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, AUGUST 2013, 64, 4, 291—304 doi: 10.2478/geoca-2013-0021
Introduction
Blattaria (i.e., cockroaches), important as ecological decom-
posers, are one of the phylogenetically old insect groups. Fos-
sil cockroaches occurred for the first time in the Late
Carboniferous (Brongniart 1885; Zhang et al. 2012), and gave
birth to two different orders Isoptera and Mantodea (Vršanský
2002, 2010). They adapted to multiple types of ecosystems
and reached a high level of ecological and morphological di-
versity (disparity) over the course of their evolution, includ-
ing aquatic (Shelford 1909), pollinating (Nagamitsu & Inoue
1997), jumping (Vršanský 2007; Bohn et al. 2010; Picker
et al. 2012), light-mimicking (Zompro & Fritzsche 1999;
Vršanský et al. 2012), translucent (Vidlička et al. 2003),
predatory (Vishniakova 1973; Liang et al. 2009, 2012), beetle-
like (Vršanský 2003a), and eusocial (Vršanský 2010) forms.
Cockroaches are composed of 27 families in their
320 Myr-long history. The oldest reproductively and sensory-
advanced family of cockroaches, the Mesoblattinidae,
evolved during the Early Jurassic (Vršanský 2000, 2003b;
Vršanský & Ansorge 2007). Nevertheless, their diversity at
the genus level is low, and only 12 genera are known in their
history. The morphological differences among genera are
mostly based on wings or on characters preserved in amber
(Mesoblattina Geinitz, 1880, Praeblattella Vršanský, 2003b,
Archimesoblatta Vršanský, 2003b, Brachymesoblatta Vršan-
ský, 2003b, Breviblattina Vršanský, 2004a, Mongolblatta
Vršanský, 2004a, Turoniblatta Vršanský, 2004b, Gondwa-
blatta Vršanský, 2004b, Nymphoblatta Vršanský & Grimaldi,
2004, Sivis Vršanský, 2009, Hispanoblatta Martínez-
Delclòs, 1993, Artitocoblatta Meunier, 1914) (Geinitz 1880;
Completely preserved cockroaches of the family
Mesoblattinidae from the Upper Jurassic-Lower Cretaceous
Yixian Formation (Liaoning Province, NE China)
DANDAN WEI and DONG REN
College of Life Science, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing, 100048, P.R. China;
rendong@mail.cnu.edu.cn
(Manuscript received January 1, 2013; accepted in revised form March 14, 2013)
Abstract: Although cockroaches were the dominant insects in various Paleozoic and Mesozoic insect assemblages,
their general morphology was extremely conservative. One of the most common of them, the Jurassic-Cretaceous
family Mesoblattinidae, is described here for the first time on the basis of completely preserved specimens. Ninety-two
specimens of Perlucipecta aurea gen. et sp. n. reveal details of head, mandible, male tergal glands and terminal hook;
cercal, leg and antennal sensilla. Its congener, P. vrsanskyi is described from the same sediments of the Yixian Forma-
tion (Upper Jurassic—Lower Cretaceous). The forewing venation variability of P. aurea, analysed for the first time in
this family is nearly identical (CV = 6.23 %) with variability of two species of family Blattulidae that occur at the same
locality (CV = 6.22 %; 5.72 %). The transitional nature of morphological characters represented by asymmetry between
left and right wings (simple/branched forewing SC and hind wing M) in P. aurea documents the phylogenetic relation
between the families Mesoblattinidae and Ectobiidae.
Key words: Jurassic, Cretaceous, Liaoning Province, fossil insects, Blattaria, Ectobiidae, Mesoblattinidae, new genus, new
species.
Meunier 1914; Martínez-Delclòs 1993; Vršanský 2003b,
2004a,b, 2009). A large numbers of taxa described and
placed in this family in the past were transferred into Calo-
blattinidae (Vršanský 2000).
Well-preserved complete fossil specimens with ultrastruc-
ture details and colouration recovered from the Yixian For-
mation, belonging to Perlucipecta gen. n. and described in
this study, are significant because they represent the first
complete Mesozoic cockroach recovered from sediments
with such details, and also correspond to the first members
of the Mesoblattinidae preserved in the Yixian Formation.
The aim of this paper is to describe the morphological struc-
tures of the specimens, and compare them with representa-
tives of their descendants, the Cretaceous—Recent family
Ectobiidae ( = Blattellidae).
We test whether their morphology and microstructures are
similar to the morphology of another family Ectobiidae, and
compare coefficients of variation in the number of veins with
other Jurassic cockroaches. The venation variability coeffi-
cients observed in these specimens, comparable to variability
of species belonging to the family Blattulidae preserved at
the same locality is surprising because variability trends are
traceable in respective phylogenetic lineages and not in re-
spective localities (and possibly in specific environments).
