Alienoptera a new insect order in the roach - mantodean twilight zone.

Ming Bai, Rolf Georg Beutel, Klaus-Dieter Klass, Weiwei Zhang, Xingke
Yang, Benjamin Wipfler

PII: S1342-937X(16)30004-1
DOI: doi: 10.1016/j.gr.2016.02.002
Reference: GR 1582

To appear in: Gondwana Research

Received date: 20 November 2015

Revised date: 29 January 2016
Accepted date: 2 February 2016

Please cite this article as: Bai, Ming, Beutel, Rolf Georg, Klass, Klaus-Dieter, Zhang,
Weiwei, Yang, Xingke, Wiper, Benjamin, Alienoptera a new insect order in the roach
- mantodean twilight zone., Gondwana Research (2016), doi: 10.1016/j.gr.2016.02.002

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 ACCEPTED MANUSCRIPT

Alienoptera a new insect order in the roach -

mantodean twilight zone

Ming Bai1, Rolf Georg Beutel2,*, Klaus-Dieter Klass3, Weiwei Zhang1,4, Xingke Yang1 &

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Benjamin Wipfler1, 2

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Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology,
Chinese Academy of Sciences, Box 92, Beichen West Road, Chaoyang District,

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Beijing, 100101, China;
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Entomology Group, Institut fr Spezielle Zoologie und Evolutionsbiologie mit

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Phyletischem Museum, Friedrich-Schiller-Universitt Jena, Erbertstrasse 1, D-07743
Jena, Germany
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Senckenberg Natural History Collections Dresden, Knigsbrcker Landstrasse 159,
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D-01109 Dresden, Germany
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P.O. Box 4680, Chongqing, 400015, P. R. China;
* Corresponding author: rolf.beutel@uni-jena.de
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ABSTRACT
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A new insect species (Alienopterus brachyelytrus Bai, Beutel, Klass, Wipfler et

Zhang gen. et sp. nov.) of a new order and family is described, based on a single
male embedded in Cretaceous Burmese amber (ca. 99 Ma). Unusual characters are
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shortened forewings combined with fully developed, operational hindwings, similar as

in Dermaptera, and specialized attachment pads otherwise only found in
mantophasmatodeans (heelwalkers). A cladistic analysis suggests a placement as
sister to Mantodea, supported by a profemoral brush and other characters. The male
genitalia show the same pattern in both groups. Specialized features are the unusual
flight apparatus, attachment structures adapted for locomotion on leaves, and a
dense profemoral setation suitable for catching small prey. Alienopterus was
apparently able to fly and likely a predator of small arthropods in bushes or trees. An
impressive radiation of Mantodea started in similar habitats at least 35 Ma later in the
early Cenozoic. In contrast, Alienopterus was an evolutionary dead end in the
roach-mantis transition zone.

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Keywords: mantis, fossil, amber, Cretaceous, Polyneoptera, Dictyoptera, Mantodea,

Alienopterus, raptorial leg

Research highlights
A new insect order, Alienoptera ord. nov., is described from Burmese amber.

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It displays a bizarre combination of characters occurring in different insect orders.
Alienopterus was an evolutionary dead end in the roach-mantis transition zone.

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1. Introduction
In this study we describe a new extinct insect species in a new monotypic order. Its
potential role as a link between roach-like insects and the praying mantises is

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evaluated. The single specimen with a highly unusual character combination is
embedded in Cretaceous Burmese amber, which has turned out as a very rich source
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of insect fossils (Grimaldi et al., 2002).
The earliest roach-like insects known from the Upper Carboniferous were
already remarkably diverse (Labandeira, 1994; Grimaldi and Engel, 2005; Legendre
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et al., 2015). Similar to extant cockroaches, Palaeozoic forms with a preserved body
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had a discoid pronotum partly covering the head, flattened posteriorly slanting coxae,
and leathery forewings (tegmina). In Carboniferous coal swamps they were abundant
and arguably the dominant group of insects (Grimaldi and Engel, 2005). The
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phylogenetic relationships among these Palaeozoic roachoids are unresolved as yet

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(Legendre et al., 2015). However, it is widely accepted that one of these lineages
gave rise to Dictyoptera, most likely in the Jurassic (Grimaldi and Engel, 2005: fig.
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7.60; Misof et al., 2014; Legendre et al., 2015; Tong et al., 2015). This clade
comprises the modern roaches (Blattodea) including termites (Isoptera) and the
praying mantises (Mantodea) (Grimaldi, 2003; Grimaldi and Engel, 2005; Legendre et
al., 2015). Despite a general similarity, the distinctly shortened ovipositor
distinguishes modern dictyopterans from the roachoids, which persisted up to the
end of the Mesozoic (Grimaldi and Engel, 2005; Legendre et al., 2015).
In the typical case roaches are saprophagous and solitary with a
ground-oriented lifestyle (Bell et al., 2007), and the same is likely true for most of the
"roachoids". However, during their history of more than 300 Ma, more or less highly
specialized groups have evolved several times, for example the Mesozoic
Umenocoleidae, a group strongly resembling beetles, with heavily sclerotized
tegmina and a small pronotum (Vransk, 2003; Grimaldi and Engel, 2005; Nel et al.,

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2014), or the Jurassic and Cretaceous Raphidiomimidae (Liang et al., 2009),

predacious forms with raptorial forelegs (Grimaldi and Ross, 2004).
In contrast to these lineages, which vanished from the scene in the late
Mesozoic, modern Dictyoptera are a quite successful group with almost 10,000
described species. Aside from the more or less generalized roaches (Blattaria), the

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group contains two highly specialized lineages: the wood- or fungus-feeding and

