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A.G. Kirejtshuk: essay on the family Nitidulidae from the monograph

Family Nitidulidae (incl. Cybocephalinae)


Body very diverse, from 0.9 up to 15.0 mm in length; 
frequently moderately convex dorsally and somewhat flattened 
or weakly convex ventrally, sometimes strongly convex 
dorsally and flattened ventrally, or subhemispheric and able 
to roll up in a ball (exclusively Cybocephalinae); usually 
of oval or somewhat elongate outline from above. Surface 
usually with uniform punctation, but sometimes with 
punctures of different sizes arranged in not quite regular 
rows, but elytra not infrequently more or less striate and 
with longitudinal rows of larger or smaller punctures. 
Pubescence usually moderately dense, fine, unicoloured and 
moderately conspicuous, although sometimes it is completely 
reduced or consists of groups of different sizes, shape and 
coloration, and not infrequently pronotal and elytral sides 
more or less ciliate. Head partly retracted into prothoracic 
segment, more or less prognathous; labrum usually bilobed, 
sometimes fused with inner surface of frons (exclusively 
Cryptarchinae); mandibles with acute apices, usually with 
well developed mola and prosteca; maxillae unilobed with 
normally raised palpi; labium with 3-segmented palpi. 
Antennae usually 11-segmented, with 3 or 4 segmented compact 
club, clearly dorsoventrally depressed, sometimes they 
consist of 10 or less segments and their club is reduced up 
to 2 segments, or includes one or more additional segments - 
up to 5-8 segments (completely comprising a compact or 
partly loosed club); however, species of the genus 
Calonecrus J. Thomson, 1857 have 10-segmented antennae with 
1-segmented and undepressed club. Pronotum and elytra with 
sides almost always distinctly bordered and usually more or 
less explanate. Elytra rarely complete, but usually more or 
less shortened, with clearly separated epipleura sharply 
turned ventrally and becoming obsolete towards their apices 
(especially in the forms with more shortened elytra); 
elytral epipleura in Calonecrus species having no lateral 
fold at curvature from dorsal side. Fore coxae strongly 
transverse and with a well exposed trochantin, separated by 
a moderately developed prosternal process; their cavities 
not completely or completely closed. Mesosternum, as a rule, 
somewhat deepened (excavate) in comparison with a remainder 
of underside and frequently with a medial carina swollen in 
the middle; mid coxae transverse with an exposed trochantin 
and their cavities open externally. Metasternum transverse 
or subquadrate, usually with a more or less distinct medial 
suture and a trace of paracoxal sutures, caudal marginal 
line behind mid coxae deviated from hind edge of cavities 
forming an "axillar space", and in some cases there is also 
an intercoxal line in fore part of metasternum; hind coxal 
cavities strongly transverse, moderately separated each from 
other, or closer together, but never contiguous. Tergite 
VIII transformed into an anal sclerite, heavily sclerotized, 
well raised and large in males and reduced and usually 
submembranous in females, forming together with the remains 
of 9th sternite ("ventral plate") and "spiculum gastrale" in 
males or "spiculum ventrale" in females, a genital capsula. 
Forms with complete elytra have small oval and uniform 
spiracles, usually between tergites and laterosternites of 
1-6 segments; most forms with shortened elytra have largest 
spiracles on segments before elytral apices (not 
infrequently very transverse and located on tergites), 
spiracles on uncovered segments are rather or extremelly 
small and sometimes elongate. Venation of hind wing reduced; 
radial and anal cells, as well as subcubital fleck absent; 
medial vein sometimes absent and there are never more than 
three anal veins. Tibiae more or less flattened, fore 
usually with a crenulate outer edge, mid and hind ones with 
two borders, or rarely one border, bearing a row of setae or 
spinae. Femora with excavations for reception of base of 
tibiae. Tarsi 5-5-5, or rarely 4-4-4 (exclusively in 
Cybocephalinae), tarsomeres 1-3 bilobed or more rarely 
simple, tarsomere 4 (if present) smallest, a bisetose 
empodium developed between the claws (not always visible 
with usual optics). In the male genitalia, the tegmen 
consists of two lobes with a deep medial excision, and the 
penis trunk is more or less membranous and not quite 
dorsoventrally compressed (Carpophlin-lineage: Epuraeinae, 
Carpophilinae, Amphicrossinae, Calonecrinae); or the tegmen 
consists of a single plate, with or without a short apical 
excision, and the penis trunk is usually heavily sclerotized 
and dorsoventrally compressed (Nitidulinae, Meligethinae, 
Cillaeinae, Cryptarchinae, Cybocephalinae). 
Body elongate, subcylindrical or somewhat dorsoventrally 
compressed, slightly sclerotized, except hard sclerotized 
epicranium with appendages, legs, pronotal plate and small 
places on other tergites. Dorsum even or frequently with 
tubercles, processes, protuberances or prominences of 
different configuration, sometimes more or less sclerotized. 
XIth abdominal tergite with pregomphi and urogomphi 
(especially raised in the forms inhabiting enclosed 
substrates and reduced in free-living ones), lacking in 
Cybocephalinae. Head with frons fused with clypeus but 
usually with raised frontoclypeal suture, 2-4 stemmata on 
each side (if visible) and lack of endocarina; hypostomal 
roads absent, but hypostomal ridges strongly convergent 
posteriorly (except for Cybocephalinae with divergent ones); 
labroclypeal epipharynx furnished with a medial ridge and 
many small stripes, lacking in some anthophagous Epuraeinae, 
Meligethinae and Cybocephalinae; mandibles with raised mola 
and prosteca; maxillae with a mala (fused lacinia and galea) 
bearing a well developed membraneous or more or less 
sclerotized appendix and 3-joined palpus, cardines distinct; 
labial palpi 1-segmented; hypopharynx with a sclerome and 
bracons (except Cybocephalinae). Spiracles biforous and 
disposed on top of spiracular tube (if raised), abdominal 
ones situated dorsolaterally or rarely under lateral 
extensions of each segment (most Cybocephalinae have annular 
spiracles with 2 lateral air tubes). Legs rather short with 
sparse, fine, short and pointed setae, tarsungulus of free- 
living larvae sometimes with a subapical sensilar vesicle or 
capitate seta (Meligethinae; Nitidulinae, Mystropini and 
General shape more variable in comparison with that in 
larva, correlating with shape of imago. Head 
opisthognathous, completely covered with anterior part of 
pronotum and with few supraorbital tubercles bearing a fine 
long and pointed seta; frons usually with a separated and 
inflated clypeus. Pronotum along its fore edge has a pair or 
2-3 pairs of small and sharp tubercles with one long and 
pointed seta situated apically, subapically and basally, but 
in some cases tubercles reduced or disloged by comparatively 
long setae. Mesonotum, elytra and metanotum usually 
glabrous; medial part of hind edge of meso- and metanotum 
arcuately or angularly projecting. Hind femora and not 
infrequently mid ones with 1-3 subapical setae. Each 
abdominal tergites glabrous or often tergites I-IV with a 
paramedial pair of tubercles, but each laterosternite (of I- 
VIII) with 1-2 pairs of setose tubercles or setae between 
spiracles and lateral edge (one of them usually at each 
spiracle). 8th and 9th abdominal segments partly retracted, 
but caudal apex with a pair of long urogomphi (even in 
species with larvae lacking urogomphi at all - Meligethinae 
and Cybocephalinae). 
