6(1):9-13, jan/jun 2010
© Copyright 2009 by Unisinos - doi: 10.4013/gaea.2010.61.02
New Blattoptera from Early Cretaceous
of Santana Formation (Araripe Basin, NE
Brazil) and a review of Arariplebatta
Mendes, 2000
Rafael Gioia Martins-Neto
Universidade Federal do Ceará (UFC), Campus Cariri/Sociedade Brasileira de Paleoartropodologia – SBPr. Av. Tenente Raimundo Rocha,
S/N, Cidade Universitária, 63000-000, Juazeiro do Norte, CE, Brasil. martinsneto@terra.com.br
Cynthia de Oliveira Assis
Centro de Ensino Superior de Juiz de Fora. Rua Dr. Constantino Paleta, 100/202 Centro, 36.015-450, Juiz de Fora, MG, Brasil.
cynthia.assis@gmail.com
Lara Vaz Tassi
Área Paleontología (Centro de Ecología Aplicada del Litoral – Consejo Nacional de Investigaciones Científicas y Técnicas), Casilla de Correo
128, 3400 Corrientes, Argentina. larinhatassi@hotmail.com
ABSTRACT. Arariplebatta Mendes is here reviewed taking into account its previous diagnosis in Portuguese, the proposal of a new species, A. cesi,
and an emended diagnosis of Blattulopsis beckeri Mendes. To include the genus and the new related forms the new family Araripeblattidae is presented and Elisama americana Vršanský is considered a synonym of Araripeblatta brevis Mendes. Those Blattoptera came from the same laminated
limestones from the basal part of Santana Formation, Ceará State, Northeast Brazil, Late Aptian in age, where the original samples were collected.
Key words: Blattopera, Araripeblatta, Araripeblattidae n. fam., Santana Formation, Lower Cretaceous, Brazil.
RESUMO. NOVOS BLATTOPTERA NO CRETÁCEO INFERIOR DA FORMAÇÃO SANTANA (BACIA DO ARARIPE, NE DO BRASIL) E A REVISÃO DE ARARIPLEBATTA
MENDES, 2000. O gênero Araripeblatta Mendes é aqui revisado considerando-se sua diagnose original em português, a atribuição de uma nova
espécie, A. cesi, e os novos elementos propostos para a diagnose de Blattulopsis beckeri Mendes. Para incluir o gênero e as novas formas relacionadas uma nova família é apresentada, Araripeblattidae, e Elisama americana Vršanský, é entendida como sinonímia de Araripeblatta brevis
Mendes. Essas formas de Blattoptera provêm dos mesmos níveis de calcário laminado das previamente descritas, correspondentes à parte basal da
Formação Santana (Membro Crato), Bacia do Araripe, estado do Ceará, com idade Neoaptiano.
Palavras-chave: Blattopera, Araripeblatta, Araripeblattidae n. fam., Formação Santana, Cretáceo Inferior, Brasil.
INTRODUCTION
The Blattoptera are an ancient group
of insects which dominate the Late
Carboniferous, which became rare
during the Triassic and greatly diversified again in Jurassic and Cretaceous
times (Schneider, 1983; Vršanský, 2002;
Grimaldi and Engel, 2005). Like other
Dictyoptera, their phylogenetic relation-
ships remain controversially discussed in
regard of some discrepancies between
the stratigraphic record and the proposed
phylogeny (Grimaldi, 2001). Following
the recent molecular studies of Lo et al.
(2002) the living roaches are a paraphyletic group of insects.
The blattopteran fauna from Santana
Formation was firstly investigated by
Pinto and Purper, (1986), Pinto (1989),
and in the following by Marcio Mendes,
who has produced ten short communications in scientific meetings, apart
from his master and doctoral thesis
about the Araripe Basin blattopterans.
The material described by Mendes
(2000) was housed at Guarulhos University (São Paulo, Brazil) and quite all
were collected by the senior author of
the present contribution. The single
New Blattoptera from Early Cretaceous of Santana Formation (Araripe Basin, NE Brazil) and a review of Arariplebatta Mendes, 2000
comprehensive publication (Mendes,
2000) was written in Portuguese, including the diagnostic features, contrary the
rules of the International Code of Zoological Nomenclature, which was one of
the motives to the study presented here.
The Santana fossil insects, including
Blattoptera forms, were also focused
in foreign researchers, especially in
the 1990s, based on samples stored
in collections outside Brazil. In resulting publications new forms were
described, sometimes ignoring the
Brazilian contributions.
In this work, fossil specimens of
Blattoptera from a small collection of
the Paleontological Laboratory from
Centro de Ensino Superior de Juiz de
Fora (CES-JF), Minas Gerais State,
Brazil, are reviewed in the light of the
foregoing commentaries.
The quarry location and the geological setting of Araripe Basin are shown
in Figure 1, and the stratigraphic data
follow the conceptions from Ponte and
Ponte Filho (1996), and Assine (2007).
