JURASSIC/CRETACEOUS COCKROACH (INSECTA, BLATTARIA) FROM MONGOLIA 457
GEOLOGICA CARPATHICA, 55, 6, BRATISLAVA, DECEMBER 2004
TRANSITIONAL JURASSIC/CRETACEOUS COCKROACH
ASSEMBLAGE (INSECTA, BLATTARIA) FROM THE SHAR-TEG
Arthropoda Laboratory of the Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117868 Moscow, Russia
Department of Zoology, Faculty of Natural Sciences, Comenius University, Mlynská Dolina, 842 15 Bratislava, Slovak Republic
Present address: Geological Institute, Slovak Academy of Sciences, Dúbravská cesta 9, 840 05 Bratislava, Slovak Republic;
(Manuscript received September 16, 2003; accepted in revised form December 16, 2003)
Abstract: A unique Upper Jurassic assemblage of cockroaches with 7 new and one unidentified species occurs in the
Shar-Teg in Mongolia (Shartegoblattina elongata gen. et sp. nov., Elisamoides mantiformis gen. et sp. nov., Breviblattina
minor gen. et sp. nov., Mongolblatta accurata gen. et sp. nov., Blattula mongolica sp. nov., Blattula vidlickai sp. nov.,
Elisama pterostigmata sp. nov.). The assemblage is the most progressive among Jurassic sites worldwide and displays
taxa characteristic for the warm and dry Lower Cretaceous assemblages: a mantid species, two species of Blattulidae
represented by BlattulaElisama complex (in the Cretaceous replaced by two sister and additional species of Elisama)
including small representatives which probably occupied the niche of the Cretaceous HolocompsidaeVitisminae, and
diverse Mesoblattinidae. Additional aberrant species preliminarily placed in the Liberiblattinidae display strong
synapomorphies with the mantises. Mesoblattinidae are also known from the Lower Jurassic of Europe and Upper
Jurassic of Asia and Africa (generally less diverse assemblages). These assemblages are characterized by the diverse
Caloblattinidae and also the Blattulidae, which are also present in Shar-Teg. The assemblage contains a similar ratio of
fore- and hindwings probably resulting from an active decomposition. The presence of frequent malformations vein
fusions in different independent taxa, which indicates some ecological stress factors in the paleoenvironment is unusual.
Key words: Jurassic, Mongolia, systematics, taxonomy, taphonomy, phylogeny, new genera, new species, Blattaria,
Fossil plants, gastropods, pelecypods, ostracods, conchostra-
cans, insects, chelycerates, fishes, labyrintodont amphibians,
turtles, crocodiles, dinosaurs and mammals have been found at
the locality, indicating the uppermost Jurassic age of the local-
ity. Over 5000 specimens of insects (under 1000 species) have
been collected so far (Gubin & Sinitza 1996).
The uppermost Jurassic fossil site Shar-Teg has yielded
some one hundred and fifty mostly fragmentary specimens of
Blattaria, which are of high significance, since they include
some transitional taxa leading to contemporary cockroaches
and praying mantises (Vranský 2002). In addition some char-
acters are reported, which are new for the fossil Blattaria. The
assemblage itself is important in providing a picture of the Ju-
rassicCretaceous transitional taxa. The Cretaceous warm and
dry assemblage of Blattaria (Vranský 1999b) is found already
formed, with the niches of the Cretaceous species already
formed and filled with their ancestral taxa.
The complete assemblage allows its comparison with the
worlds most significant sites known in Kazakhstan (Vishniako-
va 1968), Europe (Vranský & Ansorge in print) and Africa.
Material and methods
About a hundred and fifty specimens of Blattaria housed in
the Paleontological Institute, Russian Academy of Sciences,
Moscow, Russia (PIN) are studied. The material comes from
the Shar-Teg locality in Mongolia. Shar-Teg is a diverse Late
Jurassic insect assemblage (some 5000 fossils of more than
200 families with more than 300 genera and 1000 species)
collected in mudstone of the Shar-Teg Beds at the Shar-Teg
Ula Mt SW of Adzh-Bogdo Mt, Gobi-Altai Aymag of Mongo-
lia (Gubin & Sinitza 1996; Ponomarenko 1998) (Fig. 1).
