Download Béthoux和Wieland - 2009 - Evidence for carboniferous origin of the order mantodea (insecta Dictyoptera) gained from forewing

Zoological Journal of the Linnean Society, 2009, 156, 79–113. With 23 figures
OLIVIER BÉTHOUX1,2* and FRANK WIELAND3
1
Freiberg University of Mining and Technology, Institute of Geology, Department of Palaeontology,
Bernhard-von-Cotta Str. 2, D-09596 Freiberg, Germany
2
State Natural History Collections of Dresden, Museum of Zoology, Königsbrücker Landstraße 159,
01109 Dresden, Germany
3
Georg-August-Universität, Johann-Friedrich-Blumenbach-Institut für Zoologie und Anthropologie
und Zoologisches Museum, Abt. Morphologie, Systematik und Evolutionsbiologie, Berliner Str. 28,
37073 Göttingen, Germany
Received 26 February 2008; accepted for publication 30 May 2008
Homologies of the forewing venation pattern of the order Mantodea (Insecta: Dictyoptera) consistent with the
accepted insect wing venation groundplan are proposed. A comparative morphological analysis was carried out
based on a broad taxonomic sample of extant taxa. Besides macromorphological aspects, focus is given to the
pattern of the tracheal system as a basis for establishing primary homologies. All extant praying mantids exhibit
a composite stem composed of the posterior radius (RP) and the media (M) and most praying mantids exhibit a
fusion of the anterior branch of RP + M with the anterior radius (RA). The wing venation of the species †Mesoptilus
dolloi, previously assigned to the polyphyletic fossil assemblage ‘Protorthoptera’, is re-interpreted in the light of the
new homology statement. Our interpretation suggests that it is a putative stem-Mantodea, as are some other
‘protorthopterous’ taxa. This hypothesis implies that the total-group Mantodea arose as soon as the Late
Carboniferous, i.e. about 175 million years earlier than previously estimated. This analysis contributes to the view
that most of the Late Carboniferous ‘Protorthoptera’ are stem-representatives of the major polyneopteran clades
(e.g. cockroaches, grasshoppers and crickets, rock-crawlers), suggesting a survivorship of several main Pterygota
lineages at the end-Permian extinction event higher than previously expected. © 2009 The Linnean Society of
London, Zoological Journal of the Linnean Society, 2009, 156, 79–113.
ADDITIONAL KEYWORDS: Chaeteessa – evolution – Mantoida – Mesoptilus – Metallyticus – Palaeozoic –
phylogeny – Protorthoptera – Pterygota – wing venation.
INTRODUCTION
As a major component of continental biodiversity in
terms of ecological interactions, species richness, and
biodisparity, winged insects (Pterygota) constitute a
pertinent model for studying biotic macroevolutionary
patterns, and determining the nature of the main
processes that framed the observed extant diversity.
In that respect, the identification of fossil taxa as
stem groups of extant major clades is essential.
However, this identification is difficult to achieve
*Corresponding author. E-mail: [email protected]
for taxa that evolved during (pseudo)radiation events
such as Pterygota, because appropriate outgroups are
unknown or are not readily identified. Attempts to
improve our knowledge of the timing of early events
in Pterygota evolution are impeded by the lack
of well-documented insect faunas older than the
Late Carboniferous, when the group was already
represented by taxa as phylogenetically distant
as Odonatoptera (damselflies, dragonflies, and
stem-relatives) and Archaeorthoptera (grasshoppers,
crickets, and stem-relatives) (Wootton, 1981;
Belayeva et al., 2002; Grimaldi & Engel, 2005). This
lack of data concurs with the difficulties in assigning
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Evidence for Carboniferous origin of the order
Mantodea (Insecta: Dictyoptera) gained from forewing
morphology
80
O. BÉTHOUX and F. WIELAND
MATERIAL AND METHODS
We use the insect wing venation nomenclature
proposed by Lameere (1922, 1923), modified by
Kukalová-Peck (1991), and discussed by Béthoux
(2005c). Following the latter contribution, we consider
the assumption that main vein sectors were distinct
in the pterygotan groundplan (Kukalová-Peck, 1991;
among other contributions) as based on insufficient
evidence. The M5 pattern favoured by Rasnitsyn
(2007; among other contributions) is discussed and
discarded by Béthoux (in press).
Unless specified, vein and vein sector will be
referred to as ‘vein’. Wing venation abbreviations
relevant for this contribution are repeated for convenience: ScP, posterior subcosta; RA, anterior radius;
RP, posterior radius; M, media; CuA, anterior cubitus;
CuP, posterior cubitus; AA, anterior analis; AA1, first
anterior analis; AA2, second anterior analis. The
labels ‘CuA1’ and ‘CuA2’ account for the anterior and
posterior branches of CuA, respectively. The two last
labels do not imply a hypothesis of homology with the
veins CuA1 and CuA2 as usually referred to in the
Grylloblattida sensu Storozhenko (2002). We introduce the abbreviation RP*, accounting for the anterior branch of RP. We follow the vein system colourcoding proposed by Kukalová-Peck & Lawrence
(2004): blue for the radius system (R); red for the
media system (M); green for the cubitus system (Cu).
The term ‘arculus’ refers to a strongly sclerotized
structure, cross-vein like, free of trachea, and located
between two main veins, such as the one occurring
between M and CuA in Plecoptera (see Haas &
Kukalová-Peck, 2001; Béthoux, 2005c). The term
‘translocation’ refers to the fusion of a vein with
another from the base of the latter vein (see Béthoux,
2007c for evidence of this type of transformation in
the fossil record; and below). Veins involved in a
translocation exhibit no evident independent origin.
Specimen preparation follows Béthoux (2005c)
except in the following points. Wings were removed
from specimens by cutting the body cuticle surrounding the wing base using Vanna’s scissors. Unless
specified, right and left wings were mounted on their
ventral and dorsal side, respectively. The generic
acronym IWC OB accounts for Insect Wing Collection
Olivier Béthoux. The acronyms NHM IWC OB and
MNCN IWC OB account for insect wing collections
prepared by one of us and housed at the Entomology
Department, Natural History Museum (London) and
the Entomology Department, Museo Nacional de
Ciencias Naturales (Madrid), respectively. Specimens
referred to as IWC OB only (i.e. without institutional
acronym) belong to the personal collection of one of us
(OB), temporarily housed in the Museum of Zoology,
State Natural History Collections (Dresden). Some
specimens involved in this contribution are housed in
the collection of the Museum of Zoology, State Natural
History Collections (Dresden), in the Oxford University Museum of Natural History (Oxford), and
of the Entomology Department, Muséum National
d’Histoire Naturelle (Paris). These institutions are
referred to as MTD, OUMNH, and MNHN, respectively. For all institutional collections, slides are
housed next to the bodies from which wings were
removed, or information is provided allowing the
rest of the body to be identified. Regarding IWC OB,
bodies are housed either in OB’s or FW’s collection.
Specimens belonging to institutional collections and
mentioned in this contribution are not all illustrated,
but our observations can be checked thanks to specimens numbers. Photographs of these specimens can
be provided on request.
Photographs were taken using a Canon EOS 400D
digital camera coupled with a 50 mm macro lens
and an extension tube, and driven by corresponding
Canon software. High magnification photographs
were taken with a Zeiss AxioCam MRc 5 installed on
a microscope Nikon Eclipse 600, driven by the soft-
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known Late Carboniferous insect species to natural
groups.
This issue resulted in the long-lasting existence of
‘wastebasket groups’, the most important of which is
unarguably the ‘Protorthoptera’ as lately defined by
Carpenter (1992). The ‘Protorthoptera’ ranges from
the Late Carboniferous to the Permian, with fewer
representatives in the Mesozoic. It has recently been
demonstrated that a number of ‘Protorthoptera’
are genuine Archaeorthoptera (Béthoux & Nel, 2002,
2004, 2005; Béthoux, 2005b, 2007a, b; among others).
Béthoux (2005a) suggested that the Archaeorthoptera
experienced an unexpected extinction of moderate
importance at the end of the Carboniferous (but see
Béthoux, 2007a). Therefore, determining the affinities
of the remaining ‘Protorthoptera’ might significantly
modify our understanding of the early steps of Pterygota evolution, as currently recorded.
Hennig (1981) and Carpenter (1992) predicted that
the stem-group of the distinctive praying mantids
(Mantodea) was to be found within the problematic
Palaeozoic ‘Protorthoptera’ group. However, virtually
all recent authors have hypothesized that the origin
of Mantodea was not earlier than the Early Cretaceous or Late Jurassic (with the oldest material being
135 Myr old; Zherikhin, 2002; Grimaldi, 2003; Lo,
2003). We suggest that the former guess is correct,
based on an investigation of the forewing morphology
of extant praying mantids, allowing putative ‘protorthopteran’ relatives of Mantodea to be identified.
Character states relevant for the inner phylogeny of
Mantodea are discussed throughout the paper.
CARBONIFEROUS MANTODEA IDENTIFIED
MORPHOLOGICAL COMPARATIVE ANALYSIS
WING VENATION OF EXTANT MANTODEA
There are two main points of the wing venation of
extant Mantodea to be investigated. The first point
relates to the nature of the vein interpreted as M by
Smart (1956), MP by Ragge (1955), and MA by Sharov
(1962) (hereafter referred to as the composite stem),
and the second being the nature of the vein interpreted as a posterior branch of R (Smart, 1956) or RP
(Rs in Ragge, 1955). Further on we will investigate a
peculiar case relevant for a few genera within Mantodea, as well as other relevant aspects of the forewing venation of Mantodea.
Nature of the composite stem
Typically, the first fork of the composite stem [i.e. the
stem located between the anterior radius sector (RA)
and the most anterior branch of the anterior cubitus
sector (CuA)] is located distally, as observed in
Metallyticus spp. (Figs 2, 6A, 20A; Wieland, 2008:
fig. 14) and other mantids (Ragge, 1955; Smart, 1956;
Ramsay, 1990). Regarding the homologization of this
composite stem, one of the most illuminating forewings we investigated belongs to Metallyticus violaceus
(Burmeister, 1838; Fig. 1). In this atypical wing, the
composite stem forks at the first quarter of the wing.
The resulting anterior branch, which is significantly
stronger than the posterior one, is directed towards
RA, runs parallel to it, and departs from it a few
millimetres further (Fig. 1B). The location of the point
of divergence of this vein from RA is similar to that of
the point of divergence of RP from RA, as observed in
polyneopteran insects such as Orthoptera (grasshoppers & crickets; Ragge, 1955; Sharov, 1971; Béthoux
& Nel, 2002), Plecoptera (stoneflies; Needham &
Claassen, 1925; Béthoux, 2005c), Grylloblattida
(i.e. putative winged fossil relatives of rock-crawlers;
Storozhenko, 1998, 2002), and stem-Dictyoptera
(Bolton, 1922; Laurentiaux, 1958; Laurentiaux &
Laurentiaux-Vieira, 1980), among others. This similarity of position suggests that the anterior branch of
the composite stem, as observed in the atypical forewing of Met. violaceus, is RP.
The posterior branch of the composite stem is unarguably provided with a trachea, as indicated by
annulation (Fig. 1B, C). This trachea runs within
a moderately sclerotized structure, from which
it slightly diverges, and runs towards the anterior
branch of CuA. According to our observations it ends
before reaching CuA. Another narrow trachea
diverges from RP whereas this vein runs side by side
with RA (Fig. 1B, D). Distal to the apparent end
of this narrow trachea, another trachea ‘begins’
(Fig. 1D). The latter runs for a long distance in a
weakly sclerotized structure. It forks after some distance (Fig. 1E), with its anterior branch directed
towards RP (diverging from RA), but the trachea
vanishes before reaching it. The posterior stem of this
trachea runs for some distance in a weakly sclerotized
structure and vanishes in the area between RP and
the anterior branch of CuA.