The methodology and terms of the
insect descriptions are that presented in
Martins-Neto et al. (2005).
Diagnosis. Small-sized blattopteran
with tegmen length around 9 mm.
Females with recurrent ovipositor.
Contact points of first R branch with
the anterior wing border of MA origin
(oMa), and contact of CuP with the
posterior wing border are arranged in
one line slightly transverse to the wing
long axis. M and R fused close to the
tegmen base. Paired small spines on the
left fore tibia (in ventral view), and not
paired long spines in the right fore tibia
(in ventral view).
Discussion. Araripeblattidae n. fam.
differs from Blattulidae Vischniakova
(1982), the closest family, by having
RA, oMA, and CuP transversely aligned.
In Blattulidae tegmen, the secondary
branches of RP and MA are restricted
to the apical area, above the middle of
the apex. A line between the contact
of RA with the anterior wing border
and the point of contact of CuP with
the posterior wing border are nearly
perpendicular arranged to the wing long
axis. The point of M bifurcation (oMA)
is situated slightly backward of this line
(lcm=aml< oMA, see Martins-Neto et
al., 2005).
SYSTEMATIC PALEONTOLOGY
Araripeblatta Mendes, 2000
Order Blattoptera Brunner, 1882
Araripeblattidae fam. nov.
Type genus. Araripeblatta Mendes, 2000
here designated.
Type species. Araripeblatta brevis Mendes, 2000
Emended diagnosis. Apart the family characteristics, anal field occupying
around 1/3 of the tegmen length and
80% of the tegmen width.
Figure 1. Geological and stratigraphical setting of Araripe Basin (modified from Assine,
2007), and Pedra Branca Mine location, where it is exposed the calcareous beds.
10
Araripeblatta brevis Mendes, 2000
(Figures 3A-B)
Synonym. Elisama americana Vršanský
2002, fig. 11, 21–26, 30.
Emended diagnosis. Apart the generic
diagnosis, tegmen 9 mm long and 2.7
mm wide.
Holotypus. A female specimen, housed
at the Laboratory of Geosciences, Guarulhos University, Guarulhos, Brazil, collection number UnG/1T-20 (Figure 3A).
Paratypus. A female specimen, housed
at the same Institution, collection number UnG/1T-21 (Figure 3B).
DISCUSSION
Araripeblatta was firstly proposed by
Mendes (2000), based on a single and
relatively complete specimen, preserved
in ventral view. Vršanský (2002) created
a new species (E. americana) of the
genus Elisama Giebel 1856, based on
fragmentary material from the Santana
Formation, housed at the American
Museum (USA, AMNH XX; stated this
way by Vršanský, 2002, probably due
to a lapse). He ignored or omitted the
description of A. brevis Mendes (2000),
although both type specimens are identical in the preserved characteristics. In
his summary of Blattoptera remains
from Araripe Basin, Bechly (2007)
refers the work of Mendes (2000) but
does not mention Araripeblatta brevis
neither other published blattopterans from there. Obviously based on
Figure 2. Blattullopsis popovi Pinto
(Blattullidae), modified from Pinto
(1990). Abbreviations: cua-cup, cross
veins between CuP and CuA; CuP,
posterior cubitus; oMA, origin of
anterior Media; RA, anterior Radius.
Scale bar: 5 mm.
Rafael Gioia Martins-Neto, Cynthia de Oliveira Assis and Lara Vaz Tassi
Figure 3. A-B. Araripeblatta brevis
Mendes, 2000, holotype and paratypus,
respectively, redrawn from Mendes
(2000); C. Araripeblatta beckeri n.
comb., redrawn from Mendes (2000).
Scale bar: 1 mm.
Vršanský (2002), Bechly (2007) figured a
specimen as Elisama Giebel 1856 (Figure
11.23f). Nevertheless, the figured specimen exhibits the same morphology of
Araripeblatta brevis, and it is clearly not
related to Elisama, a Blattulidae with
species known from European Jurassic
and Jurassic/Cretaceous transition in
Asia (Wang et al., 2007). Elisama americana
is here considered a junior synonym of
Araripeblatta brevis.
Araripeblatta beckeri n. comb.
(Figure 3C)
Araripeblatta cesae n. sp
(Figures 4, 5)
Mendes (2000) has erected Blattulopsis beckeri Mendes (2000) for a distinct
specimen from Santana Formation
(UnG/1T-025, housed at Guarulhos
University), considering it as Blattulidae. This specimen coincides perfectly
with Araripeblatta as well as with the
new family here proposed, leading to
the proposition of the new combination. Common feature is a virtual
straight line, arranged slightly transverse to long axis of the wing, which
connects the point where the RA (first
R branch) ends, at the anterior wing
border, with the point of oMA, as well
as with the point where the CuP ends,
at the posterior wing border. Those
aspects lead to the proposition of the
new combination.
