Epiproctophora BECHLY,
1996
- Included taxa: Isophlebioptera,
and Euepiproctophora.
-
Autapomorphies:
- Wing venation: costal margin not indented at
nodus, since the costal margin basal of the nodus is in
line with the costal margin distal of the nodus instead
of being curved on it; arculus shifted basally in a
position between the two primary antenodal brackets ax1
and ax2 (convergent to the coenagrionid genus
Metathemis and the Epallagidae and
Heliocharitidae); discoidal cell distally distinctly
widened in hindwing, correlated with a much less
oblique distal side MAb (transverse or even of reversed
obliquity) than in the forewing; male hindwings with
distinct anal angle (secondarily absent in some higher
Anisoptera, e.g. Anacina and Libellulidae; a secondary
absence in Zygoptera, correlated with the evolution of
a long petiolus, is not plausible, since those
Epiproctophora with a petiolus have retained the anal
angle, e.g. Stenophlebiidae); cubito-anal field
secondarily expanded in hindwings (reversed in
Epiophlebiidae and Stenophlebioptera); distinct
membranule (maybe correlated with the broadened
hindwings); secondary branching of the CuA into an
anterior longitudinal branch CuAa and a posterior
transverse branch CuAb (secondarily suppressed in
Epiophlebiidae and indistinct in
Stenophlebioptera).
- Other characters: adult males with
development of a secondary epiproctal projection as
single appendix inferior (maybe a plesiomorphy), which
is bifurcate in the groundplan (apomorphy); thorax and
abdomen are relatively strong and stout (maybe a
symplesiomorphy); globular vertex with the 3 ocelles
arranged in a "triangle" (dubious character
state, which is absent in some Anisoptera, but also
present within Zygoptera).
-
Taxonomy:
- Anisopteroides LAMEERE, 1936 ?
- ["Anisozygoptera" & Anisoptera
sensu auct.]
- [Epiprocta sensu LOHMANN, 1995
(unpublished manuscript name according to Art. 8
IRZN)]
- [partim: "Anisozygopteria"
BECHLY, 1995 (taxon nov.)]
- Epiproctophora BECHLY, 1996a (taxon nov., based on
a suggestion of Lohmann)
- Epiprocta LOHMANN, 1996 (taxon nov.)
Isophlebioptera BECHLY, 1996
- Included taxa: Euthemistidae,
Parazygoptera,
Selenothemistidae,
and Isophlebiida.
-
Autapomorphies:
- Wing venation: in hindwings the subdiscoidal
cell is enlarged and has a bulged posterior margin,
correlated with a unique course of the anal vein AA
("pseudo-anal-loop") which is strongly bent
towards the posterior wing margin at the CuP-crossing
(= anal crossing sensu Fraser) (NEL et
al., 1993); RP3/4 rather parallel to IR2, thus the
space between RP3/4 and MA is distinctly expanded and
traversed by several pektinate convex "secondary
branches" of RP3/4; postdiscoidal space between MP
and MA very narrow (NEL et al., 1993;
convergent to Erichschmidtiidae), with only one
row of cells between them in the groundplan; CuAa
shortened and postero-distally indistinct (zigzagged),
thus the distal space between MP and CuAa strongly
expanded (NEL et al., 1993); distal of ax2 all
antenodal crossveins between the costal margin and ScP
are suppressed in the groundplan (convergent to
Erichschmidtiidae and Heterophlebioidea;
apparently reversed in several species of
Isophlebioptera, and maybe plesiomorphically retained
in Euthemistidae).
- Other characters: not yet known.
-
Taxonomy:
- [partim: "Anisozygoptera"
HANDLIRSCH, 1906 (taxon nov.)]
- [partim: "Heterophlebiina"
PRITYKINA, 1980 (taxon nov., suborder, not
subtribe)]
- [Isophlebioidea sensu NEL et al.,
1993 (stat. et sens. nov.)]
- [partim: "Euryoptera" BECHLY,
1996a (taxon nov.)]
- Isophlebioptera BECHLY, 1996a (taxon nov.)
Euthemistidae PRITYKINA, 1968
(Type genus: Euthemis PRITYKINA, 1968.)
- Included taxa: only including the type
genus Euthemis PRITYKINA, 1968.
-
Autapomorphies:
- Wing venation: several long intercalary
veins between IR1 and RP1, and between IR1 and RP2, as
well as between RP3/4 and IR2, and between IR2 and RP2
(these intercalaries are parallel to the main
longitudinal veins and have no apparent origin on them,
but originate in the cross-venation); extremely narrow
postdiscoidal area (at least in the forewings since the
hindwings are not yet known).
- Other characters: not yet known.
-
Taxonomy:
- Euthemistidae PRITYKINA, 1968 (fam. nov.)
Comment: Both mentioned putative autapomorphies are also
present in Mesoepiophlebia veronicae, either
by convergence, or as synapomorphy. Furthermore the
characteristic pattern of intercalary veins is convergently
also present in Erichschmidtiidae and
Stenophlebiidae, but Euthemistidae are differing from
the former by the presence of secondary antenodal crossveins
between costal margin and ScP, and from both groups by the
lack of any intercalaries in the extremely narrow
postdiscoidal area, and by the presence of pektinate convex
"secondary branches" of RP3/4 and MP. The presence
of secondary antenodal crossveins between the costal margin
and ScP could indicate a sistergroup relationship of
Euthemistidae and all other Isophlebioptera, but on
the other hand such crossveins are also present in a few
subgroups of Isophlebioptera as obvious reversals, so
that an autapomorphic reversal cannot be totally excluded for
Euthemistidae either.
Parazygoptera BECHLY, 1997
- Included taxa: Sphenophlebiidae
and Euparazygoptera.
Triassophlebia stigmatica TILLYARD, 1922
could be a member of Parazygoptera, too, but this is
very uncertain because of the fragmentary state of the
holotype; consequently this species is better regarded as a
Stigmoptera incertae sedis.
-
Autapomorphies:
- Wing venation: RP2 arising distinctly distal
of subnodus (only reversed in Cyclothemis
sagulica PRITYKINA, 1980).
- Other characters: not yet known.
-
Taxonomy:
- Parazygoptera BECHLY, 1997i (taxon nov.)
Comment: the petiolation of the hindwing most probably
represents a plesiomorphy, since this state is also present
in Campyloptera, Triadophlebiomorpha,
Protozygoptera, Zygoptera, Tarsophlebiidae,
Epiophlebiidae, Heterophlebioptera, and
Stenophlebioptera. The secondary loss of the lestine oblique
vein in many Parazygoptera is clearly due to a
multiple convergence (since this veinlet is retained in
Mesoepiophlebia,
Ensphingophlebia, Cyclothemis and
Shurabiola, and maybe also in
Pseudotriassothemis and
Mesophlebia), probably caused by the shifting of the
origin of RP2 distal of the subnodus, as a
pre-disposition.
Sphenophlebiidae BECHLY, 1997
(Type genus: Sphenophlebia BODE, 1953.)
