Content uploaded by Zoya Alexandrovna Fedotova
Author content
All content in this area was uploaded by Zoya Alexandrovna Fedotova on Jan 27, 2017
Content may be subject to copyright.
ISSN 00310301, Paleontological Journal, 2016, Vol. 50, No. 9, pp. 1001–1026. © Pleiades Publishing, Ltd., 2016.
1001
INTRODUCTION
Fossil gall midges are very poorly understood,
although they are widespread in amber. The highest
diversity of gall midges is characteristic of Late Eocen e
Rovno and Baltic ambers. We have identified 87 spe
cies of gall midges of 19 genera in samples of the above
ambers (Fedotova and Perkovsky, 2009; Fedotova and
Perkovsky, 2015, including the list of new species).
Information on Mesozoic gall midges is very scarce;
therefore, the gall midges recently identified in the
Late Cretaceous amber of the Taimyr Peninsula are of
great interest for comparative studies involving previ
ously characterized faunas. Eight new species and
eight new genera of gall midges are described in the
present article.
The only presently known Jurassic gall midge,
Catotricha mesozoica
Kovalev (Kovalev, 1990), is rep
resented by an imprint in Late Jurassic deposits
(Glushkovo Formation, Transbaikalia); this species
was later attributed to a separate genus,
Mesotrichoca
Jaschhof et Jaschhof. Four monotypic gall midge gen
era were described from fossil resins of the Late Creta
ceous (Campanian, Canadian amber) (Gagné, 1977)
and two more monotypic genera were described from
Early Cretaceous (Albian) amber of Spain (Arillo and
Nel, 2000); where Lestremiidae and Porricondylinae
(Cecidomyiidae) occur. New material shows that the
Taimyr amber assemblage is dominated by Lestremi
idae. Genera and species of Mesozoic gall midges are
listed in Table 1.
Gall midges from the Late Cretaceous Taimyr
amber are very diverse. A total of ten gall midge species
(eight have already been described) of eight genera,
ten tribes, one supertribe, five subfamilies belonging to
the families Lestremiidae and Cecidomyiidae (two
Porricondylinae fragments are not identified to genus)
have been recorded. Species belonging to the subfam
ily Micromyinae of the family Lestremiidae prevail in
the fauna. The recently developed systematics of this
subfamily (Jaschhof, 2009; Gagné and Jaschhof,
2014) includes nine tribes, six of which occur in the
Taimyr amber. The tribe Strobliellini is represented by
three species of three genera. Species of this tribe are
very rare in the modern fauna, with only seven species
of five genera characterized to date. Another mono
typic genus,
Eltxo
Arillo et Nel, 2000 was recorded in
the Early Cretaceous amber from Spain. Interestingly,
the antennae of all fossil Strobliellini from amber
include 2+11–13 flagellomeres, in contrast to that of
modern species with more than 18 flagellomeres.
Representatives of the pedogenetic supertribe Het
eropezidi have not yet been recorded in the Taimyr
First Gall Midges (Diptera, Cecidomyioidea)
from Late Cretaceous Amber of the Taimyr Peninsula
Z. A. Fedotova
a
and E. E. Perkovsky
b
a
AllRussia Research Institute of Plant Protection, Russian Academy of Sciences,
sh. Podbel’skogo 3, Pushkin, St. Petersburg, 196608 Russia
email: zoyafedotova@mail.ru, zoyafedotova@gmail.com
b
Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine,
ul. Bogdana Khmel’nitskogo 15, Kiev, 01601 Ukraine
email: perkovsky@fromru.com
Received August 10, 2015
Abstract
—Gall midges from Santonian amber of Yantardakh (Taimyr Peninsula) are investigated for the first
time. Eight new genera and eight new species belonging to five tribes of the subfamily Micromyinae
(Lestremiidae) are described. These are
Caputmunda yantardakhica
gen. et sp. nov. (Catochini),
Cretoper
omyia taimyrica
gen. et sp. nov. (Peromyiini),
Palaeostrobliella dmitrii
gen. et sp. nov.,
Yantardakhiella pusilla
gen. et sp. nov., and
Zherikhiniella pedicellata
gen. et sp. nov. (Strobliellini),
Menssana norilsk
gen. et sp. nov.
(Micromyini), and
Cretomycophila ekaterinae
gen. et sp. nov., and
Corporesana khatanga
gen. et sp. nov.
(Aprionini). A representative of Porricondylinae (Cecidomyiidae, Porricondylinae, Holoneurini) is
recorded. The diagnoses based on morphometric parameters of tribes and previously established genera are
emended. The species composition of gall midges from three Cretaceous amber faunas of different ages are
analyzed.
Keywords:
Diptera, Cecidomyioidea, gall midges, new taxa, Late Cretaceous Amber, Taimyr Peninsula
DOI: 10.1134/S0031030116090033
1002
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
Table 1.
Mesozoic Cecidomyioidea
Systematic status
Daya,
Upper Jurassic,
Transbaikalia
(Kovalev, 1990)
Spanish amber
(Alava), Albian
(Arillo and Nel,
2000)
Taimyr amber
(Yantardakh),
Santonian (orig.)
Canadian amber
(Medicine Hat),
Campanian
(Gagné, 1977)
LESTREMIIDAE
Catotrichinae
Mesotrichoca mesozoica
(Kovalev, 1990) +
Micromyinae
Catochini
Cretocatocha mcalpinei
Gagné, 1977 +
Caputmunda yantardakhica
gen. et sp. n. +
Strobliellini
Eltxo cretaceus
Arillo et Nel, 2000 +
Palaeostrobliella zherikhini
gen. et sp. n. +
Yantardakhiella pusilla
gen. et sp. n. +
Zherikhiniella pedicellata
gen. et sp. n. +
Campylomyzini
Cretocordylomyia quadriseries
Gagné, 1977 +
Micromyini
Menssana norilsk
gen. et sp. n. +
Peromyiini
Cretoperomyia dmitrii
gen. et sp. n. +
Aprionini
Corporesana khatanga
gen. et sp. n. +
Cretomycophila ekaterinae
gen. et sp. n. +
CECIDOMYIIDAE
Porricondylinae
Winnertziini
Cretowinnertzia angustata
Gagné, 1977 +
Diallactini
Cretohaplusia ortunoi
Arillo et Nel, 2000 +
Incertae generis
Porricondylinae sp. +
Holoneurini sp. +
Lasiopterinae
Heteropezidi
Miastorini
Cretomiastor ferejunctus
Gagné, 1977 +
Total 1 genus,
1 species 2 genera,
2 species 8 genera,
10 species 4 genera,
4 species
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1003
amber, in contrast to other nontropical amber faunas,
including Canadian amber (Fedotova and Perkovsky,
20 09) . At the sa me t ime , Taim yr a mbe r ha s yiel ded gal l
midges of the genus
Cretomycophila
gen. nov. similar to
the modern genera
Tekomyia
Möhn and
Mycophila
Felt of the tribe Aprionini, which show pupal pedo
genesis (Möhn, 1960).
CLASSIFICATION OF GALL MIDGES
The traditional concepts of gall midge systematics
reflected in the catalogue and key to Palearctic
Diptera (Skuhravá, 1986, 1997) include the division of
the gall midge family into three subfamilies: Lestremi
inae, Porricondylinae, and Cecidomyiinae. The more
archaic Lestremiinae differ from closely related and
advanced Porricondylinae + Cecidomyiinae in a com
plex of plesiomorphic features (such as welldeveloped
wing venation and the first tarsomere being consider
ably longer than the second). In two issues of the cat
alogue of the world gall midge fauna, Gagné (2004,
2010) divided Cecidomyiidae into four subfamilies:
Lestremiinae, Catotrichinae (following Jaschhof
(2001), who distinguished it from the former, assign
ing to it the only genus Catotricha), Porricondylinae,
and the higher gallfly subfamily Cecidomyiinae).
Fedotova (2000) attributed Lestremiinae and
Catotrichinae to a separate family, Lestremiidae. Clas
sification of this family (ranked as the Lestremiinae
subfamily) has then been under active development
(Jaschhof, 2009); however, significant changes with
out appropriate justification was introduced into the
classification in the third appended
Catalogue
(Gagné
and Jaschhof, 2014). Some Lestremiinae taxa (the
tribe Lestremiini and the supertribe Micromyidi) were
ranked subfamilies and, hence, became equal to the
previously established subfamilies Lestremiinae and
Catotrichinae. Thus, the previously recognized sub
family Lestremiinae was disintegrated to form several
subfamilies. The taxon Porricondylinae was also disin
tegrated, since the tribe Winnertziini was regarded as a
subfamily, becoming equal in rank to Catotrichinae,
Lestremiinae, Micromyinae, Porricondylinae proper,
and the higher gall midges Cecidomyiinae. However,
the traditional attribution of Porricondylinae + Ceci
domyiinae to a common independent branch is based
on the presence of a common apomorphy (the first
tarsomere being much shorter than the second) con
trasting with the plesiomorphic state of the same trait
in Lestremiinae (the first tarsomere always being
longer than the second). Specific structural features of
the genitalia, with the characteristic archaic traits in
Lestremiinae, represent additional distinctive features
of the derivates. At present, the subfamilies identified
are only subdivided into tribes, while more detailed
generic and subgeneric classification has not yet been
developed, since the existing tribes reflect the entire
diversity of the taxonomic groups identified, with
many genera lowered in rank to subgenera. As a result,
a relatively small group of Lestremiinae + Porri
condylinae gall midges, including 1260 species of 148
genera, is subdivided into five subfamilies, while the
major group of biologically and phylogenetically
diverse higher gall midges (Cecidomyiinae) is con
fined to a single subfamily, including 4819 species of
565 genera. The present classification of this subfamily
(regarded here as a separate family of the superfamily
Cecidomyioidea) includes all the suprageneric taxa
(subtribes, tribes, and supertribes) (Fedotova, 2013,
2014).
