Eur. J. Entorno!. 98: 445-509, 2001
ISSN 1210-5759

Dermaptera hindwing structure and folding: New evidence for familial,ordinal and superordinal relationships within Neoptera (Insecta)
Fabian HAAS1and Jarmila KUKALOVÂ-PECK2
1Section ofBiosystematic Documentation, University ofUlm, Helmholtzstr. 20, D-89081 Ulm, Germany;e-mail: fabian.haas@biologie.uni-ulm.de
2Department ofEarth Sciences, Carleton University, Ottawa, ON K1S 5B6, Canada; e-mail:jkpeck@ccs.carleton.ca
Key words. Dermaptera, higher phylogenetics, Pterygota, insect wings, wing folding, wing articulation, protowing
Abstract. The Dermaptera are a small order of insects, marked by reduced forewings, hindwings with a unique and complicatedfolding pattern, and by pincer-like cerci. Hindwing characters of 25 extant dermapteran species are documented. The highly derived
hindwing venation and articulation is accurately homologized with the other pterygote orders for the first time. The hindwing base of
Dermaptera contains phylogenetically informative characters. They are compared with their homologues in fossil dermapteranancestors, and in Plecoptera, Orthoptera (Caelifera), Dictyoptera (Mantodea, Blattodea, Isoptera), Fulgoromorpha and Megaloptera.
A fully homologized character matrix of the pterygote wing complex is offered for the first time. The wing venation of the Coleoptera is re-interpreted and slightly modified. The all-pterygote character analysis suggests the following relationships: Pterygota:
Palaeoptera + Neoptera; Neoptera: [Pleconeoptera + Orthoneoptera] + [Blattoneoptera + (Hemineoptera + Endoneoptera)]. Blattoneoptera share at least 15 wing apomorphies with the sistergroup Hemineoptera + Endoneoptera and none with the Orthoneoptera
and Pleconeoptera; Blattoneoptera: (Grylloblattodea + (Dermaptera + Dictyoptera)); Dictyoptera: (Mantodea + (Blattodea +Isoptera). Dermaptera share 13 wing apomorphies with the sistergroup Dictyoptera. In order to document the intra-ordinal relation­
ships of Dermaptera, 18 new characters of venation and articulation are added to an existing data set and analyzed cladistically. The
following relationships are suggested (43 characters, tree length 72, CI 0.819 and RI 0.935). Dermaptera: Karschiellidae +(“Diplatyidae” + (“Pygidicranidae” + (AHostethus indicum + (Anisolabididae + (“Labiduridae” + [Forficulidae + (Chelisochidae +Spongiphoridae)]))))). The taxa in quotation marks are probably paraphyletic. Fossil Dermaptera and “Protelytroptera” show thatwing-folding characters were already present in Permian ancestors. The evolution of the dermapteran wing-folding mechanism is
discussed and the hindwing is presented as a working “origami” model, which will fold as in living earwigs. The functional role of
the wing base in wing folding is examined. Characters in orders and other higher taxa are not independent and cannot be analyzed
out of context with their groundplans. Higher systematics is dealing with diametrally different problems than species-level systematics. The necessity of using a different methodology for species-level and higher-level phylogenetics is discussed and recommenda­
tions are made.
CONTENTS
Introduction........................................................................
Materials and methods ......................................................
Nomenclature and homology ......................................
Techniques ..................................................................
Species examined ........................................................
Characters of extant Dermaptera taxa ........................
Characters of fossil Dermaptera taxa ..........................
Analysis ......................................................................
Movement of wing articulation ..................................
Results ................................................................................
Diagnosis of the hindwing ..........................................
Description of the hindwig ..........................................
Folds and areas in Dermaptera hindwings ..................
Hindwing articulation ..................................................
Phylogenetic analysis of extant Dermaptera ..................
Charactersandtheirstates ..........................................
Character analysis ........................................................
Phylogenetic analysis of fossil Dermaptera ....................
Charactersandtheirstates ..........................................
Character analysis ........................................................
Discussion ..........................................................................
Pterygota: Protowing and the split into Palaeoptera
and Neoptera ................................................................

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Neoptera: Basal split into Pleconeoptera + Orthoneoptera
and Blattoneoptera + Heminoptera + Endoneoptera . . 483
The relationships of Dermaptera within Blattoneoptera 484
The relationships of extinct “Archidermaptera” ........ 484
The relationships of extant Dermaptera “families” . . . 485
Are the Zoraptera member of the Blattoneoptera ? . . . 486
Function and evolution of the dermapteran hindwing . . . 487
Dermaptera wing folding mechanism ........................ 487
Wing articulation and folding ...................................... 489
Wing venation and the evolution of fo ld in g ................ 490
Higher taxa and species require different methodologies
in character assessment .................................................. 490
Conclusion ........................................................................ 492
Table 6. The character table of the pterygote wing complex.................................................................................... 492
Acknowledgements ............................................................ 507
References .......................................................................... 507
INTRODUCTION

The Dermaptera (earwigs) are a small insect order con­taining about 2000 species (Sakai, 1996). The primarydermapteran characteristics are a unique folding of thehindwings, and the pincer-like cerci. The hindwings con­tain an enormous fan supported by anal branches, which445

unlike other pterygote orders, radiate from the centre of
the wing rather than from the wing base. The wing fan isfolded twice transversely with a folding ratio of about 10(Haas & Wootton, 1996), compared to 4 in Coleoptera(Haas, 1998). The wing articulation has a large membra­
nous area in its centre, which in the other pterygote
insects is used for transmitting major forces during flight.
Furthermore, the hindwing is folded by intrinsic elasticity
and uniquely unfolded with the help of the cerci(Kleinow, 1966; Haas et al., 2000; this paper).
The evolution and diversification of the hindwing
formed a major set of characters, which show the phylo­
genetic relationships of Dermaptera and between the dermapteran families. The crucial requirement for using thewing complex in higher systematics is its accurate
homologization at the levels of Pterygota, Palaeoptera,
Neoptera, superorder and order. Idiosyncratic, “conven­ient” wing nomenclatures, which were developed throughtime for each pterygote order, have practical use for the
lower taxa, but are useless for the inter-ordinal relation­
ships. We employ an all-pterygote homologous nomen­
clature, and for Neoptera the reference scheme
reconstructed by Kukalova-Peck (1983, 1985, 1991,
1997) and improved here (Fig. 1). The Dermaptera
hindwing is now accurately homologized, newly inter­
preted and compared in detail with the plesiomorphic re­presentatives of the monophyletic Dictyoptera (theMantodea, Blattodea and Isoptera), which was found tobe the sistergroup. The phylogenetic relationship of Blattoneoptera to Hemineoptera, Endoneoptera, Orthoneoptera and Pleconeoptera is examined and documented in
the hindwings of plesiomorphic representatives. For the
first time, the fully homologized characters of the ptery­gote wing complex are presented in a table, to provide a
broad evolutionary setting and to offer a fast and concise
overview of the many characters of the neopterous super­ordinal lines.
Our analysis of the hindwing character complex sup­ports the sistergroup relationship between the Dermaptera
and Dictyoptera with 13 characters, and provides the rea­soning for the rejection of alternatives offered by otherauthors. We did not include Zoraptera in our analysis,
because convincing wing characters are not yet available.
For further discussion of Zoraptera see Boudreaux, 1979;
Stys & Bilinski, 1990; Kukalova-Peck & Peck, 1993;
Kristensen, 1995; Rasnitsyn, 1998; and a critical review
here.
By including fossil “Protelytroptera” and Dermaptera,
we are able to support the hypothesis that the “Protely­
troptera” are paraphyletic and that some of their represen­tatives are indeed in the stem line of Dermaptera. The
study of stem-line Dermaptera was found to be necessary
for understanding the evolution of the autapomorphic
hindwing folding in extant Dermaptera. Only with theclues from fossils was it possible to reconstruct the evolu­
tionary steps between the cockroach-like hindwing(folding in a relatively simple way) and the highly
derived modern dermapteran hindwing (folding in a very
complex way).
446

Further diversification of the dermapteran wing com­plex enabled us to re-assess the intra-ordinal relationshipsof the Dermaptera. In contrast with previous studies(Giles, 1963; Popham, 1985) we found many differencesamong dermapteran families and recorded 18 new phylogenetically informative characters, thus almost doubling
the existing data set ofHaas (1995).
Morphological complexes and their functions are pro­foundly interwoven. Many wing character states
described here cannot be interpreted without considering
their function. They were examined with the help of
paper models, which duplicate the natural movements ofsclerites and fields during folding and unfolding. We
offer figures of these models with folding instructions,which make it possible for an interested reader to repeat
our experiments. Working paper models are the best way
to demonstrate and learn the movements involved in thewing folding mechanism. They also reveal functional rea­sons for many unusual characters of Dermaptera, such as
the large membranous area in the centre of the articula­tion. A paper model of the hindwing made to our instruc­tions will fold and unfold with some help, the same way
as do the hindwings ofliving earwigs.
MATERIALS AND METHODS
Nomenclature and homology
The neopterous wing characters analyzed here (Table 6) sug­gest a grouping of the related higher taxa (orders) into fivesuperordinal lines. To avoid confusion, we follow the proposalof JKP (in Kukalová-Peck & Brauckmann, 1990) to use the pre­fixes of the traditional and generally known names, plecopteroids, orthopteroids, blattoids, hemipteroids, endopterygotes
combined with the ending -neoptera, to identify the superordinallines:
Pleconeoptera: Plecoptera, Embioptera.
Orthoneoptera: Ensifera, Caelifera, Phasmatodea.
Blattoneoptera: Grylloblattodea, Zoraptera?, Dermaptera,
Mantodea, Isoptera, Blattodea.
Hemineoptera: Psocodea, Thysanoptera, Phthiraptera, Sternorrhyncha, Coleorrhyncha, Heteroptera, Fulgoromorpha, Cicadomorpha.
Endoneoptera: Hymenoptera, Coleoptera, Strepsiptera, Neuroptera, Megaloptera, Raphidioptera, Mecoptera, Diptera,
Siphonaptera, Trichoptera, Lepidoptera.
The all-pterygote homologization of the wing structures was
based on comparisons between many representatives of all
pterygote orders, extant and extinct. The previously publishedhypothetical pterygote protowing (Kukalová-Peck, 1983, 1997;
see characters listed in Table 6A) was recently found preserved
in Palaeozoic fossils (JKP, unpublished observation). It isretained in the prothoracic wing of extinct Palaeodictyoptera
and Geroptera (the most primitive extinct order ofOdonatoptera) (Kukalová-Peck, 1978, Fig. 14; Wootton et al.
1998: 599; Wootton & Kukalová-Peck, 2000). The prothoracic
wings in both orders are similar, in spite of the fact that they are
distantly related and their wings and flight are highly dissimilar.
Since the prothoracic wing pair was never involved in powered
aerial flight, this similarity very probably indicates that bothprothoracic wings are in a near original protowing condition.
The stunning resemblance between the hypothetical protowing
based on characters shared by all Pterygota and published in
1983, and the protowing-like prothoracic wings identified 15

years later in Carboniferous fossils, provides a proof, which isvery significant for higher phylogenetics. Namely, that the mor­
phological ground patterns, usually missing, can be reliably
reconstructed based on existing material. The key issue here isthat the study must be sufficiently broad. For a short review ofprotowing morphology, nomenclature and use in the relation­
ship of orders see Table 6. We use the all-pterygote wing
nomenclature developed by Kukalova-Peck (1983, 1985, 1991,
1997). The neopterous articular sclerites, unique folds, wing
lobes, veinal stems and braces are presented in a special Neoptera reference scheme inFig. 1.
Neopterous superorders and orders relate to each other in thesimilar transformations of their wing complexes. Typically ineach order, several groups of wing characters show the same
relationship (Table 6 B-H). Thus, true synapomorphies can berecognized by treating the wing complex in each order as aseparate entity, and by always judging its characters in thismutual context (see examples in Table 6). This context is neces­sary to overcome rampant homoplasies. Most wing characters
are extremely homoplasious in many unrelated higher taxa.
When extracted from their special, integrated wing complexes
and assessed as individual “neopterous” characters, most wing
characters loose their phylogenetic information and instantly
turn homoplasious. This explains why, in systematic practice,
the direct application of a successful species-level methodology(no matter how sophisticated) to order-level phylogenetics,
never worked in the past and never will in the future.
For determining derived character states we use the following
criteria, verified by a century of studies of the wing structure infossil and living insects.
(1) The pterygote principal veins and branches, once lost, do notreappear.
(2) The principal veins, once fused into veinal stems or one withanother near the wing base, do not separate.
(3) The pterygote articular sclerites, once reduced, do not reap­pear.
(4) The articular sclerites, once fused, do not unfuse.
For an unreduced wing system and accurately homologizednomenclature in Neoptera, see the reference scheme in Fig. 1.
For the arrangement and nomenclature of veins and folds in the
Dermaptera hindwing see Fig. 2. The abbreviations used in the
text and figures are given in Table 1.
A reviewer proposed to us that a distinction between a fold
and a flexion line should be recognized. The distinction was
introduced by Wootton (1979) to differentiate between folding
lines, which are important for folding, and flexion lines, whose
main function is in flight. We do agree with this distinction andsee the usefulness of it. However, in order to keep our text andfigures consistent, we here use claval fold instead of the moreexact expression claval flexion line or claval furrow.
Techniques
To examine the wing venation and articulation, allspecimens, either previously preserved in alcohol or pinned,
were softened with hot water. After about 10 minutes they weresoft enough to allow unfolding of the hindwings, which were
not removed from the body. Wings were fixed on black foamusing insect pins to prevent automatic folding of the wing. Afterdrying they were examined using dissecting microscopes atmagnifications of 10x to 70x. A camera lucida was used fordrawings. It was not possible to present all parts of wings andwing articulations in exact proportions because of their irregular
topography. In some cases, especially for the wing articulation,
alcohol-preserved specimens with folded wings were used forthe undistorted shape of the costal area and some articular sclerites.

Species examined
Original observations were made on the hindwing venation
and articulation of the species listed in Table 2. The fossil taxa
used are listed in Table 3.
Characters of extant Dermaptera taxa
Autapomorphies are known for only four out of nine“families” (Haas, 1995). We assume that the currently used“families” of the Dermaptera are monophyletic. The assumptionof monophyly is a working hypothesis, which we followed in
the present account as long as the character states within a “fam­ily” were homogeneous; otherwise the species were listed sepa­
rately. Although many differences in wing articulation andvenation are described below, not all of them are included in thephylogenetic analysis. Only those are used which appeared inclearly distinguishable character states, independent of bodysize and general sclerotization of the specimens.
Many Dermaptera are wingless, such as the Karschiellidae,
and in some cases not all characters for winged species wereavailable for our study because the specimens were previously
damaged or a dissection was not allowed. The “collection
dilemma” emerged, whether rare specimens should be kept
intact for future generations, or examined and partiallydamaged. Fortunately for this paper, the policy of the Academyof Natural Sciences, Philadelphia was in favour of scientificadvancement and dissection.
Apart from the venation and articulation characters described
here, we include additional characters presented by Haas (1995).
These characters from thorax, legs and genitalia were assessedwith the available species. Only changes in characters used by
Haas (1995), such as character states for specific taxa, are indi­cated. For a detailed discussion of these characters we refer tothat publication. Three species of Blattodea (Leucophaea mad­erae, Pofyphaga aegyptiaca, Periplaneta americana) are used
as an outgroup. However, for many wing characters there is nosuitable outgroup, because the dermapteran hindwings are
unique in their structure. The character state distribution is listed
in Table 4.
Characters of fossil Dermaptera taxa
Fossil Dermaptera are often grouped in the taxon “Archidermaptera”. The relationships of fossils to extant Dermaptera is
re-assessed, predominantly with body characters, because wing
characters are almost never preserved. The characters of thefossil taxa (Table 5) have been extracted from the literature(Martynov, 1925; Tillyard, 1931; Carpenter, 1933, 1992; Car­penter & Kukalova, 1964; Jun-feng, 1994; Vishnyakova, 1980).
It was not possible to re-assess the characters because the fossils
themselves were not available to us. Therefore, the characters
are not discussed and we follow the original descriptions. In
cases when a fossil Dermaptera genus contains several species,
only one representative species was selected to keep the numberof taxa low for the computer analysis. The other species were
not included. Only one species of extant Dermaptera, Diplatys
jacobsoni, was included to reduce calculation time. Periplaneta
americana was used for outgroup comparison. The character
state distribution is listed in Table 5.
Analysis
The data sets for extant and fossil taxa are analysed separately
because the wing venation and articulation, on which most char­acters in extant taxa are based, are not preserved in fossil Der­maptera and “Protelytroptera”. A parsimony analysis for eachdata set was conducted with PAUP 3.1 (Swofford, 1993) using
the “Heuristic Search Option” and assuming “Accelerated
Transformation” (ACCTRAN). Multiple character states wereuncertainties (not polytomies), and “Outgroup Rooting” was447Table 1. The following abbreviations are used in the text and the figures. Small initial letters indicate a cross vein, cross-vein
associated structure, of fold.
Abbreviation

Full spelling

1, 2, 3, 4Ax
AA,AP
abr
af
ap3-ap4
ap4-ja1+2

First, second, third, fourth axillary
Anal anterior, anal posterior
Anal brace
Anal fold
Cross-vein between AP3 and AP4
Cross-vein between AP4 and JA1+2
Anterior wing process (PRSc + PRR)AWP
AX, AXA, AXAJ
Axalare, anal axalare, anojugal axalare (3Ax saucer)
AXC, AXCu, AXJ
Costal axalare (convenient abbreviation for precosto-costal axalare AXPCC, used in this paper), cubital
axalare (3Ax goblet and heel),jugal axalare
Precosto-costal axalare
AXPCC
AXM, AXR, AXSc,
Medial axalare, radial axalare, subcostal axalare
B, BA, BAA, BAA1+2, Basivenale, anal basivenale, anal anterior basivenale, anal anterior 1+2 basivenale (anal brace), anal anteBAA3+4
rior 3+4 basivenale (articulation point between 3Ax anal arm and anal wing lobe)
BAP, BC, BCu
Anal posterior basivenale, costal basivenale (abbreviation for BPCC used in this paper), cubital basivenale
BJ, BM, BMA
Jugal basivenale, medial basivenale, medial anterior basivenale (a subdivision of medial basivenale)
BMP, BR, BSc
Medial posterior basivenale, radial basivenale, subcostal basivenale
BScA, BScP, BPCC
Subcostal anterior basivenale, subcostal posterior basivenale, precosto-costal basivenale (abbreviated as
BC in this paper)
BT
Basitarsus
Costa, costa anterior, costa posterior
C, CA, CPcf
Claval foldclf
Concave longitudinal fold
Cu, CuA, CuP
Cubitus, cubitus anterior, cubitus posterior
Coxa
CX
excl, exc2
Extension cross-vein 1, extension cross-vein 2
F, FA, FAJ, FC, FCu
Fulcalare, anal fulcalare (3Ax anal arm), anojugal fulcalare (3Ax anojugal arm), costal fulcalare (precostocostal fulcalare, abbreviation for FPCC used in this paper), cubital fulcalare (3Ax cubital arm)
FJ, FM, FMCu, FPCC Jugal fulcalare (3Ax jugal arm), medial fulcalare (part of medial plate), medio-cubital fulcalare (part ofmedial plate), precosto-costal fulcalare (abbreviated asF C in this paper)
FR, FSc
Radial fulcalare, subcostal fulcalare
FE
Femur
HP
Humeral plate (composed ofFC + BC)
Intercalary branchib
J, JA, JP

Jugal,jugal anterior,jugal posterior

jbrJugal bracelfLongitudinal fold

M, MA, MPMedia, media anterior, media posteriormfMedial fold

mp-cuaMedio-cubital cross-vein, also brace or arculus in this paper

PAT
PC
PFE
PR, PRA, PRC

PatellaPrecosta or precostal strip
Pefemur
Proxalare, anal proxalare, costal proxalare (abbreviation for PRPCC used in this paper)

PRCu, PRJ, PRM

Cubital proxalare,jugal proxalare, medial proxalare

PRR, PRSc, PRPCC

Radial proxalare, subcostal proxalare, precosto-costal proxalare (abbreviated as PRC in this paper)

PWP

Posterior wing process (PRA + PRJ fused with the tergum)

R, RA, RP

Radius, radius anterior, radius posterior

448

Table 1 (continued).
Abbreviation

Full spelling

rf, rbc
rc, rfc
Sc, ScA, ScP
SCX
tf
TI

Ring fold, ring brace
Ring cross-vein, ring fold cross-vein
Subcosta, subcosta anterior, subcosta posterior
Subcoxa (flattened into thoracic pleuron in pterygote thorax)
Transverse foldTibia

used. In phylogenetic reconstruction the characters are equallyweighted, non-additively coded and not polarized. Charactertransformations were examined using MacClade 3.04 (Maddison & Maddison, 1992).
Movement of wing articulation
The articulation sclerites were observed during experimental
promotion and unfolding of the wings in alcohol-preserved
specimens of Labidura riparia, manipulating them under a dis­secting microscope. The results were compared with those of
Kleinow (1966) on Forficula auricularia.
RESULTS
OrderDermapteraKirby, 1815.
Occurrence: Jurassic to Recent, worldwide.

Diagnosis of the hindwing
See characters in Table 6 and Figs 2-10, 18, 19. Char­
acters of the higher taxa (order and up) are dealt with
only at the ground pattern level. Character states below
the order-level are irrelevant for the phylogenetic positionof Dermaptera. Only the characters of dermapteran fami­lies are included. Dictyoptera, whose wing complex isclosely related to Dermaptera, are treated here as the sistergroup. Zoraptera, which may be related to Dictyoptera,
have petiolate wings and their wing articulation is notknown; they could not be included into this analysis (see“Discussion”).
The hindwings of Dermaptera differ from those of thesistergroup Dictyoptera (Mantodea + (Blattodea + Isoptera)) (Table 6E) mainly in the character states accom­
modating the quadruple wing folding. This adaptation
includes a very narrow remigium with condensed andsimplified radial and medial sectors; anojugal lobe excep­
tionally enlarged and supported by AP branches radiatingfrom its centre; arculus (mp-cua) present in both wingpairs; and AA1+2 absent. The wing articulation differs in
PRR fused with 1Ax tail; 2Ax body reduced to a thick­ened proximal margin; 3Ax goblet and saucer long,
slender and disconnected; 3Ax anal and jugal arm fused
into a large, posteriorly extended anojugal arm; medialplate desclerotized; BSc very long, thin in the middle, pli­able; BM proximally membranized; anojugal basivenalia
fused into a plate; and, in a unique anal brace overriding
the cubital basivenale (see below).
Dermaptera share with their sistergroup Dictyoptera a
distinctive, very complex flight apparatus (see Table 6 E).
This includes wing position, shape and lobes, flight mus­culature, and fusions, reductions and braces in the wing
venation and articulation. This flight apparatusbelongs to
the ground pattern of these taxa and is phylogeneticallyinformative (Figs 1, 4-13, 20-22; see discussion below).
Please note that individual characters usually make sense
only if used within the context of their own order; if
extracted from the order and compared anonymously as
neopterous characters, they would turn homoplasious anduninformative.
The basal flight adaptations in Dermaptera and Dicty­
optera are related to those of Hemineoptera + Endoneoptera (Figs 16, 17; Table 6, clade 5). In contrast,
Pleconeoptera + Orthoneoptera (Figs 14, 15, 21, 22)(Table 6, clade 2) have a very different flight-generating
system and probably represent the sistergroup of Blattoneoptera + (Hemineoptera + Endoneoptera). Note thatthe same deep split is repeated in the construction of thethird ovipositor valve (Sharov, 1969, Fig. 77, 78; Hennig,1981). The Dermaptera hindwing characters are offeredin Table 6, documented in Figs 2, 4-10, and discussed in
more detail in a separate section below.
Description of the hindwing
For plesiomorphic character states at the Neoptera levelsee the reference scheme in Fig. 1. For character coding:
0 plesiomorphic; 1, 2, 3 apomorphic, unordered, see
Table 6. The ancestral character states of Dermaptera are
best preserved in Apachelytron transversum, Lower Per­mian, Czech Republic (Fig. 3), which we consider to be a
representative of the extinct dermapteran stem group andclose to the stem line.
Fustis. The fustis (Latin: club-shaped) is a unique, dis­tinctive, sclerotized part of the dermapteran remigium
delimited anteriorly by RA, distally by the apical marginof the ancestral remigium and by the transverse fold, andposteriorly by CuA and CuA1+2 (Figs 2, 4). The fustis isdivided into base and head, and bisected by a groove inSpongiphoridae, Chelisochidae and Forficulidae (Fig. 4).
The head is the visible part of the fustis when the hindwing is at rest and covered by the tegmen. The groove inthe fustis marks the posterior margin of the tegmen andseemingly gives the wing package additional flexibility tofollow the movements of the abdomen. The fustis often
contains variously long RP, MA and a strong MP, and abroad to very broad CuA1+2.
Stem line homologue. The portion of the wing in
Apachelytron transversum, which evolved into the fustis
in extant Dermaptera, is marked by striation (Fig. 3).
Precostal strip (PC). The precostal strip is broadened
anteriorly into a soft flap in Apachyidae, Anisolabididae,“Labiduridae”, Forficulidae, Chelisochidae and Spongi­
phoridae (Fig. 4).
449

Table 2. Extant taxa examined in this study.

Table 3. Fossil taxa used in this study.

Taxon

FigurePleconeoptera: PlecopteraBurmeister, 1838
Eustheniaspectabilis Gray, 1832
Orthoneoptera: CaeliferaAnder, 1936
Valanga irregularis (Walker, 1870)
Trimerotropis suffusa Scudder
Blattoneoptera:
Isoptera Brullé, 1832
Mastotermes darwiniensis Froggatt, 1896
MantodeaBurmeister, 1838
Polypsilota aeruginosa (Goeze, 1765)
BlattodeaBrunner, 1882
Leucophaea maderae (Fabricius, 1775)
Periplaneta americana (Linnaeus, 1758)
Polyphaga aegyptiaca (Linnaeus, 1758)
Dermaptera Kirby, 1815
AnisolabididaeVerhoeff, 1902
Carcinophora americana (Palisot deBeauvois, 1817)
Apachyidae Verhoeff, 1902
Apachyusfeae Bormans, 1894
Chelisochidae Burr, 1907
Chelisoches morio (Fabricius, 1775)
Chelisochellasuperba (Dohrn, 1865)“Diplatyidae” Verhoeff, 1902
Diplatysgerstaeckeri (Dohrn, 1863)
Diplatysjacobsoni (Burr, 1911)
Haplodiplatys severus (Bormans, 1893)
Haplodiplatys bhowmiki (Srivastava
Sava, 1975)
Forficulidae Stephens, 1829
Allodahliascabriuscula (Serville, 1839)
Ancistrogasterglobalis Steinmann, 1993
Ancistrogasterspinax Dohrn, 1862
Forfícula auricularia (Linnaeus, 1758)“Labiduridae” Verhoeff, 1902
Allostethus indicum (Burmeister, 1838)
Forcipula clavata Liu, 1946
Labidura riparia (Pallas, 1773)
Nala lividipes (Dufour, 1829)“Pygidicranidae” Verhoeff, 1902
Echinosoma sp.
Echinosoma micropteryx Gunther, 1929
Pyragrafuscata Audinet-Serville, 1831
Tagalina burri Hincks, 1915
SpongiphoridaeVerhoeff, 1902
Labia minor (Linnaeus, 1758)
Marava arachidis (Yersin, 1860)
Sparattapulchra Borelli, 1906
Spongiphora crocreipennis AudinetServille, 1831
Vostox brunneipennis (Audinet-Serville,
1839)
Hemineoptera: FulgoromorphaLeach, 1815
Copidocephala sp.
Endoneoptera: MegalopteraLatreille, 1802
Protochauliodes sp.

14, 22IS, 21, 22

1S“Protelytroptera” Tillyard, 19S1
Acosmelytron delicatum Tillyard, 19S1
Apachelytron transversum
Carpenter & Kukalová, 1964
Protelytronpermianum Tillyard, 19S1

Figure andreference
1SS
19, 2S; Til­
lyard, 19S1

Dermaptera Kirby, 1S1S
12
11,20,21,22

2S, 26, 2V
4E, SE, SC

4D, SD, SB
4H, 6D, 9C

4A, SA, VA

4G, 6B,9B

4F, 6A, 9A
4C, SC, SA

4B, SB, VB-D

41, 6C, 10

16
1V

Sistergroup homologue. In Dictyoptera, the precostal
strip is not broadened, but is incorporated into the costal
margin (Figs 11-13).
450

Taxon

Archaeosomaserratum Jun-feng, 1994
Archidermapteron martynovi
Vishnyakova, 1980

Jun-feng,
1994
Vishnyakova,
1980

Asiodiplatys speciosus Vishnyakova, 1980 Vishnyakova,
1980
Dermapteron incertae Martynov, 1925
Martynov,
1925
Longiceratus mesozoica Jun-feng, 1994
Jun-feng,
1994
Microdiplatys campodeiformis
Vishnyakova,
Vishnyakova, 1980
1980
Protodiplatysfortis Martynov, 1925
Martynov,
1925
Semenoviola obliquotruncata
Martynov,
Martynov, 1925
1925
Semenovioloides capitatus
Vishnyakova,
Vishnyakova, 1980
1980
Turanoderma sepultum
Vishnyakova,
Vishnyakova, 1980
1980
Turanovia incompleta Vishnyakova, 1980 Vishnyakova,
1980

Comments. In Hemineoptera + Endoneoptera (Figs 16,17), the precostal strip is in many forewings of Hemiptera
and Endoneoptera broadened ventrally (Coleoptera); inthe hindwings, it is sometimes broadened anteriorly into amembranous lobe similar to that in Dermaptera(Kukalova-Peck & Lawrence, 1993). In contrast, Pleconeoptera + Orthoneoptera (Figs 14, 15) have the precostal
strip fused with the costa (C) in a uniform, not broadened
costal margin.
Costal margin (C, CA + CP), outer apical area. The
costal margin is unique, pliable, running to the end of thefustis. Apical area is a small, triangular area immediately
distal to the fustis (Figs 2, 7-10).
Stem line and sistergroup homologue. In Apachelytrontransversm, the costal margin is not pliable. The marginof the outer apical area beyond the fustis in Dermaptera(Fig. 2) belongs to an area between the anal fold and AP(Fig. 3). In Dictyoptera, the costal margin is also firm, notpliable.
Comments. In the forewing of Hemineoptera andEndoneoptera, CA and CP run parallel to each other (insome fossil and modern Hemiptera and in the elytra of theprimitive Coleoptera: Kukalova-Peck, 1991). In thehindwing, CA + CP form a well sclerotized costal margin.
Costal margin also occurs in both wing pairs of Pleco-Table 4. The character state distribution in extant taxa. “?”: Character not applicable or missing.
1
2
4
0
3
123456789 0123456789 0123456789 0123456789 0123

Taxon/ Character
Blattodea
Leucophaea maderae
Periplaneta americana
Pofyphaga aegyptiaca
Dermaptera
Karschiellidae“Diplatyidae”
Diplatys jacobsoni
Diplatys gerstaeckeri
Haplodiplatys bhowmiki? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?10??00000 0000000101 0???
? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?10??00000 0001000101 0???
? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?10??00000 0000000101 0???
? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ?? ?? 00 0 0 0 0000100111 1300
000000000 0100000000 0100011110 0000100111
000000000 0100000000 0000011110 0000100111
000000000 0100000000 0100001100 0000100111
000000000 0100000000 0100001100 0000100111

Haplodiplatys severus“Pygidicranidae”
Echinosoma micropteryx
Pyragrafuscata
Tagalina burri
Apachyidae
Apachyus feae
“Labiduridae”
Allostethus indicum
Forcipula clavata
Labidura riparia
Nala lividipes
Anisolabididae
Carcinophora americana
Spongiphoridae
Chelisochidae
Forficulidae

110202100 1101111101 1010111110 0010101010 1111
110211200
110221210
110221210
110221210

0100000100
1000111111
1000111111
1000111111

0010111110
1010111110
1010111110
1010111110

0011100010
0011100010
0011100010
0011100010

1111
1111
1111
1111

110210200
121231031
121231031
121221021

1110111111
2011121111
2011121111
1011121111

1010111110
1011111111
1011111111
1011111111

0011100010
1111100010
1111110010
1111120010

1110
1200
1201
1201

4
5
456789 01234

Extant Blattodea
Periplaneta americana

000000 00010

Fossil “Protelytroptera”
Acosmelytron delicatum
Apachelytron transversum
Protelytronpermianum

011000 010?0???000 0?010???001 1? ?? ?

