GeolCarp_Vol60_No6_449

www.geologicacarpathica.sk

GEOLOGICA CARPATHICA, DECEMBER 2009, 60, 6, 449—462 doi: 10.2478/v10096-009-0033-0

Advanced morphology and behaviour of extinct earwig-like

cockroaches (Blattida: Fuziidae fam. nov.)

PETER VRŠANSKÝ

1,2,3

, JUN-HUI LIANG

1,4

and DONG REN

College of Life Science, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing, 100048, P.R.China;

rendong@mail.cnu.edu.cn

Arthropoda Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117868 Moscow, Russia;

lab@palaeoentomolog.ru

Geological Institute, Slovak Academy of Sciences, Dúbravská cesta 9, P.O. BOX 106, 840 05 Bratislava 45, Slovak Republic;

geolvrsa@savba.sk

Tianjin Museum of Natural History, 206 Mangchang Road, Hexi District, Tianjin, 300074, P.R.China;

liangjh0602@126.com

(Manuscript received May 29, 2009; accepted in revised form October 2, 2009)

Abstract: We describe the extinct cockroach family Fuziidae fam. nov., represented by Fuzia dadao gen. et sp. nov.
from the ?Bathonian (168 Ma) Middle Jurassic sediments of Daohugou, Inner Mongolia, China. Males are character-
ized by unique, long and narrow bodies with a notch on forceps of earwig-like cerci, which attaches to the long external
ovipositor during courtship. In a combination with the presence of male tergal glands, it appears the most advanced
form of reproduction in the nearly 300 Myr history of long external ovipositor-bearing cockroaches. Its advanced mor-
phology significantly supports attribution of living and fossil cockroaches within a single order Blattida.

Key words: Middle Jurassic, Bathonian, China, Inner Mongolia, Daohugou, Insecta, Blattida ( = Blattaria = Blattodea),
fossil cockroaches.

Introduction

Along  with  the  stunning  pale-
ontological  discoveries  from
the world-renowned fossil beds
in  western  Liaoning  Province
of  China,  recent  findings  from
the  lacustrine  deposits  exposed
at

Daohugou

village

near

Ningcheng  (Fig. 1),  Inner  Mon-
golia, include superbly preserved
insect, pterosaur, salamander and
plant  fossils  (Gao  &  Shubin
2003;  Rasnitsyn  &  Zhang  2004;
Gao & Ren 2006).

Cockroaches, with an evolu-

tionary  history  extending  over
320 Myr, and with over 100,000
fossil specimens collected so far,
form  the  most  complete  group
of fossil insects. They appear to
be  a  varied  and  diverse  group,
giving rise to social termites and
predatory mantises. This plastic-
ity includes enormous structural
variation  of  male  genitalia
among  the  major  clades  (see
Klass 1997) and can be contrast-
ed with the conservation of gen-
eral  body  structures  and  most

Fig. 1. Localization of Daohugou area at the juncture of three provinces of Liaoning, Hebei, and Inner
Mongolia (E 119°14.318’, N 41°18.979’, height 607 m). Legend and abbreviations: 1 – stratigraphic
boundary; 2 – unconformity; 3 – provincial boundary; 4 – location of geological section; 5 – strike
and dip; Chc – Changchougou Formation; Chch – Chuanlinggou Formation; Chd – Dahongyu For-
mation; Cht – Tuanshanzi Formation; Dms – Dalaiyingzi erosion surface; J2j – Jiulongshan For-
mation; J2t – Tiaojishan Formation; Ky – Yixian Formation; Q – Quaternary; Mgn – Maanshan
gneiss. Modified after Ren et al. (2002).

450

VRŠANSKÝ, LIANG and REN

characters during the phylogeny. At higher taxonomic levels,
this body plan conservation results in 15 distinct extinct
groups (corresponding to families all of which can be dis-
crimninated on the basis of both male and female terminalia).
This paper describes a new species of fossil cockroach Fuzia
dadao gen. et sp. nov., and attributes it to a separate family,
Fuziidae fam. nov.

This species is important in several respects: it represents

further  evidence  for  decreasing  variability  of  respective  spe-
cies over time (Vršanský 2000; Webster 2007) and also for oc-
currence  of  first  mass  animal  deformations  representing
mutations (Vršanský 2004, 2005) – phenomena recently ob-
served  in  the  paleobotanical  record  as  well  (Krassilov  2003;
Visscher et al. 2004; Foster & Afonin 2005).

The unique structures of this species are male copulatory

forceps, which are commonly present in many other insect ter-
minalia such as in Mantophasmatodea (Klass et al. 2003),
Dermaptera, Japygidae, Odonata, and among Embioptera and
Phasmatodea, but not previously reported in cockroaches. For
a similar, although not homological structure, see the volsella
of male hymenopteras and scorpion-fly.

Material and methods

Nineteen complete males and sixteen females are described

and were collected from the Middle Jurassic sediments of the
Jiulongshan Formation (Ren et al. 2002; Shen et al. 2003; Liu et
al.  2004;  Rasnitsyn  &  Zhang  2004;  Gao  &  Ren  2006),  at  the
Daohugou locality in Inner Mongolia, China (Fig. 1). The Dao-
hugou  fossil  beds  consist  of  a  set  of  intercalated,  fine-grained
lacustrine  deposits  and  fine  volcanic  ash  that  unconformably
overlay pre-Cambrian rocks (Ren et al. 2002; Liu & Jin 2002).

The accurate Ar-Ar and SHRIMP U-Pb dating shows that

the  age  of  intermediate-acid  volcanic  rocks  overlying  the
Daohugou fossil-bearing beds is about 164—165 Ma, and that
the  age  of  these  fossil-bearing  beds  is  older  than  or  equal  to
165 Ma  (Chen  et  al.  2004).  Therefore  the  age  of  Daohugou
biota is considered to be Middle Jurassic (Aalenian—Bathonian)
(Ren et al. 1995; Gao & Ren 2006).

We studied the material using a Leica MZ12.5 dissecting

microscope and illustrate them with attached drawing tube
and photographs were made using Leica DC 300 photographic
equipment.

The analysed specimens are listed in the table caption. The

coefficient of variation (in percent) was computated as Standard
deviation/Average.

The venial nomenclature follows the earliest studies of the

senior author (Vršanský 1997).

Institutional abbreviations: CNU–Capital Normal Universi-

ty, Beijing, China; TNP–Tianjin Museum of Natural History.

Systematic paleontology

Order: Blattida Latreille, 1810

Superfamily: Caloblattinoidea Vršanský & Ansorge in

Vršanský (2000)

Family: Fuziidae fam. nov.

T y p e g e n u s : Fuzia Vršanský, Liang

et Ren, gen. nov.

described below.

C o m p o s i t i o n : Type genus, some unidentified, but dif-

ferent genera from the Upper Triassic of the Madygen in Kir-
gizia, Middle Jurassic (?Bajocian) of Bakhar in Mongolia and
Upper Jurassic (?Kimmeridgian) sediments of the Karatau in
Kazakhstan.

R a n g e : ?Upper Triassic—Middle—?Upper Jurassic.
D i f f e r e n t i a l d i a g n o s i s : Fuziidae (Fig. 2b1) differ

from  all  Paleozoic  families  except  the  Phyloblattidae
Schneider, 1983 by the presence of intercalary branches (syna-
pomorphy)  [For  character  polarities  see  character  analysis
(list)  below],  and  from  the  Phyloblattidae  in  having  special-
ized  and  simplified  venation,  especially  simple  Sc  and  A
branches (apomorphies).