The elevation of the former suprageneric gall midge
taxa of the entire family Cecidomyiidae is undoubt
edly justified (Krivosheina and Zaitsev, 1989; Fedot
ova, 2000), since the old classification of higher gall
midge subfamily Cecidomyiinae was capable of nei
ther reflecting the entire faunal diversity nor providing
a more advanced classification compared to that for
small and less diverse Lestremiinae and Porricondyli
nae. The ranks of higher taxa of Cecidomyiinae were
considerably lower in the weight of traits than used
analogous groups of taxa used in classifications of
other insect families. Therefore, it was proposed to
resurrect the superfamily Cecidomyioidea with the
families Lestremiidae and Cecidomyiidae, which
includes the subfamilies Porricondylinae and Cecid
omyiinae + Lasiopterinae (the former supertribes of
the family Cecidomyiinae), which share a very short
first tarsomere compared to the second and a much
simpler wing venation than in Lestremiidae. Cecid
omyiinae differs from Porricondylinae in the genital
structure and the fused first and second flagellomeres.
The recognition of the superfamily Cecidomyioidea is
presently accepted by many researchers (Matile, 1997,
a review of the classification systems of Diptera; Lam
bkin et al., 2013); it was repeatedly shown to be useful
for the development of classification of higher gall
midges of the family Cecidomyiidae. In the present
study, we regard gall midges as a superfamily, including
the family Cecidomyiidae with the subfamilies Porri
condylinae, Cecidomyiinae, and Lasiopterinae, and
the separate family Lestremiidae, including the sub
family Micromyinae with nine tribes (Gagné and Jas
chhof, 2014), six of which occur in the Taimyr Penin
sula (Table 1). Gall midges of these Lestremiidae
tribes, as well as Porricondylinae are rather frequent in
amber from other deposits (Perkovsky and Fedotova,
2004, 2008; Fedotova and Perkovsky, 2004, 2005,
2007).
MATERIAL
All specimens examined come from Yantardakh
(Kheta Formation), the largest and most thoroughly
investigated insectbearing Santonian amber locality.
This locality is a cliff on the right bank of the Mai
mecha River (tributary of the Kheta River) 3 km
upstream from the mouth of the river, Yantardakh Hill
(geographical coordinates: N 71°18
′
26.54
′′
1004
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
E99°33
′
46.51
′′
) about 30 m high and 200 m along the
strike. The insectbearing resins were collected in
three steps: in 1970 and 1971 (Zherikhin and
Sukacheva, 1973) and in 2012 (the expedition of
E.A. Sidorchuk, D.S. Kopylov, and D.D. Vorontsov).
Specimens PIN, nos. 3311/3139 and 3311/3141–
3147 mentioned in the present paper were collected by
V.V. Zherikhin and I.D. Sukacheva (Zherikhin and
Sukacheva, 1973) and all others were collected in
2012. The photographs were taken by E.E. Perkovsky
and V.Yu. Nazarenko, using a Leica M 16 stereomi
croscope, by A.P. Rasnitsyn, using a Leica M 165 ste
reomicroscope equipped with a Leica DFC 425 cam
era, and by Z.A. Fedotova, using a ZEISS Imager M1
“Axio Imager” microscope equipped with an Axio
Cam Mrc5 T2CZEISS camera.
The material investigated is stored in the amber
collection of the Borissiak Paleontological Institute of
the Russian Academy of Sciences, Moscow (PIN).
SYSTEMATIC PALEONTOLOGY
S u p e r f a m i l y Cecidomyioidea Newman, 1834
Family Lestremiidae Rondani, 1840
Subfamily Micromyinae Rondani, 1856
All fossil gall midge genera from Taimyr were
attributed to the abovenamed subfamily based on the
presence of simple medial vein M and forked cubitus
CuA.
The subfamily includes 527 extant species of
39 genera and 29 fossil species of 6 genera (Jaschhof
and Jaschhof, 2009). In general, representatives of the
subfamily occur worldwide and are either mycetoph
ages or sapromycetophages.
Tr i b e Catochini Edwards, 1938
The tribe comprises six genera and sixteen species
(Gagné and Jaschhof, 2014), including a single fossil
species and genus,
Cretocatocha mcalpinei
Gagné from
the Cretaceous amber of Canada (Alberta) (Gagné
1977).
A new genus from the Taimyr amber is described
below and assigned to the tribe Catochini based on a
very long basal part of the M vein, which is longer than
the forked part of this vein, the narrow wing cell
between the
R
4+5
and C veins, the long segment of the
costal vein adjacent to the costal break near the wing
apex, and the veins simple CuA1 and CuA2.
Larvae of extant species of the genus
Catocha
Hal
iday dwell in forest litter (Mamaev and Krivosheina,
1965).
Genus
Caputmunda
Fedotova et Perkovsky gen. nov.
Type s p ecie s.
C. yantardakhica
sp. nov.
E t y m o l o g y. From Caput Mundi, a Latin
phrase taken to mean “capital of the world,” based on
a classical European understanding of the known
world as Europe, North Africa, and Southwest Asia.
D e s c r i p t i o n. Male (Fig. 1). The body is very
short, with an inflated broad abdomen, but longer than
the wing. The head is narrower than the thorax
(Figs. 1a, 1b). Antennae are thin, with partially lost
short antennomeres. Flagellomeres are barrelshaped,
broadened slightly basally, as long as wide (Figs. 1b, 1d).
Flagellomeres have a basal whorl of short setae and
medial whorl of very long setae. The labrum and label
lum are short triangle (Fig. 1i). The palpus includes
four slightly elongated, almost parallelsided pal
pomeres (Fig. 1b) and is shorter than the head height.
The wing is shorter than the body, broad, and has a
rounded anal lobe passing into a broadly rounded ven
tral edge of the wing (Figs. 1a and 1c). Vein
R
4+5
is
close to the wing margin,
M+rm
is evanescent, and
Rs
is close to the site where
R
1+2
joins the costal vein.
The
R
4+5
vein reaches the wing margin (Fig. 1c)
noticeably in front of the wing apex, joining the costal
vein, which further extends along the edge of the wing
to its apex as a long fragment rather than ends at the
point of junction with
R
4+5
. The wing is very densely
covered with long macrotrichias. The costal vein is
interrupted near the wing apex. The wing lacks a mar
ginal fringe within a rather broad part of this area adja
cent to the M
1
vein, which forms a short fork with M
2
.
CuA
1
and CuA
2
do not form a fork; CuA
2
bends
sharply in the distal part and is very broadly rounded.
Venation is poorly pronounced at the base of the wing
against a background of a very broad cell. The hind
legs are almost as long as the body. Only the hind legs
are preserved completely; in addition, there is an iso
lated protarsus located near the head in the amber
piece (Pl. 9, fig. 1c). The mesotarsus is visible against
a background of the genitalia (Pl. 9, fig. 1b). The claws
are long, simple, and slightly bent (Fig. 1e).
The abdomen is slightly inflated and slightly
shorter (excluding the genitalia) than the head and
thorax combined. The aedeagal complex is well devel
oped, reaching or almost reaching the tip of the gono
coxite (Fig. 1f). The genitalia have broad gonocoxites
and gonostyles of almost equal width at the joint. The
gonostyles are broadly oval, slightly broadened in the
middle part and shorter than the gonocoxites.
Species composition. Type species.
C o m pa r i s o n a n d r e m a r k s. In the set of
characters, the new genus clearly differs from extant
genera, displaying a mosaic of characters of several
genera of the tribe Catochini. The genus is attributed
to this tribe due to the characteristic wing venation.
The news genus differs from
Catocha
Haliday, 1833 in
shorter distance between
R
4+5
and the costal vein, the
very broad cell between veins CuA
2
and
R
1+2
, the very
strongly bent CuA
2
vein, the absence of flagellomere
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1005
necks in male, and the relatively short first tarsomere.
The last character distinguishes the new genus from all
other genera of the tribe. In addition, it differs from
Tritocyga
Loew, 1862 in the location of the symmetric
branching point of the medial vein beyond the wing
apex, the presence of a nick at the apex of the wing,
and the sharp curvature of CuA
2
, but is similar to the
latter in the close positions of the costal vein and
R
4+5
,
the presence of a nick of the wing, the symmetrical
branches of the medial vein, and the considerable
width of the apical wing part. The new genus differs
from
Cretocatocha
Gagné, 1977 from the Cretaceous
Canadian amber (Gagné, 1977) in the absence of the
anal wing lobe, very narrow cell between the costal
vein and
R
4+5
, the costal vein extending to the wing
apex, the sharply curved CuA2, the barrelshaped
flagellomeres equal in size (
Cretocatocha
has narrow
bristlelike antennae with short segments sharply
decreasing in diameter), and in the shorter first tar
somere.
Caputmunda yantardakhica
Fedotova et Perkovsky sp. nov.
Plate 9, figs. 1a–1d
E t y m o l o g y. From the Yantardakh locality.
H o l o t y p e. PIN, no. 3311/2100, poorly pre
served male with the legs partially lost and the head
and ptotarsus located in a piece detached during treat
ment; Yantardakh,Taimyr amber; Santonian.
D e s c r i p t i o n. Male (Fig. 1). The head is almost
spherical, the eyes cover entirely the lateral sides of the
head; the body (excluding genitalia) is 3.6 times as
long as the head; the head (excluding mouthparts) is
slightly wider than high. The wing is 0.77 as long as the
body. Antennae are preserved only partially, including
the scapus, pedicel, and five flagellomeres. The distal
part of the scapus is inflated and the pedicel is almost
spherical. All flagellomeres lack necks (Fig. 1i) and
almost as long as wide (Fig. 1h). The labellum is pro
truding and triangular.
The metafemora are longer than the metatibia,
which are as long as the tarsi. The leg base is invisible.
(a), (b)
(d), (e) (f), (g)
(c) (h)
(b)
(e)
(h)
(a)
(i) (c)
(f)
(d)
Fig. 1.
Caputmunda yantardakhica
sp. nov., holotype no. PIN, no. 3311/2100, male: (a) general habitus; (b, d) head, posterior
view; (c) wing ; (e) protarsus; (f) genitals; (g) metatarsomeres 2–5; (h) metatarsus; (i) scapus, pedicel, and flagellomeres 1–5.
Scale bar in Figs. 1–7, 0.1 mm.