Extant Dermaptera
Diplatysjacobsoni

111112 11213

Fossil Dermaptera
Archaeosoma serratum
Archidermapteron martynovi
Asiodiplatys speciosus
Dermapteron incertae
Longiceratus mesozoica
Microdiplatys campodeiformis
Protodiplatysfortis
Semenoviola obliquotruncata
Semenovioloides capitatus
Turanoderma sepultum
Turanovia incompleta

1020

100110000 0100000100 0100111110 0000100011 1010
100110000 0100000100 0100111110 0000100011 1020
100110000 0101100100 0100111110 0000100011 1010

Table 5. The character state distribution in fossil
Dermaptera, Diplatys jacobsoni and Periplaneta americana.
“?”: Character not applicable or missing.
Taxon/Character

1020
1020
1020

011012011102011102011002011002011102011102011012011012011012011002

11??3
11001
11001
11012
11??1
11001
11001
11113
11113
11113
11012

neoptera + Orthoneoptera. In Orthoneoptera forewings,
costa is desclerotized and pliable basally under the pro­thoracic shield. The long, branched ScA is supporting theanterior wing margin instead of costa (C). Therefore, theterm “precostal area” in Orthoptera is erroneous. The area
between the pliable costa and ScA is the costal area.
Subcostal basivenale (BSc). This sclerite is uniquely
shaped. Proximally it is strongly sclerotized, protruding,consisting of BScA and BScP primitively separated by asuture. BScA is narrow and pointed, BScP is broad andlobate, articulated in-between the prongs of lAx head.
The bulky proximal portion of BSc slims abruptly distally
into a narrow and flexible “neck” (Figs 7—l0).
Subcosta anterior (ScA). This sector forms a convex,
blunt ridge, which creates a unique, long overhang hiding
the ScP groove. The overhang occasionally contains rem­nants ofScA (Figs 7A, 8A, B).
Stem line and sistergroup homologue. ScA forms ashort, oblique, blunt ridge running between BSc and theanterior margin in Apachelytron transversum. The blunt
ScA ridge occurs also in Dictyoptera (in Blattodea +Isoptera and in blattoid stem line; it is broadened into abulge in Mantodea) (Figs 3, l l —l3).
Comments. In Hemineoptera + Endoneoptera the bluntridge of ScA (noticeable especially in the forewings)
extends into a broad, oblique bulge. In Hemineoptera, the451

Fig. 1. A - Reference scheme showing the basal character states in the wing complex at the Neoptera level, with emphasis on thehindwings; loci of folds and of future fusions and braces in venation are indicated (Kukalova-Peck, 1997). Neoptera developed sev­
eral styles of flight, each generated by a unique combination of different shape and size of wing pairs, wing lobes, flexion lines andfolds, veinal fusions, braces, reductions, enrichments, and by various fusions, reductions and extensions of articular sclerites (seeTable 6). Only the ground pattern level characters in each higher taxon are relevant in phylogenetics, but they are sometimes
obscured by younger modifications and must be detected. They are more often found in basal representatives, but may occur insome derived phenotypes. Only a very broad study of the entire higher taxon, and its prospective relatives, brings reliable results.

bulge is weak or absent. In Endoneoptera the bulge issclerotized, large, and flanked posteriorly by a prominentbrace called humeral vein. In contrast, Pleconeoptera +Orthoneoptera have a primitive, vein-like ScA. In
Orthoneoptera, ScA is very long with many branches andit supports basally the costal margin instead of the soft,
pliable costa (Fig. 15). In Pleconeoptera, ScA isexpressed in the plecopteroid stem line, but is short andinconspicuous in modern Plecoptera (Fig. 14) andEmbioptera (JKP, unpublished observation).
Subcosta posterior (ScP). This sector is absent. Only ashort, deeply incised groove runs parallel with R and RA
and ends in the membrane beyond the middle of the fustis(Fig. 7A, B).
Stem line and sistergroup homologue. Apachelytrontransversum has only slightly shortened ScP (Fig. 3). In
ground pattern of Dictyoptera, ScP is relatively long (asin extant Mantodea; ScP is shortened in the blattoid stem
line and very short in extant Blattodea + Isoptera; Figs
11-13).
Costal area (marginal area). In Dermaptera, the costal
area (narrow area between the costal margin andsubcosta) is unique in being soft and pliable (Figs 7-10).

452

Stem line and sistergroup homologue. The costal area
in Apachelytron transversum is well sclerotized (Fig. 3).
In Dictyoptera, Pleconeoptera and Orthoneoptera, it isalso well sclerotized (Figs 11-15).
Radial basivenale (BR). In Dermaptera, this sclerite is
uniquely shaped, divided by an embayment into a shorter
proximal part, and an unusually long and swollen distal
part. The latter is fused with the medial basivenale (BM),from which it is separated by a deeply incised groove.
The proximal part of BR articulates with 2Ax, and the
distal part gives rise to the stem of radius (R).
Sistergroup homologue. In Dictyoptera, BR is also
immediately followed by the swollen radius (Figs 7,
11-13).
Comments. In Hemineoptera + Endoneoptera BR is fol­
lowed by the thick base of radius, as in Blattoneoptera. In
contrast, in Pleconeoptera + Orthoneoptera the base ofradius is relatively slender and more even (Figs 14-15).
Radial sectors RA, RP, radial stem R. Sectors RA
and RP form an extremely short stem of R (Fig. 7A). RA
is swollen basally and forms the anterior margin of thefustis (Fig. 4). RP separates from RA at the sclerotizedbase of the fustis and ends within the sclerotized head of
the fustis. RP is simple, accompanied by an adjacent MA,

Fig. 1. B - Neopterous reference scheme: Articular sclerites; arrangement in rows aligned with wing veins is color-coded. Rows
in Neoptera are partly irregular but the homologous rows in Palaeoptera are regular (Kukalova-Peck, 1983, 1997).

prealar bridge

precostal strip + CA

Tf-.arculus

remigium

J AXCu

PRCu
PRA v

AA1+2
AA3+4 / Pleconeoptera and/yOrthoneoptera lobe

/ postalar/ lobe i

Neopterahwing reterence scheme 'Blattoneoptera, Hemineoptera andEndoneoptera lobe

Fig. 1. C - Neopterous reference scheme: Articular sclerites; arrangement in columns is color-coded. Columns in Neoptera arepartly irregular but the homologous columns in Palaeoptera are regular (Kukalova-Peck, 1983, 1997).

453

which soon separates and runs close and in parallel to RP(Figs 4D-G).
Stem line and sistergroup homologue. The stem of R
(i.e. RA + RP fused) in Apachelytron transversum ismuch longer than in Dermaptera (Fig. 3). The stem of R
in ground pattern of Dictyoptera is extremely short (as inextant Isoptera; R is somewhat longer in the blattoid stem
line, Blattodea and Mantodea) (Figs 11-13; note that thebase ofRP in Isoptera is crossed by a fold and destroyed).
Comments. In the hemipteroid stem line the stem of R
is absent in some species (Kukalová-Peck &Brauckmann, 1992; Figs 30-35). In modern Hemineoptera, R is relatively long. In Endoneoptera the stem of R
is extremely short only in Coleoptera + Strepsiptera (JKP,
personal observation); R is longer in neuropteroids (Fig.
17) and mecopteroids as well as in Hymenoptera(Kukalová-Peck, 1991). In the plecopteroid stem line
(Paraplecoptera), the stem of R is absent in some speci­
mens (a plesiomorphy at the Neoptera and Pterygota
level); in extant Plecoptera, the stem of R is present but
relatively short (Fig. 14). In Orthoneoptera, the stem of R
is always well developed and strong (Fig. 15).
Medial basivenale (BM). In Dermaptera, this uniquely
derived sclerite forms a long triangle with a broad
proximal base. The proximal part of BM between theproximal part of BR and BCu is bent ventrally and membranised, thus giving more space to the complex move­ments of the wing articulation during wing folding. The
distal part of BM is fused with BR and crossed by anoblique furrow. BM narrows rapidly apically in all fami­lies except in “Diplatyidae” (Figs 4A, 7-10).
Sistergroup homologue. In Dictyoptera, BM is associ­ated with BR proximally and separated from it distally bya furrow (flexion line). The basal part ofMP running very
close to R and RP (largely retained in Mantodea, Fig. 13)
is completely obliterated in Isoptera and in Blattodea(Figs 11-12).
Comments. In Hemineoptera + Endoneoptera (Figs 16,17) the furrow that obliterates the base of MP in Blat­
todea is absent. BM is robust and associated with BR,with one significant exception. In Endoneoptera: Coleo­
ptera + Strepsiptera, the radial and medial loop (engaged
in folding and pulling the wing apex under the elytra) fallapart at base, one anteriorly, the other one posteriorly, toinitiate folding. Consequently, BR and BM are separated
by a unique, wide gap (this apex-folding mechanism inparasitic Strepsiptera has decayed, but its presence is stillapparent and shown in several characters (Kukalová-Peck& Lawrence, 1993; Kukalová-Peck, 1997).
Media anterior (MA). In Dermaptera, the anterior
medial sector (MA) fuses immediately at base with theextremely short R and RP in the sclerotized base of fustis.
RP + MA are visible as a separate vein and the separationof MA from RP is weakly indicated (Figs 7A, 8B, C, 9A,
B). The stem ofM is absent.
Stem line and sistergroup homologue. In Apachelytrontransversum, MA separates from RP near apex (Fig. 3).
In Dictyoptera, MA fuses basally with R, but the entry is
inconspicuous because disrupted by the fold between BR454

and BM. The entry of MA is traceable and verifiable inserial dissection (K.D. Klass & JKP, personal obser­
vation). MA eventually separates from RP (but, this sepa­
ration is often inconspicuous or reduced; Figs 11-13). In
all Blattoneoptera the stem ofM is always absent.
Comments. In Hemineoptera + Endoneoptera the stemof M is also always absent. The entry of MA into R (animportant synapomorphy shared with Blattoneoptera) is
quite inconspicuous, and the separation ofMA from RP isvaguely noticeable only in plesiomorphic members. In
contrast, Pleconeoptera + Orthoneoptera always bear thestem of M which forks clearly into MA and MP. This
important stem ofM is probably a synapomorphy.
Media posterior (MP). The MP is a strong, straightvein, which is uniquely unbranched and ending near thepostero-distal end ofthe fustis (Figs 4D, F).
Stem line and sistergroup homologue. MP in Apachely­
tron transversum is strong, directed obliquely and posteri­orly towards the end of the apical margin, and it bears asmall terminal fork close to the anal fold (Fig. 3). This
marks this location as the equivalent of the postero-distal
end of the fustis in Dermaptera. In Dictyoptera, MP baseis separated from BM by a fold (in Isoptera by a narrowfold; in Blattodea by a broad desclerotization; in Man­
todea by a narrow, oblique fold) (Figs 1, 11-13).
Comments. MP base is present in Hemineoptera +Endoneoptera in basal members (Figs 16-17). There has
been a long-lasting confusion about the interpretation ofmedia in the neopterous orders, caused by a consistently
incomplete preservation of MA and MP basally. The
widespread misinterpretation of M, MA and MP (whichshow the basal split in Neoptera superordinal lines) liter­
ally prevented the use of the wing venation in higher phy­logenetics. Homologization revealed that Blattoneoptera+ (Hemineoptera + Endoneoptera) lack the stem of M,
and Pleconeoptera + Orthoneoptera share the stem of M
(Kukalová-Peck, 1991, 1997; Kukalová-Peck & Brauck­mann, 1992) (Fig. 22; Table 3).
Medio-cubital brace (mp-cua, cross-vein or shortfusion), the arculus. This flight-important brace occurs
repeatedly in many taxa at many levels, as well as in bothfore- and hindwing in Dermaptera.
Stem line and sistergoup homologue. Arculus occurs in
Apachelytron permianum in both pairs of wings probably
as a synapomorphy with Dermaptera. In the blattoid stem
line, arculus is absent in both wing pairs (Fig. 20; Table
6D, G). In ground pattern of Dictyoptera, arculus is alsoabsent in both wing pairs (in fossil stem line Dictyoptera
and Isoptera only; in Blattodea and again in Mantodea, it
is absent in forewings, but present in hindwings) (Figs 3,
7-13; Table 6G).
Comments. In Hemineoptera + Endoneoptera (Figs 16,17) mp-cua brace is always present, as a synapomorphy.
Arculus is clearly absent in protowing. The strong evi­
dence for this assumption is that it is absent in the Blat­
toneoptera ground pattern, in the plecopteroid stem line
(Paraplecoptera, Liomopteridae: Kukalová, 1964a; Car­penter, 1992, his Figs 63, 64), and in the Palaeoptera
ground pattern (in Palaeodictyoptera: Kukalová-Peck,broadening ring foldFig. 2. Hind wing of Forficula auricularia showing veins, areas and folds. From Haas (1995) after Kleinow (1966).

1991). But, the arculus is present in modern Plecoptera,as an autapomorphy, and it is especially strongly
expressed in Orthoneoptera, which share a very long
fusion between CuA and M&MP (CuA fuses with M atbase and separates from MP apically) (Figs 14-15; Table
6B, C).
Cubital basivenale (BCu). Dermapteran BCu is iso­lated from the combined basivenalia BR + BM by anextensive proximal desclerotization of BM (Figs 7-10).
BCu is narrow, more or less desclerotized, and its distal
end tapers into an empty groove. Consequently, the analbasivenale (BA), acting as an anal brace, overrides thegroove and articulates directly with BM (instead of withBCu) (Fig. 25).
Sistergroup homologue. In ground pattern of Dictyoptera, BCu is also separated from the associated basivenalia BR + BM (only in Mantodea; in Blattodea + Isoptera, BCu is broadly fused with BR + BM) (Figs 11-13).
Comments. In Hemineoptera + Endoneoptera BCu is
separated from BR + BM by a gap. In Pleconeoptera +Orthoneoptera, BCu is also separated from BR + BM
(Fig. 14; Caelifera do not show this plesiomorphic char­acter state: Fig. 15). As a plesiomorphy, basivenalia arehinged rather than fused. Fusions occur repeatedly, andprovide informative characters at the order, family andsubfamily level.
The stem of Cu. Dermapteran Cubital stem is proxi­
mally membranized and pointed into a short, massive
spike (Figs 7-10).
Stem line and sistergroup homologue. In Apachelytrontransversum, the stem of Cu is longer than in Dermaptera

and not pointed (Fig. 3). In ground pattern of Dictyoptera,
the stem of Cu is very short (visible only in Mantodea; inBlattodea and Isoptera, the length of Cu is obscured
because CuP is lost) (Figs 11-13).
Comments. In Endoneoptera + Hemineoptera, the stemof Cu is always present, extremely short in Hemineoptera,and significantly longer in Endoneoptera as an autapo­
morphy. In Pleconeoptera, the stem of Cu is short. Cu is
totally absent in Orthoneoptera (in all representatives,
extinct and extant). This absence of the basal fusion
between CuA and CuP into a stem is a plesiomorphy at
the protowing (Pterygota), Neoptera, and Orthoneoptera
level (Fig. 15; Table 6A, C).
Emendation. The stem of Cu in Coleoptera was errone­
ously interpreted as “CuA”, by Kukalova-Peck & Law­
rence (1993) (see the footnote to Table 6H). Only after
this mistake was emended do Endoneoptera show theimportant basal split into Hymenoptera and ((coleopteroids + neuropteroids) + mecopteroids) (JKP, own obser­
vation, Table 6).
Cubitus anterior (CuA). The CuA runs close to MP
and forks at about the middle of the fustis into CuA1+2
and CuA3+4 (Fig. 4). CuA and CuA1+2 form the poste­rior margin of the fustis. CuA3+4 divides into a broad
fork, which ends uniquely on the anal sector AP, rather
than at the posterior wing margin.
Stem line and sistergroup homologue. In Apachelytrontransversum, CuA bears pectinate branches, but they end
similar as in Dermaptera, at the anal branch AA4 rather
than at the wing margin. In the blattoid stem line and Dictyoptera, CuA is dichotomously branched and ends on the455

Fig. 3. Representative of dermapteran stem line, Apachelytron transversum from the Lower Permian of Moravia, Czech
Republic. All dermapteran veinal fusions, braces, reductions and fold arrangements are present, but their expression is plesiomorphic. Left tegmen: Length 6 mm, width 2.2 mm. After Kukalova-Peck (1991), veinal interpretation revised.

posterior margin as in the other superordinal lines (Figs11, 20, 21).
Cubitus posterior (CuP). The base of CuP is unique,strong, perpendicular to CuA, arched or almost straight,
ending at the claval fold, and functioning as a strut (Figs7-10). The rest of CuP is short, not reaching the wing
margin; it may be replaced by a weakly sclerotized strip
flanking the claval fold and ending near AP, or absent(Figs 7-10).
Stem line and sistergroup homologue. In Apachelytrontransversum, CuP is also weak and ending on AA4 farfrom the wing margin, but it is somewhat longer than in
Dermaptera (Fig. 3). In ground pattern of Dictyoptera,456

CuP is very weak (as in extant Mantodea, CuP runs at thebottom of claval fold; CuP is absent in Blattodea + Isoptera) (Figs 11-13). In blattoid stem line (Fig. 20), thepresence of CuP at the bottom of claval fold is possible
but in a fossilized insect it cannot be securely confirmed.
In Blattodea, the weak vein proximal from claval fold(Fig. 11B), which sometimes descends into the fold, starts
from the BAA blood sinus. Therefore, it is positively
AA1+2 and it can never be “CuP”. Blood sinuses andtheir veins (veinal blood canals) are a single, continuous
structure supplying blood.
Comments. In Hemineoptera + Endoneoptera and in
Pleconeoptera + Orthoneoptera, CuP runs proximallyfrom claval fold. In Orthoneoptera, CuP is richly dichoto-Fig. 4. Dermapteran fustis. A - the fustis, base and head; B
is separated from the base by a groove.

mously branched, especially in the forewing (Table 6C).
Note that a richly branched CuP also occurs in the hemipteroid stem line (in so called “Protorthoptera”: KukalovaPeck & Brauckmann, 1992) and in modern Hemiptera:
Fulgoromorpha (Fig. 16; note that CuP forks distally
beyond the figured part). CuP lost its branches independ­
ently many times.

the remnants of principal veins. In Eudermaptera (G, H, I) the headAnal basivenale (BA), subdivisions BAA1+2,
BAA3+4, BAP; and jugal basivenale (BJ). The derma­
pteran BA subdivision BAA1+2, BAA3+4, BAP and BJ
are uniquely fused, consolidated into a single basivenal
plate, and separated by sutures. BAA1+2 is a large, trian­
gular sclerite with a shallow anterior embayment.
BAA1+2 is almost seamlessly fused with a narrow
BAA3+4. BAP and BJ are fused one to another and sepa457

Fig. 5. Dermapteran anojugal lobe: The modifications in veinal support are characteristic for the different taxa. Veinal support
fully homologized at the Neoptera level (5A).

rated by a deeply incised furrow from BAA3+4 (Fig. 7A).
The boundary between BAP and BJ is either barely
noticeable or protruding. In “Diplatyidae”, BAP is only
slightly curved in the middle and BJ is sloped ventrally(Fig. 7A). In Echinosoma micropteryx both BAP and BJ
are curved and separated by a protruding suture (Fig. 7C).
In all other examined species, BJ is curved and twisted(Figs 8-10). BJ articulates with the arm of 3Ax.

458

Sistergroup homologue. In ground pattern of Dictyoptera, anal basivenale is subdivided by folds into a weak
BAA1+2, well sclerotized but short BAA3+4, and alonger, bar-like BAP; jugal basivenale is articulated withthe jugal arm of 3Ax (as in extant Blattodea; in Isoptera,
BAA and BAP+BJ form long, broad plates and jugal arm(BJ) is absent; in Mantodea, jugal veins lost theirbasivenale and also their articulation with 3Ax by BJ)(Figs 11-13; Table 6F, G).

(C) Spongiphoridae(D) Chelisochidae

Fig. 6. Dermapteran anojugal lobe: The modifications in veinal support are characteristic for the different taxa.

Comments. In Hemineoptera + Endoneoptera, anal andjugal basivenalia are similar to those in Blattodea, butthey are undergoing reduction (Figs 16, 17; Table 6B, D).
Pleconeoptera + Orthoneoptera have a very different
arrangement of folds and fields in the anojugal fan: Allbasivenalia (BAA1+2, BAA3+4, BAP, BJ) are robust andAA3+4 and AP are long (Figs 14, 15, 21; Table 6).
Anal brace (abr). In Dermaptera, the tip of BAA1+2 isarticulated to medial basivenale (BM) and functions as
highly unusual anal brace.
Sistergroup homologue. In Dictyoptera and all other
Neoptera, the anal brace is between BAA 1+2 and the
distal end of Bcu.
Anal anterior (AA), anal posterior (AP). The anal
anterior branches (AA) are strongly reduced. AA1+2
branches are lost. The arrangement of AA3+4 branches is
unique. AA3+4 divides near base into AA3 and AA4.
The short, strong and strut-like AA3 functions as a bracearticulated, across the claval fold, with the equally strut­
like base of CuP. The rest of AA3 is lost, or it flanks theclaval fold as a weakly sclerotized strip. AA3 strip andCuP strip run along the claval fold (Figs 7-10). AA4,
after a short distance, becomes fused with AP (Fig. 7A).

A suture separating AA4 and AP is actually the anal fold.
The posterior anal sector (AP) is very strongly pectinately
branched and provides the main support to the large ano­
jugal lobe. AP basally diverges away from AA3+4, forms
a loop spreading into a window and returns to fuse with
AA4. The window is broad (Figs 7, 8A, C) or narrow(Figs 9, 10). The portion of AP distal from the window
may be present (Figs 7, 8A), or membranized (Figs 8B,
C, 9, 10). AP divides into AP1+2 and AP3+4 at about one
third of the wing length (Fig. 5A).
AP1+2 base, after diverging from AP3+4, is obscuredby the transverse fold (Figs 5A, B). AP1+2 sends out
seven branches. The terminal branch has a long fork, sothat the total number of so-called radiating branches iseight (the distal branch is numbered as 1st, and the mostproximal branch as 8th). The branches of AP1+2 arecrossed by transverse fold (Figs 2, 5, 6) separating its
branches from AP1+2, either at a longer distance (Figs
5B, C, D) or at a shorter distance (Fig. 6).
AP3+4 almost immediately divides into a broad fork of
AP3 and AP4. The base of AP3+4 is separated from thebase of AP1+2 by the transverse fold in the continuationof the anal fold, causing a weakened spot in AP1+2. The

459

Fig. 7. Dermapteran hindwing base, fully homologized. For comparison with Dictyoptera, see Figs 11, 12, 13.

broad fork of AP3 and AP4 is present only in “Diplatyidae” (Fig. 5A). In the other examined Dermaptera, thebase of AP3 is superimposed on the weakened spot in

460

AP1+2, so that the fork is absent. AP3 is located at anincreasingly greater distance from AP4 and closer to the
8th radiating branch of AP1+2 (compare Figs 5B-E, 6).

Fig. 8. Dermapteran hindwing base, fully homologized. For comparison with Dictyoptera, see Figs 11, 12, 13.

AP3 and AP4 are connected by a long, oblique cross-vein
ap3-ap4, flanked by a radiating fold.
The bend in AP4 is either simple (Figs 5A-C), or it
may be extended basally by an extension cross vein
(excl), so that the basal half of AP4 with the exc1 andap3-ap4 attached, resembles the letter “H” (Figs 5D, E,
6A, B). In Spongiphoridae and Chelisochidae, ap3-ap4
can be extended to the other side of AP4 (exc2), and in
this case, AP4 resembles a buttressed letter “H” (Figs
6C, D). AP4 is strongly indented at the ring fold (Figs 5,461

Fig. 9. Dermapteran hindwing base, fully homologized. For comparison with Dictyoptera, see Figs 11, 12, 13.

6A), then mildly curved before dissecting the ringcross-vein (rc). This curve is sinusoid and less pro­nounced in Spongiphoridae, Chelisochidae and Forficulidae (Figs 6B-D).
Stem line and sistergroup homologue. In Apachelytrontransversum, the anals are quite similar to those of462

Dermaptera. The AA1+2 is absent, AA3+4 very short(not reaching the wing margin), AA3 forms a brace with
CuP, and AA4 is simple and ends on the anal fold at
about two thirds of the wing length (Fig. 3). AA4 andAP1+2 are separate and not fused. AP1+2 sends off 10
branches, the equivalent of dermapteran radiatingFig. 10. Dermapteran hindwing base, fully homologized. For comparison with Dictyoptera, see Figs 11, 12, 13.

branches, and AP3+4 is twice forked. In ground patternof Dictyoptera, the AA area is narrow and all AA
branches are in a process of strong reduction (as in extant
Blattodea + Isoptera, AA1+2 is weak, sometimes entering
the claval fold, and AA3+4 bears few branches; in Mantodea, AA1+2 is absent and AA3+4 simple) (Figs 11-13;
characters in Table 6D-G).
Comments. Note that in Blattoneoptera, Hemineoptera
and Endoneoptera the AA area is combined withremigium in an extended remigio-anal flight unit, which
ends at the anal fold. Therefore, their following partial
anojugal lobe unit is supported only by the branches ofAP and J (Figs 11, 12, 17, 21B, 22). In contrast, in Pleconeoptera + Orthoneoptera, the remigial flight unit ends
at the claval fold. Therefore, their following full anojugal
lobe unit is supported by the branches of AA, AP, JA, JP(Figs 1, 14, 15, 21A, 22).
Inner apical area. This narrow, triangular area lies
between the outer apical area and the 1st radiating AP1+2
branch. It reinforces the anterior wing margin distallyfrom the fustis (Fig. 2).
Stem line, sistergroup homologue and Comments. In
Apachelytron transversum and in Dictyoptera, thehomologous area, very small and inconspicuous is present(Fig. 3). There is no homologue in other Neoptera.
Jugal anterior (JA), jugal posterior (JP), ring foldcross-vein (rfc). The jugal sectors JA and JP are dichoto­mously branched in a unique way. JA divides into JA1+2
and a short JA3+4, which ends on JP1+2. The JA fork
and the cross vein ja1+2-ja3+4 encloses the jugal cell,which is always crossed by the 10th radiating fold (Figs5, 6). The jugal cell is an irregular triangle with an
oblique base. Two cross veins are attached: A slightly
curved ring fold cross-vein rfc and ap4-ja1+2 (Figs 5A,B, D). In Anisolabididae and “Labiduridae”, the curvatureof rfc has an arched fold in the membrane towards the
middle of ap4-ja1+2, the end of which is sclerotized (Figs

5E, 6A). In the Eudermaptera (Spongiphoridae + Chelisochidae + Forficulidae), the triangle of the jugal cell is an
open rectangle with one broad base (Figs 6B-D). The rfc
is absent, replaced by a new arch composed of JA1+2
base, a part of ap4-ja1+2, and the sclerotized end of thearched fold (Figs 6C, D). In Forficulidae, the rfc is pre­sent but ap4-jp1+2 is shifted towards rfc, so that theymeet (Fig. 6B).
A cell enclosed by AP4, JA1+2, ap4-ja1+2 and a ringcross-vein rc is called the anojugal cell. This cell islarger than the jugal cell and more or less rectangular(Figs 5A, B, E, 6A). In Forficulidae, Spongiphoridae andChelisochidae, the anojugal cell is much larger than thejugal cell and irregular in shape (Figs 6B-D).
The dermapteran JP is absent and its two branches(JP1+2 and JP3+4) start in the membrane. JP1+2 is fusedwith the short JA3+4. Only the posterior part of JP3+4 ispresent in “Diplatyidae”, “Pygidicranidae”, Allostethus
indicum and “Labiduridae” (Figs 5A-C, 6A) and it isabsent in the other dermapteran taxa. The bent, posterior
end of JA3+4 and the jugal cross vein form a zigzag pat­tern in “Diplatyidae” and Allostethus indicum (Figs 5A,C) or they are aligned (Figs 5B, E, 6B-D).
Stem line and sistergroup homologue. In Apachelytrontransversum, jugal branches divide in a plesiomorphic
way. JA divides into JA1+2 and JA3+4, and JP divides
nto JP1+2 and JP3+4 (Fig. 3). In ground pattern ofDictyoptera, jugal branches JA and JP start from the jugal
basivenale BJ and are repeatedly forked (as in stem-line
blattoids and Isoptera; in Blattodea, JA branches are quite
variable and tend to become pectinate to fill out space inan enlarged area; in Mantodea, jugal basivenale (BJ) is
membranized) (Figs 11-13).
Comments. In Hemineoptera + Endoneoptera the jugal
branches are undergoing strong reduction, probably as asynapomorphy. In Pleconeoptera + Orthoneoptera, plesiomorphic jugal branches JA and JP start from BJ, and are463

Fig. 11. Blattodean hindwing base. For comparison with Dermaptera, see Fig. 7A, Isoptera Fig. 12, and Mantodea Fig. 13. A Blaberidae: Leucophaea maderae, hindwing; B - Polyphagidae: Polyphaga aegyptiaca, hindwing.

dichotomously branched (Figs 1, 14-17, 21; KukalovaPeck, 1983) (Table 6B, C).
464

Anojugal lobe. In Dermaptera, the anojugal lobe is partial. It starts at the anal fold and is supported only by theFig. 12. Isopteran hindwing base. Isoptera: Mastotermitidae: Mastotermes darwiniensis, hindwing.

branches of the posterior anal sector AP, and of the jugal
sectors JA and JP (Fig. 2). The lobe is very broadly
attached to the metanotum, so that the wing attachment is
actually much longer than the articulation. The functional
importance of this phenomenon is discussed below.
Stem line and sistergroup homologue. The anojugal
lobe in Apachelytron transversum, and in Dictyoptera,
has the same composition as in Dermaptera (Figs 3,
11-13, 19-22.
Comments. The partial anojugal lobe in Hemineoptera+ Endoneoptera has the same composition as in Blattoneoptera (Figs 3, 11-13, 16, 17, 22), but is usuallymuch smaller. Broad, fully veined lobe occurs only in thebasal representatives of Fulgoromorpha, Homoptera,Hymenoptera, Neuroptera, Megaloptera, Coleoptera,
Strepsiptera, and Trichoptera. Pleconeoptera andOrthoneoptera have full anojugal lobe. It starts at theclaval fold and is supported by all branches of the analand jugal sector, AA1+2, AA3+4, AP1+2, AP3+4,
JA1+2, JA3+4, JP1+2 and JP3+4 (Figs 14, 15, 22;
Kukalova-Peck 1991, 1997; Kukalova-Peck & Brauckmann, 1992).
Ring cross veins (rc), ring brace (rbc). In
Dermaptera, the unique chain of ring-shaped cross veinsends proximally with the rc between AP4 and JA. One or
two rc cross veins may be added between JA1+2, JP1+2
and the attachment to the body (Figs 2, 5, 6).