Material and methods
A total of ninety-two specimens of P. aurea and ten speci-
mens of P. vrsanskyi are deposited in the fossil insect collection
of the Key Laboratory of Insect Evolution & Environmental
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Changes, Capital Normal University, Beijing, China. The
specimens were examined with a Leica MZ 12.5 dissecting
microscope and illustrated with the aid of a drawing tube at-
tached to the microscope. Line drawings were made with
Photoshop CS 8.0 graphic software. Photographs of the fos-
sils were taken by a MZ12.5 dissecting microscope (Leica,
Wetzlar, Germany), either dry or with alcohol applied to the
surface (to reveal contrast and organic matter). The coefficient
of variation was computed as the standard deviation in the
number of veins divided by the average number of veins, on
the basis of thirty-nine specimens with complete forewings,
using SPSS 16.0 Statistical software. We refer to the total
number of veins meeting the wing margin.
All specimens were found in a section exposing the Jian-
shangou Bed of the Yixian Formation at Chaomidian village,
Beipiao City, western Liaoning Province, China. This area is
situated in a volcanic sediment basin (Fig. 1). There are at
least three thick fossil-bearing beds formed by tuffaceous
shales within the Yixian Formation, including the Jianshangou
Bed located in the lower part of Yixian Formation. Three
sections that expose the Jianshangou Bed are represented by
Libalanggou-Sihetun, Jianshangou, and Huangbanjigou
(Chen et al. 2004). The material in this study came from sev-
eral unidentified beds of the Huangbanjigou section that span
dozens of meters. This section previously revealed a large
number of well-preserved insects (Liu et al. 2007a,b; Tan et al.
2007; Gao & Ren 2008). The age of the Yixian Formation is
still debated, with three opinions: the Late Jurassic (based on
insects and primitive fossil birds – Ren et al. 1997), transi-
tional Late Jurassic—Early Cretaceous (based on diverse in-
sects, pterosaurs, ostracods and fish – Wang et al. 2004,
2005), and the Early Cretaceous (based on ostracods – Pang
et al. 2002; Zhou et al. 2003). Here, we consider the age of the
Yixian Formation to be Late Jurassic or Early Cretaceous. The
climate and the approximate paleolatitude of the location dur-
ing the depositon of the Jianshangou Bed was subtropical hu-
mid as indicated by the cockroaches considered here.
The venation nomenclature used in this paper is based on the
terminology of Comstock & Needham (1898). Abbrevia-
tions used: RFW – Right forewing, LFW – Left forewing,
HW – Hind wing, Sc – Subcosta, R – Radius, Rs – Radius
sector, M – Media, Cu – Cubitus (A – anterior, P – poste-
rior), A – Anal vein.
Systematic paleontology
Order: Blattaria Latreille, 1810
Family: Mesoblattinidae Handlirsch, 1906
Genus: Perlucipecta Wei & Ren, gen. n.
T y p e s p e c i e s : Perlucipecta aurea sp. n.
C o m p o s i t i o n : Type species and P. vrsanskyi sp. n.
D i f f e r e n t i a l d i a g n o s i s : The present genus can be as-
signed to the family Mesoblattinidiae based on the following
features: branched forewing Sc; hindwing simple Sc, Rs dif-
ferentiated, simplified M, CuA with secondarily branched
veins and blind rami.
It differs from all the other representatives of the family by
the following features: expanded venation with retained in-
tercalaries and distinct cross-veins; large size; facultative
simplification of hindwing M.
D e s c r i p t i o n : Head inverted-triangular shape, extending
beyond the pronotum.
Pronotum elliptical, transversal (width longer than length),
transparent near edges and coloured in the center.
Abdomen broad, with dark coloured longitudinal stripes.
Cerci gradually pointed to the end, 9 segments and with al-
ternating dark and pale colouration.
Forewing long, with rich venation (more than 42 branches
at margin), Sc bifurcated, R rich and stem straight, M with
abundant branches, CuA sparse, reaching the posterior wing,
CuP radian concave and then convex to the posterior wing,
A rich, simple, costal space shorter and smaller than clavus.
Hind wing Sc simple, R and Rs rich and branched, M sim-
ple or simply branched, CuA with abundant dichotomized
branches and blind rami.
Character list
Head: inverted-triangular shape, extending beyond the
pronotum slightly; eyes protruding beyond the outline of
head: synapomorphic with living cockroaches;
Antennal sockets conspicuous at sides; antenna slender,
almost as long as the body; mouthparts hypognathous, man-
dible strong with sharp teeth: synapomorphic with living
cockroaches;
Sensilla throughout the antennal segment: unknown;
Palps (maxillary and labial) long: homoplastic with
Caloblattinidae;
Fig. 1. Localities of the Yixian Formation in Jinzhou-Yixian area,
western Liaoning (modified after Ren et al. 2010). Sihetun section
(thickness: 264.95 m), Jianshangou section (thickness: 79.46 m).