1006
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
Plate 9
200
µ
m
1b
1c
1a
500
µ
m
1d
2b
2c
500
µ
m2a 2e 2d
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1007
The length ratios of the tarsomeres in the protarsus are
1 : 0.9 : 0.9 : 0.6. Tarsomere 2 of the protarsus is
2.9times as long as tarsomeres 3, 4, and 5 taken
together, while in the mesotarsus, this ratio is 2.5. The
length ratios of tarsomeres of the metatarsus is 1 : 0.5 :
0.4 : 0.4 : 0.4 and tarsomere 2 is 1.8 times as long as tar
someres 3, 4, and 5 taken together. The maximum
wing width is at the wing midlength. The wing is twice
as long as wide, bordered by short hairs.
R
1+2
reaches
the wing edge noticeably anterior to the middle of the
wing; it is 0.45 of the wing length.
R
4+5
reaches the
wing edge noticeably anterior to the wing apex and
forms a very narrow almost parallelsided cell with the
costal vein. The abdomen is inflated at the base and
almost as long as the thorax. The genitalia are trans
versely expanded. Lateral sides of the gonocoxites are
rounded; and gonostyles are very long and narrow. The
gonostyle is shorter than the gonocoxite and probably
lacks an apical claw (Fig. 1f).
Female unknown.
M e a s u r e m e n t s, m m: body length (including
head and excluding genitalia), 0.88; head height with
out mouthparts, 0.17; head width, 0.19; length of the
antenna with five preserved flagellomeres, 0.16; wing
length, 0.69; wing width, 0.34; halter length, 0.17;
abdomen length, 0.33; thorax length, 0.33; protarsus
length, 0.23; metatibia, 0.22; metatarsus, 0.23; width
of genitalia, 0.10.
Material. Holotype.
Tr i b e Peromyiini Kleesattel, 1979
The tribe includes two genera, the monotypic
genus (with
Gagnea tsutaensis
Jaschhof 2001) occur
ring in Japan and a very speciose cosmopolitan genus
Peromyia,
which includes 165 species (Fedotova and
Perkovsky, 2004; Gagné and Jaschhof, 2014), with one
species (
P. (Joannisia) monilifera
Meunier, 1904)
recorded in the Late Eocene Baltic amber and
four species (
Peromyia zherikhini
Fedotova, 2004,
P. sukachevae
Fedotova, 2004,
P. miranda
Fedotova,
2004, and
P. autonoma
Fedotova, 2004) recorded only
recently in the Late Eocene Rovno amber (Fedotova
and Perkovsky, 2004).
A new genus from the Taimyr amber described
below is assigned to the tribe Peromyini based on the
presence of an elongated fragment of
R
1+2
(from
Rs
to
C
);
the absence of the costal vein fragment after the fusion
with
R
4+5
; the presence of 2+8segmented antennae in
female, distinct neck and rounded basal enlargment in
flagellomeres, with long setae of the medial whorl
located in randomly dispersed small bosses with apical
impressions rather than in crenate pores arranged in
regular rows, as in most Micromyidi genera, and the
presence of a 3segmented ovipositor.
Larvae of the majority of extant species of the genus
Peromyia
Kieffer develop in cavities under deeply
decomposed bark of fallen trees, while some larvae
dwell in the ground litter (Mamaev and Krivosheina,
1965).
Genus
Cretoperomyia
Fedotova et Perkovsky gen. nov.
Type s p ecie s.
C. dmitrii
sp. nov.
E t y m o l o g y. From Cretaceous Period and the
generic name
Peromyia
.
D e s c r i p t i o n. Female (Fig. 2). The body is
shorter than the wing, densely covered with scales; the
abdomen is broad. The head is rounded. The eyes are
considerably shifted onto the anterior part of the head.
The eye bridge is distinct, but narrow. The head is
broader than the thorax (Fig. 2a). The antennae con
sist of 2 + 8 segments and are 2.1 times longer than the
head. The scapus and pedicel are spherical and con
siderably broader than all other flagellomeres.
Flagellomeres have very short necks and carry very
dense whorls of basal and medial setae. Flagellomeres
are slightly longer than wide. The first flagellomere is
noticeably elongated. The apical flagellomere is
slightly elongated. The eighth flagellomere is slightly
ovate and separated from the seventh flagellomere
(without a trace of fusion). The palpi are twoseg
mented and the palpiger is strongly inflated and
rounded.
The thorax is almost straight, onethird as long as
the abdomen with extended ovipositor. The dorsum is
only slightly convex, almost straight. The wing is
longer than the body and legs. The wing width cannot
be determined due to poor preservation of the speci
men. The cell between the
R
1+2
and
R
4+5
veins expands
strongly distally. Rs is 0.4 as long as the fragment of
R
1+2
between Rs and the point of fusion with the costal
vein.
R
4+5
reaches the wing edge noticeably anterior to
the wing apex, and
M + rm
is sharply bent. The M
1
vein is represented by a hardly discernible short frag
ment. The Cu vein is not preserved in the wing frag
ment.
The forelegs are longer than the body, but shorter
than the wing. The protibia are slightly shorter than
the profemora, which are almost equal to the tarsi in
length. The first tarsomere of the protarsus is 1.6 times
longer than the second. The claws of protarsi are sim
Explanation of Plate 9
Fig. 1.
Caputmunda yantardakhica
sp. nov., holotype PIN, no. 3311/2100, male: (a) lateral view of body with isolated head,
×
97.2;
(b) genitalia, ventral view,
×
209.1; (c) posterior view of complete head isolated from the body,
×
131.5); (d) wing,
×
126.1.
Fig. 2
.
Cretoperomyia dmitrii
sp. nov., holotype PIN, no. 3311/1416, female: (a) lateral view of body,
×
97.2; (b) ovipositor, dorsal
view,
×
209.1; (c) head, right lateral view,
×
131.5; (d) head, lateral view, and antennae, left lateral view,
×
126.1; (e) part of the tho
rax, abdomen, and metatibia, right lateral view,
×
209.1.
1008
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
ple hooked. The end of the abdomen and the oviposi
tor are bent in the dorsal direction. The apical plates of
the ovipositor are paired, closely adjacent, and consist
of three segments.
Male unknown.
Species composition. Type species.
C om p a r i s o n. The new genus is similar to
Per
omyia
, the most abundant genus of Lestremiidae,
which is known in Late Eocene Baltic (Gagné and Jas
chhof, 2014) and Rovno ambers. The new genus is
similar to
Peromyia
in wing venation, the presence of
necks in female flagellomeres, and elongated oviposi
tor with threesegmented plates and is distinguished
by the very large head exceeding the thorax in width;
the presence of a very narrow eye bridge; the inflated
scapus, pedicel, and palpiger; 2segmented palpi;
2 + 8segmented antennae; the very long wings rela
tive to the body and legs; the angularly bent
M + rm
vein positioned very far from
R
1+2
; the presence of a
hardly discernible fragment of the M vein; the position
of
R
4+5
reaching the wing edge far anterior to the wing
apex; and the femora considerably broader than the
tibiae and tarsi.
Cretoperomyia dmitrii
Fedotova et Perkovsky sp. nov.
Plate 9, fig. 2
E t y m o l og y. Named in honor of Dmitrii Kopy
lov, who collected amber samples with gall midges in
the Taimyr Peninsula in 2012.
H o l o t y p e. PIN, no. 3311/1416, poorly pre
served female, with partially lost legs and partially pre
served wing; Yantardakh,Taimyr amber; Santonian.
D e s c r i p t i o n. Female (Fig. 2). The head has a
widely rounded protruding occipital part. The eye
bridge is very narrow. The head is slightly longer than
wide. The wing is 1.2 times as long as the body. The
antennae are 0.48 as long as the body and with 2 + 8 seg
ments. All flagellomeres, except the terminal one,
have a distinct short neck (Figs. 2a
and 2b). The first
flagellomere is slightly larger and broader than the
others. The fifth flagellomere is 1.4 times as long as
wide. The eighth flagellomere is ovate. The palp seg
ments are very short. The palpiger is larger than any of
(a) (c)
(b), (d), (e)
(a)
(e)
(b)
(c)
(d)
Fig. 2.
Cretoperomyia taimyrica
sp. nov., holotype no. PIN, no. 3311/1446, male: (a) general habitus; (b) flagellomeres 2–8;
(c) protarsus; (d) head, anterior view; (e) ovipositor.
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1009
the segments. The thorax is narrow, almost as long as
wide and the width is less than the head length. The
coxae, especially the procoxa, are very long.
The profemora are 1.1 times longer than the protibiae;
the mesofemora are shorter than mesotibiae; and the
metafemora are 1.3 times longer than the metatibiae.
The profemora are slightly narrower than the
metafemora. In each tarsus, the sum length of tarsom
eres 3, 4, and 5 exceeds the length of the first tarsom
ere.
R
1+2
reaches the wing edge far beyond the middle
of the edge; the vein is 0.625 as long as the wing.
M e as u r e m e n t s (mm): body length (including
ovipositor), 1.13; head length (without mouthparts),
0.25; head height, 0.23; antenna length, 0.53; wing
length, 1.27;
R
1+2
length,0.78; procoxa length, 0.13;
trochanter, 0.04; profemur, 0.39; protibia, 0.35; tar
somere 1, 0.13; tarsomere 2, 0.09; tarsomere 3, 0.08;
tarsomere 4, 0.04; tarsomere 5, 0.04; tarsus, 0.38, fore
leg, 1.29; mesocoxa length, 0.11; trochanter, 0.04;
mesofemur, 0.38; metacoxa length, 0.13; trochanter,
0.04; metafemur, 0.48.
Material. Holotype.
Tr i b e Strobliellini Edwards, 1938
The tribe consists of six genera and eight species
(Gagné and Jaschhof, 2014), including
Eltxo
Arillo et
Nel, 2000 recorded in the Lower Cretaceous amber of
Alava (Spain) (Arillo and Nel, 2000).