The apparent end of the second trachea diverging
from RP (Fig. 1B, D) is likely to be an artefact because
there is no other putative origin for the trachea whose
beginning is located nearby. We hypothesize that the
trachea, for some reason, was damaged at this point.
Although large insect species can have very narrow
tracheae or lacunae occurring in cross-vein-like
structures (e.g. in some phasmids and mantids, O.
Béthoux, pers. observ.), this is not the case in Metallyticus spp. In a single case we observed a narrow
annulated trachea inside a cross-vein located between
the posterior branch of CuA, and CuP (Fig. 1F). We
still cannot rule out that the trachea is a vestigial
posterior branch of CuA. In the left forewing of the
same specimen we observed no lacunae or tracheae
inside cross-veins located in the area under scrutiny.
In the atypical forewing of Met. violaceus, distal to
the divergence of RP from RA, no trachea is present
inside the cross-veins occurring between RP and CuA.
This absence stands despite their strength and their
orientation similar to that of actual RP branches.
Therefore, despite the narrowness of the posterior
branches of the composite stem, there is no reason
to consider that they belong to a tracheal network of
secondary importance.
The alternative interpretation is that this set of
tracheae belongs to a vein system distinct from RP,
with which RP could have been fused at the wing
base. Such a system located between RP and CuA
could only be M (CuA is consistently identified by
all previous researchers). If so, M emits a posterior
branch when RP diverges from its regular course
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ware AxioVision 4.4. Original photographs were processed using Adobe Photoshop 7.0. When possible,
broken fragments of wings were re-adjusted. Dust
and remains not belonging to the wings were manually erased from photographs using the clone stamp
tool. We offer to provide original files on request
(provided that a CD is sent to the corresponding
author).
Lastly, we follow Donoghue’s (2005) recommendation for naming fossil groups with respect to extant
taxa (i.e. stem-, crown-, and total-groups; see references therein). The Linnaean taxonomic system will
be followed for convenience, because no alternative
system has ever been applied to the Mantodea. By
convention, we use the term Blattodea for the group
including cockroaches and termites.
81
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O. BÉTHOUX and F. WIELAND
(Fig. 1B, C), while an anterior branch keeps fused
with RP. This anterior branch diverges from RP after
some distance (Fig. 1B) and forks (Fig. 1B, E). Under
this interpretation M is three-branched, with a simple
posterior stem and a forked anterior stem. The first
posterior stem is directed towards CuA, the posterior
branch of the anterior stem vanishes between RP
and CuA, and the anterior-most branch is directed
towards RP.
Another specimen belonging to Met. violaceus
exhibits a different atypical morphology consistent
with our interpretation (Fig. 2). In the first fifth of the
wing, a supernumerary trachea occurs between RA
and M (or composite stem; Fig. 2B). It diverges from
RA, deviates towards M, and becomes indiscernible
from M. At the second fifth of the wing a similar
trachea diverges from M, runs parallel to it, and
re-fuses with it (Fig. 2C). Supernumerary narrower
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Figure 1. Forewing venation of Metallyticus violaceus (Burmeister, 1838), atypical morphology (MNHN, Paris, 씸; right
forewing, dorsal view; abbreviations: ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus; CuP, posterior
cubitus; AA, anterior analis). A, photograph. B, detail of the wing venation as located on A. C–F, detail of the wing
venation as located on B; arrows indicate various tracheae (see text for explanations).
CARBONIFEROUS MANTODEA IDENTIFIED
83
tracheae occur between RA and M at the wing base
(Fig. 2B). A plausible interpretation is that the supernumerary main trachea is RP, or a part of it. Anyhow,
the occurrence of a supernumerary trachea suggests
that the vein occurring between RA and the anterior
branch of CuA is composite, as seen in the usual
morphology exhibited by Metallyticus spp. and other
mantids. The most plausible interpretation is that it
is composed of RP and of M.
We collected additional evidence of the fusion of (a
part of) RP with M in Mantodea. The actual fusion of
(a part of) RP with M, occurring at the wing base, was
observed in two specimens belonging to Creobroter sp.
In the specimen IWC OB 126 (Fig. 3A, B), observation
of the ventral side of the right forewing revealed that
the fusion of RP with M is not achieved at the wing
base, but distal to the divergence of CuA and CuP.
The course of RP is distinct from the sclerotized
structure in which M runs (Fig. 3A), allowing RP and
M tracheae to be distinguished and identified after
their annulated structure (Fig. 3B). This atypical
organization was also observed on the left forewing of
the same individual. In the left forewing of the specimen IWC OB 144 (Creobroter sp.), RP and M run
parallel for a short distance before they fuse (Fig. 3C,
D). Although less obvious, this was also observed on
the right forewing of the same individual. In both
cases M is visible more distinctly from the ventral
side.
A more complicated uncommon organization was
observed in the left forewing of the specimen IWC OB
249, belonging to Hymenopus coronatus (Olivier,
1792) (Fig. 3E, F). Viewed from the ventral side, the
median trachea is located below another trachea
emerging from R (i.e. RP, or a part of it). These
tracheae intersect without fusing. As a result, the
posterior radial trachea (or a part of it) takes a
position posterior with respect to that of the median
trachea. After a short distance, both tracheae fuse
into the composite stem.
The typical organization of this area in praying
mantids involves a posterior bulge of the sclerotized
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Figure 2. Forewing venation of Metallyticus violaceus (Burmeister, 1838), atypical morphology (NHM IWC OB 22,
London; left forewing, ventral view; abbreviations: ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus;
CuP, posterior cubitus; AA, anterior analis; M, media). A, photograph. B–C, detail of the wing venation as located on A;
arrows without labels indicate a supernumerary tracheae.
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O. BÉTHOUX and F. WIELAND
structure in which the radial trachea runs, with M
(and its trachea) running underneath this bulge. We
hypothesize that the fusion of RP (or a part of it) with
M occurs in this area. In Hy. coronatus and Deroplatys desiccata Westwood, 1839 the posterior bulge of
the sclerotized structure in which R runs cannot be
clearly distinguished from the sclerotized structure
in which the composite stem runs (Fig. 3G, H,
respectively).
Our interpretation that RP would be fused with M
from the wing base in all recent Mantodea implies a
translocation. This sort of transformation of the wing
venation pattern has been evidenced in the fossil group
Pantcholmanvissiida (Béthoux, 2007c). We collected
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Figure 3. Evidence for a fusion of posterior radius (RP) with media (M) at the wing base in Mantodea (abbreviations:
ScP, posterior subcosta; RA, anterior radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). A,
Creobroter sp., detail of forewing base (IWC OB 126, 씹; right forewing, ventral view, reversed); arrows without label
indicate the course of the median trachea. B, detail of forewing base, as located on A; arrows indicate the course of the
median trachea. C, Creobroter sp., detail of forewing base (IWC OB 144, 씸; left forewing, ventral view). D, detail of
forewing base, as located on C. E, Hymenopus coronatus (Olivier, 1792), detail of forewing base (IWC OB 249, 씸; left
forewing, ventral view). F, detail of forewing, base as located on E. G, Hymenopus coronatus (Olivier, 1792), detail of
forewing base (IWC OB 238, 씹; right forewing, dorsal view). H, Deroplatys desiccata Westwood, 1839, detail of forewing
base (IWC OB 155, 씹; right forewing, ventral view, reversed).
CARBONIFEROUS MANTODEA IDENTIFIED
85
additional evidence that such a transformation is
plausible. In the undetermined Mantinae specimen
MNCN IWC OB 27 (Fig. 4), the left forewing exhibits
a typical morphology with RP + M distinct from RA
(Fig. 4A), whereas R and M form a single stem in the
right forewing (Fig. 4B, C). Near the wing base, a
narrow trachea that might be the median one fuses
with R (Fig. 4C). This atypical case illustrates that
complete fusion of a vein with another one, from the
wing base (i.e. ‘translocation’ as defined above), occurs
in the wild. The atypical morphology exhibited by
forewings of the specimen NHM IWC OB 40 (Miomantis paykullii Stål, 1871; right forewing illustrated on
Fig. 5E; typical morphology of the species illustrated
on Fig. 5A, C) might be considered as an ‘intermediate’
stage, in which RP + M is fused for a long distance with
RA (and see below). This case suggests that the apparent suddenness of translocations might be the result of
incomplete documentation.
Nature of the posterior stem diverging from RA
The nature of the vein interpreted as a posterior
branch of R (Smart, 1956), or as RP (Rs in Ragge,
1955), generally occurring in Mantodea (indicated
by * on Figs 5A, C–E, 7–17), must then be discussed.
Importantly, it is absent in Metallyticus spp. (Figs 1,
2, 6A, 20A), Chaeteessa spp. (see Smart, 1956;
Figs 6C, 20B), Mantoida spp. (see Smart, 1956: fig. 3,
contra the corresponding figure caption, inverted with
fig. 4; Figs 6E, 20C), †Lithophotina floccosa Cockerell,
1908 (see Sharov, 1962), and †Arvernineura insignis
Nel & Roy, 1996 (for additional fossil taxa, see Nel &
Roy, 1996; Grimaldi, 2003), all considered as plesiotypic taxa within Mantodea. Whether this branch
belongs to RA or is an anterior branch of RP + M
translocated onto RA is to be determined. This vein
will be referred to as vein* in the discussion, later on
as RP*.
In the first instance this vein is emitted posteriorly
(i.e. the anterior stem is aligned to the parent stem
whereas the posterior stem is oblique), which would
be unusual for a branch belonging to RA. In Metallyticus spp. (Figs 1, 2, 6A, 20A), Chaeteessa spp.
(Figs 6C, 20B), †L. floccosa and †A. insignis, branches
emitted by RA are directed anteriorly, i.e. the
posterior branch resulting from a fork is aligned with
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Figure 4. Wing venation of an undetermined Mantinae (MNCN IWC OB 27, 씹; abbreviations: ScP, posterior subcosta; RA,
anterior radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RP, posterior radius; M, media). A, left
forewing (ventral view), typical morphology. B, right forewing (ventral view, reversed), atypical morphology. C, detail of the
wing base, as located on B; arrow indicates a narrow trachea directed towards radius (R), interpreted as the median trachea.
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the parent-stem. This is the typical organization of
RA in polyneopteran orders. This peculiar orientation
of the vein* probably prompted Ragge (1955) to consider that it belongs to RP. We follow this view but
argue that this branch is an anterior stem of RP
secondarily translocated from RP + M onto RA. There
are two main lines of evidence supporting our view.
First, we documented ‘intermediate’ steps between
the states ‘anterior branch of RP + M distinct from
RA’ (as observed in Metallyticus spp. and Chaeteessa
spp.), and ‘anterior branch of RP + M fused with RA
from the wing base and diverging near wing apex’
(as observed in most extant praying mantids). In
addition, individuals exhibiting atypical morphologies
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Figure 5. Wing venation of Miomantis paykullii Stål, 1871 (씹; abbreviations: ScP, posterior subcosta; RA, anterior
radius; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RP, posterior radius; M, media). A, B, specimen
NHM IWC OB 38. A, left forewing (ventral view), typical morphology. B, right forewing (dorsal view), atypical morphology.
C, D, specimen NHM IWC OB 39. C, left forewing (ventral view), typical morphology. D, right forewing (dorsal view),
atypical morphology. E, specimen NHM IWC OB 40, right forewing (dorsal view), atypical morphology.
