Cladotypic taxonomy applied: titanopterans are orthopterans

Abstract

Received 03.ix.2007, accepted 12.xi.2007. Published online at www.arthropod-systematics.de on 7.xii.2007. > Abstract The Linnaean taxon Titanoptera is a distinctive Triassic insect order the origin of which is uncertain. Forewing venation patterns of the Permian Linnaean subfamily Tcholmanvissiinae (Orthoptera) and of the Titanoptera are re-investigated. The comparative analysis supports the view that the morphology of the latter group is derived from that of the former. As a consequence, the order Titanoptera is to be included within the subfamily Tcholmanvissiinae. A cladotypic taxonomy is developed in order to avoid the confusion inherent to taxonomic rearrangements associated with rank-based taxonomy. The following hierarchy is proposed: (Archaeorthoptera nom. Béthoux & Nel, 2002a, dis.-typ.n. (Pantcholmanvissiida nom. n., dis. Béthoux & Nel, 2002b, typ.n. (Tcholmanvissiidae nom. Zalessky, 1934, dis. Sharov, 1968, typ.n. (Tcholmantitano-pterida nom.-dis.-typ.n. (Tcholmanvissiella nom. Gorochov, 1987, dis.-typ.n. (Titanopterida nom.-dis.-typ.n. (Gigatitanidae nom. Sharov, 1968, dis. -typ.n.))))))). This fi rst application of cladotypic taxonomy unveiled several practical aspects of this system. A system governing the adaptation of pre-occupied taxon names is developed based on various cases of character state formulations; the issue of the occurrence of Linnaean suffi xes and of the preservation of Linnaean binominals within a cladotypic taxonomy are discussed; the capacity to handle the ancestor 'species' vs. apomorphy-less sister-species issue by the various nomenclatural systems is discussed.

Full text

135Arthropod Systematics & Phylogeny
65
(
2
)
135 – 156 © Museum für Tierkunde Dresden, ISSN 1863-7221
Cladotypic Taxonomy Applied: Titanopterans are Orthopterans
OLIVIER BÉTHOUX
State Natural History Collections of Dresden, Museum of Zoology,
Königsbrücker Landstrasse 159, D-01109 Dresden, Germany
[obethoux@yahoo.fr & olivier.bethoux@snsd.smwk.sachsen.de]
Received 03.ix.2007, accepted 12.xi.2007.
Published online at www.arthropod-systematics.de on 7.xii.2007.
> Abstract
The Linnaean taxon Titanoptera is a distinctive Triassic insect order the origin of which is uncertain. Forewing venation
patterns of the Permian Linnaean subfamily Tcholmanvissiinae (Orthoptera) and of the Titanoptera are re-investigated.
The comparative analysis supports the view that the morphology of the latter group is derived from that of the former. As
a consequence, the order Titanoptera is to be included within the subfamily Tcholmanvissiinae. A cladotypic taxonomy is
developed in order to avoid the confusion inherent to taxonomic rearrangements associated with rank-based taxonomy. The
following hierarchy is proposed: (Archaeorthoptera nom. Béthoux & Nel, 2002a, dis.-typ.n. (Pantcholmanvissiida nom.
n., dis. Béthoux & Nel, 2002b, typ.n. (Tcholmanvissiidae nom. Zalessky, 1934, dis. Sharov, 1968, typ.n. (Tcholmantitano-
pterida nom.-dis.-typ.n. (Tcholmanvissiella nom. Gorochov, 1987, dis.-typ.n. (Titanopterida nom.-dis.-typ.n. (Gigatitanidae
nom. Sharov, 1968, dis. -typ.n.))))))). This fi rst application of cladotypic taxonomy unveiled several practical aspects of this
system. A system governing the adaptation of pre-occupied taxon names is developed based on various cases of character
state formulations; the issue of the occurrence of Linnaean suffi xes and of the preservation of Linnaean binominals within a
cladotypic taxonomy are discussed; the capacity to handle the ancestor ‘species’ vs. apomorphy-less sister-species issue by
the various nomenclatural systems is discussed.
> Key words
Pterygota, Archaeorthoptera, Orthoptera, Titanoptera, Titanopterida, cladotypic taxonomy, adaptation, priority, ancestor
species.
1. Introduction
Attempts to exhaustively inform the taxonomic posi-
tion of fossil stem groups would necessitate a surfeit
of ranks if strictly following the traditional Linnae-
an rank-based nomenclatural system. Phylogenetic
(CANTINO & DE Q UEIROZ 2006), topology-based (SERENO
2005), and cladotypic (BÉTHOUX 2007d, e) taxonomic
systems, which are all rank-less, have the advantage of
avoiding this pitfall, and avoiding the need for modi-
fi cation of taxa names if a hierarchical re-arrangement
is necessary. Therefore their use might result into more
stable taxonomies. Among the alternative systems, the
cladotypic approach is likely to be the most effi cient,
because it relies on assumptions that are more easily
falsifi able than are those involved in other rank-less
approaches. Moreover it is fully operative as rules are
provided for the species case, unlike other alternative
systems. Herein, I apply this new system to a case in-
volving fossil taxa nested within a group having mod-
ern representatives.
I will focus on the resolution of relationships of the
Linnaean order Titanoptera Sharov, 1968 (thereafter
informally referred to as titanopterans) with respect
to the Linnaean order Orthoptera Olivier, 1789 (there-
after informally referred to as orthopterans). Several
hypotheses on the origin of the very distinctive ti-
tanopterans were proposed. The Upper Carbonifer-
ous Linnaean family Geraridae Scudder, 1885, which
is currently viewed as a close relative of orthopterans
(SHAROV 1968, 1971; Gorochov 2001; BÉTHOUX & NEL
2003; in prep.), was proposed as sister-group of titan-
opterans by GOROCHOV (2001), followed by BÉTHOUX
(2005a: 405). On the other hand SHAROV (1968, 1971)
considered that titanopterans diverged from the Lin-
naean family Tcholmanvissiidae (orthopterans repre-
sented during the Permian), which he viewed as a par-
aphyletic group. This author considered geraridaeans
to be the only representatives of the Linnaean order
Protorthoptera, itself understood as paraphyletic and
‘ancestral’ to the orthopterans.
My investigations of some taxa considered by
SHAROV (1968) as geraridaeans (BÉTHOUX & NEL 2003;
in prep.) and of representatives of the family Tchol-

BÉTHOUX: Titanopterans are orthopterans
136
manvissiidae (BÉTHOUX & NEL 2002b and herein) lead
me to propose a new interpretation of the titanopterid
forewing venation, presented herein. This interpreta-
tion implies that the order Titanoptera is not directly
related to the family Geraridae, as I argued previously,
but to the Permian family Tcholmanvissiidae. This
situation implies taxonomic re-arrangement.
2. Material and Methods
Specimens referred to as PIN are housed at the Pal-
aeontological Institute of the Russian Academy of
Science (Moscow, Russia). The specimen referred to
as AM is housed at the Australian Museum (Sydney,
Australia). Specimens referred to as NHM are housed
at the Natural History Museum (London, UK). The
specimen referred to as FG is housed at the Depart-
ment of Palaeontology, Freiberg University of Mining
and Technology (Freiberg, Germany).
I use the wing venation nomenclature elaborated
by BÉTHOUX & NEL (2002a) for Archaeorthoptera (see
taxon defi nition in the systematic section), itself based
on that of orthopterans (BÉTHOUX & NEL 2001). Cor-
responding abbreviations are repeated herein for con-
venience: ScA, anterior Subcosta; ScP, posterior Sub-
costa; R, Radius; RA, anterior Radius; RP, posterior
Radius; M, Media; MA, anterior Media; MP, posterior
Media; Cu, Cubitus; CuA, anterior Cubitus; CuP, pos-
terior Cubitus; CuPa, anterior branch of CuP; CuPaα,
anterior branch of CuPa; CuPaβ, posterior branch of
CuPa; CuPb, posterior branch of CuP; AA1: fi rst anal.
The reader who is not familiar with orthopteran and
other insect wing venation nomenclature could refer
to the discussion in BÉTHOUX (2005b; and references
therein) and to BÉTHOUX & NEL (2002a: fi g. 1b). Crit-
ics expressed by Gorochov (2005) regarding this ho-
mologization hypothesis are addressed in BÉTHOUX
(2007a). Subsequent comments by RASNITSYN (2007)
are addressed in BÉTHOUX (in press).
It will be demonstrated elsewhere that CuA is sim-
ple in forewings of orthopterans and of some stem-
orthopterans. In other words, all branches of CuA +
CuPaα as understood by BÉTHOUX & NEL (2002a)
belong to CuPaα, except for the most apical branch,
which is composed of CuA and the ultimate branch
of CuPaα. This homologization is applied herein. In
order to make the comparative discussion easier to fol-
low, I use the following vein abbreviations: CuPaα°
(indicated by ° in Fig. 1) refers to the anterior branch
of CuPaα resulting from the second branching of this
vein; CuPaα* (indicated by * in Fig. 1) refers to the
posterior branch of CuPaα resulting from the second
branching of this vein; CuPaα• (indicated by • in Fig.
1) refers to the posterior branch of CuPaα resulting
from the fi rst branching of this vein.
The restoration provided in Fig. 1C is primarily
based on a high-resolution photograph of the speci-
men AM F.36274. It was complemented by drawings
drawn with a stereomicroscope and camera lucida of
the specimens NHM In. 37340, NHM In. 37341, and
NHM In. 37342 (Fig. 2A–C, respectively), belonging
to the same species. The shape of the area between the
anterior wing margin and ScA is unknown in this spe-
cies and is inferred from related taxa. The restoration
provided in Fig. 1D is based on the restoration of SHA-
ROV (1968: fi g. 52B), but is skewed by 16° in order to
present a more plausible shape of the forewing (corre-
sponding fossils were deformed during or after fossili-
sation; SHAROV 1968; RASNITSYN 1982). In other cases
venation patterns and vein widths were drawn with a
stereomicroscope and camera lucida direct from the
fossil surface, both dry and under ethanol (except for
material from Madygen, Russia, that could be dam-
aged by ethanol immersion). Drawings were readjust-
ed on photographs using image-editing software.
In the systematic section, I use the cladotypic taxo-
nomic system elaborated by BÉTHOUX (2007d, e) for
taxa other than species, and follow the suggestions of
DAYRAT et al. (2004; and references therein) for spe-
cies names. The use of the suffi x ‘Pan’ is not related to
the rules of the PhyloCode governing the use of ‘pan-
clades’ (or panclade names; CANTINO & DE QUEIROZ
2006; see also JOYCE et al. 2004). Throughout this con-
tribution, taxa understood as taken from the Linnaean
system are indicated by the mention of their rank.
3. Results
3.1. Comparative morphological analysis
SHAROV (1968) proposed a homologization of the wing
venation of titanopterans that has been followed by all
subsequent authors (CARPENTER 1992; GOROCHOV 1995,
2003). BÉTHOUX & NEL (2002a) proposed to ‘translate’
Sharov’s nomenclature into an alternative one, intend-
ed to allow the wing venation of orthopterans to be
compared to that of other winged insects. However,
the authors agreed with Sharov’s interpretation of ti-
tanopteran wing venation pattern with respect to that
known in orthopterans. Basically, between the veins
CuA + CuPaα (Sharov’s MP + CuA1) and AA1 (1A),
two concave veins occur; as in orthopterans these are
likely to be CuPaβ (CuA2) and CuPb (CuP). This is the
most parsimonious interpretation if one refers only to
the data accessible to Sharov.
This homologization is now challenged by my in-
terpretation of the forewing venation of beybienkoi
Sharov, 1968 (orthopteran assigned to the genus Ju-
bilaeus Sharov, 1968; Fig. 1A), gigantea Gorochov,