The ring brace (rbc) is a chain of ring cross veins (rc)spanning between the AP1+2 anal, jugal and intercalarybranches, which keeps them apart. It was formerly called“vena spuria” (Giles, 1963; Kleinow, 1966). The orienta­tion of the rbc, placed between the posterior jugal
branches and the end of JA3+4, varies. The rbc is zig­
zagged in “Diplatyidae” and Allostethus indicum (Figs
5A, C), but it is fully aligned in all other Dermaptera(Figs 5B, D, E, 6).
Folds and areas in Dermaptera hindwings
Claval fold (cf). The claval fold is shallow, empty, andvery short, starting at the cubital basivenale and ending
between the 8th radiating branch and AP3+4 (Figs 7-10).
Stem line and sistergroup homologue. In Apachelytrontransversum, the claval fold is somewhat longer than in
Dermaptera but it also ends far from the wing margin.
CuP probably descended to its bottom (Fig. 3). In groundpattern of Dictyoptera, the claval fold is directed towards
the posterior margin as in other Neoptera, but it isshallow, especially near the posterior margin, flanked dis­tally by CuP, and claval furrow is probably devoid ofveins. (In extant Mantodea CuP descends to the bottom
line of claval furrow; in Blattodea, the furrow is either
devoid of veins, or AA1+2 descends temporarily to its
bottom line; in Isoptera, it is devoid of veins; in blattoid
stem line it probably contained CuP (as in Carboniferous
forewings), but this observation needs additional verifica­tion).
465

Fig. 13. Mantodean hindwing base. For comparison with Dermaptera, Blattodea and Isoptera, see Figs 7-13. Mantidae: Polypsilota aeruginosa, hindwing.

Comments. In Hemineoptera + Endoneoptera thebottom line of claval fold is devoid of veins, shallow, andoften shortened, as in Dictyoptera (Figs 1, 15, 17, 21A).
In Endoneoptera the claval fold is flanked proximally by
the anal branch (Fig. 15). In Orthoneoptera + Pleconeoptera, the claval fold is deeply incised and serves as themain flexion line during flight (Figs 14, 15).
Anal fold (af). The anal fold is difficult to distinguishfrom numerous other wing folds (Figs 5-10). The anal
fold runs basally between AA3+4 and AP and crosses themembranous window, but its next section is obscured bya fusion between AA4 and AP and is expressed only as asuture. Anal fold reappears beyond the end of AA4 andruns for a short distance anteriorly close to AP andAP1+2, while becoming increasingly shallow until it dis­appears in the membrane.
Stem line and sistergroup homologue. In Apachelytrontransversum (Fig. 3), and in Dictyoptera (Figs 11-13), theanal fold separates two distinctive flight units: The
remigium combined with the largely reduced AA, and the
anojugal lobe supported by the AP and J branches. The
466

anal fold is the main fold along which the anojugal lobe
folds at rest underneath the remigium. In the most primi­
tive blattoid folding type, present in the Palaeozoic blattoid stemgroup and also in the extant Isoptera, theanojugal lobe folds flatly underneath the remigium as abook folds along a straight spine (Kukalova-Peck & Peck,1993).
Comments. In Hemineoptera + Endoneoptera (Figs 16,17) the most primitive hindwings in the early ancestral
hemipteroids (Paoliidae), with relatively small anojugal
lobe, are flexed backwards in an oblique, postero-lateralposition without crossing each other (probably somewhat
rooflike), and the anojugal lobes do not fold at all(Kukalova-Peck & Brauckmann, 1993). The “book-like”
folding of the anojugal lobe, similar to that in the blattoid
stem line and Isoptera, is found in modern Coleoptera andto a degree in Hymenoptera. Thus, the evolutionary steps
seem to proceed from “incomplete flexing, no folding” to“complete flexing, book-like folding”, which is then com­bined with irregular “fan-like folding” until the latter pre­vails. In contrast, in Pleconeoptera and Orthoneoptera, theBSc BR B

PRSc
PRR
2Ax

arculus RP+M

clavalfold

Plecoptera

Eusthenia spectabilis

full anojugal lobe
Fig. 14. Pleconeopteran hindwing base. Unlike in Dermaptera and Dictyoptera, full anojugal lobe is present. Plecoptera: Eustheniidae: Eusthenia spectabilis, hindwing.

anojugal lobe starts at the claval fold, the principal
flexion line in flight (Brodsky, 1994) (Figs 14, 15, 21,
22). The anal fold is one of several much less important
folds, and the lobe folds like a regular fan.
Following folds are autapomorphic for the Dermaptera
and cannot be compared with the stem line, sistergroup orother taxa described here.
Ring fold (rf). This autapomorphic, semicircular foldstarts at the wing articulation and ends at the anterior
wing margin distally from the inner apical area (Figs 2, 5,
6). The ring fold crosses straight or arched indentations in
AP1+2 branches (radiating branches) which are broad­ened and reinforced by sclerotized membrane. It is also
supported by broadenings in the intercalary branches (ib,Fig. 2), which are arched in the opposite direction. These
broadenings play a very important role in the automaticdermapteran wing folding (see below).
Radiating folds. These autapomorphic folds start fromthe bases of the radiating branches and end at the poste­rior wing margin (Figs 2, 5, 6). The area between two
adjacent radiating folds represents one sector of thehindwing fan, containing a radiating branch (AP1+2

branch), or an intercalary branch (ib). Radiating folds arenumbered according to the custom, starting with the most
distal fold next to the inner apical area and counting in a
proximal direction. The radiating folds alternate between
convex and concave positions, starting with the convex
fold proximal to the inner apical area.
Transverse fold (tf). This autapomorphic fold starts in“Diplatyidae” (Fig. 5A) proximally from the fork in AP
into AP1+2 and AP3+4, and cuts off the base of theAP1+2 branch. In all other Dermaptera, the base of
AP1+2 is obscured and tf crosses AP either distally (Figs
5B, C, E, 6A, C, D), or proximally (Fig. 6B) from thebase of AP3. Then, tf curves towards the distal end of thefustis. It dissects on its way the bases of the radial
branches either close to or somewhat distant from, their
AP stem and reaches the anterior wing margin between
the fustis and the outer apical area (Figs 2, 5, 6).
Longitudinal fold (lf). The autapomorphic fold lf starts
in the wing articulation and runs as a convex fold poste­rior to the squama (and posterior to AP). It flanks anteri­orly the ulnar area and crosses tf at the distal end of thefustis. Fold lf changes into a concave fold, continues run467

Orthoptera

Trimerotropis suffusa
Fig. 15. Orthoneoptera, Caeliferan hindwing base. Unlike in Dermaptera and Dictyoptera, full anojugal lobe is present. Ortho­
ptera: Caelifera: Acrididae: Trimerotropis suffusa, hindwing.

ning between the outer and inner apical area, and ends on
the anterior apical wing margin (Fig. 2).
Squama. This autapomorphic area is delimited anteri­orly by the anterior wing margin, distally by tf, posteri­orly by lf, and proximally by the wing articulation.
Squama forms the dorsal surface of the wing package atrest. The proximal part of the squama is covered by thetegmen and the uncovered distal part is the fustis head(Fig. 2).
Ulnar area. This autapomorphic area lies posterior tolf, proximal to tf, and anterior to rf. It is supported by
AP1+2 and AP4, and ap3-ap4 is close to its posterior bor­der. The ulnar area is the ventral surface of the wingpackage at rest. In the wing package the large anojugal
fan lies folded between the squama and the ulnar area
(Fig. 2).
Outer apical area. This autapomorphic area forms theanterior apical wing margin distal to the squama. It isdelimited posteriorly by lf and proximally by tf (Fig. 2).
Inner apical area. This autapomorphic area is delim­
ited anteriorly by lf, posteriorly by the anojugal fan, andproximally by the tf (Fig. 2).

468

Anojugal fan. This autapomorphic fan constitutes
almost the entire wing except the squama, ulnar area, andouter and inner apical area (Fig. 2).
Hindwing articulation
The following is the description of the articular ele­ments. The symbols for the precostal row of sclerites(PC) are left out for convenience, because in Dermaptera
the PC and C sclerites are all fused without a suture. In
contrast to the wing venation, the wing articulation is not
visible in any neopteran fossils, including Apachelytrontransversum.
Tergal fissures. The three neopteran tergal fissures,radio-medial, medio-cubital and cubito-anal (Fig. 1) arepresent in “Diplatyidae” (Fig. 7A) and occasionally inwell sclerotized Forficulidae (Fig. 9B). Most of the dermapteran specimens have only the radio-medial fissure(for example, Fig. 9C).
Sistergroup homologue. In ground pattern of Dictyoptera, the medio-cubital fissure is obscured and the radio­
median and cubito-anal fissures are weakly indicated (asin extant Mantodea and Isoptera; they are obscured in
Blattodea).

Sc BR- R+MA^M P
CuA1+2
CuA3+4Jntra"cubital fold

CuP

claval foldanal foldHemineoptera

Copidocephala merula

partialanojugal lobe

Fig. 16. Hemineopteran hindwing base. Partial anojugal lobe. Hemiptera: Fulgoromorpha: Copidocephalidae: Copidocephalamerula, hindwing.

Comments. The fissures are variably retained and also
obscured in Hemineoptera + Endoneoptera (Figs 16, 17)
and in Pleconeoptera + Orthoneoptera (Figs 14, 15).
Proxalaria, precosto-costal proxalare (PRC). The
PRC is a small sclerite near the tergum. PRC is alignedwith the tegula, the humeral plate (HP), and the costal
wing margin (all these elements in Neoptera are com­posed of the precostal and costal row fused together).
Sistergroup homologue. In Dictyoptera, and all other
Neoptera, PRC is similar as in Dermaptera (Figs 7-11).
Comments. PRC in Neoptera hindwings is
unconnected, variously reduced and insignificant. But,
the homologue of PRC is important in Palaeoptera(Kukalová-Peck, 1997, Fig. 19.7a, b; 19.8a, b) (Table
6A).
Subcostal proxalare (PRSc), radial proxalare(PRR), anterior wing process (PRScR). In Dermaptera,
the two proxalaria are widely separated one from another(rather than fused) and articulated with the tergum bymembrane (Fig. 7B). PRR is unique in being firmly
hinged or fused with 1Ax tail (PRM) (Figs 8-10).
Sistergroup homologue. In Dictyoptera, PRSc and PRR
are also primitively articulated with the tergum, but they
are fused together and separated only by a suture (Figs11-13).

Comments. In ground pattern of Hemineoptera +Endoneoptera, subcostal and radial proxalare are also
separated and articulated with the tergum by a membrane(as in extant Endoneoptera, Fig. 17; in extant Hemineo­
ptera these sclerites are variously reduced, Fig 16). In Ple­
coneoptera, PRSc and PRR are secondarily fused with the
tergum forming the anterior wing process. In Orthoneo­
ptera, the anterior wing process is reduced. Note that thesubcostal and radial proxalare were both traditionally, buterroneously, interpreted as a “tergal outgrowth”. Instead,they originate from the same anlage as the other articularsclerites and the wing itself (Stenzhorn, 1974; KukalovaPeck, 1983, 1997).
Medial proxalare (PRM). In Dermaptera, this sclerite
forms a very large tail of 1Ax. It articulates proximallywith the tergum and distally with 2Ax body (AXM).
Anteriorly it is uniquely firmly hinged with PRR. 1Ax tail(PRM) is separated by a suture from the fully fused 1Ax
waist + neck + head. The posterior margin varies fromconcave (Fig. 7A) to lobate (Figs 7B-D) and wavy (Figs
9C, 10).
Sistergroup homologue. In ground pattern of Dictyop­tera the 1Ax tail is much less massive (in extant Blattodea+ Isoptera; in Mantodea, the tail is very narrow, with alarge posterior embayment) (Figs 11-13).

469

Fig. 17. Endoneopteran hindwing base. Partial anojugal lobe. Megaloptera: Corydalidae: Protochauliodes fuscatus, hindwings.

Comments. The 1Ax tail in Hemineoptera + Endoneoptera and in Pleconeoptera + Orthoneoptera is mostly lessmassive than in Dermaptera and quite variable. No infor­
mative higher-level character states were found (Figs
14-17).
Cubital proxalare (PRCu). The dermapteran PRCu ispresent in “Diplatyidae” (Fig. 7A) as a long, narrowsclerite articulated to the tergum and to the 3Ax goblet(AXCu). In the rest of Dermaptera, PRCu is partly (Figs
7B-D, 8B, C, 9, 10) or fully (Fig. 8A) membranised,starting at its proximal end, so that 3Ax goblet sometimes
has a small remnant of PRCu attached to it (Fig. 9C).
Sistergroup homologue. In ground pattern of Dictyoptera, PRCu is membranised, only a strip of stiff mem­brane remains (as in some extant Blattodea only, Fig.
11A; PRCu is completely membranized in Isoptera andMantodea) (Figs 12, 13).
Comments. In Neoptera, PRCu is reduced or lost. It
occupies the same “window” into which 3Ax collapses
when wings are flexed over the abdomen. Besides
Dictyoptera, ribbon-like, pliable PRCu is preserved in
Hemineoptera, in Psocoptera (JKP, personal observation)
and Fulgoromorpha (Fig. 16). Disjunct fragments occur
in Endoneoptera in Megaloptera (Fig. 17) and Neuroptera(Ithonidae, Rapismatidae, Dilaridae) (JKP, unpublishedobservation). In Pleconeoptera and Orthoneoptera, PRCu
is completely lost (Figs 14, 15). The PRCu is well devel­
oped in Palaeoptera (Kukalova-Peck, 1983, 1997).

470

Anal proxalare (PRA), jugal proxalare (PRJ), poste­rior wing process (PWP or PRAJ). In Dermaptera, theposterior wing process (PWP) is membranized.
Sistergroup homologue. In Dictyoptera, the posterior
wing process is also membranised (Figs 11-13). This is
an important synapomorphy shared by all Blattoneoptera.
Comments. Membranized PWP, if found in Zoraptera,
would be very informative about their relationship. Notethat PWP is actually the fourth axillary (4Ax), whichbecame secondarily fused with the tergum. In Endoneoptera 4Ax is very rarely preserved, in Hymenoptera (JKP,
unpublished observation) and in Coleoptera: Gyrinidae(Kukalova-Peck & Lawrence, 1993). A well sclerotized
PWP was formed many times at many taxonomic levels.
In Hemineoptera PWP is highly variable (e.g., it is mem­
branized in some Fulgoromorpha, Fig. 16). In Orthoneoptera 4Ax is sometimes present; in Pleconeoptera PWP isfirmly hinged or fused with the tergum (Figs 14, 15).
Precosto-costal axalare (AXC), tegula. In all neopterous wings, including those of Dermaptera and Dictyo­
ptera, AXC is incorporated into the tegula, a sensory
organ.
Humeral plate (HP, FPCC + BPCC). The sclerites in
the composite sclerite HP are smoothly fused, HP is longand very narrow and it is extended posteriorly by a tri­
angle of stiff membrane (Figs 7, 8, 9A).
Sistergroup homologue. In Dictyoptera, the humeral
plate is similar to that in Dermaptera, also smoothly
fused, but without the posteriorly stiffened membrane(Figs 11-13).

Fig. 18. Representative of dermapteran stem line Permelytridae, Acosmelytron delicatum, Lower Permian of Kansas, USA. Basedon holotype no. 5416, Peabody Museum, Yale University, New Haven, USA. Left elytron length about 6 mm.

Comments. The neopteran HP is composed of four
fused sclerites, the precosto-costal basivenalia and theprecosto-costal fulcalaria (Fig. 1). Humeral plate some­times shows the boundaries between the original foursclerites (e.g., in the Coleoptera: Browne and Scholtz,
1996). HP is unique for the Neoptera, but other than this,not very informative for the higher-level relationships.
First axillary (1Ax). The 1Ax head (FSc) is broad,very large, and bearing two long projections, which areextended to articulate with the lobate part of subcostal
basivenale BScP (Fig. 7). The 1Ax neck (AXSc) andwaist (AXR) are slender. The head, neck and waist aresmoothly fused together, but they are still separated by asuture from the broad 1Ax tail (PRM, described above).
The important, unique feature is the firm hinge between
1Ax tail and PRR.
Sistergroup homologue. In Dictyoptera, 1Ax head issmaller than in Dermaptera, and is separated by a suturefrom a relatively longer and thinner neck and waist, and amuch smaller tail (Figs 11-13).
Comments. The 1Ax is an irregular, obliquely con­structed neopterous axillary cluster composed of sclerites
belonging to three rows (Sc, R, M) and three columns(PR, AX, F). The sclerites in 1Ax are primitively sepa­
rated by sutures (best retained in Neuroptera forewings,
in Ithonidae and Dilaridae, unpublished observation by
JKP). 1Ax always articulates with the subcostal
basivenale (Fig. 1). We did not find any characters thatwould reliably distinguish orders according to the mor­
phology of 1Ax.
Second axillary (2Ax). In Dermaptera, the 2Ax body(AXM) is reduced to a thickened proximal margin, arched

and twisted. The 2Ax arm (FR) is long, relatively thin andflexible with a small lobe in the middle (Figs 7, 8C) or
near the distal end (Fig. 9B).
Sistergroup homologue. In Dictyoptera, 2Ax body(AXM) is also curved, weakened distally, wrinkled
around muscular insertions and shaped like a human ear.
The 2Ax arm (FR) is also weakly sclerotized and flexible,
but thinner and shorter than in Dermaptera and the lobe in
the middle is absent (Figs 11-13).
Comments. In Endoneoptera + Hemineoptera, the 2Ax
body is also sigmoidally curved with a distinctly thick­ened proximal margin as an important synapomorphy.
The 2Ax arm broadens anteriorly into a prominent lobe.
In contrast, 2Ax in Pleconeoptera + Orthoneoptera isquite different. In Pleconeoptera (Figs 14) 2Ax bears alarge, triangular, well sclerotized body (AXM), and 2Ax
arm (FR) is articulated, short, slender, well sclerotized,
movable and resting on the triangle like a lid. The large,sclerotized triangle and lid also occurs in Orthoneoptera(but in Caelifera the arm is relatively broad and is par­tially fused with the body, Fig. 15). The typically neopterous 2Ax is an irregular, obliquely constructed axillarycluster composed of two sclerites belonging to two dif­ferent rows (R, M) and two columns (AX, F). The 2Ax
body (AXM) always articulates proximally with 1Ax tail.
2Ax arm (FR) always articulates distally with radialbasivenale (BR). The plesiomorphic condition is almost
certainly a triangular body and a short, movable, articu­
lated arm without any lobes. The 2Ax cluster offers sev­
eral informative characters for higher phylogenetics(Table 6 ).

471

Tillyard’s description, the fan is folded transversely along the ring fold and the veins have broadenings where they intersect the ringfold. The same is true in Dermaptera and we consider this to be a synapomorphy. After Tillyard (1931), veinal symbols according
to Fig. 1.

Medial plate (FM and FCu, separate or fused). The
medial plate is almost completely membranized. Wellsclerotized Dermaptera (Fig. 7A) show a faintly pig­
mented remnant of the medial and cubital fulcalare(FMCu) fused into a single medial plate (Figs 7B-D).
Sistergroup homologue. In ground pattern of Dictyoptera, the medial plate is composed of two separate sclerites that are less membranized (in Mantodea only; in Blattodea + Isoptera, FM and FCu are fused; Figs 11-13).
Comments. In Hemineoptera + Endoneoptera (Figs 16,17), FM and FCu are relatively well sclerotized, adjacentand separated by a suture. In Orthoneoptera, medial plateis very similar, only larger (Fig. 15). In Pleconeoptera,
FM and FCu are separate and well sclerotized, and FCu isvery broadly hinged with 3Ax saucer (Fig. 14). We foundthat the medial plate is quite variable and usually does notprovide reliable higher-level characters, but there are a
few exceptions, e.g., in Plecoptera (Fig. 14), and in
Coleoptera (Kukalova-Peck & Lawrence, 1993).
Third axillary (3Ax). In Dermaptera, the 3Ax cluster
is unique in many aspects. The 3Ax goblet (AXCu) is
extremely long and thin and completely detached fromthe 3Ax saucer (AXAJ). 3Ax heel (AXCu, distal lobe) is
separated from the goblet by a deeply incised fold, which
also traverses the saucer. The saucer bears, near the mid­dle, an anterior and posterior projection. The distal end of
the saucer is traversed by a deeply incised fold and then
expanded to provide a long articulation site, posteriorly
for the large anojugal arm (FAJ), and anteriorly for the
stiff membranous expansion (Fig. 7C). The arms of 3Ax
are transformed as follows: The cubital arm (FCu) is partof the medial plate and desclerotized. The anal and jugalarm (FA and FJ) are fused into an anojugal arm, the FAJ
head. This extends into the FAJ tail, a long, postero472

proximal extension formed by sclerotized membrane(Figs 7-10).
In “Diplatyidae”, “Pygidicranidae” and Allostethus
indicum the cap-like articulation of the saucer with FAJ isarched and bears a narrow jugal prong, which fits into a
concavity on FAJ head (Figs 7, 8A). In the remaining
Dermaptera the jugal prong is absent (Figs 8B, C, 9, 10).
The conspicuous dermapteran anojugal arm (FAJ) bears
an almost oval head and a tail. The head shows a faintsuture between FA and FJ in “Diplatyidae”, “Pygidicranidae” and Allostethus indicum (Figs 7, 8A). The tip of thehead bears an anteriorly bent head projection articulatedwith the large triangular basivenale BAA1+2. The head iseither enlarged, with a long narrow tip (Figs 8B, 9A), or
short with a narrow tip (Fig. 8C), or its articulation withthe 3Ax saucer is undulated (Figs 9B, C, 10). The tail of
FAJ is a broad, weakly sclerotized extension in “Diplatyidae”, “Pygidicranidae” and Allostethus indicum (Figs 7,
8A). In all other examined Dermaptera, it is strongly
sclerotized, compact and much narrower (Figs 8B, C, 9,10). The posterior margin of the head articulates withthree fused basivenalia, BAA3+4, BAP and BJ.
Sistergroup homologue. In ground pattern of Dictyoptera, the 3Ax goblet, heel and saucer are close to the
Neoptera reference scheme, but the anal arm FA is longand massive and the jugal arm very short (jugal arm ispresent only in Blattodea; in Mantodea the anal arm iswidened and articulating not only with the usual
BAA3+4, but also with BAP) (Figs 11-13).
Comments. In Hemineoptera + Endoneoptera (Figs 16,17), the anal arm is fused with 3Ax saucer, as a synapomorphy, and the jugal arm is lost. In Pleconeoptera andOrthoneoptera, 3Ax is close to the Neoptera reference
scheme, and the jugal arm is lost. The typically neo-Fig. 20. Palaeozoic Dictyoptera, stem line: The ancestral anojugal lobe. A - in the forewing, bar-like BAA locks wings when
flexed; reduced jugal lobe is tucked under the tegminous anal lobe; B - in the hindwing, veinal arrangement is radial, AA area is
combined with the remigium; partial anojugal lobe starts at the anal fold. Archimylacridae, Lower Permian, Kansas, USA. After
Kukalova-Peck, 1991, new interpretation.

pterous cluster 3Ax contains several phylogeneticallyinformative higher-level characters. The axillary is com­posed of six sclerites belonging to three rows (Cu, A, J)
and two columns (AX, F). In contrast with the obliquely
constructed 1Ax and 2Ax, its sclerites are arranged inregular columns. The 3Ax contains a rotating body, andthree arms. The 3Ax body is composed of three axalaria:
AXCu, AXA, and AXJ. AXCu is subdivided by a deep,
oblique fold into the proximal, large, protruding goblet,and the distal, small, sunken heel. AXA and AXJ are
fused into AXAJ to form a saucer. The goblet and theheel articulate with the saucer. The proximal end of thegoblet (AXCu) articulates with PRCu (Fig. 7A; note that
PRCu in most Neoptera is membranized). The distal endof the goblet articulates with the posterior tip of 2Ax
body (AXM). The 3Ax manipulates three arms formed bythree fulcalaria: FCu, FA and FJ. The 3Ax heel articulateswith the cubital arm FCu. The saucer (AXAJ) articulates
proximally with PRA and PRJ combined, primitively
expressed as 4Ax, but usually fused with the tergum as
the posterior wing process (PWP). The saucer articulates
distally with the anal arm (FA), and the jugal arm (FJ).
In Neoptera ground pattern, anal lobe is significantly
enlarged and anal basivenale is divided into BAA1+2,
BAA3+4 and BAP articulated one with another. The 3Ax
anal arm (FA) always articulates with BAA3+4. The
BAA1+2 serves in Neoptera as the anal brace with BCu.
The three arms of 3Ax articulate distally with three veinal473

Fig. 21. The two types of neopteran anojugal lobes in hindwings. A - in Pleconeoptera and Orthoneoptera, the full anojugal lobe
includes all pterygote branches: AA1+2, AA3+4, AP1+2, AP3+4, JA1+2, JA3+4, JP1+2, JP3+4; B - in Blattoneoptera, Hemineoptera and Endoneoptera the partial anojugal lobe starts at the anal fold and includes only AP and J branches; A - Orthoptera: Caelifera: Valanga irregularis; B - Blattodea: Periplaneta americana.

basivenalia: Cubital (BCu), anal (BAA3+4), and jugal(BJ). The pull of the wing flexor on the goblet makes the
3Ax body pivot, rotate and collapse into the membranizedarea vacated by the desclerotized PRCu, while the arms
pull at the basivenalia and simultaneously fold the ano­
jugal lobe backwards and over the abdomen. Deviations
from this generalized pattern provide numerous importantcharacter states, at many taxonomic levels. As an
example, Hemineoptera + Endoneoptera have anal arm(FA) fused with 3Ax saucer and protruding to articulate
with BAA3+4 (Fig. 16); in coleopteroids + neuropteroids,
FA changed into a narrow rim on the saucer, and the474

articulation with BAA3+4 was replaced by several
narrow strips of stiffened membrane (Fig. 17).
PHYLOGENETIC ANALYSIS OF THE EXTANT DERMAPTERA

Examination of the wing venation and articulation
added 18 new characters to the previously existing dataset (Haas, 1995). Also, more species have become avail­able to re-assess the previously used characters and theirstates. Most of the character states recognized for dermapteran taxa were confirmed, and only a few character
states had to be changed. Some of the previously used
terms are replaced here by more precise terms. Allchanges are indicated and included in Table 4.

supported by AA, AP and J branches is separated from the remigium by a concave claval fold / flexion line; B - in Blattoneoptera,
Hemineoptera and Endoneoptera the anojugal lobe contains only AP and J branches; AA area is combined with remigium into a
special, extended flight unit separated from the diminished anojugal lobe by the anal fold.