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eusocial termites (Isoptera), today recognized as a subordinate group of Blattodea

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(Klass and Meier, 2006; Inward et al., 2007), and the predacious praying mantises
(Mantodea), which are characterized by specifically modified raptorial forelegs and an

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elaborate prey catching mechanism. With about 3000 and 2300 extant species
(Ehrmann, 2002; Krishna et al., 2013), respectively, termites and praying mantises

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are slightly less species-rich than their generalized relatives, the paraphyletic roaches
(Blattaria) (Beutel et al., 2014).
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Looking at extant species, Mantodea can be easily recognized by conspicuous
modifications linked with the specialized predacious life style. Even the few known
early-diverging extinct forms (Grimaldi, 2003) differ distinctly from non-mantodean
dictyopterans, including those considered as predators (Grimaldi and Ross, 2004;
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Hrnig et al., 2013; Lee, 2014; Vransk and Bechly, 2015). In the present study we
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describe and document a Cretaceous fossil in detail using modern techniques. The
morphological features are evaluated with respect to the phylogenetic position.
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Implications for the life style and habitats of the extinct species are discussed. An
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evolutionary scenario is developed based on the phylogenetic conclusions, with a

special focus on character transformations in the roach-mantis transition zone.
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2. Material and methods

2.1 Material and photography
The single specimen, a male, was obtained from an amber deposit in the Hukawng
Valley of Myanmar. The age has been estimated as ca. 99 Ma (98.8 0.6; earliest
Cenomanian) based on U-Pb dating of zircons from the volcaniclastic matrix of the
amber (Shi et al., 2012). The mining locality is at Noije Bum, near Tanai Village
(262133.41N, 964311.88E) (Grimaldi et al., 2002; Cruickshank and Ko, 2003).
The deposit has been investigated and dated in detail by Cruickshank and Ko (2003)
and Shi et al. (2012).
The piece of amber containing the specimen was ground and polished to a
23.5x14.5x12.5 mm cube (1.58 g) and examined with a LEICA MZ 12.5 dissecting
microscope with a drawing tube attachment. Photographs were taken using a Nikon

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D610 digital camera fitted to a Zeiss Stemi 2000-C stereomicroscope and processed
in Helicon Focus 5.1 software and lastly Adobe Photoshop CS5 to deal with the
images. By merging several photographs of one series, at different focal planes, a
single final photograph was created in which the entire sample was in focus.

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2.2 Micro-CT scanning and 3D reconstruction

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The amber specimen was scanned with an X-radia 400 (Carl Zeiss X-ray Microscopy,

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Inc., Pleasanton, USA) at the Institute of Zoology, Chinese Academy of Sciences. The
scan of the entire animal (Fig. 2) was done with a beam strength of 40 KV, absorption

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contrast and a spatial resolution of 15.4732 m. The detail scans of the head (Fig. 4)
and the postabdomen (Fig. 5) were done with 60 KV, absorption contrast and a spatial

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resolution 4.1866 m. Based on the obtained image stacks, structures of the
specimen were reconstructed and separated with Amira 5.4 (Visage Imaging, San
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Diego, USA). The subsequent volume rendering and animation were performed with
VG Studiomax 2.1 (Volume Graphics, Heidelberg, Germany). For selected
illustrations parts of the animal (e.g., the wings) were virtually removed. The same
was done with air bubbles in the amber. For the reconstruction of the hindwing (Fig.
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4E) the folded part of the wing was reconstructed and rendered separately. Both parts
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of the wing were combined using Photoshop CS5 (Adobe, San Jose, USA). Final
figures were prepared with Photoshop CS5 (Adobe, San Jose, USA) and Illustrator
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CS5 (Adobe, San Jose, USA). Resolution was not sufficient for showing setae or
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spines.

2.3 Taxon sampling, characters and phylogenetic analysis

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The main aim of the cladistic analysis was to clarify the phylogenetic position of
Alienopterus. Based on the character combination wings folded back over the
abdomen, well-developed cerci, and pad-like euplantulae, Alienopterus
brachyelytrus Bai, Beutel, Klass, Wipfler et Zhang gen. et sp. nov. can be assigned
to Polyneoptera with reasonable certainty. We thus selected a taxon sample
containing extant representatives of all ten polyneopteran orders (ingroup) plus one
member of Psocodea and Hymenoptera; genera of Zygentoma, Ephemeroptera, and
Odonata were added as outgroup terminals(Kristensen, 1991; Beutel and Gorb, 2006;
Misof et al., 2014), without enforcing the monophyly of the ingroup. Additionally, we
included three extinct roaches which were considered as raptorial (Ponopterix,
Jantaropterix and Manipulator). As mantodean terminals we included four extant
genera (Chaeteessa, Metallyticus, Hymenopus and Stagmomantis) and all described
fossils with at least some accessible characters (Burmantis, Jersimantis,
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Santanmantis and Ambermantis). Supplementary Method S1 provides detailed

information about the studied taxa including the source of information. The 58
characters (head, mouthparts, pronotum, pterothorax, wings, legs, postabdomen) are
listed in Supplementary Method S2. To address the relationships of major lineages a
few selected characters were included which cannot be scored for fossil terminals

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(e.g., internal parts of the exoskeleton, muscles, mode of insemination). We analyzed

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one matrix (Supplementary Method S3) with all taxa included and a second one

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(Supplementary Method S4) with fossil taxa with many inaccessible characters
excluded (fossil roaches and mantodean nymphs). The data were gathered in a

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matrix with WinClada and parsimony analyses were conducted with NONA (ratchet,
1000 replicates) and TNT (traditional search; 99999 random seeds, 1000 replicates)

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(Goloboff et al., 2003, 2008). Bremer support was calculated with TNT.
Supplementary Table S1 provides a list with all retrieved apomorphies for both
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analyses. An extensive discussion on the various relevant character systems and
their phylogenetic evidence is provided in Supplementary Note S1.
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3. Results
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3.1 Systematic palaeontology

Order Alienoptera Bai, Beutel, Klass, Wipfler et Zhang ord. nov.

