GEOLOGICA CARPATHICA, JUNE 2008, 59, 3, 269—272
A complete larva of a Mesozoic (Early Cenomanian)
cockroach (Insecta: Blattaria: Blattulidae) from the
Sisteron amber (Alpes de Haute Provence, SE France)
Geological Institute, Slovak Academy of Sciences, Dúbravská cesta 9, P.O. Box 106, 840 05 Bratislava, Slovak Republic;
Arthropoda Laboratory, Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya 123, 117868 Moscow,
Russian Federation; email@example.com
(Manuscript received October 31, 2007; accepted in revised form March 14, 2008)
Abstract: A complete second-instar male larva of Nula sis gen. et sp.n., belonging to the cockroach family Blattulidae
Vishniakova, 1982 is described from the Early Cenomanian amber of Sisteron in France. It reveals detailed and com-
plete 3D morphology, with important presence of the central, 3
ocellus, reduced in most adults and in all living
cockroaches and termites, but present in some mantises. The modern distribution of unspecialized sensorial system of
sensilla chaetica is also notable.
Key words: Early Cenomanian, SE France, evolution, fossil insects, Blattaria ( = Blattodea), Blattulidae, new genus,
new species, Cretaceous amber.
Blattulidae was the dominant Mesozoic family of cockroach-
es during the Cretaceous, particularly in warm and humid as-
semblages. It comprises about 300 described species and
ca. 10,000 collected specimens. In spite of their richness and
cosmopolitan distribution, all individuals can be categorized
within few uniform genera without subfamilial split.
The main morphology of the adults has been well known
since Vishniakova (1968) and descriptions include the de-
tailed morphology of the head and terminalia with female
ovipositor (Vishniakova 1968, 1982; Wang et al. 2007b), the
cercal, antennal, wing sensilla and venation variability (Vr-
šanský 2000, 2002, 2003, 2005a; Vršanský & Ansorge 2007;
Wang et al. 2007a). Nevertheless, the fine details and spatial
3D habitus and distribution of respective structures were un-
known, as the only immatures from mid-Cretaceous ambers
of Lebanon (a single individual) and New Jersey (two indi-
viduals) were not described (Vršanský 2004). This specimen
so far represents a unique opportunity for such study.
The distribution of fine sensilla (a feature not preserved in
imprint fossils) has a very special consideration.
Generally, the cockroaches from the Mesozoic ambers are
extraordinarily rare. Only 3 adults (Vršanský 2003; Anisyut-
kin & Gorochov 2008) were described from the Lebanon and
New Jersey ambers.
In SE France, amber was found in the marine blue marls
(Fig. 1a), providing dark red nodules representing fragment-
ed flows, 5—30 mm in diameter. Amber from the Salignac
near Sisteron revealed 16 inclusions 12 of which represent
undetermined flies (Diptera) (Perrichot et al. 2007). Macro-
siagon ebboi Perrichot et Nel et Néraudeau, 2004, perhaps
parasitic on wasps, represents one of the oldest records of
wedge-shaped beetles (Perrichot et al. 2004); unidentified
Phryssonotus Scudder, 1885 (Myriapoda, Diplopoda)
(Nguyen Duy-Jacquemin & Azar 2004) and a bug Ebboa ar-
eolata Perrichot, Nel, Guilbert et Néraudeau, 2006 (Het-
eroptera: Cimicomorpha), were also found in Archingeay
amber (Perrichot et al. 2006).
Order: Blattaria Latreille, 1810
Family: Blattulidae Vishniakova, 1982
Type species: Nula sis Vršanský sp.n., by monotypy.
D i f f e r e n t i a l d i a g n o s i s : The present genus may be
categorized within Blattulidae on the basis of its general habi-
tus, large almost globular head with characteristic long apical
antennal sensilla distributed in two main rows, and character-
istic long marginal pronotal setae. The slender habitus exclude
the categorization of the present fossil within the families
Caloblattinidae, Raphidiomimidae, Latiblattidae and Blattel-
lidae. Skokidae may be excluded by the absence of saltatorial
hind legs, and Mesoblattinidae and Umenocoleidae by their
more flattened head. Liberiblattinidae had wider antennal seg-
ments with a different distribution of shorter sensilla.