Three new genera from the Taimyr amber
described below are assigned to the tribe Strobliellini
due to the characteristic wing venation (the presence
of simple veins
M
and
Cu
); the considerable length of
the
R
1+2
fragment (from
Rs
to
C
), which is more than
twice longer than
Rs
; and the structure of flagellom
eres, including a rounded basal inflation. The charac
ters distinguishing the new genus from other taxa of
the tribe include a very small body size, short antennae
of 2 + 12 (rather than 2 + 18 – 23) segments; and a
smoothly curved (rather than angular) vein
M + rm.
Genus
Palaeos t robliella
Fedotova et Perkovsky gen. nov.
Type s p ecie s.
P. zherikhini
sp. nov.
E t y mo l o g y. From the generic name
Strobliella
Kieffer.
D e s c r i p t i o n. Male (Fig. 3). The body is cov
ered with abundant setae and scales. The head looks cir
cular in anterior and lateral views. The body is very short
(shorter than the wing) and the abdomen is broad. The
head length is considerably less than the thorax width
(Fig. 3a). The antennae with 2 + 12 segments and are
shorter than the body or wing. The flagellomeres have
a strongly inflated (almost spherical) basal enlarge
ment, which is much longer than the neck in all
flagellomeres. The shape of the flagellomeres, the
basal enlargement, and the neck is almost invariant
along the antenna from its base to the apex. A well
developed basal whorl of short setae and the medial
whorl of very long deflected setae are apparent on the
flagellomeres. The palpi are threesegmented (Fig. 3a
and 3e) and do not exceed the height of the head; the
medial parts of the segments are uniformly broadened.
The palpiger is well developed. The wing is broad,
shorter than the body, and lacks an anal lobe. The wing
is 2.5 times as long as wide. The
R
4+5
vein is located at
a considerable distance from the costal vein and shows
a considerable distal curvature,
M + rm
is strongly
smoothly curved and forms a broad cell with
R
1+2
,;
Rs
is slightly shifted from the point of contact of
R
1+2
and the costal vein.
R
4+5
reaches the wing edge at its
apex (Fig. 3a), fusing into the costal vein, which
extends beyond the wing apex and is interrupted
before the emergence of the marginal vein. The
M
vein
does not form a fork, represented by a very short frag
ment. The Cu vein is simple and slightly bent before
reaching the wing edge. The wing edge lacks hairs.
All legs are distinctly shorter than the body, but
almost as long as the wing. The femora, tibiae, and
tarsi are very similar in length. The hind legs are
slightly longer than the forelegs. The first tarsomere is
almost twice longer than the second. Tarsal claws are
hooked; protarsal claws have a basal tooth.
The abdomen is inflated and, even excluding the
genitalia, longer than the head and thorax combined.
Female unknown.
Species composition. Type species.
C o m p a r i s o n. The new genus strongly differs
from other representatives of the tribe Strobliellini. In
the wing shape and venation, it is most similar to the
genus
Strobliella
Kieffer and differs from the latter in
the wellpronounced curvature of the vein
M + rm
located at a distance from
R
1+2
;
the welldeveloped
Cu
1
vein, threesegmented palpi; the presence of a
tooth on the claws of the protarsi; the presence of a
vein break beyond the wing apex, 2+12segmented
antennae, and in the smaller body. In the presence of
short antennae and threesegmented palpi, the new
species is similar to
Eltxo
from the Lower Cretaceous
Spanish amber (2 + 13segmented antennae, body
0.85 mm long) and differs from it in the presence of a
wide ocular bridge, the short M vein, the welldevel
oped CuA1, the shorter flagellomeres with an almost
spherical basal enlargement, and in the larger body
covered with both bristles and scales.
Palaeostrobliella zherikhini
Fedotova et Perkovsky sp. nov.
Plate 10, fig. 3
E t y m o l o g y. In honor of V.V. Zherikhin, who
discovered the holotype.
H o l o ty p e. PIN, no. 3311/3141, wellpreserved
male in amber; Yantardakh,Taimyr amber; Santonian.
1010
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
D e s c r i p t i o n. Male (Fig. 3). The head has a
roundly protruding occiput. The eye bridge is wide.
The head length is 0.77 of the head width (without
mouthparts). The wing is 0.83 as long as the body. The
antennae are 0.43 as long as the body and 0.63 as long
as the wing; the pedicel is almost round. The fifth
flagellomere is 1.3 times as long as wide; the neck is 0. 4
of the almost spherical basal enlargement; at the
antennal apex, the basal enlargements becomes wider
than long. The twelfth flagellomere has a rounded
apex. The palp segments are short; the third pal
pomere is slightly longer than the others (with the
length ratios 1 : 1 : 1.1). The palpi are short (length less
than head width), with slightly inflated segments, of
which the second and third are longer than the others.
The mouthparts only slightly project beyond the head
edge (Fig. 3a).
The thorax is broad, 1.3 times longer than wide.
The dorsum is inflated, reaches the level of the
occiput. The procoxae are considerably longer than
the others. The trochanters are rounded, less than half
as long as the coxae. The femora of the pro and mid
legs are 0.91 as long as the tibia; in the hind legs, this
ratio is 1.3. The pro, meso, and metafemora are
almost identical in width. The pro and mesotarsi are
almost as long as the tibiae; and the metatarsi are
slightly longer. The first tarsomere of the pro and
metatarsus is twice longer than the second tarsomere;
in the mesotarsus, this ratio is 1.5. The sum length of
tarsomeres 3, 4, and 5 is less than the length of the first
tarsomere of the protarsus and, in the metatarsi, these
lengths are equal. The hind legs are 1.1 times shorter
than the forelegs. The wing lacks an anal lobe and
twice as long as wide. Vein
R
1+2
reaches the wing edge
(b)
(c)
(e)
(a)
(d)
(a) (f)
(b)–(e)
Fig. 3.
Palaeostrobliella zherikhini
sp. nov., holotype PIN, no. 3311/3141, male: (a) general habitus; (b) protarsus; (c) flagellom
eres 3–6; (d) metatarsus; (e) palpus.
Explanation of Plate 10
Fig. 1.
Palaeostrobliella zherikhini
sp. nov., holotype PIN, no. 3311/3141, male: (a) general appearance ,
×
45.6; (b) metatarsus,
×
217.6; (c) head, antennae, and palpi,
×
103.2; (d) right wing with veins M and Cu,
×
100; (e) left wing with Cu vein,
×
100;
(f) partial eye and palpus,
×
228.6; (g) protarsus and tibia,
×
225; (h) antennal base,
×
160; (i) antennal apex,
×
137.
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1011
Plate 10
1b
1a1c
1d
1f
1g
1e 1i 1h
1012
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
far posterior to the middle of the wing and is 0.625 as
long as the wing. The abdomen is slightly inflated near
the middle and almost equal to the thorax in width.
The gonocoxites and gonostyles are thin and elon
gated.
D i m e n s i o n s (mm): Body length (excluding
genitalia), 1.47, (including genitalia), 1.50; head
length, 0.20; head height, 0.26; antenna length, 0.63;
wing length, 1.19; wing width, 0.60; abdomen length,
0.85; thorax length, 0.53; length of the procoxa, 0.13;
trochanter, 0.05; profemur, 0.31; tibia, 0.35; first tar
somere, 0.12; 2nd tarsomere, 0.06; 3rd, 0.05; 4th,
0.04; 5th, 0.05; tarsus, 0.32; foreleg, 1.09; mesocoxa
length, 0.09l; trochanter, 0.05; femur, 0.31; tibia, 0.35;
mesotarsus, 0.33; midleg, 1.13; metacoxa length, 0.09;
trochanter, 0.05; femur, 0.41; tibia, 0.31; tarsus, 0.34;
leg, 1.20.
Material. Holotype.
Genus
Yantardakhiella
Fedotova et Perkovsky gen. nov.
Type s p ecie s.
Ya. pusilla
sp. nov.
E t y m o l o g y. From the Yantardakh locality.
D e s c r i p t i o n. Male (Fig. 4). The body is cov
ered with abundant setae and thin scales. The body is
very short (almost equal to the wing in length) and the
abdomen is broad. The head is rounded. The eye bridge
is distinct. The head length is almost equal to the thorax
width (Fig. 4a). The antennae have 2 + 12 segments
and are shorter than the body and the wing. The spher
ical pedicel is considerably wider than other flagellom
eres. All flagellomeres include a strongly inflated bar
relshaped basal enlargement, which is considerably
longer than the neck. The flagellomeres become
slightly shorter and narrower towards the apex of the
antenna (Figs. 4a–4c). Welldeveloped basal whorls of
short setae and medial whorls composed of very long
setae are present on flagellomeres.
The thorax seems very small and narrow as com
pared to the very large head. The notum is only slightly
(b), (c)
(c)
(e)
(a)
(a)
(b)
(d)
(e)
(d)
Fig. 4.
Yantardakhiella
pusilla
sp. nov., holotype PIN, no. 3311/3144, male (a–d), and
Menssana
norilsk
sp. nov., holotype PIN,
no. 3311/3146, female (e): (a) general appearance; (b) 5th flagellomere; (c) flagellomeres 10–12; (d) part of the femur, tibia, and
protarsus; (e) antenna.
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1013
convex. The wing is broad and lacks an anal lobe
(Fig. 4a). The wing is 2.5 times as long as wide. Vein
R
4+5
is located at a distance from the costal vein and
bent slightly distally;
M + rm
is strongly bent and
forms a broad cell along with
R
1+2
;
Rs
is located at a
distance from the point of fusion of
R
1+2
and the costal
vein (this distance is more than twice as long as Rs).
Ve i n
R
4+5
reaches the wing apex (Fig. 4a), fusing with
the costal vein that extends beyond the wing apex and
forms a break with the marginal vein. The slightly
curved M
1
vein is not forked and reaches the wing edge
near the wing apex. The rather broad simple CuA
1
vein
is not forked and vanishes only near the wing edge; the
lateral outlines of this vein are very well distinct. The
CuP is well developed.
The legs are distinctly shorter than the body and
wing, but longer than the antennae. The protibiae are
shorter than the metatibiae, while the protarsi are
1.5 times longer than the metatarsi. The first tarsom
ere is almost twice longer than the second. The claws
of the protarsi are hooked. The genitalia are small and
rounded.
