Aeshnoptera BECHLY,
1996
- Included taxa: Mesuropetalidae
and Aeshnomorpha, and
Cymatophlebiella euryptera PRITYKINA, 1968
as basal taxon incertae sedis.
-
Autapomorphies:
- Wing venation: RP1 and RP2 basally parallel
up to the pterostigma, thus the area between these two
veins is basally distinctly narrowed with only one row
of cells between them in the groundplan (convergent to
many Gomphides-Lindeniidae like the genus
Cacoides; reversed in Aeschnopsis
tischlingeri and A. jurassica,
Archipetaliidae, Gobiaeshna occulta,
Valdaeshninae, Rudiaeschnidae, and some
fossil Gomphaeschninae, e.g.
Alloaeschna and
Progomphaeschnaoides,
Plesigomphaeschnaoides, and
Gomphaeschnaoides; contra LOHMANN,
1996: 362); at least a weakly defined (zigzagged) Rspl
is present; RP3/4 and MA more or less undulating
(convergent to Uropetala and a few
Eurypalpida, e.g. Macromiidae, Idomacromia,
Aeschnosoma & Libellulosoma;
reversed in Aeschnopsis tischlingeri
and A. jurassica, Archipetaliidae,
Valdaeshna and Aeshnodea).
- Other characters: abdominal terga with a
medio-dorso-longitudinal fold or keel (secondarily
reduced or suppressed within Austropetaliida and
Gomphaeschnidae); compound eyes closely approximated or
even contiguous.
-
Taxonomy:
- [partim: "Dynamophlebiae"
BUCHECKER, 1876 (taxon nov.)]
- Aeshnoptera BECHLY, 1996a (taxon nov.) (nec
Aeshnoptera CRAMPTON, 1928)
- Aeshnoptera sensu BECHLY et al.,
2001
Comment: numerous characters within Anisoptera and
Aeshnoptera are very homoplastic:
(1) Pterostigmal brace vein: in the groundplan of Anisoptera
the pterostigma is well-braced by an oblique crossvein
aligned with its basal side. This pterostigmal brace vein is
more transverse and distally displaced in
Paraliupanshania turoniensis gen. et sp. nov. and in
most Austropetaliida. It is also rather transverse in
Progobiaeshnidae , but still aligned with the basal
side of the pterostigma. In Prohoyaeshna and
Hoyaeshna, the pterostigmal brace vein is recessed
basal of the pterostigma, convergent to Hypopetalia
and the groundplan of Petalurida (e.g.
Protolindenia). It is somewhat weakly developed in
Cymatophlebiella, and seems to be completely
reduced in Paraliupanshania britannica and
Brachytron.
(2) Area between RP1 and RP2: in the groundplan of
Aeshnoptera, RP1 and RP2 are basally closely parallel or even
converging near the pterostigma with only one row of cells
basal of the pterostigma (preserved in most
Mesuropetaloidea and most Euaeshnida including
Eumorbaeschnidae and Gomphaeschnidae). Two (or more) rows of
cells basal of the pterostigma are (secondarily) present in
Cymatophlebiella, Aeschnopsis
perkinsi sp. and probably A.
tischlingeri, Liupanshania,
Austropetaliida, Progobiaeshnidae,
Cymatophlebioidea, Paracymatophlebiidae, and some
basal Aeshnodea (Baissaeshna, Allopetalia,
Boyeria, Petaliaeschna and
Cephalaeschna, but not in Basiaeschna). In
some, but not all, of these taxa, the secondary increase of
cell rows is correlated with a secondarily divergent course
of RP1 and RP2 (e.g. Aeschnopsis perkinsi and
A. tischlingeri,
Liupanshania, Archipetaliidae, Gobiaeshna
occulta, Valdaeshninae, and
Rudiaeschnidae). A slight secondary divergence of RP1 and RP2
is also present in Baissaeshna,
Petaliaeschna and Cephalaeschna, and even
in some of those Neoaeshnida that still have only one row of
cells between RP1 and RP2 basal of the pterostigma (e.g.
Alloaeschna, Progomphaeschnaoides,
Plesigomphaeschnaoides, and
Gomphaeschnaoides).
(3) Course of RP2 and IR2: RP2 is undulating in
Paramesuropetala and
Paraliupanshania, and in most Aeshnomorpha, although
only weakly so in Austropetaliida and Rudiaeschnidae.
The undulation of RP2 is most strongly developed in
Cymatophlebiidae (especially Cymatophlebiinae),
Eumorbaeschnidae, and some Gomphaeschnidae (e.g.
Paramorbaeschna and Linaeschna). Instead of
this undulation there is a characteristical curvature of RP2
beneath the pterostigmal brace in Mesuropetalidae and
Aeshnodea (reversal). IR2 is strongly undulating only in
Cymatophlebiidae. Except for their distal parts, RP2
and IR2 are more or less parallel in the groundplan of
Anisoptera, while they are distinctly non-parallel in
Rudiaeschnidae and Euaeshnida. The apparently parallel course
of RP2 and IR2 in Euaeshnodea is caused by a secondary
branch of IR2 that is developed in the area between RP2 and
IR2 (secondarily reduced in Boyeria and
Oplonaeschna). In Mesuropetalidae, RP2 and
IR2 are more closely parallel than in the groundplan, even
converging near the posterior wing margin.
(4) Lestine oblique vein: in the groundplan of Anisoptera
there are two oblique veins between RP2 and IR2. This state
is preserved Aeschnidiidae and Petalurida, and within
Aeshnoptera in Cymatophlebiella,
Mesuropetalidae, Cymatophlebioidea, and at least some
specimens of Eumorbaeschnidae. The basal true lestine
oblique vein is only reduced in Austropetaliida, while the
distal accessory oblique vein is convergently reduced in
Liupanshaniidae, Paracymatophlebiidae, some
specimens of Eumorbaeschnidae, and all Neoaeshnida
(convergent to all Exophytica).
(5) Rspl: an at least weakly defined Rspl belongs to the
derived groundplan characters of Aeshnoptera. In most
Mesuropetaloidea, it is still a concave, but zigzagged,
pseudo-Rspl. A well-defined Rspl is present in
Paraliupanshania and both pairs of wings of
Panaeshnida, convergent to Aeschnidiidae and most
Eurypalpida (except Synthemistidae and Gomphomacromiidae).