The Nitidulidae are traditionally regarded together 
with Brachypteridae (=Kateretidae) and Smicripidae either as 
members of the same family or as families of related groups. 
A comprehensive synopsis of structural characteristics of 
these groups can be found in reviews by Crowson (1955), 
Lawrence (1982, 1991), Kirejtshuk (1992) and Audisio (1993). 
The developed ventral plate fused or articulated with 
spiculum gastrale (anterior strut) in males and spiculum 
ventrale in females is a rather important structure. These 
sternal rudiments together with the anal sclerite 
(derivative of tergite VIII or partly laterosternites VIII) 
form the genital capsule, very characteristic in these 
families (Nitidulidae, Brachypteridae, Smicripidae). These 
families also have a tendency to reduce the imaginal galea 
(as in Rhizophagidae, whilst  many Polyphaga - like 
Boganiidae, Chrysomeloidea, Curculionoidea - more frequently 
show a tendendy to reduction of the lacinia); lack of 
functioning spiracles on 7th abdominal segment in imago; 
lack of lateral expansions (plates) of imaginal 
metendosternite (usual for archaic Cucujoidea, but also 
Cleroidea, Lymexyloidea, Tenebrionoidea, Chrysomeloidea and 
Curculionoidea); articulating maxillary mala (?=galea) with 
sclerotized appendix (?=lacinia) in larvae (as in many cases 
amongst Boganiiae, Helotidae, some Cleroidea, Chrysomeloidea 
and Curculionoidea, but sometimes in Lyctidae, Ptinidae, 
Peltidae, Lymexylidae, Endomychidae); 1-segmented labial 
palpi in larvae (occurs also amongst Lyctidae, Anobiidae and 
some Rhizophagidae); larval epicranium with not more than 4 
(usually 2 or 4) stemmata on each side (although this 
feature is also not unique amongst larvae of Polyphaga). 
Aedeagi of Nitidulidae can be Carpophiline or Nitiduline 
types, but always symmetrical, with fused tegmen (without 
"parameres") and dorsoventrally compressed penis trunk, 
whilst that of Brachypteridae and Smicripidae is 
asymmetrical and with "phallobase" and articulated 
"parameres" secondarily segmented. Larvae of representatives 
of Brachypteridae and Smicripidae are without distinct 
prostheca on their mandibles. In contrast to the family 
under consideration, the Brachypteridae are additionally 
characterized in imagines: by long narrow galea, reduced 
fold between elytral surface and epipleura (as only in 
Calonecrus species within Nitidulidae), large 6th abdominal 
segment and the next segment partly retracted, spiracles on 
1-6 abdominal segments of usual oval configuration and penis 
trunk well sclerotized and laterally depressed; in larvae: 
by head with developed endocarina; divergent hypostomal 
ridges; developed hypostomal rods and oval pronotum without 
sclerotized areas. The Smicripidae - in imagines: by well 
raised frontoclypeal suture, 2-segmented labial palpi, 
notosternal sutures rather distinct and last abdominal 
segment very long; in larvae: by parallel hypostomal rods 
(as in Laemophlaeidae) and without distinct cardines (as in 
Phalacridae and Cucujidae-Laemophlaeidae). 
The name Nitiduloidea proposed by Hieke (1989) is 
sometimes used in order to separate Nitidulidae together 
with Brachypteridae and Smicripidae from other groups of 
Cucujoidea (Audisio, 1993; Crowson, 1995). However, the 
families united in the Nitiduloidea sensu latter authors 
lack clear evidence of a closer common ancestor in 
comparison with other families of Cucujoidea sensu Lawrence 
and Newton, 1982 (see also Lawrence and Newton, 1995). All 
the characters used for this uniting (see above) are more or 
less due to structural simplification and could be developed 
without a close relationship, although these characters can 
be treated as forming a taxonomic syndrome. 
Keeping up the tradition mentioned above, the 
Nitidulidae were regarded as one of the oldest and most 
primitive group bearing many archaic characters from an 
ancestor of the infraorder Cucujiformia (Crowson, 1955, 
1981, 1990; Lawrence, 1982 and others). Nevertheless, there 
is some evidence (from palaeontological, morphological, 
ecological and bionomic data) for an alternative 
interpretation (Kirejtshuk, 1994a, 1996b, see also below). 
