from the Greek
in coloration, smaller and more convex, with very “carpus” (fruit, foetus).
steeply sloping pronotal and elytral unexplanate Diagnosis. This new subgenus is more similar to
sides, less compact antennal club, hypopygidium the subgenera Askocarpolus subgen. nov., Carpo-
of their males without any additional depression philus sensu str., Megacarpolus and Semocarpolus
[although the Indo-Malayan species of Mega- subgen. nov. than other groups of the subfamily.
carpolus are also without clear depression on It is very distinct from the mentioned taxa by the
this sclerite in males]. Species of Semocarpolus comparatively sparse puncturation of integument,
subgen. nov. are also similar to the representa- slightly loose antennal club between antennomeres
tives of Gaplocarpolus subgen. nov., but, except 9 and 10, spiracles of tergite VI widely transverse,
for the mentioned distinguishing characters, also medially convex and moderately widened apex
have rather distinct dorsal puncturation and only of prosternal process. This new subgenus is also
shallow punctures on mesoventrite as well as their characterized by the following characters: medium
females with simple pygidium. Also, the species body size, subvertical pronotal and elytral sides,
of this new subgenus are clearly distinguished slightly curved along procoxae to subfl attened
from Askocarpolus subgen. nov. by more robust prosternal process with strongly widened apex
and more convex body with very steeply sloping and simple mesoventrite. The members of Gap-
pronotal and elytral unexplanate sides, not project- locarpolus subgen. nov. have the similar outline
ing posterior pronotal angles, lack of raised depres- of submesocoxal line. Externally species of this
sions on pronotal and elytral disks, and also by lack new subgenus are more similar to representatives
of paramedian pockets on mesoventrite. of Semocarpolus subgen. nov. [especially to C.
This new subgenus is also characterized by a (Semocarpolus) rubescens], differing from them
somewhat loose antennal club between antenno- not only in the features listed above, but also in
meres 9 and 10, but less loose than that observed widely truncate labral lobes, transverse depression
among members of Ecnomorphus or even in on male metaventrite before anterior edge, subme-
Askocarpolus subgen. nov. Nevertheless, the socoxal line gently curving and slightly deviating
antennal club of species of this subgenus is less from posterior edge of cavities. Besides it, this
compact in comparison with that in the representa- new subgenus differs also from subgenera:
tives of all subgenera which are here regarded as – Carpophilus sensu str. also in the simple
similar and probably related to it [i.e. Megacar- mesoventrite bearing rather shallow indistinct
polus, Gaplocarpolus subgen. nov., Askocarpolus punctures;
subgen. nov. and Carpophilus]. – Megacarpolus also in the lighter, smaller
According to N. Hayashi (1978) the larvae of C. and more convex body, more distinct and very
(S.) marginellus, in contrast to other groups of the sparse puncturation of dorsum, distinct subme-
subfamily, could be characterized by the indistinct socoxal line deviating from the posterior edge
or even absent ocelli and pair of transverse oval of cavities;
patches of sclerotized microscopic asperities on – Askocarpolus subgen. nov. also in the even
abdominal terga II-VIII. pronotal disk, simple mesoventrite (without trace
Bionomy. The species of this new subgenus of paramedian pockets), distinct submesocoxal
are associated with different decaying products line deviating from the posterior edge of cavi-
of plant origin and C. (S.) marginellus is a char- ties.
acteristic pest of stored products with an almost This new taxon is proposed in order to coun-
world-wide range. terbalance the considered groups with compa-
Composition and distribution. This new sub- rable hiati in characters. The combination of its
genus is represented only with 4 species: type peculiarities is corresponding to those of other
species with subcosmopolitan range in human subgenera of Carpophilus sensu lato.
stores and under natural conditions mostly in the Bionomy. Imagines of both species of this new
areas with tropical and subtropical climate; C. (S.) subgenus were captured probably in montane
rubescens Murray, 1864 distributed in the Indo- forest at elevation of 200-2 300 m.