137
Arthropod Systematics & Phylogeny 65 (2)
1987 (orthopteran assigned to the genus Tcholmanvis-
siella Gorochov, 1987; Fig. 1B), and of titanoperans
(Fig. 1C–D). Together with some other species, the
two former species were assigned to the family Tchol-
Fig. 1. Forewing venation homologies in Pantcholmanvissiida nom.n., dis. Béthoux & Nel, 2002b, typ.n.; orange, CuA vein; purple,
CuPa vein; blue, CuPaα vein; red, CuPaβ vein; green, CuPb vein (see text for abbreviations); A: beybienkoi Sharov, 1968 (from
BÉTHOUX & NEL 2002b); B: gigantea Gorochov, 1987 (from BÉTHOUX & NEL 2002b); C: giganteus Tillyard, 1916 (restoration; see
text); D: extensus Sharov, 1968 (modifi ed from SHAROV 1968: fi g. 52B).
A
B
C
D

BÉTHOUX: Titanopterans are orthopterans
138
manvissiidae by GOROCHOV (1987) and BÉTHOUX & NEL
(2002b) (thereafter informally referred to as ‘tchol-
manvissiidaeans’). As do other tcholmanvissiidaeans,
beybienkoi and gigantea exhibit one or several poste-
rior branches of CuPaα occurring basal to the connec-
tion with CuA (BÉTHOUX & NEL 2002b). This is a strict
apomorphic character state within Neoptera. The main
difference between gigantea and other tcholmanvissi-
idaeans relies on the apparent occurrence of branches
of CuPaα* (it is simple in other tcholmanvissiidaeans).
Additionally, gigantea exhibits another important dif-
ference in that CuPaα° is apparently simple for a long
distance and emits few distal branches. However, an
important point was overlooked: apparent branches of
CuPaα* occur opposite the section of CuPaα° that is
apparently simple; and CuPaα° is apparently branched
distally to the last apparent fork of CuPaα*. In other
words, apparent branches of CuPaα° and CuPaα* do
not co-occur at the same ‘level’. Therefore I argue
that proximal branches of CuPaα°, as they occur in
beybienkoi, are homologous to branches occurring on
CuPaα* in gigantea (see double-headed arrows on
Fig. 1B). In other words, several branches of CuPaα°
were ‘translocated’ onto CuPaα* in gigantea (imply-
ing that CuPaα* is actually simple). Translocation can
be defi ned as the fusion of a vein (sector / branch) with
another from the origin of the latter, so that there is no
visible basal free part of the translocated vein. Such
translocations frequently occur as irregularities of the
wing venation pattern, as it can be seen in the anal
area of the forewing of the specimen AM F.36274 (on
which is based the restoration of the corresponding
part on Fig. 1C; see arrows on this fi gure), and in the
branching pattern of CuPaα° in the restoration given
on Fig. 1D (the fi rst posterior branch of CuPaα° is
translocated onto CuPaα*). I observed a similar trans-
location affecting CuA branches in forewings of sev-
eral mantodean taxa (occurring as an intra-individual
polymorphism; pers. obs). As observed in gigantea,
the translocation of several branches is the mere result
of multiple single vein translocations.
I propose to characterize the organization of CuA
and CuPaα exhibited by gigantea as ‘in forewing, at
least one proximal branch of CuPaα° is translocated
onto CuPaα*’ (provided that CuPaα is branched). As
defi ned, it applies to titanopterans (Fig. 1C–D; SHAROV
1968; GOROCHOV 2003; GRIMALDI & ENGEL 2005). In
the specimen AM F.36274, the point of divergence of
the fi rst posterior branch of CuPaα° emitted from CuA
+ CuPaα° (thereafter referred to as CuA + CuPaα°
in part) is located basal to the last fork of branches
CuPaα° translocated onto CuPaα* (thereafter referred
to as CuPaα° trans.), unlike in the conspecifi c speci-
mens NHM In. 37340 (Fig. 2A) and NHM In. 37342
(Fig. 2C), where the point of divergence of the fi rst
posterior branch of CuPaα° in part is located op-
posite to the last fork of CuPaα° trans. Therefore, I
assume that the condition exhibited by the specimen
AM F.36274 is due to an infra-specifi c variation. In
some taxa from Madygen (Russia) described by SHA-
ROV (1968, 1971) and GOROCHOV (2003), the point of
divergence of the fi rst posterior branch of CuPaα° in
part is located opposite to the last fork of CuPaα°
trans. seemingly ‘overlap’, but this could be due to a
skewing post-depositional deformation (deformation
during compaction and/or tectonic deformation af-
fected specimens from Madygen; SHAROV 1968; RAS-
NITSYN 1982). In conclusion I argue that the apparent
branches of CuPaα* as exhibited by titanopterans are
homologous to the proximal branches of CuPaα° as
exhibited by beybienkoi.
Once the possibility that vein branches could trans-
locate onto a surrounding vein is admitted, it can be as-
sumed that the vein designated as CuPaβ in BÉTHOUX &
NEL (2002b: fi gs. 10, 11) and considered as branched
in beybienkoi and gigantea is merely composed of a
simple CuPaβ fused with the fi rst posterior branch of
CuPaα (CuPaα•). In noinskii Zalessky, 1929 and lon-
gipes Martynov, 1940 (tcholmanvissiidaeans both as-
signed to the genus Tcholmanvissia Zalessky, 1929),
several individuals exhibit multiple posterior branches
of CuPaα emitted before the fusion of this vein with
CuA (BÉTHOUX & NEL 2002b: fi gs. 2, 7, 8). Addition-
ally, in most titanopterans, the point of divergence of
CuPaβ and CuPaα• is located basal to the section of
CuPaα° trans. + CuPaα* that is simple (SHAROV 1968;
Figs. 1C–D, 2B). This is reminiscent of and supported
by the case discussed above. It must be noticed that no
CuPaβ was identifi ed on the specimen NHM In. 37340
(Fig. 2A).
We are left with the fact that three main stems oc-
cur between CuA + CuPaα° in part and AA1 in gi-
gantea (CuPaα° trans. + CuPaα*, CuPaα• + CuPaβ,
and CuPb), while only two occur in titanopterans
(SHAROV 1968; Figs. 1C, 2A–B; only one occurs in ex-
tensus Sharov, 1968, see below). The solution can be
readily found: CuPaα• + CuPaβ and CuPb are fused at
their origin and diverge after some distance in titano-
pterans, a fact evidenced by the very oblique origin of
CuPaα• + CuPaβ (i.e. CuPaα• + CuPaβ is translocated
onto CuPb). Moreover, alike in gigantea and beybi-
enkoi, CuPaα• + CuPaβ can readily be identifi ed in
titanopterans after its fork (which is the point of diver-
gence of CuPaα• and CuPaβ; see SHAROV 1968; Figs.
1C–D, 2B). Hence the vein CuPb is simple under this
new homologization.
At this step describing the forewing venation pattern
of extensus Sharov, 1968 (see Appendix 3 for validity
of related species; Fig. 1D) is a pinnacle. Besides the
fact that M + CuA separates into MA and MP + CuA,
that the latter fuses for some distance with CuPaα° in
part [resulting into a (MP + CuA) + CuPaα° in part

139
Arthropod Systematics & Phylogeny 65 (2)
composite stem], and that all branches of CuPaα° but
one are translocated onto CuPaα*, CuPaα° trans. +
CuPaα* is fused with the composite stem (CuPaα• +
CuPaβ) + CuPb (from which it diverges after some
Fig. 2. Forewings of giganteus Tillyard, 1916, drawings of venation (see text for abbreviations). A: Specimen NHM In. 37340
(based on a positive imprint of a left forewing, reversed; paracladotype of Tcholmanvissiidae nom. Zalessky, 1934, dis. Sharov,
1968, typ.n. and Tcholmanvissiella nom. Gorochov, 1987, dis.-typ.n.). B: Specimen NHM In. 37341, (based on a positive imprint
of a left forewing, reversed; paracladotype of Tcholmanvissiella nom. Gorochov, 1987, dis.-typ.n.). C: Specimen NHM In. 37342
(based on a negative imprint of a left forewing).
A
B
C