Characters and their states
Hindwing and wing articulation
1. Fustis, shape. Head narrow, slender and elongate (0)or head broad, disc-like (1) (Fig. 4). The fustis head isnarrow, slender and elongate in “Diplatyidae”; it is broadand disk-like in the other taxa. The character is not appli­
cable to the Blattodea and Karschiellidae.
2. Fustis, anterior margin. Smooth, without embayment (0) or with embayment (1) or with a notch (2) (Fig.
4). The fustis anterior margin is smooth and without
embayment in the “Diplatyidae” and “Pygidicranidae”;

475

This character is not applicable to Blattodea and Karwith an embayment in the “Labiduridae”, Anisolabididae,
Apachyidae and Allostethus indicum; with a notch only in schiellidae.
the Spongiphoridae, Chelisochidae and Forficulidae. This
9. Course of AP4. Angular, indented at ring fold (0) or
character is not applicable to the Blattodea and Karschielsinusoidal at ring fold (1) (Figs 5, 6). The curvature of thelidae.
AP4 is sinusoidal in the Spongiphoridae, Chelisochidae
3. Fustis, head. Not separated (0) or separated (1) by a and Forficulidae. This character is not applicable to Blat­groove from the fustis base (Fig. 4). The fustis head is todea and Karschiellidae.
separated by a groove only in the Spongiphoridae, Cheli­
10. AP4, ap3-ap4, excl and exc2. “Y”-shaped (0) or
sochidae and Forficulidae. This character is not appli­ “H”-shaped (1) or buttressed “H”-shaped (2). Haas
cable to the Blattodea and Karschiellidae.
(1995) and (Figs 5, 6). This composite of veinal portions
4. Costal area. Long and slender (0) or short and broad is “Y”-shaped in the “Diplatyidae”, “Pygidicranidae” and(1) or long and broad (2) (Haas, 1995) (Fig. 4). The costal Allostethus indicum, and “H”-shaped in the Apachyidae,
area is long and slender in the “Diplatyidae”, short and “Labiduridae” and Forficulidae. The buttressed “H”
broad in the “Pygidicranidae”; and long and broad in the appears only in the Chelisochidae and Spongiphoridae.
remaining taxa.
This character is not applicable to Blattodea and KarThis character replaces the formerly used character schiellidae.
Closer examination showed that the former distinction“marginal area” (Haas, 1995), because examination of
additional species indicates that the distinction between between two character states was insufficient. Besides the“present” and “absent” cannot be upheld, and a more “Y”-shaped situation, there are two other states and notdetailed distinction is possible now. This character is not only one (“4”-shaped), as assumed before (Haas, 1995).
In each of the new states, a small cross vein is added.
applicable to Blattodea and Karschiellidae.
5. End of CuA3+4. Between the 8th and 9th branch of
11. End of JA3+4 and two proximal rc cross veins.
AP1+2 (0) or between 7th and 8th AP1+2 (1) or between Fully aligned and straight (0) or zigzag patterned (1)6th and 7th AP1+2 (2) or between 5th and 6th AP1+2 (3).
(Figs 5, 6). The JA3+4 and rc are fully aligned and
The distal end of CuA3+4 lies between the 8th and 9th straight in “Labiduridae” (excluding Allostethus indicum),branch of AP1+2 in the “Diplatyidae” and Apachyidae; it Chelisochidae, Forficulidae and Spongiphoridae. Theylies between the 7th and 8th AP1+2 branch in Allostethus
form a zigzag pattern in all other taxa examined. This
indicum, the Anisolabididae and “Pygidicranidae”; it lies character is not applicable to Blattodea andbetween the 6th and 7th AP1+2 in the “Labiduridae” and Karschiellidae.
Forficulidae; and it lies between the 5th and 6th AP1+2
12. JP3+4. The veinal portion posterior to rc, present
branch only in the Spongiphoridae and Chelisochidae.
(0) or absent (1) (Figs 5, 6). The posterior portion ofThis character is not applicable to the Blattodea and KarJP3+4 is absent in the Anisolabididae, Chelisochidae,
schiellidae.
Spongiphoridae and Forficulidae; it is present in all other
6. Size of jugal and anojugal cells. About equal (0) or taxa examined. This character is not applicable to Blat­
anojugal cell larger than jugal cell (1) or anojugal cell todea and Karschiellidae.
smaller than jugal cell (2) (Figs 5, 6). Both cells are of
13. 9th intercalary branch (between AA3 and AA4).
equal size in the “Diplatyidae”, “Pygidicranidae” and Forked (0) or simple (1) (Figs 5, 6). This vein is forked in
Anisolabididae. The anojugal cell is larger than the jugal the “Diplatyidae”, “Pygidicranidae”, Allostethus indicum,
cell in all “Labiduridae”, Spongiphoridae, Chelisochidae “Labiduridae” and in Anisolabididae; it is simple in all
and Forficulidae; the anojugal cell is smaller than the other taxa. This character is not applicable to the Blat­
jugal cell in the Apachyidae. This character is not appli­ todea and Karschiellidae.
cable to the Blattodea and Karschiellidae.
14. 3Ax saucer, jugal prong. Large (0) or small (1) to
7. Bending zones of the AP1+2 branches. At the base absent (2) (Figs 7-10). Jugal prong is pronounced in the(0) or far from the base (1) or close to the base (2) (Figs “Diplatyidae”, “Pygidicranidae” except Tagalina burri,5, 6). The bending zones of the AP1+2 branches are and in Allostethus indicum; it is small in the Apachyidae,
located at the base in “Diplatyidae”, “Pygidicranidae”, Anisolabididae, and absent in the “Labiduridae”, Forficu­
Spongiphoridae, Forficulidae and Cheliosochidae; they lidae, Chelisochidae and Spongiphoridae.
occur at a small distance from the base in all
15. Anojugal arm FAJ, head. Almost elliptical (0) or“Labiduridae” and Anisolabididae; they occur at a great narrow and very long (1) or narrow, undulated and shortdistance from the base only in Apachyidae. This character (2) (Figs 7-10). The head of FAJ is almost elliptical withis not applicable to the Blattodea and Karschiellidae.
a narrow neck posteriorly in the “Diplatyidae”, “Pygidi­
8. Position of rfc. At jugal cell (0) or forming a com­ cranidae” and Allostethus indicum; narrow and very longplete arch (1) or in between jugal and anojugal cell (2) or in the Apachyidae, “Labiduridae” and Anisolabididae;“U”-shaped (3) (Figs 5, 6). The ring fold cross-vein rfc is and short, narrow and undulated in the Forficulidae,
attached directly to the anojugal cell in the “Diplatyidae”, Spongiphoridae, and Chelisochidae. This character is not“Pygidicranidae” and Apachyidae. It forms a complete applicable to Blattodea and Karschiellidae.
arc in the “Labiduridae” (except Allostethus indicum). It
16. Anojugal arm FAJ, neck and tail. Neck narrow,lies between the jugal and anojugal cell in the Forficuli­ tail broad and weakly sclerotized (0) or neck broad, taildae. It is “U”-shaped in Chelisochidae and Forficulidae.
narrow and strongly sclerotized (1) (Figs 7-10). The neck476

is narrow, the tail broad and weakly sclerotized in the“Diplatyidae”, “Pygidicranidae” and Allostethus indicum;
the neck is broad and the tail narrow in all other exam­
ined taxa. This character is not applicable to the Blattodea
and Karschiellidae.
17. Anojugal arm FAJ, tail. Posterior part of tail con­cave and strongly three-dimensional (0) or flat (1) (Figs7-10). FAJ is strongly three dimensional only in the“Diplatyidae”; it is flat in all other taxa. The character is
not applicable to Blattodea and Karschiellidae.
18. Combined basivenalia BAA3+4, BAP, BJ. Not orslightly twisted (0) or strongly twisted (1) (Figs 7-10).
Basivenalia BAA3+4, BAP, BJ are not or only slightly
twisted in the “Diplatyidae”, “Pygidicranidae”, Apachyidae and Allostethus indicum. They are strongly
twisted in the other taxa examined. This character is notapplicable to Blattodea and Karschiellidae.
19. Position of AA3. Far from BAA1+2 (0) or close to
BAA1+2 (1), Haas (1995) and (Figs 7-10). The AA3
diverges far distally from BAA1+2 in the “Diplatyidae”,“Pygidicranidae” and Allostethus indicum. The AA3
diverges close to BAA1+2 in the other taxa.
This character was formerly called “a cross vein: infront of or after bend” (Haas, 1995), which is a non­homologous term. The homologized terminology is now
employed, but no changes in the character assessment
were necessary. This character is not applicable to Blat­
todea and Karschiellidae.
20. Anal window, closure. Window closes proximal to
AA3 (0) or distal to AA3 (1) or window is open (2) (Figs7-10). Anal window is formed by a divergence and a sub­sequent fusion, of AA3+4 and AP; it closes proximally tothe base of AA3 in the “Diplatyidae” and“Pygidicranidae”; it closes distally to the base of AA3 in
Allostethus indicum, Apachyidae, Anisolabididae and“Labiduridae”. AP is membranized at the point of fusion
with AA3 in the Forficulidae, Chelisochidae and Spongiphoridae. This character is not applicable to Blattodea andKarschiellidae.
21. CuP. Distinct (0) or indistinct (1) (Haas, 1995). The
CuP is distinct in the Blattodea, “Diplatyidae” excluding
Diplatys gerstaeckeri, “Pygidicranidae” and Allostethus
indicum; CuP is indistinct in all other taxa examined (Figs7-10).
CuP was previously called Cu2. The Apachyidae are
now considered to have a short CuP. The character is notapplicable to the wingless Karschiellidae.
22. Concave longitudinal fold (clf). Median (0) or lat­eral (1) (Haas, 1995). The clf runs almost mediallybetween CuA and AP in the “Diplatyidae” and “Pygidi­
cranidae”; it runs closer to AP in all other taxa (Figs7-10). The clf is not present in Blattodea and Karschielli­dae; the character is not applicable.
As pointed out below, the clf is a system of two or threefolds. The functional term is used here for the combina­
tion of claval fold, anal fold and CuP.
23. Broadenings of radiating and intercalarybranches. Connected (0) or separated (1) (Haas, 1995).
The broadenings are separated only in the Chelisochidae,

Spongiphoridae and Forficulidae. This character is notapplicable to the Blattodea and Karschiellidae.
The broadenings were formerly called “broadened
areas”. The examination showed that the distinction
between the connected and separated broadenings is mostevident in the first three to four pairs of radiating andintercalary branches; no changes in character states werenecessary.
24. Branching of AP1+2. AP1+2 proximal branches
diverge very close together (0) or are broadly separated(1) (Figs 5, 6). Two proximal branches of AP1+2 diverge
close together in the “Diplatyidae”; they are well dis­
tanced in all other dermapteran taxa. AP1+2 branches areeither pectinate or dichotomously branched in the Blat­
todea (Figs 11A, B); the character is not applicable to the
Karschiellidae.
Thorax and tegmina
25. Tegmina. Symmetrical (0) or asymmetrical (1)(Haas, 1995). The tegmina are symmetrical only in the
Blattodea, all examined species of Haplodiplatys andKarschiellidae. Apart from those taxa they are asymmetri­cal.
26. Metanotum. Flat (0) or with median longitudinalgroove (1) (Haas, 1995). The metanotum is flat in the
Blattodea and Karschiellidae; it has a median groove inall other examined Dermaptera.
27. Spiny crest on tegmina. Absent (0) or present (1)(Haas, 1995). The spiny crest is absent in Blattodea andKarschiellidae, but present in the other examined Dermaptera.
28. Spiny ridge on metanotum. Absent (0) or present(1) (Haas, 1995). The spiny ridge is absent in the Blatto­dea, Karschiellidae and all examined species of Haplodi­
platys; it is present in all other examined Dermaptera.
The spiny ridge was formerly referred to as a tegminal
locking device. This term is now considered to be inap­
propriate because the tegminal locking device consists ofseveral structures not confined to the metanotum. The
combination of structures described here as characters 25
to 28 form the tegminal locking device.
29. Mesonotal ratio. High, over 0.55 (0) or low, under
0.54 (1) (Haas, 1995). It is calculated as the ratio ofmesonotal length divided by its width. A high ratio isfound in Blattodea and all dermapteran taxa except the
Chelisochidae, Spongiphoridae and Forficulidae, whichhave a low ratio.
The measurements for the Anisolabididae (Haas, 1995)
were based on a single specimen of Carcinophora ameri­cana. More specimens have become available and theratio was measured and calculated again. These newmeasurements (mean: 0.63; standard deviation: 0.03;
sample size: 5) show that the situation in the Aniso­
labididae was formerly misinterpreted as low and that
their mesonota have in fact a high ratio.
30. Median posterior tip. Lightly sclerotized and notpronounced (0) or heavily sclerotized and pronounced,
well marked (1) (Haas, 1995). A lightly sclerotised andunpronounced tip is found in all examined Dermaptera
except Chelisochidae, Forficulidae and Spongiphoridae,
477

which have a heavily sclerotised and pronounced tip. The
character state for Anisolabididae was changed from 1 to
0 after more material became available (see character 29).
31. Cranial margin of tegmen. Curved (0) or straight(1) (Haas, 1995). The cranial margin is straight in the
Chelisochidae, Forficulidae and Spongiphoridae; it iscurved in the other examined Dermaptera and Blattodea.
The character state for Anisolabididae is changed from 1
to 0 after more material became available (see character
29).
Neck, legs and abdomen
32. Neck. Blattoid-type (0) or forficuloid-type (1)(Haas, 1995). The Blattodea, Karschiellidae, “Diplatyidae” and “Pygidicranidae” posses a blattoid-type neck;
all other examined Dermaptera possess a forficuloid-type
neck.
33. Femur. Carinate (0) or rounded (1) (Haas, 1995).
The femur is carinate in Periplaneta americana, Kar­schiellidae, “Diplatyidae”, “Pygidicranidae” and Apachyidae. It is rounded in Leucopheae maderae,
Polyphaga aegyptiaca, Allostethus indicum, “Labiduridae”, Anisolabididae, Spongiphoridae, Chelisochidae
and Forficulidae.
34. Number of tarsomeres. Five (0) or three (1) (Haas,
1995). All examined Dermaptera have tarsi with three tar­someres, whereas the Blattodea have tarsi with five tarsomeres.
35. 2nd tarsomeres. Normal, not elongated (0) or longand slender (1) or heart-shaped (2) (Haas, 1995). All
examined Blattodea and Dermaptera have normal, notelongated 2nd tarsomeres, except the Chelisochidae pos­sessing a long and slender tarsomere and the Forficulidae
possessing a cordiform 2nd tarsomere.
3 6 .10th abdominal segment. Normal (0) or dilated (1)(Haas, 1995). The 10th abdominal segment is normal in
all taxa examined, except in the Apachyidae, which pos­sess a dilated 10th abdominal segment.
37. Larval cerci. Annulated (0) or smooth, not annulated (1) (Haas, 1995). The larval cerci are annulated in
Blattodea, Karschiellidae and “Diplatyidae”. They aresmooth and not annulated in all other Dermaptera exam­
ined.
38. Adult cerci. Annulated (0) or smooth, not annu­
lated (1) (Haas, 1995). The adult cerci are annulated in
Blattodea. They are smooth, not annulated in all Derma­
ptera examined.
39. Division of telson. Fused (0) or subdivided (1)(Haas, 1995). The telson or pygidium consists of threeparts which are fused in the Apachyidae, Allostethus indi­cum, “Labiduridae”, Anisolabididae, Chelisochidae, For­
ficulidae and Spongiphoridae.
40. Abdominal tergites and sternites. Abutting (0) or
overlapping pleurally (1) (Giles, 1963; Rentz & Kevan,
1991; Roth, 1991). The abdominal tergites and sternites
abut in Blattodea and overlap pleurally in all Dermaptera
examined.

478

Male genitalia
41. Number and direction of genital lobes. Twogenital lobes unidirected (0) or two genital lobes bidirected (1) or one genital lobe (2) or karschiellid-type (3)(Haas, 1995). The two genital lobes of “Diplatyidae” and“Pygidicranidae” are unidirected, whereas the two genital
lobes of Apachyidae, Allostethus indicum and Anisol­
abididae are bidirected. The Spongiphoridae, Cheliso­
chidae and Forficulidae possess only one genital lobe.
The karschellid-type penes is present exclusively in Kar­schiellidae. Due to the structure of the genitalia, this char­acter is not applicable to the Blattodea.
42. Total number of virgae. One (0) or two (1) or four(2) (Haas, 1995). The Karschiellidae, Spongiphoridae,
Chelisochidae and Forficulidae possess one virga. The“Pygidicranidae” except Pyragra fuscata, Apachyidae,“Labiduridae”, Allostethus indicum and Anisolabididae
possess two virgae. The “Diplatyidae” and Pyragra fus­
cata possess four virgae. This character is not applicable
to Blattodea due to the structure of the genitalia.
43. Basal vesicle. Absent (0) or present (1) (Haas,
1995). The basal vesicle is absent in Karschiellidae,“Diplatyidae”, “Pygidicranidae”, Anisolabididae andSpongiphoridae. It is present in all other Dermaptera
examined. This character is not applicable to Blattodea
due to the structure of the genitalia.
Character analysis
The analysis of the data set (Table 4) results in four
equally parsimonious trees with a tree length of 72, CI
0.819 and RI 0.935. Fig. 24A depicts the preferred tree.
Ambiguous characters are marked with an asterisk (*).
The Karschiellidae come out as monophyletic (char­acter 41). The “Diplatyidae” are not a subtaxon of the“Pygidicranidae” and appear to be paraphyletic. The
monophyly of a taxon “Pygidicranidae” (excluding Kar­schiellidae and “Diplatyidae”) is not supported. In con­trast, all examined species of Echinosoma and Tagalinaburri share an apomorphy (character 42) with the morederived taxa.
Allostethus indicum is usually placed in the “Labiduri­dae” (Brindle, 1965a; Steinmann, 1989) but appears to be
more closely related to the “Pygidicranidae” andApachyidae in our analyses. This is due to eight apomorphic character states in the “Labiduridae” in which
Allostethus indicum shows the plesiomorphic character
state (characters 5, 10, 11, 15, 16, 18, 19, 20). The
Apachyidae appear to be monophyletic (characters 5, 6, 7,
13, 33*, 36), however only one species was examined.
They are never a subtaxon of the “Labiduridae”.
The monophyly of “Labiduridae” (excluding Allo­stethus indicum and Apachyidae) is not supported bystrict consensus tree (not shown). However, in two offour trees they appear monophyletic (characters 8*, 12*
as ambigous apomorphies). If monophyletic, they arealways the sistergroup to the Eudermaptera (Spongipho­
ridae + Chelisochidae + Forficulidae).
The Eudermaptera are monophyletic, and there are 14
characters to support this view (characters 2, 3, 7, 9, 12*,

13, 15, 20, 23, 29, 30, 31, 41, 42). A sistergroup relation­
ship of the Chelisochidae and Spongiphoridae is sug­gested by characters 5, 8* and 10. The Spongiphoridae
are probably monophyletic (character 43 state 0) The
Chelisochidae are probably monophyletic (character 35
state 1). The Forficulidae are probably monophyletic(character 8*, 35 state 2).
PH YLOGENETIC ANALYSIS OF FOSSIL DERM APTERA

A close relationship between Dermaptera and “Protelytroptera”, or a sub-group of the latter, has often been
assumed (Tillyard, 1931; Hennig, 1969, 1981;
Boudreaux, 1979; Kukalová-Peck, 1991; Carpenter,
1992). However, no cladistic analysis has been conductedso far. We present a cladistic analysis for this relationshipincluding Protelytron permianum (Fig. 19), Acosmelytron
delicatum (Fig. 18) and Apachelytron transversum (Fig.
3). This selection was made because all other species of“Protelytroptera” are only known by their tegmina andshow no wing folding, wing venation, or body characters(Carpenter, 1992).
Apachelytron transversum (Fig. 3) shares with extant
Dermaptera many autapomorphies in the wing venation:
Veinal sector MP long and simple for most of its course;
arculus (mp-cua) present; the stem of Cu relatively long;
CuA branches end on anal vein instead of reaching poste­rior margin; CuP short, directed towards and ending onanal vein; AA1+2 completely lost (remnants present inother Blattoneoptera); and AP1+2 sending off a series (8to 10) of pectinate branches. These characters are notincluded below because they are not available either inthe other “Protelytroptera” or in fossil Dermaptera.
Characters and their states
In order to avoid ambiguities, the character numbers
continue below with 44. Apachelytron transversum shares
derived characters in the wing venation with the Dermap­
tera, Protelytron permianum and Acosmelytron delicatum
and, therefore, it is included in the analysis below.
44. Number of ocelli. Two (0) or none (1). All exam­
ined taxa apart from Diplatysjacobsoni have two ocelli.
45. Head. Opisthognathous (0) or prognathous (1).
Periplaneta americana has an opisthognathous head,
while all other examined taxa possess a prognathoushead. The character state is not known in Acosmelytron
delicatum and Apachelytron transversum.
46. Pronotum. Disc-like, large, Blattodea-type (0) or
disc-like, small, Dermaptera-type (1). The disc-like pro­notum is small in fossil and extant Dermaptera, while
being large in Periplaneta americana. The character state
is not known in Acosmelytron delicatum and Apachely­
tron transversum due to poor preservation.
47. Spines on femoral carina. Present (0) or absent(1). Spines are present in Diplatys jacobsoni, Archidermapteron martynovi, Asiodiplatys speciosus, Microdiplatys campodeiformis and Protodiplatys fortis. They areabsent in all other examined taxa.
48. Number of tarsomeres. Five (0) or three (1). There
are tarsi with three tarsomeres in Diplatys jacobsoni,
Semenoviola obliquotruncata, Semenovioloides capitatus

and Turanoderma sepultum. All other examined taxa
have tarsi with five tarsomeres.
49. Hindwing. Long, folded fan-like (0) or with one
transverse fold (1) or with two transverse folds (2), wingpackage. A wing package with two transverse folds ispresent in all fossil and extant Dermaptera. Amongst theexamined “Protelytroptera”, Protelytron permianum, hasa wing with one transverse fold. All other examined taxa
have a simple fan-wise folded wing.
50. Broadenings. Absent (0) or present (1). The broadenings are present in all fossil and extant Dermaptera andProtelytron permianum. They are absent in all otherexamined taxa.
51. Tergites and sternites overlapping in abdominal
segments. Not overlapping (0) or overlapping (1). These
plates are overlapping in all fossil and extant Dermaptera
and Acosmelytron delicatum. The characters state is notknown in Apachelytron transversum and Protelytron per­mianum due to poor preservation. There is no overlap in
the tergites and sternites of Periplaneta americana.
52. 8th and 9th abdominal tergite in females. Distinct
and separate from 10th tergite (0) or narrowed, but sepa­
rate from 10th tergite and not covered by 7th tergite (1)orfused to 10th tergite and covered by 7th (2). The tergites
are distinct and separate in Periplaneta americana,
Acosmelytron delicatum, Apachelytron transversum,
Archidermapteron martynovi, Asiodiplatys speciousus,
Dermapteron incertae, Microdiplatys campodeiformes,
Protodiplatys fortis and Turanovia incompleta. The ter­gites are narrowed in Semenoviola obliquotruncata,
Semenovioloides capitatus, Turanoderma sepultum andfused only in Diplatysjacobsoni.
53. Ovipositor. Straight, long (0) or short or absent (1).
Character state 1 is found in Diplatys jacobsoni, Derma­
pteron incertae, Semenoviola obliquotruncata, Semeno­
violoides capitatus, Turanoderma sepultum and Tura­
novia incompleta. Character state 0 is found in all otherexamined taxa, except in Acosmelytron delicatum, Prot­
elytron permianum, Archaeosoma serratum and Longiceratus mesozoica due to poor preservation or because
the specimen is a male.
54. Adult cerci. Annulated, short (0) or annulated, long(1) or annulated, but basal article is much larger thanothers (2) or smooth (3). Smooth cerci are present in
Diplatys jacobsoni and Archaeosoma serratum, Semeno­
viola obliquotruncata, Semenovioloides capitatus, Tura­noderma sepultum. They are annulated and long in Archi­
dermapteron martynovi, Asiodiplatys speciousus, Longiceratus mesozoica, Microdiplatys campodeiformis andProtodiplatys fortis. The basal article is much larger than
the following ones only in Dermapteron incertae andTuranovia incompleta. The adult cerci are annulated andshort in Periplaneta americana, Acosmelytron delicatum
and Apachelytron transversum. The character state ofProtelytron permianum is not known due to poor preser­
vation.
Character analysis
The analysis of the data set (Table 5) results in 588
equally short trees with tree length 15, CI 1.0 and RI 1.0.
479

The preferred tree is shown in Fig. 24B. The high numberof taxa together with the low number of characters results
in a large number of equally short trees. However, there is

Apachelytron transversum does not differ in charactersgiven in the matrix from the Blattodea Periplaneta ameri­cana. However, it is included in the ingroup on thethe relationships of fossil and extant Dermaptera to their stem line. Asterisk (*) - ambiguous characters.
480derived wing characters (characters 49, 50) with thehigher fossil Dermaptera.
The “Archidermaptera”, comprising the genera Semenoviola, Semenovioloides, Turanoderma, Protodiplatys,
Archidermapteron, Asiodiplatys, Dermapteron, Microdiplatys, Turanovia, Archaeosoma, and Longiceratus areparaphyletic and they are not the sistergroup of extant
Dermaptera, represented by Diplatysjacobsoni.
The monophyletic taxon within the “Archidermaptera”,
comprising the genera Protodiplatys, Archidermapteron,
Asiodiplatys and Microdiplatys, shares the possession ofspines on the femoral carina (characters 47). The genera
Dermapteron and Turanovia share the type of cerci (char­acter 54).
Extant Dermaptera constitute a monophyletic taxon,
sharing the reduction of ocelli and the fusion of the 8th
and 9th abdominal female tergite to the 10th tergite,
including a covering of the former by the 7th tergite(characters 44, 52).
DISCUSSION
As documented above and in the characters in Table 6,transformations in the ground patterns of the wing com­plex proceed step-wise from the Pterygota protowing, to
Neoptera, to Blattoneoptera, to Dermaptera and finally tothe dermapteran families. We are convinced that the phy­logenetic relationships of any pterygote order, and espe­
cially of Dermaptera (because of their highly unusual
wing pattern), can be seen only in the broad evolutionary
context of all other pterygote orders. As an example, toshow that the prominent anojugal lobe in Dermaptera istypically blattoid rather than endopterygote or orthopteroid, it is first necessary to document the diversification of
all three lobes from the shared Neoptera ground pattern.
This can be recognized only after distinguishing it fromits homologue in the ex-group, the palaeopterous groundpattern. The palaeopterous lobe must be interpreted byresearch on all palaeopterous orders. Finally, both neopterous and palaeopterous anojugal lobes must be fully
homologized, i.e., must be derived from a single, sharedpterygote protowing pattern (Table 6: Introduction).
Thus, preparing in a pterygote order a wing ground pat­tern for higher phylogenetic analysis, snowballs instantly
and inevitably into a need to know the ground patterns of
all other pterygote orders.
In a quest for presenting results based on sufficient
documentation, we had no other choice but to trace all
step-wise changes in wing character states, from the pro­towing to Dermaptera. Furthermore, dermapteran wings
are so unusually altered that to show clearly their under­
laying blattoid pattern, they had to be studied in conjunc­tion with a detailed functional analysis.
The most difficult hurdle to be overcome in using thewing complex in higher phylogenetics is dealing with thefact that the apomorphies are largely limited to reductions
and fusions. Veins and articular sclerites are getting con­stantly fused or reduced at all systematic levels. Thus, thesynapomorphies usually cannot be empirically distin­
guished if inspected as separate characters. However, inspite of this problem, the underlying ground pattern usu­
ally instantly identifies each wing and articulation as aparticular order or family. This dilemma suggests that to
avoid problems with homoplasies, it is necessary to keep
the integrated ground pattern characters together. It isessential to compare one wing ground pattern withanother wing ground patterns, and to avoid completely
comparing one separate (neopterous) character withanother separate (neopterous) character. The way to rec­ognize synapomorphies is to search for them within thecontext of their ground pattern. In other words, the step­
wise transformations in integrated character complexes,
which separate one ground pattern from another, also
show the phylogenetic relationships. We are convincedthat this method is repeatable and, therefore, an objective
method of character assessment in higher phylogenetics.
Our illustration of this method in Table 6 shows theamazing number of homoplasies that accompany and par­allel almost every wing synapomorphy. Under these cir­
cumstances, we consider selecting and assessing thehigher-level wing characters as separate entities (i.e., out­side the context of their particular ground pattern andwithout full homologization with the other ground pat­terns) as a haphazard and unreliable method. Note that
under the circumstances, subsequent processing simply
cannot improve the quality of the basal data set. Count­less morphologically unsupportable higher phylogenetic
schemes result.
As background to the wing data set in Table 6, we areoffering here a brief overview of the main phylogenetic
events, which lead to the Dermaptera. The events corre­
spond to the main nodi in the phylogenetic tree on Fig.
23.
Pterygota: Protowing and the split into Palaeoptera
and Neoptera
Pterygota are monophyletic and all wing characters(venation and articulation) evolved from a single pro­towing ground pattern. Morphological evidence indicatesthat ancestral pterygotes had three pairs of small,
movable, articulated protowings that were incapable ofpowered aerial flight, but almost certainly were flappingand generating some kind of forward movement. Only theprothoracic pair of protowings is recorded in fossils. They
were retained in a partly immobilized and slightly
reduced condition in the adults of Carboniferous Palaeodictyoptera and Geroptera (odonatoid stem line). Threepairs of small, homonomous, mobile wings were present
in the nymphs of the plesiomorphic palaeopterous
Diaphanopterodea. These nymphal wings could be simply
folded (warped) and pulled towards the body, in a waythat quite probably occurred in oldest protowings (Brod­sky, 1994; Kukalova-Peck, 1978, 1983, 1991, 1997; Willmann, 1997; Wootton & Kukalova-Peck, 2000) (Table 6).
The fundamental split of the Pterygota into Palaeoptera
and Neoptera (Martynov, 1923) probably took placewhen the three pairs of protowings were homonomous,their venation was radially arranged, there were no folds,
and the wing veins lacked fusions and braces. Some ofthese features are retained only in Palaeoptera, others
481