The photographs were taken using a NIKON camera and
line drawings were made using ROTRING 0.180.5 mm pens.
Abbreviations used: B body; P pronotum; Veins: Sc
Subcosta, R Radius, RS Radius Sector, M Media,
Cu Cubitus (A anterior, P posterior), A Anal; c
clavus; * specimen with teratologic fusion of wing veins.
Phyloblattoidea Schneider, 1983
Caloblattinidae Vranský et Ansorge in Vranský, 2000
Shartegoblattina gen. nov.
Type species: S. elongata sp. nov.
Composition: Type species.
Diagnosis: Sc richly branched, R slightly curved with as
much as 17 veins at margin, M and Cu both richly branched
(5+). Clavus as much as three times as long as wide with ex-
traordinary branchings of A (Fig. 6.5). Hindwing with differ-
Fig. 1. a Localization of the Shar-Teg locality (white rectangle)
in the framework of Mongolia. b Schematic lithological log of
the Shar-Teg profile. Arrows denote the presence of insect hori-
entiated RS, M weakly branched, CuA with secondary
branched veins, and additional blind branches. A
near the basis.
Remarks: The new genus is most closely related to Rhipi-
doblattina Handlirsch, 1908. Shartegoblattina differs only in
that clavus is narrow and R more curved (plesiomorphy).
Etymology: The name is after Shar-Teg. Gender: femi-
Shartegoblattina elongata sp. nov.
(Figs. 3.45, ?5.8, 6.56, 7.56)
Holotype: PIN 4270/1846±. Forewing.
Additional material: Forewings: PIN 4270/20c,
1785±, 1800c, 1818c, 1875, 1907, 1913; Hindwings: PIN
4270/1803±, 1804±, 1807±, 1814±, 1853±, 1865, 1889, 7194.
Body: PIN 4270/1863±.
Description: Forewing elongate, length about 14 mm,
width 3.7 mm. Sc 34 branched, R, M, and Cu all richly
branched (16+, 6+, 5). Clavus 4.24.4 mm long with peculiar-
ly branched anal veins.
Hindwing length ca. 1415 mm. Intercalaries and cross
veins both apparent. A
straight but branched near the basis
(3); R differentiated into R
(5) and RS (7+); M branched (3+);
Cu with about 7 veins plus descending additional blind
Etymology: The name refers to the elongate wing form.
Character of preservation: Forewing 8 (clavus 3);
hindwing 8; complete specimen 1.
Polyphagoidea Walker, 1868
?Liberiblattinidae Vranský, 2002
Elisamoides gen. nov.
Type species: E. mantiformis sp. nov.
Composition: Type species.
Diagnosis: R, M (simplified) and Cu (rich) all sharply
curved, intercalaries not straight but joined with cross-veins.
Reticulation abundant, mostly in area of joining M, Cu and R.
In this area, dark macula present. Hindwing pterostigma reach-
ing intercalary vein between R
Remarks: Curved main veins present a plesiomorphic
character. The presence of characteristic macula in the forew-
ing is similar to that of Blattulidae. Character of hindwing ve-
nation (intercalaries joined with cross-veins) resemble that of
early mantises. The species most probably represent the
Liberiblattinidae (Vranský 2002), the abberant Blattulidae or
a new family.
If placed in the Caloblattinoidae, Elisamoides would be
unique there in the wing form, in having forewing macula and
in simplified hindwing venation. Therefore it is placed within
The new genus can be placed within the Liberiblattinidae
with reservations, because of the poor state of preservation of
the material (Sc and A are not visible). The hindwing is similar
to that of Liberiblattina Vranský, 2002, but the pterostigma of
Elisamoides reaches, but does not overlap the RS. The charac-
ters indicating trends leading to mantises support the member-
JURASSIC/CRETACEOUS COCKROACH (INSECTA, BLATTARIA) FROM MONGOLIA 459
Fig. 2. 1 Blattula vidlickai sp. nov. Reconstruction based on specimens 4270/1850 (complete specimen), 1815 (hindwings), 1919
(forewing on the left), 1834 (hind leg) and 1837 (head-slightly enlarged to display the colouration); 23 Elisama pterostigmata sp.
nov. 2 Forewing (4270/1916). 3 Hindwing (4270/1948); 4 Blattula mongolica sp. nov. Forewing (4270/1798).
ship of the genus in the Liberiblattinidae. Liberiblattina dif-
fers in having an extensively coloured forewing (but without
dark macula). It probably represents a sister taxon to Elisam-
oides (synapomorphies of Elisamoides and Liberiblattina in-
clude simplified forewing M, colouration and hindwing
pterostigma). The forewing of Gurvanoblatta Vishniakova,
1986 is without reticulations (apomorphy), and has rich M.