BÉTHOUX: Titanopterans are orthopterans
140
distance). In other words, the correct homologization
for the vein occurring between (M or MP +) CuA (+
CuPaα° in part) and AA1 is (CuPaα° trans. + CuPaα*)
+ [(CuPaα• + CuPaβ) + CuPb].
From illustrations provided by TILLYARD (1925) and
SHAROV (1968), the degree of vein fusions and trans-
locations is variable in hind wings of the group. From
the morphology exhibited by the forewing, and with
respect to the putative ancestral state as exhibited in
Permian orthopterans, I assume that CuPb and CuPaβ
are simple in hind wings of titanopterans.
I (BÉTHOUX 2005a) suggested that titanopterans and
species assigned to the Linnaean family Geraridae are
relatives on the basis of the following character states:
in forewings, vein CuPaβ branched; in hind-wings,
vein CuPb branched. From the comparative analysis
carried out above, CuPaβ is simple in titanopteran
forewings, and so is CuPb in hind wings. This hypoth-
esis was based on an erroneous interpretation of titan-
opteran wing venation and is no longer supported. The
lack of the character states ‘in forewing, fi rst posterior
branch of CuPaα (CuPaα•) occurring basal to the con-
nection of CuPaα with CuA’, and ‘in the distal half of
the forewing, RP and MA not fused’, characteristic of
the titanopterans (see below) but lacking in geraridae-
ans, support the view that both groups are not closely
related: geraridaeans are stem-orthopterans, while ti-
tanopterans are nested within the taxon including or-
thopterans.
3.2. Systematic implications and
taxonomic systems
Following his hypothesis of a close relationship bet-
ween the family Geraridae and the order Titanoptera,
and a Linnaean rank-based nomenclatural system, GO-
ROCHOV (2001: 18) included the family Geraridae with-
in the order Titanoptera and erected two suborders,
Gerarina and Mesotitanida. He provided neither diag-
nosis nor formal defi nition that could allow assignment
of species to these taxa. Additionally he mentions (p.
18) that “a less specialized, putative group of Gerarina,
or collateral lineage, may be a possible ancestral group
for the Mesotitanina and all other Or tho pteroidea”. In a
collegial contribution RASNITSYN (2002) and GOROCHOV
& RASNITSYN (2002) consider the family Geraridae as
stem-‘Polyneoptera’, distinct from the order Mesoti-
tanida, itself considered as equi valent to Titanoptera
(BELAYEVA et al. 2002). As a result, one is puzzled with
the sense to be given to the taxon name ‘Titanoptera’.
Additionally, the sub-order Mesotitanida are viewed by
GOROCHOV (2001) and GOROCHOV & RASNITSYN (2002)
as a paraphyletic group including stem-orthopterans.
Ultimately GOROCHOV (2003, 2004) reiterates the use
of the taxon name ‘Titano ptera’.
Indeed, as suggested by SHAROV (1968, 1971), the
order Titanoptera is closely related to taxa previously
assigned to the subfamily Tcholmanvissiinae, itself in-
cluded in the order Orthoptera. In other words the sub-
family Orthoptera-Tcholmanvissiinae must include
the order Titanoptera. Strictly following a rank-based
approach would necessitate an in depth reorganization
of corresponding taxa ranks.
In order to avoid issues inherent to the Linnaean
approach I follow the taxonomic system the develop-
ment of which is initiated in BÉTHOUX (2007d) and
implemented in BÉTHOUX (2007e). For convenience,
main aspects of this procedure are repeated herein.
Each taxon defi nition is set up with the designation of
two cladotypes that are specimens exhibiting a desig-
nated type-character-state. Cladotypes must belong to
different species. A name designates a monophyletic
group until one of the following assumptions is falsi-
fi ed: (1) the character state typifi ed by cladotypes is
homologous in cladotypic species, (2) the character
state typifi ed by cladotypes is derived, and (3) indi-
viduals exhibiting the type character state evolved
from an isolated (segments of) metapopulation line-
age. Taxa are assemblages for which monophyly is
objectively defi ned, testable, and emendable.
For convenience, a taxonomic application consist-
ent with the ICZN is provided in Appendix 1. The ap-
plication is designed with the aim of maximizing the
hierarchical content or names the suffi x of which is
associated to a rank. For that purpose, each supra-ge-
neric taxon is composed of only two taxa of inferior
rank. This application retrieves the same phylogenetic
information as the cladotypic application performed
below (and see Appendix 3; the genus Mesotitan Till-
yard, 1916 as newly understood might not be mono-
phyletic), plus hierarchical information based on suf-
fi xes associated to ranks. This application is left with
the problem of the authorship of the taxon name Ti-
tanoptera (see below).
Prior to the redefi nition of taxa including the spe-
cies previously assigned to the family Tcholmanvissii-
dae by BÉTHOUX & NEL (2002b) and to the order Titan-
optera by SHAROV (1968) and GOROCHOV (2003), I take
the opportunity of adapting a more inclusive taxon in
which these species are nested. This should avoid mix-
ing Linnaean and cladotypic taxon names further in
the discussion. Provisional taxa compositions are pro-
vided in Appendices 2–3. Presumed hierarchy of taxa
defi ned below is summarized on Fig. 3.

141
Arthropod Systematics & Phylogeny 65 (2)
Archaeorthoptera nom. Béthoux & Nel, 2002a,
dis.-typ.n.
Defi nition. Species that evolved from the (segments
of) metapopulation lineage in which the character
state ‘in forewings, CuA (fused with M or diverging
from it) connected to CuP or one of its branches’, as
exhibited by fi sheri Brongniart, 1885 and schneideri
Béthoux, 2005c, has been acquired (venation designa-
tions as in BÉTHOUX & NEL 2002a).
Cladotypes. Specimens MNHN-DHT-R51164 (belong-
ing to fi sheri Brongniart, 1885; see BÉTHOUX & NEL
2002a: fi gs. 13–14; BÉTHOUX & NEL 2003: fi g. 4) and
ROM 45568 (holotype of schneideri Béthoux, 2005c;
see BÉTHOUX 2005c: fi gs. 1–3).
Paracladotypes. Specimens MNHN-DHT-R51269
and MNHN-DHT-R51139 (belonging to fi sheri Brong-
niart, 1885; see BÉTHOUX & NEL 2003: fi gs. 2, 3, re-
spectively).
Discussion. The word ‘connected’ as used in the char-
acter formulation encompasses a short contact of the
two veins to a long fusion. BÉTHOUX & NEL (2002a:
14) provided another character (state) formulation for
one of the autapomorphies of the taxon Archaeortho-
ptera they list, referring to the same structure: “convex
CuA emerging from convex M + CuA […] distally
fused with anterior branch (CuPa or CuPaα) of CuP”.
At the time BÉTHOUX & NEL (2002a) named the taxon
Archaeorthoptera, all known species exhibiting a fu-
sion of CuA (distal to its divergence from M) with CuP
involved the anterior branch of the latter. A condition
was identifi ed which I considered as plesiomorphic in
the species dumasii Brongniart, 1879, which exhib-
its, in hind wings, a brief connection of CuA with the
stem of CuP, before the latter vein branches (BÉTHOUX
2003). It is clear that states regarding the branching
pattern of CuP actually belong to different character(s)
from those character states regarding the connection
of CuA with CuP. The character (state) formulated by
BÉTHOUX & NEL (2002a) includes two different char-
acters.
The new formulation of the type-character-state is
modifi ed in order to avoid ambiguity and minimize the
need of future emendations. The new formulation is
not subsumed in the character (state) formulation of
BÉTHOUX & NEL (2002a). The situation is rather op-
posite: the original character state necessarily occurs
if the new character state occurs. Additionally these
authors cannot be granted as the authors who fi rst des-
ignated a single diagnostic character state of the taxon
Archaeorthoptera because they list several autapomor-
phies in the diagnosis of the taxon. The taxon name
Archaeorthoptera is then not preoccupied.
The putative ancestral state is ‘in forewings, CuA
(fused with M or diverging from it) distinct from CuP’.
There is no argument in favour of the hypothesis of a
convergent origin of the type-character-state among
cladotypic species. This character state is assumed to
be derived, although close adelphospecies and ami-
taspecies are unknown. At least the defi ning character
state is absent in all other polyneopteran taxa. I as-
sume that individuals exhibiting the type character
state evolved from a (segments of) metapopulation
lineage isolated from other such lineages by cohesion
mechanisms.
The taxon Archaeorthoptera is nested within an un-
named taxon which type-character-state is ‘CuA fuses
with M at the wing base’. However, there is no direct
evidence of this fusion (see BÉTHOUX & NEL 2002a;
BÉTHOUX 2007a for support of this hypothesis). This
taxon is not cladotypically defi ned because appropri-
ate cladotypes are unknown. However, the fusion of
CuA with M is implicit in the defi nition of the Archae-
orthoptera.
The species elongata Brongniart, 1893: 433, listed
in the composition list (see Appendix 2), is referred
to as Ctenoptilus elongatus (Brongniart, 1893) by
BÉTHOUX & NEL (2004). The authors coordinated the
original specifi c epithet, elongata, according to a new
generic attribution (according to the ICZN, articles
31.2, 34.2). There is no reason to follow this proce-
beybienkoi Sharov, 1968
gigantea Gorochov, 1987
noinskii Zalessky, 1929
elongata Sharov, 1968
minuta Sharov, 1968: 159
Tcholmanvissiidae
Tcholmantitanopterida
Tcholmanvissiella
Pantcholmanvissiida1
2
3
4
5
longipes Martynov, 1940
giganteus Tillyard, 1916
Gigatitanidae
Titanopterida
6
Fig. 3. Scheme of presumed hierarchy in Pantcholmanvissiida;
defi ning character-states (see text for abbreviations): 1: in
forewing, fi rst posterior branch of CuPaα (CuPaα•) occurring
basal to the connection of CuPaα with CuA; 2: in the distal
half of the forewing, RP and MA distinct from each other; 3:
in forewing, CuPaβ and CuPaα• have the same point of origin
from CuPaα; 4: in forewing, at least one branches of CuPaα°
has the same point of origin as CuPaα*; 5: in forewing, CuPaα•
+ CuPaβ and CuPb have the same point of origin; 6: in forewing,
M + CuA separates into MA and MP + CuA.

