GEOLOGICA CARPATHICA
, OCTOBER 2016, 67, 5, 433 – 450
doi: 10.1515/geoca-2016-0027
www.geologicacarpathica.com
Taxonomic diversity of cockroach assemblages
(Blattaria, Insecta) of the Aptian Crato Formation
(Cretaceous, NE Brazil)
SHIH-WEI LEE
National Museum of Marine Science & Technology, No. 367 Pei-Ning Road, Zhongzheng District, Keelung 20248, Taiwan;
leesw@mail.nmmst.gov.tw
(Manuscript received November 11, 2015; accepted in revised form September 22, 2016)
Abstract: A comprehensive revision of 981 specimens of fossil cockroaches from the Lower Cretaceous laminated
limestones of the Crato Formation of Northeast Brazil shows that they belong to eleven taxa, including Piniblattella
limai, P. magna sp. n., Perlucipecta santanensis. sp. n., Raptoblatta waddingtonae; Ocelloblattula santanensis sp. n.,
Elisama brevis (= E. americana, syn.n.), E. hindwingnii sp. n., Ponopterix axelrodi (= P. maxima syn.n.), Umenopterix
burkhardi comb. n., and Cratovitisma oldreadi (Umenocoleidae = Cratovitismidae syn.n. = Ponopterixidae syn.n.).
The family Ectobiidae is numerically most abundant in the assemblage of cockroaches of the Crato Formation
(83 % of cockroaches), followed by Blattulidae (13 %) and Umenocoleidae (4 %). 79.2 % of specimens are complete
and fully articulated. Members of the family Alienopteridae are probably also present. Representatives of a relatively
common Mesozoic superfamily Caloblattinoidea are missing. With the exception of the endemic genera Cratovitisma
and Raptoblatta and the exclusively Gondwanan genus Ocelloblattula, all other genera were cosmopolitan.
Taxonomic richness of cockroaches of the Crato Formation is thus rather low, and consists of geologically long-
ranging and geographically-widespread genera, genera restricted to Gondwana, and short-ranging endemic genera
found in the Crato Formation only.
Keywords: Fossil insects, new species, Blattodea, variability, taphonomy, Mesozoic, Early Cretaceous, Gondwana.
Introduction
Cockroaches, mantises and termites represent the tradi-
tionally monophyletic Dictyoptera (Hennig 1981; Grimaldi
& Engel 2005; Djernaes et al. 2012, 2015; Bai et al. 2016),
originating in the Late Carboniferous (Zhang et al. 2012) and
being numerically abundant during the Palaeozoic (Schneider
1977; 1978a, b; 1980a, b: 1983; 1984; Schneider & Werne-
burg 1993). In the Mesozoic, both termites (Isoptera) and
mantises (Mantodea) evolved by the pattern (Mantodea +
(“Blattaria”+ Isoptera)) (Grimaldi 2007) or independently
originated from the cockroach family Liberiblattinidae
(Vršanský 2002; Vršanský & Aristov 2014; Vršanský &
Bechly 2015). Cockroaches possessed diverse life habits
(Barna 2014) and played an important ecological role as
decomposers in the present and past (Zherikhin 1993;
Vršanský et al. 2013).
Exceptionally-preserved insects dominate both in abun-
dance and taxonomic diversity in the fossils of the Crato
Formation. More than 350 species representing over 20
insect orders have been described, with terrestrial adult
orthopterans, hemipterans, ephemeropterans and odonatas
(Menon & Martill 2007; Bechly 2007 b; Heads & Leuzinger
2011; Barling et al. 2015). Three mantises and six termite
species have been described so far (Bechly 2007 c; Hörnig et
al. 2013; Lee 2014), but about a quarter of the arthropod
specimens from the Crato Formation are cockroaches
(Bechly 1998; Menon & Martill 2007; Dittman et al. 2015;
n = 3,651), with only dozens of species described. The aim of
this study is the revision of cockroaches from the Crato For-
mation, including older descriptions.
Geological setting
The Crato Formation is situated in the north central part of
the Chapada do Araripe (Fig. 1), a large plateau in north-
eastern Brazil (Martill & Bechly 2007). The formation is an
up to 60 m-thick, mixed carbonate-sicliclastic sequence
dominated in the middle part by laminated micritic lime-
stones, interbedded at the base with claystones, siltstones,
and sandstones (Martill & Heimhofer 2007; Barling et al.
2015). The limestones from this location contain gymno-
sperms and angiosperms, invertebrate fauna with Chilopoda,
Arachnida, Crustacea and Insecta, and vertebrate fauna
including fish, anurans, turtles, lizards, crocodiles, pterosaurs
and birds (Menon & Martill 2007). These fossils occur in the
lowermost member of the Crato Formation, namely in the
Nova Olinda Member, with an age of ~120 million years,
which corresponds to the Aptian stage (Martill 2007; Martill
& Heimhofer 2007). Chapada do Araripe was located at
10° to 15° S within the tropics during the Early Cretaceous
(Martill 2007). The depositional environment of the Nova
Olinda Member was a lake/lagoon water body with water-
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column stratification and bottom-water anoxia
allowing exceptional fossilization, with still unclear sali nity
conditions. Hypotheses range between a freshwater
and a brackish lake up to a hypersaline lagoon (Martill &
Bechly 2007). Among the fossils of the Crato Formation,
insects dominate. According to a large-scale investigation
(Bechly 1998; Menon & Martill 2007; Dittman et al. 2015;
n = 3651), about 26 % of arthropod specimens are cock-
roaches. However, only a dozen cockroach species were
reported in the Crato Formation, including some unnamed
species and species based on a few specimens (
e.g.,
Crato
vitisma oldreadi Bechly, 2007; “Ponopterix” burkhardi
Nel, Prokop et Kirejtshuk, 2014;
Raptoblatta waddingtonae
Dittman, Hörnig, Haug et Haug, 2015
). Thousands of
specimens belong to three dominant species, Ponopterix
axelrodi Vršanský
et Grimaldi, 1999a, Elisama brevis
(= E. americana) Mendes, 2000, and Piniblattella limai
(Pinto et Purper, 1986; Pinto, 1989) (Vršanský 2004;
Bechly 2007a). Using the taphonomic categories of speci-
men preservation by Martins-Neto & Gallego (2006), about
79.2 % of specimens described here belong to Stage I,
18.8 % belong to Stage II, and 2 % belong to Stage IV. These
pro portions are in contrast to the results of Martins-Neto
(2006), Martins-Neto & Gallego (2006), and Martins-Neto
& Tassi (2009) who found that different genera show
various preservation states on the basis of large scale
field data.
