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А.Г. Кирейчук: очерк сем. Nitidulidae из монографии

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.

                     TAXONOMIC COMPOSITION

        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).

                      HISTORIC DEVELOPMENT

        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