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http://zoobank.org/A4D93942-79D7-4B45-8E24-1E6CFFB4FB1A
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Family Alienopteridae Bai, Beutel, Klass, Wipfler et Zhang fam. nov.

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http://zoobank.org/11ACE691-DCB4-4FC5-AB5B-EFEE5C3EB4BA
Type genus: Alienopterus Bai, Beutel, Klass, Wipfler et Zhang gen. nov.

Alienopterus Bai, Beutel, Klass, Wipfler et Zhang gen. nov.

http://zoobank.org/04CAB2C7-3E25-4FD4-BC21-562330C7AEB1
Type species: Alienopterus brachyelytrus Bai, Beutel, Klass, Wipfler et Zhang
sp. nov.

Alienopterus brachyelytrus Bai, Beutel, Klass, Wipfler et Zhang sp. nov.

http://zoobank.org/8D5503C4-2368-49D2-A950-926AA174641B

Material

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Currently only one nearly complete specimen is known (No. BU-001057). This
adult male is designated as the holotype of the new species. The (type) specimen is
currently on long-term loan in the Institute of Zoology, Chinese Academy of Sciences
(IZAS) (specimen available for study by contacting MB or WWZ). From 2026 it will be
deposited in the currently established Three Gorges Entomological Museum,

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Chongqing, China.

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Etymology
The genus name "Alienopterus" refers to the unusual (Latin "alienus") combination of

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characters including the wings (New Latin "-pterus" = -winged, from Greek "ptern" =
wing) of the new species. The species epithet refers to the short (Greek "brachys")

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and apparently hardened forewings ("elytron", as generally used for the hardened
forewings of beetles).
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Diagnosis
The description of a new species placed in a new taxon of higher rank requires the
erection of a series of new taxa, in this case a new genus, family, and order. In the
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ideal case each taxonomic level should be associated with a diagnosis. However, in
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this case it is impossible to determine which character is diagnostic at which rank.

This depends on the character combination in the closest relatives of the new
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species, which are unknown or may not exist. We avoid this procedural conflict by
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providing a diagnosis currently valid for the entire series of newly erected taxa.
Elements of this diagnosis can be appropriately associated with a particular taxon as
soon as further species of the new order will be discovered.
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The new species (and the newly defined taxa of higher rank) can be recognized
by the following combination of characters: orthognathous head with biting
mouthparts, long filiform antennae, three ocelli, large and strongly convex compound
eyes; pronotum with very deep transverse groove in posterior third, lateral parts bent
downward to cover pleural elements; forewings short and apparently hardened,
covering anterior part of hind wings; hind wings well-developed, surpassing tip of
abdomen, folded longitudinally but not transversely, resting on abdominal dorsum in
repose; profemora with antero- and posteroventral row of densely placed setae; patch
of setae inserted on distal anterior surface (profemoral brush); meso- and metafemora
lacking comparable setal rows and patches; all tibiae with only two ventral terminal
spines, of similar size and shape on all legs; tarsi five-segmented, four proximal
tarsomeres each with paired pad-like euplantulae; large pretarsal arolium present
between claws; cerci well-developed, multisegmented; male subgenital plate
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undivided, bearing small styli; male genitalia asymmetric (mantodean type, see
below). Differing from roachoids (stemgroup Dictyoptera) and Blattodea
(including termites [Isoptera]) by the presence of a profemoral brush and
interantennal and parietal ridges. Differing from Blattodea by the presence of three
ocelli (plesiomorphy) instead of two or zero, and from Mantodea by the similar

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terminal spines on all tibiae (plesiomorphy) instead of a distinctly enlarged anterior

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terminal protibial spine, and by protibia lacking a dorsal curvature (see list of

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characters in Supplementary Method S2). The new taxon differs from both
dictyopteran orders by the combination of short sclerotized forewings and long hind

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wings, and by the deep posteriorly located transverse pronotal groove.

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Description (adult male) (Figs. 1-5). Slender, body length 14.5 mm (anterior face of
head to tip of tergite X); maximum width 3.2 mm (at forewing bases, the "shoulder"
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region). Entire cuticular surface glossy, with well-developed vestiture of long, erect
and semi-erect, yellowish setae.
Head (Fig. 4): Orthognathous head wider than pronotum, nearly as wide as
mesothoracic shoulder region; distinctly triangular, widest at dorsolateral ocular area;
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posterior cephalic region completely exposed, strongly constricted in postoccipital