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Pronotum: large, wide (as wide as the body), almost 3
times wider than the head; elliptical, parallel edge; transparent
margin and with dark markings at center: symplesiomorphic
at the level of Phyloblattoidea;
Body wide (with markings at margin), slightly shorter
than forewing: autapomorphic;
Tergal glands present: synapomorphic at the level of
Caloblattinidae or earlier;
Cerci: with reduced number of segments (under 10) and
alternating colouration: synapomorphic at the level of the
family;
Cercal sensilla small circle, present at the margin of each
segment: synapomorphic at the level of the family;
Legs: slender, with rich spines: symplesiomorphic at the
level of Phyloblattoidea;
Forewing with distinct markings only from the center to
the posterior end: autapomorphic;
Forewing with distinct intercalaries: symplesiomorphic
at the level of advanced Phyloblattoidea;
Forewing with distinct crossvein-like reticulations: sym-
plesiomorphic at the level of advanced Phylloblattoidea;
Forewing costal area narrow: synapomorphic with ad-
vanced Liberiblattinidae and Blattellidae, homoplastic with
Blattulidae;
Forewing Sc short and branched: symplesiomorphic
(level of earliest cockroaches);
Forewing Rs indistinctly differentiated: unknown;
Forewing venation rich: symplesiomorphic (level of ear-
liest cockroaches);
Forewing medial area wide and expanded: symplesio-
morphic (level of earliest cockroaches);
Forewing cubital area narrow and reduced: autapomorphic;
Forewing clavus long (longer than Sc) and wide: not dia-
gnostic;
Forewing A simple and rich: synapomorphic with Blat-
tellidae, simplicity is homoplastic with Blattulidae;
Hindwing Rs differentiated: symplesiomorphic at the
level of Phyloblattoidea;
Hindwing M facultatively simple: synapomorphic with
Blattellidae;
Hindwing CuA multiply branched: symplesiomorphic at
the level of Liberiblattinidae.
E t y m o l o g y : Perlucipecta is a combination of perlucidus
(Latin for transparent) and pectus (Latin for thorax), refer-
ring to its specially transparent (in the edge) and coloured (in
the middle) pronotum; gender feminine.
Perlucipecta aurea sp. n.
(Figs. 2—7, 10a,c,e,g,i,k; Table 1)
H o l o t y p e : CNU-B-NN-2011610pc (part and counter-
part), a well-preserved complete specimen (Figs. 2a, 3b).
T y p e l o c a l i t y : Huangbanjigou, Chaomidian village,
Beipiao City, Liaoning Province, North-east China.
T y p e h o r i z o n : Yixian Formation, Upper Jurassic or
Lower Cretaceous.
A d d i t i o n a l
m a t e r i a l : CNU-B-NN-2011600, 601,
603—606, 609, 611—613, 615, 617, 622, 627, 628, 631—635,
638, 640, 641, 643, 644, 646—649, 652, 657, 658, 660—666,
668—671, 673pc—674, 675, 677—680, 683—692, 694, 697—699,
701, 703, 706, 712, 715, 732, 733, 737, 739, 742—744pc,
747pc, 748, 749, 754, 757, 759, 774, 784—790, 795.
E t y m o l o g y : The specific name is derived from the Latin
word “aureus” – (meaning “golden”), for the colour of the
Yixian Formation sediment.
D i f f e r e n t i a l d i a g n o s i s : The species differs from its
congener P. vrsanskyi by distinct markings under forewing
R region; the shape of a more simple tergal gland (convex to
the abdominal segment) (Fig. 2e), larger size and the shape
of dark markings in pronotum.
Description: Middle-sized, with body length 8.9—13.2 mm
(excluding pronotum), length of abdomen 9.0—12.5 mm,
width of abdomen 3.5—6.0 mm. Head small, inverted-trian-
gular shape, slightly longer than wide, slightly extending
beyond the pronotum (length/width 1.9—2.2/2.4—2.8 mm)
(Figs. 2h, 3a); antennal socket and compound eye obvious
at sides, antenna length 12.4—13.5 mm, with 76—77 very
short segments (Figs. 2b, 10k); antenna sensilla throughout
the antenna segment (Figs. 2c, 3g); mandibles strong
(Fig. 2i, Fig. 3d).
Pronotum breadthwise elliptical, broad (length/width
2.9—4.5/4.5—5.5 mm), transparent by the edge and with dark
markings in the middle; abdomen broad, with 6—8 segments
distinct, with dark markings at margin; gland length
0.4—0.7 mm, width 1.2—1.5 mm, convex to the abdominal
segment (Figs. 2e, 10a,c,e,g,i); terminalia smooth, with cerci
gradually to the end (9 segments) and alternating colouration
(Fig. 2f), cercus sensilla preserved as small circles at the
margin of each segment (Figs. 2g, 3e).