Material and methods
The drawings were made from dry specimens directly
from the fossil surface using a camera lucida attached to
a Wild M5 and a Leica MZ125 stereo microscope and scan-
ning with a Canon CanoScan 4400F flatbed scanner. Fossils
were photographed with a Leica DFC490 digital macro
camera on a Leica Z16-Apo microscope with Synchroscopy
Auto-Montage™ software. All figures have been sub-
sequently edited with the Adobe Photoshop™ CS3 imaging
software.
Differences between the right and left side of the wings
within the same individual were examined. The nomen-
clature for fore- and hind wings is based on Kukalová-Peck
(1991) and Carpenter (1992). Abbreviations used: C — costa,
CA — costa anterior, CP — costa posterior, ScA — subcosta
anterior, ScP — subcosta posterior, RA — radius anterior,
RP(= RS) — radius posterior, MA — media anterior, MP —
media posterior, CuA — cubitus anterior, CuP — cubitus
posterior, AA — anal anterior, AP — anal posterior, JA —
jugal anterior, JP — jugal posterior and the nomenclature
used for cockroaches, RA + RP + MA would be R + RS in the
system of Comstock & Needham (1898).
The total number of species is interpreted from the two
collections, including (1) MSF, Fossils Worldwide, Annesuse
Raquet-Schwickert Hebelstrasse 4 D-67734 Sulzbachtal,
Germany, with 933 specimens, and (2) SMNS, Staatliches
Naturkunde Museum Stuttgart, Germany, with 48 specimens
that were used in the systematic analysis.
Results
Systematic palaeontology
Order: Blattaria Latreille, 1810
= Blattodea Brunner von Wattenwil, 1882
Superfamily: Umenocoleoidea Chen et T’an, 1973
Family: Umenocoleidae Chen et T’an, 1973
2007 Cratovitismidae syn.n. — Bechly, 239–249.; syno nymization
is based on the revision of the Cratovitisma below.
2014 Ponopterixidae syn.n. = derived as Ponopterixiidae — Nel,
Prokop et Kirejtshuk, 2014 (= Vitisminae Vršanský et Ansorge,
2001 eventual jun. syn., in which case derived as Vitismidae
Vršanský et Ansorge, 2001 according to ICZN principle of
coordination for family-group names); syno nymization is
based on the description of Umenopterix (see below) and its
affinities to Umenocoleus.
Monophyletic beetle-like lineage with families Umeno-
coleidae and Alienopteridae Bai et al., 2016, stemming from
Vitisminae. The representatives of this family are known
since the earliest Cretaceous of Siberia (Vršanský 2003a).
Emended description (after Vršanský (2003a)): Small
(mostly under 20 mm), beetle-like cockroaches with head
Fig. 1. Location of the Crato Formation (7°19’00” S and 39°40’40” W).
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which can be derived and globular (eventually ortho-
gnathous); antenna with wide segments and very long setae
in transversal rows; pronotum with paranota tending to
reduce; with legs eventually short and strong. The most
charac teristic are heavily sclerotized forewings with reduced
venation and with the presence of cup-like cells. Hind wing
venation generally of early polyphagoid form (that is simpli-
fied venation with CuA without tertiary branches, CuP
simple, A1 curved, with short branches), but with reduced
venation of R (with few branches — possibly with
pterostigma), M branched. Cerci with numerous long hairs;
females with short external ovipositor. Species are most
probably diurnal with similar flight activity of both males
and females because of the same body size, with sexual
dimorphism not much pronounced.
Genus: Cratovitisma Bechly, 2007
Cratovitisma oldreadi Bechly, 2007 (Fig. 2)
Diagnosis (improved after Bechly 2007a): Body length
of 6.9 mm, forewing length 5.3 mm. As in Vitisma, the fore-
wings are broader than in Ponopterix (max. 2 mm wide),
strongly sclerotized. The vein CuP seems to be strongly
curved and delimits a typical cockroach anal field in the fore-
wings. The hind wing venation is similar to that of Vitisma,
thus more dense and round than in
Ponopterix, and with
pterostigma. The transverse head seems to place this new
genus and species closer to standard Umenocoleidae than
Vitisma (which is also now proved to belong to Umeno-
coleidae based on the amber record).
Emended description: The head is transverse as in Umeno-
coleidae (different from Vitisma type) and the antennae are
also similar to Umenocoleidae, thus shorter than the body.
The pronotum is flat and disk-like as in Jantaropterix
Vršanský et Grimaldi in Vršanský (2003a), Vitisma and true
cockroaches, very different from Ponopterix and Umeno
coleus. Forewing, Sc bifurcated, Radius with eight branched,
M with five to six branched, CuA similar with Genus
Ponopterix to have a single terminal fork, CuP strongly
curved and delimits a typical cockroach’s anal field. Hind
wing, Sc alone and less than half of whole wing length, RA
with six branched, RS with a total of four branches and simi-
lar to Ponopterix axelrodi, MP alone and CuA with probably
seven branched (or six CuA plus one CuP). The ovipositor
from this single specimen is 0.6 mm long (about 7.89 % of
body length), is tendentious shorter than that in Umeno-
coleoidae species in the Crato Formation, also clearly shorter
than that in Elisama brevis (= E. americana) (Blattulidae).
Genus: Ponopterix Vršanský et Grimaldi in Vršanský
(1999a)
Ponopterix axelrodi Vršanský et Grimaldi in Vršanský
(1999a); 2003a (Fig. 3)
1999 Ponopterix axelrodi. — Vršanský et Grimaldi, 167–176.
2007 Ponopterix maxima. — Bechly, 239–249 syn. n.; synonymiza-
tion is based on the revision below.
Emended description: Body 5.0 to 8.0 mm long. Head
spherical, 1.8 to 3.0 mm wide. Antenna about the same length
with body, 7.1 to 8.3 mm long, the first three segments elon-
gate, the others very short. Forewing 4.9 to 9.1 mm, usually
about 6.5 mm long, 1.7 to 2.8 mm wide. Radius rich (up to
10 branches), M and Cu variable but usually with single ter-
minal fork. Anal veins numerous. Hind wing pterostigma
present, 5.2 to 6.3 mm long, is almost the same length as the
forewing. Sc alone, shorter than half of the whole wing
length, RA with four branched, RS with two stem branches
and that is two to four terminal branches totally, MP with two
to three branched, CuA with six branched, CuP alone. Legs
cursorial, heavily mounted with spurs. All tarsi 5-segmented.
Female with external ovipositor at least 0.67 to 0.9 mm long,
is approximately 10 % – 12.12 % of body length. Male with
distinct styli and with phallomere of a primitive type. Cerci
placed closer to each other than in the female. Cercus 0.7 to
1.5mm long, with 11–13 segments.