The Rspl is strongly curved with several rows of cells
between it and IR2 in Paraliupanshania,
Cymatophlebioidea (especially
Cymatophlebiidae), and Aeshnidae (including
Oplonaeschna), convergent to Aeschnidiidae
and some Libellulidae. The Rspl is strictly parallel to IR2
with only one row of cells between these two veins in most
Mesuropetaloidea (except
Paramesuropetala and
Paraliupanshania) and most of the basal taxa of
Euaeshnida (e.g. Eumorbaeschnidae and most
Gomphaeschnidae, Brachytronidae, and Telephlebiidae). It is
more or less parallel to IR2, but with two or three rows of
cells between these two veins in
Cymatophlebiella, Austropetaliida,
Progobiaeshnidae, Paracymatophlebiidae, and a few
basal taxa of Neoaeshnida (e.g. Allopetaliidae).
(6) Mspl: a well-defined Mspl is present by convergence in
Paraliupanshania, some
Cymatophlebiinae, and all Paneuaeshnida, convergent to
Aeschnidiidae and higher Eurypalpida (Corduliidae,
Macrodiplacidae, and Libellulidae). It is still somewhat less
well-developed in Paracymatophlebiidae and
Eumorbaeschnidae, but very well-defined in all Neoaeshnida.
In Paracymatophlebiidae and Eumorbaeschnidae
the course of Mspl is somewhat irregular with up to two rows
of cells between Mspl and MA, while in the groundplan of
Neoaeshnida (Gomphaeschnidae, Brachytronidae, and
Telephlebiidae) the Mspl is parallel to MA with only one row
of cells between these two veins (two rows in
Allopetaliidae). The Mspl is strongly curved with several
rows of cells between it and MA only in Aeshnidae (including
Epiaeschna and Oplonaeschna), convergent to
Aeschnidiidae and a few Libellulidae.
(7) Course of RP3/4 and MA: in the groundplan RP3/4 and MA
are distally diverging, while they are closely parallel up to
the wing margin in Mesuropetaloidea,
Valdaeshninae, and Euaeshnida (triple convergence).
Furthermore, in the groundplan of Aeshnoptera, RP3/4 and MA
are more or less undulating (convergent to the petalurid
genus Uropetala and a few "corduliine"
Eurypalpida, e.g. Macromiidae, Idomacromia,
Aeschnosoma and Libellulosoma). This
condition is convergently strongly reduced or completely
suppressed (reversals) in Aeschnopsis (except
the type species), Archipetaliidae,
Valdaeshna, and all Aeshnodea. A stronger undulation
is convergently present in Cymatophlebiidae,
Paracymatophlebiidae, and Eumorbaeschnidae.
(8) Postdiscoidal area: the presence of at least three rows
of cells in the basal part of the postdiscoidal area of both
pairs of wings seems to be a symplesiomorphy of many basal
Aeshnoptera with Liassogomphidae, Aeschnidiidae
and Petalurida that is reversed to two rows in Exophytica,
Aeschnopsis perkinsi and A
jurassica (and in the hindwing of
Mesuropetala muensteri),
Araripeliupanshania, Austropetaliida, and in the
groundplan of Neoaeshnida (retained in Gomphaeschnidae,
Brachytronidae and Telephlebiidae, but again reversed to
three rows of cells in Aeshnidae). The presence of three rows
of cells in a few Lindeniidae (e.g. Cacoides and
Melanocacus) and many Libellulidae (except
Tetrathemistinae) probably is due to reversal, too. More than
three rows of cells are present in
Paraliupanshania, most Cymatophlebiinae
(except Cymatophlebia purbeckensis,
C. pumilio, and C. herrlenae),
Prohoyaeshna, Rudiaeschna
limnobia, and very few Aeshnidae with very dense wing
venation (e.g. Heliaeschna).
(9) MAb and trigonal planate: in the groundplan of Anisoptera
the distal side MAb of the discoidal triangle is straight,
and there is no convex secondary vein (trigonal planate)
originating on MAb in the basal postdiscoidal area. An angled
MAb and trigonal planate is present by convergence in
Liupanshaniidae, Valdaeshna, and Neoaeshnida
(slightly indicated already in Eumorbaeschnidae), as
well as in Cretapetaluridae and in many Gomphides
(e.g. Hageniidae and Lindeniidae).
(10) Discoidal triangle: in the groundplan of Anisoptera the
forewing discoidal triangle is distinctly more transverse
than the hindwing discoidal triangle which is more or less
equilateral. This is indicated by the condition in
Aeschnidiidae, Liassogomphidae, Petalurida,
Mesuropetaloidea, Gomphides, while the referring state in
Eurypalpida seems to be due to a reversal. Distinctly
longitudinal elongate discoidal triangles in both wings
represent a convergence of Aeshnomorpha taxon nov. and
Cavilabiata (reversed in the hindwing of Chlorogomphidae and
the forewing of Eurypalpida). A discoidal triangle that is
divided into two cells by a single crossvein in both pairs of
wings seems to represent the plesiomorphic state within
Anisoptera, while unicellular discoidal triangles or
multicellular discoidal triangles represent alternative
apomorphic states that have been realized by multiple
convergence (e.g. the unicellular hindwing discoidal triangle
of Mesuropetala muensteri, or the
multicellular discoidal triangles of Liupanshaniidae,
Cymatophlebiella euryptera, Hypopetalia
pestilens, and Panaeshnida, reversed in Gomphaeschnidae
except the most basal genus Oligoaeschna).
(11) Hypertriangle: the number of crossveins in the
hypertriangle is very homoplastic within Anisoptera, but the
most parsimonious interpretation suggests that an unicellular
hypertriangle belongs to the groundplan of Anisoptera and
Aeshnoptera (retained in Mesuropetaloidea), while a
two-celled hypertriangle belongs to the groundplan of
Aeshnomorpha, and a multicellular hypertriangle belongs to
the groundplan of Panaeshnida (probably autapomorphies of the
two mentioned groups), reversed in Cymatophlebia
purbeckensis and C. pumilio,
Paracymatophlebiidae, Gomphaeschnidae (except the most basal
genus Oligoaeschna), and a few other taxa (maybe
including Eumorbaeschna jurassica ?).