only in Neoptera. Fusions and braces are crucial for pow­
ered forward fligh. Without exception, they occur in allwings capable of powered flapping flight; insects withhighly reduced venation are very small and depend on air
currents. Therefore, the absence of shared fusions andbraces in Neoptera shows that the neopterous ancestor
could not fly in the air. In other words, flapping aerial
flight is not a neopterous (or pterygote) synapomorphy.
Lack of the shared flight adaptation in Neoptera is consis­
tent with the fact that Neoptera and Palaeoptera have fun­damentally different wing pair symmetry, costal margin,
anojugal lobes, veinal fusions, veinal braces, and arrange­
ment of the wing articulation (Fig. 1A, B). In Table 6A,
an astonishing 65 different wing characters are listed in
the ground pattern of Palaeoptera and Neoptera. Pterygote
protowing is shared by all Pterygota but the functional
adaptations, which converted the flapping appendage toan aerial flying appendage, evolved in several directions.
Like in aeroplanes, there is a multiplicity of forms andpatterns that can become flight-worthy.
Palaeoptera bear forewings and hindwings which arehomonomous in size, contain a radially arranged andregularly corrugated wing venation, are membranous, andwith a strong irregular inter-veinal network. The
remigium, anal lobe and jugal lobe are all at one plane.
The jugal area is always strongly reduced. There are noshared stabilized wing folds between wing areas. Twoveinal stems are always present, the medial and cubital.
The radial stem in modern Ephemeroptera and Odonatoptera has RA & RP fused side-by-side (in the radial stemof Neoptera, RA & RP are always superimposed). The
sclerites in the palaeopterous wing articulation are always
arranged in regular rows and columns and are positioned
close together (clustered) centrally near the wing base(Kukalova-Peck & Brauckmann, 1990). This arrangement
is predisposed for an easy fusion between fulcalaria (F)sclerites on the body side and basivenalia (B) sclerites on
the wing side. The resulting articular plate rests on thepleural wall and creates a lever rocking on the pleural ful­crum point. But, the wings become fixed in a permanently
outstretched position, are incapable of being flexed back­ward over the abdomen, and prevent escape from preda­
tors by hiding in narrow spaces. In exchange, the insect
can glide effortlessly with minimum energy expenditure,and may “outgrow” predators by gigantic increase in size.
Mayflies and damselflies can hold their wings lifted
upwards when at rest. Only a single, most primitive fossil
order, the Diaphanopterodea, retained the ancestral ability
to flex their wings backward over the abdomen. In therest of Palaeoptera the fusion of centrally positioned
sclerites into an articular plate happened twice differently:(i) in Palaeodictyoptera + Megasecoptera, and again (ii)in Ephemeroptera + Odonatoptera (Kukalova-Peck, 1985,
1997; in progress). All Palaeoptera have fragile membra­
nous forewings and hindwings. The data above indicatethat the early survival tactics in Palaeoptera was not
hiding from predators, but rather flying away. The
resulting speedy and highly sophisticated perfection of482

flight is richly documented in the Palaeozoic fossil record(Wootton & Kukalova-Peck, 2000, and for references).
Neoptera bear in their ground pattern a pair of fore andhindwing, which are heteronomous in appearance, sizeand function. The forewings are tegminous, narrower andthicker than the hindwings. They participate in flight andprotect the hindwings, which are the main flying pair
(Brodsky, 1994). In forewings, the remigium and anal
lobe are at one plane, but the jugal lobe is at anotherplane. The jugal lobe is membranized, strongly reduced
and is folded under the anal lobe. Anal lobe bears a long,strong anal bar composed of BAA + BAP, adapted tolocking the wing at rest. Anal branches are not radiallyarranged, but are distributed irregularly. Principal branch
AA1+2 is distanced from AA3+4, and AA3+4 is always
fused basally with AP.
The hindwings are much broader than the forewings,
are membranous, with a sparse inter-veinal network, andbear plesiotypic radially arranged veins as in Palaeoptera.
The anal lobe and jugal lobe are at the same plane. Bothwing pairs have four stabilized folds: Medial, claval, analand jugal. They share an anal brace between BAA and
BCu engaged in wing flexing. The veinal corrugation(convex: +, and concave: -) is permanent in the veins
ScA+, ScP- and RA+, but the remaining veins have avariable corrugation (convexity and concavity is often
diminished or even reversed) (Fig. 1).
Neopterous wing articulation is typical and unique,and it presents Neoptera as a convincingly monophyletic
group. The asymetrical composite sclerites 1Ax and 2Ax,
and the rotating composite sclerite 3Ax (Fig. 1) aredesigned for flexing of the wings backward and locking
them securely in a flexed position. The characters mostinformative in higher phylogenetics occur in 3Ax (goblet+ saucer) and its arms, 2Ax body and arm, subcostal
basivenale BSc, PWP, and in the veinal basivenalia. The
same features are highly autapomorphic in Dermaptera.
As an important fact, Neoptera share a highly efficient
flexing mechanism, but do not share any veinal stems andbraces necessary for aerial powered flight. This is consis­
tent with the neopterous ancestor whose main survival
strategy was probably flexing the wings backward andhiding from predators in narrow spaces, while flight
seems to have been initially of a somewhat lesser impor­tance. Indeed, the high manoeuverability of wings andhigh speed in flight comparable to those in Carbonifer­
ous Odonata and Megasecoptera (Wootton & KukalovaPeck, 2000) came to the Neoptera only in the Mesozoic,
in Diptera and some Hymenoptera. The most informative
higher level characters in neopterous venation occur inveinal stems and braces near the wing base (Table 6B-H).
In the Neoptera ground pattern, the forewing is protec­
tive as well as flying. But, the main flying wing is thehindwing with an enlarged anojugal lobe (Brodsky,
1994). This concept, which is originally based on func­
tional analysis of the flight motor, is confirmed here bywing articulation. The neopterous hindwing ground pat­tern includes a triple subdivision of the anal basivenale, toaccommodate the many veins needed to support theenlarged anojugal lobe. Neopterous anal basivenale BA(=the sclerotized blood sinus of anal branches) is subdi­
vided into three mutually articulated sections (BAA1+2,
BAA3+4, BAP: Figs 1, 11-17), which service numerous
anal branches supporting the enlarged anal lobe. This
triple subdivision is absent in the Palaeoptera, well pro­nounced in the lower Neoptera and is present or indicatedin the higher Neoptera even when the anal lobe was
reduced (see Figs 11-17; Table 6). The enlarged lobe
occurs in plesiomorphic Pleconeoptera, Orthoneoptera
and Blattoneoptera, in some basal representatives of
Hemineoptera (Fulgoromorpha, some Auchenorrhyncha)
and of Endoneoptera (Hymenoptera: Xyelidae, Coleoptera, Megaloptera, Ithonidae (Neuroptera), Philopotamidae (Trichoptera). Later changes in the anojugal lobe
include further secondary enlargement (e.g., in Dermaptera), abrupt reduction (in petiolate wings), and gradualreduction (in Hemineoptera, Endoneoptera). The changes
occur independently, randomly and in parallel, simultane­
ously at the superorder, order, suborder and family level,
and are responsible for many homoplasies. In Pleconeo­
ptera, the basal anojugal lobe was initially large and thendiminished - gradually in derived Plecoptera and abruptly
in Embioptera. In Orthoneoptera, the prevalent tendency
was to retain and further enlarge the anojugal lobe and to
add secondary anal branches. In Blattoneoptera(including Dermaptera) the AA area is combined with theremigium, and removed from the partial anal lobe. But,
the remaining AP + J areas became enlarged, and AP
developed secondary anal branches to fill up the newspace. In Hemineoptera and Endoneoptera the AA area isalso removed, but instead of compensating for the loss,
the entire hindwing started to diminish and the anojugal
lobe lost even more anal branches, and the forewings
became the main flying pair.
An important fact is that Neoptera share all articularsclerites, but do not share any veinal stems or braces near
the wing base, which are crucial for aerial powered flight.
This indicates that the wing flexing evolved before theseadaptations were established in the neopterous groundpattern. There are two different patterns of fusions andbraces important in aerial flight, but neither can be ances­tral to the other: In Pleconeoptera + Orthoneoptera and in
Blattoneoptera + (Hemineoptera + Endoneoptera). The
differences include: (1) fusions and braces of the medialvein; (2) composition of the anojugal lobe; (3) composi­tion of the flight units; and (4) the use of the flightimportant folds. Brodsky (1994) proposed that bothgroups also independently evolved two types of flightmotor. These two patterns probably evolved from acommon ancestor (as summarized here in Figs 22, 23)
and represent sistergroups.
Neoptera: Basal split into Pleconeoptera +Orthoneoptera and Blattoneoptera + Hemineoptera +Endoneoptera
The most profound morphological difference between
the two basal groups of Neoptera is in the composition of
the anojugal lobe. This lobe is particularly important inflight. Pleconeoptera (Fig. 14) and Orthoneoptera (Fig.

15) have a full anojugal lobe, which starts at the clavalfold (the main flexion line in flight) and contains two anal
and two jugal sectors (AA, AP, JA, JP), a symplesiomorphy (Figs 21, 22). The remigium represents a flight
unit of its own (Table 6B). Contrary to this, Blattoneo­
ptera, Hemineoptera and Endoneoptera share a partial
anojugal lobe, which starts at the anal fold and contains
only one anal and twojugal sectors (AP, JA, JP) (Figs 11,
12, 13, 17, 21, 22). The anterior anal sector (AA) is pro­
gressively reduced and combined with the remigium into
a composite flight unit, an autapomorphy (Figs 20, 21B,
17) (Table 6D). The claval fold, which lost its function as
the main flexion line, is gradually reduced, firstshallower, then shorter or lost. The anal fold became themain flexing fold (an autapomorphy). It separates the par­tial anojugal lobe, which is supported only by thebranches ofAP and J. Further splitting occurs as follows.
In Blattoneoptera the original main flying role of thehindwings is sustained in spite of the partial anojugal
lobe. This is possible by adding some autapomorphies.
Diminished veinal support is compensated for by the sec­
ondary terminal branches, which are added to AP1+2.
This phenomenon is especially well pronounced in
Dermaptera and in related Dictyoptera (see Table 6E).
In contrast, partial anojugal lobe in Hemineoptera +Endoneoptera gradually becomes smaller and anal
branches shorter and more impoverished. Eventually,
most endoneopterous hind wings end up being smaller
than the forewings, their role in flight is reversed, andforewings become the main flying pair. Sometimes thefore- and hindwing hook together and are used as a single
flying wing. Even this complex flight adaptation is a
homoplasy, occurring in parallel in Hemiptera, Hymeno­
ptera, and Lepidoptera.
The morphological split of the anojugal lobe into two
types accompanies an equally important split of thebraces in the medial vein into two types. In Orthoneo­
ptera + Pleconeoptera the medial sectors MA and MP
fuse basally into a long stem of M (a synapomorphy) andmedial branches belong to both MA and MP. In contrast,
in Blattoneoptera + Hemineoptera + Endoneoptera,
MA is captured immediately at the basivenale by RP (orR), and is released again from RP apically after a long
fusion. Thus, medial branches belong only to MP (Figs11, 12, 17, 21B, 22B) (Kukalova-Peck, 1991, 1997;
Kukalova-Peck & Brauckmann, 1992). In Dermaptera,
the media is highly reduced and not informative.
Note that the fusion between MA and RP (or R) near
medial basivenale (BM) as well as the separate base ofMP, are almost always macroscopically obscured. This
obscurity instigated decades of endless debates, errone­ous homologizations, and unfounded criticism. Neverthe­less, the states of MA and MP described above are very
easily verified in large, freshly killed Blattodea or
Megaloptera, by sectioning the veinal bases under water(K.D. Klass & JKP, personal observation).
The wings are flexed backward in the resting position
in Pleconeoptera + Orthoneoptera and are wrappedaround the body dorsally and laterally (especially near the483

wing base). In contrast, in Blattoneoptera + Hemineoptera+ Endoneoptera the wings are flexed in a roof-like posi­tion (low or high), or are dorsally flat. The position offlexed wings in Dermaptera and Dictyoptera is identical:
Dorsally flat.
The internal folding of the anojugal lobe in Pleconeoptera + Orthoneoptera is regularly fan-like, with equal
distances between the folds. In Blattoneoptera + Hemi­
neoptera + Endoneoptera the folding is irregularly fan­like: Some folds may be added, and they may cross someanal veins. Dermaptera share with Dictyoptera AP1+2
bearing added secondary branches; Dermaptera also con­tain several added apomorphic folds (see Table 6E).
The oldest wings of Blattoneoptera are known from theMiddle Upper Carboniferous (Westphalian). Some belong
to the blattoid stem line (probably related to Isoptera, see
Table 6F, G), and others to the living order Blattodea.
Protelytroptera, the polyphyletic stem line including thedirect ancestors of Dermaptera, is known from the Early
Permian, but is almost certainly older than the dictyopteroid line (some dermapteroid and Dermaptera characters
are more plesiomorphic than in Dictyoptera, see Table
6D-G).
Contrary to the opinion of some, the wing complex, if
properly used, is actually convincingly and exceptionallyrichly informative at higher systematic levels. And, thefluctuation of size, amount of sclerotization, the numberof small branches, on top of unpredictable variability andreversals, often makes wing characters uncertain and notvery useful at the species and genus level.
With a fully homologized wing complex, the Pleconeoptera and Orthoneoptera share several synapomorphies with each other, but none with Blattoneoptera,
Hemineoptera, or Endoneoptera (see Table 6B). Conse­
quently, the taxon Polyneoptera of Martynov (1940) (plecopteroids + orthopteroids + blattoids) and its alleged
stem lines (cumulatively called “Protorthoptera”) are bothpolyphyletic and should be abandoned (Kukalova-Peck,1991; Kukalova-Peck & Brauckmann, 1992; KukalovaPeck & Peck, 1993).
Accurately homologized wing veins, wing lobes andwing articulation convincingly put Dermaptera into
Blattoneoptera (Table 6D, E). Dermaptera share withDictyoptera the following hindwing character states. (1)A partial anojugal lobe, (2) a shortened, shallow clavalfold, (3) a well pronounced, long and deeply incised anal
fold, (4) a significant narrowing of the AA field, (5) atypically blattoid (very strongly) extended AP field, and(6) the presence of secondary, serial, terminal branches
on AP1+2. For additional character states see Table
6B-H
The relationship of Dermaptera within
Blattoneoptera
The relationship of Dermaptera to other neopterous
orders was more recently discussed by Stys & Bilinski(1990), and by Kristensen (1995). Those authors con­cluded that, because of a similar ovarian structure,
Dermaptera are a prospective sistergroup of Hemineo­

484

ptera + Endopterygota. This implies that Dermaptera areeither not a member of the Blattoneoptera, or that Blat­
toneoptera are paraphyletic.
We do not share either view. The autapomorphies of
the Blattoneoptera, occurring in the Grylloblattodea(Notoptera), Dermaptera, and in Dictyoptera, are as fol­lows. The pronotum is disc-like and flat. The flexed
wings lie horizontally on the abdomen and they arearticulated far forward at the anteriormost corners of themetanotum. The postnota are reduced. The flight motor
system is highly modified in the same, complex way(Boudreaux, 1979, his Fig. 71, p. 207; CSIRO, 1991, p.
320; Brodsky, 1994). Another alleged autapomorphy of
the Blattoneoptera, the strongly slanted pleural suture(Boudreaux, 1979), appears to be a later development
because it is not visible in Acosmelytron delicatum (Fig.
18).
All members of Blattoneoptera (except the predaceous
Mantodea) live on the ground, in litter, crevices or otherhidden places. It seems that running away from predatorsand hiding in narrow places, rather than flying away, wasthe main survival strategy of ancestral Blattoneoptera.
Grylloblattodea, Dermaptera and Dictyoptera show very
distinctive adaptations for hiding, such as the dorsoventrally flattened body (Blattodea), accompanied by later­
ally overlapping abdominal tergites and sternites (Der­maptera), the strongly slanted pleura (Grylloblattodea,
Dermaptera, Dictyoptera), and the large coxae with strong
muscles for running (Grylloblattodea and Dictyoptera)(Boudreaux, 1974; this paper).
In the wing structure, Dermaptera and Dictyoptera
share important blattoneopteran characters, which areclearly recognisable, even if sometimes differently modi­
fied (Table 6E). The blunt ridge replacing ScA in Dictyo­
ptera is also present in Dermaptera, but is changed into an
overhang. The radial basivenale (BR) is also long andswollen, but variously shaped. The medial basivenale isalso right-triangular, enlarged and separated from BR bya conspicuous furrow, but its proximal margin is membranized. The 2Ax body (AXM) is also wrinkled andbears an arched proximal margin, but it is narrower andmore twisted. The 3Ax anal arm (FA) is also very promi­
nent, but differently shaped and extended by incorpo­
rating FJ. The posterior wing process (PWP) is also
desclerotized, but much narrower. The wing attachment isalso broader than the wing articulation. These wing char­acter states are absent in the ground patterns of
Orthoneoptera, Pleconeoptera, Hemineoptera, and Endo­
neoptera (Figs 14-17, Table 6B-H).
The presence of several “dictyopteroid” character statesin the fossil stem line of Dermaptera, such as tarsi withfive tarsomeres and annulated cerci, clearly shows thatthe tarsi with three tarsomeres and forceps-like cerci inextant Dermaptera are not part of the ancestral dermapteran ground pattern and evolved later.
The relationships of extinct “Archidermaptera”
The results obtained in this study partly confirm andpartly reject the previous hypotheses on the relationships
of fossil Dermaptera. The “Archidermaptera” are clearly aparaphyletic assemblage of those fossil species thatcannot be referred to extant families, as was also con­cluded by Willmann (1990). Their status as a separate“suborder” (Carpenter, 1992; Sakai, 1996; and others) is
not supported by any autapomorphy and should be aban­
doned.
Vishnyakova (1980) used the “phenetic-similarity”
approach to “archidermapteran” relationships. In spite ofthis different method, the results are to some extent
similar to ours. Vishnyakova proposed a close relation­
ship between Dermapteron and Turanovia and grouped
them into one subfamily. The sistergroup relationshipbetween the genus Dermapteron and Turanovia is also
supported here (character 54), but analysing the genus
level is beyond the scope of this contribution. According
to Vishnyakova (1980), the genera Archidermapteron,
Asiodiplatys, Microdiplatys and Protodiplatys belong to
the Protodiplatinae. They indeed appear also to be monophyletic in our analysis.
The results presented here contradict Vishnyakova(1980) in that the genera Semenoviola, Semenovioloides
and Turanoderma are members of the extant “Pygidicranidae”. Extant Dermaptera share as derived characters the
reduction of ocelli and the special structure of the female
abdomen. In contrast, the fossil genera show the plesiomorphic states of these characters. It is quite possible that
Archaeosoma martynovi and the genera mentioned above
represent the sistergroup of extant Dermaptera.
Extant Dermaptera, in spite of being quite markedly
different from the rest of extant insect taxa, share only
two ambiguous apomorphies: The reduction of ocelli, andthe covered 8th and 9th abdominal tergites in females(characters 44*, 52*). All other “typical” dermapterancharacters, such as the loss of cercal annulation, overlap­
ping tergites and sternites, and highly specialized wingfolding, are already present in Palaeozoic and Mesozoic
fossils. Thus, Jurassic fossils are easily recognized as
Dermaptera (Vishnyakova, 1980) even if the wing vena­tion and articulation is not visible. In extant Dermaptera,
there is remarkably little morphological and ecologicalvariation. A similar close adherence to an ancient groundpattern can be observed in Blattodea.
The evolution of characters is not presented in detail,because of the great number of equally parsimonious
trees. However, some aspects in evolution of the charac­
teristic wing folding are discussed below.
The relationships of extant Dermaptera “families”
Extant Dermaptera clearly constitute a monophyletictaxon, as suggested by a highly autapomorphic wingarticulation, venation, and folding pattern. However,these characters are not applicable to the large number of
wingless earwig species, and to the Hemimerina (African,
epizootic on giant rats; Rehn & Rehn, 1935) and Arixeniina (Malaysian, epizootic on bats; Jordan, 1909). Their
inclusion into the Dermaptera is justified by the posses­
sion of smooth, not annulated adult cerci (character 38),
holocentric chromosomes (White, 1971, 1972) and by themost striking dermapteran body character, the lateraloverlapping of abdominal tergites and sternites (charac­ters 40, 51). Their tarsi have only three tarsomeres (char­acters 34, 48). None of the living, closely related groups,
such as Blattodea share these derived character states.
Hemimerina and Arixeniina are most often considered
as “suborders” of the Dermaptera, because they are con­spicuously different in many features. However, both taxa
have acquired a high number of autapomorphic characters
due to their peculiar life styles, as is common in epizootic
or parasitic taxa. These autapomorphies obscure the char­acters showing the relationships with existing derma­
pteran taxa. We assume that Arixeniina and Hemimerina
are indeed the sistergroup or a subtaxon of a dermapteran“family”, as was proposed by Popham (1985) for Arix­
eniina (called Arixeniidae) and recently by Klass (2001).
The latter author found detailed similarities in theabdomen of Hemimerus and other forficuline taxa.
Because the new characters presented here are con­cerned with the wing venation and articulation, the sys­
tematic position of the Arixeniina and Hemimerina is notdiscussed in this account.
The above results partly confirm and partly contradict
the findings of Haas (1995). The exclusion of the Karschiellidae and “Diplatyidae” from the “Pygidicranidae”
is confirmed, as is the paraphyly of the “Diplatyidae”.
The monophyletic state of the “Pygidicranidae” has been
doubted by Haas (1995) on grounds of the four virgae in
Esphalmeninae and Pyragrinae. The results here support
this view because the reduced number of virgae (to two)is shared by all species of Echinosoma, Tagalina burri,and Allostethus indicum, by Apachyidae, Anisolabididae,“Labiduridae”, and by Eudermaptera (character 42). In
contrast, the last common ancestor of extant Dermaptera
probably had two unidirected genital lobes with four vir­gae. Later, the two genital lobes became bidirected (char­acter 41) and finally one genital lobe was reduced. The
latter apomorphic condition is found in the Eudermaptera(Forficulidae + Spongiphoridae + Chelisochidae).
The present results contradict the view of Popham(1985) on the evolution of male genitalia. Popham con­sidered the “Pygidicranidae” to be monophyletic becauseof the structure of the male genitalia, interpreted here as
plesiomorphic. Popham further assumed that the reduc­tion of one genital lobe occurred twice, in Spongipho­
ridae and again in Chelisochidae + Forficulidae. We
assume that it occurred only once.
In the present contribution, Apachyidae and Aniso­
labididae appear to be monophyletic families. This result
is not definitive, because only one winged species of eachtaxon was available to us for study. Most species of theAnisolabididae are wingless and their classification islargely based on body characters. Their sistergroup rela­tionship with the Eudermaptera (Haas, 1995) is not sup­ported here, because the assumed shared derived charac­ters (characters 29-31; or, characters 5-7 of Haas, 1995),
were found here to be erroneously coded.
The monophyly of the “Labiduridae” (excluding
Allostethus indicum) is not unambiguously supported
because they seem to constitute an assemblage of superfi­
cially similar Dermaptera. This notion is further strength­485

ened by the taxonomic history of the “Labiduridae”.
According to Burr (1911), “Labiduridae” contain all Protodermaptera except the “Pygidicranidae”, namely:
Esphalmeninae, Psalinae, Labidurinae, Parisolabinae,
Brachylabinae, Platylabinae and Allosthetinae. Later on,
more and more groups were removed from this taxon. For
example, the Esphalmeninae are now recognized as“Pygidicranidae” (Steinmann, 1986). Some authors, like
Brindle (1965b), included the Apachyidae into the“Labiduridae”. In contrast to “Labiduridae”, Apachyidae
and Forficulidae are taxonomically almost unchanged
since their study by Burr (1911), suggesting a much more
distinctive character set than in “Labiduridae”.
For the first time, the sistergroup relationship of the“Labiduridae” to the Eudermaptera is suggested (charac­ters 5, 6*, 8*, 11, * ambiguous character). Popham (1985)
took the view that “Labiduridae” are the sistergroup of
Forficulidae + Chelisochidae, based on an elongatedsecond tarsomere and the presence of the basal vesicle(characters 35, 43). As pointed out earlier (Haas, 1995),
the elongation of the second tarsomere is very variable inthe “Labiduridae”. So it was not possible, in contrast to
Popham (1985), to establish a clear distinction between
the “Labiduridae” + Forficulidae + Spongiphoridae andthe other dermapteran “families”. According to the
hypothesis presented here, the presence of a basal vesicle(character 43(1)) is an apomorphy shared by Allostethus
indicum, Apachyidae, Anisolabididae, “Labiduridae” andEudermaptera, with two independent secondary reduc­tions, in the Anisolabididae and Spongiphoridae. How­ever, the sistergroup relationship of “Labiduridae” andEudermaptera might explain the similarities noticed by
Popham (1985).
In contrast with Popham (1985), the Eudermaptera areconsidered here to be monophyletic. There are 14 charac­ters from different body regions supporting the monophyly. In fact, Eudermaptera (Forficulidae +Spongiphoridae + Chelisochidae) are better supported
than any other dermapteran taxon discussed in thisaccount.
In the present contribution, the sistergroup relationshipbetween the Chelisochidae and Spongiphoridae is sug­gested (characters 5, 8*, 10). Previously, Zacher (1912)
grouped Chelisochidae and Spongiphoridae under the“Labiales”, but this proposition was not followed by later
authors. Haas (1995) considered Forficulidae and Cheli­sochidae to be sistergroups, but this relationship is notsupported by new evidence.
The above results show that most Dermaptera“families” are either not monophyletic, or only one autapomorphy is known. Future research should focus on thisproblem and on the autapomorphic characters states for
all genera and families, which need to be identified. Thistask is especially urgent in the species-rich families Ani­
solabididae and Spongiphoridae. An improved under­
standing of their phylogeny will considerably improveinsight into the evolution ofDermaptera.

486

Are the Zoraptera member of the Blattoneoptera?
The position of Zoraptera remains uncertain. The
Zoraptera are hemimetabolous, which rules out a close
relationship with the Endoneoptera. They have a general­ized head, chewing mouthparts, and lack the enlarged
cibarial sucking pump, domed clypeus or any otherhemineopteran mouthpart adaptations. This rules out aclose relationship with the Hemineoptera. So, the Zora­
ptera remained plesiomorphic regarding to development
and mouth parts. However, they bear a reduced postnotum of the blattoid type, large cylindrical blattoid
coxae, and the highly derived flight motor as in Dictyoptera (consisting of wing attachments shifted to theutmost-anterior corners of thoracic terga); reduced phragmata; and highly reduced indirect flight musculature(Boudreaux, 1979, p. 215; Rasnitsyn, 1998). The male
genitalia appear to be blattoid (Smithers, 1991). The wing
venation is compatible with blattoids but the decisivewingbase characters are destroyed by petiolation(Kukalova-Peck & Peck, 1993). A membranized posterior
wing process (PWP), if present, should provide a con­vincingly blattoid character, but it has not yet been avail­able for observation.
The “non-blattoid” characters of Zoraptera include
tarsi with only two segments, cerci that lost their annula­tion, and a pronotum that is not disc-like. But these char­acters may easily be apomorphies due to the small bodysize and a special life cycle. The thoracic pleural sulci are
not slanted, but neither were those in the dermapteranancestor Acosmelytron delicatum (Fig. 18).
Rasnitsyn (1998) rejected the association of Zorapterawith Blattoneoptera: Dictyoptera for two reasons. (1)Zoraptera bear on the pterothorax an invaginated sternum
forming an internal ridge, which also occurs in
Hemineoptera and Endoneoptera but not in Blattoneo­
ptera. But this inner ridge is not a convincing characterbecause it occurs convergently in groups with very broad
coxae and narrow sternum, such as in unrelated
damselfly-like extinct Megasecoptera (Palaeoptera) (per­sonal observation by JKP). Recently, Rasnitsyn & Novokshonov (1997) removed Hemineoptera from Neoptera
and combined them with extinct Palaeodictyopteroidea
under Palaeoptera. This assumption breaks the convinc­ingly monophyletic Neoptera ground pattern (see massive
evidence supporting Neoptera monophyly in Table 6A),
and also contradicts Rasnitsyn’s (1998) interpretation of
the internal sternal ridge as a possible synapomorphy for
Zoraptera, Hemineoptera and Endoneoptera. (2) The
identical wing motor system in Dictyoptera and Zoraptera
is rejected as a possible synapomorphy because Rasnitsyn
found reduced indirect flight musculature both in Zora­
ptera and in Xyelidae (Hymenoptera). But the muscle
reduction and the Dictyoptera motor system are two dif­ferent character sets. The muscle reduction in Zoraptera ispart of a coevolved, integrated flight motor system, which
also occurs in Dictyoptera (Boudreaux, 1979; see descrip­tion above). In contrast, the muscle reduction in Xyelidae
is limited to this family. Endoneoptera including
Hymenoptera share the plesiomorphic neopterous flightmotor system (Brodsky, 1994). Therefore, the reductionof indirect flight muscules in Xyelidae is not part of theidentical and complex flight motor system as in Zoraptera
and in Dictyoptera, but an unrelated homoplasy.
Rasnitsyn (1998) proposed that Zoraptera belong to theextinct Palaeozoic “superorder” Caloneurodea, believedby him to be the sistergroup of Endoneoptera. But,
Caloneurodea bear a typical, hemipteroid, domed clypeus
above the cibarial sucking pump and typical stylet-likemouthparts (Paleuthygramma tenuis: Kukalova-Peck,
1991: Fig. 6.22; note that the last character is mentioned
also by Rasnitsyn, 1998). Head characters and venational
systems identify the extinct Caloneurodea as ancestral
hemipteroids (Kukalova-Peck & Brauckmann, 1992). In
contrast, head morphology shows that Zoraptera are nei­
ther hemipteroids, nor a member of Caloneurodea. Refer­ring the controversial modern insect order to an extinct,
poorly known taxon is extremely risky (Hennig, 1969,1981).
The above review of Zoraptera characters indicates thatthe relationship with Blattoneoptera: Dictyoptera is, under
the circumstances, the most parsimonious choice. But theevidence seems insufficient for a definitive decision. For
that reason, we choose not to include Zoraptera in the cladistic analysis.
FUNCTION AND EVOLUTION OF THE DERM APTERANHINDW ING

The hindwings of Dermaptera are highly derived inseveral aspects of their venation pattern. They are theonly wings known to us, in which veins do not radiatefrom the articulation, but from a region almost in thecentre of the wings (Fig. 2). Each radiating and inter­calary anojugal branch contains a broadening where it isdissected by the ring fold. The broadenings, which look
like “weakened” joints, actually provide strong veinal
support for the fan. In comparison with Coleoptera, which
also bear transversely and longitudinally folded
hindwings (Haas, 1999; Haas et al., 2000), the dermapteran hindwings have a very broad attachment to themetanotum, which is much broader than the area occu­
pied by the articulation. Also, the hindwing base is excep­
tional in containing special bending zones, and a large
central membranized area. All these morphological pecu­
liarities accompany and make possible the special folding
pattern and folding mechanism, described below.
Dermaptera wing folding mechanism
Dermaptera hindwings are unique within the Pterygota
in being able to fold an enormous anojugal lobe, which isalmost 10 times larger than the squama (Haas, 1994).
This is achieved by folding the wing four times, as shownin Fig. 2. (1) By a fan-wise folding of the anojugal lobe
along the radiating folds. (2) By a transverse fold between
the squama and ulnar area on one side, and the inner andouter apical area, and the bases of the radiating branches
on the other side. (3) By a bending of the radiating andintercalary branches along the ring fold. (4) By the longi­
tudinal folding along the border of the squama to theulnar area. The dermapteran folded wing forms a wingpackage with the squama as the dorsal surface, and theulnar area as the ventral surface. The anojugal lobe isdensely folded between the squama and ulnar area. The
four steps of folding actually take place almost simultane­
ously and they are dependent one on another, so that they
are not clearly separable. The reader is encouraged tophotocopy and enlarge Fig. 2 and to fold it into a paper
model of an earwig hindwing. When properly manipu­
lated, this model will fold and unfold, and show thefolding mechanism.
This complex folding is achieved entirely by intrinsic
elasticity in the wing, because the wing continues folding
automatically in freshly killed specimens. The forces aremainly generated by the broadenings in radiating andintercalary branches (Kleinow, 1966; Haas et al., 2000).
In contrast to their morphological appearance, the broad­enings are not weakened portions in veins, but are active
force-generating centres. They are the key element forfolding the large fan. By bending the veins along the ringfold, the whole fan starts closing, as in the model. Finally,
the ulnar area is tucked under the squama by the elasticity
in the longitudinal fold (Kleinow, 1966; Haas et al.,2000).
The broad attachment of the wing to the thorax is ofmajor functional importance. As long as the attachment isundamaged, the wing package pulls itself automaticallytowards the thorax, as a result of forces generated along
the longitudinal fold. If the wing attachment is partly
destroyed, so that the attachment is limited to the articula­tion, the wing does not return to its resting position (Haas,
1994; Haas et al., 2000). The elasticity in the longitudinalfold pulls the anojugal fan, which is attached to themetanotum, into the wing package; this returns the wing
to its final resting position (Kleinow, 1966; Haas et al.,2000).
Since intrinsic elasticity folds the wing, a different
mechanism must be employed to unfold it (elasticity
cannot be effective in opposite directions). To unfold thewing, the wing package is erected over the thorax (Klei­
now, 1966; Haas et al., 2000). This movement will auto­
matically open the wing package: The anojugal fan ispulled out by its attachment to the thorax, the wingpackage is opened, and the squama and ulnar area
assumes an angle of about 90°. Next, the abdomen bends
over, and the cerci interact with the wing package andunfold (“unzip”) the wings. Both wings are simultane­
ously erected, but they are unfolded one after the other(Kleinow, 1966; Haas et al., 2000).
Because the wing automatically folds by its intrinsic
elasticity, counter-acting structures and mechanisms areneeded to keep the wing unfolded. The stiffening isachieved by two mechanisms: The central wing hinge(Flugelmittelgelenk), and the concave longitudinal fold(clf) (konkave Langsfalte, both terms of Kleinow, 1966).
The central wing hinge consists of four plates: Squama,
ulnar area, outer apical area, and inner apical area. These
plates are arranged so that the angles around their centre
do not add up to 360°. Therefore, this bi-stable mecha-

487

Fig. 26. Model for the wing stiffening mechanism composedof the claval and anal fold, squama and ulnar area, and CuP.
The reader is encouraged to photocopy and enlarge the figureand fold it according to instructions. For further explanations
see text.

