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, OCTOBER 2014, 65, 5, 365—374 doi: 10.2478/geoca-2014-0025
Revision of Anomopterella Rasnitsyn, 1975
(Insecta, Hymenoptera, Anomopterellidae) with two new
Middle Jurassic species from northeastern China
LONGFENG LI, CHUNGKUN SHIH and DONG REN
College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, 100048 Beijing, China; rendong@mail.cnu.edu.cn
(Manuscript received May 7, 2014; accepted in revised form October 7, 2014)
Abstract: New Anomopterella Rasnitsyn, 1975, wasps from the Jiulongshan Formation (Bathonian/Callovian) at
Daohugou (Inner Mongolia of China), with well-preserved wings, body, antennae and tarsi, document relationship
among species and enhance our understanding of the morphology of the extinct family Anomopterellidae Rasnitsyn,
1975. Based on eight well-preserved specimens, we describe, A. pygmea sp. n. and A. brevis sp. n., describe new
materials of A. ovalis Li, Rasnitsyn, Shih & Ren, 2013, and A. ampla Li, Rasnitsyn, Shih & Ren, 2013, emend the genus
in Anomopterellidae (A. pygmea sp. n. with toruli and eyes), and provide a determination key to species of the genus,
including four specimens outside the formal classification.
Key words: Fossil insect assemblage, new species, extinct parasitoids, Middle Jurassic, Jiulongshan Formation,
Daohugou, China.
Introduction
Anomopterellidae, a Mesozoic family of wasps, was origi-
nally classified as a family of the superfamily Evanioidea,
and later as a subfamily of the family Praeaulacidae (Ras-
nitsyn 1975). In 2013, Anomopterellidae was restored as a
family with three genera: Anomopterella Rasnitsyn, 1975,
Synaphopterella Li, Rasnitsyn, Shih & Ren, 2013 and Cho-
ristopterella Li, Rasnitsyn, Shih & Ren, 2013, on the basis
of new fossil specimens from Daohugou and phylogenetic
analysis of the superfamily Evanioidea (Li et al. 2013a).
These analyses also suggest that the extinct family Praeaula-
cidae is the most basal group of Evanioidea, and Anomo-
pterellidae is the second family of Evanioidea present in the
upper Middle Jurassic of China (Li et al. 2013a). Anomo-
pterella Rasnitsyn, 1975 was erected with the type species A.
mirabilis Rasnitsyn, 1975 from the Upper Jurassic of the
Karabastau Formation in Kazakhstan (Rasnitsyn 1975).
In addition to the type species A. mirabilis from the Upper
Jurassic Kazakhstan, A. gobi Rasnitsyn, 2008 occurs in the
Upper Jurassic of Mongolia, while A. huangi Zhang & Ras-
nitsyn, 2008, A. coalita Li, Rasnitsyn, Shih et Ren, 2013, A.
brachysteli Li, Rasnitsyn, Shih & Ren, 2013, A. ampla Li,
Rasnitsyn, Shih & Ren, 2013, A. divergens Li, Rasnitsyn,
Shih & Ren, 2013 and A. ovalis Li, Rasnitsyn, Shih & Ren,
2013 occur in the Upper Middle Jurassic Jiulongshan Forma-
tion at Daohugou in northeastern China (Rasnitsyn 1975,
2008; Zhang & Rasnitsyn 2008; Li et al. 2013a). The species
from Daohugou indicate that this genus varies in the wing
venation (e.g. forewing with crossvein 1r-rs is absent or rudi-
mentary like a stub; crossvein cu-a is interstitial or slightly
postfurcal) and in the shape of the first metasomal segment
(e.g. broad triangular, elongated triangular and particularly
narrow with a short petiole).
In this study, we describe eight new well-preserved fossil
specimens of Anomopterella from the Middle Jurassic Jiulong-
shan Formation at Daohugou Village in Inner Mongolia,
China, which belongs to the Yanliao Biota (Ren et al. 2010,
2012). The Daohugou locality contains abundant and diverse
insect assemblages, comprising complete specimens of
Ephemeroptera (reference missing), Plecoptera (Liu et al.
2007), (Huang et al. 2008), Mantophasmatodea (Huang et al.