(12) Subdiscoidal triangle: the number of cells in the
subdiscoidal triangle is a very homoplastic character within
Anisoptera. The two- or three-celled forewing subdiscoidal
triangles are most likely a symplesiomorphy within Anisoptera
(e.g. Liassogomphidae, Aeschnidiidae,
Petalurida and basal Gomphides - Lindeniidae), correlated
with the plesiomorphic retained transverse shape of the
forewing discoidal triangle in all these taxa. Since
Cordulegastrida, Hemeroscopidae, Chlorogomphida, and
Neopetaliidae have forewings with a longitudinal triangle and
unicellular subdiscoidal triangle, these states seem to be
derived groundplan characters of Cavilabiata that are
reversed in Eurypalpida (= Libelluloidea sensu
FRASER, 1957) which again have transverse forewing discoidal
triangles (reversal) and partly also divided forewing
subdiscoidal triangles. Within Aeshnoptera the divided
forewing subdiscoidal triangles have been reduced
Austropetaliida (reversed in Hypopetalia pestilens),
Valdaeshna surreyensis, and in the groundplan
of Euaeshnida. Contrary to the forewings, the subdiscoidal
triangles of the hindwings probably have been unicellular in
the groundplan of Anisoptera and Aeshnoptera, and became
secondarily subdivided into two or three cells by convergence
in Cymatophlebiella euryptera,
Progobiaeshna liaoningensis, Hypopetalia
pestilens, and Cymatophlebioidea (reversed in
Valdaeshna surreyensis),
Eumorbaeschnidae, and Aeshnidae. However, this hypothesis is
rather uncertain, because this character is very homoplastic,
and the alternative hypothesis of a divided hindwing
subdiscoidal triangle in the groundplan of Panaeshnida
(convergent to Cymatophlebiella and
Hypopetalia; reversed in Valdaeshna,
Gomphaeschnidae, Brachytronidae and Telephlebiidae) is nearly
equally parsimonious.
(13) Cubito-anal crossveins: in the groundplan of Anisoptera
there are no accessory cubito-anal crossveins between
CuP-crossing and PsA. They are still absent in
Cymatophlebiella, Mesuropetaloidea (except in
Aeschnopsis tischlingeri and
Liupanshania) and Austropetaliida. Such crossveins
are present in Aeschnopsis tischlingeri,
Liupanshania and in most Panaeshnida
(probably synapomorphy), but absent (reduced) in
Cymatophlebia pumilio and C.
herrlenae, Paracymatophlebiidae, Gomphaeschnidae
(except in the forewing of Oligoaeschna venusta),
Brachytron, and maybe in
Eumorbaeschna.
(14) PsA: a very well-defined pseudo-anal vein PsA is
developed in both pairs of wings of Aeschnidiidae, the
forewing of Petalurida, the forewing of
Mesuropetaloidea, both pairs of wings of
Aeschnopsis, Austropetaliida and
Cymatophlebiinae, both pairs of wings of Gomphides, and in
the forewing of Eurypalpida. This character is obviously
correlated with a transverse shape of the discoidal triangle
and a well-defined subdiscoidal triangle. Therefore, PsA is
more or less reduced towards a normal cubito-anal crossvein
in most wings with longitudinal elongate discoidal triangles,
such as the hindwing of Liupanshaniidae, both pairs of
wings of many Paneuaeshnida, Cordulegastrida, Chlorogomphida,
Neopetaliidae, and the hindwings of Eurypalpida. Within
Aeshnoptera the strongest reduction is in the hindwings of
Liupanshaniidae, and in both pairs of wings of
Valdaeshninae and Aeshnodea which have the most elongated
discoidal triangles. One of the rare exceptions from this
rule is the hindwing of Aeschnopsis perampla
(= Cymatophlebiopsis pseudobubas) which has a
very elongate and narrow discoidal triangle, but also a very
distinct PsA.
(15) Anal loop: in the groundplan of crown-group Anisoptera
(thus excluding the basal taxa Aeschnidiidae and
Liassogomphidae) there is a well-defined anal loop
that is posteriorly well-closed and divided into four to six
cells, and there is a rather short gaff. This is suggest by
the well-defined anal loop in many Petalurida, most
Aeshnoptera, most Gomphides, and nearly all Cavilabiata. In
Cymatophlebiella and in the groundplan of
Aeshnomorpha the gaff is slightly prolonged, although the
anal loop is not enlarged. The anal loop is distinctly
enlarged in Progobiaeshna liaoningensis,
Hoyaeshna, Rudiaeschnidae, and
Euaeshnida (especially in Gomphaeschnaoides
petersi and in all Aeshnidae), correlated with a strong
prolongation of the gaff. The anal loop is completely reduced
in Cymatophlebiella, many specimens of
Cymatophlebia, and most Liupanshaniidae
(except Araripeliupanshania). In
Paracymatophlebiidae the anal loop is still distinct, but the
crossvein that is forming its posterior margin is not very
strong. A unique very narrow and very longitudinal elongate
anal loop is present in Mesuropetalidae (only
Cretapetaluridae and Cordulagomphinae have a somewhat
similar structure, but their anal loop is still distinctly
different).
Some of these characters, e.g. the development of a better
defined and curved Rspl and Mspl, are so often convergently
realized within Aeshnoptera that they could even be regarded
as an underlying synapomorphy (sensu Saether) or as
a trend / tendency (sensu Brundin). Anyway, the
phylogenetic hypotheses that are built on these homoplastic
characters are only supported by relatively weak evidence.
However, there is no other choice than using these weak
characters as far as possible, since such wing venational
characters mostly are the only available and usable
characters for fossil dragonflies.
Cymatophlebiella
PRITYKINA, 1968
(Type species: Cymatophlebiella euryptera
PRITYKINA, 1968.)
- Included taxa: only including the type
species Cymatophlebiella euryptera
PRITYKINA, 1968.
-
Autapomorphies:
- Wing venation: pterostigma very short and
weakly braced; discoidal triangles three-celled;
subdiscoidal triangle two-celled; anal loop completely
reduced (correlated with a secondarily straight course
of CuAb); two rows of cells between RP1 and RP2.
