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Zoological Journal of the Linnean Society, 2008, 152, 17–24. With 4 figures
New evolutionary evidence of Grylloblattida from the
Middle Jurassic of Inner Mongolia, north-east China
(Insecta, Polyneoptera)
DI-YING HUANG1*, ANDRÉ NEL2* and JULIAN F. PETRULEVIČIUS3
1
Nanjing Institute of Geology and Palaeontology, State Key Laboratory of Palaeobiology and
Stratigraphy, Chinese Academy of Sciences, Nanjing, 210008, China
2
CNRS UMR 5202, Muséum National d’Histoire Naturelle, CP 50, Entomologie, 45 Rue Buffon,
F-75005, Paris, France
3
CONICET, Argentina & CNRS UMR 5202, Muséum National d’Histoire Naturelle, CP 50,
Entomologie, 45 Rue Buffon, F-75005, Paris, France
Received 1 March 2006; accepted for publication 13 April 2007
The small modern insect order Grylloblattida has an abundant fossil record during the Late Palaeozoic and the
Mesozoicirca. The relationships between these fossil taxa and the modern grylloblattids remain unclear because
most of them are based on isolated wings or have poorly preserved body features. Modern grylloblattids are
wingless insects. The new grylloblattid family Plesioblattogryllidae fam. nov. is erected for the new genus and
species Plesioblattogryllus magnificus gen. nov., sp. nov., from the Middle Jurassic of north-eastern China.
The well-preserved specimen provides further evidence that could support its close relationships with the modern
grylloblattids: (1) several very similar head structures, e.g. developed laciniae with inner row of setae, maxillary
palps segmented into five, labial palps segmented into three, large labrum, and morphology of antenna; (2) paired
eoplantulae on tarsomeres 1–4; (3) long ovipositor and large eggs comparable with those of modern taxa. The new
genus has strongly developed mandibles with sharp pointed apical teeth and strong marginal teeth, and strong
hook-like fore claws with basal teeth, suggesting it was carnivorous. The major differences between the extinct and
extant Grylloblattida, such as the lack of wings, the eyes and ocelli either degenerated or absent, and the thorax
degenerated in the modern forms, are probably related to their adaptation to their life under rocks and
rock-crawler habits. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152,
17–24.
ADDITIONAL KEYWORDS: evolution – Grylloblattodea – Plesioblattogryllidae fam. nov.
INTRODUCTION
The small order Grylloblattida (rock crawlers), currently considered as ‘living fossils’, comprises 26
living species within four genera (Vrsansky et al.,
2001). Its fossil record is distinctly larger with more
than 44 families, even extending to the Late Carboniferous (Storozhenko & Vranský, 1995; Storozhenko,
1997). However, many of these taxa are based on
isolated wings, with their body features unknown.
*Corresponding author.
E-mail: [email protected]; [email protected]
Extant grylloblattids are wingless insects. The definition of the Grylloblattida sensu Storozhenko (1997,
2002: 279) is far from being clear. This author indicated that the synapomorphies of the Grylloblattida
are ‘absent because of paraphyletic state of the order
in respect to other perlideans’. Storozhenko (1998,
2002) considered that the Embioptera are ‘derived’
from the Permian grylloblattid Sheimia Martynova,
1958. There is still not consensus on the relationships between Plecoptera, modern Grylloblattidae,
Embioptera, and Dermaptera. Wheeler et al. (2001)
considered that the modern Grylloblattidae are the
sister group of the Dermaptera, being together the
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24
17
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D.-Y. HUANG ET AL.
sister group of (Phasmida + Orthoptera), and that the
(Plecoptera + Embioptera) are not directly related to
this clade. Beutel & Gorb (2001) considered that the
Grylloblattidae are the sister group of the clade
{Phasmatodea + [Mantodea + (Isoptera + Blattodea)]}.
Grimaldi (2001) supposed that the extant and fossil
‘Grylloblattodea’ falls in an unresolved polytomy with
the majority of the other ‘polyneopteran’ orders. Terry
& Whiting (2005) and Cameron, Barker & Whiting
(2006) considered that the Grylloblattida could be the
sister group of the recently discovered order Mantophasmatodea, and altogether could be the sister
group of the Dictyoptera. Grimaldi & Engel (2005:
223) also supposed that the Palaeozoic and Mesozoic
alate ‘grylloblattids’ could ‘represent a stem group of
both apterous Grylloblattida and Mantophasmatodea.