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area; narrow neck suggests high movability of head; posterior side of head capsule
concave; distinct ridge originates near ventral margin, ascends dorsad (occipital ridge;
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part of parietal ridge sensu Wipfler et al., 2012), then bends anterad, extends further
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longitudinally (parietal ridge s.str.), and obliterates between compound eyes; indistinct
epistomal ridge recognizable on frontal side; antennal sockets mesally connected by
interantennal ridge; epicranial sutures not visible. Compound eyes strongly convex
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(hemispherical), with large near-circular base and small ommatidia; three large ocelli
present anterodorsally, arranged in an almost straight transverse line between
compound eyes, with median ocellus placed slightly more anterad. Antennae inserted
below lateral ocelli, very long and slender, filiform, multi-segmented; flagellum without
regionally differentiated flagellomeres; scapus distinctly longer and slightly wider than
other antennomeres; pedicellus slightly longer than flagellomeres, narrowest near
mid-length; transverse subdivision of flagellomeres (Hockman et al., 2009) not
recognizable. Labrum well-developed, free; clypeolabral suture recognizable, quite
indistinct. Mouthparts orthopteroid. Mandibles apparently unmodified but mesal parts
not visible (enclosed between labrum and maxillae). Maxillae articulate with cardines
laterad of anterior postmental region; stipes elongate, bearing fairly long
five-segmented palps; distal palpomere slightly widened distally; endite lobes not
recognizable. Labium composed of prementum and very large postmentum (about
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twice as wide as prementum); palps three-segmented; endite lobes prominent (glossa

and paraglossa not clearly discernible as individual structures); postmentum with
distinctly visible anterolateral expansions; elongate posterior part comparatively
narrow, tapering posteriorly, posterior edge reaching foramen occipitale. Posterior
tentorial grooves not recognizable. Gula not developed, at least not recognizable as

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defined unit.

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Thorax (Figs. 1-4): Paired lateral cervical sclerites and intercervical sclerites

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(Wieland, 2006) present; median unpaired ventral cervical sclerites not distinctly
recognizable. Pronotum slightly longer than wide; very deep transverse furrow

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(supracoxal furrow, Fig. 2A) separates posterior quarter of pronotum from longer
anterior part, corresponds with internal supracoxal ridge; anterior pronotal part

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strongly convex, constricted by U-shaped horizontal groove; posterior ends of
horizontal groove connected with supracoxal groove; central anterior part of pronotum
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hemispherical in lateral view, round in dorsal view; lateral pronotal region strongly
bent downward, almost vertical, reaching beyond lateral base of coxa, largely
covering pleural region; distinct lateral lamellar pronotal extensions absent.
Mesothorax distinctly shortened; dorsally with triangular scutellar shield .
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Mesosternum well-developed, flat and plate-like; paired triangular fold-like structures

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present anterior to mesocoxae (possibly the trochantin), mesofurcal origin probably

indicated by median invagination between them. Only small part of mesopleura
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exposed, partly covered by pronotum and partly by forewing; mesopleural ridge

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visible. Metathorax much larger than pro- and mesothorax; large triangular
metascutellum recognizable dorsally. Metapleuron largely exposed; metapleural ridge
very distinct, originates from posterolateral metacoxal margin. Individual metasternal
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elements not recognizable.

Legs moderately long, length slightly increasing from prothorax to metathorax.
Pro- and mesocoxae quite elongate, conical, slightly narrowing from base to apex;
distinct concavity present on lateral side of distal part, most distinctly on procoxae;
metacoxae enlarged, triangular, with basal part extending far laterad; elevated mesal
part forms even platform; straight mesal metacoxal edges adjacent; distinct laterally
directed process present at laterodistal end of metacoxal platform. Pro-, meso- and
metatrochanters small, roughly triangular. Femora larger than other leg elements,
about twice as long as coxae, very slightly tapering distally; diameter of profemur not
enlarged compared to meso- and metafemora; each profemur with distinct anterior
and posterior longitudinal rows of setae on ventral surface; each row formed by
numerous (more than 45) densely spaced setae; proximal setae longer than distal
ones; a dense group of setae (profemoral brush) located on distal half of anterior
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surface; meso- and metafemora (Supplementary Figure S1) lack ventral rows of setae
(or setae minute) and femoral brush; genicular spine on anterodorsal femoral apex
lacking on all legs. All tibiae very similar, more slender than femora, about equally
long, slightly widening distally; protibia not curved, i.e. dorsal edge straight, not
convex; tibiae with numerous setae on all sides, apparently longer on mesal surface;

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larger setae or spines arranged in two ventral rows absent; spines on dorsal side of

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tibiae also missing; two ventral terminal spines (Tt1, Tt5; Supplementary Figure S1)

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present distally on all tibiae, all of similar size and shape, not placed on tibial
projection, i.e. protibia lacking apical spur. Tarsus five-segmented; tarsomeres one to

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four of all legs with two pad-like euplantulae; size decreasing from one to four; the
small tarsomere four distinctly protruding; elongate and slender apical tarsomere five

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strongly bent upward. Pretarsus with paired claws equal, flanking a large pan-shaped
arolium.
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Two pairs of wings present, folded back on dorsum of pterothorax and abdomen
in repose. Forewings covering anterior parts of hindwings; highly modified, forming
scale-like shortened pseudoelytra with leathery cuticular structure; shoulder region
strongly pronounced, part of fore wing mesad of shoulder horizontal, lateral part
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nearly vertical; veins not recognizable in forewing; callous stigma possibly present but
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very indistinct; epipleura not present as delimited region; strongly diverging mesal
edges of the pseudoelytra fit with scutellar shield; distinct mesal interlocking device
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lacking; articulation of forewings not recognizable. Hindwings (Fig. 4E) fully