Description of other specimens: SMNS 66326 (Fig. 3D)
— Male. Hind wing well preserved. SMNS 66329 — Female.
Ventral view. Compound eyes well preserved. Abdominal
segments clearly. SMNS 66334 (Fig. 3K) — Probably male.
Hind wing only partly visible. SMNS 66335 — Female.
Ventral view. Abdomen swelled. SMNS 66336 (Fig. 3L) —
Male. Forewing venation visible, varies between right and
left sides. Hind wing RA with four-branched and RS with
two stem-branches. SMNS 66338 (Fig. 3N) — Female.
Fig. 2. Cratovitisma oldreadi Bechly, 2007, holotype; SMNS
66000-127. A — Habitus. B — Hind wing. C — Right and left
forewings.
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Fig. 3. Ponopterix axelrodi Vršanský et Grimaldi, 1999 (= Ponopterix maxima Bechly, 2007). A — SMNS 66323. B — 66324.
C — 66325. D — 66326. E — 66327. F — 66328. G — 66329. H — 66331. I — 66332. J — 66337. K — 66334. L — 66336. M — 67574.
N — 66338.
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Hind wing preserved completely. Cerci and ovipositor clearly
visible.
Remarks: Ponopterix axelrodi differs from Ponopterix
maxima Bechly, 2007 by the smaller body size and by hind
wing with slightly different RP pattern (in P. maxima RP with
sequential branches one after another and in P. axelrodi RP
with two stem-branches and each one may have more termi-
nal branches (although this variability is not significant
because it can also be found in the same species). However,
on the basis of the similarity of fore- and hind wings and on
the basis of a gradual transition in body size between
P. axelrodi and P. maxima, (Table 1), I conclude that these
two species belong to a single species.
Emended description of specimens categorized to
Ponopterix maxima Bechly, 2007: Body 7.5 to 12.5 mm
long. Antenna at least 6 mm long and about half of body
length. Head broader than in P. axelrodi, especially between
the compound eyes; pronotum comparatively larger and
broader and saddle-shaped. Forewing 6.4 to 10.5 mm long
and 1.9 to 2.7 mm wide, Sc alone, Radius with four to eight
branched, usually four to six (seven- and eight branched only
by one specimen), M and Cu variable but usually with single
terminal fork, anal veins numerous, generally all venations
more or less straight. Hind wing about the same length as the
forewing, Sc alone and less than half of the whole wing
length, RA with two to four branched, RP with three to five
branched, MP with two branched, CuA with five branched
and CuP alone. Leg has five-segments tarsi with claws,
first segment elongated, 2
nd
,
3
rd
and 5
th
segments about the
same length and the 4
th
segment extremely short. Female
with external ovipositor at least 0.8–1.3 mm long, about
7.84 % –12.26 % of body length. Cercus in roach form, 12 to
16 segments.
Description of other specimens: SMNS 66323 (Fig. 3A)
— Male. Length of the antenna is about ½ of the body length.
Large compound eyes. Hind wing only partly visible. SMNS
66324 (Fig. 3B) — Lateral side. Femur of three legs well
preserved. SMNS 66325 (Fig. 3C) — Male. Venation of both
forewings clearly preserved. Hind wing with pterostigma and
large part of venation identifiable. SMNS 66327 (Fig. 3E)
— Female. Legs well preserved, tarsus with five segments,
first segment elongated, 2
nd
, 3
rd
and 5
th
segment about the
same length, 4
th
segment extremely short. Abdomen swelled.
SMNS 66328 (Fig. 3F) — Female. The venations of both
forewings clearly visible. Ovipositor and cercus well pre-
served. SMNS 66330 — Female, ventral side. Legs well pre-
served especially in femur and coxa part. SMNS 66331
(Fig. 3H) — Female. Forewing heavily sclerotized, venation
reduced. Hind wing partly preserved. SMNS
66332 (Fig. 3I)
— holotype. Compound eyes well preserved. The venations
of forewings clearly visible. SMNS 66333 — Female. Head
broader than the other specimens. Fore and hind wings only
partly preserved. SMNS 66337 (Fig. 3J) — Male. Venation of
forewings clearly visible. Hind leg well preserved, including
coxa part. 66562 — Female. Forewing clearly. Abdomen
swelled. Cercus and ovipositor well preserved. SMNS 67574
(Fig. 3M) — Probably male, lateral side. Mouthpart well
preserved, maxillary palps and labia palps visible.
Genus: Umenopterix gen. n.
Type species: Ponopterix burkhardi Nel, Prokop et
Kirejtshuk, 2014
Diagnosis: The pronotum fully developed, standard, trans-
versal with fully developed paranotalia. CuP strongly sig-
moidally curved.
Description: as for species (see Nel et al. 2014).
Derivation of name: Named after family Umenocoleidae
and Genus: Ponopterix.
Remarks: The genus apparently represents an early off-
shoot of the family, descending from taxa close to the type
genus Umenocoleus, which also has unique sigmoid curvation
of the clavus (including U. nervosus, see Zhang 1997). This
character was the sole argument against the categorization of
Umenocoleus within cockroaches, but the present genus fully
valid categorization (of the whole family) within Blattaria.
The above inference is also supported by the “standard” mor-
phology of round, transversal pronotum, with fully developed
paranotalia — a character lost in all “Petropterix” lineages, but
retained in the second lineage represented by Vitisma and to
some extent by Jantaropterix. Therefore, the present species
must be placed in a new genus, whilst any relation with
Ponopterix is unlikely, but these characters unequivocally sup-
port the assignment of this species (and also Umenocoleus as
the type of the Umenocoleidae) within cockroaches.
Superfamily: Corydioidea Saussure, 1864
(Brues & Malander, 1932)
(= Polyphagoidea Walker, 1868) (Princis, 1960)
Family: Blattulidae Vishniakova, 1982
= Araripeblattidae Mendes, 2000 syn.n. (synonymization based on
the paragraph below)
Type genus: Elisama Giebel, 1856.
Remarks: The family originated in the Late Triassic.
It represents a dominant cockroach family at many sites
ranging from the Early Jurassic to the Early Cretaceous for
more than 100-million years (Wang et al. 2007a, b).
Genus: Elisama Giebel, 1856
= Ctenoblattina Scudder, 1886 (Vršanský 2005a–c)
= Araripeblatta Mendes, 2000 syn.n.