Lastly, even if Rasnitsyn (1976, 1980) listed several
similarities between Blattogryllus and the extant
Grylloblattidae, the potential synapomorphies of
this last group with the fossil ‘Grylloblattida’ are not
clear.
Thus, the present discovery of a new Middle Jurassic fossil insect attributable to the ‘Grylloblattida’,
with well-preserved body structures, is potentially of
great interest to solve the problem of the relationships between the extant Grylloblattidae and the
fossil ‘Grylloblattida’.
We follow the terminology of the wing venation of
Storozhenko (1998) in order to allow a more simple
comparison of our fossil with those already described,
even if we reserve our opinion on its real phylogenetic
value, especially concerning the problem of the exact
nature of the vein ‘M5’ (see Kukalová-Peck, 1991).
SYSTEMATIC PALAEONTOLOGY
ORDER ‘GRYLLOBLATTIDA’ WALKER, 1914
(SENSU STOROZHENKO, 2002)
PLESIOBLATTOGRYLLIDAE FAM. NOV.
Type genus: Plesioblattogryllus gen. nov.
Diagnosis: Wing venation nearly identical to that of
Blattogryllus karatavicus Rasnitsyn, 1976; mandibles
very strong with a sharp pointed apical tooth and a
few marginal teeth with a broad base; eyes and ocelli
present; antenna short with segments 2–7 shorter
than others; pronotum with broad lateral expansions;
tarsomeres 1–4 with a pair of rather large euplantulae; tarsi with strong claws and no arolia; fore tarsal
claw with a large base and strong basal tooth.
Remark: Some new material of grylloblattids was discovered in the same locality after recent fieldwork.
Two new forms seem to be closely related or attributable to the new family. Thus, a further cladistic
analysis of the Plesioblattogryllidae will be possible in
the near future.
PLESIOBLATTOGRYLLUS
GEN. NOV.
Derivation of the name: Named after the Greek
‘plesios’ meaning close or near, and Blattogryllus.
Type species. Plesioblattogryllus magnificus sp. nov.
Diagnosis. That of the family.
PLESIOBLATTOGRYLLUS MAGNIFICUS
GEN. NOV., SP. NOV.
FIGURES 1–3
Derivation of the name: Named after the wonderful
state of preservation of the type specimen.
Material: Only the holotype included (NIGP 133701).
The holotype is an almost complete female specimen
with brownish colour, dorsally compressed, and
without counterpart. The material is deposited in the
Nanjing Institute of Geology and Palaeontology.
Type locality: Daohugou Village, Ningcheng County,
Inner Mongolia, north-east China; Middle Jurassic,
Jiulongshan Formation.
Description: Head 10.5-mm long, 7.7-mm wide,
rather developed, upper surface covered with small
tubercles and tiny setae; eyes 3.4-mm long, 1.9-mm
wide (left one), 4.0-mm apart, in lateral position,
subelliptic in shape, contracted posteriorly; three
ocelli 0.6 mm in diameter at middle, olive shaped,
disposed in a triangle but nearly aligned between
eyes; left mandible 4.8-mm long (without tip), 2.8-mm
wide at base, right mandible 5.2-mm long and 2.6-mm
wide at base, both very robust, asymmetrical
(problem of preservation?), left mandible with a long,
curved, and sharp pointed apical tooth and three
marginal teeth, first and second being strong and
pointed, and third marginal tooth located near base
and very blunt; lacinia c. 4-mm long and very developed, pointed at apex, with a row of dense setae in
middle inner margin; preserved part of maxillary palp
4.5-mm long, developed, with five segments of distinctly increasing width forwards, covered with fine
setae; labial palp 4.3-mm long, developed, with three
elongated segments of nearly the same width, covered
with fine setae, second segment being shorter than
the other two, the last one distinctly swollen and
rounded at apex; clypeus 1.2-mm long, 2.8-mm wide,
strong, with the shape of a transverse ellipse; labrum
2.5-mm long, developed, slightly projected in median
part of anterior edge, extending to two-thirds of the
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NEW MESOZOIC GRYLLOBLATTID FAMILY
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Figure 1. Plesioblattogryllus magnificus gen. nov., sp. nov., holotype NIGP 133701. A. Photograph of general
habitus (scale bar represents 10 mm). B. Photograph of foreleg (scale bar represents 2 mm). C. Photograph of head (scale
bar represents 5 mm). D. Photograph of hind leg (scale bar represents 2 mm). E. Photograph of eggs (scale bar represents
2 mm).