2008), Coleoptera (Chang et al. 2009), Trichoptera (Wang et
al. 2009; Gao et al. 2013a,b), Megaloptera (Wang & Zhang
2010), Nakridletia (Vršanský et al. 2010), Lepidoptera (Hua-
ng et al. 2010; Zhang et al. 2013), Dermaptera (Shang et al.
2011; Zhao et al. 2011), Grylloblattodea (Cui et al. 2011,
2012), Orthoptera (Gu et al. 2012), Heteroptera (Dong et al.
2012), Odonata (Li et al. 2013d), Blattaria (e.g. Wei et al.
2013), Homoptera (Li et al. 2013c), Mecoptera (Wang et al.
2013), Neuroptera (Shi et al. 2013), Diptera (Shi et al. 2013)
and Hymenoptera (Li et al. 2013a,b). According to the Ar-Ar
and SHRIMP U-Pb dating, the Jiulongshan Formation corre-
sponds to the upper parts of the Middle Jurassic (Bathonian/
Callovian boundary, 165 Ma – Ren et al. 2012).
The specimens reported herein are from the Middle Juras-
sic Jiulongshan Formation of Daohugou Village, Shantou
Township, Ningcheng County, Chifeng City, Inner Mongo-
lia, China (Fig. 1). The age of the Daohugou fossil-bearing
beds is likely older than or equal to 165 Ma (Chen et al.
2004).
On the basis of the new characters of four of the newly
collected specimens, we established A. pygmea sp. n., A.
brevis sp. n., and re-describe A. ovalis, and A. ampla. The
additional four specimens remain outside the formal species-
level classification within the genus Anomopterella. We
emend the diagnosis of Anomopterella Rasnitsyn, 1975 and
add a taxonomic key to all known species.
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Materials and methods
This study is based on eight specimens collected near Dao-
hugou Village and housed in the Key Lab of Insect Evolu-
tion and Environmental Changes, the College of Life
Sciences, Capital Normal University in Beijing, China
(CNUB; Dong Ren, Curator). The specimens were examined
and photographed using a Leica MZ12.5 dissecting micro-
scope with a Leica DFC 500 digital camera and illustrated
with the aid of a camera lucida attached to the microscope.
The line drawings were drawn by CorelDraw 12.0 and Adobe
Photoshop CS5.
Wasp wing venation terminology is adapted from Rasnitsyn
(1975), following the general concept of Comstock and
Needham (1898): C – Costa, R – Radial, Rs – Radial sec-
tor, M – Media, Cu – Cubitus, 1A – the first Anal, 1r-rs –
the first Radial crossvein, 2r-rs – the second Radial cross-
vein, 3r-m – the second Radiomedial crossvein, 1m-cu – the
first Mediocubital crossvein, 2m-cu – the second Mediocu-
bital crossvein, cu-a – the first Anal crossvein.
The Daohugou fossil-bearing beds consist of tuffaceous
grand conglomerates, tuffaceous conglomerates, tuffaceous
siltstones, tuffaceous mudstones, tuffaceous shales, and vol-
canic breccias. In these layers, the layer of a tuffaceous con-
glomerate contains abundant fossils. The paleoclimate of
this area is usually interpreted as humid and warm-temperate
(Ren et al. 2010, 2012).
Systematic paleontology
Order: Hymenoptera Linnaeus, 1758
Suborder: Apocrita Gerstaecker, 1867
Superfamily: Evanioidea Latreille, 1802
Family: Anomopterellidae Rasnitsyn, 1975
Genus: Anomopterella Rasnitsyn, 1975
T y p e s p e c i e s : Anomopterella mirabilis Rasnitsyn, 1975.
T y p e l o c a l i t y : Karatau, south Kazakhatan. Late Jurassic
Karabastau Formation.
Differential diagnosis (improved after Rasnitsyn
1 9 7 5 ) : Body size under 8 mm. Head medium size, usually
as wide as or narrower than thorax. Antenna with 18 or more
segments. When preserved, legs long, tarsus with five seg-
ments. Mesosoma short and high. Propodeum high, strongly
convex or quite short, with posterior margin subvertical in
lateral view. Forewing venation with 1r-rs absent or rudi-
mentary like a stub, M + Cu not aligned with RS + M, 1-M
present, 2r-rs and 2m-cu basad of 3r-m. Vein cu-a interstitial
or slightly postfurcal. Metasoma broadest beyond its mid-
length, with the first segment gradually broadening distally
and longer than any other segments. Ovipositor short and
only slightly extending beyond metasomal apex, with sheaths
shorter than basal sclerite.