CARBONIFEROUS MANTODEA IDENTIFIED
87
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Figure 6. Wing venation of praying mantids exhibiting no vein*. A, Metallyticus splendidus Westwood, 1835 (IWC OB
192, 씸; left forewing, ventral view; abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus;
AA, anterior analis). B, detail of the area between anterior radius (RA) and posterior radius + media (RP + M), as located
on A; arrows without labels indicate short sclerotized structures located between RA and an irregularly sclerotized
structure, itself located between RA and RP + M. C, Chaeteessa sp. (MTD, Dresden; right forewing, dorsal view). D, detail
of the area between RA and RP + M, as located on C. E, Mantoida maya Saussure & Zehntner, 1894 (specimen IWC OB
102, 씹; left forewing, dorsal view, reversed). F, detail of the area between RA and RP + M, as located on E; arrow indicates
an isolated cross-vein.
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O. BÉTHOUX and F. WIELAND
suggesting that the vein* is homologous to a branch of
RP + M recur.
In forewings of Chaeteessa sp. (Figs 6C, 20B) and of
Mantoida maya Saussure & Zehntner, 1894 (Figs 6E,
20C) the course of the anterior branch of RP + M is
marked by an inflexion point located opposite the
origin of the vein* as in praying mantids considered
as more derived. The inflexion point and its location
constitute the first evidence of homology between the
two structures (similar course and location). Notably,
the anterior branch of RP + M is simple in these
species.
In a specimen belonging to Amorphoscelis sp. (IWC
OB 243; Fig. 7), the anterior branch of RP + M is
forked opposite of its point of inflexion. In the left
forewing (Fig. 7A), both branches remerge into a
single one, whereas in the right forewing the
anterior branch vanishes in the wing membrane
(Fig. 7B). A similar fork of the anterior branch of
RP + M was observed in the specimen IWC OB 190,
belonging to Amorphoscelis sp. (Fig. 8). However,
unlike in the specimen IWC OB 243, comparison of
the left and right forewings reveals that the anterior
branch of RP + M, or the anterior branch of the latter,
fuses with RA in the specimen IWC OB 190. In the
left forewing (Fig. 8A, B) the anterior branch of
RP + M forks, its anterior trachea fuses with RA,
diverges from it after some distance, and re-fuses
with the posterior branch of the anterior branch
of RP + M (indicated by ° on Fig. 8B). Although we
could not observe the course of the trachea along RA,
we hypothesize that the trachea fusing with RA and
the one emerging from RA after some distance are of
the same origin. Areas surrounding the free parts of
the trachea fusing with RA are weakly sclerotized.
This is unlike the posterior branch of the anterior
branch of RP + M, around which sclerotization is continuous. In the right forewing of the same specimen
(Fig. 8C–F), the sclerotization surrounding the anterior branch of RP + M stops at some point, and the
corresponding trachea fuses with RA without forking
(Fig. 8E). The course of the posterior branch of the
anterior branch of RP + M as in the left forewing is
only materialized by a line of darker setae and is free
of trachea (Fig. 8D, E). After some distance a posterior trachea is emitted from RA (Fig. 8D, F). Interestingly, this trachea is distinct from a weakly
sclerotized structure whose origin is proximal to that
of the trachea, but later joins it. A strongly sclerotized
area surrounds this trachea when it reaches the location of the posterior branch of the anterior branch of
RP + M as demonstrated in the left forewing (indicated by ° on Fig. 8D). Another trachea later diverges
from RA and joins the sclerotized structure just mentioned (Fig. 8D, G), implying that the anterior branch
of RP + M is forked while being fused with RA.
In Amorphoscelis pulchra Bolivar, 1908 the actual
course of the anterior branch of RP + M, whether
diverging from RP + M, partly fused with RA, or
seemingly diverging from RA, can only be determined
after the course of the relevant tracheae. In the right
forewing of the specimen NHM IWC OB 23 (Fig. 9A,
C), the pattern of sclerotized structures suggests that
vein* diverges from RP + M (Fig. 9A). This is contra-
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Figure 7. Wing venation of Amorphoscelis sp. (IWC OB 243, 씹; abbreviations: ScP, posterior subcosta; CuA, anterior
cubitus; CuP, posterior cubitus; AA, anterior analis). A, left forewing (ventral view); left arrow indicates the fork of the
anterior branch of posterior radius + media (RP + M), right arrow indicates the branching of RP + M and the fusion of the
resulting branches. B, right forewing (dorsal view); arrow indicates the first fork of the anterior branch of RP + M.
CARBONIFEROUS MANTODEA IDENTIFIED
89
dicted by the tracheal pattern, which is provided with
a broad trachea diverging from RA, whereas a narrower trachea diverges from RP + M and directs
towards vein*. In the left forewing of the same individual, the origin of vein* from RA is more evident
based on macroscopic observation (Fig. 9B, D),
although a trachea of secondary importance diverges
from RP + M and fuses with vein*. In the left forewing of the specimen NHM IWC OB 24, the origin of
vein* is ambiguous based on macroscopic and micro-
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Figure 8. Wing venation of Amorphoscelis sp. (IWC OB 190, 씹; abbreviations: ScP, posterior subcosta; CuA, anterior
cubitus; CuP, posterior cubitus; AA, anterior analis). A, left forewing (ventral view). B, detail of the area between anterior
radius (RA) and posterior radius + media (RP + M), as located on A; left arrow indicates the anterior branch of the anterior
branch of RP + M; right arrow indicates the anterior branch of the anterior branch of RP + M emerging from RA;
° indicates the posterior branch of the anterior branch of RP + M. C, left forewing (dorsal view). D, detail of the area
between RA and RP + M, as located on C; ° indicates the posterior branch of the anterior branch of RP + M. E, detail of
the fusion of the anterior branch of RP + M with RA, as located on D; arrow indicates the RP + M trachea. F, detail of
the divergence of the posterior branch of the anterior branch of RP + M from RA, as located on D; arrow indicates the
trachea of the posterior branch of RP + M. G, detail of the divergence of the anterior branch of the anterior branch of
RP + M from RA, as located on D; arrow indicates the trachea of the anterior branch of RP + M.
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O. BÉTHOUX and F. WIELAND
scopic observations (Fig. 9E, G): a network of narrow
tracheae occurs between RA and RP + M. In the right
forewing of the same specimen the origin of vein*
is ambiguous based on macroscopic observations
(Fig. 9F), whereas it apparently diverges from RP + M
based on the tracheal pattern (Fig. 9H).
In summary, a fork of the anterior branch of
RP + M occurs in several Amorphoscelidae [the mono-
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Figure 9. Wing venation of Amorphoscelis pulchra Bolivar, 1908; in C, D, G, H, thickness of arrows indicates strength
of indicated tracheae (abbreviations: ScP, posterior subcosta; CuP, posterior cubitus; AA, anterior analis). A–D, specimen
NHM IWC OB 23. A, left forewing (ventral view). B, right forewing (dorsal view). C, detail of the connection of the anterior
branch of posterior radius + media (RP + M) with anterior radius (RA), as located on A. D, detail of the connection of the
anterior branch of RP + M with RA, as located on B. E–H, specimen NHM IWC OB 24. E–F, arrows without label indicate
the origin of a branch of anterior cubitus (CuA). E, left forewing (ventral view). F, right forewing (dorsal view). G, detail
of the connection of the anterior branch of RP + M with RA, as located on E. H, detail of the connection of the anterior
branch of RP + M with RA, as located on F.
CARBONIFEROUS MANTODEA IDENTIFIED
branch of CuA diverges from RP + M, or from CuA, in
the left and right forewing, respectively (Fig. 10A and
B, respectively; origin of this branch indicated by an
arrow).
In the specimen MNCN IWC OB 31, assigned to
Oxypilinae, both forewings exhibit a vein* diverging
from RA (Fig. 11A, B). In the right forewing, RP + M
is provided with an anterior stem fusing with RA
(Fig. 11A, C), whereas RP + M is simple in this area
in the left forewing (Fig. 11B, D). It suggests that the
fusion of vein* with RA is incomplete in the right
forewing and complete in the left forewing. It must be
noted that the anterior-most branch of CuA diverges
from CuA in the right forewing, and from M in the left
forewing.
In the specimen IWC OB 141, belonging to Creobroter sp., vein* originates from RA in the left forewing (Fig. 12A, B), whereas it originates from RP + M
in the right forewing (homology is ascertained by a
similar location and a similar number of branches;
Fig. 12C, D). This pattern is reminiscent of that
observed in Metallyticus spp. (Figs 1, 2, 6A, 20A).
Some cases are less evident and do not allow the
homology statement we propose to be positively ascertained. It is, however, still worth mentioning them.
In the specimen NHM IWC OB 38, belonging to Mi.
paykullii, the left forewing exhibits a vein* diverging
from RA and a three-branched RP + M (Fig. 5A). The
Figure 10. Wing venation of an undetermined Mantinae, atypical morphology (MNCN IWC OB 28; abbreviations: ScP,
posterior subcosta; CuP, posterior cubitus; AA, anterior analis). A, left forewing (dorsal view); arrow without label indicates
the origin of a branch of anterior cubitus (CuA) from posterior radius + media (RP + M). B, right forewing (ventral view);
arrow without label indicates the origin of the same branch of CuA as in A. C, detail of the area between the anterior branch
of RP + M and anterior radius (RA) as located on A; arrows indicate inflexion points of the anterior branch of the
supernumerary branch of RP + M (see text). D, detail of the area between the anterior branch of RP + M and RA as located
on B; arrows indicate the points of fusion and divergence of the supernumerary branch of RP + M with RA.
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phyly of which is uncertain (e.g. Handlirsch, 1925;
Chopard, 1949a; Wieland, 2003), therefore they will
be referred to as amorphoscelidaeans hereafter]. The
anterior branch resulting from this fork approximates
RA, is connected to it via a network of tracheae, or is
almost fully fused with it. It can then be reasonably
hypothesized that a gradual reduction of the tracheal
network could have resulted in a definitive and complete fusion of the anterior branch of RP + M with RA,
from the wing base. Atypical morphologies exhibited
by derived taxa provide additional support to this
hypothesis.
In the specimen MNCN IWC OB 28, belonging to
an undetermined Mantinae, none of the forewings
exhibit a vein diverging from RA (Fig. 10). However,
the composite stem RP + M is provided with a supernumerary simple anterior stem absent in related
taxa (in Mantinae RP + M is typically posteriorly
pectinate; see Fig. 4A; see also Fig. 18, relevant for
Angelinae). The absence of co-occurrence of the
supernumerary stem and of vein* suggests that these
structures are homologous. In addition, the supernumerary branch runs parallel to RA for some distance
in the left forewing (Fig. 10A, C), whereas it is fused
with RA for some distance in the right forewing
(Fig. 10B, D). This is clearly reminiscent of the
morphology observed in the genus Amorphoscelis
Stål, 1871. It must be noted that the anterior-most
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O. BÉTHOUX and F. WIELAND
right forewing of the same specimen exhibits no vein
diverging from RA, and a two-branched RP + M
(Fig. 5B). It cannot be excluded that vein* is fully
fused with RA in the right forewing of this specimen.
In the specimen IWC OB 237, belonging to Deroplatys lobata (Guérin-Méneville, 1838), the left forewing is provided with a vein* diverging from RA,
and RP + M is two-branched (typical morphology;
Fig. 12E). This is different from the right forewing,
which has a simple RA and a three-branched RP + M
(Fig. 12F). This case would imply that vein* is
homologous to the anterior branch of the anterior
branch of RP + M, rather than to the anterior branch
of RP + M (as above). However, it could be hypothesized that, in the right forewing of the specimen IWC
OB 237, vein* is fully fused with RA, and that RP + M
produces a supernumerary branch in response to this
atypical transformation. Such ‘compensation’ is well
known in cockroach forewings, in which a reduced
number of branches from one vein can be compensated by a higher number of branches of a neighbouring vein. As a result, the ‘density’ of main
vein branches is preserved (see Schneider, 1977;
Laurentiaux-Vieira & Laurentiaux, 1980; among
others; also compare the number of branches of
RP + M and of CuA on Fig. 18A and B, respectively).