- Included taxa: including
Sphenophlebia interrupta BODE, 1953,
Mesoepiophlebia veronicae Nel & Henrotay in
NEL et al., 1993 and Ensphingophlebia
undulata BODE, 1953. Proeuthemis
pritykinae NEL & JARZEMBOWSKI, 1996 most probably
also belong to this new family, although its zigzagged MA
agrees with Asiopteridae. However, the similar
derived pattern of intercalary veins is here preliminarily
regarded as the stronger evidence.
-
Autapomorphies:
- Wing venation: several long intercalary
veins between IR1 and RP1, and between IR1 and RP2, as
well as between RP3/4 and IR2, and between IR2 and RP2
(convergent to Erichschmidtia and
Euthemis).
- Other characters: not yet known.
-
Taxonomy:
- Sphenophlebiidae BECHLY, 1997i (fam. nov.)
Euparazygoptera BECHLY, 1997
- Included taxa: Asiopteridae and
Triassolestoidea.
-
Autapomorphies:
- Wing venation: no antefurcal crossveins
present in the space between RP and MA basal of the
midfork (only reversed in Italophlebia
gervasuttii).
- Other characters: not yet known.
-
Taxonomy:
- Euparazygoptera BECHLY, 1997i (taxon nov.)
Asiopteridae PRITYKINA, 1968
(Type genus: Asiopteron PRITYKINA, 1968.)
- Included taxa: all genera formerly
included in Oreopteridae PRITYKINA, 1968 and
Asiopteridae sensu PRITYKINA, 1968, except
Sogjutella mollis PRITYKINA, 1980 and
"Sogdopteron" legibile
PRITYKINA, 1980 (see BECHLY, 1997i). Two further members
are "Mesoepiophlebia"
bexleyi NEL & JARZEMBOWSKI, 1996 and
Turanopteron pommerana (ANSORGE, 1996).
-
Autapomorphies:
- Wing venation: IR2 distally zigzagged
(convergent to "Sogdopteron"
legibile); MA distally zigzagged (convergent
to Steleopteridae, Mesophlebiinae, and
Italophlebia); hindwing discoidal cell
secondarily more quadrangle-like, thus with an oblique
distal side MAb (reversal; convergent to
Sogjutella mollis).
- Other characters: not yet known.
-
Taxonomy:
- Asiopteridae PRITYKINA, 1968 (fam. nov.)
- Oreopteridae PRITYKINA, 1968 (fam. nov. with type
genus Oreopteron PRITYKINA, 1968) (jun.
subj. syn. in BECHLY, 1997i)
- Oreopteridea sensu PRITYKINA, 1968
(superfam. nov.) (superfamily suffix at odds with
recommendation 29A IRZN)
- Metazygoptera sensu LOHMANN, 1981 (taxon
nov.)
- Oreopteroidea sensu NEL et al.,
1993 (nom. correct.)
Comment: "Sphenophlebia"
pommerana ANSORGE, 1996 was transferred to the genus
Turanopteron PRITYKINA, 1968 by BECHLY
(1997i) because of the nearly identical venation that was
already recognized by ANSORGE (1996). Oreopteron
asiaticum PRITYKINA, 1968 and Oreopterella
paula PRITYKINA, 1968 seem to be closely related, since
they uniquely share a zigzagged RP1, while
"Oreopteron" simile PRITYKINA,
1968 has to be transferred to the genus
Asiopteron PRITYKINA, 1968. Since
Oreopteridae and the genus Oreopteron are not
monophyletic, and Asiopteridae lacks any
autapomorphies if Oreopteridae are excluded,
Oreopteridae is here regarded as a synonym of
Asiopteridae. A detailed phylogenetic system of
Asiopteridae and several new genera and species will be
introduced in NEL et al. (in prep.). CARLE &
WIGHTON (1990: 58) suggested that Oreopteridae and
Asiopteridae are synonyms of Progonophlebiidae,
although they did not justify this hypothesis, which has to
be rejected according to BECHLY (1997i) (also see NEL et
al., 1993).
Sogjutella is certainly no member of
Asiopteridae (= Oreopteridae) since it shares the
unique kink in RA between nodal and subnodal veinlet as
strong synapomorphy with Triassolestoidea. Likewise
"Sogdopteron" legibile
shares a very complex synapomorphy in the anal area with
Triassolestini.
Triassolestoidea TILLYARD, 1918
(Type genus: Triassolestes TILLYARD,
1918.)
- Included taxa: Cyclothemistidae
and Triassolestidae.
-
Autapomorphies:
- Wing venation: nodal and subnodal veinlet
not aligned and separated by a short kink in RA (unique
within Odonata, but further transformed in
Triassolestinae).
- Other characters: not yet known.
-
Taxonomy:
- Triassolestoidea; PRITYKINA, 1980 (stat. nov.)
(nom.transl. ex Triassolestinae TILLYARD, 1918)
- [Progonophlebioidea sensu BECHLY, 1996a
(stat. nov.)]
- Triassolestoidea sensu BECHLY, 1997i
(sens. nov.)
Cyclothemistidae BECHLY, 1996
(Type genus: Cyclothemis PRITYKINA,
1980.)
- Included taxa: Cyclothemistinae
and Pseudotriassothemistinae.
-
Autapomorphies:
- Wing venation: the typical pattern of the
longitudinal veins in Isophlebioptera is
reversed (convergent to Selenothemistidae;
certainly not a symplesiomorphy), so that the
postdiscoidal space is not narrowed distally, and the
space between RP3/4 and MA is less expanded (RP3/4 is
even parallel to MA in
Cyclothemis).
- Other characters: not yet known.
-
Taxonomy:
- Cyclothemistidae BECHLY, 1996a (fam. nov.)
- Cyclothemistidae sensu BECHLY, 1997i
(sens. nov.)
Cyclothemistinae BECHLY, 1996
(Type genus: Cyclothemis PRITYKINA,
1980.)
- Included taxa: only including the two
species of Cyclothemis PRITYKINA, 1980, and
the mono-specific genera Shurabiola
PRITYKINA, 1980 and Sogjutella PRITYKINA,
1980.
-
Autapomorphies:
- Wing venation: cubito-anal area and
posterior branches of CuAa strongly reduced.
- Other characters: not yet known.
-
Taxonomy:
- Cyclothemistidae BECHLY, 1996 (fam. nov.)
- Cyclothemistinae sensu BECHLY, 1997i
(stat. nov.)
Comment: the sister-genera Shurabiola and
Sogjutella share the complete suppression of
all secondary antenodal and all antesubnodal crossveins as
putative synapomorphies. Autapomorphies of
Sogjutella are the oblique distal side (MAb) of the
discoidal cell (reversal), the suppression of the lestine
oblique vein between RP2 and IR2, and the strongly reduced
anal and cubito-anal area in the hindwing, with only one row
of cells, which is correlate with the generally reduced
venation and the slender shape of the wing.
Pseudotriassothemistinae BECHLY, 1997
(Type genus: Pseudotriassothemis BECHLY,
1997)
- Included taxa: including the former
taxa "Triassoneura"
okafujii FUJIYAMA, 1991,
"Triassothemis" nipponensis
FUJIYAMA, 1991 and
"Triassothemis" minensis
FUJIYAMA, 1991 that have all been classified in a new genus
Pseudotriassothemis by BECHLY (1997i).