The Fuziidae differ from all the known Mesozoic families

(for comparison see below; for character polarities see charac-
ter analysis) in the following characters: very small head cov-
ered  by  a  very  large  pronotum  (head  less  than  a  half  of  the
pronotal  width);  forewing  widest  in  the  apical  third,  costal
area very long (a third of wing) and wide (more than a third of
wing).  Wings  except  for  hindwing  (and  possibly  forewing)
apex  are  without  colouration.  Forewing  Sc  strong,  simple  or
with terminal branch; RS slightly differentiated in most indi-
viduals, even when there is an apparent trend of uniformiza-
tion of R1 and RS in the present taxon (Fig. 7i – RS can be
tracked only on the basis of more richly branched apical R1),
distinctly expressed in all the Blattulidae.

Autapomorphies (see also character analysis) of the new

family are: eyes significantly projected beyond the head out-
line; coloured labial palps; wide pronotum; shape of forewing,
wide costal area, short clavus; curved ovipositor and forceps-
like male cerci. Other significant shared apomorphies (perhaps
synapomorphies) with the Blattulidae are simplified Sc and A,
and bonded R1 with RS.

The character of venation with numerous intercalaries and

cross-veins forming comb are synapomorphic with the Calob-
lattinidae (i.e. unnamed relic caloblattinid from the Upper Ju-
rassic  of  Shar-Teg  in  Mongolia  –  see  Vršanský  2004,
fig. 6.7—6.8  p. 464,  which  also  share  simplified  A  branches).
The entire hindwing, differing from the ancestral state of the
Caloblattinidae  by  having  a  reduced  M  and  CuA  and  CuP
branches  limited  to  a  simple  vein,  is  homoplasic  with  the
Liberiblattinidae.  Fuziidae  differ  from  the  Liberiblattinidae
having  eyes  which  protrude  beyond  the  outline  of  head  (ho-
moplasic with living cockroaches), the form of palps (palps of
the Fuziidae resemble extant rather than Mesozoic cockroach-
es (except the Blattulidae), all of which have long palps), with
apical segment partially cup-like; slightly curved forewing R,
simple Sc; and richly branched hindwing CuA. It additionally
differs from all other Polyphagoidea by having a narrow body
with characteristic terminalia (autapomorphy) and in expand-
ed venation, slender veins with very slender intercalaries and
in wide costal field.

Additional families differ in possessing strong autapomor-

phies: Umenocoleidae Chen et Tian, 1973 differ in possesing
hard elytra; Raphidiomimidae Vishniakova, 1973 are elongat-
ed with prognathous head; Latiblattidae Vishniakova, 1968
are extremely widened (all additionally with plesiomorphically

451

BLATTIDA: FUZIIDAE FAM. NOV. – ADVANCED MORPHOLOGY OF EXTINCT EARWIG-LIKE COCKROACHES

secondarily branched A and Sc) and Eadiidae Vršanský, 2009
have curved leg spurs and short pronotum.

Fuzia differs from the Mesozoic Blattoidea (Mesoblat-

tinidae Handlirsch, 1906 and Blattellidae Karny, 1908) in hav-
ing a short (25 % of wing length) and fluently curved CuP,
wide costal field (autapomorphies) and by the presence of an
external ovipositor with basal ridge serving for attachment of
cerci, and double hook (plesiomorphies). The male terminal
hook may be homological to the hook hla (Klass 1997) includ-
ing sclerite L3 (see McKittrick 1964), but more probably to the
terminal hooks of the right phallomere of some living Blattidae
(McKittrick 1964; fig. 108) or terminal processes paa + pda of
the left phallomere of some living Blattellidae (Klass 1997).

According to the basal position of the new family with re-

spect to Mesozoic cockroaches, there is no need of (phyloge-
netic)  comparison  Fuziidae  with  the  living  families  (which,
for example all lack the external ovipositor). However, some
superficial similarity is seen with the family Blattellidae (ho-
moplastic  simple  forewing  Sc  and  A;  similar  pleating)  and
also  Polyphagidae  (homoplastic  or  synapomorphic  sharp  but
fluently curved CuP).

The area between CuA and CuP is irregular with the width

of this area varying along the wing length, with free space for
one CuA branch present. The clavus is very short (25 % of the
wing  length),  with  fluently  curved  CuP  and  a  diagonal  kink
shifted posteriorly. Branches of A are simple, A1 occasionally
has  blind  branches.  Hindwing  with  facultative  terminally  di-
chotomized  veins,  differentiated  R1  and  RS,  without
pterostigma;  and  with  secondary  and  tertiary  branched  CuA.
Males with cerci forceps-like by notches on their distal mesal
edges. Female with short (1/7 of total body length), rigid and
curved external ovipositor with basal ridge on right side.

D e s c r i p t i o n : As for the species.
T a x o n o m i c p o s i t i o n : Character list is provided in or-

der to identify character states (polarities):

– Shape of head: eyes protruding beyond the outline of

head:  synapomorphic  with  the  Blattulidae,  Liberiblattinidae
and  living  cockroaches  (eyes  does  not  protrude  beyond  the
head  outline  in  the  stem  family  Caloblattinidae)  (Even  the
head is well preserved, ocelli are invisible, in spite of their an-
ticipated presence. All three ocelli of extinct cockroaches were
plain and visible mostly in amber material (Vršanský 2008a;
Anisyutkin & Gorochov 2008);

– Palps very short (shorter than length of the head): synapo-

morphic with the Blattulidae (very long in ancestral Caloblat-
tinidae, comparatively longer in the related Liberiblattinidae);

– Apical segment of palps cup-like: synapomorphic with

advanced Caloblattinidae and its descendants (cup-like termi-
nation absent in early Caloblattinidae);

– Large wide pronotum (as wide as the body): autapomor-

phic (such a wide pronotum is present in some Paleozoic
groups, but absent in the stem Caloblattinidae), homoplastic
with the unrelated Latiblattidae;

– Elongated body (4 times longer than wide): autapomor-

phic (elongated body is present in some living cockroaches,
but absent in Paleozoic and Mesozoic groups except ho-
moplastic in Raphidiomimidae and some Umenocoleidae);

– Tergal glands present: plesiomorphic (according to phy-

logenetic position of Fuzia, which is not directly related to the

Liberiblattinidae, perhaps inherited from certain Caloblat-
tinidae. Glands are plesiomorphicaly present in the Liberiblat-
tinidae – stem for all living cockroaches, Skokidae, termites
and mantises);

– Cerci with reduced number of segments (under 15): syn-

apomorphic with the Blattulidae (the stem Caloblattinidae
have numerous segments, Liberiblattinidae a little less numer-
ous than the Caloblattinidae);

– Cerci with forceps: autapomorphic;
– Styli very long (more than a half of cerci): symplesio-

morphic with the early Caloblattinidae (shorter in all advanced
Caloblattinidae and also in the Blattulidae);

– Curved external ovipositor with ridge: autapomorphy;
– Sledner carinated legs: autapomorphy (in the stem

group, the Caloblattinidae, the legs are strongly carinated; in
some derived Liberiblattinidae homoplasically partially re-
duced; in Skokidae (as a jumping adaptation) and a new fami-
ly (as a social adaptation) nearly reduced);

– Forewing shape: autapomorphic (the shape of wing with

wide costal area, and the wing widest in the apical third is ab-
sent in Paleozoic and Mesozoic cockroaches, although ho-
moplasically ocurrs in some advanced living groups);

– Forewing costal area wide: autapomorphy, homoplasi-

cally present in the Latiblattidae (Latiblattidae are not directly
related, thus this character cannot be synapomorphic);

– Forewing with more or less distinct intercalaries: apo-

morphic with early Phyloblattidae, Caloblattinidae, Liberiblat-
tinidae; present, but tend to reduce in the Blattulidae;