Species composition. Type species.
C o m p a r i s o n. The new genus is similar to Early
Cretaceous
Eltxo
in the wing venation, size and shape
of the body, flagellomeres, tarsi, and threesegmented
palpi and differs from it in the wellpronounced curva
ture of
M + rm
located at a distance from
R
1+2
; the
presence of a fragment of the
M
vein; the narrow eye
bridge (eyes of
Eltxo
are completely isolated and
located at a considerable distance from each other);
the weakly convex notum (in
Eltxo,
the notum is
humpshaped); the broad very distinct CuA vein, and
in the slightly inflated spherical pedicel.
Yantardakhiella pusilla
Fedotova et Perkovsky sp. nov.
Plate 11
Etymology. From the Latin
pusillus
(tiny).
H o l o ty p e. PIN, no. 3311/3144, wellpreserved
male. Yantardakh,Taimyr amber; Santonian.
A syninclusion is a paratype of
Zherikhiniella pedi
cellata
sp. nov. (PIN, no. 3311/3145) described in the
present paper.
D e s c r i p t i o n. Male (Fig. 4). The head has a
protruding arched occipital part. The eye bridge is nar
row, but eyes cover entirely the lateral sides of the head.
The face is almost as wide as long. The wing is almost
as long as the body. The antennae are 0.77 as long as
the body. All flagellomeres, except the terminal one,
have a short neck, which is the longest in the middle
flagellomeres shorter in the basal and apical flagellom
eres (Figs. 4a–4c). The fifth flagellomere is 1.9 times
as long as wide. The neck is slightly inflated at the top
and is 2.1 times smaller than the basal enlargement.
The twelfth flagellomere is conical. The palpi are
unseen. The mouthparts slightly protrude beyond the
edge of the head.
The abdomen does not exceed the thorax in width,
but is 1.6 times longer. The coxae are very long and the
trochanters are slightly elongated and covered with
abundant setae. The femora of all legs are shorter than
the tibiae. The profemora are wider than the meso
and metafemora. The pro and metatarsi are longer
than the tibiae. The first tarsomeres of the pro and
metatarsi are almost twice longer than the second tar
someres of the same tarsi, whereas in the mesotarsus,
this ratio is 1.7. The sum length of tarsomeres 3, 4, and
5 of all tarsi exceeds the length of respective first tar
someres. The wing lacks an anal lobe and 2.6 times as
long as wide. The
R
1+2
vein contacts the wing edge far
beyond the middle of the wing and is 0.59 as long as
the wing. The genitalia are very small and hardly dis
cernible.
Female unknown.
M e a s u r e m e n t s (mm). Body length (including
genitalia), 0.91; head length, 0.24; antenna length,
0.69; wing length, 0.93; wing width, 0.36; thorax
length, 0.29; abdomen length, 0.46; protibia length,
0.34; length of anterior first tarsomere, 0.18; length of
2nd tarsomere, 0.09; of the 3rd, 0.08; of the 4th, 0.06;
of the 5th, 0.09; protarsus length, 0.50; mesocoxa
length, 0.08; trochanter, 0.05; femur, 0.29; metatibia,
0.29; first metatarsomere, 0.10; length of the 2nd tar
somere, 0.06; of the 3rd, 0.05; of the 4th, 0.05; of the
5th, 0.06; metatarsus length, 0.32.
Material. Holotype.
Genus
Zherikhiniella
Fedotova et Perkovsky gen. nov.
Type s p e cies.
Zh. pedicellata
sp. nov.
E t y m o l o g y. In memory of the outstanding
Russian paleoentomologist V.V. Zherikhin.
D e s c r i p t i o n. Male (Figs. 5a–5c). The body is
shorter than the wing, covered with abundant scales;
the abdomen is broad. The head is rounded, with the
eyes strongly shifted towards the anterior part of the
head. The eye bridge is distinct. The head length is
almost equal to the thoracic width (Fig. 5a). The
antennae are less than 1.5 times as long as the head and
have 2 + 7 segments. The pedicel is spherical and con
siderably broader than other flagellomeres. The
flagellomeres lack necks and carry thick basal and
medial whorls, while the apical whorl is hardly discern
ible. The flagellomeres are densely packed, slightly
longer than wide, becoming slightly shorter and nar
rower towards the antennal apex (Figs. 5a and 5b). The
first flagellomere is not elongated. The slightly elon
gated apical flagellomere may represent a fusion of two
flagellomeres. The threesegmented palpi are very
short, shorter than half length or width of the head.
The thorax is almost as wide as the abdomen. The
notum is only slightly convex, almost straight. The
wing is broadly rounded, 1.9 times as long as wide,
lacks an anal lobe (Fig. 5a). The proximal part of the
R
4+5
vein is at a long distance from the costal vein,
1014
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
Plate 11
0.2 mm 1
23
0.1 mm
0.1 mm
54
0.1 mm 0.05 mm
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1015
reaches the wing edge noticeably beyond the wing
apex, is arched in the distal part, extends beyond the
apex of the wing after fusion with the costal vein, and
forms a break with the marginal vein.
M + rm
is
strongly curved and forms a broad cell with
R
1+2
, and
Rs
is located at a great distance from the point of
fusion of
R
1+2
and the costal vein (the distance is
almost equal to the
Rs
length). The
M
1
vein is a weakly
developed short segment. The CuA
1
vein is simple,
broad, and distinctly outlined throughout its length
from the base to the point of contact with the wing
edge. The CuP is poorly developed.
The fore legs are shorter than the body and wing,
while the hind legs are longer. The protibia are slightly
longer than the femora and the metatibiae are slightly
shorter. The pro and metatarsi are shorter than
respective femora and tibiae. The first tarsomere is
almost twice longer than the second. The claws of
the protarsi are hooked. The genitalia are small and
round. Very long setae are seen on the posterior edge of
the tarsi.
Female unknown.
Species composition. Type species.
C o m p a r i s o n. The new genus is similar to
Yan
tardakhiella
described above in the very clear CuA
vein, body size, shape of the body and tarsi, broad and
very distinct CuA vein, a slightly inflated rounded
pedicel, and a weakly convex dorsum and is distin
guished by the shape of flagellomeres (which lack a
neck), the curvature of the
M + rm
vein located at a
(b)
(c)
(e)
(a)
(f)
(d)
(e)
(b)
(f)
(a), (c), (d)
Fig. 5.
Zherikhiniella
pedicellata
sp. nov., holotype PIN, no. 3311/3142, male (a–c), and
Menssana
norilsk
sp. nov., holotype PIN,
no. 3311/3146, female (d–f): (a) general habitus, lateral view; (b) body, anterior view, protarsus; (c) palpi; (d) ovipositor apex;
(e) wing ; (f) head, antenna, foreleg with partially closed femur, coxa, trochanter, mesofemur, and tibia.
Explanation of Plate 11
Figs. 1–5.
Yantardakhiella pusilla
sp. nov., holotype PIN, no. 3311/3144, male: (1) general appearance,
×
70.3; (2) wing,
×
121.5;
(3) protibia, protarsus, and mesotarsus,
×
194.1; (4) distal half of the antenna, trochanter, and mesofemur,
×
258.6; (5) thorax,
head, antenna, and leg bases, ×121.7.
1016
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
considerable distance from
R
1+2
, and by the presence
of a hardly discernible fragment of the
M
vein.
Zherikhiniella pedicellata
Fedotova et Perkovsky sp. nov.
Plate 12, Plate 13, fig. 2; Figs. 5a–5c
Etymology. From the wellpronounced infla
tion of the pedicel.
H o l o t y p e. PIN, no. 3311/3142, male, Yan
tardakh, Taimyr amber; Santonian. Syninclusion is
specimen PIN, no. 3311/3143, abdomen of Porri
condylinae sp.
D e s c r i p t i o n. Male (Figs. 5a–5c). The anten
nae are 1.6 times longer than the head. Flagellomeres
are simple, slightly longer than wide, lack a neck, bar
relshaped. The wing is 1.1 times longer than the body
and 2.3 times as long as wide;
R
1+2
is 0.53 as long as the
wing. The pro and metafemora are almost equal in
length. The protarsi are slightly longer than the tibiae,
while the metatarsi are shorter. The first tarsomeres of
all legs are almost twice longer than respective second
tarsomeres.
Measurements, mm. Holotype: body
length, 0.81; antenna length, 0.28; wing length, 0.92;
width, 0.40; thorax length, 0.18; width, 0.13; abdomen
length, 0.46; length of the procoxa, 0.08; trochanter,
0.03; femur, 0.21; tibia, 0.22; protarsus, 0.24; foreleg,
0.78; metacoxa length, 0.08; trochanter, 0.03; femur,
0.30; tibia 0.28; tarsus, 0.27; hind leg, 0.96. Paratype
PIN, no. 3311/3147, male: body length, 0.70; antenna
length, 0.19; thorax length, 0.19; abdomen length,
0.29; procoxa length, 0.10; trochanter, 0.03; femur,
0.20; tibia, 0.17; first tarsomere, 0.07; 2nd tarsomere,
0.04; 3rd, 0.03; 4th, 0.03; 5th, 0.04; protarsus length,
0.21; foreleg length, 0.77; length of the first metatar
somere, 0.06; 2nd tarsomere, 0.04; 3rd, 0.03; 4th,
0.03; 5th, 0.04; metatarsus 0.20.
Material. Holotype and paratypes PIN,
nos. 3311/3147 and 3311/3145 found at the same
locality.
Tr i b e Micromyini Rondani, 1856
The tribe includes five genera with 154 extant spe
cies and three fossil species (Fedotova and Perkovsky,
2007; Gagné and Jaschhof, 2014): one of which
(
Monardia submonilifera
Meunier: 39) is known from
the Late Eocene Baltic amber (male and female, both
lost now) and two fossil species are recorded in Late
Eocene Rovno amber (
Monardia impellucida
Fedot
ova et Perkovsky and
Xylopriona aristata
Fedotova et
Perkovsky). The genus
Xylopriona
Kieffer is regarded
as one of four subgenera of a large (51 species) genus
Monardia
Kieffer in the recent catalogue (Gagné and
Jaschhof, 2014).