Aktassoblatta Vishniakova, 1971 has rich hindwing branches
(plesiomorphy). Kazachiblattina Vranský, 2002 lacks the
forewing colouration and differs in having almost straight Cu
branches (apomorphy). Hindwing of Kazachiblattina is with-
out pterostigma, M is rich (plesiomorphies).
Elisamoides also might represent a taxon close to the stem
group of Blattulidae, but the more arcuated character of en-
branchements would be unique within the family. In that case,
also the presence of hindwing reticulations would be enigmat-
ic (both plesiomorphies). Nevertheless, the Blattulidae proba-
bly evolved from taxa like Elisamoides.
Until more complete specimens are collected, Elisamoides
is preliminarily placed within the Liberiblattinidae.
Etymology: The name refers to resemblence with Elisama
in the presence of macula.
Elisamoides mantiformis sp. nov.
(Figs. 3.67, 6.34)
Holotype: PIN 4270/1828. Forewing.
Additional material: Hindwings: PIN 4270/1844±, 1865.
Description: Forewing length about 8 mm (fragment
length 7 mm), width 2.75 mm. Reticulations most abundant in
front of clavus. R richly branched with about 10 branches, M
with few (about 5) veins, reaching before apex, Cu richly
branched (about 9 veins).
Hindwing with differentiated R and RS (5+68), branched
M (34) and incipient pterostigma covering intercalary be-
Etymology: The name indicates that the species appar-
ently shows a tendency that is realized in the Mantodea.
Character of preservation: Forewing 1, hindwing 2.
Fig. 3. 1 Breviblattina minor sp. nov. (4270/1843); 23 Mongolblatta accurata sp. nov. 2 Forewing (4270/1856). 3 Hindwing
(4270/1857); 45 Shartegoblattina elongata sp. nov. 4 Forewing (4270/1846). 5 Hindwing (4270/1807); 67 Elisamoides manti-
formis sp. nov. 6 Forewing (4270/1828), 7 Hindwing (4270/1838).
Blattulidae Vishniakova, 1983
Blattula Handlirsch, 1906
Blattula mongolica sp. nov. (Figs. 2.4, ?5.2, 6.2)
Holotype: PIN 4270/1798±*. Forewing.
Additional material: Forewings: PIN 4270/28,
1767±, 1826c, 1866.
Diagnosis: The largest (forewing length almost 12 mm)
blattulid of the site. Forewing characteristics: length/width:
≤12 mm/3.5 mm; R 13+; M 37; Cu 47; A 6+ (simple
JURASSIC/CRETACEOUS COCKROACH (INSECTA, BLATTARIA) FROM MONGOLIA 461
Fig. 4. Blattula vidlickai sp. nov. (Figs. 4.19: 4270/1850, 1837, 1815, 1919, 1903, 1881, 1847, 1792, 1881).
veins). Total number of veins about 30.
Description: The largest Blattula known. Sc probably
simple; R richly branched (without secondary branches),
curved; M almost straight, comparatively rich; Cu rich, reach-
ing apical quarter of the forewing.
Remarks: Because the single completely preserved wing
is generally deformed in shape (in addition to fusion between
M and R it has expanded R, which could be a result of the fu-
sion), it is very difficult to appreciate the phylogenetical posi-
tion of the species within Blattula.
The species differs from all the known Jurassic representa-
tives of the genus in being large.
Both B. mongolica and B. vidlickai described herein are with-
out characteristic dark macula in the forewing and therefore as-
signed to the genus Blattula, and not Elisama Giebel, 1856.
Etymology: The species name is after Mongolia.
Character of preservation: Forewing 5 (clavus 1).