C o m p o s i t i o n : Type species: A. huangi, A. gobi, A. co-
alita, A. brachysteli, A. ampla, A. divergens, A. ovalis, A.
pygmea sp. n. and A. brevis sp. n.
S t r a t i g r a p h i c a n d p a l e o g e o g r a p h i c r a n g e : Up-
per Jurassic Karabastau Formation in south Kazakhstan (A.
mirabilis), Upper Jurassic Shar-Teg Formation in southwest-
ern Mongolia (A. gobi); upper Middle Jurassic Jiulongshan
Formation at Daohugou in northeastern China (A. huangi, A.
coalita, A. brachysteli, A. ampla, A. divergens, A. ovalis, A.
pygmea and A. brevis sp. n.).
R e m a r k s : Anomopterella was erected based on 2 speci-
mens (Rasnitsyn 1975). After studying ten (six with part and
counterpart) well-preserved fossil specimens attributed to
Anomopterella, we added diagnostic characters of the head,
body and forewings (Li et al. 2013a). In this paper, after
studying an additional eight specimens (one with part and
counterpart), we improve diagnostic characters with well-pre-
served body, complete antenna and tarsus of legs. A. pygmea
sp. nov. is the first species of Anomopterellidae with clearly
preserved toruli and eyes.
Anomopterella pygmea sp. n.
(Figs. 2, 4A)
D e r i v a t i o n o f n a m e : From the Latin word “pygmea”
meaning “scrubby”, referring to the small body size of this
wasp.
T y p e m a t e r i a l : CNU-HYM-NN-2012-045, a well-pre-
served complete female (Fig. 2A).
T y p e h o r i z o n a n d l o c a l i t y : Near Daohugou Vil-
lage, Shantou Township, Ningcheng County, Inner Mongolia,
China. Jiulongshan, the late Middle Jurassic.
D i f f e r e n t i a l d i a g n o s i s : Antenna with 23 segments.
Propodeum short, slightly narrower than metanotum or meta-
postnotum. Forewing with 1-Rs originating more distal of
pterostigma base, 1r-rs absent, 3-Rs longer than 3r-m, cu-a in-
terstitial. Metasoma short with first metasomal segment parti-
cularly narrow like a distinct petiole, remaining part round.
D e s c r i p t i o n : A female wasp preserved in dorsal view,
body and wings well preserved and veins clearly discernible.
Small body size (4.4 mm long, excluding antennae), 1.1 mm
wide (Fig. 2A). Head and mesosoma mostly dark.
Head round, nearly as long as wide (length 0.8 mm, width
0.9 mm). Compound eyes large and symmetrically located
on both sides of the head; two ocelli round with diameter of
about 0.1 mm; antenna with 25 segments and gradually thin-
ner from base to apical; two toruli present and in contact
Fig. 1. Map showing the locality at Daohugou Village (Inner Mon-
golia, China) with the Middle Jurassic Jiulongshan Formation.
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with radicles; scape long and thickened apically, distinctly
longer than radicle, but slightly longer than pedicel (scape
length 0.25 mm; radicle length 0.12 mm, pedicel length
0.23 mm); flagellomeres with basal eleven segments nearly
as long as wide, twelfth to apex gradually becoming shorter
and narrower.
Mesosoma approximately 1.7 mm long and 1.1 mm wide,
about 1.6 times as long as wide; mesonotum long and broad,
but the boundary and shape of mesoscutum and scutellum
unknown; metanotum nearly as wide as metapostnotum,
both very short; propodeum longer than metanotum and
metapostnotum combined, but nearly as wide.
Legs partly preserved, but hind legs complete; hind femur
about 5.5 times as long as wide and nearly as wide as the
maximal width of tibia; hind tibia thin basally and gradually
swollen toward apex, nearly as long as hind femur; hind tar-
sus consisting of 5 segments, much narrower than hind tibia,
first segment distinctly longer than any other segments of
hind tarsus.