F o r e w i n g : Length 12.0—17.0 mm, width 4.0—5.6 mm;
broad, with distinct markings only from the middle to poste-
rior end (Figs. 2a, 3b, 4—7), with intercaleries and numerous
crossveins; venation rich, with 43—54 veins at margin,
forewing sensilla present in the veins (Fig. 2l); Sc short and
Sc
R
M CuA A Sc+R R+M R+Cu M+Cu SUM Total
Number
36
36
32
30
33
33
30
28
32
32
29
Minimum
1
18
7
3
6
20
26
23
14
35
43
Maximum
4
24
15
10
10
26
37
31
22
46
54
Median
2.00
20.00
11.00
5.00
7.00
23.00
32.00
26.00
17.00
39.50
47.00
Mode
2
20
11
5
7
22
31
26
17
39
46
Std. error of mean 0.13
0.29
0.37
0.27
0.20
0.31
0.49
0.44
0.37
0.47
0.55
Std. deviation
0.77
1.71
2.08
1.40
1.16
1.75
2.65
2.32
2.09
2.68
2.96
CV in %
31.68
8.28
18.48
28.21
15.88
7.53
8.3
8.72
12.13
6.23
6.67
Table 1: Perlucipecta aurea sp. n. Variability of forewing venation. CV – coefficient of variability; Sc – subcosta; R – radius; M – me-
dia; CuA – cubitus anterior; A – anal veins; SUM – A excluded; Total – total number of veins.
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Fig. 2. Drawings of Perlucipecta aurea sp. n. (collection CNU-B-NN-2011). a – Holotype 610; b – Antenna (606); c – Detail of antenna
sensilla (733); d – Legs (609); e – Ventral view with glands (600); f – Detail of terminalia with cerci (606); g – Detail of cerci sensilla
(640); h – Head (603); i – Detail of head mandible (759); j – Left forewing (length 14.0 mm) (687); k – Right forewing (length
17.0 mm) (784); l – Detail of forewing sensilla (640); m – Left forewing (length 13.5 mm) (670); n – Right forewing (length 15.0 mm)
(698); o – Right forewing (length 13.5 mm) (669); p — Right hind wing (length 13.0 mm) (789); q – Right forewing (length 13.9 mm)
(699); r – Right forewing (length 12.5 mm) (670); s – Left forewing (length 15.0 mm) (684); t – Hind wing (length 13.9 mm) (668).
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Fig. 3. Photographs of Perlucipecta aurea sp. n. (collection CNU-B-NN-2011). a – Head (603); b – Holotype 610; c – Legs (609);
d – Detail of head mandible (759); e – Detail of cercal sensilla (640); f – Detail of spines (664); g – Detail of antennal sensilla (733);
h – Detail of wing sensilla (640).
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Fig. 4. Drawings of Perlucipecta aurea sp. n. (collection CNU-B-NN-2011). a – 603 (Right forewing length 16.0 mm); b – 604 (Right
forewing length 15.5 mm); c – 605 (Left forewing length 15.5 mm); d – 691 (Left forewing length 14.6 mm); e – 703 (Left forewing
length 15.0 mm); f – 685 (Right forewing length 15.0 mm); g – 601 (Right forewing length 15.0 mm); h – 606 (Right forewing length
14.5 mm); i – 694 (Right forewing length 15.0 mm); j – 662 (Right forewing length 16.0 mm); k – 675 (Right forewing length
15.0 mm); l – 689 (Right forewing length 15.0 mm); m – 663 (Left forewing length 16.0 mm); n – 664 (Left forewing length 16.5 mm);
o – 609 (Left forewing length 15.0 mm). Scale bar = 2 mm.
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Fig. 5. Photographs of Perlucipecta aurea sp. n. (collection CNU-B-NN-2011). a—f – (703, 694, 675, 689, 664, 609).
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branched, with 1—4 veins reaching anterior wing margin at
narrow space, Sc area shorter than clavus; R stem straight
reaching wing apex, with regular branching pattern reaching
anterior wing margin, posteriorly pectinate with secondary
forks, with 18—24 branches; main stem of M parallel to R;
medial area wide, with 7—15 terminal branches; cubital area
narrow and gently curved, with 3—10 branches reaching the
posterior wing; CuP with radian convex to the posterior
wing near 40 % of wing length, terminal almost with bend-
Fig. 6. Photographs of Perlucipecta aurea sp. n. (collection CNU-B-NN-2011) a—f – (691, 604, 662, 605, 663, 603).
ing; clavus broad, more than 50 % of wing width; A gently
curved, with 6—10 veins, some reaching CuP.
H i n d w i n g : Length 11.0—13.9 mm; with intercalaries
and without pterostigma; with about 22—29 veins of rem-
igium; Sc simple; R1 (2—4) and the Rs (8—14) are differenti-
ated; M simple or bifurcated near margin, almost straight to
wing apex; CuA with about 9—11 branches.
L e g s : Slender (Figs. 2d, 3c); length of fore femora 1.16—
1.49 mm and tibiae 1.16—1.22 mm, length of mid femora
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Fig. 7. Photographs of Perlucipecta aurea sp. n. (collection CNU-B-NN-2011). a—f – (789, 669, 759, 670, 784, 683, 668, 669, 715, 684,
601, 698, 606, 685, 687). Scale bar = 2 mm.