Blattula vidlickai sp. nov. (Figs. 2.1, 4.19)
Holotype: PIN 4270/1897 = 1919. Forewing.
Additional material: Forewings: PIN 4270/21(+P),
25, 1759±, 1763±, 1764, 1771, 1773, 1777, 1780, 1782,
1783±, 1786, 1788, 1793, 1811±, 1829±, 1834±leg, 1854±,
1858±, 1861±, 1871=1903c, 1873, 1874, 1878±, 1885, 1891,
1893, 1909; Hindwings: PIN 4270/14, 1768, 1769, 1784±,
1792, 1799, 1815±, 1816±, 1819±, 1847±, 1872, 1881; Com-
plete specimens: PIN 4270/1850±, 1905.
Diagnosis: One of the smallest (forewing length under
6 mm) blattulids known. Hindwing with pterostigma indicat-
ed, and with richly branched A
(unique within Blattaria).
Forewing with simplified venation, with the total number of
veins about 25 (2327), with tip sharply curved. Massive
sheath covering ovipositor present.
Description: Forewing characteristics: length/width:
4.05.7 mm/1.31.9 mm; Sc 1; R 1012; M 27; Cu 25; A
56. Posterior anal vein may be forked.
Hindwing with the simple Sc, R
(24) + RS (26) with 68
veins and indicated pterostigma in the region of R
; M may be
occasionally richly branched, usually with 2 veins (27); CuA
with 35 veins; CuP simple; A
richly branched (67). A
to 4-branched. Cu with numerous cross-veins. Pronotum with
two parallel stripes.
Remarks: In spite of the strong autapomorphies discrimi-
nating the species among other Elisama species (richly
in the hindwing), the taxon is similar to
Vitisminae (Holocompsidae Rehn, 1951) in the general
groundplan of the hindwing and sharply curved CuP in the
forewing. The extremely small size of some specimens (up to
forewing length 4 mm in PIN 4270/1847) is also similar to
It differs from the most of the Blattulidae in its small size
such small Blattula species with the sharp tip of the forewing
are known also from the Lower Jurassic deposits of Germany
and England (Vranský & Ansorge, in print), but A
hindwing is not branched.
The absence of the cases of vein fusion in this most common
species is significant. This indicates its ability to resist the stress
factors which affected other roach species in the locality.
Etymology: The species is named after ¼ubomír Vidliè-
ka, the scientist who introduced me into the world of contem-
Character of preservation: Forewing 29 (clavus 2);
hindwing 12 (both hindwings 1); complete specimens 2.
Elisama Giebel, 1856
Elisama pterostigmata sp. nov. (Fig. 5.1,3,5,7)
Holotype: PIN 4270/1916*. Forewing.
Additional material: Forewings: PIN 4270/10, 24,
26, 1758±, 1770c, 1774*, 1775(B), 1801, 1802, 1832±,
1835±*, 1841c, 1845±, 1851±c, 1859±, 1875, 1894c, 1899.
Hindwings: PIN 4270/16, 22, 1765±(B), 1781, 1794, 1797±,
1828, 1848±, 1855±, 1920±.
Diagnosis: Pterostigma is slightly indicated in the hind-
wing. Total forewing length usually 78 mm. Macula is shift-
ed more centrally, comparing to the Cretaceous representa-
tives of the genus.
Description: Forewing characteristics: length/width:
6.59 mm/2.12.6 mm (clavus ca. 3 mm). Sc 1; R 1013; M
47; Cu 27, A 56 (simple veins).
Hindwing length 78 mm. The most stable venation charac-
ter R 6+4 (3+4 in a single specimen); M 3; Cu 45+1.
Remarks: Fusion of veins (fused are different branches
of M or of R and M) is present in at least 3 of 19 specimens
(most wings collected are incomplete) being an extraordinari-
ly high proportion within the Blattulidae and cockroaches in
The hindwing pterostigma probably represents a plesiomor-
phic character within the Blattulidae. Figure 5.3 shows sharp-
ly curved CuP in the forewing, a character typical for Holo-
It is one of the oldest (Jurassic) species of the genus. It dif-
fers from other (even Cretaceous) representatives of the genus
in having sharply curved CuP in the forewing and dark macu-
la shifted more centrally.
Etymology: The species name refers to the pterostigma
present in the hindwing.