- Other characters: not yet known.
-
Taxonomy:
- Cymatophlebiella PRITYKINA, 1968 (gen.
nov.)
- Cymatophlebiella sensu BECHLY
et al., 2001 (pos. nov.)
Comment: BECHLY et al. (2001) excluded
Cymatophlebiella from Cymatophlebiidae and
considered it as a basal Aeshnoptera incertae sedis
that might eventually be closer related to Panaeshnida.
Mesuropetaloidea BECHLY, 1996
(Type genus: Mesuropetala HANDLIRSCH,
1906.)
- Included taxa: Mesuropetalidae
and Liupanshaniidae.
-
Autapomorphies:
- Wing venation: arculus shifted very close to
the first primary antenodal ax1; RP3/4 and MA closely
parallel up to the wing margin in both pairs of
wings.
- Other characters: not yet known.
-
Taxonomy:
- Mesuropetaloidea BECHLY et al., 2001
(stat. nov.) (nom. transl.ex Mesuropetalidae BECHLY,
1996a)
Mesuropetalidae BECHLY, 1996
(Type genus: Mesuropetala HANDLIRSCH,
1906.)
- Included taxa: only including the
genus Mesuropetala HANDLIRSCH, 1906 and
Aeschnopsis HANDLIRSCH, 1939 (=
Cymatophlebiopsis HANDLIRSCH, 1939), which both
include several new species that are described in BECHLY
et al. (2001).
-
Autapomorphies:
- Wing venation: RP2 and IR2 very closely
parallel, even converging near the posterior wing
margin; characteristical shape of the discoidal
triangles, with the forewing discoidal triangle being
even more transverse than in the groundplan of
Anisoptera; anal loop longitudinal elongate (convergent
to Cretapetaluridae).
- Other characters: foliate superior
appendages (cerci) (convergent to
Cymatophlebiinae, Polycanthagynini incl.
"Aeschna" petalura, and
Petalurinae, but only known from
Mesuropetala muensteri.) ().
-
Taxonomy:
- Mesuropetalidae BECHLY, 1996a (fam. nov.)
- Mesuropetalidae sensu BECHLY et
al., 2001 (description of several new
species)
Comment: according to BECHLY et al. (2001)
the valid name for Mesuropetala koehleri
(HAGEN, 1848) is Mesuropetala muensteri
(GERMAR, 1839) since both holotypes clearly are conspecific.
Since Aeschna muensteri GERMAR, 1839 is the
type species of Morbaeschna NEEDHAM, 1907,
the latter genus has to be regarded as junior subjective
synonym of Mesuropetala HANDLIRSCH, 1906.
Consequently, Needham's new aeshnid which is not
conspecific with Mesuropetala muensteri, has
been classified as Eumorbaeschna jurassica
(CARPENTER, 1932) in a new genus by BECHLY et al.
(2001).
BECHLY (1996a) and NEL et al. (1998) already
demonstrated that Mesuropetala does not share
any strong synapomorphies with Petalurida, while it does
share several derived character states with the other
Aeshnoptera.
Liupanshaniidae BECHLY et al., 2001
(Type genus: Liupanshania HONG, 1982.)
- Included taxa: only including the
fossil genera Liupanshania HONG, 1982,
Araripeliupanshania BECHLY et al.,
2001, Paramesuropetala BECHLY et
al., 2001, and Paraliupanshania
BECHLY et al., 2001.
-
Autapomorphies:
- Wing venation: unique shape of the very
elongate and narrow hindwing discoidal triangle
(anterior side of discoidal triangle distally curved
and ending on the anterior side MA of the
hypertriangle; MAb is strongly sigmoidally curved, with
a very concave basal part and a strong angle in the
distal part); hindwing discoidal triangle divided into
several (at least three) cells by parallel crossveins;
forewing discoidal triangle divided into three cells
(but only known in Araripeliupanshania
and Paramesuropetala); both pairs of
wings (but especially the hindwing) with a strong
convex secondary longitudinal vein (trigonal planate)
in the postdiscoidal area, originating at the angle of
MAb (convergent to several other groups of Aeshnoptera
and Gomphides); the distal accessory oblique vein
between RP2 and IR2 is secondarily absent.
- Other characters: not yet known.
-
Taxonomy:
- Liupanshaniidae BECHLY et al., 2001
(fam. nov.)
Comment: BECHLY et al. (2001) demonstrated
that the few derived similarities with Aeshnomorpha and
Panaeshnida are better interpreted as convergences.
Aeshnomorpha BECHLY
et al., 2001
- Included taxa: Austropetaliida
and Panaeshnida.
-
Autapomorphies:
- Wing venation: forewing discoidal triangle
longitudinal elongate, too, therefore the discoidal
triangles of both pairs of wings are of similar shape;
hypertriangles divided by at least one crossvein
(reversed within Gomphaeschnidae and a few other taxa);
RP2 at least slightly undulating (reversed in
Archipetaliidae and some fossil Gomphaeschninae like
Alloaeschna and
Gomphaeschnaoides; modified to a
characteristical curvature in Aeshnodea,
contra LOHMANN, 1996); Rspl better defined;
gaff at least slightly prolonged (somewhat reduced in
Cymatophlebiinae, correlated with the reduction
of the anal loop).
- Other characters: presence of at least a
small intraocellar lobe in adults (convergent to
Cavilabiata, contra CARLE, 1995); male
secondary genitalia (in the groundplan) include a
lamina anterior with elongate median cleft (CARLE,
1996), a short L-shaped ligula (CARLE, 1995, 1996) with
the antero-ventral face developed into a sharp edged
valve separator (CARLE, 1996), hamuli anteriores
lamellate and directed medially (CARLE, 1996), and
hamuli posteriores strongly reduced in size (CARLE,
1995, 1996) (however, as already suggested by LOHMANN,
1996, some of these genital characters are ambiguous or
even dubious, while at least the reduced hamuli
posteriores are certainly not a symplesiomorphy,
contra LOHMANN, 1996); larval prehensile mask
with an elongated prementum (certainly not a
symplesiomorphy, contra LOHMANN, 1996, since
the prementum in larval austropetaliids and aeshnids is
distinctly longer than in Epiophlebia and in
fossil larvae of Isophlebiidae and
Aeschnidiidae); larval epiproct typically bifurcate
apically (CARLE, 1996); larval compound eyes produced
forward, being widest anterior to antennal bases
(CARLE, 1996); proventricular lobes of gizzard small
and mound-like with eight or fewer clustered teeth
(CARLE, 1996); presence of a true dorso-longitudinal
carina (not only a sharp fold) on at least some of the
adult abdominal terga (convergent to
Tarsophlebiidae and Laterocarinida; reduced in some
Archipetaliidae and in Gomphaeschna).