-
Autapomorphies:
- Wing venation: CuP-crossing looking like an
anterior secondary branch of AA (not like a short,
straight and transverse "crossvein" but
elongated, curved, and more or less parallel to the
longitudinal veins), that is ending directly beneath
the arculus (at least in hindwings); hindwing with two
strongly curved basal posterior branches of AA (unique
within Odonatoptera); IR2 originating on RP3/4; at
least the hindwings are secondarily non-petiolated;
hindwing nodus in a very basal position.
- Other characters: not yet known.
-
Taxonomy:
- Pseudotriassothemistinae BECHLY, 1997i (subfam.
nov.)
Comment: P. okafujii and P.
nipponensis at least are sister-species (synapomorphy:
unique crossvein-brace between RP and IR2 below the subnodus)
or might even represent fore and hindwing of the same
species, but this could only be positively demonstrated if a
specimen with both pairs of wings preserved together would be
found. An autapomorphy of P. minensis seems
to be the secondary presence of secondary antenodal
crossveins between costal margin and ScP distal of ax2
(convergent to Mesophlebiinae and
Italophlebia).
Triassolestidae TILLYARD, 1918
(Type genus: Triassolestes TILLYARD,
1918.)
- Included taxa: Mesophlebiinae and
Triassolestinae.
-
Autapomorphies:
- Wing venation: in the hindwing [AA &
CuP] and [MP & CuA] are partly fused basal of the
arculus (quite unique within Odonata; currently known
from Progonophlebia,
Triassolestodes,
"Sogdopteron" legibile, and
Italophlebia); in the hindwing the
distal part free part of [AA & CuP] is not
longitudinal and fusing with CuAa but perpendicular
(aligned with arculus) and directed towards the hind
margin, so that the subdiscoidal cell is posteriorly
open (convergent to Isophlebiidae);
characteristical kink in CuA between the gaff-portion
and CuAb in the hindwing.
- Other characters: not yet known.
-
Taxonomy:
- Triassolestinae TILLYARD, 1918
- Triassolestidae; TILLYARD, 1923 (stat. nov.)
- Triassolestidae sensu PRITYKINA, 1981
(sens. nov.)
- Triassolestidae sensu BECHLY, 1997i (sens.
nov.)
- Triassoneuridae RIEK, 1976 (fam. nov. with the type
genus Triassoneura RIEK, 1976) (jun.
subj. syn. in PRITYKINA, 1981, followed by BECHLY,
1997i)
- Triassoneuridae; FUJIYAMA, 1991 (sens. nov.)
- Triassoneuridae; NEL et al., 1993 (sens.
nov.)
- Triassolestidae; NEL et al., 1993 (sens.
nov.)
Mesophlebiinae TILLYARD, 1916
(Type genus: Mesophlebia TILLYARD in TILLYARD
& DUNSTAN, 1916.)
- Included taxa: only including the two
monotypic sister-genera Mesophlebia
TILLYARD, 1916 and Progonophlebia TILLYARD,
1925.
-
Autapomorphies:
- Wing venation: nodus distal of the midwing
position since the postnodal part of the hindwing
distinctly shortened (a unique character in the
"anisozygopteroid" grade); MA distally
zigzagged (convergent to Steleopteridae,
Asiopteridae and Italophlebia);
characteristical shape of the distal furcation of RP
into RP1 and RP2 (unsafe in
Progonophlebia); RP1 and RA at least somewhat
converging beneath the pterostigma (unsafe in
Progonophlebia); secondary presence of
secondary antenodal crossveins between costal margin
and ScP distal of ax2 (convergent to
Pseudotriassothemistinae and
Italophlebia, but not certain for
Progonophlebia, thus maybe rather an
autapomorphy of Mesophlebia).
- Other characters: not yet known.
-
Taxonomy:
- Mesophlebiidae TILLYARD in TILLYARD & DUNSTAN,
1916 (fam.nov.)
- Mesophlebiinae; TILLYARD, 1917 (stat. nov.)
- Mesophlebiidae; TILLYARD, 1922
- Progonophlebiidae TILLYARD, 1925 (fam. nov. with
type genus Progonophlebia TILLYARD,
1925) (jun. subj. syn. in BECHLY, 1997i)
- Mesophlebiidae; HANDLIRSCH, 1939 (sens. nov.)
- Mesophlebiidae; PRITYKINA, 1981 (sens. nov.)
- Mesophlebiidae; JELL & DUNCAN, 1986
- Mesophlebiidae; NEL & PAICHELER, 1993 (sens.
nov.)
- Progonophlebiidae; NEL & JARZEMBOWSKI
(1997)
- Mesophlebiinae sensu BECHLY, 1997i (stat.
et sens. nov.)
Comment: Mesophlebia tillyardi HANDLIRSCH,
1939 is a junior subjective synonym of Mesophlebia
antinodalis TILLYARD, 1916 according to COWLEY (1942).
Progonophlebia cromptoni ZEUNER, 1958 (nec
"cramptoni": NEL et al., 1993) is
an junior objective synonym of Progonophlebia
woodwardi TILLYARD, 1925 according to NEL &
JARZEMBOWSKI (1997), since the two referring holotypes are
indeed part and counterpart of the same specimen.
Unique autapomorphies of Mesophlebia are an
oblique vein that is obliquely slanted towards the apex
between RP and IR2 below the transverse subnodus, the
reversed obliquity of the nodal crossvein (convergent to
Triassolestodes), and a more distinct
constriction of the space between RA and RP1 beneath the
pterostigma.
Triassolestinae TILLYARD, 1918
(Type genus: Triassolestes TILLYARD,
1918.)
- Included taxa: Triassothemistini
and Triassolestini.
-
Autapomorphies:
- Wing venation: forewing distinctly
petiolated (polarity unsafe); nodus and subnodus are
perpendicular to RA, and more or less aligned
(reversal); somewhat basal of the subnodus there is a
distinctly oblique crossvein that is obliquely slanted
towards the apex between RA and RP (most distinct in
Triassolestodes asiaticus, but also
clearly present in Italophlebia
gervasuttii and Triassothemis
mendozensis); distinctly reduced number of
antesubnodal crossveins (only 2 or 3).
- Other characters: not yet known.
-
Taxonomy:
- Triassolestinae TILLYARD, 1918 (subfam. nov.)
- Triassolestinae sensu BECHLY, 1997i (sens.
nov.)
Triassothemistini FUJIYAMA, 1991
(Type genus: Triassothemis CARPENTER,
1961.)
- Included taxa: only including the two
genera Triassothemis CARPENTER, 1961 and
Italophlebia WHALLEY, 1987. The genus
Triassothemis also includes the former taxa
"Triassoneura" heidiae
RIEK, 1976 and "Triassoneura"
regularis RIEK, 1976 that have been classified by
BECHLY (1997i) in a new subgenus
Afrotriassothemis BECHLY, 1997, while
Triassothemis mendozensis CARPENTER, 1961 was
classified in the nominate subgenus
Triassothemis CARPENTER, 1961.