The specimen NHM IWC OB 36, belonging to Statilia maculata (Thunberg, 1784), exhibits an atypical
morphology occurring in both forewings, in which no
stem diverges from RA, and RP + M is forked (see
Fig. 13B, right forewing). In the typical morphology
of the species, a vein* diverges from RA, whereas
RP + M is simple (Fig. 13A). A compensation cannot
be completely ruled out in the specimen NHM IWC
OB 36. The similar location of the divergence of vein*
from RA in the typical morphology, and of the fork of
RP + M in the atypical morphology, however, suggest
that the typical vein* is homologous to the atypical
supernumerary anterior branch of RP + M.
In the specimen NHM IWC OB 33, belonging to
Amantis reticulata (De Haan, 1842), we observed that
the left forewing has a branch diverging posteriorly
from RA and a forked RP + M. The right forewing has
a three-branched RP + M. However, the respective
location of these branches does not allow a homology
statement to be positively ascertained.
In summary, the recurrence of atypical patterns
suggesting that vein* as exhibited by most mantids
is homologous to the anterior branch of RP + M as
exhibited by Metallyticus spp. (Figs 1, 2, 6A, 20A) and
Chaeteessa spp. (Smart, 1956; Figs 6C, 20B), among
others, strongly supports the conclusion drawn from
the investigation of the amorphoscelidaean wing
venation. Although a full fusion of vein* with RA,
resulting in a supernumerary compensatory branch of
RP + M, could explain some cases of the atypical
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Figure 11. Wing venation of an undetermined Oxypilinae sp. (MNCN IWC OB 31, 씹; abbreviations: ScP, posterior
subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis). A, right forewing (dorsal view). B, left forewing
(ventral view). C, detail of the connection of the anterior branch of posterior radius + media (RP + M) with anterior radius
(RA), as located on A. D, detail of the connection of the anterior branch of RP + M with RA, as located on B.
CARBONIFEROUS MANTODEA IDENTIFIED
93
morphologies we document, the most parsimonious
explanation, with respect to the transition sequence
documented in amorphoscelidaeans, is to consider
these atypical morphologies as representing rare
occurrences of a plesiomorphic condition. It is
unlikely that the median vein takes part in the vein*,
therefore we hereafter refer to it as RP*.
A further step in the homologization of RP* concerns its number of branches. We observed a number
of specimens exhibiting an atypical morphology suggesting that an anterior branch of RP* runs fused
with RA. These specimens exhibit a fork of RP* near
its point of divergence from RA (Fig. 14). The resulting anterior branch fuses with RA in the left forewing
of the specimen IWC OB 242 (Parasphendale affinis
Giglio-Tos, 1915; Fig. 14C, D; the right forewing,
exhibiting a typical morphology, is illustrated on
Fig. 14A, B). In the left forewing of the specimen IWC
OB 76 [Ameles decolor (Charpentier, 1825); Fig. 14F],
it runs parallel to RA (the right forewing of the
specimen IWC OB 76, exhibiting the typical morphology, is illustrated on Fig. 14E). We interpret the atypical morphology of the right forewing of the specimen
IWC OB 71 (A. decolor; Fig. 14G) in a similar way:
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Figure 12. Intra-individual variations in the origin of the anterior branch of posterior radius (RP*); on B, D, arrows
indicate the origin of RP* (abbreviations: ScP, posterior subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA,
anterior analis; M, media; RA, anterior radius). A–D, forewings of Creobroter sp. (IWC OB 141, 씸). A, left forewing
(ventral view). B, detail of left forewing apex. C, right forewing, atypical morphology (dorsal view). D, detail of right
forewing apex. E–F, forewings of Deroplatys lobata (Guérin-Méneville, 1838) (IWC OB 237, 씹). E, left forewing (ventral
view). F, right forewing, atypical morphology (dorsal view).
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O. BÉTHOUX and F. WIELAND
RP* apparently emits two successive branches (see
also Fig. 5C, D), although an additional translocation
of a branch of RP + M onto RA cannot be completely
ruled out in this case. A fork of RP* was observed in
Hierodula sp. (right forewing; MNCN IWC 30), in
Omomantis zebrata (Charpentier, 1843) (observed on
a single specimen housed at the NHM), and Polyspilota sp. (single specimen housed at the MNCN).
Therefore, we hypothesize that RP* is usually forked,
and that its anterior branch is usually fused with RA.
The compelling evidence we gathered strongly supports the view that in all praying mantids but
Metallyticus spp., Chaeteessa spp., Mantoida spp.,
some taxa currently assigned to the Amorphoscelidae,
and some fossil taxa, the anterior branch of RP + M
runs fused with RA from the wing base, and diverges
distally from RA (or its posterior stem). We propose
not to differentiate the stem of RP* that keeps fused
with RA, and not to differentiate the stem of RP that
keeps fused with M.
Additional evidence of the RP + M fusion
Now that the fusion of RP* with RA is assessed as a
trait of most praying mantids, and because this fusion
is absent in Metallyticus spp., Chaeteessa spp., and
Mantoida spp., it is worth examining the organization
of the area located between RA and RP + M in the
latter. As mentioned above, the divergence of RA and
RP, from the stem of R, is usually located in the distal
part of the wing in polyneopteran insects. Therefore, a proximal area between RA and RP is a trait
unknown in these insects.
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Figure 13. Wing venation of Statilia maculata (Thunberg, 1784) (씹; abbreviations: ScP, posterior subcosta; CuA,
anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, specimen NHM IWC OB 37, left
forewing (ventral view), typical morphology. B, specimen NHM IWC OB 36, right forewing (dorsal view), atypical
morphology. C, specimen NHM IWC OB 35, left forewing (dorsal view), atypical morphology. D, detail of the connection
of the anterior branch of posterior radius (RP*) with posterior radius + media (RP + M), as located on C.
CARBONIFEROUS MANTODEA IDENTIFIED
95
In Metallyticus spp. (Fig. 6A, B), short sclerotized
structures connect RA to an irregularly sclerotized
structure. This structure is free of trachea, parallel
to both RA and RP + M, and we interpret it as the
‘remnant’ of the sclerotized structure in which the RP
trachea ran before becoming completely fused with M.
Along its course, the composite vein RP + M actually
switches from this structure to the ‘regular’ one in the
aberrant specimen illustrated on Figure 2 (also in
the specimen IWC OB 126, Fig. 3A). In our opinion,
the occurrence of this ‘remnant’ constitutes by itself
additional evidence of the fusion of RP with M in
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Figure 14. Evidence of the branching of the anterior branch of posterior radius (RP*) (abbreviations: CuA, anterior
cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A–D, forewings of specimen IWC OB 242,
belonging to Parasphendale affinis Giglio-Tos, 1915 (씸). A, right forewing (dorsal view). B, detail of right forewing apex,
as located on A. C, left forewing (dorsal view, reversed). D, detail of left forewing apex, as located on B; arrows indicate
the two branches of RP*. E–G, forewings of specimens belonging to Ameles decolor (Charpentier, 1825). E, right forewing
of the specimen IWC OB 76 (dorsal view; 씹). F, left forewing of the specimen IWC OB 76 (ventral view; 씹). G, right
forewing of the specimen IWC OB 71 (dorsal view; 씹).
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O. BÉTHOUX and F. WIELAND
Metallyticus spp. In this taxon regular cross-veins
occur between RA and RP + M distal to a distinctive
point of inflexion of RP + M (Figs 1A, 2A, 6A, 20A).
We interpret this point of inflexion as homologous to
the point of divergence of RP from R as in other
polyneopteran insects (see above).
In the specimens of Chaeteessa spp. and Mantoida
spp. that we could examine (see Fig. 6C, D and
Fig. 6E, F, respectively; and O. Béthoux, pers.
observ.), very few cross-veins occur in this area, if any.
As in Metallyticus spp., regular cross-veins occur
distal to a point of inflexion of RP + M or distal to the
first fork of RP + M. We interpret the absence of
cross-veins in the basal part of the area between RA
and RP + M as evidence of RP being fully fused in the
composite stem. As a result of this fusion, an area
between RA and RP, which does not occur in putative
sister-groups of Mantodea, is present in Chaeteessa
spp. and Mantoida spp., with the absence of crossveins being indicative of this novelty.
Our opinion is supported by the fact that the
absence of cross-veins is more or less correlated with
the absence of fusion of RP* (or the anterior branch
of RP + M) with RA. In amorphoscelidaeans (lacking
this fusion or not), very few or no cross-veins occur
between RA and RP + M basal to the first fork of
RP + M (based on specimens NHM IWC OB 23–26
and IWC OB 190, 243). In Mantodea exhibiting the
fusion of RP* with RA, cross-veins occur in the area
that is posterior to RA + RP* [Fig. 13, Fig. 18 and O.
Béthoux, pers. observ. on Mantis religiosa (Linnaeus,
1758), among other taxa]. In these taxa, however, this
area separates RA + RP* and RP + M, and hence is an
area between branches of RP, not between RA and RP.
It is then consistent that this area has cross-veins,
as is usual in areas separating RP branches in polyneopteran insects (although usually located in a more
distal position).
Secondary fusion of RP* with RP + M
The most derived case in mantodean forewing morphology is probably that exhibited by the species
belonging to Schizocephala Serville, 1831, Parathespis Saussure, 1869, Hoplocorypha Stål, 1871, and
Danuria Stål, 1856 (among those genera investigated
in our survey). In these, we suggest that RP*, diverging from RA + RP*, is secondarily fused with RP + M.
This proposition is supported by several observations.
In the left forewing of the specimen MNCN IWC
OB 24, belonging to Schizocephala bicornis (Linnaeus, 1758), RP* is connected to the single branch of
RP + M via two arculuses (i.e. two strongly sclerotized
structures free of trachea), the respective tracheae
having no direct contact (Fig. 15A, C). In the right
forewing of the same specimen, the RP* trachea and
RP + M trachea share the same sclerotized structure
for some distance and run side by side (Fig. 15B, D).
The degree of connection of RP* with RP + M is
variable in Hoplocorypha nigerica Beier, 1930: RP*
can be distinct from RP + M (Fig. 16A, C), connected
to RP + M via a strong cross-vein (Fig. 16B, D), briefly
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Figure 15. Wing venation of Schizocephala bicornis (Linnaeus, 1758) (MNCN IWC 24, 씹; abbreviations: ScP, posterior
subcosta; CuA, anterior cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, left forewing (ventral
view). B, right forewing (dorsal view). C, detail of the connection of the anterior branch of posterior radius (RP*) with
posterior radius + media (RP + M), as located on A. D, detail of the connection of RP* with RP + M, as located on B.
CARBONIFEROUS MANTODEA IDENTIFIED
97
connected to RP + M (Fig. 16E, G), or exhibit no clear
origin (Fig. 16F, H).
In both forewings of the specimen IWC OB 239
(Fig. 17), assigned to Hoplocorypha sp., a short
oblique, and moderately sclerotized structure occurs.