BÉTHOUX: Titanopterans are orthopterans
142
dure under cladotypic taxonomy, because it results in
species name instability. The original specifi c epithet
is then restored.
Pantcholmanvissiida nom.n., dis. Béthoux & Nel,
2002b, typ.n.
Defi nition. Species that evolved from the (segments
of) metapopulation lineage in which the character
state ‘in forewing, fi rst posterior branch of CuPaα
(CuPaα•) occurring basal to the connection of CuPaα
with CuA’, as exhibited by noinskii Zalessky, 1929
and beybienkoi Sharov, 1968, has been acquired (ve-
nation designations as in BÉTHOUX & NEL 2002a; see
also BÉTHOUX & NEL 2002b).
Cladotypes. Specimens PIN 3353/391 (holotype of
noinskii Zalessky, 1929; see BÉTHOUX & NEL 2002b:
fi g. 6) and PIN 1700/4126 (holotype of beybienkoi
Sharov, 1968; see BÉTHOUX & NEL 2002b: fi g. 10).
Paracladotypes. Specimens PIN 117/258 & 259 and
PIN 3353/381 (see BÉTHOUX & NEL 2002b: fi gs. 7, 8,
respectively).
Derivatio nominis. Name based on the word ‘Tchol-
manvissiidae’ and the prefi x ‘Pan’, ‘all’ in Greek.
Discussion. The putative ancestral state is ‘in forew-
ing, fi rst posterior branch of CuPaα occurring distal to
the connection with CuA’. The type-character-state is
presumably synapomorphic for the Pantcholmanvis-
siida because it is absent in other Archaeorthoptera.
There is no argument in favour of the hypothesis of
a convergent origin of the type-character-state among
cladotypic species. I assume that individuals exhibit-
ing the type character state evolved from a (segments
of) metapopulation lineage isolated from other such
lineages by cohesion mechanisms. The Pantcholman-
vissiida encompasses species assigned to the Linnaean
family Tcholmanvissiidae by BÉTHOUX & NEL (2002b),
as well as those belonging to the Linnaean order Ti-
tanoptera as understood by SHAROV (1968) (see also
GOROCHOV 2003).
G
OROCHOV (1995: 86) formulated a character (state)
as a single synapomorphy of a group including his
(Linnaean) sub-families Tettoedischiinae and Tchol-
man vissiinae. Under the wing venation nomencla-
ture used herein, GOROCHOV (1995) suggested that the
branch basal to the fusion of CuA with CuPaα (indi-
cated by * on Fig. 4) actually belongs to CuA + CuPaα,
and fuses with CuPaα. His inter pre tation involves the
same structure (CuPaα) as that involved in the defi ni-
tion of the Pantcholmanvissiida. The possibility that
GOROCHOV (1995) should be granted as the fi rst author
who mentioned the type-character-state of the Pantch-
olmanvissiida must then be discussed. BÉTHOUX & NEL
(2002b) argued that GOROCHOV (1995) inter preta tion
is “hardly possible”, mainly because a branch of CuA
+ CuPaα cannot arise basal to the fusion of CuA with
CuPaα. However, there are two scenarios that could fi t
with GOROCHOV’s (1995) statement.
First, CuPaα could be branched proximally and its
anterior branch fused with M + CuA (Fig. 4A). The
composite vein CuA + (anterior branch of) CuPaα
would then diverge from M. It can be imagined that, in
an unknown ‘primitive’ taxon, the fi rst branch of CuA
+ CuPaα became successively oblique, then aligned
with the posterior branch of CuPaα, resulting into the
morphology exhibited by the Pantcholmanvissiida.
However, there is a major impossibility in this sce-
nario: CuPaα and its ‘sister-branch’ CuPaβ are emit-
ted from CuPa distal from the wing base (where the
hypothetical branching of CuPaα and fusion with CuA
could be unobservable on fossil material), and there is
no known related taxon in which an anterior branch
of CuPaα fuses with M + CuA distal to the origin of
CuPa.
The second scenario is more elaborate (Fig. 4B). It
implies that after its formation (i.e. fusion of CuA and
CuPaα), CuA + CuPaα is bent backwards, branches,
and fi nally runs towards the wing apex, following the
same path as earlier. This would imply that the vein
indicated herein as CuPaα° is composed of {[CuPaα
+ (CuA + CuPaα)] + (CuA + CuPaα)}. This would
be evidenced by a strengthening of the corresponding
structure, which does not occur in the known species.
The homology I propose instead is that the vein
indicated by * on Fig. 4 belongs to CuPaα (i.e. is
CuPaα* as mentioned above) and arises before the
anterior branch of the later (CuPaα°) fuses with CuA
(Fig. 4C). It is assumed that the branch of CuPaα oc-
curring basal to the fusion with CuA in Pantcholman-
vissiida is homologous to the fi rst branch of CuPaα
that diverges from CuA + CuPaα (i.e. distal to the fu-
sion of CuA with CuPaα) in sister-taxa of Pantchol-
manvissiida. From the available data, this is a more
plausible homology statement.
In summary, the homology statement provided by
GOROCHOV (1995) can be seen as a correct primary
homology statement within Pantcholmanvissiida (the
character state is similar in species assigned to this
taxon), a plausible secondary homology statement
within Archaeorthoptera (the character state was ac-
quired by common ancestry), but the primary homol-
ogy statement is erroneous within Archaeorthoptera
(the structure described as the character is not derived
from the structure it is supposed to). However, the
most important point is that, theoretically, the character
(state) defi ned by GOROCHOV (1995) and the character
state I use for defi ning the Pantcholmanvissiida could
co-occur (Fig. 4D). This can be viewed as characters
that fail the conjunction test (PATTERSON 1982, 1988;
see also DE PINNA 1991), hence they are not homolo-
gous. Therefore GOROCHOV (1995) cannot be granted

143
Arthropod Systematics & Phylogeny 65 (2)
as the author who fi rst designated the type-character-
state of the Pantcholmanvissiida as defi ned herein, but
BÉTHOUX & NEL (2002b), who listed the corresponding
character state as the only diagnostic character of the
family Tcholmanvissiidae.
The taxon name Tcholmanvissiidae cannot be used
for the taxon under scrutiny because it is preoccupied, as
SHAROV (1968) explicitly associated it to another char-
acter state (see below). Therefore, another name must
be searched for. GOROCHOV (1995: 86) erected no name
for the taxon including his Tettoedischiinae and Tchol-
manvissiinae. Therefore I erect a new taxon name.
Tcholmanvissiidae nom. Zalessky, 1934,
dis. Sharov, 1968, typ.n.
Defi nition. Species that evolved from the (segments
of) metapopulation lineage in which the character
state ‘in the distal half of the forewing, RP and MA
distinct from each other’, as exhibited by longipes
Martynov, 1940 and giganteus Tillyard, 1916, has
been acquired.
Cladotypes. Specimens PIN 1700/1488 (holotype of
longipes Martynov, 1940; see BÉTHOUX & NEL 2002b:
fi g. 6) and AM F.36274 (specimen attributed to gigan-
teus Tillyard, 1916; see MCKEOWN 1937: fi gs. 1–3, pl.
4; see JELL 2004: unnumbered fi gure on p. 29; GRIMAL-
DI & ENGEL 2005: fi g. 7.42).
Paracladotypes. Specimens PIN 1452/5 and PIN 1700/
1454 (belonging to longipes Martynov, 1940; see BÉ-
THOUX & NEL 2002b: fi gs. 2, 4, respectively), and MNH
In. 37340 (belonging to giganteus Tillyard, 1916;
Fig. 2A; see Zeuner, 1939: pl. LXXX, fi g. 1).
Discussion. The putative ancestral state is ‘in the dis-
tal half of the forewing, RP and MA fused for some
distance’. The type-character-state appeared more
than once among Orthoptera (BÉTHOUX & NEL 2002a).
It is a reversion of a character state acquired in stem-
ortho pterans, namely the fusion of RP with MA (or
one of its anterior branches). A connection of RP
with MA is present in successive sister-groups of the
Tcholman vissiidae. Considering the series of character
state changes that separate the Tcholmanvissiidae from
other ortho pterans exhibiting the same character state,
it is assumed that it appeared in the common ancestor
of longipes and andersoni and is locally apomorphic.
There is no argument in favour of the hypothesis of
a convergent origin of the type-character-state among
clado typic species. I assume that individuals exhibit-
ing the type character state evolved from a (segments
of) metapopulation lineage isolated from other such
lineages by cohesion mechanisms. This taxon encom-
passes the subfamily Tcholmanvissiinae as understood
by BÉTHOUX & NEL (2002b) and the order Titanoptera
as understood by SHAROV (1968).
Despite the fact that the name Tcholmanvissiidae
has a suffi x typical of Linnaean families, I adapt it un-
modifi ed in the new cladotypic taxonomy because it is
preoccupied. Neither ZALESSKY (1929), nor ZALESSKY
(1934), nor MARTYNOV (1940) mentioned a unique
Fig. 4. Possible scenarios for the homologization of the vein
indicated by *, either as a branch of CuA + CuPaα fused with
CuPaα (GOROCHOV 1995) (A, B) or as a branch of CuPaα
(BÉTHOUX & NEL 2002b) (C), and possible co-occurrence
of homologizations B and C (D) (colour coding as in Fig. 1;
posterior branches of CuA are represented by dashed lines
as CuA is considered as branched by GOROCHOV 1987, 1995
but simple in this contribution; CuPaα• is represented by
a dashed line as it is not the focus of this illustration, and it
is not fused with CuPaβ in all Pantcholmanvissiida; see text
for abbreviations). A: CuPaα is branched, its anterior branch
fuses with M + CuA, and its posterior branch fuses with CuA +
CuPaα. B: CuA + CuPaα, after its formation (fusion of CuA and
CuPaα), bends backwards, branches, and fi nally runs towards
wing apex. C: CuPaα is branched basal to its connection with
CuA. D: co-occurrence of homologizations B and C.
A
B
C
D