Larvae of extant species of the genus
Monardia
develop in decomposed wood and larval
Xylopriona
develop in soil (at depths reaching 40 cm) and decom
posed wood (Mamaev and Krivosheina, 1965).
A new genus described below from the Taimyr
amber is assigned to the tribe Micromyini based on
characteristic wing venation pattern (simple
M
vein
and forked Cu vein), the presence of 2 + 8segmented
antennae in female, the presence of large triangular
transparent sensoria on female flagellomeres, and the
considerable length of the fragment of the
R
1+2
vein
between Rs and the costal vein.
Genus
Menssana
Fedotova et Perkovsky gen. nov.
Type s p ecie s.
M. norilsk
sp. nov.
E t y m o l o g y. From the first part of the Latin
proverb
Mens sana in corpore sano
(a sound mind in a
sound body).
D e s c r i p t i o n. Female (Figs. 5d–5f). The head
is round in anterior and posterior views, slightly higher
than long. The eyes are shifted towards the anterior
part of the head and considerably narrowed in the
occipital part. The eye bridge is narrow. The antennae
have 2 + 8 segments, the pedicel is globular, and the
first flagellomere is longer than the others. All
flagellomeres have a slight constriction near the mid
dle and carry a basal whorl of short bristles, a medial
whorl of very long protruding bristles, and an apical
whorl of transparent spiked plates well discernible on
the lateral sides of the flagellomere. The antennae are
short and straight. The palpi are threesegmented. The
wings are narrow. The
R
1+2
vein reaches the wing edge
far beyond the middle of the wing and is 0.625 as long
as the wing. The
R
4+5
vein reaches the wing edge at the
wing apex, and the costal vein extends beyond the wing
apex. The medial vein is poorly developed but discern
ible over the entire length. The cubital veins form an
acute angle. The femora, tibiae, and tarsi are almost
equal in length and covered with abundant setae. The
abdomen is thin, with the basal part slightly inflated,
and 2.7 times longer than the thorax; the apical plates
are threesegmented.
Male unknown.
Species composition. Type species.
C o m p a r i s o n. The new genus is similar to the
large extant genus
Monardia
in size, body and tarsus
shape, slightly inflated rounded pedicel, and slightly
Explanation of Plate 12
Fig. 1.
Zherikhiniella pedicellata
sp. nov., holotype PIN, no. 3311/3142, male: (1a) general appearance,
×
70.4; (1b) partial head
with antenna,
×
200; (1c) legs,
×
92.9;
Fig. 2.
Zherikhiniella pedicellata
sp. nov., paratype PIN, no. 3311/3147, male: (2a) general appearance,
×
157.9; (2b) head, thorax,
and legs,
×
158.8;
(2c) palpi,
×
428.6.
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1017
Plate 12
0.2 mm
1a 2a
1b
2c
2b
1c
0.2 mm
1018
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
Plate 13
0.2 mm 0.2 mm
0.05 mm
1a 2a
2b
1b 2c 1c
Explanation of Plate 13
Fig. 1.
Menssana norilsk
sp. nov., holotype PIN, no. 3311/3146, female: (1a) head, antenna, and wings,
×
75.3; (1b) antenna,
×
269.2); (1c) legs viewed from below,
×
100.
Fig. 2.
Zherikhiniella pedicellata
sp. nov., paratype PIN, no. 3311/3145, male: (2a) general appearance,
×
96.9; (2b) protarsus,
×
174.2; (2c) head,
×
175.
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1019
convex notum, location of eyes on the anterior side of
the head, narrow eye bridge, presence of a necklike
constriction, short neck, and characteristic apical sen
sory plates on female flagellomeres; threesegmented
apical plates of the ovipositor, and foursegmented
palpi. The distinctive features of the new genus are its
very small body, elongated fragment of the
R
1+2
vein
(from
Rs
to
C
), which is more than twice longer than
Rs
; straight
M
+
rm
vein; and 2 + 8segmented anten
nae in female, in contrast to the larger number of seg
ments (not less than 2 + 9, up to 35) in most of
Monar
dia
females.
Menssana norilsk
Fedotova et Perkovsky sp. nov.
Plate 13, fig. 1; Figs. 4e, 5d–5f
Etymology. From the city of Norilsk.
H o l o t y p e. PIN, no. 3311/3146, wellpreserved
male, Yantardakh, Taimyr amber; Santonian. Synin
clusion in PIN, no. 3311/3146a, antennae of Porri
condylinae (Holoneurini) gen. indet.
D e s c r i p t i o n. Female (Figs. 4e, 5d–5f). The
antennae are 2.1 times as long as the head and 0.38 as
long as the body. The wing is 1.2 times longer than the
body, broadly rounded and broadened in the proximal
part; the anal lobe of the wing is well developed. The
wing is 2.7 times as long as wide. The length of the
foursegmented palpi is less than the head width.
The foreleg is 0.625 as long as the body and 0.53 as
long as the wing. The femora, tibiae, and tarsi are
almost equal in length and covered with abundant
bristles. The ratios of tarsomere lengths in the protar
sus are 1 : 0.4 : 0.4 : 0.3 : 0.4; the first tarsomere is
2.8 times longer than the second. The sum length of
the 3rd, 4th, and 5th tarsomeres is equal to the length
of the first tarsomere of the protarsus. The
R
1+2
vein
contacts the wing edge of the far beyond the middle of
the wing and is 0.625 as long as the wing. The abdomen
is narrow, inflated slightly basally, 2.7 times as long as
the thorax. The top apical plate is elongated oval,
longer than two other plates.
M e a s u r e m e n t s, m m. Body length, 1.08;
head length, 0.19; head height, 0.22; thorax length,
0.29; thorax width, 0.20; antenna length, 0.41; abdo
men length, 0.79; wing length, 1.28; wing width, 0.48;
R
1+2
length, 0.81; procoxa length, 0.13; trochanter,
0.04; femur, 0.32; tibia, 0.33; first tarsomere, 0.14;
2nd, 0.05; 3rd, 0.05; 4th, 0.04; 5th, 0.05; protarsus,
0.33; foreleg, 0.66.
Material. Holotype.
Tr i b e Aprionini Jaschhof, 1998
The tribe includes five genera and 25 extant species
(Gagné and Jaschhof, 2014) as well as two fossil spe
cies from the Late Eocene Rovno amber,
Aprionus
admirandus
Fedotova and
A. improvisus
Fedotova et
Perkovsky (Fedotova and Perkovsky, 2007).
Larvae of a number of extant species of the genus
Aprionus
Kieffer dwell in decomposed wood, while
others occur in soil (at depths not exceeding 20 cm)
and forest litter. Larvae of
A. terrestris
Mamaev were
found on white rot inside beech wood; those of
A. fla
vidus
(Winnertz), on honeycomb rots inside fir wood;
those of
A. similis
Mamaev, on developing mycelium
under strongly decomposed spruce bark; those of
A. dentifer
Mamaev, in thick pine forest litter rich in
mycelium; and those of
A. miki
Kieffer, in spruce tree
stumps. Some species of
Mycophila
Felt damage
champignons (Mamaev and Krivosheina, 1965).
Two new genera discovered in the Taimyr amber
(see below for description) were assigned to the tribe
Aprionini based on the considerable length of the frag
ment of the
R
1+2
vein from
Rs
to
C
; the absence of a
fragment of the costal vein after the fusion of this vein
with
R
4+5
; the presence of 2 + 8segmented antennae
in female, the presence of a wellpronounced neck and
a globular basal enlargement on the flagellomeres, and
a threesegmented ovipositor.
Genus
Cretomycophila
Fedotova et Perkovsky gen. nov.
Type s p ecie s.
C. ekaterinae
sp. nov.
E t y m o l o g y. From the Cretaceous and the
name of the extant genus
Mycophila
Felt.
D e s c r i p t i o n. Male (Fig. 6). The body is very
short (but still longer than the wing), with a broad
inflated abdomen. The head is narrower than the tho
rax (Fig. 6a). The antennae are short, 1.8 times as long
as the head, with 2 + 10 segments. The flagellomeres
are short barrelshaped. The necks are poorly pro
nounced and look like rather necklike constrictions.
The mesoflagellomeres are only slightly longer than
wide, terminal ones are almost spherical (Figs. 6a, 6f).
The flagellomeres have a basal whorl of short setae and
a medial whorl composed of very long setae (Fig. 6f).
The length of the foursegmented palpi (Fig. 6a) is less
than the head height; the palpomeres are very thin and
the palpiger is well developed. The notum is very
strongly bulged. The wing is broad, shorter than the
body, with a broad anal cell; the anal lobe is not devel
oped and the ventral edge of the wing is broadly
rounded. The
R
4+5
vein contacts the wing edge consid
erably anterior to the wing apex;
M + rm
is straight and
does not form a bend with
Rs
. The
Rs
length is almost
equal to the distance from the point of contact of
Rs
and
R
1+2
to the costal vein. The costal vein continues
to the wing apex as a short fragment after connection
to the
R
4+5
vein (Fig. 6a), where there is a break at the
wing apex. The CuA
1
and CuA
2
veins form a short
acuteangled fork. The major stem of the cubital vein
and CuA
1
are considerably thickened. Neither curved
veins nor a broad cell formed by these veins are dis
cernible at the base of the wing. All legs are longer than
the body and wing. The first tarsomere is 1.5–2.0 times
1020
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
longer than the second. The claws are long, simple,
and slightly bent (Fig. 6g).
The abdomen is slightly inflated. The abdomen
with genitalia is longer than the head and thorax taken
together. The welldeveloped aedeagal complex
reaches (or almost reaches) the gonocoxite apex
(Fig. 6e). The genitalia include broad gonocoxites and
gonostyles, which are almost equal in width at the
junction. The gonostyles are shorter than the gono
coxites, broad oval, slightly broadened in the middle;
the apical claws of the gonocoxites are very thin. The
bilobed cerci reach the gonocoxite apices; the hypo
proct is truncated at a right angle.
Species composition. Type species.