-
Autapomorphies:
- Wing venation: MA and MP distinctly
converging near the hind margin; area between RP3/4 and
MA basally rather narrow for a considerable distance;
area between IR2 and RP3/4 basally very narrow, then
expanded, and distally again narrowed; area between RP2
and IR2 basally very narrow.
- Other characters: not yet known.
-
Taxonomy:
- Triassothemidae FUJIYAMA, 1991 (fam. nov.) (nom.
imperf.)
- Triassothemidae; BRIDGES, 1994
- Triassothemistini sensu BECHLY, 1997i
(stat. et sens. nov., nom. correct. et transl.)
- Italophlebiidae WHALLEY, 1986 (fam. nov. with the
type genus Italophlebia WHALLEY, 1986)
(jun. subj. syn. in BECHLY, 1997i)
Comment: distinct autapomorphies of
Italophlebia are the distally strongly converging
veins RP1 and RP2, the distally zigzagged vein MA, and the
shortened and zigzagged IR1. Autapomorphies of the new
subgenus Afrotriassothemis BECHLY, 1997 are
the numerous cells below the elongated pterostigma, very long
and straight IR1, RP2 and IR2 basally very close together
fore a longer distance, and two rows of cells between the
distal parts of IR2 and RP3/4.
Triassolestini TILLYARD, 1918
(Type genus: Triassolestes TILLYARD,
1918.)
- Included taxa: only including
Triassolestes epiophlebioides TILLYARD, 1918,
Triassoneura andersoni RIEK, 1976 and
Triassolestodes asiaticus PRITYKINA, 1981,
and probably also "Sogdopteron"
legibile PRITYKINA, 1980, which will be classified
in a new genus Sogdopterites in NEL et
al. (in prep.).
"Triassoneura" primitiva
PRITYKINA, 1981 is an Archizygoptera that seems to
be closely related to the genus Batkenia
PRITYKINA, 1981, and thus has been transferred to a new
genus Paratriassoneura BECHLY, 1997.
-
Autapomorphies:
- Wing venation: arculus secondarily shifted
closer to ax2 than to ax1, or even distal of ax2 (not
preserved in Triassolestes).
- Other characters: not yet known.
-
Taxonomy:
- Triassolestini sensu BECHLY, 1997i (stat.
et sens. nov.) (nom. transl. ex Triassolestinae
TILLYARD, 1918)
Discussion: a strong synapomorphy of all
Triassolestini, except the most basal
"Sogdopteron" legibile, is a
unique veinal pattern in the forewing base, in which [AA
& CuP] is basally fused with the hind margin, and
distally fused with [MP & CuA], only two short portions
remained free, the basal one looking like the CuP-crossing,
and the distal one as oblique "pseudo-subdiscoidal
veinlet" beneath the tip of the discoidal cell (unique
within Odonatoptera).
A distinct synapomorphy of Triassolestodes
and Triassoneura is the short basal fusion of
MP and CuA distal of the forewing discoidal cell (unique
within Odonatoptera, but similar to the hindwing of
Tarsophlebiidae). A further synapomorphy of these two genera
might be the circumstance that ax2 is secondarily shifted
basal of the arculus (reversal), but this character is
unknown in Triassolestes.
The holotypical wing of Triassolestes looks
very similar to the forewing of Triassoneura
and Triassolestodes. The only conflicting
character would be the closed discoidal cell, but according
to NEL (pers. comm.) this character state is based on an
error of Tillyard. An unusual feature (autapomorphy) of
Triassolestes is the distally widened
postdiscoidal area with more than one row of cells in the
widened part. However, such an autapomorphic reversal
happened in Cyclothemistidae, too.
Autapomorphies of Triassolestodes are the
reversed obliquity of the nodal crossvein (convergent to
Mesophlebia), a unique type of hindwing
arculus (NEL pers. comm.), and the reduced wing venation with
only one row of cells between all the longitudinal veins,
giving the wing a zygopteroid appearance. Distinct
autapomorphies of "Sogdopteron"
legibile PRITYKINA, 1980 are the basally zigzagged
RP2 and the distal obliteration of MA. The distally
converging veins MA and MP may be a derived similarity with
Triassothemistini, but the position of the arculus
closer to ax2 than to ax1 has to be regarded as stronger
evidence for a relationship with Triassolestini, since
this character state is a very rare reversal within
Epiproctophora.
Selenothemistidae HANDLIRSCH, 1939
(Type genus: Selenothemis HANDLIRSCH,
1920.)
- Included taxa: including the three
genera Selenothemis HANDLIRSCH, 1920,
Turanothemis PRITYKINA, 1968, and
Caraphlebia CARPENTER, 1969, and maybe also
Paraliassophlebia HONG, 1982.
-
Autapomorphies:
- Wing venation: in hindwings the distal side
(MAb) of the discoidal cell is about twice as long as
the basal side (= posterior part of arculus) (NEL
et al., 1993); unique shape of the hindwing
subdiscoidal cell; the typical pattern of the
longitudinal veins in Isophlebioptera is
reversed (convergent to Cyclothemistidae; might
also be plesiomorphic absent), thus the postdiscoidal
space is not narrowed and RP3/4 (RP3/4) is not parallel
to IR2; two oblique veins between RP2 and IR2 (maybe
rather an autapomorphy of
Selenothemis.
- Other characters: not yet known.
-
Taxonomy:
- Selenothemidae HANDLIRSCH, 1939 (fam. nov.) (nom.
imperf.)
- Selenothemistidae; COWLEY, 1942 (nom.
correct.)
- Turanothemistidae PRITYKINA, 1968 (fam. nov. with
the type genus Turanothemis PRITYKINA,
1968) (jun. subj. syn. in BECHLY, 1996a)
Isophlebiida BECHLY, 1996
- Included taxa: Archithemistidae
and Isophlebioidea.
-
Autapomorphies:
- Wing venation: CuAa further shortened in the
hindwing, correlated with an distally more strongly
expanded area between MP and CuAa and the development
of numerous convex parallel "secondary
branches" of MP (already indicated in the
groundplan of Isophlebioptera); pterostigmata
elongated and basally recessed (NEL et al.,
1993; convergent to Aeschnidiidae, Petaluridae
and Anacina); MA and MP distally converging (reversed
in Isophlebiidae).
- Other characters: not yet known.
-
Taxonomy:
- Isophlebiida BECHLY, 1996a (taxon nov.)
Archithemistidae TILLYARD, 1917
(Type genus: Archithemis HANDLIRSCH, 1906; =
Diastatommites TILLYARD, 1925 nom. subst.
pro. Diastatomma GIEBEL, 1856, nec
Diastatomma BURMEISTER, 1839; =
Diastatommites HANDLIRSCH, 1920, nomen nudum.)
- Included taxa: including the three
species Archithemis brodiei (GEINITZ,
1884), Archithemis liassina (STRICKLAND,
1840), and Petrophlebia anglicana TILLYARD,
1925.
-
Autapomorphies:
- Wing venation: a single row of very long and
oblique crossveins in the postdiscoidal area of the
hindwing (maybe a synapomorphy with
Isophlebioidea since also present in some
Campterophlebiidae, e.g. Sogdophlebia,
Angaroneura, and
Sibirioneura amurica); IR2 originating on
RP3/4.