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developed, wrapped around upper part of abdomen, folded longitudinally but not
transversely; posterior edge slightly surpasses abdominal apex; wing articulation not
clearly recognizable; main longitudinal veins partly visible, especially in proximal
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region (visible parts conforming with the typical dictyopteran pattern (Smart, 1951,
1956)).
Abdomen (Fig. 5): Abdomen composed of ten distinctly developed segments and
reduced segment XI with cerci. Sternite II fully developed as anteriormost visible
ventrite, sternite I at least strongly reduced (or absent). Cerci distinctly developed,
conical, about as long as last visible sternite (IX); composed of large basal cercomere,
three distinctly shorter intermediate cercomeres, and elongate and slender distal
cercomere. Subanal lobes bearing paraproct sclerites (PP-l, PP-r in Fig. 5C) large,
slightly asymmetric (or distorted); supraanal lobe not visible. Sternite IX forms
posteriorly expanded subgenital plate, apically bearing paired, short styli positioned
close to midline; apparently uniformly sclerotized (no subdivision recognizable).
Male genitalia (phallic organs) originate from large cavity bordered by sternite IX
ventrally and subanal lobes and cercal bases dorsally; only apical projecting parts of
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genitalia visible; basal parts enclosed in genital cavity or embedded in debris in

interspaces of cavity; internalized parts (e.g., apodemes) not accessible; phallic
organs strongly asymmetric. Left part with two apical processes: ventral one (pda)
short, wide, directed towards right side, with broadly rounded apex; dorsal process
(paa) slender, curved dorsad; bases of processes pda and paa separated by

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transverse cleft (entrance of a pouch, lve); interpretation of other elements of left side

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(L1 in Fig. 5A) ambiguous, posterior part apparently wrapped in membranous

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foldings, anterior part concealed by tergite X; L1 (as suggested by visible part)
probably developed as narrow sclerite, possibly flexed ventrad along edge-like right

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extremity. Right part of genitalia mainly represented by a large compact lobe (fda in
Fig. 5B-D); mesal part of fda (labeled fdm) projects slightly beyond remainder of lobe

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fda, vertically orientated, slightly bent towards right side; lobe fda probably giving rise
to a ventral lobe or tooth (pia) (as suggested by ventral view) projecting to left side
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(i.e. lobes fda and pia diverge to left side like two halves of a slightly opened book;
white arrows in Fig. 5B indicate free posterior edge of lobe pia); dorsal and lateral
walls of lobe fda probably continuously sclerotized (sclerite R1); sclerite R1
apparently forming a mesally directed arm dorsally (white arrow in Fig. 5D);
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lateroventrally R1 apparently broadly continuous with sclerotization of lobe pia. See

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character discussion in Supplementary Note S1 for more details on phallic organs.

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3.2 Phylogenetic analyses

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In the first analysis with 58 morphological characters and 31 terminals

(Supplementary Method S1+S2; matrix provided in Supplementary Method S3) the
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strict consensus tree of 25 equally parsimonious trees (98 steps) (Supplementary

Figure S2) was poorly resolved, especially within Polyneoptera. This was obviously
due to the included fossil roach taxa (Manipulator, Ponopterix, Jantaropterix). As
many characters could not be scored for them, they switched between different
positions in the tree, thereby blurring the phylogenetic results. In the second analysis
we excluded the three fossil roaches and two fossil mantodean nymphs
(Jersimantis, Burmantis) with important inaccessible characters (matrix provided in
Supplementary Method S4). This analysis including 26 terminals yielded only three
most parsimonious trees (96 steps) (Fig. 6) (see Supplementary Table S1 for potential
autapomorphies for all nodes in both analyses).
The second analysis supported Metapterygota (= Odonata + Neoptera),
Neoptera, and tentatively Polyneoptera. All insect orders with more than one
representative included were obtained as monophyletic. Alienopterus was

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unambiguously placed as sister group of Mantodea including extinct and extant

terminals. Basal relationships in Mantodea could not be resolved. Only the "higher"
mantodean taxa Stagmomantis and Hymenopus are combined in a clade. It should be
noted that our dataset was neither composed to address deep insect relationships
(Misof et al., 2014; Wipfler et al., 2015), nor the intraordinal phylogeny of Mantodea

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(Svenson and Whiting, 2004, 2009; Wieland, 2013). The focus was on the placement

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of Alienopterus.

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4. Discussion
Alienopterus exhibits a seemingly weird combination of characters of different

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groups of insects. This mosaic of structural features results in a very unusual
appearance, which is reflected by the name "Alienoptera". The combination of short
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sclerotized forewings and fully developed, operational hindwings resembles the
condition found for instance in earwigs (Dermaptera; Haas, 2003), staphylinid beetles
(Thayer, 2005) and some tettigoniids (Orthoptera; Bailey, 1979). The large,
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pan-shaped arolium (an attachment device) and the upward lift of the ultimate
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tarsomeres are typical for heelwalkers (Mantophasmatodea (Beutel and Gorb, 2008)).
The saddle-shaped pronotum is similar to conditions found in grasshoppers and
relatives (Orthoptera (Snodgrass, 1908)). Other features place Alienopterus in a
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lineage comprising roaches, termites and praying mantises (Dictyoptera), as the sister
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group of the order Mantodea (Fig. 6; see Supplementary Note S1 for detailed
discussions).
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Unfortunately, most derived (apomorphic) features characterizing Dictyoptera

are not visible in the single fossil specimen of Alienopterus. This includes internal
structures like the inner skeleton of the head (tentorium, character 15 in
Supplementary Method S2), soft tissue (muscles; antennal vessels, character 56),
female postabdomen (subgenital plate formed by coxosternite VII, character 48),
characters related to reproduction (ootheca, character 45), and details of the
mouthparts (postmola on mandible, character 50; lacinia in cavity of galea, character
55). However, a specific detail of the leg supporting an assignment to Dictyoptera is
visible in Alienopterus: a ventral sclerite located between the margins of the tibia and
the basal tarsomere (intertibiotarsal sclerite, character 52). As far as currently known,
this sclerite occurs only in the dictyopteran orders.
Despite the incomplete accessibility of characters, a placement of Alienopterus
in Dictyoptera is also clearly implied by derived features shared with Mantodea. The