Type species. Blattidium molossus Westwood, 1854
Elisama brevis Mendes, 2000 comb.n. (Fig. 4)
2000 Araripeblatta brevis — Mendes, 23–35.
2000 Blattopteris beckeri — Mendes = Araripeblatta beckeri Mar-
tins-Neto et al., 2010
2010
Araripeblatta cesae — Martins-Neto et al., 2010
2002 Elisama americana — Vršanský, 1–16.
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Emended discussion (after Martins-Neto et al. 2010): the
family Araripeblattidae differs from Blattulidae Vischnia-
kova, 1982, the closest family, by contact points of first
RA branch with the anterior wing border (RA1-ending),
MA2-origin, and contact of CuP with the posterior wing
border (CuP-ending) are arranged in one line slightly trans-
verse to the wing long axis. In Blattulidae, the point of
MA2-origin is situated slightly backward of the line
RA1-ending and CuP-ending (see also Martins-Neto et al.
2005). However, this main difference does not apply to this
study. Apart from the lines from RA1-ending, MA2-origin
and CuP-ending of Araripeblatta brevis variant, both intra-
specific (even between right and left forewings in the same
specimen) and inter specific (Fig. 5A–K), there are no signi-
ficant differences between Araripeblatta brevis Mendes,
2000 and Blattulidae, that is, Elisama tsaganica Vršanský,
2002, E. parallela Vršanský, 2003, E. extenuata Ren, 1995,
E. cuboides Wang, Ren et Liang, 2007 and Habroblattula
drepanoides Wang, Liang et Ren, 2007 (Fig. 5L–O). Araripe
blatta is here consi dered a junior synonym of Elisama and
belongs to family Blattulidae. Indeed this RA1-MA2-CuP
lines are clearly different but between Blattulidae and
Piniblattella limai (=Mesoblattina limai) (Ectobiidae)
(Fig. 5P–S).
Differential description: Elisama brevis differs from Pini
blattella limai by smaller body size. It is similar to genus
Ocelloblattula Anisyutkin et Gorochov, 2008 (Blattulidae)
and other primary Upper Jurassic family Caloblattinidae
Vršanský et Ansorge, 2001 by head with three ocelli residing
in the forehead between the compound eyes (Anisyutkin &
Gorochov 2008; Vršanský 2008a). The ovipositor of E. brevis
(about 10.13–16.67 % of body length) is tendentious con-
vexer than that in Umenocoleoideas both species in the Crato
Formation (by
P. axelrodi: 10 –12.12 % and by P. maxima:
7.84 –12.26 %), and still clearly longer than that in Crato
vistima sp. (SMNS 66000-127 by 7.89 %).
Emended diagnosis and description: Small-sized
roaches. Antenna about the same length as body. Body length
between 7.9–9.5 mm. Head wide and clearly longer than the
fore margin of pronotum. Pronotum is about 1.5 times wide
as the head, approximately of round shape, except in the pos-
terior part where it is sharp gradually. Forewing 8.0–10.2 mm
long and 2.4–3 mm wide. Sometimes slightly sclerotized.
It is possible that coloured band pattern reduced accompanies
with the degree of sclerotization. Wing fore margin arcuate
basally, almost straight further. Costa area as wide as sub-
costa area. Sc very short, not reaching the apical third of
a wing. RA+RS with total of 13–16 branches, instead of ten.
M with three–four branches. CuA with four–six branches,
rarely seven (SMNS 67572 right side, Fig. 4J), CuP extremely
curved and Anal veins reach the hind margin. Hind wing
about the same length as the forewing and 8.5–9.8 mm long.
Sometimes in anterior margin, that is RA area, slightly scle-
rotized, or that is pterostigma. Sc alone, reaching to a little
less than ½ of the anterior margin of the whole wing length.
RA with four–five branches, instead of three, RS with six–
seven branches. MP with three branches. CuA in principle
with six branches, rarely seven (SMNS 67566 left side,
Fig. 4F), CuP should be alone and probably reaching the end
of the wing. Fore and hind wings show very low morpho-
logical differences (Tables 2, 3). Legs presented long and
strong spines. Tarsus normally has all five segments. Abdomen
in female is tendentious more rounded than in male, in male
more elongate. Cerci in roaches form, 12–18 segments.
Ovipositor convex and sharp, with sheath, 0.8–1.5 mm long
(about 10.13–16.67 % of body length). Terminalia with
unsegmented styles.
Description of other specimens: SMNS 66000-125
(Fig. 4A) — Male. Eye big, place by the side of the head and
area in-between is large. Pronotum is about 1.5 times as wide
as head, nearly rounded, except in the hind part sharp gra-
dually. SMNS 67563 (Fig. 4C) — Female. Forewings sclero-
tized. Hind leg has long and strong spines. Ovipositor present.
SMNS
67564 (Fig. 4G) — Female. Large compound eyes
well preserved. Fore and hind legs tibia with several long,
thin spines. Terminalia with unsegmented styli. SMNS 67565
(Fig. 4I) — Female. Pronotum sclerotized, nearly rounded
and about 1.5 times long as head width. Forewing sclero-
tized. Abdomen rounded. Ovipositor short, well preserved.
SMNS 67566 (Fig. 4F) — Female, with the complete and
unfold fore and hind wings. SMNS 67567 (Fig. 4H) — Female.
Hind wing in the anterior margin, that is RA area, sclerotized,
or that is pterostigma. Abdomen rounded. Ovipositor sharp.
SMNS 67568 (Fig. 4E) — Female. Antenna about the same
length as the body. Mid and hind legs tibia with several long,
strong spines. SMNS 67569 (Fig. 4D) — Female. Pronotum
is about 1.5 times as wide as the head, nearly rounded, except
in the hind point sharp. Mid leg femur with several short
spines. SMNS 67570 (Fig. 4K) — Female. Abdomen swelled.
Ovipositor and cerci well preserved. SMNS 67571 (Fig. 4B)
— Female. Abdomen rounded. Cerci with numerous sensilla.
SMNS 67572 (Fig. 4J) — Female, with the complete and
unfold fore and hind wings. Full length mid leg is preserved,
tarsus with five segments with pretarsus, the lengths are
probably 0.9, 0.6, 0.2, 0.2 and 0.3 mm, five segments with
total of 1.8 mm long.
Elisama hindwingnii sp. n. (Fig.6A)
Holotype: SMNS 67573.
Derivatio nominis: Name partially referring to well pre-
served hind wings.
Body size range (mm)
6.0–6.5
6.5–7.0
7.0–7.5
7.5–8.0
8.0–8.5
8.5–9.0
9.0–9.5
9.5–10.0 10.0–10.5 10.5–11.0 11.0–11.5
Specimen No.
1
2
1
1
2
2
0
1
2
3
2
Table 1: Body length Frequency distribution of two species, Ponopterix axelrodi: 6.4–8.0 mm long and Ponopterix maxima: 8.4–11.2 mm
long. The body size distribution in the studied specimens of this genus is rather continuous and shows no sexual dimorphism.