Malayan region; C. (S.) succisus Erichson, 1943 Composition and distribution. Three members
from Neotropical Region (Central America and of this new subgenus (type species and two species
ZOOSYST. ROSSICA Vol. 17 • A.G. Kirejtshuk: Generic classifi cation of sap beetles 115
under description) are known from the Indochi- narrower protarsus, more elongate last abdominal
nese and Malayan provinces. segment in females;
6. The subgenus Askocarpolus subgen. nov. – from Ecnomorphus in the less fl attened body
(type-species: Carpophilus oblongopunctatus and different characters of puncturation and sculp-
Grouvelle, 1903) ture of integument;
Etymology. The name of the subgenus is com- – from Myothorax in the less convex and usually
posed from the Greek “askos” (chamber, cell, sac) elongate oval body, distinct anterior and posterior
and “carpolus” (Megacarpolus and other generic angles of pronotum.
names with the end “polus”) transformed from Finally, the mentioned paramedian pockets on
the Greek “carpus” (fruit, foetus). mesoventrite of the new subgenus under consid-
Diagnosis. This new subgenus is very distinct eration have some analogy with depressions on
from all groups of the subfamily (and the family mesoventrite in Sebastianiella spp. (Nitidulinae)
as a whole) due to a pair of deep paramedian from South Africa.
pockets in the middle of mesoventrite situated at Bionomy. The species of this new subgenus
level of apex of prosternal process. It is also well seem to be associated with decaying fruits and
characterized by more or less distinct oval para- other decaying products of plant origin.
median depressions at base of the pronotum and Composition and distribution. This new subge-
very weak longitudinal depressions in anterior half nus is represented only with 5 species (4 of them
of elytra, almost simple scarcely lobed meso- and under description) known from the south of the
metatarsi, posterior edge of metaventrite with a Palaearchearctic Province of Palaearctic Region
deep excision between metacoxae, not completely and Indochinese Province of Indo-Malayan
divided fork-sclerite of tegmen and articulated Region, but the type species of the subgenus is
connection between ventral plate and spiculum described from Darjeeling.
gastrale of the male. 7. The synonymy of the names Ecnomorphus
Appearence of its representatives is also rather (type-species: Nitidula sexpustulata Fabricius,
similar to that of species in the subgenus Gaplo- 1791; designated by Jelínek & Audisio, 2003);
carpolus subgen. nov., but differs from that in the Tribrachys syn. nov. (type-species: Tribrachys cau-
more or less expressed paramedian depressions at dalis LeConte, 1859; designated by C.T. Parsons,
base of pronotum and longitudinal depressions 1943); Eidocolastus syn. nov. (type-species: Colas-
on elytra as well as in distinct anterior angles tus plagiatipennis Motschulsky, 1858; designated
of pronotum, less strongly sloping pronotal by Jelínek & Audisio, 2003); Stauroglossicus syn.
and elytral sides and in shape of the last labial nov. (type-species: Stauroglossicus terminalis Mur-
palpomeres strongly widened to apex. This new ray, 1864: 398; designated by Jelínek & Audisio,
subgenus shares a certain similarity (particularly 2003); Microxanthus syn. nov. (type species: Car-
in the body shape, characters of puncturation and pophilus tempestivus Erichson, 1843; designated by
sculpture of sclerites) to the subgenera Megacar- Jelínek & Audisio, 2003); Idocolastus (proposed
polus and Semocarpolus subgen. nov., although in for Eidocolastus) is based on wide comparison of
addition to characteristic paramedian pockets on many members of the genus.
mesoventrite, depressions on pronotum and elytra, The name Tribrachys was treated as a junior
the representatives of Askocarpolus subgen. nov. synonym of the name Carpophilus in general
differ from those of the fi rst in their somewhat (Grouvelle, 1913), however, in accordance with
longer elytra, usually more asymmetrical last the designations of type species the first of
labial palpomere, lack of sexual dimorphism in mentioned names should be regarded as a junior
structure of 3-segmented antennal club; and from synonym of Ecnomorphus. The type-species of
the second – in the less robust and less convex Tribrachys seems to be closely related to C. (E.)