– Forewing Sc simple: synapomorphic with the Blattulidae

(stem Caloblattinidae but also Liberiblattidae and all Mesozo-
ic groups have Sc branched; Sc homoplasically reduce in the
living Blattellidae);

– Forewing RS indistinctly differentiated (see Figs. 7—8):

synapomorphic with the early Blattulidae (RS is not as distinct
as in other Mesozoic families except for the Blattulidae: this
partial reduction, completely expressed in the Blattulidae, is
very unusual and sophisticated, and thus unlikely to represent
a homoplasy);

– Forewing venation rich: symplesiomorphic with the

Caloblattinidae (rich venation is generally a plesiomorphic
state – venation tends to reduce in several independent
lineages);

– Forewing CuA expanded: symplesiomorphic with the ear-

ly Caloblattinidae (CuA is reduced even in the advanced Calob-
lattinidae and in all derived Mesozoic and living families);

– Forewing clavus short (less than a third of wing): auta-

pomorphy;

– Forewing branches of A simple: synapomorphic with the

Blattulidae (A are branched in the stem Caloblattinidae and all
Mesozoic groups; A is homoplasically simplified in the living
Blattellidae);

– Hindwing terminal coloration: symplesiomorphic with

the advanced Caloblattinidae (present in some advanced Calo-
blattinidae, Liberiblattinidae, Blattulidae and Skokidae);

– Hindwing R1 and RS with standard (not comb-like

branches) dichotomisations: symplesiomorphic with the Phy-
loblattidae (present also in the stem Caloblattinidae);

– Hindwing M simplified: synapomorphic with the Blattul-

idae (M is rich in the stem family Caloblattinidae, in the

452

VRŠANSKÝ, LIANG and REN

Liberiblattinidae and Skokidae – thus homoplasy with the
Blattulidae is unlikely; M is homoplasically simplified in the
Blattellidae);

– Hindwing CuA multiply branched: symplesiomorphic

with the Caloblattinidae.

Genus Fuzia gen. nov.

T y p e s p e c i e s : Fuzia dadao sp. nov; see below.
D e s c r i p t i o n : As for the species.
C o m p o s i t i o n : Type species only.
D e r i v a t i o n o f t h e n a m e : Fuzi is Chinese for master.

Fuzia dadao sp. nov.

Figs. 2—8, Table 1

H o l o t y p e : CNU-B-NN-2006-666. A complete male.

(Figs. 2b, 4A1—3).

T y p e l o c a l i t y : Daohugou, Inner Mongolia, China.
T y p e h o r i z o n : Jiulongshan Formation (designated by

Ren et al. 2002). Middle Jurassic (?Bathonian).

P a r a t y p e : CNU-B-NN-2006-301. A complete female.

(Figs. 3A, 6B1—2)

A d d i t i o n a l m a t e r i a l : CNU-B-NN-2006-031, 035=042,

038,  044,  322,  357,  669,  670,  671,  672,  673,  674,  675,  676,
677,  681,  682;  TNP-42982  (complete  males:  Figs. 5,  7).
CUN-B-NN-2006,  305,  306,  314F,  328,  341,  348,  371,  380,
381,  383,  1001,  1002,  1003,  1004,  1005  (complete  females:
Figs. 6, 8). The same locality as the type.

D e r i v a t i o n o f t h e n a m e : After dadao (Chinese for

“perspicacious understanding of the path”; and also for sharp
knife) – alluded also to the type locality, the Daohugou.

D e s c r i p t i o n : Head small (ca. 1/8 of the total body length;

pronotum width to head width ratio over 2:1), with palps shorter
than head (Fig. 2a). Male head is almost globular (Fig. 2a),
and slightly longer than wide (length/width 1.2—2/1.2—2 mm),

Fig. 2. Males of caloblattinoid Fuzia dadao sp. nov. a – Head (CNU-B-NN-2006-677); b1-2 – Holotype CNU-B-NN-2006-666. A com-
plete male; c – Dorsal view with tergal glands (CNU-B-NN-2006-357); d – Detail of cercus with numbered cercomeres and indicated
notch (CNU-B-NN-2006-357). Notice the more sophisticated carving on the left cercus (arrow). Daohugou, Inner Mongolia, China. Middle
Jurassic (?Bathonian). Abbreviations: Sc – subcosta, R – radius, R1 – radius anterior, RS – radial sector, M – media, CuA – cubitus
anterior, CuP – cubitus posterior, A – anal veins. T – tergites, G – glands. Right forewing length 13.7 mm.

453

BLATTIDA: FUZIIDAE FAM. NOV. – ADVANCED MORPHOLOGY OF EXTINCT EARWIG-LIKE COCKROACHES

in females the head is significantly elongated (3/2.2 mm). The
pronotum  is  very  large,  especially  in  females  (2.5—3.1/4.2—
4.5 mm) (2.2—2.6/3.2—3.8 mm in males), with very wide para-
notalia (Fig. 2b1) and two coloured central stripes. Forewing
(Fig. 2b1)  widest  in  apical  third,  with  very  wide  costal  area
(nearly a third of the wing’s width), with intercalaries and nu-
merous  cross-veins  forming  comb-like  structures.  Length/
width:  10—15.5/3.6—6.5 mm  (in  females  12.5—13/4.2—4.5 mm),
with simple and very long (a third of a wing), strong partially
coloured  Sc  (occasionally  with  1  or  two  terminal  dichoto-
mies). The RS is differentiated in most individuals. R (15—25)
is  almost  straight,  the  branches  are  simple  or  secondary
branched,  apical  R1  vein  (predecessing  RS)  has  multiple
branches. M 3—10; CuA (4—13) expanded, and usually reach-
es the apical part of wing, this is expressed as narrowing to-
wards  apex.  Clavus  is  very  short  (less  than  a  third  of  the
wing’s length), with simple branches of A running parallel to
the anterior margin and to each other (A1 occasionally with

blind branches). A 4—7. The hindwing (Fig. 2b1) has faculta-
tive terminal branchelets, a simple Sc; R1 (3—4) and the RS
(9—13) is differentiated, without pterostigma, but with a
darkened R1. The M almost straight, but reduced to a few
veins (2—6); CuA (6—11) are secondarily or tertiary branched,
not reduced, and also with additional blind branches; and the
CuP simple.

The (Fig. 2c) male body is significantly longer than the

forewings, but body is very narrow (about as wide as prono-
tum) even in females, in contrast to all other cockroaches with
external ovipositor, which have a wide body. Male body sig-
nificantly elongate, very narrow, with cerci forming forceps
(with notches). Tergal glands are present in at least three seg-
ments. The external ovipositor is very rigid (preserved 3D
unlike rest of the fossils) curved, and of the long type.

All legs (Fig. 2b1) very short and comparatively slender and

with slender carination. Fore femur length/width similar in
males and females (2—2.22/0.67 mm – holotype and paratype

Fig. 3. Females of caloblattinoid Fuzia dadao sp. nov. a – Paratype (CNU-B-NN-2006-301); b – Dorsal view on the ovipositor and num-
bered cercomeres (CNU-B-NN-2006-383). Daohugou, Inner Mongolia, China. Middle Jurassic (?Bathonian). Abbreviations: V I. – Ventral
valves I (gonapophyses 8 or 1

valves); V II. – Dorsal valves (gonapophyses 9 or 2

valves); V III. – Outer valves III (gonoplacks or 3

valves); lam – Ovipositor plates 1 and/or 2 (all homologized according to Vishniakova 1968). Alternatively (see Klass 1998), the median parts
(V I and V III) represent the 1

valves (ga of Klass 1998), left and right, but not down to their bases. The lateral parts (V II and lam) represent

the 2

(gp of Klass 1998) and 3

valves (gl of Klass 1998), together with coxite 9 (the anterolateral parts; lateral parts of aa of Klass 1998;

= “lam 2” of Vishniakova 1968). In this part the origin of 2

valves should be anteromesally (lam) and that of 3

valves posterolaterally

(V II), but the proportions do not really fit with this interpretation. Right forewing length 12.5 mm.