mandibles; antenna c. 15-mm long, as long as
head + pronotum, its base located close to the mandible; about 24 antennomeres, first one large, second
shortest, third rather long, segments 4–7 distinctly
shorter than the others, segments 8–24 with a
notable decreasing width, last one with rounded end,
all antennomeres covered with fine setae; ecdysial
distinctly visible, Y-shaped.
Pronotum 5.2-mm long medially, 5.7-mm long
laterally, 9.3-mm wide, broader than head, covered
with tiny setae, with middle line and large rounded
lateral expansions, anterior margin slightly projected, with pronounced angles, posterior margin
rounded; mesonotum 4.2-mm long, about 4.7-mm
wide, contracted backwards, distinctly narrower than
pronotum.
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D.-Y. HUANG ET AL.
Figure 2. Plesioblattogryllus magnificus gen. nov.,
sp. nov., holotype NIGP 133701. Drawing of general
habitus.
Legs all similar; femora, tibiae, and tarsi densely
covered with distinct setae; fore and hind coxae
visible, and rather elongate; forelegs rather sturdy,
femur 6.8-mm long, 1.8-mm wide, tibia 5.7-mm long,
0.8-mm wide; median legs rather short, femur 9.2-mm
long, 1.7-mm wide, tibia 6.5-mm long, 1.1-mm wide;
posterior legs slender, femur 9.6-mm long, 1.9-mm
wide, tibia 9.7-mm long, 0.8-mm wide, tarsi 5.0-mm
long; inner margin of median tibia with three strong
aligned spines; inner margin of posterior tibia with
four strong aligned spines, apical end with three
strong spines; all tarsi with five segments,
tarsomeres 1–4 with a pair of rather large euplantulae, fifth segments rather elongate with increased
width; no arolia but two claws, hind tarsal claws
0.7-mm long, strong, and rather straight, fore tarsal
claws 0.9-mm long, with a large base and a strong
basal tooth.
Wings partly overlapping and difficult to interpret,
in particular for the anal areas; wings dark infuscate
with darker spots in distal halves; preserved parts of
Figure 3. Plesioblattogryllus magnificus gen. nov.,
sp. nov., holotype NIGP 133701. Drawing of head.
fore and hindwings similar, but the possible differences in the anal areas are not preserved; wings
hyaline; forewing 43-mm long, about 11-mm wide;
costal area between subcosta (Sc) and anterior wing
margin narrow, 1.0-mm wide, in all wings, with about
20 more or less sigmoidal cross-veins between Sc and
anterior wing margin; costal vein reaching wing apex;
vein Sc ending on anterior wing margin 3.7 mm to
wing apex; radius (R) ending very close to wing apex;
Sc and R closely parallel; several cross-veins in area
between Sc and R, the most distal ones being oblique;
base of Rs 22 mm from wing base, in basal half of
wing; area between R and radial sector (Rs) rather
narrow, 1.2-mm wide in its broadest part, with one or
two rows of cells; Rs posteriorly pectinate, with five
main branches; cubitus apex (CuA) emerging from Cu
at wing base and basally fused with common stem
R + media (M); R and M + CuA separating about
4 mm from wing base; M + CuA anteriorly pectinate,
with first branch media anterior (MA) emerging 7 mm
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24
NEW MESOZOIC GRYLLOBLATTID FAMILY
from its base; MA posteriorly pectinate, with three
branches in its distal fourth; media posterior (MP)
emerging 10 mm from base of M + CuA and apparently simple; CuA posteriorly pectinate with three
long and simple branches; cubital area between
M + CuA and cubitus posterior (CuP) broad, with long
sigmoidal cross-veins; CuP simple; anal vein 1 (A1)
posteriorly pectinate with three simple branches; A2
forked; hindwing about 40-mm long.