-
Taxonomy:
- [Aeschnina sensu SELYS, 1850]
- [Aeschnidae sensu BUCHECKER, 1876]
- [partim: Aeschninae & Cordulegasterina
sensu KIRBY, 1890]
- [Aeschnidae sensu JACOBSON & BIANCHI,
1905]
- [Aeschninae sensu RIS, 1909]
- Palanisoptera PFAU, 1991 (nec Palanisoptera
sensu LOHMANN, 1995, 1996)
- Aeshnata BECHLY, 1996a (taxon nov.) (nec Aeshnata
LOHMANN, 1996)
- Aeshnomorpha BECHLY et al., 2001
(taxon nov.)
Comment: LOHMANN (1996) mentions several alleged
symplesiomorphies that should indicate a position of
austropetaliids and aeshnids basal of Petalurida and
Exophytica ("ektoflexate" hind tibiae, no sexual
dimorphism in the armature of the mid and hind tibiae, and
terminal segment of male vesicula spermalis less fused and
without processus dorsales, correlated with smaller female
spermathecae). We regard all these states as ambiguous or
even dubious characters that are not yet sufficiently
investigated and documented. If they are correct at all, they
could rather represent reversals, since a more basal position
of Petalurida was convincingly demonstrated by BECHLY (1996)
and NEL et al. (1998).
The non wing venational characters are mostly unknown in
Mesuropetaloidea and therefore might also represent
autapomorphies of Aeshnoptera.
The name Aeshnomorpha BECHLY et al. (2001)
should be preferred for this monophylum, since the previous
name Aeshnata BECHLY, 1996a could lead to confusion with the
junior homonym Aeshnata LOHMANN, 1996, which was used by the
latter author for a very different monophylum (Aeshnodea).
This risk of confusion would be aggravated by Lohmanns
use of the suffix "-ata" as a standardised suffix
for his high-level sistergroups, and his rejection of the
monophyly of Aeshnomorpha. The name Palanisoptera PFAU, 1991
should be avoided because of its conjunction with the
probably erroneous hypothesis that all other extant
Anisoptera do form a monophyletic group (Neanisoptera PFAU,
1991). Furthermore the name Palanisoptera was recently used
by LOHMANN (1996) for a very different monophylum (here named
Euaeshnida BECHLY, 1996a), too, which could again lead to
considerable confusion.
Austropetaliida BECHLY
et al., 2001
- Included taxa: Archipetaliidae
BECHLY, 1996 and Austropetaliidae
(sens. nov.).
-
Autapomorphies:
- Wing venation: series of five to eight
reddish costal spots (convergent to Neopetaliidae)
(CARLE & LOUTON, 1994; CARLE, 1995, 1996),
including an apical spot and a spot in the middle of
the postnodal space (contrary to Neopetaliidae);
pterostigmata secondarily shortened (BECHLY, 1994), and
the pterostigmal brace vein not aligned with its basal
side (LOHMANN, 1996); IR1 very long (convergent to
Petalurida and Valdaeshninae); the insertions of
the CuP-crossing and PsA on the anal vein AA are very
close to each other; basal true lestine oblique vein
reduced or completely suppressed (there is only one
distinct oblique vein between RP2 and IR2 in a very
distal position, probably homologous to the distal
accessory oblique vein).
- Other characters: larval labrum strongly
widened distally (CARLE, 1995, 1996); massive
ventro-lateral development of larval occipital ridge
massive (CARLE, 1995, 1996); larval femora dorsally
excrescent, supplied with tubercles (SCHMIDT, 1941;
CARLE, 1995, 1996); larval transverse abdominal muscles
completely suppressed (CARLE, 1995, 1996), therefore
larvae secondarily unable of jet-propulsion; larvae
with extensively granulate body surface (CARLE, 1996);
larva with lateral abdominal lobes on all abdominal
segments (CARLE, 1995, 1996); larval cerci shortened,
being shorter than half of the length of the ventral
margin of the 10th abdominal segment (LOHMANN, 1995,
1996); terminal segment of vesicula spermalis pendulous
with paired flagellae being sickle-like (CARLE, 1995);
epiproct of adult males developed as a very broad and
apically trifid plate (FRASER, 1933), with the median
lobe being much larger than the lateral lobes; cerci of
adult males short and foliate (FRASER, 1933); males
with the ventral margin of the second abdominal tergite
expanded as genital lobes (LOHMANN, 1996, based on a
personal information by Bechly); leaf-like lateral
expansions of abdominal terga VII and VIII (CARLE,
1996; but maybe not belonging to the groundplan).
-
Taxonomy:
- [partim: "Fissilabres" SELYS,
1854 (taxon nov.)]
- [{partim: "Petalinae"
MUTTKOWSKI, 1910}]
- [{partim: "Petaliini"
sensu TILLYARD, 1917}]
- [partim: "Fissilabioidea"
FRASER, 1929, 1933 (taxon nov.)]
- [Austropetaliidae sensu CARLE &
LOUTON, 1994]
- [Austropetaliata sensu LOHMANN, 1995
(unpublished manuscript name according to Art. 8
IRZN)]
- Austropetaliata LOHMANN, 1996 (taxon nov.)
- Austropetaliida BECHLY et al., 2001
(taxon nov.)