C o m p a r i s o n. The features distinguishing the
new genus from the modern genera of the subfamily
Micromyinae include contact of the vein
R
4+5
with the
wing edge noticeably anterior to the wing apex and the
presence of a nick exactly at the wing apex. The char
acters distinguishing the new genus from
Mycophila
include even shorter necks of male flagellomeres, a
shorter eye bridge consisting of 5 or 6 facets (rather
than 1–3 facets in
Mycophila
), the presence of a broad
cell between the
R
1+2
and
R
4+5
veins, a slightly curved
CuA
2
vein, a relatively long first tarsomere, and four
segmented palpi.
Cretomycophila ekaterinae
Fedotova et Perkovsky sp. nov.
Plate 14; Fig. 6
E t y m o l o g y. In honor of Ekaterina A. Sidor
chuk, who collected the material.
(b)
(c), (d) (e), (g)
(a)
(f)
(g)
(e)
(d)
(b)
(c)
(a)
(f)
Fig. 6.
Cretomycophila ekaterinae
sp. nov., holotype PIN, no. 3311/2588, male: (a) general habitus; (b) metatibia and tarsus; (c)
protibia and tarsus; (d) mesotibia and tarsus; (e) genitalia; (f) antenna; (g) metatarsus (shape variability).
Explanation of Plate 14
Figs. 1–8.
Cretomycophila ekaterinae
sp. nov., holotype PIN, no. 3311/2588, male: (1) general appearance,
×
55.7; (2) head, palpi,
genitalia, antennae, and four tarsomeres of protarsus,
×
160.8; (3) mesotibia and tarsus,
×
136.7; (4) hind leg and tarsus, meso and
profemora and tibia,
×
111.4; (5) wing and mesotibia and tarsus,
×
104.3; (6) notum, neck, partial head, and palpus,
×
123.7;
(7) genitalia, ventral view,
×
283.3; (8) genitalia, dorsal view,
×
283.3.
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1021
Plate 14
500
µ
m
4
1500
µ
m
8
200
µ
m
200
µ
m
7
3
2
5
6
500
µ
m500
µ
m
1022
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
H o l o ty p e. PIN, no. 3311/2588, wellpreserved
male, Yantardakh, Taimyr amber; Santonian.
D e s c r i p t i o n. Male (Fig. 6). The head is round,
slightly elongated, with the eyes covering the entire
lateral sides of the head. The head is 0.67 as long as
wide and 0.29 as long as the body (genitalia not
included) as (Fig. 6a). The wing is 0.91 as long as the
body. The antennae are 1.8 times longer than the head.
The pedicel is globular and slightly broader than
flagellomeres. The antennae have 2 + 10 segments;
flagellomeres 1–7 have very short necks, while others
lack a neck (Figs. 6a and 6f). The 5th flagellomere is
1.4 times as long as wide, with a neck 7.5 times smaller
than the basal enlargement. The medial whorl consists
of very long abundant bristles (Fig. 6f). The length of
the foursegmented palpi is less than the head height.
The midlegs are longer than the forelegs, but
shorter than the hind legs. The hind leg is 1.3 times
longer than the foreleg. The pro and mesofemora are
longer than the tibiae, while the metafemora are equal
in length to the metatibiae. The tarsi of all legs are
longer than the tibiae. The protarsi and metatarsi are
considerably longer than the femora, while the meso
tarsi are only slightly longer than the femora. The
metatarsi are 1.3 times longer than the pro and meso
tarsi. The length ratios of tarsomeres of the protarsus
are 1 : 0.6 : 0.5 : 0.4 : 0.4, the first tarsomere is 1.7 times
longer than the second. The sum length of the 3rd,
4th, and 5th tarsomeres of the protarsus is 1.4 times
greater than the first tarsomere. The length ratios of
mesotarsomeres are 1 : 0.5 : 0.9 : 0.8 : 0.8 and the sum
length of the 3rd, 4th, and 5th tarsomeres of the meso
tarsus is 1.2 times greater than the first tarsomere. The
length ratios of metatarsomeres are 1 : 0.5 : 0.4 : 0.3 : 0.3
and the sum length of the 3rd, 4th, and 5th tarsomeres
of the metatarsus is 1.8 times greater than the first tar
somere length. The wing width is maximum near the
middle of the wing. The wing is 1.9 times as long as
wide; the ventral wing edge is fringed by long hairs.
The
R
1+2
vein contacts the wing edge far beyond the
middle of the wing and is 0.55 times as long as the
wing. The
R
4+5
vein contacts the wing edge far before
the middle of the wing and forms a broad cell along
with the costal vein. The medial vein is indiscernible.
The abdomen is inflated at the base and slightly longer
than the thorax. The genitalia are broadened trans
versely. The lateral sides of the gonocoxites are
rounded. The gonostyles are shorter than the gono
coxites, strongly inflated, with small apical claws
(Fig. 6e).
Female unknown.
M e a s u r e m e n t s, m m. Body length, 0.85
(genitals not included), 0.95 (genitals included); head
length, 0.23; head height (mouthparts not included),
0.15; antenna length, 0.42; length of the 5th flagellom
ere, 0.12; wing length, 0.75; wing width, 0.39; abdo
men length, 0.39 (genitalia not included); 0.44 (geni
tals included); thorax length, 0.36; thorax width, 0.32;
foreleg length, 0.90; coxa 0.08; trochanter, 0.03;
femur, 0.26; tibia, 0.22; tarsus, 0.31; midleg length,
0.95; coxa, 0.07; trochanter, 0.03; femur, 0.29; tibia,
0.26; tarsi, 0.30; hind leg length, 1.19; coxa, 0.08; tro
chanter, 0.05; femur, 0.33; tibia, 0.33; tarsus, 0.40; first
tarsomere, 0.16; 2nd, 0.07; 3rd, 0.07; 4th, 0.05;
5th, 0.05; width of the genitalia, 0.31.
Material. Holotype.
Genus
Corporesana
Fedotova et Perkovsky gen. nov.
Type s p ecie s.
C. khatanga
sp. nov.
E t y m o l o g y. From the second part of the Latin
proverb
Mens sana in corpore sano
(a sound mind in a
sound body).
D e s c r i p t i o n. Male (Figs. 7a–7i). The body is
covered with narrow scales. The occipital part of the
head is protruding and broadly rounded. The eye
bridge is narrow. The wing is only slightly longer than
the body. The antennae have 2 + 11 segments. The first
and second flagellomeres are longer than the others.
Necks of all flagellomeres are short. The length of the
threesegmented palpi is less than head width. The
mouthparts slightly protrude beyond the head edge
(Fig. 7a).
The thorax is broad. The inflated notum protrudes
over the occipital part of the head. The pro, meso,
and metacoxae are almost equal in length. The tro
chanters are rounded and less than half as long as the
coxae. The femora of the forelegs and midlegs are as
long as respective tibiae, while the metafemora are
1.2 times as long as the metatibiae. The pro, meso,
and metafemora are almost equal in width. The tarsi of
all legs are longer than the tibiae. The first tarsomere of
the pro and metatarsi is twice longer than respective
second tarsomere, whereas in the midleg, this ratio is
1.5. The hind legs are 0.91 times as long as the forelegs.
The wing lacks an anal lobe. The costal vein ends at the
point of fusion with the R
4+5
vein. The abdomen is
slightly and uniformly inflated and much narrower
than the thorax. The genitalia are very large, almost
equal in width to the abdomen in lateral view and the
gonocoxites and gonostyles are large and strongly
inflated.
C o m p ar i s o n. The new genus is similar to
Apri
onus
Kieffer in the presence of strongly elongated
gonocoxites diverging in an almost conical pattern,
narrow eye bridge, location of the eyes (shifted
towards the anterior side of the head), 2 + 11seg
mented antennae, curved
M
+
rm
vein, and narrow
scales covering the body. The distinctive features of the
new genus include the presence of a long fragment of
the
R
1+2
vein (from Rs to the point of fusion with the
costal vein), thickened marginal and cubital veins of
the wing, termination of the costal vein at the point of
fusion with
R
4+5
, elongated flagellomeres 1 and 2, very
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1023
large genitalia, and elongated forelegs (as compared to
hind legs).
Corporesana khatanga
Fedotova et Perkovsky sp. nov.
Plate 15; Fig. 7
Etymology. From the settlement of Khatanga.
H o l o t y p e. PIN, no. 3311/2085, wellpreserved
male. Yantardakh, Taimyr amber; Santonian.
D e s c r i p t i o n. Male (Figs. 7a–7i). The head is
1.3 times as long as wide (mouthparts not taken into
account). The wing is 1.1 times longer than the body.
The antennae are 0.91 times as long as the body and
0.83 times as long as the wing; the pedicel is almost
spherical. The 5th flagellomere is 3.6 times as long as
wide, and the oval neck is 2.8 times smaller than the
basal enlargement. The 12th flagellomere is ovate. The
third palpomere is slightly longer than the others.
The mouthparts slightly protrude beyond the head
edge (Fig. 7a).
The thorax is 1.2 times as long as wide. The inflated
notum protrudes over the occipital part of the head.
The sum length of the 3rd, 4th, and 5th tarsomeres
exceeds the length of the first tarsomere, except for the
metatarsi, in which the lengths are equal. The wing is
2.0 times as long as wide. The
R
1+2
vein contacts the
wing edge far beyond the middle of the wing and is
0.59 times as long as the wing. The abdomen is slightly
and uniformly inflated and much narrower than the
thorax. The strongly inflated gonocoxites show a con
ical protrusion from the base. The length of the gono
styles slightly exceeds the gonocoxite width.
Measurements, mm. Body length (genitalia
not included), 0.66, (genitalia included), 0.74; head
length, 0.18; head height, 0.14; antenna length, 0.59;
wing length, 0.73; wing width, 0.36; procoxa length,
0.19; trochanter, 0.04; femur, 0.26; tibia, 0.26; first
tarsomere, 0.12; 2nd tarsomere, 0.06; 3rd, 0.07;
4th, 0.05; 5th, 0.05; protarsus, 0.35; foreleg, 1.10;
mesocoxa length, 0.10; trochanter, 0.03; femur, 0.23;
(b)
(c)
(a)
(g)–(i)
(a)
(b)–(f)
(h)
(d)
(g)
(i)
(e)
(f)
Fig. 7.