456

VRŠANSKÝ, LIANG and REN

spective  orders  has  caused  broad  discussion.  Vishniakova
(1968)  studied  and  described  Mesozoic  ovipositors  in  detail
and  homologized  the  outer  ovipositor  valves  with  the  inner
vales of living cockroaches, thus allowing the classification of
extant and extinct cockroaches within a single order (Hennig
1981,  with  reservations).  The  most  significant  difference  re-
mained  the  presence  of  the  ootheca  (egg-case)  in  all  living
forms, except for some derived species (Roth & Willis 1958).
Nevertheless, an unnamed living relic cockroach bears exter-
nal  ovipositor  ( .  Vidlička,  P.  Vršanský  in  preparation)  and
apparently  Melyroidea,  in  spite  of  its  alternative  categoriza-
tion as a terminal blattellid taxon also has a similar terminalia
structure (Vršanský 2003).

Grimaldi (1997) divided the monophyletic Dictyoptera into

living cockroaches without external ovipositor (Blattaria), ter-
mites  and  mantises;  and  extinct  Blattodea  –  cockroaches
with  external  ovipositor.  However,  the  Liberiblattinidae  –
stem  group  of  the  living  Dictyoptera,  Skokidae,  Mesoblat-
tinidae  (precursors  of  the  Blattellidae,  Blaberidae  and  Blatti-
dae),  and  also  some  living  cockroaches  possess  rudimentary
external ovipositor (Vršanský 2002). Thus the loss of the ex-
ternal ovipositor is an evolutionary trend and the Blattaria sen-
su  Grimaldi  would  appear  polyphyletic.  Inward  et  al.  (2007)
simply place termites within the order of cockroaches, but this
opinion  is  not  supported  (see  also  Lo  et  al.  2007)  as  living
cockroaches and termites (as well as mantises) evolved from
Mesozoic cockroaches independently (Vršanský 2008b).

Thus, the taxon Blattida ( = Blattaria) is of paraphyletic

(non-cladistic) nature (see also Vršanský et al. 2002; Lo et al.
2007). The hypothesis of Béthoux & Wieland (2009) suggest-
ing that the holophyly of mantises started from the non-blat-
tarian  Carboniferous  ancestor  cannot  be  followed  for  the
reason  of  the  well  traced  transition  of  the  Mesozoic  cock-
roaches  Liberiblattinidae  into  the  Baissomantidae  mantises
(Vršanský 2002).

Among fossil cockroaches, three families (Latiblattidae,

Eadiidae; a new family) appear endemic and limited to a few
specimens from the Upper Jurassic Lagerstätte of Karatau and
from the Mesozoic Archingeay amber, suggesting that indige-
nous families were not uncommon.

Complete, well preserved specimens of the new family

Fuziidae, are only abundant at the present locality. However,
some  poorly  preserved  wings  from  the  slightly  older  sedi-
ments  from  the  Bakhar  in  Mongolia,  some  complete  speci-
mens  from  the  ?Kimmeridgian  Upper  Jurassic  of  Karatau  in
Kazakhstan, and also some specimens from the Upper Triassic
locality Madygen in Kirgizia, show identical venation pattern
and elongated body. Thus, the family had perhaps originated
during the Triassic, and persisted until the Upper Jurassic.

Evidence for behaviour in extinct insects from amber com-

prise a wide variety of patterns including mating, egg-laying,
progeny care, food carriage, parasitism, mutualism and phore-
sy  (Arillo  2007),  and  distinct  morphology  such  as  preserva-
tion of metapleural glands in primitive ants (Engel & Grimaldi
2005) indirectly indicates a competent behaviour. In the sedi-
mentary  record,  evidence  for  distinct  behaviour  is  also  com-
mon,  such  as  sound  apparatus  preserved  in  grasshoppers
(Sharov  1968;  Martins-Neto  1999),  evidence  for  parasitism
(Brauckmann et al. 2007), pollination (Krassilov & Rasnitsyn

1982; Rasnitsyn & Krassilov 1996a,b; Labandeira 2000), feed-
ing on plants (see Labandeira 2002) and insects (Durden 1988),
and  oviposition  (Béthoux  et  al.  2004;  Labandeira  2006).  The
presence of male tergal glands which are used for pre-copula-
tion  attraction  of  females  in  the  extant  cockroach  species  is
unique,  even  the  glands,  indicated  as  plesiomorphic  for  living
taxa,  must  have  been  present  in  most  Mesozoic  cockroaches
(but also in termites and mantises – see below).

This character, supported here to be plesiomorphic for all

Mesozoic-originating cockroaches is thus not an autapomor-
phy of living lineages of cockroaches and contributes to the
attribution of Mesozoic and living families within a single or-
der, the Blattida (typified ordinal name of Blattaria = Blattodea,
paraphyletic in respect to Isoptera and Mantida), supported ad-
ditionally by modern morphology as shown here.

All Mesozoic (except Subioblattidae, Poroblattinidae,

Spiloblattinidae  originating  in  the  Paleozoic),  Cenozoic  and
living  cockroaches  (together  with  mantises  and  termites)  are
monophyletic,  originating  from  the  Phyloblattidae-Caloblat-
tindae  lineage  –  see  Vršanský  (2002)  and  Vršanský  et  al.
(2002).  The  present  taxon,  the  Fuziidae,  may  be  categorized
within Blattida (all Dictyoptera other than termites and man-
tises,  that  is,  the  Blattaria)  on  the  basis  of  general  habitus
(Fig. 2b1), of a large pronotum with distinct paranotalia, mul-
tisegmented  cerci  (Fig. 2c)  with  numerous  sensilla  chaetica,
and carinated legs. Synapomorphies with the Blattida are a hy-
pognathous head directed backwards (Fig. 2a), and character-
istic venation pattern of forewings (with pectinated regularly
branched veins, and with distinct clavus) and hindwings (sim-
ple Sc; R1 somewhat comb-like, RS differentiated, simplified
M and expanded CuA, simple CuP).

We now briefly designate ovipositor types of cockroaches.

The external ovipositor of Fuzia is assigned to the “Mesozoic
long  type”  (found  in  the  Caloblattinidae,  Umenocoleidae,
Latiblattidae) based on the general robust shape with more or
less parallel linear dorsal and ventral outer valves (V1 and V2)
and the sharp apex, and with distinct external ovipositor plates
(Lam 1, 2 according to Vishniakova 1968) and V3. This type
can be contrasted with the two alternative types of ovipositor.
The slender “short-type” external ovipositor of the Blattulidae
characterized  by  a  dominant  U-shaped,  sharply  cut  at  apex
parallel outer ventral and dorsal (V1, 2) valves, with internal-
ized rudimentary valves V3. The “rudimentary-type” external
ovipositor  of  the  Skokidae,  Raphidiomimidae  and  Mesoblat-
tinidae, is characterized by short, triangular dorsal and ventral
valves, with merged slender short ovipositor plates (Lam 1 ac-
cording  to  Vishniakova  1968),  terminated  by  small,  but  dis-
tinct and hardly sclerotised V3.

The fore wings are widest in the apical third. This is fre-

quently found in several extant Blattida families and should
not normally be used to diagnose on family level. Neverthe-
less this character state was absent in all the Mesozoic cock-
roaches, and thus at that stage of the evolution of cockroaches
was a significant character transformation.