Wings well preserved, forewing 3.6 mm long and 1.5 mm
wide, with 1-Rs (0.3 mm) longer than 1-M (0.2 mm) and
much longer than its distance (0.1 mm) to pterostigma;
pterostigma long, about 4.5 times as long as wide (length
0.9 mm, width 0.2 mm); 1r-rs absent; 2r-rs (0.4 mm) issuing
from pterostigma nearly at the apex of it, about 2 times as long
as the width of pterostigma; 3-Rs longer than 3r-m (3-Rs
length 0.3 mm, 3r-m length 0.2 mm); cell 3r broad, 2.4 times
as long as wide (length 1.2 mm, width 0.5 mm); cell 2 + 3rm
wider than 2mcu, in contact with 1mcu by a point; cell 1mcu
nearly rectangle, 2 times as long as wide (length 0.4 mm,
width 0.2 mm); cu-a interstitial (0.3 mm long) and curved,
nearly as long as 1-Rs; cell cua broad, 2 times as wide as
1mcu. Hind wing 1.8 mm long and 0.6 mm wide, with 1-Rs,
1-M and Rs + M forming “Y-shape”, 1-Rs nearly as long as 1-M
(both length about 0.1 mm), cu-a present and curved.
Metasomal short with 6 segments (excluding propodeum),
first metasomal segment particularly narrow like a distinct
petiole, elongate triangular, 1.5 times as long as wide (length
0.6 mm, maximum width 0.4 mm); remaining part round in
dorsal view, the second segment transversally ellipse, dis-
tinctly shorter and wider than first; the third segment slightly
shorter and wider than the second, the fourth to sixth seg-
ments gradually becoming shorter and narrower.
D i m e n s i o n s ( i n m m ) : Body length 4.4; length of head
0.8, width 0.9; length of antenna 2.8; length of mesosoma
1.7, width 1.1; length of forewing 3.6, width 1.5; length of
hind wing 1.8, width 0.6; length of first metasomal segment
0.6, width 0.4, length of second metasomal segment 0.4,
width 1.1; length of remaining metasomal segments 1.1,
maximal width 1.4; length of hind leg: femur 1.2, tibia 1.2,
tarsomeres I—V: 0.24, 0.22, 0.17, 0.17, 0.15.
Anomopterella brevis sp. n.
(Figs. 3A—B, 4B)
Derivation of name: From the Latin word “brevis”
meaning “short”, referring to the forewing with 3-Cu very
short.
Type material: CNU-HYM-NN-2012-043P/C (Fig. 3A—B).
T y p e h o r i z o n a n d l o c a l i t y : Near Daohugou Village,
Shantou Township, Ningcheng County, Inner Mongolia,
China. Jiulongshan, the late Middle Jurassic.
D i f f e r e n t i a l d i a g n o s i s : Forewing with 1-Rs origin
quiet away from pterostigma base, 1r-rs rudimentary like a
short stub, cu-a postfurcal, 3-Cu short, about 1/2 length of
1m-cu. First metasomal segment short, comparatively thin
basally but broadened apically.
D e s c r i p t i o n o f n e w m a t e r i a l : Body mostly dark. A
wasp preserved in dorsal view, sex unknown, body and
wings partly preserved but veins clearly discernible. Small
body size (3.7 mm long as preserved, excluding antennae),
1.6 mm wide (Fig. 3A—B).
Head round, nearly 0.6 times as long as wide (length
0.5 mm, width 0.8 mm). Antenna with about 23 segments
and gradually thickened from base to tenth; scape short and
thick, distinctly longer and wider than radicle; flagellomeres
with basal ten segments much longer than wide, the maxi-
mum length about 2.7 times as long as the maximum width,
eleventh to apex gradually becoming shorter and narrower.
Mesosoma broad, nearly as long as wide. Legs partly pre-
served, fore tibia distinctly shorter and narrower than hind
tibia, hind femur slightly wider than hind tibia, hind tibia
thin basally and gradually swollen toward apex.
Wings well preserved, forewing 4.0 mm long and 1.6 mm
wide, with 1-Rs (0.4 mm) distinctly longer than 1-M
(0.1 mm); 1r-rs present, like a very short stub on Rs, Rs gen-
iculate; 2r-rs (0.4 mm) issuing from pterostigma nearly at the
apex, about 2 times as long as the width of pterostigma; 3-Rs
nearly as long as 3r-m (lengths of both 0.2 mm); cell 3r broad,
2.3 times as long as wide (length 1.4 mm, width 0.6 mm);
cell 2 + 3rm nearly as wide as 2mcu, in contact with 1mcu by
a point; cell 1mcu nearly rectangle, about 2.5 times as long
as wide (length 0.5 mm, width 0.2 mm); cu-a postfurcal
(0.3 mm long) and curved. Hind wing with 1-Rs, 1-M and
Rs + M forming “Y-shape”, 1-Rs (0.2 mm) longer than 1-M
(0.1 mm), cu-a present and curved.