body with more gently sloping pronotal and ely- ligneus Murray, 1864, while the type-species
tral sides to moderately explanate edges, strongly of the Ecnomorphus is very distinct from other
widened last labial palpomere and projecting groups of the subgenus in some characters, which
posterior pronotal angles. scarcely could make possible to regard it as a
Besides the characteristic paramedian pockets member of separate subgenus.The type-species
on mesoventrite as well as depressions on prono- of Stauroglossicus and Eidocolastus belong to
tum and elytra, Askocarpolus subgen. nov. differs the groups closely related each to another as well
from other subgenera of the genus Carpophilus as quite similar to the rest groups of the subge-
sensu lato: nus and, therefore, the last two names should be
– from Carpophilus sensu str. in the absence regarded in composition of the same subgenus
of isolated distal plate of mesoventrite divided together with Tribrachys and Ecnomorphus.
by median ridge; less steeply sloping pronotal V. Motschulsky (1858) proposed the name Ec-
and elytral sides, wider last labial palpomeres, nomorphus in connection with the description of
less widely separated of meso- and metacoxae, E. fulvipes which he compared with the European
116 A.G. Kirejtshuk: Generic classifi cation of sap beetles • ZOOSYST. ROSSICA Vol. 17
Nitidula sexpustulata Fabricius, 1792. A. Murray murrayi Grouvelle, 1892 (=hebetatus Grouvelle,
(1864) removed E. fulvipes from the subgenus 1908, syn. nov. proposed for fulvipes Grouvelle,
Ecnomorphus, because he decided that the char- 1894, non Motschulsky, 1858); C. (E.) plagiati-
acters listed by V. Motschulsky for this subgenus pennis (Motschulsky, 1858) (=bosschai Grouvelle,
fi tted rather to the European than Indo-Malayan 1892; dilutus Motschulsky, 1858, non Murray,
representatives. However, neither A. Murray nor 1864; nigricans Grouvelle, 1897, syn. nov.);
V. Motschulsky designated the type-species of C. (E.) proximus Grouvelle, 1906; C. (E.) senex
the taxon properly. V. Motschulsky included in Murray, 1864; C. (E.) sexpustulatus (Fabricius,
Ecnomorphus also E. biguttatus, described by him 1791) (=abbreviatus Panzer, 1793; bimaculatus
in the same publication. A. Murray interpreted Marsham, 1802); C. (E.) sibiricus Reitter, 1879; C.
this name for species of the subgenus Myothorax, (E.) sinuatus Grouvelle, 1917; C. (E.) subplanus
supposing that V. Motschulsky under this name Grouvelle, 1917; C. (E.) tempestivus Erichson,
described a variety of C. (M.) vittiger Murray, 1843; C. (E.) terminalis Murray, 1864 (=gentilis
1864. The recent examination of the type series Murray, 1864, syn. nov.); C. (E.) ustulatus Mur-
of E. biguttatus showed that the species with ray, 1864; C. (E.) variabilis Grouvelle, 1897;
this type series indeed should be regarded as a C. (E.) waterhousei Dobson, 1993; C. (E.) zuni
member of the genus Platyarcha Kirejtshuk, 1987 Casey, 1884; and also some dozens of species
[Platyarcha biguttata (Motschulsky, 1858), comb. which will be described in a future monograph
nov.: Cryptarchinae, Platyarchini]. of the author devoted to the fauna of the Indo-
The groups which could be regarded as Ec- Malayan Region.
nomorphus and Microxanthus do not show any 8. Because of variability in extent of the de-
clear hiatus. The alone character which can be velopment of tarsal claws among some groups
used to discriminate them could be the distinctness of related species the names Pria and Allopria
of posterior angles of the pronotum. However this syn. nov. would be reasonable to consider as
character sometimes is rather variable even within synonyms.