It possesses a trachea, diverges from RA, and fuses
with RP + M. After some distance, the composite stem
resulting from the fusion of this ‘vein’ with RP + M
forks. Based on the observations mentioned above, we
interpret this organization as RP* fusing with RP + M
for some distance and then diverging. This is arguably a derived state within Mantodea. Interestingly,
this organization occurs as a rare atypical case in
S. maculata (Fig. 13C, D).
Branching pattern of CuA
Another distinctive feature of forewings of most
mantids is the branching pattern of the posterior
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Figure 16. Wing venation of Hoplocorypha nigerica Beier, 1930 (씹; abbreviations: ScP, posterior subcosta; CuA, anterior
cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, C, specimen NHM IWC OB 29, right forewing
(dorsal view). B, D, specimen NHM IWC OB 31, right forewing (dorsal view). C, detail of the origin of the anterior branch
of posterior radius (RP*), as located on A. D, detail of the origin of RP*, as located on B. E–H, specimen NHM IWC OB
30. E, left forewing (ventral view). F, right forewing (dorsal view). G, detail of the origin of RP*, as located on E. H, detail
of the origin of RP*, as located on F.
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O. BÉTHOUX and F. WIELAND
Figure 18. Wing venation of Euchomenella heteroptera (De Haan, 1842) (NHM IWC OB 28; 씹; abbreviations: CuA,
anterior cubitus; CuP, posterior cubitus; AA, anterior analis; M, media; RP, posterior radius); arrows without label indicate
the alternative origin of a branch of anterior cubitus (CuA), presumably homologous in both forewings (apparent fusion
of posterior subcosta (ScP) with anterior radius (RA) is a result of artefacts of preparation). A, left forewing (ventral view).
B, right forewing (dorsal view).
branch of CuA (referred to as CuA2), which is generally anteriorly pectinate (i.e. successive anterior
branches emitted by CuA2 are simple; see Figs 1A,
2A, 4A, B, 6A, 10B, 13A–C, 14A, C, E, F, 15A, B, 16B,
E, F, 18A, 20; but see below). The anterior branch of
CuA (CuA1), when branched, is generally posteriorly
pectinate (Figs 1A, 2A, 4A, B, 6A, 13A–C, 18A). This
branching pattern was noticed by Ragge (1955: 135)
and is referred to as internally pectinate. Whether
this pattern is diagnostic of Mantodea as a whole
must be discussed, as several mantids do not exhibit
it, and it can be variable at the levels of individuals
and species.
In Metallyticus spp., anterior branches of CuA2 can
be branched distally, but typically distal to the fork of
the corresponding posterior ‘sister-branch’ (i.e. anterior branches of CuA/CuA2 are branched distal to
posterior branches of CuA2; Figs 1A, 2A, 6A, 20A; see
scheme in Fig. 19A). This is a constant feature
of Metallyticus spp. forewing venation pattern (see
figures; eight forewings in IWC OB; and 18 observed
forewings of OUMNH, of which 16 exhibit an
anteriorly pectinate CuA2, 2 whereas two exhibit an
ambiguous pattern).
In the specimen NHM IWC OB 28 belonging to
Euchomenella heteroptera (De Haan, 1842), the first
anterior branch of CuA2 in the left forewing
(Fig. 18A) is arguably homologous to the first posterior branch of CuA1 as in the right forewing (Fig. 18B; compare also Fig. 16A, B). Similar
‘switches’ occur relatively commonly in mantids.
Four out of 13 forewings of Ameles decolor (Charpentier, 1825) present in IWC OB and relevant for
comparison exhibit an anteriorly pectinate CuA2 (and
CuA) as schematized in Figure 19B (see Fig. 14E, F).
Conversely, the remaining nine forewings exhibit
a dichotomously branched CuA as schematized in
Figure 19C (see Fig. 14G; a single individual can
exhibit both morphologies). The point of origin of the
branch indicated by ‘c’ in Figure 19B is located opposite the point of origin of the branch indicated by ‘c’
in Figure 19C. It suggests that these branches are
homologous, and that a dichotomously branched CuA
could derive from a posteriorly pectinate CuA.
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Figure 17. Wing venation of specimen IWC OB 239, belonging to Hoplocorypha sp. (씹; abbreviations: CuA, anterior
cubitus; CuP, posterior cubitus; AA, anterior analis; RA, anterior radius). A, left forewing, ventral view. B, detail of the
origin of the anterior branch of posterior radius (RP*), as located on A; arrow without label indicates the origin of (the
posterior branch of) RP*.
CARBONIFEROUS MANTODEA IDENTIFIED
A
C
a
b
c
d
D
a
b
c
d
E
a
b
c
d
a
b
c
d
Figure 19. Possible transition from an anteriorly pectinate branch of the posterior branch of anterior cubitus
(CuA2) to a posteriorly pectinate anterior cubitus (CuA).
A, branching pattern of CuA as in Metallyticus spp. B, one
of the branching patterns of CuA exhibited by Ameles
decolor (Charpentier, 1825). C, most frequent branching
pattern of CuA as in A. decolor (Charpentier, 1825). D,
branching pattern of CuA as in Amorphoscelidae. E,
branching pattern of CuA as in Chaeteessa spp.
In amorphoscelidaeans, CuA is typically dichotomously branched, with a very basal first fork of the
posterior branch, as schematized in Figure 19D (see
Figs 7, 8, 9A, B, E). This morphology could have
equally derived (1) from the pattern presented in
Figure 19A, as a consequence of the reduction of
the number of CuA2 branches; or (2) from the pattern
presented in Figure 19C, as a consequence of a relocation of the point of origin of the ‘a’ and ‘b’ branches,
and the point of origin of the ‘c’ and ‘d’ branches.
Interestingly, in the right forewing of the specimen
NHM IWC OB 24 (Fig. 9F), belonging to A. pulchra,
the branch indicated by ‘b’ in Figure 19D and E
diverges from the stem that gives rise to branches
‘c’ and ‘d’, resulting in the pattern presented in
Figure 19E (whereas the typical morphology of the
species is schematized in Fig. 19D). This pattern
is identical to that exhibited by Chaeteessa spp.
(Figs 6C, 20B; see also Nel & Roy, 1996: fig. 3), in
which CuA as a whole is posteriorly pectinate. From
the available data, in Chaeteessa spp., the second fork
of CuA is consistently closer to the first fork than to
the third fork. This suggests that the branch indicated by ‘b’ in Figure 19E is homologous to the branch
indicated by ‘b’ in Figure 19D, and that the branching
pattern of CuA exhibited by Chaeteessa spp. is likely
to be derived from a dichotomously branched CuA.
In summary, all branching patterns of CuA as
observed in Mantodea could have derived from an
anteriorly pectinate pattern. However, the polarity of
character state transformation is not evident. Ascertaining the position of Metallyticus Westwood, 1835
with respect to other mantids is essential in this
situation. At this step, it can only be considered
as plausible that CuA is anteriorly pectinate in the
groundplan of Mantodea forewing venation.
A similar pattern is present in Cnemidolestodea
(see Béthoux & Nel, 2004, 2005; Béthoux, 2005a,
2007a), but this condition is considered as derived
within Archaeorthoptera, as plesiotypic representatives of the latter taxon exhibit a posteriorly pectinate
CuA (as it is, fused with CuPa) (Béthoux, 2003, 2006;
see also Béthoux, 2005b; Prokop & Ren, 2007).
Regarding Plecoptera, the families Eustheniidae,
Diamphipnoidae, and Austroperlidae have an anteriorly pectinate CuA (see Zwick, 1979; McLellan, 1996;
Béthoux, 2005c). According to the phylogenetic
scheme of Zwick (2000), this is likely to represent the
ancestral state of Antarctoperlaria (in Gripopterygidae CuA is forked or simple, therefore the character
describing the branching pattern is not applicable). In
Euholognatha the character is not applicable as CuA
is forked or simple (Béthoux, 2005c). The ancestral
condition in Systellognatha cannot be ascertained:
whereas Pteronarcyoidea exhibit an anteriorly pectinate CuA, Perloidea exhibit a posteriorly pectinate
CuA (when applicable; see Needham & Claassen,
1925; Béthoux, 2005c). Thus, although an ancestral
plecopteran condition ‘CuA anteriorly pectinate in
forewings’ can be drawn from the available information (and based on the phylogenetic scheme in Zwick,
2000), the robustness of this inference is low. It is
equally plausible that the plecopteran groundplan
involves a simple or forked CuA, hence the character
describing the branching pattern of CuA would be
inapplicable. Regarding putative stem-Dictyoptera
(Bolton, 1922; Laurentiaux, 1958; Laurentiaux &
Laurentiaux-Vieira, 1980) and stem-Blattodea (see
Rehn, 1951; Schneider, 1977, 1978, 1980a, 1980b,
1982, 1983), virtually all of them exhibit a posteriorly
pectinate CuA. Within Blattodea, some taxa such
as the Polyphagidae exhibit an anteriorly pectinate
CuA2 (see Rehn, 1951), but the wing venation of the
members of this family is extremely variable. The
available data on extant taxa (morphology and phy-
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B
99
100
O. BÉTHOUX and F. WIELAND
logeny) do not allow the ancestral Blattodean condition to be ascertained. Anyhow, a posteriorly pectinate
CuA2 could be diagnostic of Mantodea, or of (some)
Dictyoptera.
Summary
A synthesis of our comparative morphological analysis is presented in Figure 20, with a sample of various
mantodean forewings. The left forewing of the specimen IWC OB 193 (Met. splendidus; Fig. 20A) is atypical in that RP + M has two main stems (RP + M is
typically posteriorly pectinate in Metallyticus spp.,
see Figs 1A, 2A, 6A). Branches resulting from the fork
of RP + M might be RP and M diverging distally. If so,
M would be three-branched. This atypical morphology
is presented as a matter of record.
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Figure 20. Homologization of forewing venation in Mantodea (abbreviations: CuA, anterior cubitus; CuP, posterior cubitus;
AA, anterior analis; M, media; RA, anterior radius; RP, posterior radius). A, Metallyticus splendidus Westwood, 1835 (based
on specimen IWC OB 193, left forewing, 씸). B, Chaeteessa filata Burmeister, 1838 (based on specimen described by Smart,
1956, housed at OUMNH). C, Mantoida maya Saussure & Zehntner, 1894, first interpretation (based on specimen IWC OB
97, left forewing, 씹). D, Tarachodula pantherina (Gerstaecker, 1869) (based on specimen IWC OB 96, left forewing, 씹).
CARBONIFEROUS MANTODEA IDENTIFIED
FOSSIL MATERIAL
This section is based on literature data (see below)
and new investigations. We reinvestigated material
assigned to †Mesoptilus dolloi Lameere, 1917, a Late
Carboniferous species yielded by the famous French
deposit of Commentry (Lameere, 1917; Fig. 21;
c. 310 Mya). The holotype specimen MNHN DP
R51159 (Fig. 21) and the specimen MNHN A27122
(Fig. 22) were observed. Although the wings are
poorly preserved on the specimen MNHN A27122, it
is assigned to the same species as MNHN DP R51159
based on: the similar size; identical wing coloration;
similarly long, oblique, and numerous branches of
ScP; similar (fore-)leg morphology. A new restoration of
the wing venation of †Mes. dolloi is provided on
Figure 21C. The area where the origin of RP (diverging
from R) is likely to have occurred is poorly preserved
on the holotype, and is not visible on the specimen
MNHN A27122. The location of the origin of RP is
inferred from related taxa (see below). We also consider
the leg morphology of the specimen MCZ 5875,
paratype of †Homocladus grandis Carpenter, 1966.