Metasomal short with 5 segments as preserved (excluding
propodeum), first metasomal segment short, comparatively
thin basally but broadened apically (length 0.6 mm, maxi-
mum width 0.6 mm); remaining part round in dorsal view,
the second segment transversally triangular, distinctly wider
than first; the remaining segments distinctly shorter than the
second.
Di m e n s i o n s ( i n m m ) : Body length 3.7; length of head
0.5, width 0.8; length of antenna 3.4; length of mesosoma
1.6, width 1.6; length of forewing 4.0, width 1.6; length of
first metasomal segment 0.7, maximum width 0.6.
Anomopterella ovalis Li, Rasnitsyn, Shih & Ren, 2013
(Figs. 3C, 4C)
2013 Anomopterella ovalis Li, Rasnitsyn, Shih & Ren, p. 12, fig. 8
N e w m a t e r i a l : CNU-HYM-NN-2012-044, collected
near Daohugou Village, Shantou Township, Ningcheng
County, Inner Mongolia, China, the Middle Jurassic. The new
material is considered as A. ovalis Li, Rasnitsyn, Shih & Ren,
2013 mainly due to the following characters: forewing with
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Fig. 2. Anomopterella pygmea sp. n., Holotype CNU-HYM-NN-2012-045. A – Habitus; B – Interpretation of head examined under alcohol;
C – Interpretation of head examined in the absence of alcohol. Scale bars: 1 mm.
1r-rs rudimentary, like a very short stub on Rs, 3-Rs long, cu-a
postfurcal and first metasomal segment elongate triangular.
D e s c r i p t i o n o f n e w m a t e r i a l: A wasp preserved in
dorsal view, body and wings partly preserved but veins
clearly discernible. Small body size (5.2 mm long, excluding
antennae), 1.1 mm wide (Fig. 3C).
Head round, 2/3 times as long as wide (length 0.6 mm,
width 0.9 mm). Antenna with about 23 segments and gradu-
ally thickened from base to tenth; scape short and thick,
nearly as long and wide as radicle; flagellomeres with basal
ten segments with nearly equal width, eleventh to apex grad-
ually becoming shorter and narrower.
Mesosoma long and narrow, approximately 2.0 mm long
and 1.1 mm wide, about 1.8 times as long as wide. Legs partly
preserved, hind femur slightly wider than hind tibia, hind
tibia thin basally and gradually swollen toward apex.
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Fig. 3. Anomopterella brevis sp. n., Holotype, CNU-HYM-NN-2012-043P/C. A – Part; B – Countpart; C – Anomopterella ovalis Li, Ras-
nitsyn, Shih & Ren, 2013, CNU-HYM-NN-2012-044; D – Anomopterella ampla Li, Rasnitsyn, Shih & Ren, 2013, CNU-HYM-NN-2012-047.
Anomopterella spp.; E – CNU-HYM-NN-2012-029; F – CNU-HYM-NN-2012-046; G – CNU-HYM-NN-2012-048; H – CNU-HYM-
NN-2012-049. Scale bars: 1 mm.
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Wings well preserved, forewing 4.0 mm long and 1.8 mm
wide, with 1-Rs (0.4 mm) longer than 1-M (0.2 mm); 1r-rs
rudimentary, like a very short stub on Rs; 2r-rs (0.4 mm) is-
suing from pterostigma nearly at the apex; 3-Rs nearly as
long as 3r-m (both with lengths of about 0.3 mm); cell 3r
broad, 2.3 times as long as wide (length 1.4 mm, width
0.6 mm); cell 2 + 3rm slightly wider than 2mcu, in contact
with 1mcu by a point; cu-a postfurcal (0.3 mm) and curved.
Hind wing 2.3 mm long and 0.8 mm wide, with 1-Rs, 1-M
and Rs + M forming “Y-shape”, 1-Rs nearly as long as 1-M
(both about 0.1 mm long), cu-a present and curved.