one species or within one series of the same spe- 9. The name Axyrodes was proposed for one
cies and, therefore, it can not be used for a reliable undescribed Neotropical species (nomen nudum)
diagnosis. This is a reason to synonymize the taxa and compared with the described African and
Ecnomorphus and Microxanthus. Himalayan species of Axyra. Later A. Grouvelle
Composition of the subgenus (in all cases new put one species to this taxon [Axyra (Axyrodes)
synonymy is based on re-testing type series): nitida Grouvelle, 1898], which indeed is a mem-
C. (E.) acutangulus Reitter, 1884 (=cingula- ber of Pleoronia (comb. nov.). At the same time,
tus Reitter, 1884, syn. nov.; ? cingulatoides the characters mentioned by Murray (1867: 170)
Na kan e, 1959 – the latter was proposed taken make possible to assume that having proposed the
in to consideration the studied variability of the “Axyrodes” was considered by him as one elongate
species); C. (E.) alticola Sharp, 1889; C. (E.) member of Pleoronia or Lobiopa.
antiquus Melsheimer, 1844; C. (E.) apicipennis 10. The synonymy of the names Megauchenia
Fairmaire, 1869; C. (E.) bakeweli Murray, 1864 and Orvoenia syn. nov. (described among Co-
(=planatus Murray, 1864, syn. nov.; aterrimus lydiidae) is evident because the type species of
Macleay, 1864, syn. nov.); C. (E.) brachypterus the latter, Orvoenia borneensis Dajoz, 1980, is
(Say, 1825); C. (E.) comatus Sharp, 1889; C. (E.) certainly very similar to Megauchenia gracilis
compressus Murray, 1864; C. (E.) confertus Sharp, Kirejtshuk, 1990. T.K. Pal & J.F. Lawrence (1986)
1889; C. (E.) corticinus Erichson, 1843; C. (E.) wrote on similarity of the species of these taxa
debilis Grouvelle, 1897 (=opaculus Grouvelle, when they excluded the tribe Orvoenini from the
1897, syn. nov.); C. (E.) deplanatus Boheman, family Colydiidae.
1854; C. (E.) discoideus LeConte, 1858; C. 11. The synonymy of the names Tetrisus and
(E.) dubitabilis Grouvelle, 1897; C. (E.) dufaui Pseudoischaena syn. nov. is quite clear, because
Grouvelle, 1908; C. (E.) elaterinus Sharp, 1889; the type species of both (Tetrisus cholevoides
C. (E.) epuraeoides Sharp, 1889; C. (E.) extensus Murray, 1864, by monotypy and Pseudoischaena
Grouvelle, 1908 (=trapezicollis Kirejtshuk, 1995); longula Grouvelle, 1897, by nomotypy) are close-
C. (E.) ferrugineus Murray, 1864; C. (E.) frivolus ly related and belong to the same species group.
Murray, 1864; C. (E.) fulvipes Motschulsky, 1858 The taxa Trimenus and Tetrisus were regarded as
(=fusculus Motschulsky, 1858); C. (E.) humilis subgenera of the same genus in the composition of
Erichson, 1843; C. (E.) inconspicuous Murray, the subfamily Epuraeainae (Kirejtshuk, 1998), as
1864; C. (E.) lepidus Murray, 1864; C. (E.) ligneus the type species of them have females looking like
Murray, 1864 (=decipiens Horn, 1879); C. (E.) rather similar because of probable convergences
luridipennis Macleay, 1873 (=loriai Grouvelle, of external structures. Nevertheless, the males of
1906, syn. nov.); C. (E.) marginatus Erichson, both are rather different and show a clear attri-
1843; C. (E.) mcnamarai Dobson, 1993; C. (E.) bution of different subfamilies (Epuraeinae and