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profemoral brush (character 21), a cleaning device, was hitherto only known from
mantises (including all fossils where this body part is visible). Several other derived
features are shared by both taxa, but are not unique to them: a notch and lobe at the
tibial apex (character 51), and interantennal (character 5) and parietal (character 2)
ridges on the head. The presence of a callous stigma (a thickened area of the

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forewing; character 34) would be an additional argument, but the condition in

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Alienopterus is ambiguous. The strongest argument for Alienopterus + Mantodea

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comes from the similar structure of the male genitalia. Male genital structures are
highly divergent between insect orders and also often on lower taxonomic levels (e.g.,

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Zoraptera (Dallai et al., 2014), Blattodea (Klass, 1997)). The visible projecting parts of
Alienopterus (ca. 30% of the genital apparatus) fully conform with conditions found in

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Mantodea. This is a rare case where the genital morphology of an extinct taxon can
be reconstructed to an extent allowing a reliable phylogenetic interpretation. Another
potential synapomorphy of Mantodea and Alienopterus is a feature of the legs: the
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ventral femoral rows of setae (or spines) are more strongly developed on the forelegs
(Fig. 3A) than on the middle and hind legs (Fig. 3E-F). Another character set shared
by both taxa is likely linked with raptorial habits, the triangular, very movable head
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with large, strongly protruding compound eyes. Similar conditions occur in extinct
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predacious roaches (Vransk, 2003; Hrnig et al., 2013; Lee, 2014; Vransk and
Bechly, 2015), possibly as a result of parallel evolution.
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Compared to all known extant and extinct mantodean species, Alienopterus

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has retained a considerable number of plesiomorphies. One of them is the presence

of an arolium, an attachment structure occurring in many groups of Neoptera (winged
insects able to fold back the wings) and arguably a groundplan apomorphy of this
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lineage (Beutel and Gorb, 2001, 2006; character 27). The arolia are exceptionally
large in Alienopterus, similar to the condition in Mantophasmatodea (character 28),
but absent in all extant mantodeans, and also missing (or perhaps vestigial) in
Santanmantis (Grimaldi and Engel, 2005) and other extinct species of this order. The
most conspicuous differences concern the forelegs, which are distinctly modified in
Mantodea as specialized organs for catching prey: the thickness of the spines in the
ventral rows on the femur and tibia is increased (e.g., compared to Blattodea), the
anterior terminal tibial spine (Tt1) is distinctly enlarged and placed on a projection,
and the entire tibia is curved (at least in the ground plan of the group; characters 22,
23, 47). Due to these modifications, which are apparently completely missing in
Alienopterus, the mantodean forelegs form an efficient grasping and holding device.
We follow Klass and Ehrmann (2003) and others (e.g., Wieland, 2013) by considering
this specialized raptorial foreleg as a key innovation and defining apomorphy of
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Mantodea. Alienopterus does not have this specialization (Fig. 3A). Thus we place it
in a new order, Alienoptera. This is also in agreement with our cladistic analysis
including the fossil roaches (Supplementary Figure S2) where Alienopterus is not
unambiguously placed with Mantodea.
Aside from preserved plesiomorphic features, Alienopterus displays an entire

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series of apomorphies unknown in Mantodea, including modifications of the forelegs:

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the setae of the ventral profemoral rows are delicate compared to the corresponding

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spines of Mantodea and most blattodean groups (thin setae also occur in corydiid
cockroaches). Additionally, the ventral rows of protibial spines are missing in

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Alienopterus, in clear contrast to nearly all groups of Mantodea and Blattodea.
Conspicuous apomorphies of Alienopterus are the strongly shortened forewings (in

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combination with fully developed hindwings), the distinctly enlarged arolium, elevated
terminal tarsomeres, and the saddle-shaped pronotum. Even though these features
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occur in other insect groups, suggesting possible phylogenetic affinities, the analyses
presented here show that they evolved independently in Alienopterus as
autapomorphic conditions, contributing to a weird appearance of this extinct insect.
Alienopterus is embedded in 99 Ma old amber. The age of the clade Mantodea is
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discussed controversially (Grimaldi, 2003; Bethoux and Wieland, 2009; Svenson and
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Whiting, 2009; Misof et al., 2014; Legendre et al., 2015), but based on the fossil
evidence, the minimum age is 130 million years. Consequently, the last common
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ancestor of Mantodea and Alienoptera must have lived at least 30 million years
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earlier than Alienopterus brachyelytrus. In this (minimum) time this unique

combination of characters and adaptations evolved in the lineage including the
species described here.
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All in all, the characters evaluated here and the results of our cladistic analysis
place Alienopterus as the sistergroup to Mantodea, with retained ancestral
conditions, but also an array of specialized features. Both phylogenetically and in its
structural features, Alienopterus is a connecting link in the narrow space between
roach-like ancestors and the successful praying mantises as we know them today.
The reconstruction of the lifestyle of extinct species is greatly impeded by the
complete lack of direct observations. However, the accessible morphological features
of Alienopterus allow several conclusions.
A flight apparatus with large hind wings and shortened forewings is unknown in
extant Dictyoptera, where both fore- and hind wings are either well developed or
reduced to varying degrees. The large hind wings suggest that Alienopterus could fly
well, while the forewings played a minor role in this functional context if at all.
Alienopterus is a case of functional dipterism, with flight mechanics likely differing
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considerably from other dictyopterans, but similar to conditions occurring in earwigs

(Dermaptera) and beetles (Coleoptera). These groups fold their membranous flight
organs under the forewings for mechanical protection, with a derived transverse
folding mechanism (Haas, 2006). This is obviously not the case in Alienopterus,
where the hind wings are largely exposed, and only folded longitudinally (as in most