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Stratum typicum: Lower Cretaceous, Upper Aptian,
Nova Olinda Member of Crato Formation.
Locus typicus: Chapada do Araripe, vicinity of Nova
Olinda, southern Céara, North-East Brazil.
Differential diagnosis: Forewing at least 12 mm long.
Hind wing, except with probably slightly larger size
otherwise very similar to E. parallela Vršanský, 2003 by
venations (in E. parallela: Sc alone, R+RA four to seven, MP
three to five, CuA six to eight branches); differs from
E. tsaganica Vršanský, 1999b by hind wing larger size and
similar in most parts of venations (in
E. tsaganica hind wing
8–9 mm long and Sc simple, RA three to seven, RS three to
Fig. 4. Elisama brevis Mendes, 2000. A — SMNS 66000-125. B — 67571. C — 67563. D — 67569. E — 67568. F — 67566. G — 67564.
H — 67567. I — 67565. J — 67572. K — 67570.
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seven, MP two to four, CuA four to seven, CuP simple and A1
two to four branches). Later two morphological differences
between E. parallela and E. tsaganica are not diagnostic, and
can present sexual dimorphism (Vršanský 2003b).
Description: Forewing 12 mm long and much close to
E. parallela both in size and form. Forewing detail unknown
and is not less than 12 mm long and 4.2 mm wide. Hind wing
Sc alone, reaching less than ½ length of the anterior margin,
Fig. 5. The line RA1-ending, MA2-origin and CuP-ending. A–K — Elisama brevis. L — Elisama parallela. M — E. extenuate.
N — E. cuboides. O — Habroblattula drepanoides. L–O: After Vršanský 2003b; Wang et al. 2007a, b. redrawn here; P–S: Piniblattella
limai (Pinto et Purper, 1986). Not to the same scale.
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RA with five branches, RS with seven branches, MP with
three branches, CuA with eight branches, CuP alone and A1
with about four (to five) branches. According to the wing
characters and its large size, this specimen is a representative
of the genus Elisama, but does not belong to E. brevis, very
similar is also the hind wing of genus Kridla Vršanský, 2005.
Remarks: E. hindwingnii sp. n. is similar to E. parallela
both in size and form. Generally the hind wing is larger than
in other species of the genus.
Differential description: It differs from Kridla stastia
Vršanský, 2005 by hind wing clearly larger size (in K. stastia
8.1 mm long) and wing venation (in K. stastia: Sc alone, RA
seven, RS four, MP two, CuA six and CuP simple branches)
(Vršanský 2005a). It differs from Elisama fragmentaria
Vršanský, 2005 by larger size of hind wing (in E. fragmen
taria ca. 10–11 mm long) and with generally more branches
(in E. fragmentaria Sc simple, RA five, RS two to three, MP
two and CuA about six branches) (Vršanský 2005b,c). It dif-
fers from Elisama brevis Mendes, 2000 by larger size of hind
wing (in
E. brevis 8.5–10 mm long) and wing venations (in
E. brevis Sc alone, RA four to five, RS six to seven, MP three
and CuA six branches). It differs from Habroblattula dre
panoides Wang, Liang et Ren, 2007 from Liaoning, China by
hind wing of slightly larger size (in H. drepanoides 10 to
11 mm long) and wing venation (in H. drepanoides Sc sim-
ple, RA five to six, RS nine to eleven, MP two to five and
CuA six to nine branches) (Wang et al. 2007a). Elisama
extenuata Ren, 1995 by hind wing 7–8 mm long and Sc
simple, RA four to six, RS four to seven, MP three to four
and CuA five to six branches (Wang et al. 2007b). Elisama
cuboides Wang, Ren et Liang, 2007 by hind wing 10 mm
long and Sc simple, RA six, RS seven, MP six and CuA
seven branches (Wang et al. 2007b). Macaroblattula
ellipsoids Wang, Ren et Liang, 2007 by hind wing 10.5 mm
long and Sc simple, RA four to five, RS six to eight, MP six
and CuA six to eight branches (Wang et al. 2007b).
Character of preservation: Only one pair complete and
nearly unfolded hind wing preserved.
Genus: Ocelloblattula Anisyutkin et Gorochov, 2008
Type species: Ocelloblattula. ponomarenkoi Anisyutkin
et Gorochov, 2008.
Stratigraphic range: Hauterivian – Aptian.
Geographic range: Gondwana.
Ocelloblattula santanensis sp. n.
Holotype: SMNS 66558
Derivatio nominis: Named after the city of Santana
do Cariri near the type locality.
Stratum typicum: Lower Cretaceous, Upper Aptian,
Nova Olinda Member of Crato Formation.
Locus typicus: Chapada do Araripe, vicinity of Nova
Olinda, southern Céara, North-East Brazil.
Diagnosis: This species is very similar to Elisama brevis
both in size and form, it differs from the forewing preserved
with a distinctly band colour pattern (Figs. 6, 7), these
discontinuous markings suggest membership of the genus
Ocelloblattula Anisyutkin et Gorochov, 2008 (family
Blattulidae). Head with very large and globular compound
eyes, probably three ocelli (SMNS 66558, Fig. 6C) reside in
the forehead between the compound eyes.
Description: Small size roaches. Antenna about the same
length as body. Body length between 7.9–8.2 mm. Head wide
and clearly longer than the fore margin of pronotum. Pronotum
is about 1.5 times wide as the head, approximately of round
shape, except in the posterior part where it is sharp gradually.
Forewing is morphologically very similar to Elisama brevis
(Table 4), 8.0–8.9 mm long and 2.4–2.7 mm wide and
shows rust red colour. Foreleg tibia is about the same length
as tarsus.
Superfamily: Blattoidea Latreille, 1810
Family: Ectobiidae Brunner von Wattenwyl, 1865
(=Blattellidae Karny, 1908)
Genus: Piniblattella Vršanský, 1997
Type species: Mesoblattina vitimica Vishniakova, 1964
Composition: Piniblattella sharingolensis, Vršanský,
2005; P. minuta Vršanský, 1997.
Stratigraphic range: (probably Berriasian) Hauterivian
– Albian.
Geographic range: Cosmopolitan.