Metasoma nearly as long as mesosoma, with 5 segments
as preserved (excluding propodeum), first metasomal seg-
ment distinctly longer than other segments, comparatively
thin basally and gradually broadened apically forming an
elongate triangular (length 0.9 mm, maximum width
0.5 mm); remaining part oval in dorsal aspect, the second
segment transversally trapezoid; the remaining segments dis-
tinct shorter than the second.
D i m e n s i o n s ( i n m m ) : Body length 5.2; length of head
0.6, width 0.9; length of antenna 3.7; length of mesosoma
2.0, width 1.1; length of forewing 4.0, width 1.8; length of
Fig. 4. A – Anomopterella pygmea sp. n. Line drawing of holotype, CNU-HYM-NN-2012-045; B – Anomopterella brevis sp. n. Line draw-
ing of holotype, CNU-HYM-NN-2012-043P/C; C – Anomopterella ovalis Li, Rasnitsyn, Shih & Ren, 2013, Line drawing of CNU-HYM-
NN-2012-044; D – Anomopterella ampla Li, Rasnitsyn, Shih & Ren, 2013, Line drawing of CNU-HYM-NN-2012-047; Scale bars: 1 mm.
hind wing 2.3, width 0.8; length of first metasomal segment
0.8, width 0.5.
Anomopterella ampla Li, Rasnitsyn, Shih & Ren, 2013
(Figs. 3D, 4D)
2013 Anomopterella ampla Li, Rasnitsyn, Shih & Ren, p. 10, fig. 6
N e w m a t e r i a l : CNU-HYM-NN-2012-047, collected
near Daohugou Village, Shantou Township, Ningcheng
County, Inner Mongolia, China, the Middle Jurassic. The new
material is considered to be A. ampla Li, Rasnitsyn, Shih &
Ren, 2013 mainly due to the following characters: forewing
with 1r-rs absent, 3-Rs distinctly longer than 3r-m, cu-a inter-
stitial, and first metasomal segment broad-triangular.
D e s c r i p t i o n o f n e w m a t e r i a l : A wasp preserved in
dorsal view, body and wings partly preserved but veins
clearly discernible. Small body size (4.3 mm long, excluding
antenna), 1.3 mm wide (Fig. 3D).
Head transversely ovate, distinctly wider than long (length
0.5 mm, width 0.8 mm). Compound eyes large and symmet-
rically located on both sides of the head. Antenna with about
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21 segments as preserved, and gradually thickened from
base to tenth; scape thick, wider than pedicel; flagellomeres
with basal ten segments with nearly equal width, eleventh to
the last gradually becoming shorter and narrower.
Mesosoma approximately 1.5 mm long and 1.0 mm wide,
about 1.5 times as long as wide; pronotum short, covered by
mesonotum; mesonotum long and broad, transscutal suture
present; metanotum with small metascutellum and narrower
than metapostnotum; metapostnotum very short; propodeum
shorter and narrower than metapostnotum.
Legs partly preserved, but hind legs complete; hind femur
(0.9 mm) shorter than hind tibia (1.1 mm); hind tibia thin ba-
sally and gradually swollen toward apex; hind tarsus consisting
of 5 segments, much narrower than hind tibia, the first seg-
ment distinctly longer than any other segments of hind tarsus.
Wings well preserved, forewing 3.4 mm long and 1.3 mm
wide, with 1-Rs (0.3 mm) longer than 1-M (0.2 mm); 1r-rs not
distinct present and 2-Rs angular at its place; 2r-rs (0.3 mm)
issuing from pterostigma nearly at the apex, about 2 times as
long as the width of pterostigma; 3-Rs longer than 3r-m (3-Rs
length 0.3 mm, 3r-m length 0.2 mm); cell 3r broad, 2.4 times
as long as wide (length 1.2 mm, width 0.5 mm); cell 2 + 3rm
wider than 2mcu, in contact with 1mcu by a point; cell 1mcu
nearly rectangle, 2 times as long as wide (length 0.4 mm,
width 0.2 mm); cu-a interstitial (0.2 mm long) and curved.
Hind wing with 1-Rs, 1-M and Rs + M forming “Y-shape”,
1-Rs shorter than 1-M (1-Rs length 0.1 mm; 1-M length
0.2 mm), cu-a present and curved.