Nula differs from Elisama Giebel, 1856; Eublattula Hand-
lirsch, 1939; Svabula Vršanský, 2005a; Xonpepetla Cifuent-
es-Ruiz et Vršanský, 2006; Habroblattula Wang, Liang et
Ren, 2007 and Macaroblattula Wang, Ren et Liang, 2007 in
being slim. Tarakanula Vršanský, 2003 differs in having a
comparatively smaller head and in being more slender. Blat-
tula Handlirsch, 1906 has a comparatively narrow pronotum
and smaller head and is not recorded after the Tithonian –
thus its occurrence late in the Mesozoic is unlikely. Nula
also has an irregular carination of tibiae, small articulation of
the foreleg, and irregular ridge in hind femora (autapomor-
phies). Of a similar age is Kridla Vršanský, 2005b, known
from a single isolated hindwing coming from an unusual
find from a drilling core from East Siberia, which also, un-
like Nula, appears to belong to a more robust species. Vrtula
Vršanský, 2008 is a robust and significantly elongated aber-
D e s c r i p t i o n : Head large, with very large compound
eyes and three distinct discrete, plain ocelli, of which the
median ocellus is slightly smaller. Antenna fine and long,
with very distinct scape and pedicel, and with at least 44 seg-
ments in the second immature stage. Mandible comparative-
ly small, with sharp asymmetrical teeth. Pronotum of this
immature stage small, not concealing the head. Pro-, meso-
and metanotum with strong setae at margin. Legs cursorial,
fore and hindlegs long, mid-legs comparatively shorter. Fore
coxa short, femora long. Fore tarsus with numerous fine cha-
eta. All tibiae with numerous carination and with a lot of
chaeta. Hind femur with broken longitudinal ridge. Abdo-
men comparatively narrow, terminal plate with few chaeta.
R e m a r k s : The extreme conservativeness of the family
comprises immature stages. Additionally all the amber fos-
sils represent genera different from the genera known from
the imprint fossils, pointing to a different ecology of the am-
ber and imprint fossil source areas. Thus the erection of a
new genus appears safe.
D e r i v a t i o n o f n a m e : After nula (zero), sustaining
suffix – la for the representatives of this family. Alluding to
the incompleted life cycle of the present individual.
gen. et sp.n.
H o l o t y p e : SIS-17.2. Deposited in the amber collection
of the Earth History Department, National Museum of Natu-
ral History, Paris. A complete male immature.
T y p e l o c a l i t y : Salignac, near Sisteron, Department
Alpes de Haute Provence, SE France.
S t r a t i g r a p h i c h o r i z o n : Upper Cretaceous, Lower
Cenomanian fide Perrichot et al. (2004).
D e s c r i p t i o n : Head width 1.44 mm, eye width 0.29 mm.
Width of the antennal socket 0.17 mm. Scape and pedicel
length 0.13, 0.15 mm respectively; antennal width ca.
0.07 mm. Each segment with row of 5—7 sensilla. Segments
3—17 (22) with one row, more terminal segments with two
rows. Mandible 0.13 mm long, 0.38 mm wide (Left
0.17 mm; right 0.21 mm).
Pronotum ca. 1.2 mm wide, with four marginal seta, me-
sonotum little wider (ca. 1.4 mm) also with four marginal
seta long at least 0.19 mm; metanotum long, with at least 6
shorter marginal seta.
Articulation of fore leg very short, with coxa length
0.3 mm; trochanter small (0.19/0.12 mm); femur, tibia and
tarsus comparatively long 0.75/0.17 mm; 0.54/0.12 mm;
0.75 mm. Femur with doubled ridge. Pretarsus with two
symmetrical claws. Three terminal, four median (two rows)
and a single basal long chaeta are present. Arolium round.
Mid leg robust, trochanter large (0.29/0.13 mm); femur
wide (1.04/0.35 mm) with at least five chaeta on distal, 15
on proximal, 9 + 9 in two rows of median side (all ventral);
tibia (0.87/0.12 mm) with numerous chaeta at margin, and
with sporadic series of strong spurs long up to 0.29 mm.
Hind legs comparatively fine, with long femur (1.44/
0.42 mm) bearing a strong, irregular longitudinal ridge, with
6 chaeta. Anterior margin with 8, posterior with 5 chaeta.