Corporesana khatanga
sp. nov., holotype PIN, no. 3311/2085, male: (a) general habitus; (b) wing; (c) foreleg; (d) hind leg;
(e) midleg; (f) protarsus; (g) flagellomeres 4–7 (shape variability); (h) head, scapus, pedicel, and flagellomeres 1 and 2;
(i) antenna.
1024
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
Plate 15
500
µ
m
7
6
5
1
2200
µ
m
34
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FIRST GALL MIDGES (DIPTERA, CECIDOMYIOIDEA) 1025
tibia, 0.23; first tarsomere, 0.09; 2nd tarsomere, 0.06;
3rd, 0.06; 4th, 0.06; 5th, 0.04; mesotarsus, 0.31; mid
leg, 0.90; metacoxa length, 0.07; trochanter, 0.03;
femur, 0.38; tibia, 0.31; first tarsomere, 0.14; 2nd tar
somere, 0.07; 3rd, 0.06; 4th, 0.06; 5th, 0.04; tarsus,
0.37; leg, 0.97; genitalia length, 0.12.
M a t e r i a l. Holotype.
ACKNOWLEDGMENTS
We are grateful to A.P. Rasnitsyn (PIN) for help
during the study and photographing gall midges in the
amber, to V.Yu. Nazarenko and S.A. Simutnik
(Schmalhausen Institute of Zoology, National Acad
emy of Sciences of Ukraine) for help with photo
graphing gall midges, to A.P. Vlaskin (Schmalhausen
Institute of Zoology, National Academy of Sciences of
Ukraine) for primary polishing and labeling the sam
ples, to E.A. Sidorchuk, D.S. Kopylov (PIN) and
D.D. Vorontsov (Institute of Developmental Biology,
Russian Academy of Sciences), who collected Taimyr
amber samples with gall midges in 2012, to V.V. Mar
tynov (Slavyansk), who recorded all gall midges in the
samples collected in 2012, and to E.A. Sidorchuk and
A.P. Rasnitsyn for sample polishing.
REFERENCES
Arillo, A. and Nel, A., Two new fossil cecidomyiids flies
from the Lower Cretaceous amber of Alava (Spain)
(Diptera, Cecidomyiidae),
Bull. Soc. Entomol. Fr.,
2000,
vol. 105, pp. 285–288.
Fedotova, Z.A.,
Gallitsyfitofagi (Diptera, Cecidomyiidae)
pustyn’ i gor Kazakhstana: morfologiya, biologiya, raspros
tranenie, filogeniya i sistematika
(Phytophagous Gall
Midges (Diptera, Cecidomyiidae) of Deserts and Moun
tains of Kazakhstan: Morphology, Biology, Distribution,
Phylogeny, and Taxonomy), Samara: Samar. Gos. Sel.
Khoz. Akad., 2000.
Fedotova, Z.A., Classification of gall midges of the tribe
Aphidoletini (Diptera, Cecidomyiidae, Aphidoletidi) and
description of new genus and species from the Kuril Islands,
Entomol. Obozr.,
2013, vol. 42, no. 4, pp. 823–848.
Fedotova, Z.A., Classification of gall midges of the super
tribe Brachineuridi (Diptera, Cecidomyioidea, Cecidomyi
idae, Lasiopterinae) and description of new genera and spe
cies the Kuril Islands,
Entomol. Obozr.,
2014, vol. 93, no. 3,
pp. 666–739.
Fedotova, Z.A. and Perkovsky, E.E., New gall midges
(Diptera, Cecidomyiidae) from the Rovno amber: Subfam
ily Lestremiinae, tribes Strobliellini, Campylomyzini; sub
family Porricondylinae, tribes Diallactini, Asynaptini,
Paleontol. J.,
2004, no. 5, pp. 538–547.
Fedotova, Z.A. and Perkovsky, E.E., New gall midges
(Diptera, Cecidomyiidae) from the Rovno amber: Subfam
ily Porricondylinae, tribes Bryocryptini and Winnertziini;
subfamily Lasiopterinae, tribes Brachineurini and Oligotro
phini,
Paleontol. J.,
2005, no. 1, pp. 41–51.
Fedotova, Z.A. and Perkovsky, E.E., New taxa of gall
midges of the subfamily Lestremiinae (Diptera, Cecid
omyiidae) from the Rovno amber,
Paleontol. J.,
2007, no. 4,
pp. 437–450.
Fedotova, Z.A. and Perkovsky, E.E., New gall midges of the
tribe Leptosynini (Diptera, Cecidomyiidae) from the Late
Eocene ambers and the classification of the supertribe Het
eropezidi,
Paleontol. J.,
2009, vol. 43, no. 9, pp. 1101–1179.
Fedotova, Z.A. and Perkovsky, E.E., New gall midges
(Diptera, Cecidomyiidae, Stomatosematidi, Brachineu
ridi) from Late Eocene amber of Gulyanka (Zhitomir
Region),
Paleontol. J.,
2015, no. 3, pp. 270–278.
Gagné, R.J., Cecidomyiidae (Diptera) from Canadian
amber,
Proc. Entomol. Soc. Washington,
1977, vol. 79,
pp. 57–62.
Gagné, R.J., A catalog of the Cecidomyiidae (Diptera) of
the World,
Mem. Entomol. Soc. Washington,
2004, vol. 25,
pp. 1–408.
Gagné, R.J., Update for a catalog of the Cecidomyiidae
(Diptera) of the world: Digital version 1.2010;
http://www.ars.usda.gov/SP2UserFiles/Place/12754100/G
agne_2010_World_Catalog_Cecidomyiidae.pdf. (accessed
23 Nov. 2010).
Gagné, R.J. and Jaschhof, M.,
A Catalog of the Cecidomyi
idae (Diptera) of the World,
3rd ed.; Digital version 2.
ZooBank registration (1/1/2014): urn:lsid:zoobank.
org:pub:2FC82C5E40FD47EDB6F1BEC0DFFB776D
Jaschhof, M., Catotrichinae subfam. n.: A reexamination
of higher classification in gall midges (Diptera: Cecidomyi
idae).
Entomol. Sci.,
2001, vol. 3, pp. 639–652.
Jaschhof, M. and Jaschhof, C., The wood midges (Diptera:
Cecidomyiidae: Lestremiinae) of Fennoscandia and Den
mark,
Stud. Dipterol. Suppl.,
2009, vol. 18, pp. 1–333.
Kovalev, V.G., Diptera: Muscida,
Tr. Paleontol. Inst. Akad.
Nauk SSSR,
1990, vol. 239 (Late Mesozoic Insects of East
ern Transbaikalia), pp. 123–177.
Krivosheina, N.P. and Zaitsev, A.I., Phylogeny and evolu
tionary ecology of Diptera,
Itogi Nauki Tekhn. Ser. Ento
mol.,
1989, vol. 9, pp. 1–144.
Lambkin, C.L., Sinclair, B. J., Pape, T., et al., The phyloge
netic relationships among infraorders and superfamilies of
Diptera based on morphological evidence,
Syst. Entomol.,
2013, vol. 38, pp. 164–179.
Mamaev, B.M. and Krivosheina, N.P.,
Lichinki gallits
(Diptera, Cecidomyiidae). Sravnitel’naya morfologiya,
biologiya, opredelitel’nye tablitsy
(Larval Gall Midges
(Diptera, Cecidomyiidae): Comparative Morphology, Biol
ogy, Keys), Moscow: Nauka, 1965.
Explanation of Plate 15
Figs. 1–7.
Corporesana khatanga
sp. nov., holotype PIN, no. 3311/2085, male: (1) general appearance,
×
75.7; (2) wing,
×
300;
(3) abdomen, genitalia, and legs,
×
125.2; (4) palpi, foreleg, and midleg,
×
169.2; (5) flagellomeres 4–6,
×
305.9; (6) antennae,
×
123.7; (7) genitalia, caudal view,
×
75.
1026
PALEONTOLOGICAL JOURNAL Vol. 50 No. 9 2016
FEDOTOVA, PERKOVSKY
Matile, L., Phylogeny and evolution of the larval diet in the
Sciaroidea (Diptera, Bibionomorpha) since the Mesozoic,
Mém. Mus. Nat. d’Hist. Natur.,.
1997, vol. 148 (The Origin
of Biodiversity in Insects: Phylogenetic Tests of Evolution
ary Scenarios), pp. 273–303.
Möhn, E., Studien an paedogenetischen Gallmückenarten
(Diptera, Itonididae): Teil 1,
Stuttgarter Beitr. Naturk.,
1960, no. 31, pp. 1–11.
Perkovsky, E.E. and Fedotova, Z.A., New species of gall
midges (Diptera, Cecidomyiidae) from the Rovno amber:
Subfamily Lestremiinae, tribes Micromyiini and Peromyi
ini,
Paleontol. J.,
2004, no. 4, pp. 396–406.
Perkovsky, E.E. and Fedotova, Z.A., New taxa of gall
midges of the subfamilies Porricondylinae and Lasiopteri
nae (Diptera, Cecidomyiidae) from the Rovno amber,
Paleontol. J.,
2008, no. 2, pp. 166–175.
Skuhravá, M., Family Cecidomyiidae, in
Catalogue of
Palaearctic Diptera: SciaridaeCecidomyiidae,
Budapest:
Acad. Kiado, 1986, vol. 4, pp. 72–297.
Skuhravá, M., 2.7. Family Cecidomyiidae, in
Contribution
to a Manual of Palaearctic Diptera (with Special Reference to
Flies of Economic Importance): Nematocera and Lower
Brachycera,
Budapest: Science Herald, 1997, vol. 2,
pp. 71–204.
Zherikhin, V.V. and Sukacheva, I.D.,
O melovykh naseko
monosnykh yantaryakh (retinitakh) severa Sibiri
(On Creta
ceous Insectbearing Ambers (Retinites) of Northern Sibe
ria), Leningrad: Nauka, Leningr. Otd., 1973.
Translated by S. Semenova