Forewing morphology
†Mes. dolloi is affiliated with a number of Palaeozoic
genera such as †Strephoneura Martynov, 1940, †Spargoptilon Kukalová, 1965, †Homocladus Carpenter,
1966, †Paracladus Carpenter, 1966, and †Graticladus
Novokshonov & Aristov, 2004. Most of these genera
have been assigned to the family †Strephocladidae by
Carpenter (1992). As the monophyly of this family is
uncertain (see below), the corresponding species will
be referred to hereafter as strephocladidaeans. All of
them exhibit an anteriorly pectinate CuA or CuA2,
suggesting that they have dictyopteran or mantodean
affinities (see above). Carpenter (1966) noted the difficulty in identifying the median system in these taxa.
In most of them, at least a branch of M is connected
to RP, and it cannot be ruled out that some anterior
branches of CuA fuse with M for some distance.
However, in his interpretation of the wing venation of
†Ho. grandis, Carpenter (1966) consistently identified
M as a system constituted by a forked anterior branch
and a simple posterior branch. In †Graticladus
severus Novokshonov & Aristov, 2004, in which there
is no connection of M with RP, M is also composed of
a forked anterior branch and a simple posterior
branch, just as in the atypical forewing of Met. violaceus (Fig. 1). We hypothesize that this is also the
case in †Mes. dolloi. If so, in this taxon, the most
anterior stem of M is fused with RP, the second stem
of M (the first posterior branch of the anterior branch)
runs free, and the posterior stem of M is fused for
some distance with the anterior branch of CuA (such
a fusion of the anterior branch of CuA with M has
been observed in extant Mantodea, see Fig. 10A).
Whether the anterior branch of CuA is fused with M
from the wing base or fuses with it later, via an
arculus (as indicated on Fig. 21C), cannot be determined with certainty.
Although the location of the origin of RP and the
exact number of branches belonging to M in †Mes.
dolloi could be debated, the anterior-most branch of
M unarguably fuses for some distance with RP. This
is evidenced by the long distance separating the point
of divergence of the anterior branch of M from more
distal branches (as it is, of RP). It is noteworthy that
the location of actual RP branches is similar to that
observed in forewings of Metallyticus spp. This is also
the case in †Ho. grandis and †Homocladus ornatus
Carpenter, 1966 (see Carpenter, 1966; figs 4, 5,
respectively).
An incomplete fusion of M with RP in forewings
was also acquired within Archaeorthoptera (= totalOrthoptera) (see Béthoux & Nel, 2003, 2004, 2005),
but the most likely plesiotypic members of this taxon
lack this fusion (Béthoux, 2003, 2006). It is absent in
forewings of Plecoptera (Needham & Claassen, 1925;
Béthoux, 2005c), occurs sporadically in Grylloblattida
in which it is likely to be a derived character state
(see Storozhenko, 2002: fig. 393, nodes 30, 64; atypical morphologies exhibiting a fusion of an anterior
branch of M with R occur sporadically, see Tillyard,
1928a: fig. 14; Tillyard, 1928b: fig. 5), and is absent in
stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958;
Laurentiaux & Laurentiaux-Vieira, 1980), among
others. Therefore, coupled with the branching pattern
of CuA reminiscent of that observed in Metallyticus
spp., the character state ‘anterior branch of M fused
with RP for a long distance’ is strong evidence for
considering †Mes. dolloi and its relatives as stemMantodea.
Leg morphology
Several legs are preserved on the holotype of †Mes.
dolloi, but none can positively be identified as a
foreleg from their respective position, because of the
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In conclusion, crown-Mantodea are characterized
by the following forewing character state: RP and M
fused from the wing base (as opposed to: M and RP
distinct). A putative diagnostic character state is: CuA
internally pectinate (hence CuA2 anteriorly pectinate;
as opposed to: posterior branch CuA simple, or
without consistent branching pattern). Within Mantodea, plausible plesiomorphic character states of the
forewing morphology are as follows (mainly based
upon the morphology exhibited by Metallyticus spp.,
typical and atypical, Chaeteessa spp., and fossil taxa
mentioned above): RA with anterior branches; and M
branched.
101
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O. BÉTHOUX and F. WIELAND
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Figure 21. Mesoptilus dolloi Lameere, 1917 (holotype specimen MNHN DP R51159; abbreviations: CuA, anterior
cubitus; CuP, posterior cubitus; AA, anterior analis; M, media; R, radius; RA, anterior radius; RP, posterior radius). A,
habitus (under ethanol, polarized light). B, detail of a leg, as located on A (dry-ethanol composite photograph); arrows
indicate remains of spines. C, drawing of the right forewing. D, right forewing (negative imprint, reversed; dry-ethanol
composite photograph, polarized light).
CARBONIFEROUS MANTODEA IDENTIFIED
103
disarticulation of the individual (Fig. 21A). However,
one leg preserves evidence of stout spines located
on the femur (Fig. 21B). They are incompletely
preserved as they are broken off along the line of
splitting (the opposite imprint of the specimen is
unfortunately not available). Body parts are also
partly disconnected in the specimen MNHN A27122
(Fig. 22A). In this specimen, however, stout spines
with a bent acron and a broad base are well preserved on the femur of one leg, and two spines are
preserved on the tibia (Fig. 22B). The leg is overall
poorly preserved (the opposite imprint of the specimen is not available). The shape of the spines
located on the femur as observed on the specimen
MNHN A27122 are indicative of a raptorial leg,
most probably a foreleg. Connection with the morphology exhibited by extant praying mantids is
made difficult by the poor preservation of the fossil
material, but it is likely that †Mes. dolloi had raptorial forelegs.
Regarding †Ho. grandis, Carpenter (1966: 62) mentioned that an incomplete leg, with two rows of heavy
spines on the tibia and smaller spines on the femur, is
preserved next to the wing of the paratype. This leg is
likely to belong to the same individual (Fig. 23A).
Examination of photographs of the specimen revealed
that two parallel rows of spines occur on the ventral
side of the tibia, without additional spines between
these rows (Fig. 23B). The spines are sturdy, pointed,
and variable in length. About 20 spines are discernible
on each row. They stand close together, point slightly
apicad, and are relatively long with respect to the
diameter of the tibia. These characters (number, closeness, and length of the spines) can be taken into
account for a comparison with other polyneopteran
taxa.
In extant Mantodea the fore tibia similarly carries
two parallel rows of long spines. Some other groups of
Polyneoptera display spines along the ventral side of
their fore tibiae, arranged in rows, often in correlation
with a carnivorous lifestyle. Among them, for instance,
are the Mantophasmatodea (see Klass et al., 2003:
fig. 3; Klass, Zompro & Adis, 2003: fig. 10.1A, B;
Zompro et al., 2003: fig. 4), and some Ensifera, e.g.
Listroscelidinae (Beier, 1955: fig. 45; Groll & Günther,
2003), Saginae (Kaltenbach, 1967: fig. 127, tab. 7,
Kaltenbach, 1990: figs 2–5), and Gryllacrididae
(Karny, 1937: pl. 1: figs 5, 9, pl. 3: figs 5, 9; Chopard,
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Figure 22. †Mesoptilus dolloi Lameere, 1917 (specimen MNHN A27122). A, habitus (under ethanol, polarized light). B,
detail of a leg, as located on A (dry-ethanol composite photograph). C–D, details of a leg, as located on B (dry-ethanol
composite photographs); arrows indicate spines.
104
O. BÉTHOUX and F. WIELAND
1949b: fig. 297). However, we observed no such
arrangement of spines in the Blattodea we investigated (Blattidae, Blaberidae, Blattellidae, Lamproblattidae, Cryptocercidae, Polyphagidae). Instead, in
cockroaches the spines are arranged around the tibia
in an unordered manner (see Roth, 1991: figs 19.1,
19.2B, 19.8).
The number of sturdy spines in a single row on the
tibia of †Ho. grandis was probably about 20. Exhibiting more than 15 spines is a state shown by some
Mantodea, e.g. some Hymenopodinae, Mantidae,
Toxoderidae, and Empusidae (Roy, 1999; F. Wieland,
pers. observ.). These taxa, however, are generally considered to be derived within Mantodea (Beier, 1968;
Grimaldi, 2003; Svenson & Whiting, 2004; Grimaldi
& Engel, 2005). Most extant species exhibit fewer
than 15 spines per row. This is inclusive of all putative basal Mantodea, in addition to the fossil species
identified so far (see Grimaldi, 2003). The same is
true for other Polyneoptera. In Mantophasmatodea
the number of tibial spines is lower (see Klass et al.,
2003: fig. 3D; Engel & Grimaldi, 2004: fig. 3; Arillo &
Engel, 2006: fig. 5). In Ensifera, the number of tibial
spines is often much lower than 15, with an exception
being Saga Charpentier, 1825, in which they may
reach a number of up to 15 spines per row (Kaltenbach, 1967: tab. 7). Some Ensifera exhibit strongly
elongated spines, although in smaller numbers (e.g.
Listroscelidinae and Gryllacrididae; see Karny, 1937:
pl. 1: figs 5, 9, pl. 3: figs 5, 9; Chopard, 1949b: fig. 297;
Beier, 1955: rig. 45; Hale & Rentz, 2001: fig. 6.2A;
Groll & Günther, 2003). Otherwise, the spines are
smaller than those of †Ho. grandis. As far as we are
aware, among Polyneoptera, the high number and
closeness of spines present on the tibia of †Ho.
grandis is displayed only by Mantodea.
A number of important features could not be
observed on the specimen. For example, foreleg spines
in extant Mantophasmatodea are mere cuticular out-
growths (F. Wieland, pers. observ. in Karoophasma
biedouwensis Klass et al., 2003; and K.-D. Klass, pers.
comm.), whereas in Ensifera, Mantodea, and Blattodea we observed distinct sutures between the spines
and the tibia, indicating that the spines are of setal
origin. Unfortunately, the occurrence of sutures separating the spines from their bases cannot be assessed
in †Ho. grandis. In addition, an apical spur on the
fore tibia, formed by the enlarged distal anteroventral
spine, is present in all extant (and most fossil) Mantodea except Chaeteessa spp. (e.g. Beier, 1968; Roy,
1999; Grimaldi, 2003; Klass & Meier, 2006). Whether
this trait is plesiomorphic (Beier, 1968) or represents
a secondary loss in Chaeteessa spp. (Klass & Meier,
2006; also implicated by the phylogenetic trees by
Grimaldi, 2003 and Grimaldi & Engel, 2005), is still
debated. Unfortunately, the fore-tibial apex of †Ho.
grandis is not preserved, therefore neither the insertion of the tarsus on the tibia, nor the occurrence of
an apical spur, can be assessed.
In conclusion, the arrangement of spines on the fore
tibia excludes close relationships with Blattodea, and
the number and closeness of spines suggest that the
known preserved leg of †Ho. grandis is a raptorial
foreleg mostly similar to that of some extant Mantodea. It must be acknowledged that important traits
of the tibial morphology that would help to diagnose
†Ho. grandis as a Mantodea are unknown (presence
or absence of tibial spur, lateral or terminal tarsal
insertion, setal origin of tibial spines). If a close
relationship with Mantodea is assumed, the high
number of tibial spines would be homoplasic, as it is
shared by derived extant Mantodea, the wing venation of which is far more derived than that of †Ho.
grandis.
Summary
As a result of this comparative analysis, †Mes. dolloi
and its Palaeozoic relatives can be considered as
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Figure 23. †Homocladus grandis Carpenter, 1966 (paratype specimen MCZ 5875, courtesy of the Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA). A, left forewing and associated leg remain. B, detail of the leg,
as located on A (dry-ethanol composite photograph).