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Fig. 8. Drawings of Perlucipecta vrsanskyi sp. n. (collection CNU-B-NN-2011). a – Holotype 704; b – Detail of terminalia with hook
(794); c – Ventral view with glands (704).
Fig. 9. Photographs of Perlucipecta vrsanskyi sp. n. (collection CNU-B-NN-2011). a – Holotype 704; b – Head (727); c – Detail of ter-
minalia with hook (794).
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Fig. 10. Photographs of glands and antenna (collection CNU-B-NN-2011). a – Detail of P. aurea gland (600); b – Detail of P. vrsanskyi
gland (704); c – Detail of P. aurea gland (661); d – Detail of P. vrsanskyi gland (626); e – Detail of P. aurea gland (627); f — Detail of
P. vrsanskyi gland (654); g – Detail of P. aurea gland (706); h – Detail of P. vrsanskyi gland (770); i – Detail of P. aurea gland (671);
j – Detail of P. vrsanskyi gland (791); k – Detail of a complete antenna (606).
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1.89—1.93 mm and tibiae 1.47—1.56 mm, length of hind fem-
ora 2.12—2.19 mm and tibiae 2.59—3.45 mm; legs gradually
get longer from the front to the hind legs; mid and hind leg
with spines on the tibiae, and spines strong and hard, with
helical ridges (Fig. 3f).
Perlucipecta vrsanskyi sp. n.
(Figs. 8—9, 10b,d,f,h,j)
H o l o t y p e : CNU-B-NN-2011704, male, nearly complete
specimen (Figs. 8, 9).
T y p e l o c a l i t y : Huangbanjigou, Chaomidian village,
Beipiao City, Liaoning Province, North-east China.
T y p e h o r i z o n : Yixian Formation, Upper Jurassic or
Lower Cretaceous.
A d d i t i o n a l
m a t e r i a l : CNU-B-NN-2011626, 630,
654, 656, 716, 727, 770, 791, 794.
E t y m o l o g y : The species name vrsanskyi is after Peter
Vršanský, for the help he has given me for years.
D i f f e r e n t i a l d i a g n o s i s : Differing from P. aurea in
small size; pronotum colouration; shorter forewing; with
markings only in the centre (Fig. 8a); hind wing with
pterostigma; the shape of the gland (trapezoidal, parallel to
the abdominal segment) as shown in Fig. 8c.
Description: Small-sized, with body length 8.7—10.0 mm
(excluding pronotum), width of abdomen 3.1—3.8 mm. Head
small, inverted-triangular shape (length/width = 1.5 mm/
1.3 mm) (Fig. 9b), extending beyond the pronotum slightly;
pronotum length 1.9—3.0 mm, width 2.9—3.5 mm, with dark
markings at center; abdomen with 7—8 segments preserved,
wide in the middle of segments; gland length 0.6—0.7 mm,
width 0.9—1.0 mm, trapezoidal, parallel to the abdominal
segment (Figs. 8c, 10b,d,f,h,j); terminalia with cerci alternat-
ing coloration.
F o r e w i n g : Length 9.0—11.0 mm, width 2.9—3.0 mm, with
distinct markings in the center; with intercaleries and venation
rich, with 46 veins at margin; Sc short and with 2 branches, Sc
area shorter than clavus; R stem parallel reaching wing apex,
with regular branching pattern reaching anterior wing margin,
with 23 branches; main stem of M parallel to R; medial area
wide, with 10 terminal branches; cubital area narrow and gen-
tly curved, with 3 branches reaching the posterior wing; CuP
with radian convex; clavus broad, more than 50 % of wing
width; A gently curved, with 7 veins, some reaching CuP.
H i n d w i n g : Length 8.1 mm (as preserved); with interca-
laries and with pterostigma; with about 26 veins of rem-
igium; Sc simple; R1 (4) and the Rs (22) are differentiated;
M bifurcated near margin.
Discussion
With the exception of the Mesoblattinidae, the Blattulidae,
Liberiblattinidae and Caloblattinidae (Wang & Ren 2013)
families, each represented by a few genera and species, occur
in the deposits of the Yixian Formation (only the below-
mentioned Blattulidae were described). Mesoblattinidae with
ca. 200 specimens and 7 species (the two most common spe-
cies are described here) numerically dominate in the cock-
roach assemblage from the Yixian Formation. Such domi-
nance represents a standard pattern in all the Jurassic assem-
blages worldwide (Vršanský 2004a, 2008). The proportion
of the representative species P. aurea relative to all cock-
roaches approaches 10 % – a similar proportion as in the
Tithonian of the Shar-Teg locality in Mongolia and Cher-
novskie Kopi in Russia (Barna in print).