Character of preservation: Forewing 19 (body 1),
hindwing 10 (body 1).
Mesoblattinoidea Handlirsch, 1906 (= Blattoidea)
Mesoblattinidae Handlirsch, 1906
Breviblattina gen. nov.
Type species: B. minor sp. nov.
Composition: Type species.
Diagnosis: The genus with the smallest known species of
the family (wing length 7 mm or less). Pronotum very large,
as wide as half of wing length; forewing margins not parallel
(in general, small species have margins less parallel), venation
simplified, without terminal forks, Sc simple, hindwing R
Remarks: The genus differs from Hispanoblatta Mar-
tínes-Delclóz, 1993 in having non-parallel wing margins
(apomorphy depending on the diminished size of the insect).
JURASSIC/CRETACEOUS COCKROACH (INSECTA, BLATTARIA) FROM MONGOLIA 463
Fig. 5. 1 Elisama pterostigmata sp. nov. Forewing (4270/1916); 2 ?Blattula mongolica sp. nov. Pronotum (4270/1860); 3 Elisa-
ma pterostigmata sp. nov. Forewing (4270/1774); 4 ?Pronotum (4270/1760); 5 E. pterostigmata sp. nov. Forewing (4270/1899);
6 ?Pronotum (4270/1822); 7 E. pterostigmata sp. nov. Hindwing (4270/1848); 8 Shartegoblattina elongata sp. nov. Pronotum
Fig. 6. 1 Juramantis initialis Vranský, 2002 (4270/1842); 2 Elisama mongolica sp. nov. Forewing (1798); 34 Elisamoides
mantiformis sp. nov. Forewing (4270/1828); 5 Shartegoblattina elongata sp. nov. Forewing (4270/1818); 6 Shartegoblattina elon-
gata sp. nov. Forewing (4270/1846); 78 gen. et sp. indet. 7 Forewing (4270/1825). 8 Hindwing (4270/1795).
JURASSIC/CRETACEOUS COCKROACH (INSECTA, BLATTARIA) FROM MONGOLIA 465
Similarly non-parallel wing margins (plesiomorphic de-
rived from Caloblattinidae which are large in size) are also
known in Archimesoblatta altera (Vranský, 1997), Artito-
coblatta colominasi (Meunier, 1914), and partially in Me-
soblattina Geinitz, 1880.
The genera Archimesoblatta Vranský, 2003 and Artito-
coblatta colominasi differ from Breviblattina in rich vena-
tion, large size and terminal branchelets (all plesiomorphies).
Mesoblattina differs in having partially irregular venation
(According to Vranský 2000, the above-mentioned are all
representatives of the family Mesoblattinidae. Other Mesob-
lattinidae are placed in Caloblattinidae Vranský et Ansorge,
2000 or belong to other families.)
Etymology: The genus name is after brevis, the Latin for
short, and genus Blatta. Gender: feminine.
Breviblattina minor sp. nov. (Figs. 3.1, 7.4)
Holotype: PIN 4270/1843. Complete specimen.
Additional material: Forewings: PIN 4270/1801,
1868±c, 1906, 1912±.
Description: Wing length/width 5.47 mm/ca. 2.7 mm.
Sc branched, R expanded, with as much as 14 veins at margin,
M with 24 branches, Cu with 5 veins. Hindwing with richly
Etymology: The species name refers to the small size of
Taphonomy: Forewing 4 (clavus 1), body 1.
Mongolblatta gen. nov.
Type species: M. accurata sp. nov.
Composition: Type species.
Diagnosis: Forewing leatherous, venation regular, Sc
branched, RS differentiated, M and Cu richly branched, anal
veins simple, intercalaries punctuated in anal area.
Remarks: In general appearance the genus resembles
Hispanoblatta (and also Piniblattella Vranský, 1997 from
Blattellidae, which differs in having Sc simple), but the wing
is leatherous and anal intercalaries punctuated.
Artitocoblatta colominasi and Archimesoblatta altera
differ in having wing margins not parallel.
The new genus differs from Mesoblattina in having fully
The taxon appears transitional between Mesoblattinidae
and Blattellidae, it is placed in Mesoblattinidae because of the
branched hindwing M.