- Other characters: not yet known.
-
Taxonomy:
- Architheminae TILLYARD, 1917 (subfam. nov.) (nom.
imperf.)
- Archithemidae; HANDLIRSCH, 1920 (nom.transl.)
- Archithemidae; TILLYARD, 1925
- Archithemistidae; COWLEY, 1942 (nom.
correct.)
- [Diastatommidae HANDLIRSCH, 1906 (objectively
invalid fam. nov. based on a homonym type genus)]
- Petrophlebiidae TILLYARD, 1925 (fam. nov. with type
genus Petrophlebia TILLYARD, 1925)
(jun. subj. syn. in BECHLY, 2003e)
Comment: Petrophlebia anglicana was
already correctly placed by FRASER (1957), and later
transferred to Campterophlebiidae by NEL et
al. (1993), but in both cases without explication. Thus
Petrophlebiidae is not a synonym of
Campterophlebiidae (contra NEL et al.,
1993). Although there are only few putative synapomorphies
known for Archithemistidae, this taxon very probably
is monophyletic since the three included species are so
similar that they could easily be classified in the same
genus.
Isophlebioidea HANDLIRSCH, 1906
(Type genus: Isophlebia HAGEN, 1866.)
- Included taxa: Campterophlebiidae
and Isophlebiidae, as well as
Euphaeopsis HANDLIRSCH, 1906 as genus in familia
incertae sedis.
-
Autapomorphies:
- Wing venation: hindwing subdiscoidal cell further
enlarged and transversely elongated (NEL et
al., 1993), correlated with an very long and
straight gaff (= basal CuA before its branching) (NEL
et al., 1993); in male hindwings the area of
the "anal triangle" is hypertrophied and
subdivided into numerous cells.
- Other characters: not yet known.
-
Taxonomy:
- Isophlebioidea NEL et al., 1993 (stat.
nov.) (nom. transl. ex Isophlebiidae HANDLIRSCH,
1906)
- Isophlebioidea sensu BECHLY, 1996a (sens.
nov.)
Campterophlebiidae HANDLIRSCH, 1920
(Type genus: Campterophlebia BODE, 1905.)
- Included taxa: including the genera
listed in NEL et al. (1993), except
Shurabiola PRITYKINA, 1980.
-
Autapomorphies:
- Wing venation: male hindwing with a very
acute or even hook-like projecting anal angle; the
hindwing longitudinal veins (especially RP3/4) are
distinctly undulate, and the space between MA and MP is
distally constricted by an opposite curvature of these
two veins.
- Other characters: not yet known.
-
Taxonomy:
- Campterophlebiidae HANDLIRSCH, 1920 (fam.
nov.)
- Karatawiidae MARTYNOV, 1925 (fam. nov. with type
genus Karatawia MARTYNOV, 1925)
(uncertain subjective synonymy proposed by NEL et
al., 1993)
Comment: the only Triassic (Raetian) species
Sogdophlebia singularis PRITYKINA, 1970 was
originally described in Heterophlebiidea -
Liassophlebiidae (sensu Pritykina), but transferred
to Isophlebioidea - Campterophlebiidae by NEL
et al. (1993). This placement is also endorsed by
BECHLY (1997i).
Isophlebiidae HANDLIRSCH, 1906
(Type genus: Isophlebia HAGEN, 1866.)
- Included taxa: including the genera
listed in NEL et al. (1993).
-
Autapomorphies:
- Wing venation: the primary antenodal
brackets ax1 and ax2 are hypertrophied and of distinct
and converging obliquity (NEL et al., 1993);
the anal vein AA is apparently not reaching CuA in both
pairs of wings, so that the subdiscoidal cell is
posteriorly open, only delimited by the hind margin of
the wing (NEL et al., 1993; convergent to
Triassolestidae); basal closure of discoidal
cell in forewings (NEL et al., 1993;
convergent to Caloptera, Lestida, Coenagrionomorpha,
Epiophlebiidae, and Anisopteromorpha), correlated with
a shifting of the distal side (MAb) of the discoidal
cell distinctly distal of the arculus (thus not a
plesiomorphy; contra NEL et al.,
1993); the distal side (MAb) of the discoidal cell and
the gaff (basal CuA before its furcation) are
orientated in one transverse axis and the gaff is
further prolonged, so that the basal field between MP
and CuA is twice as wide as the basal field between MA
and MP (NEL et al., 1993); CuAa is strongly
bent at the furcation of the gaff and running more or
less parallel to MP for some distance (NEL et
al., 1993); the typical pattern of the
longitudinal veins in Isophlebioptera is
reversed (probably not plesiomorphic absent), the
postdiscoidal space is somewhat less narrowed and RP3/4
is not parallel to IR2.
- Other characters: very large size,
correlated with an extremely strong thorax and abdomen;
female cerci very elongated and developed as
"accessory ovipositor blades" (the true
ovipositor is not hypertrophied, contra NEL
et al., 1993 and NEL &
MARTÍNEZ-DELCLÒS, 1993.
-
Taxonomy:
- Isophlebiidae HANDLIRSCH, 1906 (fam. nov.)
- Isophlebiinae; TILLYARD, 1917 (stat. nov.)
Euepiproctophora
BECHLY, 2003e
- Included taxa: Epiophlebiidae and
Anisopteromorpha.
-
Autapomorphies:
- Wing venation: strong "tendency"
towards a basal closure of discoidal cell in forewings,
including the development of a dorsal arcular bracket
(convergent to Caloptera, Lestida, Coenagrionomorpha,
and Isophlebiidae), but the forewing discoidal
cell is still open or incompletely closed in many
specimens of Heterophlebioptera (apparently even
variable within the same species, especially in
Liassophlebia); hindwing subdiscoidal cell at
least traversed by one crossvein, which in
Pananisoptera becomes the pseudo-anal vein PsA;
groundplan with distinct pattern of alternating width
of wing spaces between the longitudinal veins, at least
in hindwings, with broad spaces between RP1 and RP2,
between IR2 and RP3/4, between MA and MP, and between
CuA and hind margin, and with narrow spaces between RP2
and IR2, between RP3/4 and MA, and between MP and CuA
(LOHMANN, 1995, 1996; this character state is here
regarded as derived state that is correlated with the
evolution of the discoidal cell towards a discoidal
triangle).