Fig. 25. Movements of the hindwing articulation sclerites
during folding. Arrows indicate the movement of single scler­ites, while thick lines indicate the turning or pivoting axes.
Scheme based on observation of Labidura riparia.

Fig. 28. Photocopy and enlarge this figure twice and fold it
according to the following instructions. Fold one model so thatthe solid straight lines are convex and the dashed lines are con­cave. Now bend the fan beyond the circle to the left, so that theconvex/concave relation of the circle line results. Fold thesecond model so that the dashed lines are convex and thestraight solid lines are concave. In this second case, the partbeyond the folding line has to be bent to the right in order toproduce the convex/concave relation of the circle line. Thesemodels demonstrate that the bending direction of the partbeyond the circle line can be changed without changing thebending directions of individual veins.
488

nism is used to stiffen the wing (Haas & Wootton, 1996;
Haas et al., 2000).
The second essential stiffening mechanism is the con­cave longitudinal fold (clf). The wing is kept unfolded bycounteracting the elastic forces along the longitudinal fold(lf) (Kleinow, 1966). The clf is snapped through during
the wing unfolding by the cerci, and probably released by
the action of the wing articulation; we assume it is amovement of 3Ax. A paper model demonstrates quite
well the function of the clf. The reader is encouraged tofold a sheet of paper longitudinally, so that a convex
crease (i.e., the lf) appears, and afterwards obliquely, sothat a concave crease (i.e., the clf) intersects the firstcrease at an angle. As long as the oblique, concave creaseis not snapped through, the two halves, separated by theconvex crease, are freely movable. The sheet becomeslocked if the concave crease is snapped through.
Kleinow (1966) believed that the concave longitudinalfold (clf) is only one fold. Our research shows that it actu-

Fig. 27. Model of the dermapteran wing articulation, to
explain the straightening of the bending zones in the anterior
wing margin, and the probable release mechanism of the stiff­
ened wing. The reader is encouraged to photocopy and enlarge
the figure and fold it according to instructions. For further
explanations see text.

ally consists of two, and sometimes three, folds: A shortclaval fold between CuP and AA3, a somewhat longer
anal fold just in front of AP, and a broad, flat CuP rem­nant, sometimes expressed as a concave fold. The clavalfold is concave when active, and it begins close to the tipof BA. It never crosses, though it often comes close to,the longitudinal fold (lf) between the squama and ulnar
area. The anal fold is convex and it runs close to AP. The
part of the anal fold involved in the stiffening of the wing
begins without a definitive origin near the base of AP. It
extends distally, crosses AP, andjoins the transverse fold.
Although the morphological interpretation of the folds
is different from that described by Kleinow (1966), hisfunctional considerations are correct. It is the crossing ofa convex fold by a concave fold, which keeps thehindwing unfolded along the border of the squama andulnar area. However, this is achieved by a combination ofseveral folds, and not by a single fold. The (concave)claval fold ends just in front of the (convex) longitudinalfold, but yields a second fold as a continuation (Fig. 26).
If the claval fold would cross the folding line between thesquama and ulnar area, veins CuP and AA would have to
be flexible at this line, but flexibility is probably not rec­oncilable with their stiffening function.
In some species of Dermaptera, especially in the moresclerotized ones, a third concave fold is found. It some­times begins as a continuation of the claval fold, andsometimes just anterior to it, probably as a small remnant
of CuP. It ends by fading in the ulnar area between
CuA3+4 and AP. No experiments were done with living
specimens, but the presumed function of this fold is toprovide an additional folding line, strengthening the con­cave longitudinal fold (clf). Clf is now a strip of sclero­
tized wing membrane, which crosses the longitudinal fold
and stiffens the wing by its almost vertical position (Fig.
26). This induces a stronger corrugation, necessary tocounteract the folding forces in the longitudinal fold.
Wing articulation and folding
The dermapteran hindwing articulation contains many
unique characters. Compared with Dictyoptera (Figs11-13), the dermapteran median plate is desclerotized
and replaced by a large membranous area (Brodsky,
1994) (Figs 7-10; median plate, i.e. medial and cubital
fulcalare, FMCu). Note that a weakly preserved medianplate is still retained in Pyragra fuscata (Fig. 7B). The
part of the medial basivenale BM adjacent to the medianplate is also desclerotized. The combined radio-medialbasivenale BR + BM is enlarged and has two bendingzones. Axillary sclerite 1Ax is large compared to Blattodea and Mantodea (Brodsky, 1994) (Figs 7A, 11, 12).
All these deviations from the Blattoneoptera ground pat­tern are due to the movements of articular sclerites during
wing folding.
The large membranous area in the centre of the articula­tion, previously occupied by the median plate, is where
several folds intersect (Fig. 25A). It is crumpled in thefolded wing, but outstretched in the unfolded wing. One
fold originates in each of the two bending zones of BR +BM; three other folds begin close to the FAJ and project

into this area; another fold starts between BAA and BM(Fig. 25A). This arrangement of folds is only possible in asoft, pliable membrane; stiff material, such as paper, is
not flexible enough. Therefore, it is impossible to build an
accurate paper model with all folds present in this area
(Fig. 27). The specific arrangement of folds reflects the
movements of the articular sclerites during folding. The
function of the membranous area is to allow crumpling,
and to provide space into which BAA and the anterior
part of FAJ are turned. During wing folding, combined
basivenale BR + BM is flexed in two bending zones.
BAA turns about its pivot with the BM, which makes theproximal corner of BAA draw near BR. FAJ is turned
upside down and it lies underneath AA3 + BAP in thefolded wing (Figs 25, 27).
The presence of two bending zones (proximal anddistal) in the combined basivenalia BM + BR issurprising, because dermapteran basivenalia are loaded
during flight and any bending seems counterproductive.
However, Kleinow (1966) noticed the straightening of theproximal bending zone. This is achieved through low­
ering the metanotum by a contraction of tergo-sternal
muscles. This pulls the median margin of 1Ax ventrally,and pushes its lateral margin dorsally. This tilt promotes
the wing, erects the wing package over the body, andsimultaneously straightens the proximal bending zone in
BR + BM. But, the distal bending zone is unaffected bythis movement. We assume, based on observations by
Kleinow (1966) and the model (Fig. 27), that the distalbending zone is also straightened (but indirectly) by thecerci.
As described above, the abdomen bends over, the cerci
interact with the wing package, and they unfold (“unzip”)
the wing. This action simultaneously stretches the mem­brane of the articulation, and the sclerites are passively
pulled apart and into their flight position. The 3Ax, FAJ
and BAA are expanded to form a rod in the flight-readywing. As inferred from our own observations on
Labidura riparia, the paper model, and Kleinow’s data,
the rod keeps the metanotum and BM at a distance andthe bending zones in BR + BMA straight (Fig. 27). Thisaction prevents the wing from folding automatically. The
arrangement is apparently stable enough to allow the
movements of the wing base during flight, and to with­stand the aerodynamic forces. After landing, the rod is
unlocked by turning 3Ax about its long axis, which is theusual way of folding wings in the Neoptera (Snodgrass
1935).
The differences between the wing articulation of Blattodea and Dermaptera mostly reflect the differences inwing folding mechanisms. In Blattodea, the hindwings
are unfolded simply by promoting them, even if theirapical part is transversely folded (Haas & Wootton, 1996;
Haas, 1998). In Dermaptera, the wing is unfolded by thecerci (Kleinow, 1966; our own observation), which elimi­nates the need for a mechanical system of unfolding thewing from its base. The use of cerci also provides
freedom for further transformations in folding pattern andmechanism. By using the cerci to unfold their wings, Der-489

maptera evolved a folding pattern, which is far too com­plex to be unfolded from the wing base.
Wing venation and the evolution of folding
The wing folding and venation patterns in extant Dermaptera are almost perfectly uniform (Haas, 1995). The
complexity of these patterns suggests that they evolved
only once. The fossil record of the Dermaptera and theirstem line, the “Protelytroptera”, gives some clues to theevolution of folding and venation patterns, even if mostfossils do not have unfolded wings (Carpenter, 1933,
1939; Carpenter & Kukalova, 1964; Kukalova, 1964b,
1966, 1969). The only known exception is dermapteranstem line representative Protelytron permianum, withhindwings showing a relatively simple folding pattern.
The comparison of folding patterns in Dermaptera andProtelytron permianum (Figs 2, 19) shows that the former
is folded twice transversely, whereas the latter has only
one transverse fold. The transformation could have hap­
pened in the following way.
The hindwing of Protelytron permianum has four prin­cipal folding lines. (1) The anal fold, which is the foldrunning between AA4 and AP, and between RP+MA andAP (this folding line is not indicated by Tillyard (1931)
but is necessary to fold the fan under the remigium). (2)The folding line termed “nf” (or ring fold) running fromthe posterior wing margin to point “k”. At the intersectionof nf and the radiating branches, broadenings termed“chitinous patches” are shown (Tillyard, 1931). (3) The
folding line running from point “k” to the anterior wing
margin, and reaching it at point “n”. The latter two folds(nf and fold from point k to n) are certainly new in Pterygota; they allow the wing to fold ventrally - transversely.
And finally (4) the folds located between the radiating
branches of AP (all dotted lines in Fig. 19).
Later in evolution, the fold “nf” shifted distally, accom­
panied by a change in shape and relative proportions ofwing areas, and reached the anterior wing margin inde­
pendently from the fold “k-n”. Consequently, the anal
fold was retained in functional relation with the “k-n”
fold, but lost its connection to “nf” fold.
Simultaneously, fold “nf” had to change from convex
to concave. This step might seem improbable but for thecomplex function of the broadenings in extant Derma­
ptera (Kleinow, 1966; Haas et al., 2000). There, forces
are generated which bend the radiating branches dorsally
along the ring fold and the intercalary branches ventrally.
Considering the specialized structure of the broadenings,
a reversal of the bending direction would imply majorchanges in these structures. However, these are not neces­sary when the convex-concave pattern of the radiating
folds is reversed. Then, the bending direction of the trans­verse fold can change without changing the bending
direction ofthe broadenings (Fig. 28).
In addition to this folding, the dermapteran hindwing
also folds longitudinally. Simple geometric reasoningshows that the longitudinal fold evolved after the trans­verse folding, which is also suggested by the fossil Prote­lytron permianum. A sheet of paper will clarify this
immediately: First, fold the sheet longitudinally and then,490

transversely. After unfolding the paper, the longitudinalfold will be convex throughout its length, whereas thetransverse fold changes from convex to concave at theintersection point with the transverse folding line. Folding
the paper first transversely and then longitudinally results
in a reverse situation, when the transverse fold is convex
throughout, and the longitudinal fold changes fromconvex to concave.
Exactly the same succession as in the paper folds above
occurs in the dermapteran wing in the central wing hinge(Flugelmittelgelenk of Kleinow, 1966), where the innerand outer apical areas, squama and ulnar area meet. Here,the longitudinal fold changes its orientation from proxi­
mally convex to distally concave, but the transverse foldremains convex. The same configuration of folds con­firms that the anojugal lobe has been folded fan-wise
before being folded transversely. However, it is not pos­sible to deduce whether the folding line “nf” moved dis­tally before or after the longitudinal fold evolved, because
the intersection area is reduced. It should also be notedthat the new longitudinal fold is running across both theremigium and the anojugal lobe.
The fossil “Archidermaptera” give no information
about the evolution of hindwing folding in the Derma­
ptera. The oldest known Dermaptera are Jurassic in age(Vishnyakova, 1980) and already had a small wing pack­age. The hindwings were probably already folded by theautapomorphic dermapteran folding pattern and folding
mechanism. Because Jurassic Dermaptera possessed long,annulated cerci, and a small wing package, the cerci were
probably used to unfold the hindwings the same way as inextant Dermaptera (for other cercal functions in extant
Dermaptera see Briceno & Eberhard, 1995). Conse­
quently, short and smooth cerci are not an adaptation tohindwing unfolding. Incidentally, all known “Protelytro­
ptera” with no or simple hindwing folding bear short
cerci.
H IG H ER TAXA AND SPECIES REQUIRE DIFFEREN T
M ETHOD OLOGIES IN CHARACTER ASSESSMENT

Phylogenetically informative morphological charactersof insect species are selected and assessed by direct
observation and comparison. They are usually (1) notvery difficult to recognize, (2) numerous, and (3) not very
difficult to homologize. The major drawback is that theyare (4) frequently unstable and open to unrecognizable
reversals. Phylogenetic methods using parsimony andconsensus are designed to deal with back mutations andother complicating factors.
In contrast, phylogenetically informative, stabilized
characters in higher taxa of insects are exclusively only
those embedded in their ground patterns. The characters
are invariably extremely old (minimum 100-350 my;
Marshall et al., 1994; Raff, 1996). The problems withhigher taxa are very different from those encountered in
species. (1) Ground pattern characters are relatively few.
Therefore, each error potentially has a large impact. (2)They are sometimes difficult to find in extant insects
because they have been hidden under later (suborderthrough-species level) reductions, fusions and extensions.

(3) They must be very carefully and fully homologized,
because the putative “homologue” is too often a homoplasy. But, full homologization must include the relatedorders and superorders, a very broad and sometimes
unmanageable task. (4) The modes of character transfor­
mation are actually very few (reductions, fusions, braces
and extensions) but they occur in parallel at very many
unrelated taxonomic levels. The paucity of choices versus
frequent co-occurrences creates a huge web of homoplasies. (5) The unrelated complexes are frequently adapted
to a similar function. This also generates homoplasies. (6)Most of the higher-level characters are integrated in oldmorphological complexes (wing complex, mouthparts,
pleura, genitalia, terminalia, etc.), which generate func­tion by working together, and which are readily identifi­
able only as a complex. Such characters neither function
nor evolve individually. Handling them as if they wereindependently functioning and evolving, means imposing
on nature a process that does not exist. Not surprisingly, it
is only after a character complex is disassembled and its
characters are evaluated at too high a level (e.g., under
Pterygota instead under Blattodea) that higher character
states turn homoplastic. Note that the same observation isvalid also for the limb-derived arthropod appendages on
the head (e.g., the much disputed “synapomorphic” man­
dible in Odonatoptera and Neoptera), the thorax, and theabdomen.
Assessing order-level characters outside their orders
always creates homoplasies. As a pertinent example, thestem of M (= a derived fusion of MA + MP) is an apomorphy. M occurs in the representatives of Pleconeoptera, Orthoneoptera, Ephemeroptera and Odonata.
Under Pterygota, M shows these groups as related, which
is a homoplasy unsupported by other wing characters. In
contrast, under Palaeoptera M supports palaeopterous
monophyly (all Palaeoptera share the stem of M and thestem of Cu; these two stems are absent from the groundpattern of Neoptera). This monophyly and sistergrouprelationship is supported here in Table 6A by 63 addi­
tional characters. Under Neoptera, the rise of M supports
the basal division of Neoptera into two groups: Pleconeoptera + Orthoneoptera (M is formed and shared) andBlattoneoptera + Hemineoptera + Endoneoptera (M did
not form). This division is supported in Table 6B by 28
additional characters.
Most important, no character complex will yield goodresults in character analysis if some characters are leftout. As an example, the previously overlooked vein ScA+
turned out to be very important in analyzing Orthoneo­
ptera, Blattoneoptera and Endoneoptera. The equally
overlooked precostal strip (PC) polarizes Palaeoptera and
Neoptera, defines orders in Hemineoptera and superfami­lies in Coleoptera (Table 6A, H). The precise nomencla­
ture of anal veins, which were previously randomly
serially numbered as 1A, 2A, 3A, provides documenta­tion that Neoptera contain two groups of superorders withdifferently composed anojugal lobes (Fig. 22). The wing
function (wing folding, flight flexion lines), previously
seen as separate topic, became very important inrevealing higher- level homoplasies and transformations,etc.
The coding of wing characters in the ground patterns ofhigher taxa is a quite different process than in species.
Importantly, coding is by no means intuitive. Neither
does it use the ancestral or democratic method (sensu
Prendini, 2001). Instead, it observes evolutionary steps, inwhich the character complexes evolved. Principal wingveins and articular sclerites evolved from being present,
to becoming gradually reduced and then lost; and, frombeing independent (separate), to becoming braced andthen progressively fused. A principal vein or articularsclerite in a ground pattern when fused, does never unfuse
and when completely reduced and lost, does not reappear.
There are no reversals in the higher ground pattern char­acters in any representatives (see Table 6). This observa­
tion so far has never been falsified. Thus, the character
coding in Table 6 is based on the predictability of evolu­
tionary changes in the descendants. We know that sistergroups share similar fusions, extensions, reductions andlosses. We also know that each next step in wing evolu­tion will bring further fusions, reductions and losses inthe descendants. If fusions and losses are “removed” froma ground pattern, the ancestral condition is revealed. If we
continue long enough in “removing” derived characters,
we reach the hypothetical protowing, without braces,
reductions or fusions.
The phylogenetic tree lends itself to reconstruction only
because the ground patterns of higher taxa are composedof stabilized character states (i.e., expressed in all repre­
sentatives and without any reversals). If they were notstabilized, any higher taxon would sooner or later dissi­
pate and could not be described or visually recognized inliving species. Only the stability of ground pattern charac­ters over hundreds of millions of years makes it possible
to have higher taxa and to arrange them into phylogenetic
trees.
Thus, higher phylogenetics is completely impracticablewithout first finding the ground pattern character states inall potentially related higher taxa. This is achieved byvisually recognizing, in each taxon, the character states
shared by all its representatives. If a character is highly
specialized and transformed, this transformation must be
smoothly derivable from the shared ground pattern state.
This search is obviously a long-term undertaking.
In summary, the only phylogenetically informative
morphological character states in the higher taxa are thosecontained in their ground pattern. The ground patterncharacters must be carefully identified, fully homo­logized, and none must be left out. They must be judged
at all times within a double context: (1) of the own mor­
phological character complex and its function, and (2) ofthe ground pattern of the indigenous taxon. In the neopterous and palaeopterous higher taxa, the fusions, reduc­tions, braces and extensions are added step-wise. They
define the superordinal lines, superorders, orders, subor­ders and sometimes families, and reflect theirrelationship. The process of tracing all steps (nodal trans­
formations) is shown here only in Dermaptera. Further

491

information is available in the files of JKP and will be
published in monographs. Note that below the familylevel a parsimony analysis usually brings better results
than the method described above.
CONCLUSION

The relationship of Dermaptera to the other pterygoteorders, as analyzed in this account and presented in Table
6, offers the following results.
Neoptera: [Pleconeoptera + Orthoneoptera] + [Blattoneoptera + (Hemineoptera + Endoneoptera)].
Blattoneoptera: (Grylloblattodea + (Dermaptera +Dictyoptera)).
Dictyoptera: (Mantodea + (Blattodea + Isoptera)).
Dermaptera: Karschiellidae + (“Diplatyidae” +(“Pygidicranidae” + (Allostethus indicum + (Anisolabididae + (“Labiduridae” + [Forficulidae + (Chelisochidae + Spongiphoridae)]))))). Taxa in quotation marks
are probably not monophyletic.
The step-wise assembly of the dermapteran hindwing
venation and articulation is much easier to understand if
we offer the broad overview of nodal evolutionarychanges in the Pterygota. Only a character table of theentire pterygote wing complex provides, in Dermaptera,
the necessary amount of order-level information, which
separates synapomorphies from overwhelming, omnipres­ent, and otherwise “unrecognizable” homoplasies. As
another advantage, a table renders the wing charactersmore user-friendly for study than mere description.
Constructing a complete and flawless character table ofthe pterygote wing complex at the higher level willprobably take another decade. Nevertheless, we proposethat starting the conceptualization now will contribute to
the clarity of this paper, open the topic to contributions,and make easier any future use of the wing structure inhigher phylogenetics. For these reasons, we offer below
the first version of the fully homologized pterygote wing
character table.
Table 6. The character table of the pterygote wing
complex.
Introduction. The character table of the fully homolo­
gized pterygote wing complex (with emphasis on the
Neoptera) is based on a long-term study of the wing com­plex (wing venation and articulation) by Kukalova-Peck.
Symbols for veins, sclerites, cross veins and brace are as
defined by Kukalova-Peck, 1983, 1991, 1997; KukalovaPeck & Lawrence, 1993; and as presented in an improved
version in this account (Figs 1A, 1B; Table 6). Note thatthe higher-level characters of wings as mentioned in thetable cease being informative at about the family leveland below.

Ta b l e 6A. Character state distribution in Protowing, Neoptera and Palaeoptera.
Protowing characters(ancestral to clades 1, 0) all coded as 0

Clade 1. Neoptera (Fig. 1)

Clade 0. Palaeoptera

Kukalova-Peck, 1978; Kukalova-Peck&
Brauckmann, 1990; Wootton & KukalovaPeck, 2000

Kukalová, 1964a; Kukalová-Peck, 1978,
1983, 1991, 1997; Brodsky, 1994

Kukalová-Peck, 1978, 1983, 1985, 1991,
1997; Brodsky, 1994

Flight, Folds, Wing Flexing
Protowings, as recorded on fossil prothora­ 1 - prothoracic wings immobilized,
ces: mobile, membranous with dense
veins reduced; later fused with tergum
irregular cross-veins, anterior margin
into prothoracic shield, many times informed, veinal sectors fluted, articularparallelsclerites form short, arched welt

0 - present (in fossil odonatoids Geroptera& Palaeodictyoptera); later veined, dimin­
ished, in Ephemeroptera stem line, or fusedwith tergum into prothoracic shield, or lost

Meso- & metathoracic protowings: bothpairs almost equally wide

1 - heteronomous: forewings narrow,
hindwing with broad anojugal lobe; later
hindwings diminish repeatedly, in par­allel

0 - homonomous; later hindwing somewhat
broader near base or reduced, or petiolate,
repeatedly in parallel

Meso- & metathoracic protowing pairs,arrangement of veins: homonomous &radialMembrane: rather thick with dense,
irregular cross-veins, in both protowing
pairs

1 - heteronomous, anal area inforewings asymmetrical, adapted forlocking them in flexed position
1 - heteronomous, forewings tegminous,
hindwings thinly membranous withweak irregular cross-veins

0 - wing pairs homonomous, with radiallyarranged wing venationFore- & hind protowing pair: similar func­tion

1 - heteronomous function: forewings
protecting & flying, hindwings principal
flying pair

0 - homonomous function, both pairs fly­ing

Wing flexing: protowings are pulled closer
to body & warped (in nymphs ofDiaphanopterodea only)

2 - 3Ax pivots, collapses into a mem­
branous window, pulls on three arms
articulated with BCu, BA, BJ & foldwing veins

1 - gaps between rows of sclerites Cu, A, J
close, pliable veinal bases bend, wigs flex
(in adult Diaphanopterodea only)492

0 - homonomous, both wing pairs membra­
nous with dense, irregular cross-veins

Table 6A (continued).
Protowing characters(ancestral to clades 1, 0) all coded as 0

Clade 1. Neoptera (Fig. 1)

Position of protowings flexed backward:
half-closed in all three thoracic wing pairs(as in nymphs of Diaphanopterodea)

0 - flexed wings held mildly roofwise in 0 - mildly roofwise, half-closed (in basal
half-closed position (in fossil hemipteradult Diphanopterodea only); later overlap
oids: Paoliidae only); later overlap api­
in derived representatives (Kukalova-Peckcally, partly, or completely& Brauckmann, 1990)
2 - folds run in membrane between
1 - wing flexing folds cross veinal bases (inveinal sectors
Diaphanopterodea only); later wing flexing
lost in all other Palaeoptera, (twice inde­
pendently)

Folds: uncertain

Clade 0. Palaeoptera

Medial fold absent
Claval fold absent
Anal fold absent
Jugal fold absent

1 - medial fold present
1 - claval fold present
1 - anal fold present
1 - jugal fold present

0 - absent
0 - absent
0 - absent
0 - absent

Venation in protowing

Venation in forewing

Venation in forewing

PC strip: thick, separated from CA by asuture0 - thick; later PC expands into epipleuron (in Hemiptera, again in Coleoptera)
0 - same (in Hemiptera, Coleoptera);later CP- fuses with CA+ close to base

1 - PC strip narrow, sclerotized, serrated;
later serration lost in many living Ephemerida
1 - CA+ and CP- run in parallel in basal
third of the wing (in many fossils); later
they fuse into C close to base0 - same
0 - homonomous, radially arranged veins inboth wing pairs

Costa (C): CA+ and CP- run in parallel toeach other, fuse into C beyond mid-wing
Anal lobe not enlarged
Anal veins, arranged: radially, spaced inregular intervals, in both protowing pairs

Locking devices to hold protowings inwarped position: probably absent
Jugal lobe: regular size in both protowing
pairs
Jugal lobe membrane: not different fromother areas
Remigium & anal &jugal lobe: all posi­tioned at one plane

0 - same
1 - heteronomous: AA1+2 shifted ante­riorly, distanced from AA3+4; AA3+4& AP fused basally (radially arranged
veins in hindwings)
1 - long anal bar locks on itself; later it
locks on scutellum or on articular sclerites
2 - heteronomous size:jugal lobe much
shorter than anal lobe (regular size inhindwings)
1 - thinner than in the rest of forewing

0 - locking device absent

1 - strongly diminished, homonomous inboth wing pairs
0 - not different

1 - heteronomous position: remigium & 0 - homonomous position: at one planeanal lobe at one,jugal lobe at another
plane

Venation in protowing

Venation in hindwing

Venation in hindwing

Veins arranged: radially
PC strip: thick, separated from CA+ by asuture0 - radially
0 - same; later PC strip narrow, fused
into anterior margin, or expanded into
anterior lobe in Dermaptera & Coleoptera

0 - radially
1 - PC strip narrow, sclerotized, serrated,
separated by suture from anterior margin;
later serration lost in most Ephemerida

Veinal areas: placed at one plane

0 - same

0 - same

CuP: dichotomouslybranched

0 - present (in Orthoneoptera, hemipteroid stem line, Fulgoromorpha, Neuroptera); later simple, or lost

0 - present (in palaeodictyopteroids only);
later CuP simple in Ephemeroptera + Odonatoptera

Anal basivenale (BA): not elongated or
subdivided (anal lobe regular, notenlarged)

1 - BA subdivided into three articulated
subdivisions: BAA1+2, BAA3+4, BAP(anal lobe enlarged)

0 - BA not subdivided (anal lobe notenlarged)

Remigium and anojugal area: not forming
two flight units (protowing does not fly)

1 - form flight units separated by claval 0 - not forming flight units
flexion line (in Pleconeoptera + Ortho­
neoptera); later by anal fold in Blattoneoptera + Hemineoptera + Endoneoptera

493

Table 6A (continued).
Protowing characters(ancestral to clades 1, 0) all coded as 0

Clade 1. Neoptera (Fig. 1)

Clade 0. Palaeoptera

Anal area, size: regular, not enlarged

1 - anal (anojugal) lobe strongly
enlarged, undulating in flight; later
diminished many times, in mostEndoneoptera & many other groups
0 - enlarged;
0 - same

0 - regular (in Diaphanopterodea); laterslightly enlarged in Palaeodictyoptera orstrongly diminished in Ephemeroptera +Odonatoptera
1 -highlyreduced
0 - same

1 - anal brace formed by contact
between two basivenalia (BCu & BAA)
connected at an angle