The simpler appearance of some Epuraeinae, Carpophilinae, 
Meligethinae and others in contrast to the opinion of 
Crowson (1988) is connected rather with a faster ontogenetic 
development than with the archaic character of their 
structures. It can be recognized that the following 
characters seem to have been inherited from ancestral forms 
of the group under consideration: 
IMAGO: oval body slightly convex dorsally and ventrally with 
sculpture, punctation and pubescence somewhat like those in 
Soronia Erichson, 1843, Ericmodes Reitter, 1877/1878, 
Lophocateretes Olliff, 1883, Zimioma des Gozis, 1886, Ostoma 
Laicharting, 1781 and Thymalus Latreille, 1802; prognathous 
head with bilobed free labrum, raised mandibular mola and 
prostheca, unilobed maxilla; widely explanate pronotal and 
elytral sides; elytra complete, with wide and complete 
epipleura; fore coxae widely separated by comparatively wide 
prosternal process, which is far projecting as a fold on 
mesosternal surface; metasternum with well raised medial and 
paracoxal sutures; metacoxae transverse, but not medially 
inclined as those in Cleroidea (including probable 
Peltoidea); all trochanters of the cucujoid (normal) or 
nearly tenebrionoid ("heteromeran") types; fore tibia with 
crenulate outer edge, but mid and hind tibiae with 2 borders 
bearing setae along outer edge; apices of all tibiae with a 
pair of spurs; tarsi with tarsomeres 1-3 lobed and tarsomere 
5 longest and ending with distinctly bisetose empodium 
between claws; anal sclerite completely retracted into 8th 
abdominal segment in both sexes; ventral plate in males 
divided into 2 parts joined by spiculum gastrale; slightly 
sclerotized tegmen considerably surrounding the slightly 
sclerotized and dorsoventrally compressed penis trunk (as in 
Axyra Erichson, 1843, Megauchenia Macleay, 1825, Prometopia 
Erichson, 1843, Platychora Erichson, 1843 and so on); fork- 
sclerite articulated with tegmen; penis trunk with unpaired 
apodema at base and paired lobes closing subapical orifice. 
LARVA: body elongate, slightly sclerotized, except for hard 
sclerotized epicranium with appendages, pronotal plate and 
small areas on other tergites; head with frons fused with 
clypeus; labroclypeal epipharynx furnished with a medial 
ridge and many small strips; mandibles with raised mola and 
prostheca; maxillae with a mala and 3-joined palpi; labial 
palpi 1-segmented; hypopharynx with a sclerome and bracons; 
spiracles disposed on top of spiracular tube. 
The family has at least 3 000 published names for 
presumably valid species ranged into 271 genera and 
subgenera, although the expected number of species in the 
recent fauna of this group should be estimated at over 7000 
- 8000, including not less than 2 000 from the Indo-Malayan 
and 500 from the Palaearctic regions. The family is supposed 
to consist of 9 subfamilies united in 2 lineages represented 
by groups probably with a common phylogenetic ancestry. Some 
of the subfamilies are divided into tribes, and the latter 
correspondingly into complexes of genera. Almost all groups 
with a rank of subfamily and tribe are represented in the 
treated region (except Cychramptodini Kirejtshuk, Lawrence, 
1992  and Lawrencerosini Kirejtshuk, 1990c - from the 
Nitidulinae, endemic for the Australian region; Arhinini 
Kirejtshuk, 1987b - from Cryptarchinae, endemic for 
Afrotropical region; and Mystropini Murray, 1864 - from 
Nitidulinae, endemic for Neotropical region), although 
Calonecrinae are as yet known only from the southern part of 
Indochina southwards. A more detailed explanation of the 
composition of the family and the historic development of 
the lineages and subfamilies are given in Kirejtshuk, 1994a 
(and also in Kirejtshuk, 1982, 1986c, 1992; notes on 
suprageneric taxa and respective references: Pakaluk, 
S'lipin'ski, Lawrence, 1994; Lawrence, Newton, 1995). The 
following group designations with a taxonomic fixation as 
subfamily or tribe will be used in the present monograph 
(number of genera and subgenera represented on the territory 
under consideration given in parentheses): 
1. Epuraeinae Kirejtshuk, 1986c: 27 [Epuraeini (- 11) and 
Taenioncini new tribe (- 4)]; 
2. Carpophilinae Erichson, 1842: 148 (- 10); 
3. Amphicrossinae Kirejtshuk, 1986c: 28 (- 1); 
4. Calonecrinae Kirejtshuk, 1982: 117 (- 1); 
5. Nitidulinae Latreile, 1802: 132 [Nitidulini (- 29), 
Strongylini Sturm, 1844: 7 (- 8), Cychramini Lacordaire, 
1854/1855: 318 (- 2)]; 
6. Meligethinae C.G. Thomson, 1859: 67 (- 7); 
7. Cillaeinae Kirejtshuk and Audisio in Kirejtshuk, 1986d: 
219 (- 4); 
8. Cryptarchinae C.G. Thomson, 1859: 69 [Cryptarchini (- 3), 
Platyarchini new tribe (- 1), Eucalosphaerini Kirejtshuk, 
1987b: 63, 80 (- 1)]; 
9. Cybocephalinae Jacquelin du Val, 1858 (- 2). 
The Nitidulidae consist of groups which are rather 
diverse not only in structure but also in their ecology, 
trophics and mode of life. Diversity of mode of life in 
different groups is less developed on territories with 
temperate climate and high mountain elevation but becomes 
extremely wide in regions with subtropical and tropical 
climates. Such regularity will be analyzed in detail in the 
last part of this monograph. 
Nevertheless it should be here emphasized that the most 
groups of the family have close connections with woody 
ecosystems adhering to trees and bushes. In particular it 
can be observed in characteristic of the Nitidulid fauna 
revealed on the territory under consideration. The 
subfamilies Epuraeinae, Amphicrossinae, Calonecrinae and 
Cryptarchinae are specific forest groups, whilst only most 
Carpophilinae, Nitidulinae and Cillaeinae live in forests of 
different types and are connected with fungi developing on 
tissues of trees and bushes, oozing tree sap and substrates 
like that. Not infrequently species of Pocadius Erichson, 
1843; Thalycra Erichson, 1843; Quadrifrons Blatchley, 1916; 
Thalycrodes Blackburn, 1891 from Nitidulinae connected with 
subterraneous fungi live also in forest. Even the groups 
inhabiting on flowers or fruits often occur on alive or dead 
trees and bushes (tribe Mystropini and others). The 
parazitoid Cybocephalinae and Cychramptodini from 
Nitidulinae are mostly represented in ecosystems with trees 
and bushes where colonies of coccids and white flies exist. 
Finally, many of forms bred on herbaceous plants prefer to 
exist within forest communities (subfamily Meligethinae and 
others). However, necrophagous species of Nitidula 
Fabricius, 1775 and Omosita Erichson, 1843 sensu lato 
ussualy are more common beyond forests. And only few genera 
can be regarded as groups characteristic for open grass 
ecosystems (such as many desert and steppe groups of the 
genus Meligethes Stephens, 1832, sensu lato from 
Meligethinae and cactivorous species of the genus Camptodes 
Erichson, 1843 from Nitidulinae]. 