The RS is indistinct and reduced also in the Blattulidae,

but the presence of the long external ovipositor of caloblat-
tinid type, the hindwing lacking pterostigma, and branched
CuA (strong plesiomorphies) exclude direct ancestry of the
Blattulidae.

458

VRŠANSKÝ, LIANG and REN

The hindwing (Fig. 2b1) is similar to the Liberiblattinidae

Vršanský (2002), but the ancestry of Liberiblattinidae in re-
spect to Fuziidae might be excluded on the basis of the apo-
morphically curved R in the forewing and the advanced
external ovipositor of the “rudimentary-type”.

A globular head with short, as long as the head, maxillary

palps, a fluently curved CuP, and simple A branches running
parallel  to  the  anterior  margin  are  autapomorphic,  whilst  R
reaching the apex is plesiomorphic. This combination of fea-
tures  prevents  Fuzia  being  classified  within  the  known
Caloblattinidae.  Instead,  the  Fuziidae + Blattulidae  lineage
probably diverged from the ancestors of Caloblattinidae or an
earlier, yet undescribed representative of that family.

Enlargement of the pronotum combined with the large

forewing costal area, especially in females, is a protective ad-
aptation indicating a cryptic way of life. A coefficient of varia-

tion (CV) of 6.3 (for the total number of veins meeting mar-
gin) characterizes advanced modern taxa rather than Mesozoic
cockroaches, which typically have a CV of 10—13. This CV
might be caused by the conservative pattern of A, which var-
ies in males, and more seriously by conservative female size.

Forewings are not symmetrical, but radial veins had a very

low  CV  (14.44),  which  support  comparatively  good  flight.
The high total CV 8.51 for the total number of veins of both
sexes indicates that the species, may have been the most spe-
cialized  taxon  of  its  age.  Deformities  (vein  fusions)  in  the
wing  appear  relatively  common  and  distinct  with  seven  vein
fusions in 54 wings of 36 specimens suggesting that the spe-
cies was undergoing rapid evolution.

The lack of coloration in both sexes may support the cryptic

way of life (Jarzembowski 1994) of this large species, but it
may indicate at least locally open environments (see below).

Fig. 7. Explanatory drawings of males of caloblattinoid Fuzia dadao sp. nov. a – CNU-B-NN-2006-677 (R)(right forewing length 14 mm);
b – CNU-B-NN-2006-357 (right forewing length 10.6 mm); c – TNP42982; d – CNU-B-NN-2006-670; e – CNU-B-NN-2006-676
(R)(right forewing length 14.5 mm); f – CNU-B-NN-2006-672 (right forewing length 10 mm); g – CNU-B-NN-2006-322 (right forewing
length 10 mm); h – CNU-B-NN-2006-669 (right forewing length 13 mm); i – CNU-B-NN-2006-671 (R)(right forewing length 15.5 mm);
j – CNU-B-NN-2006-042. Daohugou, Inner Mongolia, China. Middle Jurassic (?Bathonian). R – reverse to actual position. Short line indi-
cates position of R1 and RS. Scales 1 mm.

459

BLATTIDA: FUZIIDAE FAM. NOV. – ADVANCED MORPHOLOGY OF EXTINCT EARWIG-LIKE COCKROACHES

The relative rarity of the species (ca. 3 % of all cockroaches)
suggests that these “open-patch“ environments were rare. It is
notable that unlike in extant cockroaches (Vidlička 2001), the
large Middle Jurassic cockroaches did not make themselves
conspicuous.

The only coloured part of the present species is the hind-

wing apex (and possibly the forewing apex), which serves as
terminalia-protection, as in fossil cockroaches apex hindwings
are not covered by forewings. The presence of apex coloration
also excludes unpreserved coloration of other body parts.

The non-segmented styli reaching the length of cerci is not

recorded in any extinct cockroach families. The extruded pres-

ervation of male copulatory organs, including the hook, is
unique and indicates that the terminalia were significantly pro-
truded from the body at least during copulation.

Paleoethology and paleoecology of

Fuzia dadao

sp. nov.

Our samples allow some data on the paleocological prefer-

ences of Fuzia dadao sp. nov. to be inferred. The most distinct
structure of the present species is the forceps of males, and the
ovipositor of the females. The forceps are formed by asymmet-

Fig. 8.  Explanatory  drawings  of  females  of  caloblattinoid  of  Fuzia  dadao  sp.  nov.  a  –  CNU-B-NN-2006-306  (R)(right  forewing  length
13 mm);  b  –  CNU-B-NN-2006-348  (R)(right  forewing  length  12 mm);  c  –  CNU-B-NN-2006-314  (right  forewing  length  12 mm);  D  –
CNU-B-NN-2006-328 (right forewing length 13 mm); e – CNU-B-NN-2006-371 (right forewing length 12 mm); f – CNU-B-NN-2006-341
(right forewing length 12.5 mm); g – CNU-B-NN-2006-383 (right forewing length 13 mm); h – CNU-B-NN-2006-380 (R)(right forewing
length  12.5 mm);  i  –  CNU-B-NN-2006-1001(R)(right  forewing  width  4.2 mm);  j  –  CNU-B-NN-2006-1005  (R)(right  forewing  length
12.5 mm);  k  –  CNU-B-NN-2006-381  (R);  l  –  CNU-B-NN-2006-1003  (R)(right  forewing  length  13 mm);  m  –  CNU-B-NN-2006-1002
(R)(right forewing length 12 mm); n – CNU-B-NN-2006-1002 (R); o – CNU-B-NN-2006-305 (right forewing length 13 mm). Daohugou,
Inner Mongolia, China. Middle Jurassic (?Bathonian). R – reverse to actual position. Short line indicates position of R1 and RS. Scales 1 mm.

460

VRŠANSKÝ, LIANG and REN

rical cerci, with narrow notches in strong, fused and widened
cercal segments 8—13. The female ovipositor is comparatively
short, extremely rigid (in contrast to other, also comparatively
hard body structures, was preserved in 3D) and curved. The
male notches fit to the ridge formed by the base and the strong
tubercle of narrowed lateral margins of the female ovipositor
outer valves – suggesting that the forceps served primarily
for attaching to females during courtship and copulation.

Such a strong adaptation suggests fusion during copulation

and, perhaps protection against extra mate copulation, howev-
er, such a protection is extraordinarily rare among extant in-
sects. The presence of a long external ovipositor, suggest that
the only possible position during courtship was with heads op-
posed, as observed in living cockroaches lacking external ovi-
positor (face-to-face position appear unrealistic because of the
eventual reverse angle of male terminalia with respect to the
body in such a pose).

The presence of male tergal glands also indicates an elabo-

rated  reproductive  behaviour.  Such  glands  are  used  for  pre-
copulation  attraction  of  females  in  the  living  cockroach
species.  The  female  is  posed  over  males  first,  attracted  by
male  pheromones  and  feeding  on  its  secretions  (male  with
outstretched wings – Vidlička 2001). Prior to this study, the
tergal  gland  was  not  recorded  in  any  of  thousands  of  fossil
specimens  of  external  ovipositor-bearing  cockroaches,  and
thus it was thought the secret-feeding phase of courtship was
absent. Nevertheless, the presence of these glands in such an
old lineage is a direct indication of the plesiomorphy of glands
and also this kind of behaviour.

The position of glands and their development vary among

living cockroaches, and have different position (T2—T6) and

number (5 in Fuzia). According to the frequent action of ho-
meotic genes it can be concluded the origin of the tergal
glands is single, and this character plesiomorphic at the level
of Mesozoic-originating Dictyoptera.