Piniblattella limai (Pinto et Purper, 1986) (Fig. 8)
1986 Mesoblattina limai. — Pinto et Purper, 6–10.
Elisama brevis
Forewing
(Right/Left)
SMNS
SC
RA+RP
M
CuA CuP
A
Total
66000-125
1/1
13/12
3/3
4/6
1/1
7/6
29/29
67564
1
15
4
6
1
6
33
67565
1
14
3
4
1
6
29
67566
1/1
13/14
3/3
5/6
1/1
7/8
30/33
67567
1/1
14/14
4/3
6/5
1/1
5/5
31/29
67572
1
16
3
6
1
6
33
Elisama brevis
Hind wing
(Right/Left)
SMNS
SC
RA
RS
MP
CuA
Total
66000-125
1/1
4/4
6/6
3/3
6/6
20/20
67564
1
5
7
3
6
22
67565
1/1
4/4
6/6
3/3
6/6
20/20
67566
1/1
6/5
6/6
3/3
6/7
22/22
67567
1/1
5/4
7/7
3/3
6/6
22/21
67571
1
5
6
3
6
21
67572
1
4
7
3
6
21
Table 2: Forewing venation variability of Elisama brevis Mendes,
2000.
Table 3: Hind wing venation variability of Elisama brevis Mendes,
2000.
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Diagnosis (improved after Pinto & Purper (1986), Bechly
(2007a)): Body length 11–14 mm; antenna about as long as
body; pronotum subcircular, very broad (about 2–2.33 times
of head width) with very broad lateral lobes; forewing vena-
tion typically ectobiid with costa area short, Sc bifurcated,
R 8–11 branches (some of them bifurcated), MA 4–6
branches (some of them bifurcated), CuA generally bifur-
cated, several anal veins simple and ending on CuP, inter-
calary veins and crossveins present (not as few as mentioned
in the original description). It differs from Piniblattella
magna sp. n., by significantly smaller size, forewing CuA
with fewer branches (in P. magna sp. n. three to five) and
A-veins not fused. It differs from P. vitimica by generally
smaller size (forewing, 13.5 to 15.5 mm long and 4 to 4.9 mm
wide; hind wing 15.2 mm long); by forewing M with fewer
branches than P. vitimica (2–5 dichotomizing branches
and 12–22 veinlets) and CuA with only two main branches
(P. vitimica one to seven veinlets). Compared to the
other species of genus Piniblattella differs by less
veins-venations.
It differs from P. sharingolensis Vršanský, 2005 by fore-
wing without intercalaries. Hind wing unknown.
Description: Pronotum subcircular, hind margin rela-
tively straight, about twice the width of the head, fore
Fig. 6. A — Elisama hindwingnii sp. n.; SMNS 67573. 1. Fore- & hind wings. 2. Hind wings. B–D: Ocelloblattula santanensis sp. n.
B — SMNS 66558. C — Forehead having probably three ocelli. D — 66316.
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margin of pronotum reaches nearly the same length as
the top of head. Head with two large compound eyes, with
two ocelli in the forehead. Thorax with coxa in roaches
form, mid-and hind leg tarsus about 4/5 length of tibia,
tarsus with five segments. Forewing: 10.3–13.0
mm
long and 3.5–4.2 mm wide, Sc typically blattellid with
bifurcated, RA+Rs with a total of 15–23 branches, M with
six–nine branches in total (except SMNS 66315 left side
with five, Fig. 8C), CuA with two branches (except SMNS
66315 left side with four). Forewings show very low
morphological differences (Table 5). Hind wing: about
12–13 mm long, detail still unknown. Cerci in typical
roaches form and about 14 segments. Terminalia with unseg-
mented styli.
Description of other specimens: SMNS 66312 (Fig. 8A):
Ventral side. Head with two well preserved compound eyes,
near the scape and in the fore head side exist two ocelli,
mouthparts well preserved. Pronotum about twice the width
of the head. Thorax with coxa in roaches form. Terminalia
near cerci with one pair styli, unsegmented. SMNS 66314
(Fig. 8B): Antenna shorter than body length. Mid- and hind
leg tarsus long and well preserved. SMNS 66315 (Fig. 8C):
The venation between left and right forewings clearly diffe-
rent, especially in M and CuA. In the right side, CuA with
Fig. 7. Ocelloblattula santanensis sp. n. Uncatalogued specimens from diverse private collections.
F
B
G
K
C
H
D
I
E
J
A
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O. santanensis
Forewing
(Right/Left)
SMNS
SC
RA+RP
M
CuA CuP
A
Total
66316
1/1
14/14
3/3
4/4
1/1
7/8
30/31
66558
1/1
15/15
3/3
5/5
1/1
6/6
31/31
Fig.7A
1/1
15/13
3/3
5/5
1/1
6/7
31/30
Fig.7B
1/1
13/14
4/4
5/4
1/1
8/7
32/31
Fig.7C
1/1
13/15
3/4
5/4
1/1
8/7
31/32
Fig.7D
1/1
14/14
3/3
4/3
1/1
6/6
29/28
Fig.7E
1/1
16/–
3/–
4/4
1/1
7/7
32/–
Fig.7F
1/1
15/13
4/4
4/4
1/1
6/6
31/29
Fig.7G
1/–
15/–
–
–
–
–
–
Fig.7H
1/1
15/13
4/5
5/5
1/1
7/7
33/32
Fig.7I
1/–
13/–
4/–
5/–
1/–
7/–
31/–
Fig.7J
–/1
–/13
–/5
–/4
–/1
–/6
–/30
Fig.7K
1/1
16/15
3/3
4/5
1/1
7/7
32/32
two- and M with a total of nine branches, that is more in
“normal” form. On the left side, the CuA with four- and M
with “only” five branches, is rather unusual. Left side is also
a little smaller (9 mm long, 3 mm wide) than the right side
(10.3 mm long, 3.5 mm wide) and the “normal” size. It is pos-
sible, that the left side is still in nymph phase, but the sclero-
tization in both sides is quite equal. SMNS 66319 (Fig. 8E):
Pronotum about double width of head. Its fore margin is
almost the same length as the head. SMNS 66322 (Fig. 8D):
Hind leg tibia with several spines. Terminalia with one pair
well preserved cerci, 14 segments.
Piniblattella magna sp. n. (Fig. 9)
2007 Unnamed new genus and species A. — Bechly, 248–249.
Holotype: SMNS 66000-116.
Paratypes: SMNS 66310; 66313; 66317; 66318; 66320.
Derivatio nominis: Named after the large size.
Stratum typicum: Lower Cretaceous, Upper Aptian,
Nova Olinda Member of Crato Formation.
Locus typicus: Chapada do Araripe, vicinity of Nova
Olinda, southern Céara, North-East Brazil.