Most Nitidulidae are more regularly collected in 
conditions of temperate and subtropical climate in the North 
Hemisphere (including usually in mountain forest of the 
territory under consideration) during middle and late spring 
or very early summer (March-June). However, in the mentioned 
circumstances, some of species are more frequent rather 
within or at autumn, but the groups with both type activity 
have, as a rule, intervals in occurance during summer and/or 
winter. Another pattern of activity can meet in tropical 
fainforest or in condition without sharply expressed 
differences in seasons, although at the present it is 
impossible to trace more or less regular types of this 
pattern to do any grouping of different faunistic 
components. Lowland rainforest is inhabited by some 
Nitidulid forms showing imaginal activity the year round, 
usually these forms are associated with such habits as under 
bark of trees, flowers and soft fruits. 
The most ancient habits or those similar to them appear 
to be amongst the fungivorous unspecialized forms from the 
different groups of both Nitidulidae and the infraorder 
Cucujioformia in general (Peltidae, Lophocateridae, 
Phloiophilidae from superfamily Cleroidea; Derodontidae and 
Nosodendridae from superfamily Dermestoidea; many families 
from superfamilies of Cucujoidea and Tenebrionoidea). In 
particular an archaic mode of life is probably characterized 
by the nitidulid species, inhabiting exuding tree sap in the 
genus Epuraea sensu lato (Epuraeinae); most Amphicrossinae; 
Calonecrinae; genus Soronia (Nitidulinae) and subgenus 
Glischrochilus (Librador) Reitter, 1884 (Cryptarchinae). 
These features appear also to be attributed to many 
representatives of different subgenera of the genus Epuraea 
sensu lato (subgenera Epuraea sensu stricto; Epuraeanella; 
Aphenolia Reitter, 1884; Africaraea Kirejtshuk, 1989a; 
Amedanyraea Kirejtshuk et Pakaluk, 1996) from Epuraeinae; 
many or all representatives of subgenera Carpophilus 
(Carpophilus) Erichson, 1843 and C. (Ecnomorphus) 
Motschulsky, 1858 from Carpophilinae; as well as to most 
Strongylini (except for species of Camptodes) from 
Nitidulinae and most Cryptarchini from Cryptarchinae. Many 
mycetophagous forms from Nitidulini and Cillaeinae maintain 
to a greater or lesser extent some elements of an archaic 
mode of life and appropriate habits. It is particularly true 
for many arboricolous nitidulins from genera Prometopia; 
Parametopia Reitter, 1884; Lobiopa Erichson, 1843; Axyra; 
Megauchenia; Ipidia Erichson, 1843; Platychora; Taracta 
Murray, 1867; Psilotus Fischer, 1829; Perilopa Erichson, 
1843; Gaulodes Erichson, 1843; Ussuriphia Kirejtshuk, 1992; 
as well as some representatives of genera Pocadites Reitter, 
1884; Hebasculinus Kirejtshuk, 1992; Atarphia Reitter, 1884 
and subgenera of Aethina (Aethina) Erichson, 1843 sensu 
stricto; A. (Circopes) Reitter, 1873 and Lordites (Phenolia) 
Erichson, 1843 from Nitidulini; and probably species of the 
genera Ecnomaeus Erichson, 1843; Cillaeus Castelneau, 1835; 
Cillaeopsis Grouvelle, 1899; Platynema Ritsema, 1885; 
Ithyphenes Murray, 1864; and at least part of members of 
genera Colopterus Erichson, 1842; Brachypeplus Erichson, 
1842; Conotelus Erichson, 1843 from Cillaeinae. To a lesser 
extent it is true for the forms completely or partly 
attached to litter and decomposing substrates near or in the 
soil and sometimes to decaying fruits and seeds, such as 
representatives of Urophorus (Urophorus) Murray, 1864 sensu 
stricto; U. (Anophorus) Kirejtshuk, 1990b; Stelidota 
Erichson, 1843; Pocadius; Lordites (Lordites) sensu stricto; 
L. (Aethinodes) Blackburn, 1891 and L. (Plesiothina) 
Kirejtshuk, 1990a; Thalycra; Quadrifrons; Thalycrodes; and, 
perhaps, some Physoronia Reitter, 1884  from Nitidulini. 
Most fungivorous Nitidulidae are recorded as breeding 
in fermenting substrates with yeasts and might well serve as 
vectors of the yeasts. However, many representatives of 
different subfamilies prefer or are connected with 
basydiomycete fruiting bodies [such as some Epuraea 
(Epuraea) sensu stricto; E. (Aphenolia); E. (Epuraeanella); 
Lordites (Phenolia); Pocadius; Pocadites; Thalycra; 
Cyllodes; Pallodes Erichson, 1843; Neopallodes; Tricanus 
Erichson, 1843; Oxycnemus Erichson, 1843]. Some Epuraea 
(Haptoncus), Carpophilus sensu lato, Colopterus, 
Glischrochilus sensu lato and others are involved in 
transmission of Ceratocystis (Ascomycetes) or Fusarium 
It is possible to trace some different and regular of 
changes in ecology, trophics and mode of life, and 
appropriate transformations of structure (Kirejtshuk, 1989b, 
1996b). The most expressive correlation between ecological, 
bionomical and structural traits is in the groups which 
independently became anthophagous and using pollen as a main 
food resource (Epuraeinae, Carpophilinae, Nitidulini, 
Strongilini, Cychramini, Cillaeinae, all Mystropini and 
Meligethinae - Kirejtshuk, 1994a, 1996a). Not infrequently, 
inflorescence gives an intermediate inhabitation for the 
forms with a tendency to phytophagy [as that amongst some 
recent representatives of Epuraea (Micruria) Reitter, 1875a 
(Hayashi, 1978); E. (Haptoncurina) and Parepuraea Jeli'nek, 
1977 (considered below and Kirejtshuk, unpublished) from 
Epuraeinae; different subgenera of Carpophilus (Connell, 
1956) from Carpophilinae; Neopocadius Grouvelle, 1906c 
(Bruch, 1923) and Camptodes from Nitidulinae; and some 
Brachypeplus sensu lato (Kirejtshuk, 1994a, 1996a) from 
Cillaeinae, although other relatives of the mentioned groups 
yet became completely anthophagous]. Many nitidulids, having 
overcome this stage of ecological change and become 
completely anthophagous at both larval and imaginal instars, 
remain at this stage of regular ecological change until now 
[Propetes sensu lato and Epuraea (Apria) Grouvelle, 1919 
(Jeli'nek, 1992) from Epuraeinae; subgenera Carpophilus 
(Caplothorax) Kirejtshuk, 1996b and C. (Plapennipolus) 
Kirejtshuk, 1996b; as well as subgenus Urocarpolus 
Kirejtshuk, 1996b of genus Nitops Murray, 1864 from 
Carpophilinae; all Meligethinae; subgenera Aethina (Ithyra) 
Reitter, 1873; A. (Olliffura) Jeli'nek and Kirejtshuk in 
Kirejtshuk, 1986a; A. (Cleidorura) Kirejtshuk and Lawrence, 
in press from Nitidulini (Nitidulinae); probably, all 
Mystropini (Nitidulinae) and Macrostola from Cillaeinae]. 