This hypothesis is additionally supported by a recent dis-

covery of similar sexual pheromones in mantises (Hurd et al.
2004) and male accessory glands are found also in the lower
termites (Weesner 1969).

Some additional inferences can be drawn from the available

fossil record. The morphology of female ovipositor suggests
that it served for oviposition into hard substrate, such as roots,
straws or, less likely – because of the lack of wing coloration
– into decaying wood (wood is rare in open environments).
Hard wood as an oviposition substacted can be excluded be-
cause newborn cockroaches would be unable to emerge from
the wood.

Transfer of nutritional packets from the female may have

played a more prominent role in the ancestors of current cock-
roaches (R. Brossut personal communication to Nalepa 1994).

The longest observed fusion of cockroaches during copu-

lation lasted 180 minutes (in the German cockroach – but
usually 72—115 minutes in this species).

Long-lasting attachment of sexes during courtship is a com-

mon adaptation of insects, as with the Southern green stink
bug Nezara viridula (Linnaeus 1758) may be attached for up
to 80 days (P. Štys, personal communication 2008).

An alternative to grasping directly on the ovipositor would

be merely to use it for grasping onto the abdomen itself, as of-
ten occurs in tettigoniids where males grasp the tegumentary
foldings at the abdomen tip with their specialized cerci. Nev-
ertheless,  the  specialized  ridge,  absent  in  all  other  external
ovipositors strongly suggests attachement directly to the ovi-
positor. The tubercle defining the ridge is quite apart from the
base  excluding  its  ovipositor-strengthening  function.  A  con-
siderable gap left between the male and female genital orifices
would  make  copulation  problematic,  but  male  terminalia  are
preserved quite far from abdomen, suggesting they were capa-
ble of protrusion.

In addition, the strongly sclerotized forceps at the end of an

elongated plastic body may have served as a protective device
against common predatory cockroaches and/or other carnivores,
as in some living beetles such as the Staphilinidae mimicking
wasp behaviour (their abdomen apex bears no threat to preda-
tors) and, to a lesser extent it may have been used in male com-
bat. Notwithstanding, most living insects protect themselves by
body movements, or through the mimicking of stinging insects,
which did not became abundant until the Cretaceous (Rasnitsyn
2002) – therefore the origin of this form of protection in the
present species remains obscure, also contradicted by the fact
that females lack them. Anyway, once evolved, a structure may
be secondarily used in this derived purpose.

The preferred habitat of Fuzia dadao sp. nov. is probably a

more open shrub and/or woody habitat rather than forest land-
scape,  because  large  species,  such  as  those  studied  from  the
canopy forests in Laos, Borneo and Equador, are rarely unco-
loured  in  rainforests  ( .  Vidlička,  P.  Vršanský,  in  prepara-
tion).  Open  habitat  is  also  supported  by  the  improved  flight
abilities of the both sexes – the roughly similar ratio of both
sexes  and  low  CV  of  female  forewing  venation  indicate  fe-

length width Sc

CuA CuP A

total

males/n 12

15 15

15 14

max

15.5

6.5

1 25

1 7

min

3.6

1 15

1 4

dev

2.07 0.77 0 2.58 1.99 2.16

0 1.06 4.46

ave

12.15 4.33 1 18.4 6.46 8.93

1 5.85 40.93

17.07 17.73 0 13.75 30.33 24.24

0 18.25 10.90

females/n 12

21 20

21 19

max

4.5

1 23

1 7

min

12.5

4.2

1 11

1 6

dev

0.45 0.15 0 2.50 2.32 1.63

0 0.37 2.55

ave

12.54 4.38 1 17.65 6.85 7.43

1 6.16 40.44

3.59 3.47 0 14.15 33.92 21.94

0 6.08 6.30

all/n

36 34

36 31

max

15.5

3.6

1 25

1 7

min

4.7

1 11

1 5

dev

1.466 0.53 0 2.50 2.15 1.96

0 0.74 3.46

ave

12.35 4.36 1 17.33 6.69 8.06

1 6.03 40.66

11.87 12.17 0 14.44 32.18 24.27

0 12.27 8.51

Table 1: Fuzia dadao sp. nov. Variability of forewing venation. n –
number  of  wings  (one  individual  occasionally  reveal  data  for  both
wings (LR)); min – minimum;  max – maximum; dev – standard
deviation; ave – average; CV – coefficient of variation; Sc – sub-
costa; R – radius 1+radius sector (when differentiated); M – media;
CuA – cubitus anterior; CuP – cubitus posterior; A – anal veins;
L – left, R – right. Total – total number of veins (all veins, and CV,
are measured when meeting margin). Length and width in mm. Analy-
sed  males:  042,  TNP42982LR,  CNU-B-NN-2006-322LR,  357L,
666LR, 669R, 671R, 672LR, 676LR, 677L; analysed females: CNU-B-
NN-2006-305R,  306LR,  314LR,  328LR,  341LR,  348LR,  371LR,
380R, 381R, 383R, 1001R, 1002LR, 1003R, 1005R.

461

BLATTIDA: FUZIIDAE FAM. NOV. – ADVANCED MORPHOLOGY OF EXTINCT EARWIG-LIKE COCKROACHES

males were active in flight. Conversely, the large pronotum, a
highly  protective  structure  suggests  cryptic  habits  and  the
presence  of  litter  (Mesozoic  cockroaches  with  such  a  large
pronotum  are  extremely  rare,  even  when  many  living  cock-
roaches  have  pronota  much  more  pronounced.).  An  entirely
open landscape of the Daohugou source area might also be ex-
cluded based on the presence of an abundant flora.

Another phylogenetic trend is also apparent – the reduction

of leg carination, indicating the stress in the protection was pro-
gressively changed from a passive, morphological, to an active,
behavioural defence. It should have been connected with the ra-
diation of effective predators such as beetles, but also the car-
nivorous cockroaches during the Early and Middle Jurassic.

It can be argued that in living cockroaches reduction of leg

carination  relates  more  to  the  kind  of  substratum  the  cock-
roaches live in (bark, leaf litter, plants leaves, etc.), but in all
Mesozoic  cockroaches  carination  is  extensive.  It  is  reduced
only  in  actively  moving  jumping  cockroaches  (Vršanský
2007),  in  the  blattulid  Tarakanula  shcherbakovi  Vršanský,
2003, adapted to rapid running, and in extinct eusocial cock-
roaches (P. Vršanský, submitted).

Nevertheless, cryptic habits (in leaf litter for instance) can-

not be definitely excluded for Fuzia, because its short legs are
not a good defence against an agile predator.

The dietary specialization of Fuzia dadao sp. nov. can, to

some  extent,  be  inferred  from  the  mouthpart  morphology.
Short  palps,  and  long  and  wide  mandibles  exclude  the  pa-
lynivory  [a  single  known  palinivorous  extinct  cockroach
Skok  svaba  Vršanský,  2007  has  long,  cup-like  palps  and
small  mandibles  (Vršanský  2007)]  as  well  as  carnivory  [car-
nivorous  cockroaches  have  elongated  mandibles  with  sharp
teeth (Vishniakova 1973; Liang et al. 2009); early carnivorous
mantises have wide but short and robust mandibles (Vršanský
2002)]. This suggests saprophagous diet analogical to those
of  most  cockroaches,  particularly  to  Blattulidae,  that  also
have short palps.

Conclusions

– Fuzia dadao gen. et sp. nov. from the Bathonian Middle

Jurassic of the Daohugou in Inner Mongolia, China, represents
a new family Fuziidae. It was a species with cryptic habits in
shrub source-area, with females active in flight. The presence
of male forceps attaching to the notch in the female external
rigid ovipositor is unique. Male and female body size is simi-
lar, but a peculiarity is the variability range of females, falling
within the variability range of males;

– According to the basal position of the present species with

respect to Mesozoic cockroaches, the presence of male tergal
glands is a plesiomorphic character for most Mesozoic and all
living cockroaches (excluding the very basal Phyloblattidae
and Caloblattinidae);

– Mesozoic cockroaches had a modern morphology, super-

ficially similar to living cockroaches.