The Jurassic specimens of this family were predominantly
represented by tegmina. The specimens described here are
complete and thus reveal many details about the morphology
of the Mesoblattidae. The presence of two sister species dif-
fering mostly in the male tergal gland can imply that they oc-
cupied different source habitats. A similar set of sister species
is known in the Blattulidae (Vršanský 2003b), but nowhere
else in cockroaches. Cockroaches with sophisticatedly co-
loured parts imply that the habitats of the Yixian Formation
inhabited by these cockroaches were humid (Ding et al. 2003).
Dry habitats are rather characterized by monochromatic and
pale cockroach individuals.
Morphology (including microstructures, sexual glands and
terminalia) is extremely similar to Piniblattella vitimica
Vishniakova, 1964 of the family Ectobiidae ( = Blattellidae),
which were descendants of the Mesoblattinidae (see Vršanský
1997). Mesoblattinidae differ only in the following plesio-
morphies: rich venation with local irregularities and with
richly branched forewing Sc, specialized hind wing R1,
branched hind wing M and in primitive reproduction with
primitive ootheca.
The total number of forewing veins in Perlucipecta differs
between the left and right wings in the same specimens. For
example, it varies between 50 (right forewing) and 47 (left
forewing) in the specimen CNU-B-NN-2011606 and be-
tween 43 (right forewing) and 44 (left forewing) in the
specimen CNU-B-NN-2011610. These patterns show that
cockroaches are often asymmetrical in their venation, as ob-
served also in living individuals (Roth 1991), amber speci-
mens (Vršanský et al. 2011) as well as in other fossil
specimens preserved in sediments (Liang et al. 2012).
Intraspecific variability in wings (P. aurea) is revealed for
the first time in this family (Hispanoblatta as a representative
of the Blattellidae is not relevant). The coefficient of variation
(CV) in the total number of veins (per wing) of P. aurea is rel-
atively low (6.23 %). This can be associated with the higher
age of the group at that time (since it originated in the earliest
Jurassic), and fits well with the trend of the decreasing vari-
ability of insect species over time (Vršanský 2000). For com-
parison, this value is relatively similar to the CV of similar
Piniblattella vitimica from the Valanginian of Siberia of the
Ectobiidae (4.11 %). In contrast, the CV of species from the
Early Jurassic is 11.24 % (Vršanský & Ansorge 2007). These
coefficients of variation are also highly similar to those of spe-
cies belonging to a different cockroach family, the Blattulidae,
at the same locality (Elisama extenuata CV
sum
= 6.22 %;
Habroblattula drepanoides CV
sum
= 5.72 %) (Wang et al.
2007a,b).
To summarize, these exceptionally well preserved repre-
sentatives of the ecologically significant family Mesoblat-
tinidae extend the appearance of the advanced cockroach
characters such as details of reproductive tract and micro-
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structures. These specimens document the transitional char-
acter of the species between the two well-defined cockroach
families Mesoblattinidae and Ectobiidae (which had origi-
nated in or near the J/K boundary). Notably, this between-
family transition did not influence the taxonomic diversity
and/or ecological dominance of both groups during the Late
Jurassic—Early Cretaceous.
Acknowledgments: We sincerely thank Dr. ubomír Vidlička
and Dr. Peter Vršanský for revision of the manuscript, and
are grateful to Yingying Cui, Dr. Yunzhi Yao, Dr. Junjie Gu,
Qiang Yang, Taiping Gao, Chaofan Shi and other students of
our laboratory for help. This research is supported by the Na-
tional Basic Research Program of China (973 Program)
(2012CB821906), the National Natural Science Foundation
of China (No. 41272006, 31230065), the Project of Great
Wall Scholar and Key Program (KZ201310028033), and
China Geological Survey (1212011120115).
References
Barna P. in print: Low diversity cockroach assemblage from Cher-
novskie Kopi in Russia confirm wing deformities of insects at
J/K boundary. Biologia, Bratislava (in print).
Bohn H., Picker M. & Klass K.D. 2010: A jumping cockroach from
South Africa, Saltoblattella montistabularis, gen. nov., sp. nov.
(Blattodea: Blattellidae). Arthropod Systematic & Phylogeny 68,
1, 53—69.
Brongniart C. 1885: Les insectes fossiles des terrains primaires.
Coup d’oeil rapide sur la faune entomologique des terrains
paléozoïques. Bull. Soc. Amis Sci. Nat. Rouen 1885, 50—68.
Comstock J.H. & Needham J.G. 1898: The wings of insects. Amer.
Naturalist 32, 376, 231—257.
Chen P.J., Wang Q.F., Zhang H.C., Cao M.Z., Li W.B., Wu S.Q. &
Shen Y.B. 2004: Disussion on the stratotype of Jianshangou of
Yixian Formation. Sci. China Ser. D, Earth Sci. 34, 883—895
(in Chinese).
Ding Q.H., Zhang L.D., Guo S.Z., Zhang C.J., Peng Y.D., Jia B.,
Chen S.W. & Xian D.H. 2003: Paleoclimatic and palaeoenvi-
ronmental proxies of the Yixian Formation in the Beipiao area,
western Liaoning. Geol. Bull. China 22, 3, 186—191.