BÉTHOUX: Titanopterans are orthopterans
144
character (state) diagnostic of the family Tcholman-
vissiidae, but SHAROV (1968) mentioned that “the spe-
cies of Tcholmanvissiidae differ from the Oedischiidae
mainly in the absence of an anastomosis between the
anterior branch of MA and RS [RP]” (translation from
SHAROV 1971: 29); he mentioned no other “main” dia-
gnostic character state. This is a homology statement
synonymous to that given for the type-character-state
of Tcholmanvissiidae as herein, although under a dif-
ferent wing venation nomenclature. Therefore priority
is given to SHAROV (1968) as the author who fi rst des-
ignated the type-character-state of this taxon.
The holotype of the species longipes Martynov,
1940 is selected as cladotype because several fore-
wings belonging to this species are described and
they consistently exhibit an MA distinct from RP (see
BÉTHOUX & NEL 2002b).
After BÉTHOUX & NEL (2002b), the genus Tchol-
manvissia (erected by Zalessky, 1929) includes two
species (noinskii Zalessky, 1929, and longipes Mar-
tynov 1940). In the diagnosis provided by these au-
thors, not a single diagnostic character state that could
have allowed the name ‘Tcholmanvissia’ to be adapted
is mentioned. I found none in the literature. The taxon
Tcholmanvissiidae is the least inclusive taxon includ-
ing noinskii and longipes that is cladotypically defi ned,
therefore it should be used as the taxonomic address
(CANTINO et al. 1999; DAYRAT et al. 2004) for the spe-
cies previously assigned to the genus Tcholmanvissia.
Correct taxonomic combinations are then Tcholman-
vissiidae noinskii Zalessky, 1929 and Tcholmanvissii-
dae longipes Martynov 1940.
There is some uncertainty regarding the specifi c
assignment of the specimen AM F.36274, to which the
status of cladotype is given in various places herein.
Until the “argument” mentioned by JELL (2004: 8)
is elucidated, I follow SHAROV (1968), CARPENTER
(1992), and GOROCHOV & RASNITSYN (2002) who con-
sidered that it belongs to the species giganteus Till-
yard, 1916. In any case, under cladotypic taxonomy,
names of species and of taxa other than species are
defi ned independently. A supra-specifi c taxon name
defi nition can be emended if the specifi c identity of a
cladotype provided in an early defi nition is incorrect.
If so, the cladotype identity prevails over the species
name given in the defi nition (BÉTHOUX 2007d).
Tcholmantitanopterida nom.-dis.-typ.n.
Defi nition. Species that evolved from the (segments
of) metapopulation lineage in which the character
state ‘in forewing, CuPaβ and CuPaα• have the same
point of origin from CuPaα’, as exhibited by gigantea
Gorochov, 1987 and giganteus Tillyard, 1916, has
been acquired (venation designations as in BÉTHOUX &
NEL 2002b and herein).
Cladotypes. Specimen PIN 3353/78 (holotype of
gigantea Gorochov, 1987; see BÉTHOUX & NEL 2002b:
fi g. 11) and AM F.36274 (specimen attributed to gi-
gan teus Tillyard, 1916; see MCKEOWN 1937: fi gs. 1–3,
pl. 4; JELL 2004: unnumbered fi gure on p. 29; GRIMALDI
& ENGEL 2005: fi g. 7.42).
Derivatio nominis. Name based upon the words
Tcholmanvissia and Titanopterida.
Discussion. The putative ancestral state is ‘in fore-
wing, CuPaβ and CuPaα• with distinct origins’. The
type-character-state is presumably apomorphic of the
Tcholmantitanopterida because it is absent in oth er
Archae orthoptera, Pantcholmanvissiida, and Tchol-
man vissiidae. There is no argument in favour of the
hypothesis of a convergent origin of the type-character-
state among cladotypic species. I assume that indi-
viduals exhibiting the type character state evolved
from a (segments of) metapopulation lineage isolated
from other such lineages by cohesion mechanisms.
The genus Jubilaeus Sharov, 1968 includes the
species beybienkoi only. As far as I am aware there is
no single diagnostic character state that could allow
its ‘association’ to another taxon within the Tchol-
mantitanopterida. The adaptation of the name ‘Jubi-
laeus’ into cladotypic taxonomy is then currently
impossible. For the same reason as above, the correct
taxonomic combination for this species is Tchol-
mantitanopterida beybienkoi Sharov, 1968.
Tcholmanvissiella nom. Gorochov, 1987, dis.-typ.n.
Defi nition. Species that evolved from the (segments
of) metapopulation lineage in which the character
state ‘in forewing, at least one branch of CuPaα° has
the same point of origin as CuPaα*’, as exhibited
by gigantea Gorochov, 1987 and giganteus Tillyard,
1916, has been acquired (venation designations as in
BÉTHOUX & NEL 2002a and herein).
Cladotypes. Specimens PIN 3353/78 (holotype of
gigantea Gorochov, 1987; see BÉTHOUX & NEL 2002a:
fi g. 11) and AM F.36274 (specimen attributed to
giganteus Tillyard, 1916; see MCKEOWN 1937: fi gs.
1–3, pl. 4; JELL 2004: unnumbered fi gure on p. 29;
GRIMALDI & ENGEL 2005: fi g. 7.42).
Paracladotypes. Specimens NHM In. 37340 and NHM
In. 37341 (specimens attributed to giganteus Tillyard,
1916; Fig. 2A,B, respectively; see ZEUNER 1939: pl.
LXXX, fi gs. 1, 2, respectively).
Discussion. The putative ancestral state is ‘in forewing,
all branches of CuPaα° have a point of origin distinct
from that of CuPaα*’. The type-character-state is
pre sumably apomorphic of the Tcholmanvissiella be-
cause it is absent in other Archaeorthoptera, Pan-
tchol manvissiida, Tcholmanvissiidae, and Tcholman-
titanopterida. There is no argument in favour of the
hypothesis of a convergent origin of the type-charac-

145
Arthropod Systematics & Phylogeny 65 (2)
ter-state among cladotypic species. I assume that in-
di viduals exhibiting the type character state evolved
from a (segments of) metapopulation lineage isolated
from other such lineages by cohesion mechanisms.
G
OROCHOV (1987: 79) provided a diagnosis of the
genus Tcholmanvissiella which is: “on forewing, the
stem of MP + CuA1 [CuA + CuPaα°] has almost
no branches, and main ridge of branches of MP +
CuA1 [CuA + CuPaα°] is located proximally to
anastomosis of MP [CuA] with CuA1 [CuPaα°]”.
Once again, it is diffi cult to understand how branches
of CuA + CuPaα° could occur proximal to the fusion
of the constituents of this composite vein (CuA
and CuPaα°). If one considers GOROCHOV’s (1987)
diagnosis as composed of a single character state (but
see below), this character state is distinct from that
used to defi ne the Tcholmanvissiella as herein: for
the same reasons as detailed above (see discussion on
Pantchomanvissiida), the character states ‘main ridge
of branches of CuA + CuPaα° located proximally to
anastomosis of CuA with CuPaα°’ could co-occur with
the character state ‘in forewing, at least one branches
of CuPaα° has the same point of origin as CuPaα*’.
Therefore GOROCHOV (1987) cannot be granted as the
fi rst author who designated the type-character-state of
the taxon Tcholmanvissiella as defi ned herein.
If one considers GOROCHOV’s (1987) diagnosis as
composed on a single character state, the taxon name
Tcholmanvissiella is preoccupied. However, there
is no argument supporting this view. In the same
paper, several diagnoses in which distinct characters
(states) are listed end with “, and [last character
state]”. Therefore, in “the stem of MP + CuA1 [CuA
+ CuPaα°] has almost no branches, and main ridge of
branches of MP + CuA1 [CuA + CuPaα°] is located
proximally to anastomosis of MP [CuA] with CuA1
[CuPaα°]”, “main ridge of branches of MP + CuA1
[CuA + CuPaα°] is located proximally to anastomosis
of MP [CuA] with CuA1 [CuPaα°]” appears as a
second character (state) distinct from the former.
Therefore, I suppose that GOROCHOV’s (1987) diagno-
sis refers to two different characters (states), and that
the name Tcholmanvissiella is not preoccupied. Hence
I can freely adapt it in cladotypic taxonomy.
The only species assigned by GOROCHOV (1987) to
the genus Tcholmanvissiella has no known diagnostic
character state on its own, and cannot be associated to
any known species apart from those assigned to the
Titanopterida (defi ned below). In other words, this
species is the only member of the Tcholmanvissiella
that is not a Titanopterida. The least inclusive taxon
including gigantea that is cladotypically defi ned is
Tcholmanvissiella. Incidentally the Linnaean bino mi-
nal is preserved: the correct combination is Tchol man-
vissiella gigantea Gorochov, 1987.
One could have noticed that cladotypes of taxa
Tchol man titanopterida and Tcholmanvissiella are iden-
tical. It is not an issue under the taxonomic procedure
used herein because typifi cation is based upon a pair of
individuals and a character state (BÉTHOUX 2007d).
Titanopterida nom.-dis.-typ.n.
Defi nition. Species that evolved from the (segments
of) metapopulation lineage in which the character
state ‘in forewing, CuPaα• + CuPaβ and CuPb having
the same point of origin’, as exhibited by giganteus
Tillyard, 1916 and vulgaris Sharov, 1968, has been
acquired (venation designations as in BÉTHOUX & NEL
2002a and herein).
Cladotypes. Specimens AM F.36274 (specimen attrib-
uted to giganteus Tillyard, 1916; see MCKEOWN 1937:
fi gs. 1–3, pl. 4; JELL 2004: unnumbered fi gure on p. 29;
GRIMALDI & ENGEL 2005: fi g. 7.42) and PIN 2240/4593
(holotype of vulgaris Sharov, 1968; see SHAROV 1968:
fi g. 50B).
Derivatio nominis. Based on the word ‘Titanoptera’.
Discussion. The putative ancestral state is ‘in forew-
ing, CuPaα• + CuPaβ and CuPb having distinct points
of origin’. The type-character-state is presumably apo-
morphic of the Titanopterida because it is absent in
other Archaeorthoptera, Pantcholmanvissiida, Tchol-
manvissiidae, Tcholmantitanopterida, and Tcholman-
vissiella. I assume that cohesion mechanisms isolated
individuals exhibiting the type-character-state from
those that do not. The occurrence of the type-charac-
ter-state on the specimen PIN 2240/4593 was assessed
based upon examination of photographs provided by
A.P. Rasnitsyn (pers. comm. 2007).
Adaptation of a name for this taxon is a tricky case.
The composition of the taxon matches that given by
SHAROV (1968: 123) to the order Titanoptera. This
author (p. 123) mentioned two character states that
differentiate the order Titanoptera from the order Or-
thoptera, and none are formulated precisely enough
to be eligible as type-character-state. Indeed, SHAROV
(1968) is not the author of this taxon name, but BRONG-
NIART (1885: 379), who erected it as a genus name for
a fragmentary fossil specimen I regard as belonging to
stem-odonatans. BRONGNIART (1885) did not explicitly
associate this name to a single character state.
T
ILLYARD (1916) fi rst described a species belong-
ing to the order Titanoptera as understood by SHAROV
(1968). He assigned it to a new genus, Mesotitan, but
did not provide a single diagnostic character (state),
nor any state convincingly diagnostic. Later on TILL-
YARD (1925) described a new species (which is actu-
ally a hind wing of the former species) he assigned
to the same genus Mesotitan, and erected the family
Mesotitanidae. However, none of the character states
he mentioned are strictly diagnostic of the taxa he
created. In the same vein CRAMPTON (1928) erected
the order Mesotitanoptera on the basis of the family