- Other characters: the three caudal gills of
larvae secondarily absent, and replaced by complex
rectal gills; larval epiproct and paraprocts strongly
shortened and forming a so-called "anal
pyramid" together with the cerci (the anal pyramid
is still rather short and blunt in the groundplan, but
always functioning as anal valves in connection with
the improved rectal respiration); larval abdomen
shortened and distally broadened (instead of very
slender, elongate, and equilateral); larvae with
enlarged abdominal latero-sternites (E.L. SMITH,
unpubl.; somewhat dubious character with uncertain
polarity); proventriculus located anterior of the
second abdominal segment (CARLE, 1995; polarity
unclear); the posterior tergal sclerite of the
metathorax is suppressed and the anterior sclerite
enlarged (PFAU, 1986, 1991; convergent to Caloptera and
Coenagrionomorpha); adult synthorax with the dorsal
portion of the interpleural suture suppressed (retained
in all Zygoptera, except the Anisoptera-like genus
Philoganga; ASAHINA 1957; also known from the
fossil genus Paraheterophlebia, but
apparently absent in Isophlebioptera -
Parazygoptera); characteristical colour pattern of
adult thorax and abdomen in the groundplan, with
alternating yellow and dark-brown stripes (a rather
weak character of uncertain polarity, which is
furthermore mostly unknown in the fossil taxa); females
do grab the male abdomen instead of their own abdomen
with their legs during the formation of the
pairing-wheel (only a diagnostic difference to
Zygoptera, because the polarity of this character is
unknown; since Hemiphlebia shows a similar
behaviour, it might even be a symplesiomorphy); in
flight the forelegs are held perpendicular to the other
two pairs of legs ("paraflexate" position
sensu LOHMANN, 1995, 1996; only a diagnostic
difference to Zygoptera, since the polarity of this
character is unknown).
-
Taxonomy:
- [partim: "Euryoptera" BECHLY,
1996a (taxon nov.)]
- Euepiproctophora BECHLY, 2003e (taxon nov.)
Comment: the hypothesis that Epiophlebia is
closer related with Anisoptera than Isophlebioptera is
convincingly demonstrated by the mentioned larval
synapomorphies. The presence of a synlestine type of larva
(with three caudal gills of characteristical shape including
a median gill that is formed by a long epiproct; no anal
pyramid; abdomen very elongate and slender) in basal extant
Zygoptera (Hemiphlebiidae, Synlestidae) and fossil
isophlebioid larvae, as well as the oldest known odonate
larva from the Triassic of Australia, is such a distinct and
complex similarity that it cannot be reasonably regarded as
convergence, but only as a symplesiomorphy. Therefore the
strongly Anisoptera-like-larvae (with a shortened and
broadened abdomen with well-developed rectal gills and with
suppressed caudal gills) represent a very strong synapomorphy
for Epiophlebiidae and Anisoptera (but not
Isophlebioptera). The attribution of the referring fossil
larvae (e.g. "Samarura"
gigantea) from the Lower Cretaceous of Sibiria to
Isophlebioptera is absolutely certain, since FLECK and
NEL (pers. comm. 1998) could recently identify the
synapomorphic characters of the isophlebioid wing venation on
the wing sheath of one of the referring fossil larvae at PIN
in Moscow. Furthermore, the only fossil adult odonates in the
same layers are large isophlebioids, while the only occuring
fossil odonate larvae are these very large
"Samarura" larvae (length about 12.5
cm!).
FRASER (1957) proposed that a tendency towards a division of
the hindwing discoidal cell into an anterior hypertriangle
and a posterior discoidal triangle might indicate a close
relationship of Epiophlebia and Anisoptera, too.
However, the division of the hindwing discoidal cell is very
variable Epiophlebia, and the formation of a true
hypertriangle is even still variable in
Liassophlebia. Furthermore I regard the alleged
division of the hindwing discoidal cell in some specimens of
Epiophlebia as based on an erroneous interpretation
anyway, since the referring crossvein is clearly developed in
the median space, forming a second "arculus", while
the discoidal cell is never divided at all.
Epiophlebiidae
MUTTKOWSKI, 1910
(Type genus: Epiophlebia CALVERT, 1903 nom. subst.
pro Palaeophlebia SELYS, 1889, nec
Palaeophlebia BRAUER, 1889; =
Neopalaeophlebia HANDLIRSCH, 1906, jun. obj.
syn.)
- Included taxa: only including the two
relic species Epiophlebia superstes (SELYS, 1889)
and E. laidlawi TILLYARD, 1921.
-
Autapomorphies:
- Wing venation: both pairs of wings well
petiolated; terminal kink of CP and nodal furrow
reduced and nodus ventrally traversed by the costal row
of spines (NEL et al., 1993); unique double
development of the posterior part of the hindwing
arculus (NEL et al., 1993) which looks like
being formed by two closely twisted crossveins.
- Other characters: discal node
(sensu Carle, 1982) secondarily absent; male
secondary copulatory apparatus with a derived type of
vesicula spermalis, a paedomorphotic ligula, a
secondary absence of a median epression in the lamina
anterior, hook-like hamuli anteriores, and strongly
enlarged hamuli posteriores that are developed as
intromittent organs (ASAHINA, 1954; CARLE, 1995); adult head with a large
transverse crest (interocellar lobe) of the vertex
between the ocelli (ASAHINA, 1954; CARLE, 1995); adult
antenna with flattened elongate-ovoid pedicel with a
lateral hairy fringe (ASAHINA, 1954; CARLE, 1995);
unique hairy tubercle beneath the compound eye
(ASAHINA, 1954); larvae with a stridulating organ on
the abdominal segments III-VI (CARLE, 1995); male
occipital region with "anti-tandem" lobes
(CARLE, 1995); male ventral epiproctal spurs engaging
anterior lamina (CARLE, 1995); adults with a fringe of
long hairs along the ventral margin of the abdominal
tergites.
-
Taxonomy:
- [Palaeophlebinae NEEDHAM, 1903 (objectively invalid
subfam.nov. based on a homonym type genus) (nom.
imperf.)]
- [Palaeophlebiinae; JACOBSON & BIANCHI, 1905
(objectively invalid subfam.nov. based on a homonym
type genus) (nom. correct.)]
- [partim: "Anisozygoptera"
HANDLIRSCH, 1906 (taxon nov.)]
- Neopalaeophlebidae HANDLIRSCH, 1906 (fam. nov.)
(jun. obj. syn.)
- Epiophlebinae MUTTKOWSKI, 1910 (subfam. nov.) (nom.
imperf.)
- Epiophlebiinae; TILLYARD, 1917 (nom.
correct.)
- Epiophlebioidea; TILLYARD & FRASER, 1940 (stat.
nov.)
- Epiophlebioidea; ASAHINA, 1954 (sens. nov.)
- [partim: Anisoptera sensu
PRITYKINA, 1980 (sens. nov.)]
- [Epiophlebiomorpha PRITYKINA, 1980 (taxon
nov.)]
- [Epiophlebiina PRITYKINA, 1980 (taxon nov.,
suborder, not a subtribus]
- Epiophlebioidea; PRITYKINA, 1980 (sens. nov.)
- [Epiophlebioptera sensu LOHMANN, 1995
(unpublished manuscript name according to Art. 8
IRZN)]
- Epiophlebioidea; NEL et al., 1993 (sens.
nov.)
- [Epiophlebioptera BECHLY, 1996a (taxon nov.)]
- [Epiophlebioptera LOHMANN, 1996 (homonymous taxon
nov.)]
- [Pan-Epiophlebioptera LOHMANN, 1996 (trivial
name)]
Comment: according to LINDEBOOM (pers. comm.)
the secondary male genital apparatus of Epiophlebia
has several apparently plesiomorphic features that could be
regarded as evidence for a sistergroup relationship of
Epiophlebia with the remaining extant Odonata, since
Zygoptera and Anisoptera share the putative apomorphic states
(ligula detached and lifted from sternum, plate-like hamuli
anteriores, and a lamina anterior with a median depression).