Etymology: The genus is named after Mongolia. Gender:
Mongolblatta accurata sp. nov. (Figs. 3.23, 7.13)
Holotype: PIN 4270/1849. Forewing.
Additional material: Forewings: PIN 4270/1796±,
1827±*, 1831±, 1833±, 1856±*, 1864c, 1877, 1898, 1900,
1908c. Hindwings: PIN 4270/11, 1761±, 1766±, 1806,
1838±, 1839±, 1857±.
Description: Forewing leatherous, anal intercalaries
with typical punctuation; hindwing with narrow remigium.
Forewing length/width: 10 mm/3.9 mm. Forewing Sc 2 bran-
ched; R with 13 or more branches (R branches and first
branch of M with dark colouration), in the preserved speci-
mens M and Cu with 5 branches each. Clavus ca. 5 mm long.
with 57, RS with 912 veins, M with 4 branch-
es. CuA secondarily branched.
Remarks: It is of interest that the fore- and hindwings
collected are almost equal in number, in spite on the fact that
forewings are leatherous and stiff, and yet many of them are
fragmented indicating a kind of damage. Normally the hind-
wings, being thin and so more vulnerable to mechanical dam-
age, experience considerable loss before and during the burial
Colouration is similar to Piniblattella vitimica (Vishniako-
va, 1964), but with coloured first branch of M.
Etymology: The species name (accurata is the Latin for
accurate) alludes to the intermediate size of the species.
Character of preservation: Forewing 11; hindwing 7.
Family, genus et species incertae sedis (Fig. 6.78)
Material: Forewings: PIN 4270/1808±, 1825; hindwing:
Description: Forewing length about 20 mm. Venation is
unique in possessing numerous comb-like connections of
veins to the apparent intercalaries. Veins more or less straight,
Systematic remarks: The poor state of preservation
disallows identification of the position of the fossil within the
current system. The species may well represent a Paleozoic or
even a new family. A similar but more sophisticated pattern of
the forewing venation is present in Paleozoic Phyloblattidae.
Nevertheless, the straight veins and modern character of the
branches indicate rather advanced taxon. The character of vena-
tion similar to Archimesoblatta altera (Vranský, 1997) (Me-
soblattinidae), except for comb-like interconnection of veins.
Character of preservation: Forewing 2; hindwing 1.
The cockroach diversity of some Upper Jurassic environ-
ments (Karatau as partly described by Vishniakova 1968:
1750 specimens with 25 species of 15 genera) is thought to be
comparable to the diversity of contemporary tropical forests
(Panfilov 1968) (nevertheless such comparison is not signifi-
cant because of taphonomical differences). The same diversity
is known for the Lower Cretaceous sites (Baissa described by
Vranský 1997, 1998, 1999a,b: 583 specimens with 24 spe-
cies of 9 genera in 2 diferrent assemblages; Bon Tsagaan ac-
cording to Vranský (2003): 651 specimens with 20 species
of 12 genera).
Diversity of the Shar-Teg assemblage (171 specimens with
8 or 9 species in 6 genera) is considerably lower, even richer
than other Lower Jurassic sites in Europe (120 specimens with
6 species in 4 genera in the Toarcian of Germany and En-
gland, as described by Vranský & Ansorge, in print).
Moderate in diversity, the Shar-Teg fauna is important for the
presence of modern family Mesoblattinidae and advanced Blat-
tulidae with two species and also Mantids (Fig. 6.1
Juramantis initialis described by Vranský (2002)). This char-
acterizes the Shar-Teg assemblage as Cretaceous in essence. In
addition to praying mantises, mesoblattinids and blattulids, the
modernization is represented by Elisamoides (probably Liberi-
blattinidae; the family otherwise known from the MiddleUp-
per Jurassic of Kazakhstan and Mongolia and Lower Creta-
ceous of Mongolia).
According to the presence of the most of the modern ele-
ments in the lowermost Cretaceous of Mongolia and Siberia
(where they occur for the first time), it is possible that the ad-
vanced Shar-Teg taxa (Juramantis sp., Elisama spp., Brevib-
lattina sp. etc.) are directly ancestral for them.
Fig. 7. 13 Mongolblatta accurata sp. nov. Forewings (4270/1856, 1864). Hindwing (4270/1857); 4 Breviblattina minor sp. nov.