Metasomal short with 6 segments (excluding propodeum),
first metasomal segment short, comparatively thin basally
but broadened apically (length 0.5 mm, maximum width
0.5 mm); remaining part ovate in dorsal view, the second
segment transversally ellipse, distinctly wider than the first;
the remaining segments distinctly shorter than the second.
D i m e n s i o n s ( i n m m ) : Body length 4.3, width 1.3;
length of head 0.5, width 0.8; length of antenna 2.8; length of
mesosoma 1.5, width 1.0; length of forewing 3.4, width 1.3;
length of first metasomal segment 0.5, maximum width 0.5;
length of hind leg: femur 0.9, tibia 1.1, tarsomeres I—V: 0.37,
0.18, 0.12, 0.08, 0.10.
Key to species of Anomopterella Rasnitsyn, 1975, modi-
fied and improved according to Li et al. (2013a):
1. Forewing with cu-a interstitial......................................2
– Forewing with cu-a postfurcal......................................5
2. Forewing with Rs originating close to pterostigma, Rs
very short between 2r-rs and 3r-m..........................................
......................................................A. gobi Rasnitsyn, 2008
– Forewing with Rs originating far from pterostigma,
Rs (3-Rs) long between 2r-rs and 3r-m, 1r-rs absent ..............3
3. Both propodeum and metasoma short, first metasomal
segment particularly narrow like a distinct petiole (length/
width ratio 1.7), remaining part round....................................
..................................................................A. pygmea sp. n.
– Both propodeum and metasoma long, first metasomal
segment triangular................................................................4
4. First metasomal segment elongate triangular (length/
width ratio between 1.2 and 1.6).............................................
.........................A. coalita Li, Rasnitsyn, Shih & Ren, 2013
– First metasomal segment broad triangular (length/
width ratio 1.1).......................................................................
..........................A. ampla Li, Rasnitsyn, Shih & Ren, 2013
5. 1-RS originating from pterostigma for a distance com-
parable with length of 1m-cu. Ovipositor long, far surpassing
metasomal apex. First metasomal segment broad triangular
(length/width ratio 1.0)..........................................................
...............................................A. mirabilis Rasnitsyn, 1975
– 1-RS originating far from pterostigma. Ovipositor short,
weakly extending beyond metasomal apex...........................6
6. Rudimentary 1r-rs about as long as 1m-cu. First metaso-
mal segment particularly narrow with distinct petiole
(length/width ratio 1.7)..........................................................
.................A. brachystelis Li, Rasnitsyn, Shih & Ren, 2013
– Rudimentary 1r-rs much shorter or lost. First metasom-
al segment elongate triangular..............................................7
7. Rudimentary 1r-rs absent................................................
....................A. divergens Li, Rasnitsyn, Shih & Ren, 2013
– Rudimentary 1r-rs dot-like small.................................8
8. First metasomal segment elongate triangular (length/
width ratio between 1.4 and 1.6). Legs shorter and slightly
wider......................................................................................
..........................A. ovalis Li, Rasnitsyn, Shih & Ren, 2013
– First metasomal segment short (length/width ratio be-
tween 0.9 to 1.2 )..................................................................9
9. Hind legs much longer and narrower, hind femur thin.
Forewing with 3-Cu long, nearly as long as 1m-cu................
...................................A. huangi Zhang & Rasnitsyn, 2008
– Hind legs longer and thicker, hind femur thicker.
Forewing with 3-Cu short, nearly 1/2 length of 1m-cu ..........
.....................................................................A. brevis sp. n.
Discussion
Presently, nine species of Anomopterella are recorded
(Table 1), suggesting (1) in total, 21 fossil specimens were
found in the Jurassic sediments, with 2 specimens from the
Upper Jurassic of Kazakhstan and Mongolia and 19 speci-
mens from the upper Middle Jurassic of China. The rarity of
Anomopterella specimens (only 19 specimens from a very
large Daohugou fossil insect collection at the CNU of more
than 250,000 specimens (Ren et al. 2010, 2012)) can imply
that source area population sizes of species belonging to
Anomopterella during the late Middle Jurassic were low rela-
tive to population sizes of other species. We did not find any
species of Anomopterella in the CNU’s similarly vast collec-
tion from the Lower Cretaceous Yixian Formation. These
differences in species richness and abundance between the
Middle Jurassic and Lower Cretaceous suggest that, Ano-
mopterella was more diverse and abundant during the late
Middle Jurassic than during the Early Cretaceous. (2) The 19
specimens from the late Middle Jurassic have a narrow range
of body lengths from 3.7 to 7.8, forewing lengths from 3.4 to
6.3, and the ratios of wing length/body length ranging from
0.7 to 1.1. Compared to the large body size of female pele-
cinids (Shih et al. 2010) in the Middle Jurassic ranged be-
tween 9 and 27 mm, the small body size of Anomopterella
highlighting that this group had a narrow range of these mor-
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Table 1:
A
list
of
species
of
Anomopterella
Rasnitsyn,
1975,
with
morphometric
characters,
and
information
about
their
sex,
age,
and
locality.