BÉTHOUX: Titanopterans are orthopterans
146
name Mesotitanidae without providing any diagno-
sis.
Although they state that the names Mesotitanida
(fi rst coined by GOROCHOV 2001 as that of an order
synonym of Gerarida) and Titanoptera refer to the
same taxa, GOROCHOV & RASNITSYN (2002) preferred
the former, and refer to TILLYARD (1925) as the person
who erected the former name. However, this is not the
case: this reference is based on a rule-free coordina-
tion of a Linnaean familial name [as it is, Mesotitani-
dae, erected by TILLYARD (1925)] into a Linnaean or-
dinal name, and following a rule of priority (TILLYARD
1925 rather than SHAROV 1968). This procedure is not
followed here. Neither GOROCHOV (2001) nor GORO-
CHOV & RASNITSYN (2002) provided a unique character
state diagnostic of the order Mesotitanida. GOROCHOV
(2001) also erected the subordinal name Mesotitanina,
without mention of a single diagnostic character state.
Therefore, as far as I am aware, there is no previous
association of a single character state to a taxon name
including the species assigned to the order Titano-
ptera by SHAROV (1968). All available Linnaean names
refer to the great size of most known species, but this
is hardly a reliable character for defi ning a taxon. Ad-
ditionally, the situation with names erected under the
Linnaean system is confusing. Therefore, I erect a new
name, designate a new type-character-state, and desig-
nate cladotypes accordingly.
Regarding the composition of the group, the po-
sition of the species vladimiri Gorochov, 2004 (as-
signed to the Linnaean genus Permotitan Gorochov,
2004) must be discussed. An anomaly of the hypo-
thesis stating that the geraridaeans and titanopterans
are close relatives, defended by GOROCHOV (2001)
and followed by BÉTHOUX (2005a), was the absence
of both groups during the whole Permian period. This
was before GOROCHOV (2004) assigned vladimiri, from
the Permian Vorkuta coal basin (Russia), to the order
Titanoptera, or closely related to this taxon. This as-
signment was based upon (1) the large size of the spec-
imen (estimated forewing length about 140 mm), (2)
the occurrence of regular cross-venation between ScP
branches, and (3) the area between the anterior wing
margin and ScP that does almost not taper proximal to
its end. Size (1) can hardly be viewed as a character of
defi nitive phylogenetic interest. Character (2) is not an
obvious trait of Titanopterida as it is not occurring in
libelluloides Sharov, 1968. This character varies great-
ly among Permian orthopterans and occurs in many
Archaeorthoptera. The validity of the character (3) is
diffi cult to assess in Titanopterida yielded by the de-
posit of Madygen (Trias; Russia) because of the effect
of post-depositional deformation that affected fossils
(SHAROV 1968; RASNITSYN 1982). Unfortunately, this
material is the basis for most of our knowledge on Ti-
tanopterida. SHAROV (1971: 210) described specimens
probably belonging to the species extensus Sharov,
1968 as having a “costal fi eld gradually tapering to
the apex of the wing”, suggesting that character (3)
is either diffi cult to appreciate or not diagnostic of Ti-
tanopterida, or both.
Additionally vladimiri exhibits ScP branches mak-
ing a 40° angle with the main stem of ScP. When
present in Titanopterida, such branches usually make
a more oblique angle in forewing, especially in the
distal area. Additionally, the point of divergence of
MA and MP (free part of MP under the nomenclature
used by Gorochov) is located in a very distal position,
unlike in known Tcholmanvissiidae. Finally, it must
be noticed that the restoration of vladimiri provided
by GOROCHOV (2004), based upon a very incomplete
and single specimen, is highly speculative. It cannot
be ruled out that a fusion of the anterior branch of MA
with RP, commonplace among Permian orthopterans
but absent in Tcholmanvissiidae, actually occurred in
this species. Finally, critical review of data on vladimiri
lead me to conclude that it cannot be conclusively as-
signed to the Pantcholmanvissiida.
Gigatitanidae nom. Sharov, 1968, dis.-typ.n.
Defi nition. Species that evolved from the (segments
of) metapopulation lineage in which the character
state ‘in forewing, M + CuA separates into MA and
MP + CuA’, as exhibited by vulgaris Sharov, 1968 and
extensus Sharov, 1968, has been acquired (venation
designations as in BÉTHOUX & NEL 2002a).
Cladotypes. PIN 2240/4593 (holotype of vulgaris
Sharov, 1968; see SHAROV 1968: fi g. 50B) and PIN
2240/4503 (paratype of extensus Sharov, 1968).
Paracladotypes. Specimens PIN 2240/4526 and PIN
2555/1541 (see SHAROV 1968: pl. XII fi gs. 2 and 5,
respectively), and FG/596/IV/1 (see Figs. 5–6), all at-
tributed to vulgaris Sharov, 1968.
Discussion. The putative ancestral state is ‘in forew-
ing, M + CuA separates into M (= MA + MP) and CuA’.
The type-character-state is present in grylliformis Sha-
rov, 1968, which is a genuine cricket. The type-charac-
ter-state is presumably apomorphic of the Gigatinati-
dae because it is absent in other Pantcholmanvissiida,
Tcholmanvissidae, Tcholmantitanopterida, Tcholman-
vissiella, and Titanopterida, all taxa from which grylli-
formis Sharov, 1968 can be readily excluded. I assume
that individuals exhibiting the type character state
evolved from a (segments of) metapopulation lineage
isolated from other such lineages by cohesion mecha-
nisms. The occurrence of the type-character-state on
specimens PIN 2240/4593 and PIN 2240/4503 was as-
sessed after examination of photographs provided by
A.P. Rasnitsyn (pers. comm., 2007)
S
HAROV (1968: 131) mentioned the selected type-

147
Arthropod Systematics & Phylogeny 65 (2)
character-state as one of the diagnostic character (state)
of the family Gigatitanidae: “the base of MA2 [MP] is
displaced to MP [CuA] or even to MP + CuA1 [CuA +
CuPaα°] ”. The character state is not mentioned in the
determination key provided by SHAROV (1968: 157),
where he cited two characters (states). Hence there is
no known preoccupation of the name. My decision re-
garding the choice of the character-state is based upon
my opinion that it can be more sharply defi ned than
other characters mentioned by Sharov, hence minimiz-
ing the risk of future emendations.
S
HAROV (1968: 202) distinguished the genus Na-
notitan Sharov, 1968, to which he assigned extensus,
from the genus Gigatitan Sharov, 1968, to which he
assigned vulgaris, based upon the following characters
(states): “absence of a differentiated proximal branch
of Sc [ScP], [...] fusion of the base of MA2 [MP] and
MP + CuA1 [CuA], [...] and fusion of the bases of
CuA2 [CuPaα° trans. + CuPaα*] and CuP [(CuPaα• +
CuPaβ) + CuPb]”. Although extensus in known from
few specimens, all these characters suggest that they
belong to a species distinct from vulgaris.
Fig. 6. Specimen FG 596/IV-1, assigned to vulgaris Sharov,
1968, paracladotype of Gigatitanidae nom. Sharov, 1968,
dis.-typ. n.: detail of the wing base (negative imprint of a right
forewing, reversed, light-mirrored; see text for abbreviations).
Fig. 5. Specimen FG 596/IV-1, assigned to vulgaris Sharov, 1968, paracladotype of Gigatitanidae nom. Sharov, 1968, dis.-typ. n.:
drawing of the venation and photograph (negative imprint of a right forewing, reversed; see text for abbreviations).

BÉTHOUX: Titanopterans are orthopterans
148
4. Discussion
4.1. Origin and evolution of the Titanopterida
Based on a new comparative analysis and interpreta-
tion, the wing venation pattern of the titanopterans / ti-
tanopteridans is homologised with respect that of oth-
er orthopterans / archaeorthoperans. The evolutionary
history of the former group is then less puzzling than
previously thought: rather than suddenly radiating and
disappearing during the Triassic, Titanopterida arose
from a set of large-sized Permian orthopterans. This
scenario implies that raptorial forelegs, known in Tri-
assic Titanopterida (SHAROV 1968), were acquired in
orthopterans, and questions the view that ancestral
orthopterans were herbivorous. Orthopterans gain an
additional and unique stridulatory apparatus, occur-
ring in both sexes, and Titanopterida experienced a
reduction or loss of hind leg structures related to jump
(SHAROV 1968). As now known the Pantcholmanvis-
siida represents a lineage that survived the Permian /
Triassic biocrisis. However an accurate estimation of
the impact of this event on the taxon diversity is out-
of-reach, due to the incompleteness of our record.
4.2. Fossil taxa and saturated morphologies
The paper highlights the contribution of fossil mate-
rial for assessing homologies in derived taxa. The
species gigantea Gorochov, 1987 exhibits a fusion
of CuPaα• with CuPaβ, which is an apomorphic state
among orthopterans, but no translocation of CuPaα°
branches onto CuPaα*, a plesiomorphy within Tchol-
manvissiella, to which it belongs. This interpretation
resulted into a new homologization of the wing vena-
tion of Titanopterida, itself resulting into a strong sup-
port for one of the available phylogenetic hypotheses
regarding the origin of the group.
It is worth mentioning that SHAROV (1968, 1971),
despite his visionary statement that Titanopterida de-
rived from Tcholmanvissiidae, did not achieve a cor-
rect homologization. In my opinion, this is arguably
related to the fact that gigantea Gorochov, 1987, with
its unique character states combination, was unknown
to him. Although this study concerns fossil taxa only,
I consider this example as a demonstration of the use-
fulness of fossils for determining primary homolo-
gies and character states polarity, hence relationships,
among taxa exhibiting saturated morphologies.
4.3. Practical cases of cladotypic taxonomy
This fi rst application of cladotypic taxonomy unveiled
several practical aspects of this nomenclatural system.
In the following section I discuss a proposition govern-
ing the adaptation of previously erected taxon names
based on various cases of character state formulations,
the issue of the occurrence of Linnaean suffi xes within
a cladotypic taxonomy, the issue of Linnaean binomi-
nals within a cladotypic taxonomy, and the capacity
to handle the ancestor ‘species’ vs. apomorphy-less
sister-species issue by the various nomenclatural sys-
tems. First, I discuss the question of accuracy of type
character state formulations.
4.3.1. Character state formulations and antonyms
Formulation of type character states is a matter of
semantics when coming to emendation of previous
defi nitions. Although cladotypic taxonomy has the es-
sential advantage of allowing defi nition emendations
to be performed thanks to reference to type-speci-
mens (BÉTHOUX 2007d), formulations should avoid
polysemic words, so that the risk of confusion in fu-
ture emendations is lowered. I suggest that antonyms
could be mentioned in the discussion associated to the
defi nition, or in the formulation even. The formula-
tion of the putative ancestral state, which should be
an antonym of the type character state formulation,
fi ts these requirements. This mention is desirable as
it could help to circumvent the meaning of the defi ni-
tion. Explicit references to fi gures are also desirable,
as they are character state ostentations.
4.3.2. Adaptation and character state formulation
Earlier I insisted on the fact that a “taxon [...] name
[...] is to be permanently anchored to the original ho-
mology assumption” (BÉTHOUX 2007d). It results in
the notion of name pre-occupation, which can be ap-
plied to previously erected Linnaean taxa. Therefore
the capacity to determine whether a former character
state formulation includes, equates to, or is subsumed
into a later character state formulation is essential for
the adaptation of previously erected Linnaean names,
as well as for emendation of cladotypically defi ned
taxa. Three cases are possible: (1) a latter character
state formulation is subsumed in a former one if the
latter character state necessarily occurs if the former
character state occurs, but not reciprocally; (2) a
former character state formulation is equivalent to a
latter one if the latter character state necessarily occurs
if the former character state occurs, and reciprocally
(i.e. the former character state necessarily occurs if the
latter character state occurs); and (3) a latter character
state formulation subsumes a former one if the former
character state necessarily occurs if the latter character
state occurs, but not reciprocally. In other words, (1)
the latter character state formulation is a hyponym of
the former character state formulation; (2) the latter