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dictyopterans). However, the sclerotized forewings form a cavity containing the

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proximal part of the hind wings. It is conceivable that the resulting protection of the

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wing joint is an advantage, as it is also the case in earwigs, beetles, and some stick
insects (Phasmatodea) with shortened leathery forewings (Bradler, 2003). Apparently

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this is a trade-off between easy and rapid unfolding (without transverse folding
mechanism) and full protection of the membranous flight organs in repose. Beetles

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and earwigs are adapted to penetrate narrow crevices and usually dorsoventrally
flattened (Beutel, 1997). This is apparently not the case in Alienopterus and
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Phasmatodea, which are characterized by a more or less cylindrical body, suggesting
a more exposed lifestyle.
The three pairs of legs of Alienopterus are structurally very similar and were
probably used for normal hexapod walking. However, a rather unspecialized raptorial
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function of the forelegs is conceivable, as long and densely arranged ventral setae
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are present on the profemora but lacking on the middle and hind legs. The delicate
condition of the profemoral vestiture and the absence of counteracting tibial spines
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(both arguably apomorphic conditions) suggest that prey-grasping was distinctly less
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powerful than in Mantodea. The forelegs were likely suitable for grasping small items
from the substrate in front of the insect, perhaps with movements similar to those of
mantodean forelegs. This may have included small and delicate arthropods such as
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for instance midges or aphids. A raptorial lifestyle is also suggested by the high
movability of the head. Beyond this, the large, widely separated compound eyes and
the presence of a profemoral brush, which like in extant Mantodea may have served
as a cleaning device for the eyes, indicate that as in Mantodea the raptorial habits of
Alienopterus were mainly supported by vision. Regardless of the precise target and
technique of food acquisition, it is likely that an early stage of raptorial prey-catching
had evolved in the common ancestor of Alienopterus and Mantodea.
The enlarged arolium of Alienopterus is a unique feature in Dictyoptera, with
parallel evolution in heelwalkers (Mantophasmatodea) and "basal" stick insects
(Timematodea) (Beutel and Gorb, 2006, 2008). Timematodeans spend their lives on
leaf surfaces and show cryptic behaviour to a degree that certain species or morphs
are adapted to leaves of specific plant species (Nosil and Crespi, 2004). Additionally
they rely on defensive glands. As an almost unique feature, heelwalkers can lift or
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lower the terminal tarsomere, thus either keeping the arolium off the substrate
(walking on the heels) or in touch with it, depending on the specific situation (Beutel
and Gorb, 2006; Roth et al., 2014). The arolia are only used for walking on smooth
surfaces such as leaves, for striking prey, when additional grip is required (e.g.,
transportation of larger prey), or for walking in an upside down position (Roth et al.,

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2014). Like in Mantophasmatodea, the large arolium of Alienopterus could be lifted

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with the last tarsomere. This suggests that it was used in a similar way and probably

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also in similar habitats.
The complex genitalia of Dictyoptera are bilaterally symmetric in females (Klass,

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1998) but strongly asymmetric in males (Klass, 1997). A particular architecture of
complex male genitalia usually requires a defined way of how to connect them with

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the female organs. This usually correlates with a particular posture (or series of
postures) for initiating copulation. In Mantodea the male climbs or jumps on the back
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of the female and clings to the thorax with his forelegs. Then the male curves the
postabdomen to the right (or to the left in the rare cases of mirror-inverted genitalia;
see Supplementary Note S1) and pushes his phallic organs into the genital pouch of
the female (Kumar, 1973). The great similarity of the male apparatus of Alienopterus
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strongly suggests a similar pattern in this extinct taxon.

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Based on our structural observations, we conclude that Alienopterus

brachyelytrus was a predator with a preference for trees or bushes, with suitable
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foliage and open spaces. Similar environments are preferred by many recent species
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of Mantodea. However, early mantises probably hunted on bark, a lifestyle still

observed in Recent branches of the order (Wieland, 2010). The transition to an open
three-dimensional environment such as trees, bushes or larger herbaceous plants
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occurred in the Tertiary, thus at least 35 million years after Alienopterus lived
(Grimaldi and Engel, 2005). Alienoptera can thus be considered as an early lineage
in dictyopteran evolution, a highly specialized but unsuccessful predator in the
roach-mantis transition zone.

5. Conclusions
The detailed morphological investigation of the new species Alienopterus
brachyelytrus from Burmese amber and the phylogenetic analyses (including extant
and extinct taxa) revealed that it belongs to Dictyoptera. The profemoral brush, male
genitalia, and other features suggest that it is the sistergroup of the order Mantodea,
despite the seemingly unorthodox combination of features (e.g., shortened tegmina,
enlarged arolium). A placement within mantodeans (incl. the stemgroup) can be

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excluded. Alienopterus was characterized by a specialized efficient flight apparatus,

attachment structures adapted for locomotion on leaves, and a dense profemoral
setation suitable for catching small prey. These features also suggest that it lived in
bushes or trees. However, it failed to succeed in this environment, which was later
successfully populated by the praying mantises with their distinctly modified raptorial

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forelegs. Alienopterus was thus an evolutionary dead end in the

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roach-mantis-transition zone.

