GeolCarp_Vol59_No3_269

GEOLOGICA CARPATHICA, JUNE 2008, 59, 3, 269—272

www.geologicacarpathica.sk

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A complete larva of a Mesozoic (Early Cenomanian)
cockroach (Insecta: Blattaria: Blattulidae) from the

Sisteron amber (Alpes de Haute Provence, SE France)

PETER VRŠANSKÝ

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

geolvrsa@savba.sk

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

Russian Federation; lab@palaeoentomolog.ru

(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.

SHORT COMMUNICATION

Introduction

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).

Systematics

Order: Blattaria Latreille, 1810

Family: Blattulidae Vishniakova, 1982

Nula gen.n.

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.

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VRŠANSKÝ

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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-
rant taxon.

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.

Nula sis

gen. et sp.n.

(Fig. 1b)

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.

Discussion

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).

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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).

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