Comment: although the taxon name
"Austropetaliata" was already proposed for this
clade by LOHMANN (1996), BECHLY et al. (2001)
suggested not to use this name because of the following two
reasons: (1) Lohmann is using the suffix "-ata"
explicitly to give equal suffixes to sistergroups, which is
rather unscientific and formalistic; (2) since this clade was
previously addressed under the family-group name
Austropetaliidae a scientific name which sounds similar and
which allows the further use of the vernacular expression
"austropetaliids" would be preferable. Out of the
same reasons BECHLY et al. (2001) preferred the
names Aeshnida, Petalurida, and Gomphides, instead of
Lohmann's names Palanisoptera (including Gomphaeschnata
and Aeshnata), Petalurata, and Gomphata.
Archipetaliidae
BECHLY, 1996
(Type genus: Archipetalia TILLYARD, 1917.)
- Included taxa: only including
Archipetalia auriculata TILLYARD, 1917.
-
Autapomorphies:
- Wing venation: anal loop secondarily absent
(convergent to Phyllopetaliinae); RP2 secondarily less
undulating and secondarily more divergent from RP1
(reversal; contra LOHMANN, 1996); RP3/4 and MA
secondarily non-undulate; costal area of wings with 7-8
brownish spots (maybe a plesiomorphy, contra
CARLE, 1996 and LOHMANN, 1996).
- Other characters: see CARLE (1996).
-
Taxonomy:
- Archipetaliinae BECHLY, 1996a (subfam. nov.)
- Archipetaliidae LOHMANN, 1996 (jun. obj. syn.;
could be regarded as stat. nov. of Archipetaliinae
Bechly ?)
- Archipetaliinae CARLE, 1996 (jun. obj. syn.)
- Archipetaliini CARLE, 1996 (jun. obj. syn.; could
be regarded as stat. nov. of Archipetaliinae Bechly
?)
Comment: LOHMANN (1996) cites "four spots between
wing base and nodus" as further potential autapomorphy,
but this is rather the plesiomorphic state since also present
in Hypopetalia.
Austropetaliidae
CARLE & LOUTON, 1994
(Type genus: Austropetalia TILLYARD, 1916.)
- Included taxa: Austropetaliinae
and Phyllopetaliinae.
-
Autapomorphies:
- Wing venation: not yet known.
- Other characters: compound eyes medially in
contact with each other (for further characters see
CARLE, 1996).
-
Taxonomy:
- Austropetaliidae CARLE & LOUTON, 1994 (fam.
nov.)
- Austropetaliinae sensu BECHLY, 1996a
(stat. et sens. nov.)
- Austropetaliidae sensu LOHMANN, 1996
(sens. nov.)
Comment: BECHLY et al. (2001) demonstrated
that the inclusion of Cymatophlebiidae in crown-group
Austropetaliata LOHMANN, 1996, as well as the different
phylogeny of Austropetaliidae in LOHMANN (1996, first part),
are based on an insufficient character analysis with numerous
errors, and a neglect of conflicting evidence, and therefore
have to be dismissed.
Austropetaliinae
CARLE & LOUTON, 1994
(Type genus: Austropetalia TILLYARD, 1916.)
- Included taxa: only including the
genus Austropetalia TILLYARD, 1916 with the type
species A. patricia TILLYARD, 1910 and A.
tonyana THEISCHINGER, 1995 (= A. victoria
CARLE, 1996, jun. subj. syn. according to LOHMANN, 1996
postscipt).
-
Autapomorphies:
- Wing venation: see CARLE (1996).
- Other characters: see CARLE (1996).
-
Taxonomy:
- Austropetaliinae; BECHLY, 1996a (stat. et sens.
nov.) (nom. transl. ex Austropetaliidae CARLE &
LOUTON, 1994)
- Austropetaliini sensu BECHLY, 1996a (stat.
et sens. nov.)
- Austropetaliinae sensu LOHMANN, 1996
postscript (nec Austropetaliinae sensu
LOHMANN, 1996, first part)
- Austropetaliinae sensu CARLE, 1996
- Austropetaliini sensu CARLE, 1996
Phyllopetaliinae
BECHLY, 1996
(Type genus: Phyllopetalia SELYS, 1857)
- Included taxa: Hypopetaliini and Phyllopetaliini.
-
Autapomorphies:
- Wing venation: anal loop secondarily absent
(convergent to Archipetalia); apical spot
transversely enlarged.
- Other characters: see CARLE (1996).
-
Taxonomy:
- Phylopetaliini BECHLY, 1996a (trib. nov.)
- Eurypetaliinae CARLE, 1996 (subfam. nov.) (subj.
jun. syn.)
- [Hypopetaliinae sensu LOHMANN, 1996
postscript]
- Phyllopetaliinae sensu BECHLY, 2003e (stat.
nov.)
Comment: LOHMANN (1996) cites "thorax laterally with
yellow or green stripes" as potential synapomorphy of
Austropetalia and Phyllopetalia, although
this clearly represents a symplesiomorphy.
Hypopetaliini LOHMANN,
1996
(Type genus: Hypopetalia MCLACHLAN, 1870.)
- Included taxa: only including the type
species Hypopetalia pestilens MCLACHLAN,
1870.
-
Autapomorphies:
- Wing venation: wing spots enlarged and
reddish (for further character see LOHMANN (1996) and
CARLE (1996)).
- Other characters: see LOHMANN (1996) and
CARLE (1996).
-
Taxonomy:
- Hypopetaliinae LOHMANN, 1996 (subfam. nov.)
- Hypopetaliinae CARLE, 1996 (subfam. nov.) (jun.
obj. syn.)
- Hypopetaliini CARLE, 1996 (trib. nov.) (jun. obj.
syn.; could be regarded as stat. nov. of Hypopetaliinae
Lohmann ?)
Phyllopetaliini
BECHLY, 1996
(Type genus: Phyllopetalia SELYS, 1857.)
- Included taxa: Rheopetaliina and Phyllopetaliina.
-
Autapomorphies:
- Wing venation: see CARLE (1996).
- Other characters: see CARLE (1996).
-
Taxonomy:
- Phyllopetaliini BECHLY, 1996a
- Eurypetaliinae sensu CARLE, 1996
- Eurypetaliini CARLE, 1996 (trib. nov.) (jun. subj.
syn.)
- Phyllopetaliini sensu LOHMANN, 1996
postscript (sens. nov.)
Rheopetaliina CARLE,
1996
(Type genus: Rheopetalia CARLE, 1996.)