Acknowledgments: We thank Alexandr P. Rasnitsyn (Paleon-
tological Institute, Russian Academy of Sciences, Moscow),
Klaus-Dieter Klass (Senckenberg Naturhistorische Sammlun-

gen  Dresden),  Barry  Lomax  (The  University  of  Nottingham),
Pavel Štys (Charles University) seven anonymous reviewers for
revision;  and  Chungkun  Shih  (Capital  Normal  Univeristy,
Beijing) for donating material. We also thank Tian-Tian Wang
(Capital  Normal  Univeristy,  Beijing)  for  providing  photo-
graphs and drawings of females for this publication.

Supported by UNESCO-Amba, UNESCO-IGCP 458,

VEGA 6002 and 2/0125/09, MVTS, Literárny fond, Schwarz
stipend,  National  Natural  Science  Foundation  of  China
(Nos. 30430100,  40872022),  the  Nature  Science  Foundation
of  Beijing  (No. 5082002),  Scientific  Research  Key  Program
(KZ200910028005)  and  PHR  Project  of  Beijing  Municipal
Commission of Education.

References

Anisyutkin L.N. & Gorochov A.V. 2008: A new genus and species of the

Cockroach Family Blattulidae from Lebanese Amber (Dictyoptera,
Blattina). Paleontol. J. 42, 1, 43—46.

Arillo A. 2007: Paleoethology: fossilized behaviours in amber. Geol.

Acta 5, 159—166.

Béthoux O. & Wieland F. 2009: Evidence for Carboniferous origin of the

order Mantodea (Insecta: Dictyoptera) gained from forewing mor-
phology. Zool. J. Linn. Soc. London 156, 1, 79—113.

Béthoux O., Galtier J. & Nel A. 2004: Earliest evidence of insect endo-

phytic oviposition. Palaios 19, 4, 408—413.

Bier K. & Müller W. 1969: DNA-Messungen bei Insekten und eine Hy-

pothese über retardierte Evolution und besonderen DNA-Reichtum
in Tierreich. Biologisches Zentralblatt 88, 425—449.

Brauckmann C., Schöllmann L. & Gröning E. 2007: Haemolymph-suck-

ing on Carboniferous insects: presumed parasitic mites (Acarina) on
Vorhalle Neoptera. Verh. Naturwiss. Vereins, Hamburg (NF) 43,
57—63.

Chen W., Ji Q., Liu D., Zhang Y., Song B. & Liu X. 2004: Isotope geo-

chronology of the fossil-bearing beds in the Daohugou area,
Ningcheng, Inner Mongolia. Geol. Bull. China 23, 12, 1165—1169
(in Chinese with English abstract).

Durden C.J. 1988: Hamilton insect fauna. In: Mapes G. & Mapes R.H.

(Eds.): Regional geology and paleontology of Upper Paleozoic
Hamilton Quarry area in Southeastern Kansas. Geol. Soc. Amer.,
Lawrence, KS, 117—124.

Engel M.S. & Grimaldi D.A. 2005: Primitive new ants in Cretaceous am-

ber from Myanmar, New Jersey, and Canada (Hymenoptera: Formi-
cidae). Amer. Mus. Novit. 3485, 1—23.

Foster C.B. & Afonin S.A. 2005: Abnormal pollen grains: an outcome of

deteriorating atmospheric conditions around the Permian—Triassic
boundary. J. Geol. Soc. 162, 4, 653—659.

Gao K.-Q. & Ren D. 2006: Radiometric dating of ignimbrite from Inner

Mongolia provides no indication of a post-Middle Jurassic age for
the Daohugou Beds. Acta Geol. Sin. 80, 1, 42—45.

Gao K.-Q. & Shubin N. 2003: Earliest known crown-group salamanders.

Nature 422, 424—428.

Grimaldi D. 1997: A fossil mantis (Insecta, Mantodea) in Cretaceous am-

ber of New Jersey, with comments on the early history of the Dicty-
optera. Amer. Mus. Novit. 3204, 1—11.

Handlirsch A. 1903: Zur Phylogenie der Hexapoden. Sitz.-Ber. Math.-

Naturwiss. Kl. K. Akad. Wiss. 112, 1, 716—738.

Handlirsch A. 1906: Die fossilen Insecten und die Phylogenie der rezent-

en Formen. Bd. I. Engelmann, Leipzig, 1—1340.

Hennig W. 1981: Insect phylogeny. John Wiley and Sons, New York,

1—514.

Hurd L.E., Prete F.R., Jones T.H., Singh T.B., Co J.E. & Portman R.T.

2004: First identification of a putative sex pheromone in a praying
mantid. J. Chem. Ecol. 30, 1, 155—166.

Inward D., Beccaloni G. & Eggleton P. 2007: Death of an order: a com-

prehensive molecular phylogenetic study confirms that termites are

462

VRŠANSKÝ, LIANG and REN

eusocial cockroaches. Biol. Lett. 3, 3, 331—335.

Jarzembowski E.A. 1994: Fossil Cockroaches or Pinnule Insects? P. Ge-

ologist Assoc. 105, 301—311.

Karny H. 1908: Die zoologische Reise des naturwissenschaftlichen Vere-

ines nach Dalmatien im April 1906. B. Spezieller Teil. etc. 6. Or-
thoptera und Blattaeformia. Mitt. Naturwiss. Ver. Univ. Wien 6,
101—113.

Klass K.D. 1997: The external male genitalia and the phylogeny of Blat-

taria and Mantodea. Bonner Zoologische Monographien 42, 1—341.

Klass K.D. 1998: The ovipositor of Dictyoptera (Insecta): Homology and

ground-plan of the main elements. Zoologischer Anzeiger 236, 2—3,
69—101.

Klass K.D., Picker M.D., Damgaard J., van Noort S. & Tojo K. 2003:

The taxonomy, genitalic morphology, and phylogenetic relation-
ships of southern African Mantophasmatodea. Entomologische Abh.
61, 3—67.

Krassilov V.A. 2003: Terrestrial palaeoecology and a global change.

Pensoft, Sofia, 1—464.

Krassilov V.A. & Rasnitsyn A.P. 1982: A unique finding: pollen in the

intestine of Early Cretaceous sawflies. Paleontol. Zhurn. 4, 83—96
(in Russian, English translation: Paleontol. J. 16, 4, 80—94).

Labandeira C.C. 2000: The palaeobiology of pollination and its precur-

sors. In: Gastaldo R.A. & DiMichele W.A. (Eds.): Phanerozoic ter-
restrial ecosystems. Paleont. Soc. Pap. 6, 233—269.

Labandeira C.C. 2002: The history of associations between plants and

animals. In: Herrera C.M. & Pellmyr O. (Eds.): Plant-animal in-
teractions: An evolutionary approach. Blackwell Science, London,
26—261.

Labandeira C.C. 2006: Silurian to Triassic plant and hexapod clades and

their associations: new data, a review, and interpretations. Arthro-
pod Systematics & Phylogeny 64, 1, 53—94.

Latreille P.A. 1810: Considérations générales sur l’ordre naturel des ani-

maux composant les classes des Crustacés, des Arachnides et des
Insectes avec un tableau méthodique de leurs genres disposés en
familles. Schoell, Paris, 1—444.