Abbreviations
of
material
ID
numbers:
CNU
–
Capital
Normal
University,
Beijing,
China;
HYM-NN
–
Hyme
noptera-Ningcheng,
Inner
Mongolia;
BL.
–
Body
length
(mm);
FL
.
–
Forewing
length (mm); S
1
L
. – First metasomal segment length (mm); S
1
W. – First metasomal segment width (mm); F – female; M – male;
J
2
– Middle Jurassic; J
3
– Late Jurassic; NK – unknown.
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phological differences and smaller body size may have some
advantages, such as agility, flexibility, stealth and energy
conservation when they find and deposit eggs onto defence-
less surface larval hosts (Shih et al. 2010).
New and complete specimens show that Anomopterella
significantly varied in wing venation and in the shape of the
first metasomal segment. Nearly complete wing venation, in-
cluding C, Rs, M, Cu, 2r-rs, 3r-m and 2m-cu is always
present. All species of Anomopterella have 2r-rs and 2m-cu
basad of 3r-m, indicating that this vein character is very sta-
ble for this genus and can be considered diagnostic and con-
servative. On the other hand, veins of all species of
Anomopterella show many differences, such as cu-a postfur-
cal (vs. cu-a interstitial), 1r-rs absent (vs. rudimentary 1r-rs,
dot-like and small), 3-Rs long (vs. 3-Rs short), which can be
considered as interspecific differences. In addition to the dif-
ferences in veins, the first metasomal segment of Anomo-
pterella is morphologically diverse, with a very broad range
of the ratio of length/width from 0.9 to 1.7. Also, the first
segment of the metasoma presents different shapes, such as
broad triangular, elongate triangular and particularly narrow
with a distinct short petiole.
We stress that toruli and compound eyes in Anomopterel-
lidae are reported for the first time in our study, namely in A.
pygmea sp. n. In general, insects have compound eyes and
ocelli. Compound eyes are the most important visual organ
of insects, which can mostly identify moving objects. How-
ever, ocelli do not have the imaging function, and are only
used to sense the strength and direction of the light (Cai et al.
2001). The main function of ocelli is to control the swing
and tilt in the process of flight, and sense the change of the
light cycle associated with circadian behaviour (Cai et al.
2001). The central ocellus is known to recognize the position
of insects, and is especially well-developed in predatory
mantodeans and extinct cockroaches (Vršanský 2008). De-
formities (Vršanský 2005) are rarely reported in hy-
menopteran wings at Daohugou, including a single deformed
specimen of Anomopterellidae (CNU-HYM-NN-2012-022;
Li et al. 2013a).
Conclusion
A. pygmea sp. n., A. brevis sp. n., and six other new fossils
emend the records of the Mesozoic family Anomopterel-
lidae, and enhance our understanding of the morphology of
the extinct family Anomopterellidae. New and complete
specimens show that Anomopterella significantly varied in
wing venation and in the shape of the first metasomal seg-
ment. The differences in species richness and abundance be-
tween the Middle Jurassic and Lower Cretaceous suggest
that, Anomopterella was more diverse and abundant during
the late Middle Jurassic than during the Early Cretaceous.
Acknowledgments: We sincerely thank three anonymous
reviewers and the editor for their useful comments and im-
provements of this manuscript. Research was supported by
the National Basic Research Program of China (973 Pro-
gram) (2012CB821906), the National Natural Science Foun-
dation of China (Nos. 31071964, 31172143, 31230065,
41272006) and China Geological Survey (1212011120116);
Great Wall Scholar and KEY project of the Beijing Municipal
Commission of Education (KZ201310028033), Program for
Changjiang Scholars and Innovative Research Team in Uni-
versity (IRT13081).
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