- Included taxa: only including the type
genus Rheopetalia CARLE, 1996.
-
Autapomorphies:
- Wing venation: not yet known.
- Other characters: see CARLE (1996).
-
Taxonomy:
- Rheopetaliini CARLE, 1996
- Rheopetaliina sensu LOHMANN, 1996
postscript (stat.nov.)
Phyllopetaliina
BECHLY, 1996
(Type genus: Phyllopetalia SELYS, 1857.)
- Included taxa: only including the type
genus Phyllopetalia SELYS, 1857.
-
Autapomorphies:
- Wing venation: see CARLE (1996).
- Other characters: see CARLE (1996).
-
Taxonomy:
- Phylopetaliini BECHLY, 1996a (trib. nov.)
- Eurypetaliini sensu CARLE, 1996
(trib.nov.)
- Phylopetaliina sensu LOHMANN, 1996
postscript (stat. nov.)
Panaeshnida BECHLY et al., 2001
- Included taxa: Progobiaeshnidae
and Aeshnida.
-
Autapomorphies:
- Wing venation: Rspl very well-defined (not
zigzagged) in both pairs of wings; hypertriangles
divided by several parallel crossveins (reversed in the
Gomphaeschnidae except the most basal genus
Oligoaeschna, and maybe in
Cymatophlebia purbeckensis and C.
pumilio); discoidal triangles divided into more
than two cells (convergent to
Cymatophlebiella euryptera and Hypopetalia
pestilens; reversed in the Gomphaeschnidae except
the most basal genus Oligoaeschna); submedian
space divided by one or more accessory cubito-anal
crossveins between CuP-crossing and PsA (reversed in
Gomphaeschnidae and Brachytron, and maybe in
Eumorbaeschna).
- Other characters: not yet known.
-
Taxonomy:
- Panaeshnida BECHLY et al., 2001 (taxon
nov.)
Progobiaeshnidae BECHLY et al., 2001
(Type genus: Progobiaeshna BECHLY et
al., 2001)
- Included taxa: currently only
including Progobiaeshna liaoningensis
BECHLY et al., 2001, and preliminarily also
Gobiaeshna occulta PRITYKINA, 1977.
-
Autapomorphies:
- Wing venation: pterostigmata relatively
short, and pterostigmal brace vein transverse, not
oblique like the basal side of the pterostigma;
pseudo-IR1 strongly reduced (very short and originating
distinctly distal of the pterostigma); anal loop
enlarged and divided into about nine cells, correlated
with a more pronounced elongation of the gaff
(convergent to Hoyaeshna and Aeshnidae,
and a few other Aeshnida; but unknown in
Gobiaeshna); several rows of cells between IR2
and Rspl which are more or less parallel; hindwing
subdiscoidal triangle two-celled (convergent to
Cymatophlebiella euryptera, Hypopetalia
pestilens, Eumorbaeschna
jurassica, and Cymatophlebioidea, except
Valdaeshna surreyensis; but only known in
Progobiaeshna liaoningensis;).
- Other characters: not yet known.
-
Taxonomy:
- Progobiaeshnidae BECHLY et al., 2001
(fam. nov.)
Discussion: the characters clearly demonstrate that this
taxon belongs to the Aeshnoptera-Aeshnomorpha, as sistergroup
of Aeshnida (see synapomorphies of Panaeshnida). As a
consequence of this proposed phylogenetic position,
Gobiaeshna certainly cannot be regarded as a synonym
of Baissaeshna, contrary to the speculations
of WIGHTON & WILSON (1986: 520), since
Baissaeshna shares the apomorphic characters of
Aeshnida-Euaeshnida-Neoaeshnida-Aeshnodea that are absent in
Progobiaeshnidae. The incorrect placement of
Gobiaeshna within the more derived portion of the
"gomphaeschnine" grade by WIGHTON & WILSON
(1986) was based on several insufficiencies of the data
matrix (homoplastic characters, unsafe polarities, unknown
states), and the exclusion of Cymatophlebia
from the analysis which led to partly incorrect polarities.
The significant similarities between the wing venation of
Progobiaeshnidae and the wing venations of
Rudiaeschnidae and Austropetaliida are due to numerous
symplesiomorphies that are of considerable importance for the
reconstruction of the groundplan of Aeshnomorpha.
Aeshnida BECHLY, 1996
- Included taxa: Cymatophlebioidea
and Paneuaeshnida.
-
Autapomorphies:
- Wing venation: presence or a Mspl (still
weakly developed in the groundplan; a well-developed
Mspl obviously evolved parallel in some species of
Cymatophlebia and in all
Euaeshnida).
- Other characters: compound eyes enlarged and
medially contiguous; presence of a very distinct
dorso-longitudinal carina on the abdominal terga 3-8 of
adults (convergent to Tarsophlebiidae and
Laterocarinida; reduced in Gomphaeschna;
contra LOHMANN, 1996); adult anal appendages
elongated (especially the cerci).
-
Taxonomy:
- Aeshnida BECHLY, 1996a (taxon nov.)
- Aeshnida sensu BECHLY et al., 2001
Comment: the non wing venational characters are unknown in
Progobiaeshnidae, and therefore might as well
represent autapomorphies of Panaeshnida.
Cymatophlebioidea HANDLIRSCH, 1906
(Type genus: Cymatophlebia DEICHMÜLLER,
1886.)
- Included taxa: Cymatophlebiidae
and Rudiaeschnidae.
-
Autapomorphies:
- Wing venation: Rspl distinctly curved and
separated by at least three rows of cells from IR2
(convergent to Progobiaeshnidae and Aeshnidae,
incl. Oplonaeschna) (the same might be true
for the Mspl, although its occurrence is quite
homoplastic); one to three convex oblique and
undulating secondary veins anastomosing between IR2 and
RP3/4 directly basal of the origin of Rspl, at least in
the hindwings (somewhat reduced in
Valdaeshna; apparently present by convergence
in Aktassia, but different); hindwing
subdiscoidal triangles divided into two or three cells
(convergent to Cymatophlebiella
euryptera, Progobiaeshna
liaoningensis, Hypopetalia pestilens, and
Eumorbaeschna jurassica; reversed in
Valdaeshna surreyensis).
- Other characters: not yet known.