(4270/1843); 56 Shartegoblattina elongata sp. nov. Hindwing (4270/1807, 1804).
The evolution of mantids specialized carnivores dur-
ing the latest Jurassic, represents an important phenomenon.
The rapid radiation of mantids which are known from all the
paleoentomologically well explored Early Cretaceous regions
of Laurasia and Gondwana (starting from the Turonian also in
North America (New Jersey)) is interesting.
In comparison with Shar-Teg, the less diverse European
Lower Jurassic sites as well as the diverse Upper Jurassic of
Karatau possess more ancient fauna with archaic lineages
In the Shar-Teg assemblage, the presence of the two blat-
tulid species complexes, represented by Blattula vidlickai and
JURASSIC/CRETACEOUS COCKROACH (INSECTA, BLATTARIA) FROM MONGOLIA 467
Mongolia with ca. 5:1, and Baissa with ca. 6:1. It is interest-
ing that in the Baissa warmer assemblage, a much lower ratio
(2:1, like in the Shar-Teg) is observed (Vranský 1998). In
the warmer water, the wings may be more easily disarticulated
from bodies because of decomposition, and thus the partuition
of bodies in warmer assemblages of Baissa is also low, like in
Shar-Teg. The role of decomposition is additionally con-
firmed by the high proportion of isolated forewing clavi. Thus
the preservation pattern observed in Shar-Teg indicates the
predominantly autochthonous preservation with the low me-
chanical damage, possibly in contrast to assemblages with a
higher rate of forewings collected. In these circumstances, the
possible explanation of the underrepresentation of roach bod-
ies is the activity of predators such as fish.
The high participation of teratologic vein fusion is also no-
table. It clearly indicate some stress factor in the paleoenvi-
ronment. However, Blattula vidlickai, the most common spe-
cies at the site is free of such fusions, possibly because a kind
of adaptation to the stress agents.
The cockroach assemblage of Shar-Teg shows moderate
taxonomic diversity and is composed of both archaic (pos-
sibly even Paleozoic) as well as advanced (transitional to con-
In spite of the Jurassic age and persistence of a possible
Paleozoic relict, the Shar-Teg assemblage is of Cretaceous
type in essence, as the presence of diverse Mesoblattinidae
and Blattulidae (2 abundant and additional infrequent species)
The assemblage probably presents a fragmentary orycto-
cenosis which resulted from active decomposition (and possi-
bly predation) rather than from transport by running water.
Some extraordinary ecological stressing factor was prob-
ably operating in the Shar-Teg biocenosis, resulting in malfor-
mations (in form of vein fusions) during the development of
Acknowledgments: I thank the Arthropoda Laboratory of the
Paleontological Institute of the Russian Academy of Sciences
for very kind help during the study. I thank Professor Alexandr
P. Rasnitsyn (PIN RAN, Moscow), Dr. Jozef Michalík (GlU
SAV), Dr. ¼ubomír Vidlièka, Dr. Duan itòan (IZ SAV,
Bratislava), Mgr. Peter Majchrák (FÚ SAV) and Mgr. Michal
Kubi (GlÚ SAV) for fruitful consultations, reviewing the
manuscript and for technical help. Partly supported by RFFI
010564741 and the Literárny Fond.
Gubin Yu.M. & Sinitza S.M. 1996: Shar-Teg: a unique Mesozoic
locality of Asia. In: Morales M. (Ed.): The continental Jurassic.
Mus. South Arizona Bull. 60, 311318.
Panfilov D.V. 1968: Ecological and landscape characteristic of the
Jurassic insect fauna of Karatau. P. 722. In: Rohdendorf B.B.
(Ed.): Jurassic Insects of Karatau. Nauka Press, Moscow, 1
252 (in Russian).