Gao T.P. & Ren D. 2008: Description of a new fossil Anthoxyela
species (Hymenoptera, Xyelidae) from Yixian Formation of
Northeast China. Zootaxa 1842, 56—62.
Geinitz F.E. 1880: Der Jura in Mecklenburg und seine Ver-
steinerungen. Z. Dtsch. Geol. Gesell. 22, 510—535.
Handlirsch A. 1906: Die Fossilen Insekten und die Phylogenie der
rezenten Formen: Ein Handbuch für Paläontologen und Zoolo-
gen. Engelmann, Leipzig, 1—640, 36 plates.
Liang J.H., Vršanský P. & Ren D. 2012: Variability and symmetry of
a Jurassic nocturnal predatory cockroach (Blattida: Raphidio-
mimidae). Rev. Mexicana Ciencias Geol. 29, 2, 411—421.
Liang J.H., Vršanský P., Ren D. & Shih C.K. 2009: A new Jurassic
carnivorous cockroach (Insecta, Blattaria, Raphidiomimidae)
from the Inner Mongolia in China. Zootaxa 1974, 17—30.
Liu M., Lu W.H. & Ren D. 2007a: A new fossil mordellid (Coleoptera:
Tenebrionoidea: Mordellidae) from the Yixian Formation of
Western Liaoning Province, China. Zootaxa 1415, 49—56.
Liu Y.S., Sinitshenkova N.D. & Ren D. 2007b: A new genus and
species of stonefly (Insecta: Plecoptera) from the Yixian For-
mation, Liaoning Province, China. Cretaceous Research 28,
322—326.
Martínez-Delclòs X. 1993: Blátidos (Insecta, Blattodea) del Cretá-
cico Inferior de Espan
~a. Familias Mesoblattinidae, Blattulidae
y Poliphagidae. Bol. Geol. Minero 104, 5, 52—74.
Meunier F. 1914: Un blátido y una larva de odonato del Kimerid-
gense de la Sierra del Montsech (Lérida). Mem. Real Acad.
Ciencias y Artes de Barcelona 11, 9, 121—124.
Nagamitsu T. & Inoue T. 1997: Cockroach pollination and breeding
system of Uvaria elmeri (Annonaceae) in a lowland mixed-
dipterocarp forest in Sarawak. Amer. J. Botany 84, 208—213.
Pang Q.Q., Li P.X., Tian S.G. & Liu Y.Q. 2002: Discovery of ostra-
cods in the Dabeigou and Dadianzi Formations at Zhangjiagou,
Luanping County, northern Hebei Province of China and new
progress in the biostratigraphic boundary study. Geol. Bull.
China 21, 6, 329—336.
Picker M., Colville J.F. & Burrows M. 2012: A cockroach that
jumps. Biol. Lett. 8, 3, 390—392.
Ren D., Shih C.K., Gao T.P., Yao Y.Z. & Zhao Y.Y. 2010: Silent
stories – insect fossil treasures from dinosaur era of the
Northeastern China. Sci. Press, Beijing, 1—17.
Ren D., Guo Z.G., Lu L.W., Ji S.A., Tang F., Jin Y.G., Fang X.S. &
Ji Q. 1997: A further contribution to the knowledge of the Up-
per Jurassic Yixian Formation in western Liaoning. Geol. Rev.
43, 5, 449—460 (in Chinese with English abstract).
Roth L.M. 1991: New combinations, synonymies, redescriptions
and new species of cockroaches, mostly Indo-Australian Blat-
tellidae. Invertebrates Taxon 5, 953—1021.
Shelford R. 1909: Note on some amphibious cockroaches. Record
Indian Mus. 3, 125—127.
Tan J.J., Ren D. & Shih C.K. 2007: New ommatids of Ommatinae
(Coleoptera: Archostemata: Ommatidae) from the Yixian For-
mation of western Liaoning, China. Progress Nat. Sci. 17, 7,
803—811.
Vidlička L., Vršanský P. & Shcherbakov D.E. 2003: Two new tro-
globitic cockroach species of the genus Speleoblatta (Blattaria:
Nocticollidae) from North Thailand. J. Nat. Hist. 37, 107—114.
Vishniakova V.N. 1964: Additional characters of wing venation in
forewings of a new Upper Jurassic cockroach. Paleontol. Zh.
1, 82—87 (in Russian).
Vishniakova V.N. 1973: New cockroaches (Insecta: Blattodea)
from the Upper Jurassic sediments of Karatau. In: Narchuk
E.P. (Ed.): Problems of the insect palaeontology (in Russian):
Lectures on the XXIV Annual Readings in Memory of N.A.
Kholodkovsky (1—2 April, 1971) [Voprosy paleontologii
nasekomykh. Doklady na 24-m Ezhegodnom chtenii pamyati
N.A.Kholodkovskogo, 1971.] Nauka, Leningrad, 64—77 (in
Russian).