Differential diagnosis (improved after Bechly 2007a):
The species is categorized within Piniblattella, which differs
from Mesoblattina by forewing with less M branches and
A with less numerous reticulations. Pronotum shield-like,
forewing with M strongly branched and CuA with just two
major branches. Large size roaches. Body length
17.8–25.0 mm; antennae distinctly longer than body (31 mm
in a specimen with 25 mm body length); pronotum very large
with 2.15–2.33 times of head width and with very broad
lateral lobes; pronotum broadest in the middle of the poste-
rior half; forewing generally less than body in length, vena-
tion typically blattellid. It differs from P.
limai by its dis-
tinctly larger size and in having forewing CuA three–five
branches (in
P. limai the CuA is with two branches), and the
A-veins fused often especially in the base part. It differs from
P. vitimica by generally larger size and forewing M with
fewer branches (by P.
vitimica with 2 to 5 dichotomizing
branches and 12 to 22 veinlets). Larger compared with other
Piniblattella spp.
Description: Antenna shorter than double the body length
(2 ×17.8–27.0 mm). The top of head slightly protrudes from
the fore margin of pronotum. Pronotum broad, approximately
oval shape, posterior half much wider and hind margin rela-
tively straight. Forewing: 16–21 mm long and 4.8–6.2 mm
wide, Sc with two major branches, each one sometimes has
several (two–four) terminal branches, RA+RS with 18–20
branches, M with 8–12 major branches, some branches with
more terminal branches, CuA with three–five branches,
A-veins fused often especially in the base part. Forewings
show very few morphological differences (Table 6). Hind
wing is about the same length as forewing and is 16–19.5 mm
long. Hind leg tarsus is about ¾ of the length of the tibia and
with five or four (SMNS 66000-116, 66317; see Fig. 9A–B)
segments, the first segment is longest and about half of the
length of the whole tarsus. Segments continue to be shorter
from the second to the fourth tarsal segment, the fifth seg-
ment is elongated again, with two claws and pretarsus. Cerci
with 17–20 segments.
Description of other specimens: SMNS 66000-116
(Fig. 9A): Body size (17.8 mm) relatively smaller than in the
other specimens of this species. Pronotum broad and well
preserved. The male external genitalia presented with hook-
like structure. Unusual hind leg tarsus, left side with five but
right side with four segments. The right side tarsus (totally
4.7 mm) is shorter than the left side tarsus (totally 5.1 mm).
On the basis of the length of each segment, the “third” seg-
ment should be lost by right side. This “four segments
tarsus-” (Vršanský 2002 reported it from a Cretaceous
mantodean Jantarimantis zherikhini), which reflects incom-
plete regeneration (Bohn 2003), is not rare, in specimen
SMNS 66317 both sides of hind leg tarsus with only four
segments. SMNS
66310 (Fig. 9C): Body size relatively large.
Antenna (41 mm long) is about shorter than double of body
length (25 mm). Cerci well preserved, about 20 segments.
SMNS 66313 (Fig. 9F): Mid and hind leg tarsus with five
segments. SMNS 66317 (Fig. 9B): Hind leg tarsus with four
segments. SMNS 66318 (Fig. 9E): Forewing Sc with several
branches. Abdomen swelled. SMNS 66320 (Fig. 9D): Fore-
wing Sc with two major branches, the Sc2 has further termi-
nal branches. Hind wing is only partly preserved.
Family: Mesoblattinidae Handlirsh, 1906
Genus:
Perlucipecta Wei et Ren, 2013
Perlucipecta santanensis sp. n. (Fig. 10)
2007 Unnamed genus and species B. — Bechly, 249.
Holotype: SMNS 66308.
Paratypes: SMNS 66309; 66311; 66321.
Derivatio nominis: Named after the city of Santana
do Cariri near the type locality.
Table 4: Forewing venation variability of Ocelloblattula
santanensis sp. n.
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Stratum: Lower Cretaceous, Upper Aptian, Nova Olinda
Member of Crato Formation.
Locality: Chapada do Araripe, vicinity of Nova Olinda,
southern Céara, North-East Brazil.
Diagnosis: Small-sized roaches. Body about 8.7–11.9 mm
length; shape of body longish oval; antennae about as long as
body; pronotum much broader than head (width 3.0–3.7 mm,
thus 1.8–2 times head width), posteriorly broader than ante-
riorly, but with narrower lateral lobes than blattellid
species mentioned above; forewing thin, venation with
a broad costal margin.
Description: First time reported by Bechly (2007a).
Antenna a little longer than body length. Body about
8.7–11.9 mm long. The top of head slightly protrudes from
the fore margin of pronotum. The hind margin of pronotum is
relative straight. Forewing, costal margin broad, Sc probably
bifurcated, Sc1 extra short and except the base part not very
distinct. Hind wing Mp at least three branches, CuA at least
five branches. Fore-, mid-, and hind legs show similar pat-
terns, that is, tibia always slightly longer than femur, tarsus
with five segments and all together about 70–86% of tibia
length, the first segment is longest, about a half of the whole
tarsal length, the second segment is the second longest, third
and forth segment shorter, the fifth segment elongated again
and with claws and pretarsus. Cerci in roaches form, 13–14
segments. Treminalia with unsegmented styli.
Fig. 8. Piniblattella limai; A — SMNS 66312. B — 66314. C — 66315,
variations different between right- & left forewings from the same individual.
D — 66322. E — 66319.
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Description of other specimens: SMNS 66308 (Fig. 10A):
Antenna slightly longer than the body length. Fore- and hind
wings overlap and only partly identifiable, those are: fore-
wing Sc with two branches, CuA at least two branches, hind
wing Mp at least three branches, CuA at least five branches.
Foreleg tarsus well preserved. Cerci in typical roaches form,
14 segments. SMNS 66309 (Fig. 10B): Forewing probably
thinner, only the base part slightly sclerotized. Hind leg well
preserved. SMNS 66311 (Fig. 10C): Hind leg well preserved.
Terminalia with unsegmented styli. SMNS 66321 (Fig. 10D):
Ventral side. Head with a pair of large compound eyes.
Mouth parts, including labrum, maxillary- and labial palps
partly visible. In thorax, the coxal and the trochanters were
well preserved.
Genus: Raptoblatta Dittmann, Hörnig,
Haug et Haug, 2015
Remarks: Predator regarded as a relative of praying man-
todeans, with venation with characteristic reticulations visible
on the apical parts of both left and right forewings as well as
hind wings. This taxon is difficult to discriminate from the
dominant Laurasian as well as Gondwanan Archi mesoblatta,
and is very closely related to Gondwanan Gondwablatta,
with identical forelegs (spines were not preserved), but with
a different pronotum. Pronotum of Raptoblatta is apparently
misinterpreted and has a standard, vaulted, small and trans-
versal shape. Nevertheless, it is different from Gondwa
blatta. The carnivorous mode of life might explain the lack
of other predatory lineages of cockroaches in nume rous
localities with occurrence of Archimesoblatta. Nevertheless,
categorization within Mesoblattinidae excludes the manto-
dean relation.