149
Arthropod Systematics & Phylogeny 65 (2)
character state formulation is a synonym of the former
character state formulation; and (3) the latter character
state formulation is a hypernym of the former char-
acter state formulation. This test can be viewed as a
derivation of the conjunction test (PATTERSON 1982,
1988; see also DE PINNA 1991). Adaptation of a preoc-
cupied erected name explicitly related to a single di-
agnostic character state is possible in the second case
only. Emendation of a previous taxon defi nition is ac-
ceptable only in the fi rst case. If a previously erected
Linnaean name is associated to a character state which
is actually composed of several character states (e.g.
Archaeorthoptera in BÉTHOUX & NEL 2002a), emenda-
tion is impossible because cladotypes are not available
(see BÉTHOUX 2007d for emendation procedure). As a
result, the taxon name is not preoccupied.
4.3.3. Preoccupation and Linnaean suffi xes
Taxa for which names end with a Linnaean suffi x could
be preoccupied. This is the case of the taxon Tchol-
manvissiidae, for which a previous author explicitly
associated the corresponding taxon to a single char-
acter state. Two options are possible: either the taxon
name is adapted unchanged, or a new name is created,
considering that the previous one was not erected un-
der cladotypic taxonomy. If one desires the work of
previous researchers to be acknowledged, the former
option is to be preferred. It must be noticed that the
name of the type-genus of a given family could alter-
natively be selected. However, it is possible that both
names are preoccupied, based on different diagnostic
character states (as well as names of the corresponding
subfamily, tribe, subtribe, etc., all derived from the ge-
nus name). Therefore, I suggest that Linnaean names
to which a suffi x has been assigned could be adapted
unchanged under cladotypic taxonomy.
If so, and as a result of the adaptation process, a
taxon, the name of which ends with a given Linnaean
suffi x, could include taxa for which names end with
the suffi x of an equivalent or higher rank. This is the
case of the taxon Tcholmanvissiidae, including the tax-
on Gigatitanidae. In fi rst instance this is confusing for
taxonomic practitioners used to Linnaean ranks. How-
ever, this case happens when any Linnaean ranked tax-
on is found to be paraphyletic (which is the case of the
family Tcholmanvissiidae). The treatment of this prob-
lem is merely different under Linnaean and cladotypic
taxonomies: under the former, suffi xes of taxon names
are modifi ed according the modifi cation of their rank;
under the latter, no modifi cation is necessary, which I
believe is to be preferred, as it renders taxonomy more
stable. All in all, the occurrence of rank-based suffi xes
within a cladotypic taxonomy can be viewed as a lega-
cy of the Linnaean framework, but some might prefer
to strictly exclude corresponding names.
4.3.4. Linnaean binominals, taxonomic addresses,
and preoccupation
Under rank-less uninominal species nomenclature,
taxon names listed in a taxonomic address have the
function of providing information about phylogenetic
relationships of species (LANHAM 1965; CANTINO et
al. 1999; DAYRAT et al. 2004): names of taxa of vari-
ous inclusiveness precede the specifi c epithet (e.g.
Archaeorthoptera Pantcholmanvissiida Tcholmanvis-
siidae longipes Martynov, 1940). If a Linnaean genus
name is adapted, species belonging to this genus could
be designated by the same combination as under Lin-
naean nomenclatural system (e.g. Tcholmanvissiella
gigantea Gorochov, 1987). It implies that the adapta-
tion of taxon names previously understood as those of
genera is critical if one’s desire it to preserve continu-
ity between Linnaean taxonomy and a new cladotypic
one.
However, as a result of an expectable improvement
of phylogenetic relationships resolution within Lin-
naean genera as currently understood, it is likely that
new taxa nested ‘between’ adapted Linnaean genera
and species will be defi ned. The rule stating that the
taxonomic address of a species should end with the
name of the least inclusive taxon implies that Linnae-
an binominals, as we now know them, will be lost at
some point (but see below). This will result in discon-
tinuity between a Linnaean and a mature cladotypic
taxonomy. Nevertheless, it must be acknowledged that
under the Linnaean nomenclatural system, new infor-
mation on intra-generic relationships are taken into
account by authoritatively and arbitrarily modifying
the composition of the previous genus, authoritatively
and arbitrarily erecting new genera, and, if necessary,
modifying the specifi c epithet in accordance with the
gender of the new genus to which a species is assigned
(and, if necessary, authoritatively and arbitrarily modi-
fying higher taxa ranks, hence their name). If the con-
cern is about a stable binominals database, the Lin-
naean generic nomenclatural system is not more stable
than a (least inclusive taxon name + uninominal) cla-
dotypic system.
Still, if one’s opinion is that authoritative and arbi-
trary selection of a name to be applied as ‘pre-epithet’
before a species ‘epithet’ would guarantee stability of
a species database and allow an easier use of it, it can
be envisioned that one of the taxon names of a spe-
cies taxonomic address could be selected as a main
‘pre-epithet’. However, I believe that the cladotypic
procedure, coupled with the rank-less uninominal spe-
cies proposition (LANHAM 1965; DAYRAT et al. 2004),
is governed by quite intuitive principles. After all, a
mature cladotypic taxonomy is, in a simplistic view,
no less than a determination key in which dichotomies
leave the choice between a (hypothetical) plesiomor-
phy and an (hypothetical) synapomorphy, each step

BÉTHOUX: Titanopterans are orthopterans
150
being preceded by modifi cations that arose earlier in
the history of a taxon. This applies up to the species
level, for which diagnoses could be provided (see
BÉTHOUX 2007d for the species defi nition). Therefore a
mature cladotypic taxonomy would be highly practical
to taxonomy users, besides the crucial introduction of
the concepts of biological evolution (through evidence
of historical modifi cation of character states) and of
derived character state (rather than similarity) to non-
professionals.
The rule of priority given to preoccupied names is
in confl ict with a possible rule of priority given to pre-
vious Linnaean genera (aimed at preserving Linnaean
binominals). My opinion is that the former takes pre-
cedence over the latter, as I allocate more importance
to the author who fi rst identifi ed a unique diagnostic
character state of a taxon rather than to the author who
fi rst erected a name and authoritatively and arbitrar-
ily (or by allegiance to one of the International Codes
of Nomenclature) assigned it as genus name to a spe-
cies.
4.3.5. Ancestor ‘species’ or apomorphy-less
sister-species
The new nomenclatural system has the capacity to
classify species that have no known apomorphy on
their own but that of the least inclusive taxon to which
they belong (e.g. Tcholmanvissiella gigantea Goro-
chov, 1987). These ‘species’ are putatively composed
of individuals belonging to the ‘ancestor lineage’, or
are apomorphy-less sister-species of the other member
of the taxon they belong to (in our example, the taxon
Titanopterida). Cladotypic taxonomy relies on the as-
sumption that cohesion mechanisms isolated individu-
als exhibiting a type-character-state from those that do
not, or individuals exhibiting the type character state
evolved from a (segments of) metapopulation lineage
isolated from other such lineages by cohesion mech-
anisms. This is the null hypothesis. In our example,
individuals of the taxon Titanopterida are supposed
to have been isolated from those belonging to Tchol-
manvissiella gigantea Gorochov, 1987. Therefore, the
latter taxon is automatically assumed to be an apomor-
phy-less (or ‘apomorphy-unknown’) sister-species
rather than an ancestral species. If one proves that no
isolation occurred between individuals that acquired
the type-character-state of the Titanopterida and those
belonging to Tcholmanvissiella gigantea Gorochov,
1987, the isolation assumption is not respected, the
former taxon is invalidated, and the latter taxon is a
stem-lineage ‘species’ indeed. Several approaches ex-
ist that could demonstrate the absence of isolation (see
DAYRAT 2005; and references therein). Nevertheless,
in my opinion, the distinction between stem-lineage
‘species’ (or ancestor ‘species’) and apomorphy-less
sister-species is a question that has little practical im-
plications on taxonomy. If palaeontological research is
aiming at unveiling unknown character state combina-
tions and providing data on the minimal divergence
dates, cladotypic taxonomy is fully capable of dealing
with these two main inputs: species name combina-
tions convey phylogenetic information at the highest
degree of precision, and the minimal age of a taxon
is automatically that of its apomorphy-less sister-spe-
cies: by defi nition, if this species is indeed an ancestral
‘species’, the taxon does not exist.
The Linnaean nomenclatural system necessitates
the creation of a surfeit of ranked taxa for ancestral
‘species’ / apomorphy-less sister-species. The Phy-
loCode, as currently developed, in unable to deal with
the question because it does not take into account the
species case, this being due to the fact that it lacks the
isolation assumption. With respect to these nomenclat-
ural systems, the cladotypic one arguably handles the
question in the most simple and informative way.
5. Conclusion
Comparative investigations carried out in this con-
tribution demonstrate that the Triassic Titanopterida
derived from Early Permian Pantcholmanvissida. Our
view of titanopteridans evolution is greatly modifi ed
once the earlier relatives are taken into account: rather
than suddenly radiating and disappearing during the
Triassic, titanopteridans existed for at least 50 My
(million years). Our poor record of this group does not
allow any accurate estimation of the impact of the Per-
mian / Triassic biocrisis on the group.
Forewing venation patterns are found to be highly
complex, hence phylogenetically informative. I argue
that more facilities should be involved in comparative
morphological studies dedicated to the wing venation
character system of palaeopteran and polyneopteran
taxa, including extant ones. Putative results encompass
improvement of the resolution of pterygotan phylo-
geny, rigorous defi nition of extinct and modern groups,
altogether resulting in an improved knowledge of the
early evolution of a taxon that represents more than
half of the extant biodiversity, namely the winged in-
sect.
Regarding the use of cladotypic taxonomy, avoid-
ing mandatory ranks is found to be particularly re-
levant for fossil taxa, which endlessly exhibit original
character state combinations informing us of the his-
torical order of additive modifi cations that led to mod-
ern taxa. All these ‘intermediate’ taxa simply do not
fi t within a strictly ranked taxonomy, essentially based
on modern taxa, themselves assumed to be of high
rank. Based upon the practical case investigated in this