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Alienoptera is the fourth newly discovered insect taxon of recent years assigned
ordinal rank, following the Mantophasmatodea (with extant and extinct taxa; Klass et

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al., 2002), Nakridletia (extinct; Vransk et al., 2010) and Coxoplecoptera (extinct;
Staniczek et al., 2011). However, an improved resolution of phylogenetic relationships

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among early (Palaeozoic) pterygote insects may require the erection of further new
insect orders. This will likely concern the multitude of groups assembled in
Grylloblattida (see Cui et al., 2015), but also the paraphyletic stem-Dictyoptera, the
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roachoids (see Grimaldi and Engel, 2005: fig. 7.60), and possibly also other groups.
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Acknowledgments

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We thank Sven Bradler (Georg-August-Universitt Gttingen) for his helpful

comments concerning stick insects and all curators who hosted the first author or

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loaned or supplied material for this study. This research was supported by the

National Basic Research Program of China (973 Program) (No. 2011CB302102), the
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National Natural Science Foundation of China (Nos. 31172143, 31350110218 (B.W.)),

the National Science Fund for Fostering Talents in Basic Research (Special Subjects
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in Animal Taxonomy, NSFC-J1210002), Research Equipment Development Project of

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Chinese Academy of Sciences (YZ201509), by a Humboldt Fellowship (M.B.) from

Alexander von Humboldt Foundation, by the BIG4 project (MSCA-ITN-2014-ETN:

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Marie Skodowska-Curie Innovative Training Networks [ITN-ETN],

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642241BIG4H2020-MSCA-ITN-2014), and by the Deutsche

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Forschungsgemeinschaft DFG (KL1162/7-1; WI 4324/1-1).

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Figure legends

Fig. 1. Photographs of Alienopterus brachyelytrus Bai, Beutel, Klass, Wipfler et

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Zhang gen. et sp. nov., holotype specimen (No. BU-001057). (A) Lateral view of

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specimen. (B) Dorsal view of specimen. (C) Complete view of the amber cube with

scale. Scale bars: 1 mm (violet).

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Fig. 2. 3D reconstruction based on a -CT scan of Alienopterus brachyelytrus Bai,

Beutel, Klass, Wipfler et Zhang gen. et sp. nov. (A) Lateral view; left foreleg partly
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missing due to scan limitations. (B) Dorsal view. (C) Ventral view, with larger parts of

legs removed. Abbreviations: an = antenna, ar = arolium, ce = compound eye, cer =

cercus, CS9 = abdominal coxosternum IX, cx1 = procoxa, cx2 = mesocoxa, cx3 =
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metacoxa, fe = femur, fw = forewing, hw = hind wing, lp = labial palpus, mp = maxillary

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palpus, mss = mesoscutellum, mts = metascutellum, pn = pronotum, sf = supracoxal

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furrow, ti = tibia. Scale bars: 1 mm (violet).

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Fig. 3. Morphological details of Alienopterus brachyelytrus Bai, Beutel, Klass,

Wipfler et Zhang gen. et sp. nov. (A) Forelegs. (B) Head. (C) Arolium. (D) Forewing.

(E) Mesofemur and mesotibia. (F) Metafemur and metatibia. Abbreviations: ce =

compound eye, cla = pretarsal claw, fe = femur, feb = femoral brush, Fva = anterior

row of setae on ventral surface of profemur, Fvp = posterior row of setae on ventral

surface of profemur, lp = labial palpus, mp = maxillary palpus, ta = tarsus, ti = tibia,

Tt1 =tibial terminal spine 1, Tt5 = tibial terminal spine 5. Scale bars: 1 mm (violet); 0.5

mm (blue).

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Fig. 4. Details of Alienopterus brachyelytrus Bai, Beutel, Klass, Wipfler et Zhang

gen. et sp. nov. A-C+E: 3D reconstructions based on -CT data; D: line drawing. (A)

Head in lateral view. (B) Head in posteroventral view. (C) and (D) Head in frontal view.

T
(E) Unfolded hind wing. Abbreviations: A1 = anal vein, an = antenna, as = antennal

IP
socket, ce = compound eye, cl = clypeus, cs = cervical sclerite, CuA = anterior cubital

vein, CuP = posterior cubital vein, er = epistomal ridge, fo = frontal ocellus, ias =

R
SC
interantennal ridge, lb = labrum, lo = lateral ocellus, lp = labial palpus, M = medial

vein, md = mandible, mp = maxillary palpus, pm = prementum, pn = pronotum, po =

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postmentum, pr = parietal ridge, RA = anterior radial vein, RP = posterior radial vein,

ScP = posterior subcostal vein. Scale bars: 1 mm (violet).

MA
D

Fig. 5. 3D reconstruction of postabdomen of Alienopterus brachyelytrus Bai, Beutel,

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Klass, Wipfler et Zhang gen. et sp. nov. (A) Ventral view. (B) Dorsal view. (C)
Posterior view. (D) Lateral view. Abbreviations: cer = cercus, CS9 = coxosternite of
P

abdominal segment IX, PP-l, PP-r = left and right paraproctal sclerites (on subanal
lobes), sl9 = stylus of abdominal segment IX, TG9, TG10 = tergites of abdominal
CE

segments IX and X. Genitalic sclerites (L1, L4, R1), projections (afa, fda, fdm, loa, paa,
pda, pia, and vla), and pouches (lve) are labeled according to the terminology of Klass
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(1997). Scale bars: 1 mm (violet).

Fig. 6. Result of the cladistic analysis. Strict consensus tree of three most
parsimonious trees with 96 steps. Cladistic analysis based on 58 characters (SI
Appendix, Method S2) for 26 taxa (SI Appendix, Method S1). Polyneoptera (excl.
following groups) in dark blue, Dictyoptera stem in light blue, Blattodea in green,
Alienoptera in orange, Mantodea in brown. Numbers on branches indicate Bremer
support above 1. Potential autapomorphies for all clades are provided in SI Appendix,
Table S1. Insect (sub-)orders are represented by icons that do not necessarily show
the sampled species.

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Figure 1

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Figure 3

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Figure 5
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Figure 6
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Graphical abstract

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