Tibia wide (0.17 mm), with spurs long up to 0.37 mm. Seven
distinct sterna present.
R e m a r k s : One chaeta on the mid-tibia apparently origi-
nates deeply under the cuticle.
Antennae are broken at segments 43 and 44 respectively,
so it is not possible to assume the total number of segments
and their total length.
D e r i v a t i o n o f n a m e : sis is after Latin “if you like it”,
also a suffix and shortened locality Sisteron.
The head unconcealed by the pronotum is a common fea-
ture of the first two instar stages of primitive living cock-
roaches (of Mesozoic origin), and since there are no
significant differences in morphology among Blattulidae and
these living cockroaches, there are no contraindications for
such assignment of the present species. Thus, if the second
Fig. 1a. Geological scheme of the locality (modified after Haug et
al. 1964 and Perrichot et al. 2004).
INSECTA: BLATTARIA: BLATTULIDAE FROM THE CENOMANIAN SISTERON AMBER (SE FRANCE)
Fig. 1b. Four different projections of Nula sis Vršanský, sp.n. (all ventral views). Holotype SIS-17.2. (deposited in the amber collection
from the Earth History Department, National Museum of Natural History, Paris). A complete larva of second instar. Salignac, near Sister-
on, Alpes de Haute Provence, SE France. Early Cenomanian Late Cretaceous. b1 – total ventral view; b2 – detail of head in transmitting
light; b3 – sketch abstraction; b4 – reconstruction. Arrow shows a spine originating deep below the cuticle. Total length of the individual
(without legs): 3.5 mm.
instar is 3.5 mm long, applying the Dyar’s rule of succeeding
instars sizes multiplied by 1.4
×, the first instar should be
2.5 mm long, which are similar sizes of first two immatures
of Blattula brevicaudata Vishniakova, 1968 from the Late
Jurassic of Karatau in Kazakhstan. B. brevicaudata adults
have a wingspan of ca. 7.5 mm. If we continue, the third in-
star would be 4.9 mm long (around 5 mm in B. brevicauda-
ta), the fourth 6.9 mm and the fifth 9.7 mm. Thus, the
number of instars can be predicted to be 4—6 (similar to the
studied but undescribed Blattulidae from the sedimentary
record) and the size of the adult 6—10 mm, with the most
probable wing length about 7 mm.
The presence of the central ocellus is of special consider-
ation, since this structure has been discovered in cockroach-
es of the family Caloblattinidae Vršanský et Ansorge in
Vršanský (2000) and considered to be plesiomorphic only
recently (Vršanský 2008a). It was though to have reduced in
the Blattulidae, since in the rich Late Jurassic (Vishniakova
1968) and Early Cretaceous record from China it has not
been observed (Wang et al. 2007a,b). Nevertheless, the cen-
tral ocellus is present in the present larva and also in at least
one derived species (Anisyutkin & Gorochov 2008) and thus
this structure must be reduced during the ontogenetic devel-
opment of most Blattulidae. Ocelli of Nula were probably
plain, as in the adult of Ocelloblattula ponomarenkoi Ani-
syutkin & Gorochov, 2008.
The precise level of recognition of distribution of numer-
ous sensilla indicates that the senses of the Mesozoic cock-
roaches were modern, virtually indistinguishable from the
living species. The sensilla were certainly different, unspe-
cialized (Vršanský et al. 2001), but their number and distri-
bution was identical with modern ones.
Acknowledgments: I thank Vincent Perrichot, Didier
Néraudeau (Univ. Rennes 1), Dany Azar (Lebanese Univer-
sity and NMNH Paris), Leonid Anisyutkin (Zoological Insti-
tute, St. Petersbourg) and Jozef Michalík (Geological
Institute, Bratislava) for extensive help during the study and
for revision of the manuscript. I also thank ubka Puškelová
for technical support. Supported by VEGA 6002, MVTS;
Literary Fund. This paper is a contribution to the French
ANR Project AMBRACE (No. BLAN07-1-184190).
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.
Haug E., Kilian W., Bertrand L., Zürcher P., Martin D., Gubler Y.,
De l’Apparent A.F., Goguel J., Lanteaume M. & Lemoine M.