The next stage in the mentioned regularity of changes in 
mode of life and trophics is a transition from anthophagy to 
carpophagy (Epuraeinae and Carpophilinae) or complete 
phyllophagy (Anister Grouvelle, 1901; Xenostrongylus 
Wollaston, 1854; Strongyllodes Kirejtshuk, 1992 from 
Cychramini, Nitidulinae). 
Species of Nitidula Fabricius, 1775 and Omosita 
Erichson, 1843 sensu lato are associated with fungi growing 
on carcasses of vertebrates and became rather usual 
inhabitants of places with garbage, agricultural and 
industrial refuse with remains of animals. Few species of 
Epuraea (Epuraea) sensu stricto are accustomed to live in 
conditions of deserts and other arid territories finding 
fungi to eat under faeces of mammals or in burrows of 
rodents. Some mycetophagous Amphicrossus and Amphotis not 
infrequently have symbiotic relations with ants (African 
Amphicrossus parallelus Grouvelle, 1912 described in genus 
Nitidopecten Reichensperger, 1913; and european species of 
Amphotis often occur in nests of Lasius Fabricius, 1805 - 
Hymenoptera, Formicidae). However, development of Amphotis 
marginata (Fabricius, 1781) is also recorded in galls with 
Biorhiza pallida (Olivier, 1791) (Hymenoptera, Cynipidae) 
(Lengerken, 1941). The tribe Lawrencerosini, as far as 
known, is completely myrmecophilous (Kirejtshuk, 1990c and 
unpublished). Some African representatives of Aethina 
(Aethina) sensu stricto regularly live in nests of bees [A. 
(A.) tumida Murray, 1867 is recorded in nests of honey bees 
- Lundie, 1940 and its relatives are collected in nests of 
other Apidae (Kirejtshuk, unpublished)]. Australian Onicotis 
Murray, 1864 and some Australian Brachypeplus 
(?Brachypeplus) sensu stricto are also associated with 
Some arboricolous forms, being related to mycetophagous 
ones, became facultative or obligatory predators of insect 
larvae and other soft invertebrates living under bark and 
wood, sometimes becoming as regular inhabitants of holes of 
Scolytidae [some Epuraea (Epuraea) sensu stricto; 
Glischrochilus (Glischrochilus) Reitter, 1873 sensu stricto; 
Pityophagus Shuckard, 1839 and others]. However, only for a 
few groups the can predation be regarded as a taxonomic 
peculiarity. Amongst them the predators on scale insects - 
many Cychramptodini from Nitidulinae and most Cybocephalinae 
(without trace of close kinship or evident similarity) have 
particular placements (Kirejtshuk, Lawrence, 1992). Some 
species of subgenus Cybocephalus (Cybocephalus) Erichson, 
1844 sensu stricto are recorded as predators of whiteflies 
(Kirejtshuk, James, Heffer, in press and unpublished). 
The Nitidulidae are characterized by a rather short 
larval development and comparatively long-lived imagines, 
but both instars are very shortly active in contrast to many 
coleopterous groups, except for some groups from the 
superfamily Cucujoidea. Namely this circumstance allows to 
the Nitidulidae to master and to be accustomed to extremely 
different types of substrate, frequently existing within a 
comparatively short period (as oozing tree sap, flowers with 
short duration of blossoming and so on). 
Ponomarenko (1983) mentioned that the appearance of 
different Cucujoid families of beetles began at the end of 
the Lower Cretaceous. The mesozoic Nitidulidae, as well as 
other members of the superfamily Cucujoidea have been 
recorded exclusively from different layers of the 
Cretaceous, increasing in number to the end of this period 
(Ponomarenko, 1983; Dmitriev and Zherikhin, 1988; Kirejtshuk 
and Ponomarenko, 1990). The fossil data on Nitidulidae 
remains poorly known because of difficulty of investigation. 
Many references on Kainozoic deposits of Nitidulidae should 
be restudied to clarify their position, whilst some 
references on Mesozoic deposits (Martynov, 1926; Medvedev, 
1969; Audisio, 1993) should be recognized as erroneous, and 
only the forms listed in Kirejtshuk and Ponomarenko (1990); 
Kirejtshuk (1994a) can be considered as true representatives 
of Nitidulidae [only palaeoendemic genera Crepuraea 
Kirejtshuk, 1990, Cyllolithus Kirejtshuk, 1990 (both in 
Kirejtshuk & Ponomarenko, 1990)]. At the same time the 
author knows many remains from the Cretaceous deposits of 
Kazakhstan in which it is easy to find the characteristic 
traits of the subfamily Epuraeinae, but there is no 
character to propose for them a taxon with both generic and 
species names. Unfortunately, no record of Nitidulid fossils 
has been published from the territory under consideration. 
Therefore the author can outline only the recent historic 
development which can be traced mainly after a study of 
modern distribution of the groups (which will be reviewed in 
the last part of the present monograph). 
Diversification of the Cucujoidea seemed to arise and 
to be proceeding when the characteristic mesozoic groups of 
plants became more and yet more rare, until they were 
dislodged by the newly appeared Angiosperm plants. Thus, 
evolution of the Cucujoidea is, perhaps, associated with 
development of the Kainophytic flora, even though this 
coleopterous group could take its origin somewhat earlier. 