151
Arthropod Systematics & Phylogeny 65 (2)
contribution, I see no particular aspect for which the
nomenclatural system as governed by the PhyloCode
(CANTINO & DE QUEIROZ 2006), a topology-based sys-
tem (SERENO 2005), and the Linnaean one, overcome
the cladotypic approach. Importantly, the cladotypic
nomenclatural system deals with ‘ancestral’ species
/ apomorphy-less sister-species, comparatively in the
simplest way.
It is clear that a code will have to be developed
if cladotypic taxonomy becomes accepted and used.
Current International Codes of Nomenclature provide
a suitable pre-existing framework for holotypes and
cladotypes designation and curation, and regarding
starting dates for taxon names priority, among other
aspects. Rules governing the adaptation of preoccu-
pied names could be inspired from those governing
names conversion in the PhyloCode. It is clear how-
ever that some practice will be necessary before all the
aspects of cladotypic taxonomy will be unveiled, and
a suitable code developed.
6. Acknowledgments
Two anonymous reviewers provided comments on an ear-
lier version of this contribution. Their comments were
taken into account. The editorial board of Arthropod Sys-
tematics & Phylogeny provided insightful comments. I
gratefully thank S. Butts (Yale University) and R.J. Beck-
emeyer (University of Kansas) for their comments on this
contribution and improvement of language and grammar
accuracy. I also thank Ilja Kogan (Freiberg University) for
translation of parts of Russian literature. I thank Carl Bento
(Australian Museum) who photographed the specimen AM
F.36274, and R. Jones (collections manager, Australian Mu-
seum), who provided this photograph to me. I thank A.P.
Rasnitsyn for providing photographs of the specimens PIN
2240/4593 and 2240/4598. This work was supported by a
SYS-resource grant (2002) from the NHM (London). Col-
lecting of the specimen FG/596/IV/1 was supported by a
‘Deutschen Forschungsgemeinschaft’ grant, project number
VO 1466/1-1 ‘Ecosystem reconstruction of an extraordinary
lake basin – the Triassic Madygen Formation’.
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BÉTHOUX: Titanopterans are orthopterans
154
8. Appendices
Appendix 1
Taxonomic treatment in accordance to the ICZN
Taxon Archaeorthoptera Béthoux & Nel, 2002b
Sub-order Titanoptera Brongniart, 1885 / Titanoptera Sharov, 1968 ?
Super-family Tettoedischioidea Gorochov, 1987
Family Tettoedischiidae Gorochov, 1987
Subfamily Tettoedischiinae Gorochov, 1987
Genus Tettoedischia Sharov, 1968
Tettoedischia minuta Sharov, 1968
Tettoedischia elongata (Sharov, 1968) comb.n.
Super-family Tcholmanvissioidea Zalessky, 1934
Family Tcholmanvissiidae Zalessky, 1934
Genus Tcholmanvissia Zalessky, 1929
Tcholmanvissia noinskii Zalessky, 1929
Tcholmanvissia longipes (Martynov, 1940)
Family Mesotitanidae Tillyard, 1925
Sub-family Jubilaeinae subfam. n. (type-genus: Jubilaeus Sharov, 1968)
Genus Jubilaeus Sharov, 1968
Jubilaeus beybienkoi Sharov, 1968
Sub-family Mesotitaninae Tillyard, 1925
Tribe Tcholmanvissiellini trib. n. (type-genus: Tcholmanvissiella Gorochov, 1987)
Genus Tcholmanvissiella Gorochov, 1987
Tcholmanvissiella gigantea Gorochov, 1987
Tribe Mesotitanini Tillyard, 1925
Genus Mesotitan Tillyard, 1916
Mesotitan giganteus Tillyard, 1916
Mesotitan libelluloides (Sharov, 1968) comb.n.
Mesotitan ovalis (Sharov, 1968) comb.n.
Mesotitan primitivus (Sharov, 1968) comb.n.
Mesotitan superior (Sharov, 1968) comb.n.
Genus Gigatitan Sharov, 1968
Gigatitan vulgaris Sharov, 1968
Gigatitan extensus (Sharov, 1968) comb.n.
Gigatian magnifi cus (Sharov, 1968) comb.n.

155
Arthropod Systematics & Phylogeny 65 (2)
Appendix 2
Provisional taxa composition of Archaeorthoptera
Archaeorthoptera nom. Béthoux & Nel, 2002b, dis.-typ.n.: all species assigned to the Linnaean taxa Ortho-
ptera (see EADES et al. 2007; including the cladotypic taxon Pantcholmanvissiida), Caloneurodea (species listed
in BÉTHOUX et al. 2004 and RASNITSYN et al. 2004), Cnemidolestodea (Linnaean genera listed in BÉTHOUX 2005a;
see also BÉTHOUX 2007b), and:
carbonis Handlirsch, 1904: 16 (see BÉTHOUX & NEL 2004, 2005)
carpentieri Pruvost, 1919 (see BÉTHOUX 2007c)
cubitalis Handlirsch, 1911 (see BÉTHOUX 2005c)
dumasii Brongniart, 1879 (see BÉTHOUX 2003)
elongata Brongniart, 1893: 433 (see BÉTHOUX & NEL 2004, 2005)
fi scheri Brongniart, 1885 (see BÉTHOUX & NEL 2003)
lecrivaini Pruvost, 1919 (see LAURENTIAUX & LAURENTIAUX-VIEIRA 1980)
limburgica Pruvost, 1927 (see BÉTHOUX & NEL 2002a, KUKALOVÁ 1958)
macroptera van Beneden & Coemans, 1867 (see BÉTHOUX & NEL 2004, 2005)
mazonus Béthoux, 2005c
mirifi cus Carpenter & Richardson, 1971
onzii Pinto, 1990
palmiformis Bolton, 1922 (see BÉTHOUX & NEL 2004, 2005)
radialis Handlirsch, 1911 (see BURNHAM 1983)
ramosa Béthoux & Nel 2004
robusta Brongniart, 1893: 431 (see BÉTHOUX & NEL 2004, 2005)
rochacamposi Pinto & Pinto de Ornellas, 1978
ruhrensis Brauckmann & Koch, 1982 (see BRAUCKMANN et al. 1985)
sanguinettiae Pinto & Adami-Rodrigues, 1995
schneideri Béthoux, 2005c
splilopterus Handlirsch, 1911 (see BÉTHOUX 2006)
sylvatica Laurentiaux & Laurentiaux-Vieira, 1980
trecwithiensis Kukalová-Peck & Brauckmann, 1992 (see BÉTHOUX & NEL 2002a; BRAUCKMANN & HERD 2005)
vetus Scudder, 1885 (see BURNHAM 1983)
zeilleri Langiaux & Parriat, 1974
Species of uncertain validity:
- martinsnetoi Pinto in Würdig et al., 1998, velizensis Pinto & Pinto de Ornellas, 1981, amosi Pinto, 1992,
kurtzi Pinto, 1980, all probable synonyms of rochacamposi Pinto & Pinto de Ornellas, 1978
- ornellasae Pinto, 1996, a probable hind wing of rochacamposi Pinto & Pinto de Ornellas, 1978
- danielsi Handlirsch, 1906, rossae Richardson, 1956, collaris Handlirsch 1911, validum Scudder, 1885, all
probable synonyms of vetus Scudder, 1885

BÉTHOUX: Titanopterans are orthopterans
156
Appendix 3
Provisional taxa composition of Pantcholmanvissiida
Pantcholmanvissiida nom.n., dis. Béthoux & Nel, 2002a, typ.n.: all species assigned to the Tcholmanvissiidae
(see below), and:
elongata Sharov, 1968
minuta Sharov, 1968: 159
Tcholmanvissiidae nom. Zalessky, 1934, dis. Sharov, 1968, typ.n.: all species assigned to the Tcholmanti-
tanopterida (see below), and:
longipes Martynov, 1940
noinskii Zalessky, 1929
Tcholmantitanopterida nom.-dis.-typ.n.: species assigned to the Tcholmanvissiella (see below), and:
beybienkoi Sharov, 1968
Tcholmanvissiella nom. Gorochov, 1987, dis.-typ.n.: species assigned to the Titanopterida (see
below) and:
gigantea Gorochov, 1987
Titanopterida nom.-dis.-typ.n.: species assigned to the Gigatitanidae (see below) and:
giganteus Tillyard, 1916
libelluloides Sharov, 1968
ovalis Sharov, 1968
primitivus Sharov, 1968
superior Sharov, 1968
Species of uncertain validity:
- tillyardi Sharov, 1968, and similis Sharov, 1968: 197 (see also GOROCHOV 2003), all probable
synonyms of primitivus Sharov, 1968 (all diagnostic characters could be due to affi ne defor-
mation and intra-specifi c variation)
- sharovi Gorochov, 2003, probable synonym of primitivus Sharov, 1968 (all diagnostic charac-
ters could be due to affi ne deformation and intra-specifi c variation)
- zerichini Gorochov, 2003, and longispeculum Gorochov, 2003, all probable synonyms of libel-
luloides Sharov, 1968 (all diagnostic characters could be due to affi ne deformation and intra-
specifi c variation)
- reductus Gorochov, 2003, venosus Gorochov, 2003, intermedius Gorochov, 2003, latispeculum
Gorochov, 2003, bispeculum Gorochov, 2003, and modestus Gorochov, 2003, all probable
synonyms of ovalis Sharov, 1968 (all diagnostic characters could be due to affi ne deforma-
tion and intra-specifi c variation).
Gigatitanidae nom. Sharov, 1968, dis.-typ.n.:
extensus Sharov, 1968
magnifi cus Sharov, 1968
vulgaris Sharov, 1968
Species of uncertain validity:
ovatus Sharov, 1968, similis Sharov, 1968: 201, curtis Sharov, 1968, all probable synonyms
of vulgaris Sharov, 1968 (all diagnostic characters could be due to affi ne deformation and intra-
specifi c variation).