Table 1: Taphonomy of the site. Unidentifiable Blattaria are: PIN
4270/15, 27, 1757±*,1787, 1789=1790, 1791, 1810±, 1813, 1869,
1883, 1887, 1888, 1910, 1914, 1921 (forewings); PIN 4270/1812±,
1821±, 1884, 1890, 1892, 1896, 1915, 1923, 1924 (hindwings); PIN
4270/17, 1809±, 1830±, 1876, 1904 (bodies and/or pronota); PIN
4270/12, 13, 23, 1772, 1778, 1779, 1801, 1852±*, 1862, 1867,
1870*, 1882, 1886, 1879, 1895, 1917, 1918 (others); PIN 4270/
1824, 1836, 1880, 1911 are found to not belong to Blattaria; PIN
4270/19 is missing in the collection. I assume that at least one addi-
tional species is hidden within the unidentified specimens. If num-
bers are in parentheses, the specimens are included under numbers
Forewings Hindwings Bodies
gen. et sp. indet
Elisama pterostigmata is considerable (there is an additional,
but rare species of Blattula present, which could not play an
important role in the ecosystem). The ecological importance
of the presence of the two morphologically similar species is
not known, but is present in contemporary Ectobius Stephens,
1835 (two related species usually inhabit different levels of
the vegetation), and in the CretaceousTertiary Elisama
(closely related to Blattula). It is interesting that Elisama
which is recorded starting from the Jurassic, and during the
lowermost Cretaceous spread all over the globe (the close po-
sition of continents in the Berriasian did not form barriers) re-
placed a niche of Blattula in the Cretaceous, when the sister
species complex is formed exclusively by Elisama (Blattula is
The presence of dark macula in the forewing presents an in-
teresting phylogenetic problem. The macula is present in Eub-
lattula Handlirsch, 1939 possibly an ancestral taxon with
strong plesiomorphies within the family Blattulidae, and also
in Elisamoides, a genus probably belonging to another family
(Liberiblattinidae). Therefore the macula most probably repre-
sent a plesiomorphic character within the Blattulidae, which
was independently lost in Blattula, Rithma Giebel, 1856, Tara-
kanula Vranský, 2003, etc.
Derived characters such as branched A
in the hindwing
(probably supporting the wing in flight) and the sheath cover-
ing the ovipositor in a blattulid species (Blattula vidlickai) ap-
point to the developed evolutionary stages of some niches.
The taphonomy of the site (Table 1) is notable, since the
majority of the specimens are isolated wings with the very
low partuition of bodies. Most surprising is low ratio of the
preserved fore- and hindwings (ca. 2:1). The usual ratio is
much higher as in the Early Cretaceous of Bon Tsagaan in
Ponomarenko A.G. 1998: Paleoentomology of Mongolia. First Pale-
oentomological Conference. 30. Aug.4. Sept. 1998. Moscow,
Russia. Abstracts. Palaeontological Inst., Russian Acad. Sci.,
Vishniakova V.N. 1968: Mesozoic cockroaches with the external
ovipositor and features of their reproduction. P. 5586. In: Roh-
dendorf B.B. (Ed.): Jurassic Insects of Karatau. Nauka Press,
Moscow, 1252 (in Russian)
Vranský 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, 6779.
Vranský P. 1998: The Blattaria fauna of the Lower Cretaceous of
Baissa in Transbaikalian Siberia. Diploma Thesis, Comenius
University, Faculty of Natural Sciences, Department of Zoolo-
Vranský P. 1999a: Two new species of Blattaria (Insecta) from the
Lower Cretaceous of Asia, with comments on the origin and
phylogenetic position of the families Polyphagidae and Blattul-
idae. Entomol. Probl. 30, 2, 8591.
Vranský P. 1999b: Lower Cretaceous Blattaria. In: Proceedings of
the First Palaeoentomological Conference, Moscow 1998.
AMBA/AM/PFICM98/1.99, Bratislava, 167176.
Vranský P. 2000: Decreasing variability from the Carboniferous
to the Present! (Validated on Independent Lineages of Blattar-
ia). Paleontological J. Vol. 34, Suppl. 3, 374379.
Vranský P. 2002: Origin and the early evolution of mantises.
AMBA Projekty 6, 1, 116.
Vranský P. 2003: Unique assemblage of Dictyoptera (Insecta
Blattaria, Mantodea, Isoptera) from the Lower Cretaceous of
Bon Tsagaan Nuur in Mongolia. Entomol. Probl. 33, 12,
Vranský P. & Ansorge J. in print: Lower Toarcian Blattaria (Insec-
ta) of Germany and England. Greiswalder Geowischenshafli-