Interconnections between the Cucujoidea and Kainophytic 
plants were initially mediated through fungi. At the end of 
the Cretaceous, the closer and more intimate 
interconnections seemingly began to establish at first with 
generative organs of both Gymnosperms and Angiosperms and 
further on with other plant organs. This process of 
ecological changes in the Kainozoic Coleoptera is clearly 
reflected amongst the Nitidulidae. Having admitted the 
mentioned argumentation Nitidulidae should be regarded as 
rather advanced and one of the youngest coleopterous groups 
of all (Kirejtshuk, 1994a and others), not archaic as it was 
treated in many previous interpretations. 
Fossil faunas of Nitidulidae at the beginning of the 
Eocene are better documented, but mainly for Europe. In the 
Baltic amber there are represented recent genera from 4 
subfamilies (Hieke, Pietrzeniuk, 1984): Epuraeinae (Epuraea, 
Epuraeanella), Carpophilinae (Carpophilus), Nitidulinae 
(Cyllodes), Cryptarchinae (Cryptarcha Shuckard, 1839), and 
only the Omositoides Schaufuss, 1891 was described as a new 
genus from amber records. Species from the Holarctic 
deposits of Oligocene and Miocene have been attributed to 
recent genera, except for Epanuraea Scudder, 1892, 
Cychramytes Wickham, 1913, Miophenolia Wickham, 1916 and 
Oligamphotis Theobald, 1937. The only quartenary 
Protocarpophilus macgillavryi De Jong, 1953 was described 
from Sumatra. 4 subfamilies represented amongst the forms 
from Baltic amber, the Cillaeinae are known from the 
Oligocene (Wickham, 1913) and Cybocephalinae - from the 
Miocene (Palmer, 1957). However, most records of Kainozoic 
Nitidulidae should be most thoroughly revised before a more 
detailed interpretation of the chronicles of this family is 
The family under consideration has world wide 
distribution, but ranges of its groups have some 
restrictions. As a general rule there is an asymmetrical 
distribution of the Carpophilin- and Nitidulin-lineages, the 
first showing most diversity and fullest representation in 
the Eastern Hemisphere (without any generic endemism in 
South America and Hawaii), whilst the second has a much more 
raised diversification in the Western Hemisphere (with 
highest diversity in the Neotropical region and Hawaii). A 
characteristic of the territory and fauna under 
consideration, lies in the fact that the Palaearctic and 
Indo-Malayan faunas meet here (in many groups between 
elevations 2 000-3 000 m above sea level). Except some 
endemic suprageneric taxa listed above, the following 
features of distribution of the fauna can be noted before a 
detailed analysis of distribution in the last part of this 
monograph (including new taxa which will be described in 
further parts of the monograph): 
I. Exclusively endemic Indo-Malayan supraspecific taxa: 
- Epuraeinae, Epuraeini - Epuraea (Ommoraea) new subgenus, 
E. (Ceroncura), Grouvellia, Tetrisus (Tetrisus) sensu 
- Carpophilinae - Ctilodes Murray, 1864, Vulpixenus 
Kirejtshuk, 1990a; 
- Meligethinae - Cryptarchopria Jeli'nek, 1975b; Meligethes 
(Cyclogethes) Kirejtshuk, 1979a; Kabakovia Kirejtshuk, 1979a; 
- Nitidulinae, Nitidulini - Parametopia; Taraphia; 
Pseudoischena Grouvelle, 1897=Megauchenioides Audisio et 
Jeli'nek, 1993; 
- Nitidulinae, Strongulini - Viettherchnus Kirejtshuk, 1985; 
- Cryptarchinae, Cryptarchini - Glischrochilus 
(Gymnoparomius) Kirejtshuk, 1987b; 
- Cybocephalinae - Taxicephomerus Kirejtshuk, 1994c; 
II. Taxa with principal distribution in the Indo-Malayan 
region and East Chinese (Palaearchearctic) province of the 
Palaearctic region: 
- Epuraeinae, Epuraeini - Epuraea (Micruria); 
- Meligethinae - Meligethes (Meligethes) sensu stricto; 
- Nitidulinae, Nitidulini - Ipidia (Hemipidia) Kirejtshuk, 
1992; Ussuriphia; Physoronia; Atarphia; Pocadites; 
- Nitidulinae, Strongulini - Neopallodes; 
III. Taxa with wide distribution and which are the most 
abundant taxa in the territory under consideration: 
- Carpophilinae - Carpophilus (Ecnomorphus); 
- Nitidulinae, Nitidulini - Soronia (Soronia) sensu stricto; 
- Nitidulinae, Strongylini - Cyllodes; 
- Cryptarchinae, Cryptarchini: Glischrochilus (Librodor); 
IV. Taxa and groups of species with endemism or widest 
distribution in the Eastern Hemisphere (mostly 
tropicopolitous, except for Cybocephalus sensu lato): 
- Carpophilinae: obsoletus-group and hemipterus-group of 
Carpophilus (Carpophilus) sensu stricto; 
- Nitidulinae, Nitidulini - Aethina (Circopes); 
- Nitidulinae, Cychramini - Strongyllodes; 
- Cybocephalinae - Cybocephalus sensu lato; 
V. Taxa sharing endemism or most diversity in the Indo- 
Malayan, Papuan, Australian and Novacaledonian regions: 
- Epuraeinae, Epuraeini - Epuraea (Haptoncurina), E. 
(Haptoncus), Propetes sensu lato, Tetrisus (Trimenus); 
- Epuraeinae, Taenioncini new tribe - Taenioncus, 
Taeniolinus new genus, Carpocryraea new genus; 
- Nitidulinae, Nitidulini - Megauchenia, Lordites 
(Plesiothina), Aethina (Olliffura); 
- Nitidulinae, Strongylini - Pallodes sensu lato; 
VI. Taxa sharing endemism or most diversity in the Indo- 
Malayan, Palaearctic and Nearctic regions: 
- Epuraeinae, Epuraeini Epuraea (Epuraea) sensu stricto, E. 