A. beckeri n. comb. has a slightly
longer tegmen (10 mm) and exhibits
intercalary veins (plesiomorphic), also
present in both A. brevis and A. cesae.
For otherwise, both A. brevis and A.
beckeri have not the area between CuP
and the first anal vein filled by cross
veins. Another interesting fact to note
is the confirmation that the left forelegs
(in ventral view) have relatively small
paired spines in all of the three Araripeblatta species, as well as the right foreleg
exhibiting longer and not paired spines
(see Figures 3B-C).
Etymology. Latinized from CES, Centro de Ensino Superior de Juiz de Fora
(CES/JF), Minas Gerais, Brazil.
Holotypus. CES-JF – I – 001, housed
at Centro de Ensino Superior de Juiz e
Fora, CES/JF.
Diagnosis. Area between CuP and the
first anal vein filled by numerous and
perpendicular cross veins; CuP with a
distal secondary branch relatively parallel
to both Medial ones and the anal margin.
Type locality. Pedra Branca Mine, on
the road that connects Nova Olinda and
Santana do Cariri municipalities, 4 km
far from the first city, Ceará State, Brazil.
Stratigraphy. Upper part of Crato
Member, Santana Formation.
Age. Late Aptian.
Description. Forelegs armed with spaced
paired short spines at the left fore tibia, and
longer ones, not spaced on the right one
(at right in Figure 4A). Tegmen strongly
sclerotinized, ornamented secondary
veins consisting of small homogeneous
granules (see detail in Figures 4B-C and
5A-B), 8.83 mm long and 2.91 mm wide.
Basal width as narrow as apical one. Distal
end of RA (first branch of R) close to the
tegmen mid length. Ten RP secondary
branches, the last four dichotomous, and
the last one reaching the apical area. M origin close to the tegmen base, MA simple,
parallel to RP; MP distally dichotomous.
Figure 4. Araripeblatta cesi n. sp. (holotype). A. Fore and hind legs; B. Right tegmen.
Abbreviations: A, anal vein; CuA, anterior cubitus; CunP, posterior cubitus; MnA,
anterior media; MP, posterior media; oMA, Ma origin; RA, anterior radius; RP, posterior
radius. Scale bars: A, 2.5 mm; B, 5 mm.
Gaea - Journal of Geoscience, vol. 6, n. 1, jan/jun 2010, p. 9-13.
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New Blattoptera from Early Cretaceous of Santana Formation (Araripe Basin, NE Brazil) and a review of Arariplebatta Mendes, 2000
Figure 5. Araripeblatta cesi n. sp., holotype. A. Detail of the left tegmen; B. Right tegmen. C. Habitus; Scale bar: 3 mm.
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Rafael Gioia Martins-Neto, Cynthia de Oliveira Assis and Lara Vaz Tassi
CuA five-branched, originating from the
Sc+R+M basal trunk, close to the tegmen
base. CuP strongly curved, reaching the
anal area at 1/3 of the tegmen length. A
virtual connection of the point where the
RA (first R branch) ends, at the anterior
wing border, with the point of oMA, as
well as with the point where the CuP ends,
at the posterior wing border, which forms
a straight line arranged slightly transverse
to the wing long axis. Five parallel anal
veins oblique to the anal margin. Space
between CuP and the first anal vein filled
by cross veins.
Discussion. Araripeblatta cesae n. sp. differs
from the closest species, Araripeblatta brevis
Mendes, in having the area between CuP
and the first anal vein filled by cross veins,
and the secondary CuP branch parallel and
close to the posterior wing margin.
CONCLUSION
Over the last years, the taxonomy
of Blattoptera from the Lower Cretaceous of Santana Formation became
increasingly confused. It happened due
to descriptions of isolated samples,
sometimes badly preserved, that were
housed in foreign museums or in private
collections. Furthermore, it is effected
by the inadequate approaches of local
researchers, who published the diagnoses
of new taxa in Portuguese, contrary to
the rules of the International Code of
Zoological Nomenclature. Here it is
aimed at a revision of so far published
taxa from Santana Formation, and it is
proposed a new taxon of family rank that
joints some of the previously described
genera and species. The Araripeblattidae
n. fam. includes the genus Araripeblatta
Mendes and, at least, three species. Some
of the conspicuous characteristics of
this blattopteran group are the small size,
the presence of a recurrent ovipositor in
females (plesiomorphic) and a not constant color pattern (none of the material
here analyzed exhibits it).
ACKNOWLEDGEMENTS
We would like to express our deep
thanks to Jöerg Schneider, from the
Institute of Geology, Department of
Palaeontology and Stratigraphy, TU
Bergakademie Freiberg, Germany, and
Karen Adami, from Universidade Federal de Pelotas, Brazil, for the review
of this manuscript, suggestions and
friendly help.
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Submitted on: 20/03/2010
Accepted on: 07/07/2010
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