Abdomen 15.8-mm long, segmentation not clearly
visible, first visible abdominal segment shorter than
half of the second, and other visible segments with
distinct lateral lobes; last segment very small with
slight lateral angles; two long, parallel-sided, and
straight structures corresponding to the ovipositor,
c. 8.0-mm long; right cercus (?) with three visible
elongate segments, second segment distinctly longer
than the first one; at least 11 visible large oliveshaped eggs, c. 3-mm long, 1.5-mm wide, with several
strongly developed longitudinal ridges, first egg at
level of second abdominal segment, but smaller than
the more distal eggs.
COMPARISON WITH FOSSIL
GRYLLOBLATTIDA
After the available phylogenetic analysis of the
fossil taxa currently attributed to the Grylloblattida,
P. magnificus gen. nov., sp. nov., would fall in the
Blattogryllidae Rasnitsyn, 1976 because of the following characters, after Storozhenko (1998, 2002): (1)
forewing with sigmoidal cross-veins in area between
CuP and M + CuA/CuA. Storozhenko (2002) interpreted them as branches of CuA terminating on CuP;
(2) costal area of fore and hindwings narrow; (3)
cross-veins of forewing area between Sc and C simple,
not H- or Y-shaped. Storozhenko (2002: 284) indicated
that the Blattogryllidae have ‘Sc veinlets H- or
Y-shaped’, which is absolutely not the case in all the
taxa currently included in the family; (4) forewing
veins CuA and M basally fused (no veinlet ‘M5’
between them); (5) forewing cubital area with a series
of branches of CuA directed to posterior wing margin;
(6) forewing CuA with a strong anterior flexure at
base. In fact, its forewing venation is closely similar
to that of B. karatavicus Rasnitsyn 1976.
However, Storozhenko (1988a, b, 2002: 284) indicated that both B. karatavicus and Blattogryllulus
mongolicus have tarsi with a large arolia and lacking
claws. Plesioblattogryllus has strong claws and no
arolia. These two characters are sufficient to separate
Plesioblattogryllus from the Blattogryllidae, as the
type species of this last family is B. karatavicus.
Another possible difference is the thorax as broad
as pronotum in the Blattogryllidae, unlike the thorax
21
with broad lateral expansions of Plesioblattogryllus.
But it is necessary to be prudent with this character
because of the taphonomic deformations of the body
structures in these insects with large thoraces and
heads. Interestingly, the thorax of Parakhosara
nasuta Storozhenko, 1993 (Megakhosaridae Sharov,
1961; sister group of Blattogryllidae + Mesoblattinidae, after Storozhenko, 2002) is similar to that of
Plesioblattogryllus.
We also have to compare Plesioblattogryllus with
the taxa currently attributed to the Blattogryllidae
because some of these taxa are based on isolated
wings:
(i) Baharellus Storozhenko, 1988a (two species:
Baharellus lineatus Storozhenko, 1988a, Middle
Jurassic, and Baharellus madygensis Storozhenko,
1992; Triassic). Both have the forewing Rs with only
two main branches and with a corresponding area
relatively reduced (Storozhenko, 1988a, 1992, 1998).
(ii) Blattogryllulus Storozhenko, 1988a (two species:
B. mongolicus Storozhenko, 1988a, and Blattogryllulus lucidus Storozhenko, 1988a, both Upper Jurassic).
Both have a forewing MA and CuA simple, and an
Rs not pectinate, with a double fork (Storozhenko,
1988a).
(iii) Parablattogryllus Storozhenko, 1988a (one
species: Parablattogryllus obscurus Storozhenko,
1988a, Lower Cretaceous). This taxon is based on a
hindwing, even if it was labelled as a forewing in
Storozhenko (1988a: fig. 1.i). Thus, it is difficult
to compare with our specimen but its Rs is not
pectinate.
(iv) Dorniella Bode, 1953 (two species: Dorniella
pulchra Bode, 1953; Lower Jurassic, Dorniella primitiva Storozhenko, 1992, Middle or Upper Triassic).
Both have Rs with only two branches (Storozhenko,
1988a, 1992). D. primitiva is based on a hindwing,
which is very difficult to compare with the known
forewings of D. pulchra. Thus the generic identity of
these two species is questionable.
(v) Costatoviblatta Storozhenko, 1992 (one species:
Costatoviblatta aenigmatosa Storozhenko, 1992;
Middle or Upper Triassic).