- Carpophilinae - Carpophilus (Megacarpolus) Reitter, 1919; 
- Nitidulinae, Strongulini - Oxycnemus; 
- Cryptarchinae, Cryptarchini - Glischrochilus (Librodor); 
VII. Taxa sharing endemism or most diversity in the Indo- 
Malayan, Afrotropical, Capean and Madagascarean regions: 
- Epuraeinae, Taenioncini new tribe - Raspinotus; 
- Carpophilinae - Urophorus (Urophorus); 
- Meligethinae - Meligethinus Grouvelle, 1906c; 
- Nitidulinae, Nitidulini - Axyra, Lordites (Lordites) sensu 
stricto, Aethina (Aethina), Anister; 
- Cillaeinae - Ecnomaeus; 
VIII. Taxa with wide distribution (including the Indo- 
Malayan region), but comparatively weakly represented in the 
territory under consideration: 
- Meligethinae - the subfamily in general, including Pria; 
Meligethes (Clypeogethes) Scholtz, 1932; 
- Nitidulinae, Nitidulini - Nitidula, Omosita, Thalycra- 
complex of genera; 
- Cillaeinae - the subfamily in general and, in particular, 
Colopterus, Cillaeus, Platynema; 
IX. Taxa with wide distribution (including surrounding 
areas), but absent in the territory under consideration: 
- Carpophilinae - Urophorus (Anophorus); 
- Meligethinae - Meligethes (Astylogethes) Kirejtshuk, 1992; 
- Nitidulinae, Nitidulini - Amphotis; 
- Cillaeinae - Ithyphenes; 
- Cryptarchinae, Cryptarchini - Pityophagus. 
1 a. Antennae 10-segmented with 1-segmented club and a lens- 
like preceding segment dorsoventrally undepressed; elytral 
epipleura having no lateral fold at curvature from dorsal 
side; elytra with subapical excision at outer corner; 
pygidium and part of preceding segment remaining uncovered 
by elytra; body somewhat flattened dorsally with 
particularly flattened hexagonal pronotum and convex 
ventrally; head elongate with rather projecting mandibles; 
body bright reddish with black elytra and with partly 
darkened antennae and legs, comparatively large - 5.5-11.7 
mm. Known only from Malacca peninsula and island systems of 
the Indo-Malayan region . . . . . . . subfamily Calonecrinae 
(genus Calonecrus J. Thomson, 1857 - C. wallacei J. Thomson, 
1857: Malaysia, Perak and Sarawak; Indonesia, Java) 
1 b. Antennae usually 11-segmented or rarely 9-10-segmented 
with 2-8-segmented club more or less dorsoventrally 
depressed; elytral epipleura with a distinct fold at least 
in basal part of elytra; elytra without any subapical 
excision at outer corner, complete or remaining part of 
tergites (up to 4) uncovered; body usually moderately convex 
dorsally and ventrally with more or less rounded pronotal 
sides; infrequently less than 5.5 mm.  . . . . . . . . . . . 
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 
2 (1) a. Tarsi 4-4-4; body small (usually 0.8-1.5, rarely up 
to 2.6 mm), hemispheric, rolling up in a ball; dorsum 
usually smooth and shiny with a reduced fine punctation; 
elytral epipleura reduced, downwardly sloping laterally 
(almost vertically); abdomen with 5 pairs of acting 
spiracles . . . . . . .. . . . . . . . . Cybocephalinae 
2 (1) b. Tarsi 5-5-5; body, as a rule, more than 1.0 mm, 
normally elongate or oval, if hemispheric (small forms never 
hemispheric), incapable of rolling up in a ball; dorsum 
usually with clear punctation, infrequently rather shallow 
but quite coarse; elytral epipleura different (normally 
upwardly sloping laterally), but never almost vertically 
sloping; abdomen with 6 pairs of developed spiracles . . . . 
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 
3 (2) a. Elytra sharply shortened 1-3 tergites uncovered 
before pygidium; body elongate, pronotum and elytra jointly 
almost always transverse; all uncovered tergites heavily 
sclerotized  . . . . . . . . . . . . . . . . . . . . . . . 4 
3 (2) b. Elytra complete or shortened, leaving uncovered at 
most the pygidium and part of preceding tergite; body more 
or less oval, rarely rather elongate; all tergites, except 
pygidium, membranous or slightly sclerotized (Taenioncini 
trib.n. from Epuraeinae characterized by rather sclerotized 
tergite preceding pygidium and in some representatives this 
tergite completely uncovered by elytra; representatives of 
this tribe have rather convex body with normally both 
pronotum and elytra jointly or at least elytra, longer than 
combined width and not explanate at sides)  . . . . . . . 5 
4 (3) a. Tergites uncovered by elytra flat, their combined 
length (with pygidium) usually much more than that of 
pronotum (excepting Colopterus with oval flattened body and 
uncovered tergites with their combined length subequal to 
that of pronotum); abdominal pleura bent dorsally, looking 
like wide stripes with a sharp fold to ventrites; body 
usually strongly flattened; anal sclerite of male directed 
posteriorly and tegmen unilobed or sometimes with a short 
medial excision at apex . . . . . . .  subfamily  Cillaeinae 
4 (3) b. Uncovered tergites more or less convex, their 
combined length with pygidium, as a rule, not more than that 
of pronotum; abdominal pleura narrow, frequently almost 
invisible, gently bent on ventral side; body frequently 
moderately convex dorsally and ventrally [with some 
exceptions mainly amongst species of the subgenus C. 
(Ecnomorphus)]; anal sclerite of male turned ventrally and 
tegmen bilobed with an excision dividing it almost along 
whole length . . . . . . . . . . . . subfamily Carpophilinae 
5 (3) a. Labrum fused with frons, usually with a trace of 
this fusion as a suture or remains of it (subfamily 
Cryptarchinae) . . . . . . . . . . . . . . . . . . . . . . 6 
5 (3) b. Labrum free, sometimes concealed under fore part of 
frons . . . . . . . . . . . . . . . . . . . . . . . . . .  8 
6 (5) a. Prosternal process strongly widened before apex and 
at least half as wide as head; coxae in each corresponding 
pair widely separated each from other, distance between mid 
coxae not less than that between hind ones, comprising about 
half width of first ventrite; fore corners of mentum sharply 
projecting forwards; all femora strongly widened at 
trochanter; pronotal base without any trace of border and 
looking like a fold covering most of scutellum and elytral 
base; body oval and strongly convex dorsally, almost 
hemispheric . . . . . . . . . . . . . . . . . tribe 
Eucalosphaerini (genus Eucalosphaera Jelinek, 1978) 
6 (5) b. Prosternal process not widened, slightly or 
moderately widened before apex (only in Platyarcha the 
process rather strongly widened before apex); coxae in each 
corresponding closer to each other, distance between mid 
coxae much less than that between hind ones; fore corners of 
mentum blunt, not projecting forwards (only in Platyarcha 
are the fore corners sharply acute and projecting forwards); 
femora weakly or moderately widened at trochanter; pronotum 
almost always with a border (sometimes reduced medially) and 
not looking like a fold covering scutellum and elytral base; 
body flattened or moderately convex dorsally, or rarely 
rather strongly convex  . . . . . . . . . . . . . . . . . 7 
7 (6) a. Moderately and weakly convex ventrally and 
dorsally; antennal club well and normally developed (quite 
compact); elytra with narrowly explanate sides and rounded 
(not truncate) apices frequently forming a continuous arc; 
mentum with fore corners not projecting forwardly; 
prosternal process of a usual outline; distance between fore 
coxae less than that between both mid and hind coxae . . . . 