Vršanský P. 1997: Piniblattella gen. nov. – the most ancient genus
of the family Blattellidae (Blattodae) from the Lower Creta-
ceous of Siberia. Entomol. Probl. 28, 1, 67—79.
Vršanský P. 2000: Decreasing variability – from the Carbonifer-
ous to the present! (Validated on independent lineages of Blat-
taria). Paleont. J. 34, 3, 374—379.
Vršanský P. 2002: Origin and the early evolution of mantises. AMBA
Projekty 6, 1, 1—16.
Vršanský P. 2003a: Umenocoleoidea – an amazing lineage of aber-
rant insects (Insecta, Blattaria). AMBA Projekty 7, 1—32.
Vršanský P. 2003b: Unique assemblage of Dictyoptera (Insecta-
Blattaria, Mantodea, Isoptera, Mantodea) from the Lower Cre-
taceous of Bon Tsagaan Nuur in Mongolia. Entomol. Probl.
33, 1—2, 119—151.
Vršanský P. 2004a: Transitional Jurassic/Cretaceous cockroach as-
semblage (Insecta, Blattaria) from the Shar-Teg in Mongolia.
Geol. Carpathica 55, 6, 457—468.
Vršanský P. 2004b: Cretaceous Gondwanian Cockroaches (Insecta:
Blattaria). Entomol. Probl. 34, 49—54.
Vršanský P. 2007: Jumping cockroaches (Blattaria, Skokidae fam.n.)
304
WEI and REN
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA
GEOLOGICA CARPATHICA, 2013, 64, 4, 291—304
from the late Jurassic of Karatau in Kazakhstan. Biologia, Bra-
tislava 62, 5, 588—592.
Vršanský P. 2008: New blattarians and a review of dictyopteran as-
semblages from the Lower Cretaceous of Mongolia. Acta
Palaeont. Pol. 53, 1, 129—136.
Vršanský P. 2009: Albian cockroaches (Insecta, Blattida) from
French amber of Archingeay. Geodiversitas 31, 1, 73—98.
Vršanský P. 2010: Cockroach as the earliest eusocial animal. Acta
Geol. Sin. Engl. Ed. 84, 793—808.
Vršanský P. & Ansorge J. 2007: Lower Jurassic cockroaches (In-
secta: Blattaria) from Germany and England. African Inverte-
brates 48, 103—126.
Vršanský P., Cifuentes-Ruiz P., Vidlička L., Čiampor F.J.R. & Vega
F.J. 2011: Afro-Asian cockroach from Chiapas amber and the
lost Tertiary American entomofauna. Geol. Carpathica 62, 5,
463—475.
Vršanský P., Chorvát D., Fritzsche I., Hain M. & Ševčík R. 2012:
Light-mimicking cockroaches indicate Tertiary origin of recent
terrestrial luminescence. Naturwissenschaften 99, 9, 739—749.
Wang C.D. & Ren D. 2013: Nuurcala obesa sp. n. (Blattida, Calob-
lattinidae) from the Lower Cretaceous Yixian Formation in
Liaoning Province, China. ZooKeys 318, 35—46.
Wang T.T., Liang J.H. & Ren D. 2007a: Variability of Habroblattula
drepanoides gen. et sp. nov. (Insecta: Blattaria: Blattulidae)
from the Yixian Formation in Liaoning, China. Zootaxa 1443,
17—27.
Wang T.T., Ren D., Liang J.H. & Shih C.K. 2007b: New Mesozoic
cockroaches (Blattaria: Blattulidae) from Jehol biota of western
Liaoning in China. Ann. Zoologici 57, 3, 483—495.
Wang W.L., Zhang L.J., Zheng S.L., Ren D., Zheng Y.J., Ding
Q.H., Zhang H., Li Z.T. & Yang F.L. 2005: The age of the
Yixian stage and the boundary of Jurassic-Cretaceous – the
establishment and study of stratotypes of the Yixian stages.
Geol. Rev. 51, 3, 234—242.
Wang W.L., Zhang L.J., Zheng S.L, Zheng Y.J., Zhang H., Li Z.T.
& Yang F.L. 2004: A new study on the stratotype and bio-
stratigraphy of the Yixian stage in Yixian-Beipiao region,
Liaoning – establishment and study of stratotypes of the Yixian
stage. Acta Geol. Sin. 78, 4, 433—447.
Zhang Z.J., Schneider J.W. & Hong Y.Q. 2012: The most ancient roach
(Blattida): A new genus and species from the earliest Late Car-
boniferous (Namurian) of China, with discussion on the phylo-
morphogeny of early blattids. J. Systematic Palaeont. 11, 27—40.
Zhou Z.H., Barrett P.M. & Hilton J. 2003: An exceptionally pre-
served lower Cretaceous ecosystem. Nature 421, 807—814.
Zompro O. & Fritzsche I. 1999: Lucihormetica n. gen. n. sp., the first
record of luminescence in an orthopteroid insect (Dictyoptera:
Blaberidae: Blaberinae: Brachycolini). Amazoniana 15, 211—219.