Raptoblatta waddingtonae Dittmann, Hörnig,
Haug et Haug, 2015
Material: Monotypic, holotype only. Axelrod Institute,
University of Guelph, Canada: AI 514
Diagnosis: Medium-sized, roach-like, winged insect with
wide abdominal segments and cockroach-type cerci; most
prominent structure is the foreleg with the row of femural
and tibial spines; the median side of the distal region of the
femur is concave - this region is equipped with small spines;
and has nearly the same length as the tibia.
Discussion
The ecologically-dominant family of the order Blattaria in
the Crato Formation is represented by the family Ectobiidae,
with about 83 % of the cockroach specimens. This result is
similar to that of Vršanský (2004) who also reported that this
family accounts for 60 % of the cockroaches in the Crato For-
mation based on the American Museum of Natural History
(AMNH, New York) collection. Blattulidae represent about
13 % of the cockroaches. This study shows a surpri singly low
abundance of the family Umenocoleidae (only 4 %) com-
pared to the AMNH collection, which shows that the Umeno-
coleidae are represented by up to 15 % of all roaches
(Vršanský 2004). Although the Cretaceous cockroach assem-
blages differ from the Jurassic cockroach assemblages (this
transition was characterized by the temporal change in the
dominance of particular families), Blattulidae remain domi-
nant during the Cretaceous, together with the Ectobiidae
(Vršanský 2002). Blattulidae are also dominant in the Yixian
formation (Late Jurassic/Early Cretaceous) of northern China
(Wang et al. 2007b). The Crato dictyopterans (see also Lee
2014) show similarities with those at the Cretaceous localities
of Mongolia (e.g., Vršanský 2003b, 2008b) and Chernovskie
Kopi in Russia (Barna 2014), namely Chaeteessidae, Ecto-
biidae, Blattulidae and Umenocoleidae. The Upper Jurassic
assemblage from Shar-Teg (Mongolia) shows a more even
cockroach assemblage with Blattulidae (38 %), Mesoblatti-
nidae (25 %), Liberiblattinidae (13 %) and Caloblattinidae
(25%) (Vršanský 2005 b, c; Barna 2014). The cockroach
assemblage of the Lower Cretaceous in Montsec (Spain) is
dominated by blattulids and meso blattinids (Vršanský and
Ansorge 2001). The lack of the family Fuziidae (Vršanský et
al. 2009) in the Crato Formation supports an endemic status
of the family in Inner Mongolia (China, Daohugou).
Palaeogeographical distribution of genera reveals interes-
ting patterns. Only Ocelloblattulla is limited to Gondwana,
while Perlucipecta, Piniblattella, Elisama and possibly
Petropterix (“P.” maxima) were also present in Laurasia.
Cratovitisma, Ponopterix, Raptoblatta (possibly Archi meso
blatta) and the new genus Umenopterix are endemic.
It follows that the primitive taxa are shared with Laurasia,
while the more advanced cockroaches are either indigenous
or restricted to Laurasia (while progressive Raphidiommidae
as well as rather primitive Caloblattinidae were missing).
On the other hand, phylogenetically-advanced Laurasian
taxa such as
Praeblattella (although recorded in the
New Jersey amber), T
arakanula, Elytropterix and Petro
pterix do not occur in the Crato Formation.
Piniblattella limai
(Right/Left)
SMNS
SC
RA+RP
M
CuA CuP
A
Total
66312
2/2
21/22
9/6
2/2
1/1
7/6
42/39
66314
2/2
19/18
6/8
2/2
1/1
6/6
36/37
66315
2/2
16/14
9/5
2/4
1/1
5/5
35/31
66319
2
23
7
2
1
5
40
66322
2
18
7
2
1
6
36
Piniblattella magna
(Right/Left)
SMNS
SC
RA+RP
M
CuA CuP
A
Total
66000-116
2/2
19/20
7/13
2/2
1/1
8/6
39/44
66317
2/2
19/19
8/11
4/3
1/1
8/9
42/45
66318
3/4
19/19
8/9
3/5
1/1 10/10
44/48
Table 5: Forewing venation variability of Piniblattella limai.
Table 6: Forewing venation variability of Piniblattella magna sp. n.
447
APTIAN COCKROACH (INSECTA) ASSEMBLAGES, CRATO FORMATION, NE BRAZIL
GEOLOGICA CARPATHICA
, 2016, 67, 5, 433 – 450
Phylogenetic context is congruent with the Aptian datings
of the locality. While it is impossible to delimit the phylo-
genetic status for Elisama spp., and Piniblattella spp. because
they do not possess phylogenetically important characters,
Perlucipecta can be regarded as somewhat more advanced
than its Yixian congener Ponopterix and Ocelloblattulla are
assumed to be extremely advanced taxa in spite of some
primitive characters, the latter genus furthermore also occurs
in earlier Lebanon amber (without Plesiomorphies). On the
other hand, Cratovitisma and Umenopterix are extremely
primitive compared to all (including the stratigraphically
basalmost ones) Cretaceous localities because they possess
Fig. 9. Piniblattella magna sp. n. A — holotype habitus; SMNS 66000-116, right & left hind legs tarsus with four—five segments.
B — SMNS 66317. C — 66310. D — 66320. E — 66318. F — 66313.
448
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GEOLOGICA CARPATHICA
, 2016, 67, 5, 433 – 450
characters such as standard pronotum with fully developed
paranotalia.
Acknowledgements: I am thankful to Dr. Günter Bechly
(Stuttgart) and Prof. Dr. James Nebelsick (Tübingen) for the
patient instruction and general help. I thank Dr. Karin
Wolf-Schwenninger (Stuttgart) for technical support. I am
most grateful to Dr. Peter Vršanský (Bratislava, Moscow) for
his help and critical comments. I thank Mrs. Annesuse
Raquet-Schwickert and Mr. Michael Schwickert
Fig. 10. Perlucipecta santanensis. sp. n. A — holotype habitus: SMNS 66308. B — 66309. C — 66311. D — paratype habitus: 66321.
449
APTIAN COCKROACH (INSECTA) ASSEMBLAGES, CRATO FORMATION, NE BRAZIL
GEOLOGICA CARPATHICA
, 2016, 67, 5, 433 – 450
(Sulzbach tal) for their friendly support and providing gene-
rous access to the fossil material of their collection (Fossils
Worldwide, Sulzbachtal, Germany). I thank Mrs. Sabine
Petri, Mrs. Angelika Schäfer, Mr. Detlef Stark and Mr. Walter
Siegl for the assistance with the routine work in the library
and museum. This work was financial supported by Ministry
of Education, Taiwan. Finally I want to thank the Staatliches
Museum für Naturkunde, Stuttgart, Germany for generously
providing a working space and allowing for me to access the
studied material.
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