1964: Carte géologique au 1 : 80,000e: Digne, n° 212 (3
Latreille P.A. 1810: Considérations générales sur l’ordre naturel des
animaux composant les classes des Crustacés, des Arachnides
et des Insectes avec un tableau méthodique de leurs genres dis-
posés en familles. Schoell, Paris, 1—444.
Nguyen Duy-Jacquemin M. & Azar D. 2004: The oldest records of
Polyxenida (Myriapoda, Diplopoda): new discoveries from the
Cretaceous ambers of Lebanon and France. Geodiversitas 26,
Perrichot V., Nel A. & Néraudeau D. 2004: Two new wedge-shaped
beetles in Albo-Cenomanian ambers of France (Coleoptera: Rip-
iphoridae: Ripiphorinae). Eur. J. Entomol. 101, 4, 583—589.
Perrichot V., Nel A., Guilbert E. & Néraudeau D. 2006: Fossil Tin-
goidea (Heteroptera: Cimicomorpha) from French Cretaceous
amber, including Tingidae and a new family, Ebboidae. Zoot-
axa 1203, 57—68.
Perrichot V., Néraudeau D., Nel A. & De Ploëg G. 2007: A reas-
sessment of the Cretaceous amber deposits from France and
their palaeontological significance. African Invertebrates 48,
Vishniakova V.N. 1968: Mesozoic cockroaches with an external
ovipositor and pecularities of their reproduction (Blattodea).
55—86. In: Rohdendorf B.B. (Ed.): Jurassic Insects of Karatau.
Nauka, Moscow 1—252 (in Russian).
Vishniakova V.N. 1982: Jurassic cockroaches of the family Blattul-
idae fam. nov. (Insecta: Blattida). Paleontol. J. 2, 69—79 (in
Vršanský P. 2000: Decreasing variability-from the Carboniferous to
the Present! (Validated on independent lineages of Blattaria).
Paleontol. J. 34, Suppl. 3, 374—379.
Vršanský P. 2002: Origin and the Early Evolution of Mantises.
Amba Projekty 6, 1, 1—16.
Vršanský P. 2003: Unique assemblage of Dictyoptera (Insecta-
Blattaria, Mantodea, Isoptera, Mantodea) from the Lower Cre-
taceous of Bon Tsagaan Nuur in Mongolia. Entomol. Probl. 33
Vršanský P. 2004: Cretaceous Gondwanian Cockroaches (Insecta,
Blattaria). Entomol. Probl. 34, 1—2, 49—54.
Vršanský P. 2005a: Lower Cretaceous cockroaches and mantids
(Insecta: Blattaria, Mantodea) from the Sharin-Gol in Mongo-
lia. Entomol. Probl. 35, 2, 163—167.
Vršanský P. 2005b: Insect in a drilling core—cockroach Kridla stas-
tia sp.nov. from the Verkhne-Bureinskaya Depression in East-
ern Russia. Entomol. Probl. 35, 2, 115—116.
Vršanský P. 2008a: Central ocellus of extinct cockroaches (Blattaria:
Caloblattinidae). Zootaxa (in print).
Vršanský P. 2008b: New blattarians and a review of dictyopteran
assemblages from the Lower Cretaceous of Mongolia. Acta
Palaeontol. Pol. 53, 1, 129—136.
Vršanský P. & Ansorge J. 2007: Lower Jurassic cockroaches (In-
secta: Blattaria) from Germany and England. African Inverte-
brates 48, 1, 103—126.
Vršanský P., Quicke D.L.J., Rasnitsyn A.P., Basibuyuk H., Ross
A., Fitton M. & Vidlička . 2001: The oldest fossil insect sen-
silla. Amba Projekty 5, 1, 1—8.
Wang T., Liang J.-H. & Ren D. 2007a: Variability of Habroblattula
drepanoides gen. et. sp. nov. (Insecta: Blattaria: Blattulidae)
from the Yixian Formation in Liaoning, China. Zootaxa 1443,
Wang T., Liang J.-H., Ren D. & Shi C. 2007b: New Mesozoic cock-
roaches (Blattaria: Blattulidae) from Jehol Biota of western Li-
aoning in China. Ann. Zool. 57, 3, 483—495.