However, the polarity of the latter two character states
(which are correlated anyway) is doubtful, since
Tarsophlebia, which certainly has a more basal
position than Epiophlebia, also has plate-like
hamuli anteriores and a median depression in the lamina
anterior, which is strongly suggesting a symplesiomorphy of
Tarsophlebiidae with Zygoptera and Anisoptera.
Therefore, the hook-like hamuli anteriores and the entire
lamina anterior rather have to be regarded as autapomorphic
reversals of Epiophlebia. A sistergroup relationship
of Tarsophlebia with all extant Odonata
(incl. Epiophlebia) is well-supported by the
following unique plesiomorphies of
Tarsophlebia: ligula orimentary (same shape as in
the early ontogenesis in Zygoptera); vesicula spermalis still
very short and flat with a very wide porus; hindwing
discoidal cell basaly still open (arculus incomplete); legs
clearly still with four tarsomeres of equal length. All
extant Odonata share the referring apomorphic states as
synapomorphies. Consequently, only the "primitive"
ligula (and maybe the absence of a discal node) remains as
potential evidence for a most basal position of
Epiophlebia within extant Odonata. The
Tarsophlebia-like ligula in the ontogenesis of
extant Zygoptera (e.g. Calopteryx) indicates that
the apparently plesiomorphic ligula of Epiophlebia
could as well be explained by paedomorphosis, correlated with
the secondary loss of function of the ligula by the
autapomorphic enlargement of the hamuli posteriores,
whichtake over the function as sperm intromittent organ. The
undisputet autapomorphic enlargement of the hamuli
posteriores also implies an increased need for mechanical
stability, which might also explain the secondary loss of the
median depression of the lamina anterior.
Anisopteromorpha
BECHLY, 1996
- Included taxa: Heterophlebioptera
and Trigonoptera, and
probably also Erichschmidtiidae.
-
Autapomorphies:
- Wing venation: hindwing discoidal cell
divided by a longitudinal trigonal vein into a
posterior triangulum and an anterior hypertriangulum
(still variable in Liassophlebia); [M
& Cu] distinctly bent at the arculus of both pairs
of wings (not yet in Erichschmidtiidae); male
hindwings with an perpendicular secondary posterior
branch AA2b which is running to the anal angle and
distally delimiting the anal triangle (NEL et
al., 1993; apparently still absent in
Erichschmidtiidae, and secondarily only indistinctly
developed in Stenophlebiidae) that is divided
into three cells in the groundplan.
- Other characters: not yet known.
-
Taxonomy:
- Anisopteromorpha BECHLY, 1996a (taxon nov.)
Erichschmidtiidae BECHLY, 1996
(Type genus: Erichschmidtia PRITYKINA,
1968.)
- Included taxa: only including
Erichschmidtia PRITYKINA, 1968 and
Prostenophlebia NEL &
MARTÍNEZ-DELCLÒS, 1993.
-
Autapomorphies:
- Wing venation: numerous intercalary veins
(convergent to Stenophlebiidae and
Euthemistidae) with a unique pattern, e.g. in the
apical part of the wing (numerous strictly parallel
intercalaries) and two characteristical intercalaries
in the narrow postdiscoidal space between MA and MP; MA
and MP strictly parallel, correlated with a very narrow
postdiscoidal space (convergent to most
Isophlebioptera); very oblique nodal and subnodal
veinlet (convergent to Stenophlebiidae); IR2
apparently arising on RP3/4 (convergent to
Heterophlebioidea); antenodal crossveins between costal
margin and ScP suppressed distal of ax2 (also in
Prostenophlebia, contra NEL et
al., 1993; convergent to Heterophlebioidea
and most Isophlebioptera); characteristical
pattern of small longitudinal ly elongated cells in the
distal part of the wing.
- Other characters: not yet known.
-
Taxonomy:
- [partim: "Heterophlebiina"
PRITYKINA, 1980 (taxon nov., suborder, not
subtribe)]
- Erichschmidtiidae BECHLY, 1996a (fam. nov.)
Comment: Trigonophlebia shares with
Erichschmidtia a rather unique cell below the
subdiscoidal cell, convergent to Tarsophlebiopsis
mayi, but does have secondary antenodal crossveins
between costal margin and ScP that would conflict a
sistergroup relationship of these two genera. Most probably
the Erichschmidtiidae represent the sistergroup of all
other Anisopteromorpha, although it cannot be totally
excluded that they might either belong to
Heterophlebioptera, or even to Isophlebioptera, e.g.
as possible sistergroup of Euthemistidae.
Heterophlebioptera BECHLY, 1996
- Included taxa: Myopophlebiidae
and Heterophlebioidea.
-
Autapomorphies:
- Wing venation: unique unicellular anal loop
(NEL et al., 1993) which lies beneath the
subdiscoidal cell and is ventrally closed by CuAb that
is parallel to AA and thus directed towards the wing
base instead of the posterior wing margin (probably not
homologous with the anisopterid anal loop); forewings
shortly petiolated (NEL et al., 1993);
subdiscoidal cell of the hindwing with a convex curved
or angulated posterior margin (NEL et al.,
1993); unique shape of the forewing discoidal cell that
is very transverse and narrow; strong
"tendency" towards the development of a
second incomplete arcular veinlet in the hindwings, and
a single incomplete arcular veinlet in the forewings
(but the basal closure of the forewing discoidal cell
is still very variable, in many specimens completely
open or completely closed).
- Other characters: not yet known.
-
Taxonomy:
- [partim: "Anisozygoptera"
HANDLIRSCH, 1906 (taxon nov.)]
- [partim: "Heterophlebiina"
PRITYKINA, 1980 (taxon nov., suborder, not
subtribe)]
- [Heterophlebiomorpha PRITYKINA, 1981 (taxon
nov.)]
- [Heterophlebioidea sensu NEL et
al., 1993]
- Heterophlebioptera BECHLY, 1996a (taxon nov.)
Myopophlebiidae BODE, 1953
(Type genus: Myopophlebia BODE, 1953.)
- Included taxa: including the genera
listed in NEL et al. (1993).
-
Autapomorphies:
- Wing venation: not yet known.
- Other characters: not yet known.
-
Taxonomy:
- Myopophlebiidae BODE, 1953 (fam. nov.)
- [Paraheterophlebiidae NEL, 1991 (unpublished
manuscript name according to Art. 8 IRZN)]
Heterophlebioidea NEEDHAM, 1903
(Type genus: Heterophlebia BRODIE, 1849.)
- Included taxa: Liassophlebiidae
and Heterophlebiidae.
-
Autapomorphies:
- Wing venation: antenodal crossveins between
costal margin and ScP suppressed distal of ax2
(convergent to Erichschmidtiidae and most
Isophlebioptera); IR2 arising on RP3/4 (convergent to
Erichschmidtiidae, but maybe a convergence since
the IR2 originates on RP1/2 in the liassophlebiid genus
Grimmenopteron); in the forewing the
distal side (MAb) of the discoidal cell is shifted
somewhat distally of the arculus (apomorphic state;
contra NEL et al., 1993; but maybe a
convergence since absent in the liassophlebiid genus
Grimmenopteron).