CARBONIFEROUS MANTODEA IDENTIFIED
DISCUSSION
ATAVISM, POLYMORPHISM, AND SAMPLE SIZE
Our investigation relies in part on the occurrence of
atypical features exhibited by few individuals. In the
case of Met. violaceus, we observed two forewings on
which two tracheal systems occur between RA and
the anterior branch of CuA. We interpret these tracheal systems as RP and M. A unique trachea was
observed in other specimens we investigated (eight
forewings in IWC OB). Whether the unusual morphology exhibited by these specimens is an atavism (i.e.
resurgence of an ancestral character state), or if these
specimens are extreme cases of a regular polymorphism, is unknown. In any case, the occurrence of
these atypical features has to be explained. We interpret the anterior trachea as RP because we see no
other explanation consistent with the apparent lack
of RP in this species and in Chaeteessa spp. and
Mantoida spp. This explanation is also consistent
with the morphology exhibited by putative fossil
relatives.
In species belonging to Amorphoscelis it is clear
that the connection of the anterior branch of RP + M
(i.e. RP*) with RA is a matter of intra-individual to
intraspecific polymorphism. The apparent absence of
a connection between RP* with RA (as in Fig. 8) was
observed in Amorphoscelis singaporana Giglio-Tos,
1915, Amorphoscelis siebersi Werner, 1933, and Amorphoscelis annulicornis Stål, 1871 (three, one, and four
specimens, respectively; observed under a binocular
microscope, without preparation allowing course of
trachea to be observed; NHM, London). The formation
of an eye as illustrated on Figure 9 was observed in
Amorphoscelis orientalis Giglio-Tos, 1913 (with few
cases of complete fusion over 17 observed specimens;
NHM, London), Amorphoscelis tigrina Giglio-Tos,
1913, Amorphoscelis austrogermanica Werner, 1923
(eight and two specimens observed, respectively),
and Amorphoscelis pulchra (about 20 specimens
observed). The state of fusion of RP* with RA ranges
from absent to complete in Amorphoscelis nigriventer
Beier, 1930. The fusion is consistently complete in
Amorphoscelis grisea Bolivar, 1908 and Amorphoscelis pulchella Giglio-Tos, 1913 (four and five specimens
observed, respectively). Considering the sample size,
a typical morphology can hardly be outlined for this
group of species. This variability could be explained
by the fact that these species might be successive
‘intermediates’ between the more plesiotypic taxa in
which RP* diverges from RP + M and the more
derived taxa in which RP* diverges from RA + RP*
(like some genera currently assigned to the Amorphoscelidae, such as Perlamantis Guérin-Méneville,
1843; Paramorphoscelis Werner, 1907), or by poor
sampling.
We recurrently observed extant specimens exhibiting an RP* presumably diverging from RP + M,
although the typical morphology of their respective
species is a divergence of RP* from RA (Figs 5B, 10,
11A, 12C, D, F, 13B). Similar to the case of Met.
violaceus, it is difficult to determine whether such
morphology is an atavism or an extreme case fitting
within the range of a regular polymorphism. In any
case, the fact that these atypical morphologies recurrently occur indicates that they are the result of a
consistent phenomenon. Considering the variability of
this character state as exhibited by Amorphoscelis
spp., the most plausible explanation is to consider
the character state ‘RP* diverging from RP + M’ as a
plesiomorphic condition.
Regarding the secondary fusion of RP* (diverging
from RA + RP*) with RP + M as observed in some
species currently assigned to the Thespidae, all ‘intermediate’ conditions could be observed, relevant at the
level of intraspecific to intra-individual variability.
Again, in most cases, a typical morphology cannot be
properly outlined without a significant sample.
In summary, regarding the fusion of RP with M,
only rare occurrences of atypical morphologies are
documented (as it is, in Met. violaceus). Taxa in which
this putative fusion occurred are extinct, and a good
sample of relevant taxa is not available from the fossil
record. In the case of the translocation of RP* onto
RA, species lacking the translocation, polymorphic
species (including relevant variation at the level of
individuals), and the rare occurrence of individuals
lacking the translocation but belonging to species in
which the presence of this transformation is typical,
are all properly documented. Regarding species in
which the secondary fusion of RP* (diverging from
RA + RP*) with RP + M is the typical morphology,
the rare occurrence of specimens exhibiting a lack of
this fusion is not properly documented. We assume
that the sample we investigated was not significant
enough for sufficient documentation of this case.
It must be pointed out that the atypical morphologies documented on Figures 5B, D, E, 12C, D, and
13B, D, were picked out of a sample of about 40
specimens (NHM, London), 20 specimens (IWC OB),
and about 40 specimens (NHM, London), respectively.
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stem-Mantodea on the basis of two forewing character
states: anterior branch of M fused with RP for a long
distance (completely fused in crown-Mantodea; as
opposed to M and RP distinct); and CuA2 anteriorly
pectinate (as opposed to posteriorly pectinate, or
no consistent branching pattern). This proposition is
supported by evidence suggesting that strephocladidaeans exhibited raptorial forelegs, the morphology
of which is still, however, poorly known.
105
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O. BÉTHOUX and F. WIELAND
MODE OF VEIN FUSION
The fusion of RP* (emerging from RP + M) with RA is
probably the best documented vein fusion assessed in
winged insects. Based on our observation, successive
steps were hypothesized. The first step is a branching
of the anterior branch of RP + M (i.e. RP*), with the
possible re-fusion of branches resulting from this fork
(Fig. 7). The next step is a brief fusion of the anterior
branch of RP* with RA (Fig. 8A). The following step is
a fusion of the whole RP* with RA for a short distance, with distinct origins of its posterior and anterior branches from RA, both branches re-merging into
a single RP* (Fig. 8B). From this point, various
degrees of fusion via a network of tracheae can be
observed, (Fig. 9), and, thereafter the full fusion of
RP* with RA for a short distance (Figs 10B, D, 11A),
followed by a complete translocation of RP* with RA
from the wing base (Figs 11A, 12A, B, E, 13A, 14A, E,
15), are hypothesized. The essential part of the scenario is the formation of an ‘eye’ delimited by the
anterior and posterior branch of the vein undergoing
fusion as a preliminary step towards a complete
fusion. Future investigations in insect wing venation
might provide evidence for a recurrent pattern.
We demonstrate additionally that, at least in case
of a fusion of veins, the tracheal network pattern
could be distinct from the sclerotization pattern (i.e.
the ‘classical venation pattern’; notably Fig. 8F). This
fact is to be taken into account in future investigations, in particular for homologization of wing venation of fossil taxa, for which only information on the
sclerotization pattern is accessible (at least for those
fossils preserved as compression).
UNRESOLVED ISSUES
The number of branches of the composite stem
RP + M to be assigned either to RP or M is difficult to
assess. In one of the atypical specimens of Met. violaceus, the free part of M is forked, with its anterior
branch forked after some distance (Fig. 1). Based
on our survey, in Mantodea as a whole, RP + M can
either be simple or branched. Some specimens belonging to species generally exhibiting a simple RP + M
may have it branched (as found in Ma. religiosa).
Mapping the states of this character onto a phylogeny
of Mantodea might help to resolve the point.
Homologization of the forewing anal area is not
easy. In Metallyticus spp. the first convex vein posterior to CuP (so-called AA1) can either be simple
or forked, and always reaches the posterior wing
margin. This is unlike in Chaeteessa spp., Mantoida
spp., and †A. insignis, in which AA1 is always simple
and vanishes in the wing membrane between CuP
and AA2. In contrast, several amorphoscelidaeans
exhibit an organization similar to that observed in
Metallyticus spp. (see Figs 7–9), although these taxa
are considered as derived with respect to Chaeteessa
spp. and Mantoida spp. This leaves some doubt as
to the homologization of this vein in more derived
mantids, in which it can vanish in the wing membrane, as in Chaeteessa spp. and Mantoida spp.
(although it can also reach CuP, or fuse with another
anal vein). It cannot be ruled out that the anterior
branch of AA1 as exhibited in Metallyticus spp. and
some amorphoscelidaeans actually fuses with CuP in
more derived mantids. In Chaeteessa spp. and Mantoida spp., AA1 could be primarily simple. Alternative
homologizations should be tested by congruence with
other characters.
Finally, homologization of the hind wing anal area
is out of the scope of this contribution. A survey of
hind wing anal area of the Dictyoptera as a whole,
including fossil material, would be necessary. The
interpretation provided by Smart (1956) regarding the morphology of Chaeteessa spp. hind wings
must be reviewed.
FOREWING VENATION AND PHYLOGENY
The question of the homologization of mantodean
wing venation with respect to that of other winged
insects was unresolved because of multiple and inconsistent propositions (see Ragge, 1955; Smart, 1956;
Sharov, 1962; Balderson, 1991; Deitz, Nalepa &
Klass, 2003). This situation is a common case for
polyneopteran insects (see a summary of various
interpretations regarding Orthoptera in Béthoux &
Nel, 2002: tabs. 1, 2; for Plecoptera see Béthoux,
2005c: tab. 1; for Blattodea, see Rehn, 1951: 14–15),
and precluded a correct identification of fossil taxa.
Resulting from our comparative analysis, all crownMantodea exhibit a fusion of the main stem of RP
with M, occurring at the forewing base. It must be
regarded as an apomorphic trait of the order, as it is
absent in Blattodea (Rehn, 1951; Schneider, 1977,
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Interestingly, we recurrently observed a rare atypical morphology in which RP + M, or one of its anterior
branches, fuses for a more or less long distance
with RA + RP* [Figs 4B, 5E, and O. Béthoux, pers.
observ. on specimens belonging to Ma. religiosa, Blepharopsis mendica (Fabricius, 1775) (both in IWC
OB), Creobroter sp. and D. lobata (both in FW’s collection), specimen MNCN IWC OB 35, belonging to
Parastagmatoptera flavoguttata (Serville, 1839), and
specimens belonging to Hierodula sp. housed at the
MNCN]. The fixation of this trait, which can be seen
as part of a trend involving the fusion of RP* with RA,
might be the next step of forewing venation evolution
in Mantodea.
CARBONIFEROUS MANTODEA IDENTIFIED
SISTER-GROUP AND ORIGINATION DATE
OF MANTODEA
Our work suggests that taxa more closely related
to the modern Mantodea than to any other modern
group, i.e. belonging to stem-group Mantodea, existed
as early as the Late Carboniferous (c. 310 Mya). This
contrasts with the views of several recent authors,
(Zherikhin, 2002; Grimaldi, 2003; Lo, 2003; among
others), who stated that the origin of the order Mantodea was not to be assumed to be earlier than the
Early Cretaceous or Late Jurassic (with the oldest
material being 135 Myr old). However, the question of
a Mesozoic sister-group of Mantodea is controversial.
Vršanský, Vishniakova & Rasnitsyn (2002) indicated that the families †Blattulidae, Polyphagidae,
†Umenocoleidae, and the order Mantodea are related
on the basis of ‘venation simplified, with hind wing M
fewer than five-branched, CuP simple’. The fact that
this statement could allow the order Plecoptera to be
part of this assemblage speaks for the vagueness of
this argument. Notably, a ‘simple CuP’ is common to
all Neoptera but Archaeorthoptera. Moreover, this
proposal obliterates the plethora of character states
that differentiates the Blattodea from the Mantodea,
the most important of which is the loss of a differen-
tiation of R into RA and RP. This differentiation is a
strict apomorphy of the former group (see above),
whereas Mantodea exhibits the antonymic plesiomorphic state (i.e. RA and RP differentiated). Besides the
absence of clearly defined synapomorphies, Vršanský
et al.’s (2002) scenario implies a reversion of this
character.