2 - anal brace connects anal branches:(sclerotization crossing veins in Palaeodictyopteroidea; arched parts of veins in
Ephemeroptera + Odonatoptera
1 - branches reduced, very short, condensed

Jugal area: regular size
Remigium & anal &jugal area: position at
one plane
Anal brace: absent

JA, JP braches: regular, dichotomously
branched

1 - branches long, further divided,
widely spread

Venation in protowings

Venation in both wing pairs

Veinal corrugation mild, alternating:
ScA+, ScP-, RA+, RP-, MA+, MP-,
CuA+, CuP-, AA+, AP-, JA+, JP-;cross-vein network also corrugated

Venation in both wing pairs

2 - alternating corrugation always
retained in: ScA+, ScP-, RA+; corruga­tion reduced or reversed in RP, MA,
MP, CuA, CuP, AA, AP, JA, JP & incross-vein network
The stem ofR: absent (sectors RA, RP
0 - R absent (in plecopteroid & hemistart independently from the radial
pteroid stem lines); later R very short inbasivenale BR)
Isoptera, Coleoptera, Strepsiptera, or
repeatedly longer
The stem ofR: absent
1 - derived: when formed, RA superim­
posed on RP
The stem ofM: absent (sectors MA & MP 0 - M absent (in all Blattoneotera +start independently from medial basivenale Hemineoptera + Endoneoptera); lateroccurs in Pleconeoptera + OrthoneopteraBM)
0 - absent (in Orthoneoptera only); later
The stem of Cu: absent (sectors CuA &the stem of Cu occurs repeatedly in all
CuP start independently from cubitallines
basivenale BCu)
CuP: dichotomouslybranched
0 - branched (in Orthoneoptera, hemipteroid stem line, Fulgoromorpha); later
often simple, repeatedly in parallel
Flight-important cross-vein brace arculus
0 - absent (in both wing pair in pleco­(mp-cua or m-cua or short fusion): absent
pteroid & blattoid stem lines, Isoptera);later absent in fore wings only in Blattodea, or added in both pairs, repeatedly

1 - alternating corrugation in ScA+, ScP-,
RA+, RP-, MA+, MP-, CuA+, CuP-, AA+,
intercalated sectors & network; corrugation
reduced in AP, JA, JP

Articulation in both wing pairs: Pro­
towing condition hypothetical, coded as 0
Sclerites in the precostal row (PC) andcostal row (C): free, not fused into PCC
sclerites

Articulation in both wing pairs

Articulation in both wing pairs

2 - fused into PCC sclerites in all col­umns (PR, AX, F, B), form three com­posite sclerites: PRPCC, tegula(AXPCC), humeral plate(FPCC+BPCC)

1 - PC, C stay free in columns PR, AX;later fuse into PCC only in columns F, B,
thus forming FPCC, BPCC (in Ephemeroptera)

PRPC separate from PRC (fusion in col­umns absent)

1 - PRPC+PRC fused into composite
sclerite PRPCC proximally from tegula(often retained, small, very large in
Strepsiptera)

0 - PRPC, PRC separate (in Ephemeroptera+ Odonatoptera only); later fused into
PRPCC in Palaeodictyopteroidea

Sclerites in rows Sc, R, M, Cu, A, J articu­
lated one with another (regular arrange­
ment, no fusions, no clusters)

2 - varied: sclerites in row Sc somearticulated some fused; in rows R, M
unconnected; in rows Cu, A, J articu­
lated one with another1 - articulated, but clustered centrally atwing base (as in Diaphanopterodea); later
this triggered twice fusions into articularplates, once in Palaeodictyoptera + Megasecoptera, again in Ephemeroptera +Odonatoptera

494

0 - R absent (in fossil Geroptera only); later
long R formed in parallel, in derived
Palaeodictyopteroidea, Odonatoptera, Ephe­meroptera
2 - derived: when formed, RA & RP fusedlaterally to each other1 - M (MA superimposed on MP) always
present, long
1 - the stem of Cu always present (CuA
superimposed on CuP)
0 - branched (in Palaeodictyopteroidea);
later CuP simple in Ephemeroptera + Odonatoptera
0 - absent (in Palaeodictyoptera); later
added repeatedly in some palaeodictyopteroids and in Ephemeroptera + Odonatoptera

Table 6A (continued).
Protowing characters(ancestral to clades 1, 0) all coded as 0

Clade 1. Neoptera (Fig. 1)

Clade 0. Palaeoptera

Sclerites arranged in four regular, aligned
columns: PR, AX, F, B

1 - central sclerites highly irregular, insize, shape, alignment (oblique associa­
tion connected with flexing, locking
wings in resting position
1 - fusion present: either in anterior
wing process AWP (PRScR fused with
tergum), or in posterior wing process
PWP (PRAJ fused with tergum), or in
both, repeatedly formed
1 - present, include: 1Ax(PRM+AXR+AXSc+FSc),
2Ax (AXM, FR), 3Ax (AXCu, AXAJ,
FCu, FA, FJ)
0 - separate (in Dermaptera, neuropteroids); later repeatedly together in AWP

0 - regular, aligned columns present

Anterior and posterior wing process absent(proxalaria PR are articulated, never fused,with tergum,)

Composite sclerites selected obliquelyfrom several disturbed columns: absent

Sclerites PRSc, PRR: separate (not fused
together)
Sclerite PRCu: sclerotized

Sclerites PRA, PRJ: not fused one toanother

Humeral plate: absent

Tegula: absent
First axillary lAx, irregular oblique com­posite sclerite: absent

Second axillary 2Ax, irregular obliquecomposite sclerite: absent
Third axillary 3Ax, irregular, composite,rotating sclerite with wing flexor muscle:absentSclerite AXM: regular, not different fromother sclerites
Sclerite AXCu: regular, not subdivided

Sclerites AXA, AXJ: separate, regular, not
fused

1 - PRCu occupies membranous
window area; pliable, ribbon-like (inDermaptera, large aphids); later frag­mented in Neuroptera, or repeatedly lost
1 - fused into PRAJ (= 4Ax) (in Orthoptera, Hymenoptera, Coleoptera); later
PRAJ repeatedly fused with tergum as
PWP
1 - present, composite of four fusedsclerites (FPCC+BPCC); sutures
between sclerites retained in basal mem­bers
1 - present, a composite of two sclerites(AXPCC) bearing sensory organ
1 - 1Ax composed of
PRM+AXR+AXSc+FSc separated bysutures (in Neuroptera only); later
sutures variously repeatedly reduced
1 - 2Ax contains triangular large body
AXM, articulated with narrow arm FR
1 - 3Ax contains goblet & heel (AXCu
subdivided by hinge), saucer (AXAJ), &three arms, cubital (FCu), anal (FA),jugal (FJ)
1 - AXM triangular, enlarged
1 - AXCu subdivided by hinge, forms
two lobes, larger (goblet) & smaller(heel)
1 - elongate, fused into AXAJ (saucer)

Sclerite FR: regular

1 - FR (2Ax arm) slender

Sclerite FCu: articulated proximally with
AXCu, distally with BCu

0 - articulated; fulcalaria (F) never fusewith basivenalia (B) (it would prevent
wings from flexing backward)
1 - FA articulated with fused sclerite,saucer (AXAJ) & with part ofbasivenale (BAA3+4)

Sclerite FA: articulated with two separate
sclerites, AXA & BA

0 - fusions between PR and tergum always
absent0 - absent

0 - separate (in fossil Diaphanopterodea,
Ephemeroptera, Odonatoptera); later fusedin derived Palaeodictyopteroidea
0 - PRCu visible in fossils & basal, large
representatives of all lines; later in parallel
desclerotized, visible when stained
0 - not fused one to another; later gradu­
ally membranized in Ephemeroptera &Odonatoptera, or fused in derived Palaeodictyopteroidea
0 - humeral plate absent

0 - tegula absent
0 - oblique composite sclerite 1Ax absent;sclerites always arranged in regular rows &undisturbed columns
0 - oblique composite sclerits 2Ax absent;sclerites always arranged in regular rows &columns
0 - absent; sclerites always arranged inregular rows & undisturbed columns

0 - unmodified
0 - not subdivided

0 - AXA, AXJ separate, regular (in Diapha­nopterodea only); later repeatedly &variously combined
0 - unmodified
0 - articulated (in wing-flexing Diaphano­
pterodea only); later fused with BCu &variously incorporated into articular plates
0 - present (in Diaphanopterodea only);
later FA fused with BA twice, in articularplate of palaeodictyopteroids, again of
Ephemeroptera + Odonatoptera
495

Table 6A (continued).
Protowing characters(ancestral to clades 1, 0) all coded as 0

Clade 1. Neoptera (Fig. 1)

Clade 0. Palaeoptera

Sclerite FJ: articulated with AXJ & withBJ

1 - F J articulated with 3Ax saucer(AXAJ) & with BJ

Sclerite FM & FCu: separate (not fused
into medial plate or into articular plates)

0 - separate (in Plecoptera, Mantodea,
Coleóptera); later fused repeatedly intomedial plate

0 - present (in Diaphanopterodea only);
later FJ fused with BJ twice in articularplates
0 - separate (in Diaphanoptera only); later
incorporated twice into articular plates, inderived Palaeodictyopteroids, again in
Ephemeroptera + Odonatoptera

Basivenal column (B): straight

1 - V-shaped

Subcostal basivenale BSc: subdivided into
two similar sublobes separated by suture,
BscA & BScP

0 - BscA & BScP present (in Blattodea, 0 - present (in fossil Odonatoptera, Ephe­
Mantodea); later sublobes dissimilar, or meroptera); later sulcus lost in living repre­
suture reduced
sentatives & again in Palaeodictyopteroidea

Sclerite BSc, length: short
Basivenalia BR, BM, BCu, BA, BJ & fulcalaria FR, FM, FCu, FA, FJ: articulated
one to another1 - long,jutting proximally
0 - articulated (condition enabling wingflexing but preventing energy-saving
gliding

Anal basivenale BA: regular (not length­
ened or subarticulated)

0 - straight0 - BSc short, notjutting
1 - articulated, but B & F i n a dense, hinged
cluster centrally (in Diaphanopterodea
only); later B & F fuse twice, resulting
plates enable energy-saving gliding
1 - BA long, subarticulated into
0 - BA regular (in Diaphanopterodea only);
BAA1+2, BAA3+4, BAP (to anchor
later diminished, fused into articular plates,
long fan ofbranches supporting enlarged twice independently
anal lobe

Ta b l e 6B. Character state distribution in Orthoneoptera + Pleconeoptera and Blattoneoptera + Hemineoptera + Endoneoptera.
Characters: states ofNeoptera groundpattern coded 0 o r 1 i n clades 2o r 3

Clade 2. Orthoneoptera +Pleconeoptera (Figs 14,15, 21, 22)

Clade 3. Blattoneoptera + Hemineoptera+ Endoneoptera (Figs 4-13, 20, 21, 22)

Two flight units separated by flexion line

1 - remigium & full anojugal lobe

2 - remigium + AA & partial anojugal lobe

Anojugal lobe, veinal support

0 - full anojugal lobe supported by AA,
AP, JA, JP
1 - full, large (in basal Plecoptera,
Orthoneoptera); later reduced in derived
Plecoptera, in Embioptera, or enlarged& branches added in Orthoptera

1 - partial anojugal lobe supported by AP,
JA, JP2 - partial, moderately large (in basal Fulgoromorpha, Hymenoptera, Coleopteroids,
Neuropteroids, Trichoptera); later enlargedin Blattoneoptera, or gradually reduced inmost Endoneoptera
1 - shallow, shortened
2 - anal fold
2 - irregularly fan-like (folds unsettled,
may cross branches, secondary folds added)

Flight, Folds and Folding

Anojugal lobe, size

Claval flexion line (fold)
Principal flexion line/foldAnojugal lobe, internal folding

0 - deep, full length
1 - claval flexion line
1 - regularly fan-like (folds between
anal &jugal branches equally divergent)

Apical folding (apical field withdrawn intowing at rest)
Jugal lobe proped by secondary brace inarticular membrane

0 - never formed

Wing position at rest mildly roofwise,
incompletely flexed backward, wings do
not overlap
Wing venation
PC strip separated by suture from anterior
margin (as in protowing)
Forewing, CA & CP separate, fuse far
from base (as in protowing)496

1 - apical folding evolved several times, in
Dermaptera, some Blattodea, Coleoptera
0 - absent

1 - brace present, long, runs frompostero-distal corner of metanotum to
BJ
1 - wrapped around body dorsally & lat­ 0 - present (in Hemipteroid stem line &erally (in Plecoptera, Embioptera,some Endoneoptera); later highly roof-like
Orthoptera)or dorsally flat, wings crossed at tips or
partly or fully

2 - PC strip fully incorporated into ante­ 0 - PC separated by suture; later extendedrior margin
anteriorly into lobe in Dermaptera, Coleo­ptera
1 - CA & CP fused into C near wing
0 - present, CA forms anterior margin, CP
baseruns in parallel, fuse apically (in Hemiptera,
Coleoptera only)Table 6B (continued).
Characters: states ofNeoptera groundpattern coded O o r l i n clades 2o r 3

Clade 2. Orthoneoptera +Pleconeoptera (Figs 14,15, 21, 22)

Clade 3. Blattoneoptera + Hemineoptera+ Endoneoptera (Figs 4-13, 20, 21, 22)

ScA branched,joins anterior margin far
from wing base
The stem ofR (absent in protowing)O- present (in plecopteroid stem line &Orthoneoptera only)
0 - absent (in plecopteroid stem line
only); later R arises repeatedly in Plecoptera, Embioptera, Orthoneoptera
1 - always present, long

1 - ScA changed into convex, blunt, sclero­
tized, oblique ridge
0 - absent (in hemipteroid stem line only);
later extremely short in Dermaptera, Isoptera, coleopteroids, or lengthened
0 - always absent

Sectors MA, MP arise separately frombasivenale BM (as in protowing)
Sectors MA, basal portion

1 - fused into the stem ofM which forks
distally into MA & MP
2 - distinct in the fork

0 - present

Sector MP, basal portion2 - distinct in the fork

The stem of Cu (absent in protowing)

JA, JP dichotomously branchedO- absent (always absent in Orthoneo­
ptera; later present in Pleconeoptera)O- branches broadly diverged (in Plecoptera); later number of AA branches
increased in Orthoptera
O- present (in Pleconeoptera); later
number of AP branches increased in
Orthoptera
O- present

Embayments in posterior wing margin

0 - one, claval

The stem ofM (MA+MP) (absent in pro­towing)AA dichotomously branched several times(as in protowing)
AP dichotomously branched (as in proto­
wing)

Articulation
PRSc & PRR separate

PRCu (reducing, placed in the way of col­lapsing 3Ax)
4Ax (=PRAJ articulated with tergum);(PRAJ fused with tergum = PWP)

1 - AWP present (PRSc+PRR fused
together & with tergum into anterior
wing process)2 - lost

1 - obscured: MAjoins RP or R immedi­
ately at base, entry often membranized (evi­dent in dissections under water)
1 - faint/obscured (visible in Mantodea &neuropteroids only); later visible only in
dissections
2 - present, very short (in Dermaptera,
Isoptera; later repeatedly lengthened
1 - AA branches few, narrowly diverged;
later progressively reduced, repeatedly inparallel0 - present (in basal Hemineoptera &Endoneoptera; later number of AP branches
increased in Blattoneoptera
0 - present (in Blattoneoptera); laterbranches reduced in Hemineoptera,
Endoneoptera
1 - two, larger anal, smaller claval
0 - AWP absent (in Dermaptera & neurop­
teroids only); later often formed, in parallel
1 - present (forming a long, pliable ribbon
or fragmented); later lost, repeatedly in par­allel

3Ax anal arm (FA) articulated with 3Ax
saucer & with BAA3+4 (as in Neoptera
ground pattern)
3Axjugal arm (FJ) articulated between
3Ax saucer & BJ (as in Neoptera groundpattern)
BSc, sublobes BScA & BScP

1 - 4Ax present (in basal Orthoneoptera 1 - 4Ax present (in Hymenoptera & Coleonly); later PWP arises many times
optera: Gyrinidae only); later PWP arises
many times
1 - regular
2 - more or less modified (in Hemineoptera& Endoneoptera); later fully membranizedin Blattoneopteral - triangle large, strongly & fully
2 - triangle small, curved, distally desclerosclerotized
tizedl - lid-like, about as long as the length
2 - longer than 2Ax body, oriented at anof 2Ax bodyangle
0 - separate (in Plecoptera); later medial 0 - FM, FCu separate (in Mantodea, Coleoplate formed in Embioptera, again in
ptera); later medial plate repeatedly formed
Orthoneoptera
1 - present
1 - present (in Blattoneoptera only); laterFA always fused with 3Ax saucer (AXAJ),in Hemineoptera + Endoneopteral - F J lost, connection between 3Ax &0 - present (in basal Blattodea only); later
BJ lostFJ lost in derived Blattodea, fused withsaucer in Hemineoptera + Endoneoptera
0 - sublobes BScA & BScP equally longl - BScA distinctly longer than BScP

BJ supported proximally by a long secon­
dary strutl - present, secondary strut runs inarticular membrane

PWP when formed

2Ax body (AXM) triangular
2Ax radial arm (FR)FM, FCu separate (as in Neoptera groundpattern: fused FM+FC form medial plate)0 - B J not supported, strut absent

497

Ta b l e 6C Characters state distribution in Pleconeoptera (including Embioptera) and Orthoneoptera.
Characters: states ofthe shared ancestor
coded O o r l i n clades 8o r 9

Clade 8. Pleconeoptera including
Embioptera (JKP unpublished observa­tion) (Fig. 14)

Clade 9. Orthoneoptera (Figs 15, 21, 22)l - lobe tends to diminish in derived
membersl - runs transversely in articular mem­brane

2 - lobe mostly tends to enlarge in derived
members
2 - runs close to posterior wing marginl - sclerotized2 - desclerotized basally under tightly fit­
ted prothoracic shield2 - extra long & branched, replaced C insupporting anterior margin (C too weakunder prothoracic shield)l - present, long, strong, basally supported
by M+CuA running close

Flight, Folds and Folding
Full anojugal lobe large
Long secondary strut in articular mem­brane supportingjugal lobe
Wing venation
Forewing, anterior margin
Forewing, ScA+ before joining anterior
margin (long, branched in protowing)O- present (in plecopteroid stem line
only); later progressively simplified &shortened
The stem ofR (RA + RP fused) (absent in O- absent (in plecopteroid stem line
protowing)only); later R formed in Plecoptera,
Embioptera
Arculus, cross-vein brace mp-cua or m-cua 0 - absent (in stem line plecopteroids:or short veinal fusion (absent in
Liomopterida only); later arculus added
protowing)in Plecoptera, Embioptera
The stem of Cu (CuA + CuP fused)
3 - Cu present, relatively long(absent in protowing)
Forewing, CuA deeply forked into long,1 - unique deep fork present (in plecop­
branched CuAl+2 & short CuA3+4
teroid stem line, Plecoptera,
Embioptera)
CuP branchedl - CuP simple, branches lost
Branch CuPl+2
l - regular branchAnal branches, number

0 - regular; later reducedl - replaced by unique, long fusion of CuA
fused since base with M, then with MP,then separating apically from MP
O- Cu always absent
O- CuA branches not modified

O- CuP branched
2 - CuPl+2 forms important veinal brace:in forewing between CuP & MP+CuA: inhindwing between CuP & M+CuA (JKP,
unpublished)O- number regular; later increased

Articulation
AWP (PRScR fused with tergum)
PRAJ (4Ax, PWP)

1 - AWP present, largel - PWP present, shortlAx, head (FSc) & neck (AXSc)l - lAx head articulated with BSc,
fused with lAx neck (suture preserved)(as in Neoptera ground pattern)
2Ax, body (AXM) & arm (FR) (articulated l - arm lid-like, articulated with robust,
triangular 2Ax bodyin Neoptera ground pattern)
3Ax cubital arm (FCu) (free in Neoptera
ground pattern)
3Ax, anal arm (FA) (articulated with
AA3+4 in Neoptera ground pattern)

0 - free (in Plecoptera); later fused inmedial plate, in Embioptera
1 - FA present, relatively short (in Plecoptera)

BSc with sublobes BScA & BScPl - BScA longer than BScP, thin, pro­truding strongly proximally

2 - AWP reduced
0 - 4Ax present (in Acridoidea only); later
PWP arises, repeatedly in parallel
2 - lAx head associated/fused with BSc,
articulated with pliable lAx neck2 - arm lid-like, partly fused with robust,
triangular 2Ax body; later arm broadened inderived Acridoidea
1 - fused in medial plate
2 - FA long, runs in parallel with extra long
BAA, connected with BAA by broad,flexible sclerotization
2 - fused into short BSc

Ta b l e 6D. Characters state distribution in Blattoneoptera and Hemineoptera + Endoneoptera.
Characters: states ofthe shared ancestor
coded O o r l o r 2 i n clades 4or 5

Clade 4. Blattoneoptera(Figs 4-13, 20-22)

Clade 5. Hemineoptera + Endoneoptera(Figs 16,17, 22)

Flight, Folds and Folding
Flight powered from both wing pairs,
forewings protective, hindwings main
flying pair (similar as in Neoptera groundpattern with full anojugal lobe)l - forewings terminous, narrower, fly­ing; hindwings main flying pair, mem­
branous, with broad anojugal lobe

2 - forewing thinning, hindwing dimin­
ishing (basal states as in Fulgoromorpha,
Neuroptera); later forewings mainly flying,coupled with smaller hindwings, inHymenoptera, mecopteroids, or only hindpair powered in Coleoptera + Strepsiptera

498

Table 6D (continued).
Characters: states ofthe shared ancestor
coded 0 o r l o r 2 i n clades 4or 5

Clade 4. Blattoneoptera(Figs 4-13, 20-22)

Clade 5. Hemineoptera + Endoneoptera(Figs 16,17, 22)

Claval fold: flanked by CuP & AAl+2

1 - CuP & AAl+2 unstable (either may
descend into bottom of claval fold)
2 - anal fold weakened but stabilized,
does not cross APl+2 branches

0 - present; later either CuP or AAl+2 orboth become reduced
1 - anal fold erratic, may cross APl+2 or
get adjacent to it (in Hemiptera & Neuroptera)

Anal fold reducing, erratic

Hindwings with partial anojugal lobe,
slightly shorter than forewings, anojugalarea

2 - slightly shorter, with anojugal lobe l - partial lobe broad (basal states in Fulsignificantly secondarily enlarged in AP goromorpha, Hymenoptera, neuropteroids,
coleopteroids, Trichoptera); latermuch
area, supported by added AP branches
diminished, frequently in parallel

Partial anojugal lobe modestly broader
than in forewing

2 - distal part of AP area secondarily
enlarged, bears secondary twigs (com­pensation for AA absent from lobe) (inDermaptera, Dictyoptera)

1 - present (in Hymenoptera, neuropteroids,
coleopteroids, Trichoptera); later lobe fre­quently strongly diminished, narrow

Folding in partial anojugal lobe: slightly
irregular fan-like, along anal, claval &other folds

3 - present (in Dermaptera, Mantodea,
Blattodea); later book-like & flat in blattoid stem line & Isoptera

Wing position at rest: mildly roofwise,
wings not crossed (as in Neoptera groundpattern)

2 - dorsally flat, wings crossed only attips (in blattoid stem line); later partiallyor fully crossed

2 - progressively reduced & less regular (inHymenoptera, neuropteroids,
mecopteroids); later book-like along anal
fold in Coleoptera
0 - mildly roofwise (in basal representa­
tives); later highly roofwise, or dorsally flat,crossed at wing tips, partially or fully

Wing venation
Forewing, PC strip: thick, separated bysuture from CA (as in protowing)3 - integrated into anterior margin

0 - present; later PC enlarged into epipleuron (in Hemiptera & Coleoptera), or
fused with anterior margin

Hindwing, PC strip: present, not integrated
into anterior margin

0 - present (PC forms anterior lobe in
Dermaptera); later repeatedly fully fusedwith C

0 - present (PC forms anterior lobe in Cole­
optera); later repeatedly fully fused with C

Forewing: CA & CP run in parallel for
long distance (as in protowing)

2 - CA+CP fused since base, form
strong anterior margin

0 - CA, CP run separately (in Hemiptera &Coleoptera only); later CA+CP fused into C
repeatedly near base

ScA+ forms a ridgel - ScA ridge narrow, oblique, blunted
on top2 - ridge expanded into broad, convex,
sclerotized bulge

The stem ofR (absent in protowing)l - R very short (in Isoptera); later
repeatedly lengthened

0 - R absent (in hemipteroid stem line
only); later very short in coleopteroids, or
repeatedly lengthened

Special, flight important cross-vein brace
rpma-mp2 - present relatively far from wingbase, short

1 - present near wing base, long (in coleop­
teroids + neuropteroids; later more distaly
in mecopteroids, again in Hymenoptera

Arculus, flight important homoplastic
cross-vein brace mp-cua or m-cua or shortfusion (absent in Neoptera ground pattern)

0 - absent in both wing pairs (in blattoid
stem line & Isoptera); later present onlyin hindwings in Blattodea & Mantodea,
or in both wing pairs, in Dermapteral - always present in both wing pairs (asynapomorphy)

The stem of Cu absent (as in protowing)

2 - present, very short

CuP (branched in protowing)

2 - simple; later lost

0 - absent (in hemipteroid stem line only);
later repeatedly formed
0 - branched (in hemipteroid stem line &Fulgoromorpha only); later simple or lost

AAl+2 simple, strong

2 - simple; later lost

1 - strong (in Hymenoptera only); later lostin Hemineoptera, again in all remaining
Endoneoptera

AA3+4 forks into AA3 & AA4

0 - fork present (only in Dermaptera);later simple (in Mantodea), or supportedby short serial twigs (in blattoid stem
line & Isoptera)

0 - AA3+4 forks into AA3 & AA4 (inColeoptera only); later repeatedly simple

499

Table 6D (continued).
Characters: states of the shared ancestor
coded 0 o r l o r 2 i n clades 4o r 5

Clade 4. Blattoneoptera(Figs 4-13, 20-22)

AP sparsely branched

3 - APl+2 enriched with serial pectinate 0 - AP not enriched (in Hymenoptera, coletwigs added apically (in Dermaptera,
opteroids, neuropteroids); later branches
Dictyoptera)often strongly reduced
0 - dichotomously branched
1 - branches short; later strongly reduced,
in parallel

JA, JP (dichotomously branched in pro­towing)Clade 5. Hemineoptera + Endoneoptera(Figs 16,17, 22)Wing articulation
AWP (absent in Neoptera groundplan,
PRSc & PRR separate, each hinged with
tergum)Composite sclerite PRAJ (as 4Ax when
hinged with tergum; as PWP when fusedwith tergum)
2Ax body (AXM) triangular, proximal
margin thickened, distal part weakened
2Ax arm (FR) narrow
FM, FCu separate (as in Neoptera groundpattern), not fused into medial plate
3Ax anal arm (FA) articulated with 3Ax
saucer (AXAJ) (in Neoptera ground pat­tern)
3Axjugal arm (FJ) (articulated withsaucer (AXAJ) in Neoptera groundplan)BSc, sublobes BScA & BScP (separated
by suture in Neoptera ground pattern)
Basivenale BMBM

0 - AWP absent (PRSc, PRR free in
Dermaptera only); later AWP often
arises
3 - PRAJ always completely membranized3 - 2Ax body curved, proximal margin
thickened, distal part membranized &wrinkled
1 - FR narrow, thin, pliable
1 - FM, FCu separate (in Mantodea
only); later medial plate, tringular, inparallel1 - FA articulated with saucer, narrow(in basal Blattodea); later broad in Blattodea, Isoptera, or fused into FAJ, in
Mantodea, in Dermaptera
1 - F J articulated with saucer (in basal
Blattodea only); later repeatedly lost, or
fused with FA into FAJ, in Dermaptera,again in Mantodea
1 - sublobes robust, suture distinct

0 - AWP absent (PRSc, PRR free in neu­
ropteroids only); later AWP often arises
0 - 4Ax present (in Hymenoptera & Coleoptera: Gyrinidae only); later PWP formed
repeatedly, often partly membranized2 - 2Ax body triagular, partly
desclerotized; later distally enlarged &domed in Hemineoptera
2 - F R broad, lobate
2 - medial plate short & broad, FM, FCu
separated by suture; later suture reduced
2 - FA fused with saucer & protruding (inHymenoptera & mecopteroids); later forms
only narrow rim on saucer, incoleopteroids+neuropteroids
2 - F J always fused with saucer

2 - sublobes slender, suture weak or absent

1 - triangular, massive
1 - BM invaded distally by broad fold,weakened

2 - elongate
0 - BM regular, well sclerotized, foldabsentPronotum: shape
Position of wings when flexed backward
over abdomen

1 - flat & horizontal
2 - dorsal & horizontal, flat over abdo­men, very slightly crossed apically (instem line blattoids); later wings overlap

Wing articulated with tergum at

1 - the anteriormost corner ofmetanotum (unique)1 - b y typically transformed muscula­ture shared by all Blattoneoptera

0 - other
1 - low roofwise, half open, anojugal lobe
not folded (in stem line hemipteroids: Paoliidae; later high roofwise or dorsal & hori­zontal or other, variously overlaped
0 - other

Other

Power for flight generated by regular setof muscles (as in Neoptera ground pattern)

0 - b y regular set of muscles

Ta b l e 6E. Character state distribution in Dermaptera and Dictyoptera.
Characters: states ofthe shared ancestor
are coded 0 o r l o r 2 o r 3 i n clades l 3 o r 6

Clade 13. Dermaptera (Figs 2, 4-10)

Flight, Folds and Folding
Remigium: length compared with anojugal l - remigium shorter than anojugal lobe,
lobe
very narrowl - present
Apical & quadruple wing folding
Partial anojugal lobe postero-distallyenlarged

500

3 - present, enlargement extra large

Clade 6. Dictyoptera (Figs 11-13)

0 - not shorter, regular width
0 - absent; later apical folding in derived
Blattodea
2 - present, enlargement rather modest (inIsoptera, Blattodea); later increased in Mantodea

Table 6E (continued).
Characters: states ofthe shared ancestor
are coded 0 o r l o r 2 o r 3 i n clades 13or6
Wing venation
Precostal strip (PC)
Anterior margin
Costal area (marginal area)
ScA+ between BSc and anterior margin
ScP
R, RA, RP, MA, MP
The stem of Cu basally at BCu
Arculus cross-vein brace (mp-cua or
m-cua or short fusion) (absent in Neoptera
ground pattern)
AA1+2 present