(vi) Mesoblattogryllus Storozhenko, 1990 (three
species: Mesoblattogryllus intermedius Storozhenko,
1990; Mesoblattogryllus conjuctus Storozhenko, 1992;
both from Middle or Upper Triassic, Mesoblattogryllus
longipennis Storozhenko, 1992; Lower or Middle
Jurassic). These two genera have a short Sc and its
first branch of Rs is pectinate, instead of being simple
(Storozhenko, 1992, 1998).
(vii) Baharellinus Storozhenko, 1992 (two species:
Baharellinus dimidiatus Storozhenko, 1992; Baharellinus pectinatus Storozhenko, 1992; both from Middle
or Upper Triassic). They have very different wing
shapes, rounded in B. dimidiatus and very elongate
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D.-Y. HUANG ET AL.
in B. pectinatus. Nevertheless, their Sc is short and
Rs has three branches (Storozhenko, 1992, 1998).
(viii) Protoblattogryllus Storozhenko, 1990 (three
species: Protoblattogryllus zajsanicus Storozhenko,
1990; Protoblattogryllus asiaticus Storozhenko, 1990;
Protoblattogryllus abruptus Storozhenko, 1990;
Middle or Upper Triassic). They have a short Sc and
an Rs with two branches (Storozhenko, 1990).
(ix) Microblattogryllus Storozhenko (1990) (one
species: Microblattogryllus variabilis Storozhenko,
1990; Middle or Upper Triassic). It has a short Sc and
a simple Rs.
(x) Anoblattogryllus Storozhenko, 1990 (one species:
Anoblattogryllus fundatus Storozhenko, 1990; Middle
or Upper Triassic). Its Sc is short.
(xi) Gryphopteron Handlirsch, 1939 (one species: Gryphopteron molle Handlirsch, 1939; Lower Jurassic).
Its MA is fused with Rs, and its Rs has two distal
branches (Ansorge, 1996).
(xii) Megablattogryllus Storozhenko, 1990 (two
species: Megablattogryllus magister Storozhenko,
1990; Megablattogryllus austerus Storozhenko, 1990;
Megablattogryllus pinguis Storozhenko, 1990; Middle
or Upper Triassic). Their CuA emerges first from
M + CuA, then MP, and lastly MA, unlike our fossil.
(xii) Blattogryllus Rasnitsyn, 1976 (two species:
B. karatavicus Rasnitsyn, 1976; Upper Jurassic, Blattogryllus rasnitsyni Storozhenko, 1990; Lower Jurassic). The only difference of Plesioblattogryllus with
Blattogryllus in the wing venation is its MP being
apparently simple. All other characters are identical
(Rasnitsyn, 1976; Storozhenko, 1990).
Remark: All the species currently attributed to the
Blattogryllidae based on isolated wings now fall into
uncertain position between the Blattogryllidae and
the Plesioblattogryllidae because these two families
can be separated only on the basis of the body
characters.
EVOLUTIONARY RELATIONSHIP
The present discovery and characters listed in Rasnitsyn (1976) show that there are numerous structures of at least some fossil ‘Grylloblattida’
(Blattogryllidae and Plesioblattogryllidae) similar to
those of extant Grylloblattidae. But these characters
have never been polarized in a phylogenetic analysis.
Thus, the establishment of a direct phylogenetic relationship between these groups remains questionable.
A well-supported phylogenetic analysis of all polyneopterous orders, including the fossil taxa, is necessary. Re-analyses of the characters available in fossils
(wing venation) could help to solve this problem, as
Béthoux & Nel (2002a, b) already showed for the
orthopteroid taxa.
Although more than 44 extinct families have been
attributed to Grylloblattida, very few fossils have
well-preserved body structures (Storozhenko &
Vranský, 1995; Storozhenko, 1997). The new fossil
material from the Middle Jurassic of north-east
China provides some new similarities with the
modern grylloblattids.
1. Some head structures of P. magnificus gen. nov.,
sp. nov. correspond to those of the living grylloblattids, namely: laciniae developed with an inner row of
marginal setae; maxillary palp segmented into five;
labial palp segmented into three; labrum developed;
antenna with a large first segment, smallest second
segment, and segments 2–7 distinctly shorter than
the others. The structure of the antennomeres, viz.
antennomere 2 smaller than 3, and antennomeres
4–7 distinctly shorter than the other distal segments,
visible in Plesioblattogryllus, is a rather typical
feature present in extant grylloblattids, but also in
the modern Embioptera Embia major (Crampton,
1917: fig. 9). The rather developed antennomere 3 and
its adjacent segments are especially similar to Galloisiana yezoensis (Asahina, 1961), and the third
antennomere is even more developed in some other
species such as Galloisiana biryongensis (Namkung,
1974). The mandibles and eyes in the new genus are
remarkably larger than those of the extant forms.