The confusion increases once the conclusions drawn
by Vršanský et al. (2002) are compared to those of
Vršanský (2002): in the former, the family †Raphidiomimidae is unrelated to the Mantodea, whereas in
the latter, it is suggested that they are sister-group
related on the basis of ‘predacious way of life,
indicated RS [RP] in the forewing, wings less rigid’
(Vršanský, 2002: fig. 27, node 24). However, strict
homology between the predacious organs involved in
the respective group has never been demonstrated.
Even Vršanský (2002: 14) states that the legs of the
†Raphidiomimidae and of the Mantodea differ in
the length of the tibia. Once again, the vagueness of
Vršanský’s (2002) ‘diagnosis’ leaves enough room
for the inclusion of the orthopteran †Mesotitanini
Tillyard, 1925 (as defined in Béthoux, 2007c). Importantly, we were unable to identify RA from RP in the
representatives of the †Raphidiomimidae.
In the same paper Vršanský (2002) suggests that
the family †Liberiblattinidae also possesses a differentiated RP in forewings, although this family is
supposedly more basal than the †Raphidiomimidae
and Mantodea (after fig. 27 therein). Our observations of the available illustration of the holotype
of the type-species of †Liberiblattina Vršanský, 2002
(Vršanský, 2002: fig. 8), the type-genus of the family,
led us to believe that there is no differentiated
RA and RP in this taxon. Other characters provided
by Vršanský (2002: 10) as indicative of a close relationship of the †Liberiblattinidae with the Mantodea
are inconsistent [notably, the character state
‘forewing long, straight’ is shared by the group
Blattoidea + Isoptera + Polyphagoidea + (†?)Umenocoleoidea (fig. 27, node 11)], not shared by basal
Mantodea, not present in †Liberiblattinidae, and/or
vague. Lastly, a branched ScP, the only putative
apomorphy of the †Liberiblattinidae + Mantodea, is
also present in the Carboniferous taxa we consider
as stem-Mantodea, and might be a plesiomorphy at
the level of Neoptera.
In summary, there is no strong positive evidence
for a sister-group of Mantodea occurring in the
Late Jurassic and Early Cretaceous. In contrast,
our proposition, which traces the origin of mantids
further 175 Myr back, offers plenty of Carboniferous
putative stem-Dictyoptera (Bolton, 1922; Laurentiaux, 1958; Laurentiaux-Vieira & Laurentiaux,
1980), whose relationships are poorly known, as
potential sister-groups to the Mantodea.
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1978, 1980a, 1980b, 1982, 1983), stem-Dictyoptera
(Bolton, 1922; Laurentiaux, 1958; Laurentiaux-Vieira
& Laurentiaux, 1980), Orthoptera (Ragge, 1955;
Sharov, 1971; Béthoux & Nel, 2002), Plecoptera
(Needham & Claassen, 1925; Béthoux, 2005c), and
Grylloblattida (Storozhenko, 1998, 2002). The main
stem of the vein sector RP diverges from R at the
wing base and passes by the sclerotized structure
usually ‘occupied’ by the median trachea.
All mantids but Metallyticus spp., Chaeteessa spp.,
Mantoida spp., and †L. floccosa, and some other fossil
taxa, have an RA + RP* composite stem, which is an
apomorphic trait for a monophyletic group within
Mantodea. Some taxa currently assigned to the Amorphoscelidae exhibit various states of this fusion,
which supports the view of earlier workers stating
that the traditional Amorphoscelidae are paraphyletic
(e.g. Handlirsch, 1925; Chopard, 1949a; Wieland,
2003).
As a result of our investigation, RA is differentiated
from RP in Mantodea, unlike in Blattodea in which
RA is fully fused with RP. The latter state is apomorphic within Neoptera, as evidenced by the morphology
exhibited by Orthoptera (Ragge, 1955; Sharov, 1971;
Béthoux & Nel, 2002), Plecoptera (Needham &
Claassen, 1925; Béthoux, 2005c), Grylloblattida
(Storozhenko, 1998, 2002), and stem-Dictyoptera
(Bolton, 1922; Laurentiaux, 1958; Laurentiaux-Vieira
& Laurentiaux, 1980).
107
108
O. BÉTHOUX and F. WIELAND
ORIGIN OF THE OOTHECA
IMPLICATIONS FOR INSECT PALAEOBIODIVERSITY
This contribution supports the view that the ‘Protorthoptera’ must no longer be considered wholly as an
extinct group. A preliminary review suggests that
about 6% of ‘protorthopteran’ genera assigned to families in the latest compilation (Carpenter, 1992; hereafter referred to ‘protorthopteran’ genera) are to be
considered as relatives of modern mantids. Other
recent taxonomic reviews (Béthoux & Nel, 2002, 2003,
2004, 2005; Béthoux, 2003, 2005a, 2006, 2007a, b; and
unpubl. data) reveal that about 16% of protorthopteran genera are stem relatives of Orthoptera (or
even genuine Orthoptera, see Béthoux & Ross, 2005).
Additionally the fossil Grylloblattida, encompassing
about 53% of protorthopteran genera, are representatives of an extant order (Storozhenko, 2002).
Finally, with 75% of protorthopteran genera to be
viewed as stem-groups of total-groups having modern
representatives, it can be argued that the estimation of a low end-Permian diversity and subsequent
extinction of the ‘Protorthoptera’ at the beginning of
the Triassic (Labandeira & Sepkoski, 1993); see also
(Ross & Jarzembowski, 1993) was based on a taxonomic artefact.
It could yet be conjectured that a number of these
stem taxa constitute extinct monophyletic clades,
within their new respective total-groups. For
example, as defined by Carpenter (1992), the taxon
†Strephocladidae is monophyletic, occurred in the
Permian, and has no Triassic representatives. Then,
we would observe an equally high level of family
extinction at the end-Permian. However, the validity
of such lower-rank groups belonging to stem-groups is
uncertain. Several taxonomic revisions have already
demonstrated that a significant number of Carpenter’s (1992) protorthopteran families are para- or
polyphyletic (Béthoux & Nel, 2004, 2005; Béthoux,
2007c). In light of the current analysis, it is uncertain
whether the taxon †Strephocladidae is monophyletic
or paraphyletic. This taxon could indeed be composed
of successively closer relatives of crown-Mantodea.
If so, considering extinction rates makes sense at the
genus or species level only.
In fact, our knowledge of phylogenetic relationships
among stem-Orthoptera, stem-Mantodea, and stemGrylloblattida taxa is minimal. As exemplified by the
present contribution, the main reason is that major
issues in insect taxonomy rest at the super-ordinal
to ordinal taxonomic levels [see Inward, Beccaloni &
Eggleton (2007) and subsequent comments (Lo et al.,
2007; Eggleton, Beccaloni & Inward, 2007, and
Béthoux (2007c) for recent updates of insect taxonomy
at the ordinal level]. Attaining a more accurate view
is still out of reach without further taxonomic revisions, and descriptive studies.
In summary, the total-group Mantodea was distinct
from the total-group Blattodea from the Carboniferous, implying that both groups survived the Permian–
Triassic biocrisis. Accordingly, the importance of
the end-Permian faunal turnover will have to be
re-evaluated in the future. This contribution suggests
that our current view of the early steps of Pterygota
evolution could yet be significantly modified by
outcomes of taxonomic revisions. Rather than families, genera and species used as a proxy are more
likely to provide accurate results on Pterygota evolution. However, attention will have to be paid to the
sensitivity of the data to the quality of the fossil
record at the level of deposits, as well as to the
relevance of the associated taxonomic literature.
CONCLUSION
Our comparative analysis demonstrates that †Mes.
dolloi is a Carboniferous relative of modern mantids.
A number of ‘protorthopteran’ taxa will have to be
re-examined and re-interpreted in light of the present
discovery. It implies that the divergence between the
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It is generally admitted that laying eggs within an
ootheca is an apomorphy of the Dictyoptera (as represented by modern taxa; Hennig, 1981; Grimaldi &
Engel, 2005; among others). The assumption that
stem-Mantodea existed in the Carboniferous implies
that this sort of oviposition existed at the time.
Although controversial (Brown, 1957; Hennig, 1981),
it cannot be completely ruled out that fossils of Carboniferous oothecae are actually recorded (Pruvost,
1919, 1930; Laurentiaux, 1960). Laurentiaux (1960)
suggested that a part of the Late Carboniferous stemBlattodea produced oothecae. The corresponding evolutionary steps highlighted by Grimaldi (1997) could
well have occurred during the Late Carboniferous,
or even earlier. Our discovery does not imply that
the ootheca was acquired twice, as stem-lineages of
crown-Dictyoptera could well have diverged since the
Late Carboniferous.
It must be noted that scenarios regarding the origin
of the ootheca and of the related morphological traits
in Carboniferous stem-Dictyoptera are highly speculative, considering that body remains (other than
wings) that are sufficiently well preserved are
extremely rare for insect species of this period, and
that the phylogeny of stem-Dictyoptera is virtually
unknown. The morphology of modern taxa alone suggests that oothecae and related morphological traits
were acquired only once (Klass, 2003). This is not
in contradiction with a Carboniferous origin of the
Mantodea.
CARBONIFEROUS MANTODEA IDENTIFIED
Chaeteessa sp., belonging to the collection of the MTD,
to be dissected by one of us (OB). The authors also
thank B. Archibald (MCZ, Harvard University, Cambridge), W. Farrum (MCZ, Harvard University, Cambridge), L. Clunie (Landcare Research, Auckland), M.
Mostovski (Natal Museum, University of KwaZuluNatal, Pietermaritzburg) who provided photographs
of fossil specimens, or material. We also thank the
President and Fellows of Harvard College for permission to use MCZ copyrighted material (photographs in
Fig. 23). The authors are grateful to J.-M. Pacaud
(MNHN, Paris) who allowed the holotype of Mesoptilus dolloi and the specimen MNHN A27122 to be
investigated. The authors are also grateful to Z.
Simmons (Oxford University Museum of Natural
History, Oxford) for allowing a loan of the specimens
belonging to the genera Chaeteessa and Metallyticus
(including the material described by Smart, 1956)
housed at the OUMNH. The authors are grateful to
R. J. Rinas (USA) for improvement of the English
grammar. The authors thank D. Bauer (Würzburg)
and C. Schwarz (University of Würzburg) for their
help during a collecting mission in Southern France
(August 2006). FW is thankful to S. Materna (Erlangen), J. Mehl (University of Erlangen), and K. Schütte
(University of Hamburg) for their help and support
during a collecting mission in Malaysia (March 2007),
and to M. Deyrup (Archbold Biological Station, Lake
Placid, Florida) for providing specimens. Finally, we
are grateful to the members of the Interessengemeinschaft Mantodea (IGM) who provided several of the
specimens examined in this study.
ACKNOWLEDGEMENTS
The first author is a postdoctoral research fellow
of the Alexander von Humboldt Foundation. This
research project received support from the SYNTHESYS Project (http://www.synthesys.info/), which is
financed by European Community Research Infrastructure Action under the FP6 ‘Structuring the European Research Area’ Program (visits to the MNCN,
Madrid, July 2007, and to the NHM, London, August
2007), attributed to one of us (OB). This study was
partly funded by the German Research Foundation
(DFG) as part of the research project DFG Wi599/12
(participation of FW in collecting missions to France
and Malaysia). One of us (OB) is grateful to I.
Izquierdo Moya (MNCN) and G. Beccaloni (NHM) for
their help and support during the visit to the collections they respectively are in charge of, and for allowing dissection of relevant material. The authors are
grateful to R. Roy (MNHN, Paris) for allowing specimens of Metallyticus violaceus, belonging to the
collection of the MNHN, to be dissected by one of
us (OB). Similarly, K.-D. Klass (MTD, Dresden) is
acknowledged for having allowed a specimen of
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