Liang J.-H., Vršanský P., Ren D. & Shih C. 2009: A new Jurassic carniv-

orous cockroach (Insecta, Blattaria, Raphidiomimidae) from the In-
ner Mongolia in China. Zootaxa 1974, 17—30.

Liu Y. & Jin X. 2002: Mesozoic biota-bearing depositional successions

in the Lingyuan basin, western Liaoning, and their ages: a discus-
sion. Geol. Bull. Chin. 21, 592—595 (in Chinese with English ab-
stract).

Liu Y., Liu Y., Li P., Zhang H., Zhang L., Li Y. & Xia H. 2004: Dao-

hugou  biota-bearing  lithostratigraphic  succession  on  the  southeast-
ern  margin  of  the  Ningcheng  basin,  Inner  Mongolia,  and  its
geochronology.  Geol.  Bull.  Chin.  23,  12,  1180—1187  (in  Chinese
with English abstract).

Lo N., Engel M.S., Cameron S., Nalepa C.A., Tokuda G., Grimaldi D., Ki-

tade O., Krishna K., Klass K.-D., Maekawa K., Miura T. & Thomp-
son G.J. 2007: Save Isoptera: A comment on Inward et al. Biol.
Lett. 3, 5, 562—563.

Martins-Neto R.G. 1999: A new subfamily of Ensifera (Insecta, Gryl-

loidea) from the Santana Formation (Lower Cretaceous), Araripe
Basin, NE Brasil. In: Proceedings of the First International Palae-
oentomological Conference, Moscow 1998, Bratislava, 91—97.

McKittrick F.A. 1964: Evolutionary studies of cockroaches. Cornell Uni-

versity Agricultural Experiment Station, Mem. 389, 1—197.

Nalepa C.A. 1994: Nourishment and the origin of termite eusociality. In:

Hunt J.H. & Nalepa C.A. (Eds.): Nourishment and evolution in in-
sect societies. Westview Press, Boulder, Colorado, 57—104.

Rasnitsyn A.P. 2002: 2.2.1.3.5. Superorder Vespidea Laicharting, 1781.

Order Hymenoptera Linné, 1758 ( = Vespida Laicharting, 1781). In:
Rasnitsyn A.P. & Quicke D.L.J. (Eds.): History of insects. Kluwer,
Dodrecht, 242—254.

Rasnitsyn A.P. & Krassilov V.A. 1996a: First find of pollen grains in the

gut of Permian insects. Paleontol. Zhurn. 3, 119—124.

Rasnitsyn A.P. & Krassilov V.A. 1996b: Sojadinielia floralis sp.nov. (In-

secta: Grylloblattida: Ideliidae) – one more pollen consumer of the
Permian Gymnosperms. Paleontol. J. 30, 6, 716—722.

Rasnitsyn A.P. & Zhang H. 2004: Composition and age of the Daohugou

hymenopteran (Insecta, Hymenoptera=Vespida) assemblage from
Inner Mongolia, China. Palaeontology 47, 6, 1507.

Ren D., Lu L.W., Ji S.A. & Guo Z.-G. 1995: Faunae and stratigraphy of

Jurassic-Cretaceous in Beijing and the adjacent areas. Seismic Pub-
lishing House, Beijing, 73—90 (in Chinese with English abstract).

Ren D., Gai K., Guo Z., Tan J. & Song Z. 2002: Stratigraphic division of

the Jurassic in the Daohugou area, Nengcheng, Inner Mongolia.
Geol. Bull. Chin. 21, 8—9, 584—591.

Roth L.M. & Willis E.R. 1958: An analysis of oviparity and viviparity in

the Blattaria. Trans. Amer. Entomol. Soc. 83, 221—238.

Schneider J. 1983: Die Blattodea (Insecta) des Paläozoikums Teil1: Sys-

tematik, Ökologie und Biostratigraphie. Freiberger Forschung-
shefte C382, 107—146.

Sharov A.G. 1968: Filogeniya ortopteroidnykh nasekomykh. Trudy Pale-

ont. Inst. Akad. Nauk SSSR 118, Nauka, Moscow 1—218 (in Russian;
English translation: Phylogeny of the Orthopteroidea. Keter Press,
Jerusalem, 1971, 1—251).

Shen Y., Chen P. & Huang D. 2003: Age of the fossil conchostracans

from Daohugou of Ningcheng, Inner Mongolia. J. Stratigraphy 27,
311—313 (in Chinese with English abstract).

Vidlička L. 2001: Blattaria – šváby, Mantodea – modlivky (Insecta:

Orthopteroidea). VEDA, Bratislava, 1—169 (in Slovak).

Vishniakova V.N. 1968: Mesozoic cockroaches with external ovipositor

and peculairities of their reproduction. In: Rohdendorf V.V. (Ed.):
Jurassic insects from Karatau. Nauka, Moscow, 55—86.

Vishniakova V.N. 1973: New cockroaches (Insecta: Blattodea) from the

Upper Jurassic sediments of Karatau ridge. In: Narchuk E.P. (Ed.):
Problems of the insect Palaeontology. Lectures on the XXIV Annual
Readings in Memory of N.A. Kholodkovsky (1—2 April, 1971). Nau-
ka, Leningrad, 64—77 (in Russian).

Vishniakova V.N. 1982: Jurassic Blattodea of the new family Blattulidae

in Siberia. Paleontol. J. 16, 67—78.

Visscher H., Looy C.V., Collinson M.E., Brinkhuis H., van Konijnen-

burg-van Cittert J.H.A., Kürschner W.M. & Sephton M.A. 2004:
Environmental mutagenesis during the end-Permian ecological cri-
sis. P. Natl. Acad. Sci. USA 10, 12952—12956.

Vršanský P. 1997: Piniblattella gen. nov. – the most ancient genus of

the family Blattellidae (Blattodae) from the Lower Cretaceous of
Siberia. Entomol. Probl. 28, 1, 67—79.

Vršanský P. 2000: Decreasing variability – from the Carboniferous to

the Present! (Validated on Independent Lineages of Blattaria). Pale-
ontol. J. 34, 3, 374—379.

Vršanský P. 2002: Origin and the Early Evolution of Mantises. Amba

projekty 6, 1, 1—16.

Vršanský P. 2003: Umenocoleoidea – an amazing lineage of aberrant

insetcs (Insecta, Blattaria). Amba projekty 7, 1, 1—32.

Vršanský P. 2004: Transitional Jurassic/Cretaceous Cockroach assem-

blage (Insecta, Blattaria) from the Shar-Teg in Mongolia. Geol.
Carpathica 55, 6, 457—468.

Vršanský P. 2005: Mass mutations of insects at the Jurassic/Cretaceous

boundary? Geol. Carpathica 56, 6, 473—481.

Vršanský P. 2007: Jumping cockroaches (Blattaria, Skokidae fam. n.)

from the Late Jurassic of Karatau in Kazakhstan. Biológia 62, 5,
588—592.

Vršanský P. 2008a: A complete larva of a Mesozoic (Early Cenoma-

nian) cockroach from the Sisteron amber. Geol. Carpathica 59, 3,
269—272.

Vršanský P. 2008b: New blattarians and a review of dictyopteran assem-

blages 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., Vishniakova V.N. & Rasnitsyn A.P. 2002: Order Blattida

Latreille, 1810. In: Rasnitsyn A.P. & Quicke D.L.J. (Eds.): History
of insects. Kluwer Academic Publishers, Dodrecht etc., 263—270.

Webster M. 2007: A Cambrian peak in morphological variation within

trilobite species. Science 317, 499—502.

Weesner F.M. 1969: The reproductive system. In: Krishna K. & Weesner

F.M. (Eds.): Biology of termites. Vol. I. Academic Press, New
York, 125—160.