Moreover, the large ocelli of our fossil, absent in all
modern grylloblattids, is an important difference. In
addition, no evidence indicates that the clypeus of
Plesioblattogryllus is divided into an anteclypeus and
a postclypeus, as in extant forms.
2. The five-segmented tarsi and the tarsal euplantulae of P. magnificus gen. nov., sp. nov. are strongly
similar to those of some modern grylloblattids
(Walker, 1938: pl. 10; Caudell & King, 1924: plate 3:
2–3; Namkung, 1974: figs 10,11; Storozhenko, 1998;
Beutel & Gorb, 2006). However, Crampton (1927:
125) indicated that ‘some Mantids have flange-like
tarsal pads very suggestive of those occurring in
Galloisiana and in certain Gryllacrids’ (see also
Crampton, 1933). Thus, this type of euplantulae
seems to be not uniquely present in Grylloblattida,
but also occurs in some Mantodea and Orthoptera.
Thus, it is not sufficient alone to attribute Plesioblattogryllus to the same clade as the extant Grylloblattidae. Nevertheless, the wing venation and body
structures of Plesioblattogryllus strongly differ from
those of the Mantodea and Orthoptera, suggesting
that it belongs to a very different order. The marginal and apical strong spines of posterior tibia in
Plesioblattogryllus also display similarities with the
modern forms.
© 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 17–24
NEW MESOZOIC GRYLLOBLATTID FAMILY
3. The ovipositor and eggs are similar to those of
modern forms. We interpret two long and parallelsided straight structures as the ovipositor bases. Such
structures are not only comparable with the ovipositor
of extant female grylloblattids, but also to the similar
structures described in some other extinct grylloblattid
taxa (Storozhenko & Novokshonov, 1994). The large
elliptic eggs are similar to those of modern grylloblattids (Crampton, 1927, 1933). They occupy a broad
space in the abdomen, suggesting the adult female of
Plesioblattogryllus had large ovaries. Similar size and
arrangement of eggs in the abdomen are also found in
some other Mesozoic grylloblattids (family Tshekardominidae) (Novokshonov & Aristov, 2002).
In addition, the body of P. magnificus is covered with
tiny setae as are those of the modern grylloblattids.
The well-developed wings of this large insect
suggest it was probably an active flyer. Its fore tarsal
claws differ from those of hind legs (the middle claws
are not visible) in their hook-like shape with a large
base and bearing a strong basal tooth. Moreover, the
fore tarsal claws are also distinctly longer than
the rather straight hind claws. After preservation, the
fore tarsal claws and the most apical tarsomeres were
probably movable structures. Such structures could
have been associated with the capture of small prey.
The very strong mandibles with a sharp pointed
apical tooth could help to kill the prey, and the strong
marginal teeth would help to destroy it. The developed eyes and ocelli suggest it was an active hunter.
Plesioblattogryllus was probably a predator.
The fossil evidence indicates that the Permian grylloblattid Tillyardembia was probably a pollen eater
(Afonin, 2000). Therefore the food habits of the early
grylloblattids were very diverse. The modern grylloblattids probably became adapted to live under rocks or
hidden in moss liver through the degeneration of their
sensitive organs and wings. Even the eyes are absent
in some cave-living species (Namkung, 1974). Their
thorax trends to be very developed in relation to their
habit of being a strong rock crawler. Their mandibles
degenerated in relation to their habit of omnivory.
Nevertheless, the modern Grylloblatta can also attack
a living insect if it is very hungry (Ford, 1926).
ACKNOWLEDGEMENTS
We are pleased to acknowledge the joint support
provided by the NSFC (grant no. 40672013,
40632010), the Major Basic Research Projects of MST
of China (2006CB806400), and the State Key Laboratory of Palaeobiology and Stratigraphy (No. 053106,
CAS). We thank Prof. Shu-Yong Wang (Institute of
Zoology, Chinese Academy of Sciences) for kind bibliographic help.
23
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