Clade 13. Dermaptera (Figs 2, 4-10)

Clade 6. Dictyoptera (Figs 11-13)

0 - PC thick, adjacent to C (PC expands
into anterior lobe)
3 - pliable
1- desclerotized, pliable
1 - ScA+ forms narrow, blunt, obliqueridge
1 - reduced
1 - veins simplified & condensed in
narrow remigium
1 - separated from BCu by a gap,
proximal end pointed
1 - present in both wing pairs

3 - PC fully fused with anterior wing
margin2 - firm
0 - regular
1 - same; later ridge broadened in Mantodea
0 - present
0 - not simplified, not condensed

3 - AA1+2 lost

AA3+4 forked

1 - AA3+4 forked into AA3 and AA4;AA3 short, AA4 fused with AP

AP1+2 & AP3+4 branches supporting par­tial anojugal lobe

3 - stabilized in derived pattern, AP1+2
8-branched, AP3 & AP4 uniquely
divided
2 - stabilized in derived branching pat­
tern

JA, JP branches

0 - Cu starts from BCu
0 - absent in both wing pairs (in blattoid
stem line & Isoptera); later formed only inhindwing, in Mantodea, again in Blattodea
1 - AA1+2 weak (in Isoptera & Blattodea);
later lost in Mantodea
2 - simple (in Blattodea); later bears secon­
dary pectinate twigs in blattoid stem line &Isoptera, or AA3+4 lost in Mantodea
2 - AP1+2 with secondary twigs inenlarged distal part of anojugal lobe,
AP3+4 regular
0 - JA, JP dichotomously branched

Articulation
PRSc & PRR, AWP

0 - PRSc, PRR separate, each articulated 1 - AWP present (suture between PRSc &with tergum
PRR retained)
PRR articulation regular (Neoptera ground 1 - PRR fused with 1Ax tail (unique
0 - PRR regular, articulated with 1Ax waist
pattern)adaptation)(AXR)
Sclerite PRCu (present in Neoptera ground 0 - PRCu present, long, pliable; later
0 - PRCu very weak (visible in basal Blat­
pattern)
membranized (like in most Pterygota)
todea only); later membranized, in parallel
0 - not massive
1Ax tail (PRM): regular
1 - massive
2Ax body (AXM) curved, proximal
4 - thickened & twisted proximal margin 3 - present, 2Ax body strongly resembles
margin thick, rest weakened & wrinkled
retained, the rest lostan ear
2Ax arm (FR) pliable, thin, with anterior
1 - arm long, lobe small
2 - arm short, lobe lostlobe
3Ax goblet (=proximal lobe of AXCu)
2 - goblet very long & slender0 - goblet short & robust
0 - heel and goblet broadly connected
3Ax goblet & heel (=distal lobe of AXCu) 1 - heel cut off from goblet
3Ax goblet versus saucer (AXAJ)
3Ax anal arm (FA),jugal arm (FJ): sepa­
rate (as in Neoptera ground pattern)
FM, FCu separate or fused into medialplate (FM + FCu)
BSc: sublobes BScA & BScP similar,
robust, separated by suture (as in Neoptera
ground pattern)
BR, shape

BM, proximal margin
BCu separate from BM

1 - saucer & goblet separated by mem­ 0 - goblet & saucer hinged togetherbrane
1 - fused into unique anojugal arm (FAJ) 0 - separate (in basal Blattodea only); laterbearing posteriorlyjugal prong
FJ reduced and FA broadened
1 - medial plate desclerotized
0 - FM, FCu separate (in Mantodea only);
later medial plate formed
0 - present (in Blattodea, Mantodea); later3 - BSc very long, thin, pliable, tiny
sublobes fused without suture in Isoptera
sublobes retained only on its proximal
tip
1 - BR subdivided by embayment into
0 - undisturbed, long (in Blattodea & Isop­
proximal & distal lobe
tera only); BR variously modified in Mantodea
1 - with broad, membranized embay­
0 - undisturbed
ment
0 - separate
0 - separate (in Mantodea only); later fused,suture retained in Blattodea, Isoptera, or lost501

Table 6E (continued).
Characters: states ofthe shared ancestor
are coded 0 o r l o r 2 o r 3 i n clades 13or6

Clade 13. Dermaptera (Figs 2, 4-10)

Clade 6. Dictyoptera (Figs 11-13)

BAA1+2, BAA3+4
BAP & BJ

2 - fused, form a unique triangular plate
1 - fused into narrow BAPJBAA, BAP, BJ

1 - BAA+BAP+BJ fused into uniquely
shaped anojugal plate BAJ

Anal brace: BAA1+2 articulated with BCu
at an angle (as in Neoptera ground pattern)

2 - anal brace different: BAA triangular,
overrides BCu, articulates by its distal
point with BM

0 - articulated
0 - BAP articulated with BJ (in Mantodea,
Blattodea only); later fused in Isoptera
0 - BAA, BAP, BJ articulated (inBlattodea); later fused into BAPJ in Isop­tera, or BJ reduced in Mantodea
1 - unmodified

Ta b l e 6F. Character state distribution in Blattodea + Isoptera and Mantodea.
Characters: states ofthe shared ancestor
are coded 0 o r 1 o r 2 o r 3 i n clades 7or 12

Clade 7. Blattodea + Isoptera (Figs 11,
13)

Clade 12. Mantodea (Fig. 12)

Claval fold reducing

2 - shortened

1 - long, shallow

Claval fold: reinforced by CuP, AA1+2
Folding of anojugal lobe: fan-like, slightly
irregular (along anal fold & other shorter
folds)

1 - claval fold flanked by AA1+2, which 2 - claval fold reinforced by CuP running
3 - present; later folding flat book-like in 3 - slightly irregular fan-likeblattoid stem line & Isoptera

Flight, Folds and Folding

1 - ridge narrow

2 - ridge broad, obtuse

ScP-

1 - short

The stem ofR (absent in Neoptera groundpattern)
Fusion ofRP+MA: entry ofMA discrete,visible in dissections, MA separates api­
cally from RP

1 - very short (in Isoptera only); later
often lengthened
0 - MA visibly separates from RP

2 - R relatively long

MP basal portion running close to radialstem

1 - faint (in some Isoptera); later not
visible in Blattodea

2 - partly visible, partly membranized by
the proximo-distal fold

Arculus, homoplastic flight importantcross-vein brace mp-cua (absent in Neoptera ground pattern)

0 - absent in both wing pairs (in blattoid
stem line & Isoptera only); later absent
in forewings but added in hindwings, inBlattodea

1 - arculus absent in forewings, added inhindwings (in parallel with Blattodea)

Boundary between distal end of Cu &base of CuA
CuP simple, runs along bottom of clavalfold

2 - boundary not discernible (veinsseems continuous)
2 - present (in blattoid stem line only;
later CuP lost in Isoptera, again in Blat­todea
1 - retained, simple, weak; later AA1+2
may descend into claval fold, in some
Blattodea
3 - area secondarily broadened, AA3+4
supported by short pectinate twigs (inblattoid stem line, Isoptera, some Blatto­dea)
1 - AP area secondarily enlarged, sup­ported by secondary twigs on AP1+2

1 - discernible (Cu can be clearly recognized from CuA)
2 - present, CuP runs along bottom ofclaval fold

2 - very weak
1 - fused into medial plate2 - goblet asymmetrical, its proximal
portion very robust & protruding

1 - regular
0 - FM, FCu separate
1 - goblet unmodified

AA1+2

AA3+4 simple, occupies narrow area

AP area enlarged

B

OQ

ScA+ near wing base forms oblique ridge01

Wing venation

1 - MA & RP branches homogenous, MA
separation discrete

3 - lost

2 - AA3+4 simple, area relatively narrow

2 - AP area strongly secondarily enlarged

Wing articulation
Humeral plate (HP)FM, FCu separate
3Ax goblet (proximal lobe of AXCu)

502

Table 6F (continued).
Characters: states ofthe shared ancestor
are coded 0 o r l o r 2 o r 3 i n clades 7 o r l 2

Clade 7. Blattodea + Isoptera(Figs 11,13)

Clade 12. Mantodea (Fig. 12)

3Ax anal arm (FA) &jugal arm (FJ) separate (as in Neoptera ground pattern)l - narrow FA & FJ present (in basal
Blattodea only); later FA broadened, FJ
lost, in derived Blattodea
0 - present (in basal Blattodea only);
later FJ lost in derived Blattodea, again
in Isoptera
0 - B S c & ScA not separated, regular

2 - FA+FJ connected by broad, sclerotizedmembrane, form extra wide FAJ arm whicharticulates with BAPl - F J fused with FA, too short to articulate
with BJ

3Axjugal arm (FJ) articulated with 3Ax
saucer & BJ (as in Neoptera ground pattern)
BSc & ScA regular
BM & BCu separate (as in Neoptera
ground pattern)l - BCu broadly adjacent to BM

BJ regular

0 - B J sclerotized, regularl - separated by deep fold which may con­tinue posteriorly & separate BR from R
0 - BCu slender, separate from BM (inMantoididae only); later adjacent in derived
Mantodeal - membranized

Ta b l e 6G. Character state distribution in Blattodea and Isoptera.
Characters: states ofthe shared ancestor
are coded 0 o r l o r 2 o r 3 i n clades l 0or
llClade 10. Blattodea (Fig. 11)

Clade 11. Isoptera (Fig. 13)

Folding in anojugal lobe

3 - slightly irregular, mainly along anal
fold

Broad fold near wing base, posterior &parallel with R
Fold running between RA and RP nearbase
Perpendicular fold between BM+BCu &M, Cul - present, invades distal part of BM

4 - lobe folds book-like flat under hindwing
so that small part overlaps anterior wing
margin (as in blattoid stem line)
0 - absent

0 - absent

l - present, destroys base of RP

0 - absent

l - present, separates bases o f M& Cu
from their basivenalia

Forewing: basal third
Forewing, costal area

0 - regular, not diminishedl - triangular (in Palaeozoic & modern
Blattodea)

The stem of R

3 - medium length, robust

RP baseBasal portion of MP before MP turns posteriorly towards wing margin (preservedin Mantodea)
Arculus, cross-vein brace mp-cua (absent
in Neoptera ground pattern)

0 - not disturbed
2 - obliterated by broad fold, MP reappears at arculusl - strongly diminished
0 - ribbon-like (in blattoid stem line); later
strogly reduced costal area in Isoptera may
be ribbon-likel - very short; later lengthened in blattoid
stem line
l - membranized by crossing fold
3 - cut off at BM by short fold, MP reappears distally as it turns posteriorly

Flight, Folds and Folding

Wing venation

The stem of Cu
AAl+2 simple, reducing
AA3+4, in narrow AA area associatedwith remigium

2 - absent in forewing, present inhindwing (in Palaeozoic & living Blattodea)
3 - short

0 - absent in both wing pairs (in blattoid
stem line & Isoptera)l - flanks closely claval fold, may
descend into its bottom
2 - AA3+4 simple, flanked & reinforced
by strong, convex intercalary veinl - flanks closely claval fold

2 - very shortl - AA3+4 supported by serial, short &broad secondary twigs

Wing articulation
Humeral plate
PRCu (in membranized area into which
3Ax colapses)
Medial plate
3Ax goblet, proximal portion enlarged

2 - weakl - faintly preserved (in basal Blattodea
only); later membranizedl - present, unmodified
2 - broader than distal portion, pro­truding

3 - lost
2 - membranized
2 - distal margin indented
3 - massive, highly protruding503

Table 6G (continued).
Characters: states ofthe shared ancestor
are coded 0 o r 1 o r 2 o r 3 i n clades 10or
11

Clade 10. Blattodea (Fig. 11)

Clade 11. Isoptera (Fig. 13)

3Ax anal arm FA: strong
FCu articulated with

1 - strong, narrow
1 - with 3Ax heel & BCu

3Axjugal arm FJ

0 - present (in basal Blattodea only);
later repeatedly lost2 - faintly preserved
1 - present, well expressed
1 - fused, suture preserved
1 - slender, convex
0 - articulated

3 - strong, broad, massive
2 - with heel & BCu & FA (articulation
with FA added)
2 - reduced or lost

Humeral plate
BSc, sublobes BScA, BScP
BM+BCu fusedAnojugal basivenalia
BAP, BJ

3 - lost
4 - fused into uniform BSc
2 - fused, suture lost2 - uniquely broad & flat
1 - broadly fused

Ta b l e 6H. Character state distribution in Hemineoptera and Endoneoptera.
Characters: states ofthe shared ancestor
are coded 0 o r 1 o r 2 i n clades 14or15

Clade 14. Hemineoptera (Figs 16, 22)

Clade 15. Endoneoptera (Figs 17, 22)(emendation* of previous interpretations at
the end)Claval fold with tendency to reduce

1 - long, shallow

Claval fold reinforced by

1 - flanked by CuP (in stem line hemipteroids & Fulgoromorpha only); later
CuP lost, in parallel
3 - folds irregular, may cross anal
branches (in Fulgoromorpha); later new
folds added, or folds & lobe reduced

1 - long (in Hymenoptera); later shortened,shallow, in parallel
2 - flanked by AA1+2 (in Hymenoptera
only); reinforcement lost in coleopteroids +neuropteroids + mecopteroids
3 - claval, anal,jugal fold active (in Hymenotera only); later lobe & claval foldreduced many times, in Coleoptera anal
fold folded book-like & new short folds
added
0 - coupling absent; later occurs repeatedly,in Hymenoptera, Mecoptera, Trichoptera,
Lepidoptera
3 - held mildly roofwise, slightly overlaped(in neuropteroids); later repeatedly roof­
wise, or dorsally flat, or slightly to fully
overlaped, or held vertically (in Lepidoptera)

Flight, Folds and Folding

Partial anojugal lobe, folds

Hindwings with tendency to diminish,couple with forewings in flight
Wing position at rest probably mildly
roofwise & not fully flexed backward

0 - coupling absent; later occurs repeat­
edly, in most Sternorrhyncha, Heteroptera, Fulgoromorpha, Cicadomorpha
0 - wings held mildly roofwise not over­lapped (in hemipteroid stem line: Paoliidae); later highly roof-wise, or dorsallyflat, variously overlaping

Wing venation
Forewing, epipleuron (PC strip when
extended) directed
Forewing, CA+, CP- long separate, run inparallel (as in protowing)

1 - ventro-posteriorly, obliquely underanterior wing margin (in Hemiptera)
0 - present (in hemipteroid stem line,
some Hemiptera); later CA, CP repeat­
edly fused into C near base2 - mainly ventrally (in Coleoptera)

Humeral vein (cross-vein brace c-scp)

0 - absent; later weakly indicated insome forewings, independently in par­allel
2 - bulge low; later lost, replaced bymembranized fissure at base of costalarea
1 - clavus present
1 - ScP shortened: later ScP divesbasally under R (in Heteroptera, Fulgoro­morpha, Cicadomorpha)
0 - absent (in hemipteroid stem line
only); later repeatedly formed

1 - present, formed distally of ScA bulge,
prominent in both wing pairs

Forewing, ScA+ forms sclerotized convex
bulge
Forewing: anal lobe forms pointed clavus
ScP with tendency to shorten, basallyclose to R
RA, RP separate, the stem ofR absent (asin protowing)504

0 - present (in the elytra of Coleoptera);later repeatedly fused into C at wing base

3 - bulge high, prominent, reinforced dis­tally by humeral vein; later reduced in
narrow costal areas
0 - absent
0 - ScP and R basally close but separate,
ScP relatively long
1 - present, extremely short (in Coleoptera+ Strepsiptera only); later repeatedly
lengthened

Table 6H (continued).
Characters: states ofthe shared ancestor
are coded 0 o r l o r 2 i n clades 14or l 5

Clade 14. Hemineoptera (Figs 16, 22)

Clade 15. Endoneoptera (Figs 17, 22)(emendation* of previous interpretations at
the end)MA fused with RP at base, separates
again apically, branches

3 - separation obscured, RP & MA
branches look alike, cannot be distin­
guished one from another

Cross-vein brace rpma-mp expressed

2 - far from wing base, as cross-veinbrace or short fusion between RP+MA
and MP

MP after rising from BM turns towards
posterior margin (as in protowing &Neoptera ground pattern)

2 - MP first approaches R+MA, thenturns posteriorly towards wing margin

MP basal portion

0 - distinct in large basal members; later
often very close t o R & almost invisible(evident in dissections under water)

Arculus, cross-vein brace mp-cua or shortfusion.

1 - always present (frequently homo­plastic outside Hemineoptera +Endoneoptera)
0 - absent

2 - evident, MA convexly fluted (+) (inhymenopteroid stem line, basal Hymenoptera, Neuroptera); later MA cannot berecognized
1 - near base, as long sinusoid brace (inNeuroptera, Megaloptera, medial bridge in
Coleoptera + Strepsiptera); later shorter &shifted distally twice, in Hymenoptera,again in mecopteroids
1 - MP turns posteriorly (in Hymenoptera
only); later MP first approaches RP+MA,
then turns posteriorly, in coleopteroids +neuropteroids + mecopteroids
0 - distinct (only in Hymenoptera); later
visible in some Coleoptera, in neuropter­oids deep under R, reduced in mecopteroids(evident in dissections under water)
1 - always present (in coleopteroids + neu­
ropteroids + mecopteroids); later extended
into long fusion in Hymenoptera
1 - present (in neuropteroids obscured);later arculus and posterior arculus form one
continuous fused brace in Hymenoptera
2 - present, very short (in Hymenoptera,neuropteroids); later much longer in cole­
opteroids, again in mecopteroids
0 - width regular: the stem of Cu does not
widen or sclerotize
3 - CuP weakly branched (inneuropteroids); later simple in Coleoptera,again in mecopteroids, or lost in Hymeno­
ptera, or simple or enriched by twigs in
Neuroptera
2 - AA1+2 simple, strong, functional (inHymenoptera only); later AA1+2 lost inneuropteroids + coleopteroids + mecopteroids
1 - AA3+4 branched (in Neuroptera +Coleoptera only); later simple in mecopter­
oids, independently also in Hymenoptera
2 - present (in basal Hymenoptera, neuro­
pteroids, coleopteroids, Trichoptera); later
lobe reducing

Posterior arculus, fused brace between
branches MP4-CuAl
The stem of Cu

2 - present, very short

Forewing: the stem of Cu, width

0 - width regular; later widens into
pointed sclerotized plate (in Hemiptera)
0 - CuP with long, richly divided
branches (in hemipteroid stem line &Fulgoromorpha); later CuP simple orlost, many times in parallelCuP dichotomously branched (as in pro­towing & Neoptera ground pattern

AA1+2 (in reducing AA area adjacent toremigium)

3 - AAl+2 completely reduced, lost

AA3+4 branches (in reducing AA areaadjacent to remigium)

3 - AA3+4 simple, branches lost

Partial anojugal lobe relatively large, APsparsely branched

2 - present (in Fulgoromorpha); later
lobe repeatedly strongly diminished

Articulation
Forewing, humeral plate & BSc separate
PRSc, PRR separate (as in Neoptera
ground pattern)l - connected by conspicuous, broadsclerotizationl - fused with tergum into small AWPPRAJ (hinged with tergum = 4Ax; fusedwith tergum = PWP)l - PRAJ modified, eroded; later
desclerotized, or PWP formed, or PWP
desclerotized

PRCu (sclerite in membranous window)

1 - present, long, pliable (in large
aphids); latermembranized
2 - head & neck strongly reduced

1Ax

0 - separate
0 - PRSc, PRR separate, hinged with
tergum (in basal Neuroptera only); AWP
repeatedly formed
0 - 4Ax present (in Hymenoptera & Cole­
optera: Gyrinidae only); later PWP in neu­
ropteroids, Coleoptera, again in
mecopteroids
2 - present, fragmented (in some Neuro­
ptera only); later lost
1 - unmodified

505

Table 6H (continued).
Characters: states ofthe shared ancestor
are coded 0 o r l o r 2 i n clades 14or l 5

Clade 14. Hemineoptera (Figs 16, 22)

Clade 15. Endoneoptera (Figs 17, 22)(emendation* ofprevious interpretations at
the end)

Forewing, 2Ax body distally weakened

1 - body distally enlarged, lobate; latersclerotized, lobes articulated with FR &FM (in Heteroptera, Fulgoromorpha,
Cicadomorpha)
6 - 2Ax body bears proximally thick,strongly curved rim, distal part veryweak
2 - FR lobed twice, towards BSc &towards medial plate
0 - FM & FCu close but separate, notmodified; later fused in medial plate,with or without dividing suture

0 - present; secondary enlargement &sclerotization of 2Ax body absent

2Ax body (AXM) proximally stronger,
distally weaker
2Ax arm (FR) broad, lobed towards BSc
FM, FCu separate (as in Neoptera groundpattern)

3Ax goblet & heel (two lobes of AXCu)

3Ax saucer (AXAJ) proximally longer
than goblet
anal arm FA fused with saucer, protrudes
to articulate with BAA3+4

1 - heel small, separated from goblet;
later enlarged in Heteroptera & Fulgoromorpha
1 - the proximally protruding part ofsaucer reduced
1 - present, FA forms strongly sclero­
tized ridge protruding from saucer &articulated with BAA3+4

2 - 2Ax body forms distorted, distally
weakened triangle
1 - FR lobed only towards BSc
1 - separate, modified (lost inHymenoptera; FM subdivided, FCu fusedwith 3Ax in Coleoptera + Strepsiptera);later medial plate formed in neuropteroids,
reduced in mecopteroids
0 - heel small, connected to goblet

FJ: fused with saucer, articulated with BJ

1 - articulation with BJ weak; later lost

BR & BM separate (as in Neoptera
ground pattern)
BM & BCu separate (as in Neoptera
ground pattern)

1 - broadly adjacent

0 - saucer strongly protruding proximally;
later it repeatedly evolved secondary hinge
1 - present (in Hymenoptera, meco­
pteroids); later FA blends into narrow rim
on saucer which is connected with basivenalia by narrow strips of tough membrane(in coleopteroids + neuropteroids)
2 - direct contact with BJ lost; later sclerites connected by strips of toughened mem­brane in coleopteroids + neuropteroids
0 - separate

0 - BCu, BM separate

1 - BM hinged/fused with BCu* Emendation. In Table 6H, important improvements in homologization in the hindwing of Coleoptera (Kukalová-Peck & Law­
rence 1993, Fig. 12) are proposed as follows. MAjoins R immediately at wing base and MA entry is membranized; medial bridge isa structurally important cross-vein brace rpma-mp, involved in apical folding; cubital stem is present, long; CuA forks into CuA1+2(marked as “CuA”) and CuA3+4 (marked as “CuA2”); CuP (marked as “CuA3+4”) is present; AA1+2 is lost; AA3+4 forks into
AA3 (marked as “AA1+2”) which is braced with CuP, and into AA4 (marked as “AA3+4”); arculus is mp-cu (marked as“mp-cua”); the veinal fusion near the posterior margin is MP4 + CuA1+2 (marked as “MP4 + CuA1”). The changes will be dis­cussed and documented in soon to be published paper by the same authors.
Other improvements of full homologizations in Neoptera previously offered by Kukalová-Peck (1991 and elsewhere), discussed
here and introduced in Table 6, are shown in figures of the representatives of living superordinal lines. Broader documentation anddiscussion will be presented later, when the pterygote superorders will be dealt with individually (in advanced preparation by JKP).

The data set is focused on interpreting the phylogenetic
position of Dermaptera within the Pterygota. Orders
within Palaeoptera and outside of Blattoneoptera are notfully characterized here. For that reason, the character
table is not complete enough for a cladistic analysis of
Neoptera. It is also open to further improvements as
research on the ordinal ground patterns proceeds.
Unless the forewing is specifically mentioned, all char­acters concern only the hindwing.
As customary, the numeral 0 is plesiomorphic; 1through 6 represent unordered apomorphic states.
Protowing ground pattern (Kukalova-Peck, 1978,
1983, 1985, 1991; Wootton & Kukalova-Peck, 2000). In
the ancestral protowing ground pattern hypothesis there506

are 8 principal veins (PC, C, Sc, R, M, Cu, A, J). All
except PC (precostal strip) are preserved as two corru­
gated sectors, anterior (A) convex (+) and posterior (P)concave (-). All sectors except CA (forming costal mar­gin) are dichotomously branched about 3 times. Articularsclerites are aligned with their veins in 8 rows (PC, C, Sc,
R, M, Cu, A, J), and are arranged in 4 columns (PR, AX,
F, B).
The pterygote protowing is almost perfectly retained in
the prothoracic wings of Palaeodictyoptera and extinct
Odonatoptera (Geroptera). It is supported by the costal
margin (a fusion of PC strip and CA) and by radiallyarranged veins. Each principal veins composed of twoveinal sectors: anterior - convex and posterior - concave.

Veinal sectors are dichotomously branched, mildly corru­
gated, and connected by numerous, irregular cross-veins.
The prothoracic protowing is slightly asymmetrical, withveins crowded anteriorly towards the costal margin.
Articular sclerites are present, but are fused into a com­posite, arched welt. Secondary reductions include dimin­
ished branching. These features show that prothoracic
protowings were previously adapted for flapping forward
movement of some sort, but became gradually less active
and were in the process of reduction.
However, powered aerial flight was definitely not afunction of protowings. This requires a much larger size;
denser anterior crowding of veins; the presence of an anal
brace; fusions of veins into veinal stems or braces near
the wing base; a twist-promoting wing articulation; and a
specialized and well-coordinated thoracic and limb
musculature, manipulating wings by pulling on articularsclerites. These adaptations took place in Pterygota in two
profoundly different styles, which are here recognized as
the pterygote clades Neoptera and Palaeoptera(Kukalova-Peck, 1978, 1983, 1991, 1997).
Pterygota and monophyly. Pterygota are monophyletic. We know of no evidence to the contrary.
Consequently, all characters in all orders must be
interpretable from a single protowing ground pattern ofvenation and articulation. Only one nomenclature must be
used for all orders to obtain valid answers about their
phylogenetic relatioship. It is very well documented that
all representatives, extinct and extant, always share an
identical ground pattern of stabilized order-level character
states. As a clade evolves, new transformation (mainly
reductions, fusions and extensions) will be added in thesubtaxa (suborders, families, genera and species). Thesemay obscure, but will not change, the ground pattern of
their order. Consequently, the ordinal character states are
predictably present, can be recognized and assembled in aground pattern. They are those characters, which areshared by all representatives of the order. The stability of
the wing complex means that the principal veins andarticular sclerites, which in ground pattern are fused
together in stems and composite sclerites, or lost, neverunfuse or reappear in any extinct or extant representativesof an order. Fusions, reductions or losses will gradually
accumulate, but not oscillate, because there is noevidence, ever, of reversals at the ordinal level (seenumerous examples below).
In the wing character complex, the ground pattern canbe convincingly presented only if compared with an ex­group: In Neoptera, with Palaeoptera; in Blattoneoptera,with other superordinal lines; in Dermaptera, with otherblattoid orders, etc. In Pterygota, the task of full homologization is multi-layered and almost overwhelming but
nevertheless absolutely necessary, because it provides theway to recognize seemingly “unrecognizable” homoplasies. As a fact, almost all (!) higher-level characters are
multi-homoplasious, simultaneously in related and unre­
lated orders and/or in their subtaxa. The only way to
avoid chaos is to use each time, in the Pterygote super­orders and orders, only the fully homologized ground pat-tems. To let synapomorphies stand out, the order-level
characters must be handled strictly within the context inwhich they occur. Table 6 presents many self-explanatory
examples of previously “unrecognizable” homoplasies,
which can be recognized instantly in a proper context.
These are mentioned after the word “later”. They show
the range of homoplastic diversifications, and the aston­ishing quantity of homoplasies in unrelated taxa, whichmust be excluded from cladistic analysis.
It is important to note that all character states in the
Table 6 are at the ground pattern level of the clade. Anylater transformations are irrelevant to phylogenetic con­sideration and must be excluded from consideration. The
character table deals with an exceptionally rich andunevenly changing character complex. We have tried topresent the characters in the most concise and manageable
way possible, while also pointing out homoplasies. The
goal is to document convincingly the theoretical thesis of
this paper, namely that any phylogenetic analysis of anorder using morphological character complexes must
include homologization of all orders of the respective
higher taxon, all the way back to the shared ancestor. In
the wing complex, it is the protowing of the ancestralpterygote. In a limb-derived appendage, it is the limb of
the ancestral Proterozoic arthropod. We expect that infuture wing character tables it will not be necessary todeal so much with homoplasies once this current problem
is out of the way.
ACKNOWLEDGEMENTS. We are very obliged to Dr.

DanielOtte, Academy of Natural Sciences in Philadelphia for his sup­port and permission to work with rare Dermaptera, Blattodea
and Mantodea in the outstanding ANSP collection. Mr. Don
Azuma and Dr. Jason Weintraub, also of ANSP, provided gen­erous help while we were studying the specimens.
Prof. S. Sakai, Daito Bunka University, Tokyo, Japan, Dr.
K.D. Klass, Copenhagen, Denmark, Dr. R.D. Cave, Escuela
Agricola Panamericana, El Zamorano, Honduras, and Mr. D.
Matzke, Leipzig, Germany, supplied some important specimens
used in this study. Our friend Joyce Cook ably assisted JKP withcomputing problems. We are much obliged to Dr. K.D. Klass
for collaboration on the dissections of pterygote wing base. Dr.
S.B. Peck, Carleton University Ottawa, Canada, Prof. D.
Waloßek and Dr. W. Ahlrichs, both of University of Ulm, Ger­
many helped by critically reading the manuscript.
Parts of this paper were researched in Tübingen and Jena, Ger­
many and Exeter, United Kingdom. FH would like to thank Dr.
R.J. Wootton, Biological Sciences Department, University of
Exeter, Prof. W. Maier, Lehrstuhl für Spezielle Zoologie, Uni­
versity of Tübingen, Prof. M.S. Fischer, Institut für Spezielle
Zoologie und Evolutionsbiologie, and Prof. R. Blickhan, Institut
für Sportwissenschaft, both of Friedrich-Schiller-Universität
Jena, for their support.
This paper was partially supported by an individual research
grant to JKP from the Natural Sciences and Engineering
Research Council in Canada.
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R eceivedA pril 12, 2 0 0 1 ;re v ise d A u g u st 3, 2001; accepted O ctober 1, 2001

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