- Other characters: not yet known.
-
Taxonomy:
- Heterophlebinae NEEDHAM, 1903 (subfam. nov.)
- Heterophlebioidea TILLYARD & FRASER, 1940
(stat. nov.) (nom. transl. ex Heterophebiidae
HANDLIRSCH, 1906)
- Heterophlebioidea; ASAHINA, 1954 (sens. nov.)
- Heterophlebiidea; PRITYKINA, 1968 (superfamily
suffix at odds with recommendation 29A IRZN)
- Heterophlebioidea sensu BECHLY, 1996a
(sens. nov.)
Liassophlebiidae TILLYARD, 1925
(Type genus: Liassophlebia TILLYARD,
1925.)
- Included taxa: including the genera
listed in NEL et al. (1993).
-
Autapomorphies:
- Wing venation: forewing subdiscoidal cell
widened, with a convex posterior margin (NEL et
al., 1993); hindwings with the unicellular anal
loop enlarged (NEL et al., 1993); no
antefurcal crossveins between RP and MA from the
arculus to the midfork (convergent to the myopophlebiid
genera Myopophlebia and
Paraheterophlebia).
- Other characters: not yet known.
-
Taxonomy:
- Liassophlebiidae TILLYARD, 1925 (fam. nov.)
Comment: the genus Grimmenopteron which is
only known by a single forewing, shares the two respective
autapomorphies of Liassophlebiidae, but on the other
hand has retained two unique plesiomorphies within
Heterophlebioidea. The very large size of most
Liassophlebiidae might represent an ingroup synapomorphy for
the referring species. Therefore the smaller size of
Grimmenopteron could rather be a plesiomorphy than
an autapomorphy (contra ANSORGE, 1996).
Heterophlebiidae NEEDHAM, 1903
(Type genus: Heterophlebia BRODIE, 1849.)
- Included taxa: including the genera
listed in NEL et al. (1993), except
Erichschmidtia PRITYKINA, 1968.
-
Autapomorphies:
- Wing venation: hindwings with the
hypertriangle somewhat elongated and narrowed (NEL
et al., 1993).
- Other characters: not yet known.
-
Taxonomy:
- Heterophlebinae NEEDHAM, 1903 (subfam. nov.)
- Heterophlebiidae HANDLIRSCH, 1906 (fam.nov.) (jun.
obj. syn.)
- Heterophlebiinae; TILLYARD, 1917
Trigonoptera BECHLY,
1996
- Included taxa: Stenophlebioptera
and Pananisoptera.
-
Autapomorphies:
- Wing venation: the forewing discoidal cell
is divided by a longitudinal trigonal vein in an
anterior hypertriangulum (= supratriangular cell) and a
posterior triangulum (= trigonal cell) which is still
quadrangular in the groundplan; the forewing
subdiscoidal cell is traversed by at least one
crossvein, too, which later becomes the pseudo-anal
vein PsA (in Pananisoptera); width of wing spaces
between the longitudinal veins very strongly
alternating in both wings pairs (broad spaces between
RP1 and RP2, between IR2 and RP3/4, between MA and MP,
and between CuA and hind margin; narrow spaces between
RP2 and IR2, between RP3/4 and MA, and between MP and
CuA) (LOHMANN, 1995, 1996; this character state is here
regarded as derived state that is correlated with the
evolution of the discoidal triangles).
- Other characters: compound eyes greatly
enlarged and rather close to each other; larval
paraprocts acute (still lobate in Epiophlebia;
ASAHINA, 1954).
-
Taxonomy:
- Trigonoptera BECHLY, 1996a (taxon nov.)
Stenophlebioptera BECHLY, 1996
- Included taxa: Gondvanogomphidae
and Stenophlebiidae.
-
Autapomorphies:
- Wing venation: IR2 and RP3/4 arising close
together, correlated with a very long and narrow bridge
space; base of RP2 not strictly aligned with subnodus
(more probably a plesiomorphy); cubito-anal field of
hindwings reduced, thus both wings of similar
shape.
- Other characters: not yet known.
-
Taxonomy:
- [Stenophlebioidea sensu NEL et
al., 1993]
- Stenophlebioptera BECHLY, 1996a (taxon nov.)
Gondvanogomphidae BECHLY, 1996
(Type genus: Gondvanogomphus SCHLÜTER
& HARTUNG, 1982.)
- Included taxa: only including the
fosil species Gondvanogomphus SCHLÜTER
& HARTUNG, 1982.
-
Autapomorphies:
- Wing venation: CuA without any posterior
branches in both pairs of wings; extremely small size
of wings (length less than 1,85 cm).
- Other characters: very small body size.
-
Taxonomy:
- Gondvanogomphidae BECHLY, 1996a (fam. nov.)
Stenophlebiidae NEEDHAM, 1903
(Type genus: Stenophlebia HAGEN, 1866.)
- Included taxa:
Stenophlebia HAGEN, 1866 and probably also
Sinostenophlebia HONG, 1984, but not
Prostenophlebia NEL &
MARTÍNEZ-DELCLÒS, 1993.
-
Autapomorphies:
- Wing venation: wings very long and slender;
nodal and subnodal veinlet extremely oblique (NEL
et al., 1993); stenophlebiid oblique vein
between RP1 and RP2; nodal furrow reduced (convergent
to Epiophlebiidae and Aeschnidiidae); discoidal
triangles of unique and similar shape in both pairs of
wings (transversely elongated, narrow, and strictly
triangular); hindwings with a short but distinct
petiolus (NEL et al., 1993); numerous
intercalary veins with a characteristical pattern, e.g.
between MA and MP (convergent to
Erichschmidtiidae); basal accessory antenodal
crossveins present between ax0 and ax1; pterostigma
shifted basally (convergent to Isophlebiidae,
Aeschnidiidae, and Petaluridae).
- Other characters: terminal part of male
abdomen dilated (NEL et al., 1993; convergent
to certain Liassophlebiidae, many Gomphidae and
some Libellulidae); female ovipositor strongly
reduced.
-
Taxonomy:
- Stenophlebinae NEEDHAM, 1903 (subfam.nov.) (nom.
imperf.)
- Stenophlebiidae sensu HANDLIRSCH, 1906
(stat. nov. et nom. correct.)
- [partim: "Anisozygoptera"
HANDLIRSCH, 1906 (taxon nov.)]
- Stenophlebiinae sensu TILLYARD, 1917
- [partim: "Heterophlebiina"
PRITYKINA, 1980 (taxon nov., suborder, not
subtribe)]
- Stenophlebioidea sensu PRITYKINA, 1980
(stat. nov.)
- Stenophlebidae sensu ROSS &
JARZEMBOWSKI in BENTON, 1993 (nom. imperf.)
- Stenophlebioidea sensu NEL et
al., 1993
Comment: not including the genus
Prostenophlebia that is most probably closely
related to Erichschmidtia, contra NEL et
al. (1993).