. . . . . . . . . . . . . . . . . . . . . tribe Cryptarchini 
7 (6) b. Rather flattened dorsally and ventrally; antennal 
club comparatively small, nearly 2-segmented with 9th 
segment weakly widened anteriorly; elytra with widely 
rounded sides and truncate apices; mentum with fore corners 
strongly projecting forwards; prosternal process strongly 
widened before truncate apex; the distance between mid coxae 
less than that between both fore and hind coxae . . . . . . 
. . . . . . . . . . . . . . . . . . . . . . . . Platyarchini 
new tribe (type-genus: Platyarcha Kirejtshuk, 1987b) 
8 (5) a. Mid and hind tibiae strongly depressed 
dorsoventrally and with one outer border bearing setae or 
marked hairs different from those on remainder of these 
structures; pygidial base with a pair of very wide arc-like 
depressions, usually partly covered by preceding tergite . . 
. . . . . . . . . . . . . . . . . . . subfamily Meligethinae 
8 (5) b. Mid and hind tibiae not so strongly depressed 
dorsoventrally and usually with 2 outer borders bearing 
setae or marked hairs different from those on remainder of 
these structures; pygidial base without a pair of very wide 
arc-like depressions (or with 8 small arc-like ones along 
its edge and usually partly covered by preceding tergite). . 
. . . . . . . . . . . . . . . . . . . . . . . . . . . . .  9 
9 (8) a. Dorsal punctation always diffuse; pubescence more 
or less visible; body elongate or, if oval, moderately 
convex dorsally; pronotum never bordered at base; male: anal 
sclerite far exposed posteriorly from under truncate or 
subtruncate pygidial apex or hypopygidium, with a large 
movable lobe before apex; tegmen deeply excised into two 
lobes . . . . . . . . . . . . . . . . . . . . . . . . . 10 
9 (8) b. Dorsal punctation, pubescence, pronotal base and 
body shape diverse; male: anal sclerite normally unexposed 
or slightly exposed from under not truncate pygidial apex 
(only in Neopallodes pygidial apex truncate and anal 
sclerite exposed comparatively far posteriorly, but body 
strongly convex dorsally and glabrous); hypopygidium without 
any distinct movable lobe; tegmen unexcised or shallowly 
excised at apex (subfamily Nitidulinae) . . . . . . . . . 12 
10 (9) a. Body widely oval and usually larger (at least 3.5 
mm), flattened ventrally, evenly and moderately convex 
dorsally; pronotal and elytral sides with long dense ciliae; 
male: anal sclerite not exposed from under pygidial apex; 
hypopygidium with a large movable lobe before apex . . . . . 
. . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . 
subfamily Amphicrossinae (genus Amphicrossus Erichson, 1843) 
10 (9) b. Body usually elongate and very rarely larger than 
3.5 mm; as a rule, moderately convex dorsally and ventrally; 
pronotal and elytral sides without distinct ciliae or very 
finely and shortly ciliate; male: anal sclerite clearly 
exposed (except few cases) from under pygidial apex; 
hypopygidium without any movable lobe before apex (subfamily 
Epuraeinae) . . . . . . . . . . . . . . . . . . . . . . . 11 
11 (10) a. Elytra with truncate apices leaving pygidium and 
almost all of preceding tergite uncovered; body elongate, 
subparallesided and rather convex dorsally, with very 
narrowly explanate pronotal and elytral sides; pubescence 
short or reduced (up to invisible) . . . . . . . . . . . . . 
. . . . . . . . . . . . . . . . . .  Taenioncini new tribe 
11 (10) b. Elytra with various configurations of apices, 
covering abdomen completely or leaving exposed only pygidium 
or in few cases also part of preceding tergite; pubescence 
moderately developed or slightly reduced . . . . . . . . . . 
. . . . . . . . . . . . . . . tribe Epuraeini sensu stricto 
12 (9) a. Body strongly convex dorsally with unexplanate 
pronotal and elytral sides and head somewhat inclined 
ventrally; dorsal surface glabrous (pygidium finely 
pubescent, exceptionally); pronotum diffusely punctate and 
with unbordered base partly covering scutellum and elytral 
bases; prosternum more or less shortened; male: tegmen with 
an unpaired long lobe, sometimes shallowly excised at apex; 
female: ovipositor with a sharply acute apex without styli, 
rarely rather long with slightly modified sclerites, which 
often have styli . . . . . . . . . . . . . tribe Strongilini 
12 (9) b. Body diverse, but never with same combination of 
all features as mentioned above; dorsal surface in most 
cases with quite visible pubescence (usually well 
developed); pronotum with various punctation and, as a rule, 
with a clear border along base; prosternum not shortened or 
a little shortened (in species of tribe Cychramini); male: 
structure of aedeagus various; female: ovipositor, if 
strongly modified, in a different way . . . . . . . . . 13 
13 (12) a. Body rather evenly convex from above with 
unexplanate pronotal and elytral sides, head somewhat 
inclined ventrally; dorsal surface strongly pubescent; 
pronotum diffusely punctate with an unbordered base partly 
covering scutellum and elytral bases; prosternum more or 
less shortened with a rather short intercoxal process; 
female: ovipositor with blunt or truncate and nearly 
membranous apex without or with clearly reduced styli . . . 
. . . . . . . . . . . . . . . . . . . . . . tribe Cychramini 
13 (12) b. Body diverse, but never with the same combination 
of all features as mentioned above; prosternum not or 
slightly shortened; female: ovipositor with various apices, 
but if styli absent, apex sharply acute or distinctly 
excised medially . . . . . . . . . . . . . tribe Nitidulini