-
Taxonomy:
- Cymatophlebioidea sensu BECHLY et
al., 2001 (stat. nov.) (nom. transl. ex
Cymatophlebiina HANDLIRSCH, 1906)
Comment: the well-defined oblique secondary veins between
IR2 and RP3/4 basal of Rspl represent a rather strong
synapomorphy of Cymatophlebiinae, Valdaeshninae
and Rudiaeschnidae. This character is currently known
from Cymatophlebia longialata, C.
herrlenae, C. zdrzaleki, C.
kuempeli, C. pumilio,
Prohoyaeshna milleri, Hoyaeshna
cretacica, Valdaeshna surreyensis
(somewhat reduced), and Rudiaeschna limnobia.
It is quite unique within Odonata, since a superficially
similar structure is otherwise only known from the genus
Aktassia, which is certainly unrelated to
Aeshnoptera (anal loop absent, Rspl and Mspl absent, RP1 and
RP2 basally not parallel, IR2 and RP2 not undulate), and most
likely is a close relative of the genus
Aeschnogomphus within Petalurida (NEL et
al., 1998).
Rudiaeschnidae BECHLY et al., 2001
(Type genus: Rudiaeschna DONG & ZIGUANG,
1996)
- Included taxa: only including the type
species Rudiaeschna limnobia DONG &
ZIGUANG, 1996.
-
Autapomorphies:
- Wing venation: the pseudo-anal vein PsA of
the hindwing is more strongly zigzagged and distinctly
weaker than that of the forewing (convergent
Valdaeshna ? kesslerae BECHLY et
al., 2001, and some specimens of
Cymatophlebia longialata); anal loop enlarged
and gaff prolonged; RP1 and RP2 secondarily divergent;
RP2 and IR2 distally distinctly diverging (convergent
Aeschnopsis tischlingeri and
A. jurassica, Archipetaliidae,
Gobiaeshna occulta, Valdaeshninae, and
some fossil Gomphaeschninae, e.g.
Alloaeschna and
Plesigomphaeschnaoides,
Progomphaeschnaoides, and
Gomphaeschnaoides).
- Other characters: not yet known.
-
Taxonomy:
- [Sinocymatophlebiidae BECHLY, 1997] (unpublished
manuscript name, only mentioned on Internet
website)
- Rudiaeschnidae BECHLY et al., 2001
(fam. nov.)
Cymatophlebiidae HANDLIRSCH, 1906
(Type genus: Cymatophlebia DEICHMÜLLER,
1886.)
- Included taxa: Cymatophlebiinae
and Valdaeshninae.
-
Autapomorphies:
- Wing venation: IR2 distinctly undulating
(apomorphy) and parallel to RP2 which is undulating,
too (plesiomorphy); the anastomosing secondary veins
between IR2 and RP3/4, directly basal of Rspl, are more
distinctly developed; Rspl is more strongly curved
(convergent to Aeshnidae, incl. Oplonaeschna);
RP3/4 and MA more strongly undulating (convergent to
Eumorbaeschnidae, the petalurid genus
Uropetala, and a few Eurypalpida of the
"corduliid" grade); the distal primary
antenodal bracket ax2 is shifted distinctly basal of
the level of the distal angle of the discoidal triangle
in the forewings (convergent to Neoaeshnida); the
second (more distal) acessory oblique vein between RP2
and IR2 is much more oblique and longer than the basal
one (maybe rather a plesiomorphy, since also present in
Progobiaeshnidae).
- Other characters: not yet known.
-
Taxonomy:
- [Cymatophlebiidae; LOHMANN, 1995: 60 (invalid stat.
nov. since unpublished according to Art. 8 IRZN)]
- Cymatophlebiidae sensu BECHLY, 1996a: 383
(stat. nov.) (nom. transl. ex Cymatophlebiina
HANDLIRSCH, 1906)
- Cymatophlebiidae; LOHMANN, 1996 (incorrectly
indicated as sens. nov., not indicated as stat.
nov.)
- Cymatophlebiidae sensu BECHLY et
al., 2001
- Cymatophlebiidae sensu NEL et
al., 1998
Comment: according to BECHLY et al. (2001)
Libellulium agrias WESTWOOD, 1954 has to be
regarded as a nomen dubium which probably represents a
Cymatophlebiidae incertae sedis (maybe
Valdaeshninae). Therefore, the previous synonymy of
Libellulium WESTWOOD, 1854 with
Cymatophlebia DEICHMÜLLER, 1886 has to be
rejected. Cymatophlebiella euryptera
PRITYKINA, 1968 almost certainly is no
Cymatophlebiidae at all but more likely a very basal
representative of Aeshnoptera (BECHLY et al., 2001).
Cymatophlebiinae HANDLIRSCH, 1906
(Type genus: Cymatophlebia DEICHMÜLLER,
1886.)
- Included taxa: currently only
including the genus Cymatophlebia, since
all other genera that were previously considered as
Cymatophlebiinae (= Cymatophlebiidae auct.) have to
be excluded from this subfamily (BECHLY et al., 2001).
-
Autapomorphies:
- Wing venation: undulation of RP2 more
strongly developed (convergent to
Eumorbaeschnidae and some Gomphaeschnidae); anal loop
reduced, not distinctly posteriorly closed (somewhat
variable in C. longialata); Mspl
usually better developed (convergent to Euaeshnida),
thus distinctly concave although often still zigzagged,
and therefore apparently indistinct if the corrugation
is disregarded, and strongly curved (convergent to
Aeschnidiidae, Aeshnidae, and some
Libellulidae).
- Other characters: unique ventro-lateral
expansions (genital lobes) at least on the male
abdominal tergum 3; superior appendages (cerci) foliate
(convergent to Mesuropetalidae, Polycanthagynini
incl. "Aeschna" petalura,
and Petalurinae).
-
Taxonomy:
- Cymatophlebiina HANDLIRSCH, 1906 (subfam. nov.)
(nom. imperf.)
- Cymatophlebiinae sensu CARPENTER, 1932:
110 (nom. correct.)
- Cymatophlebiinae sensu BRIDGES, 1993
- [Cymatophlebiidae sensu LOHMANN, 1995: 60
(invalid stat. nov. since unpublished according to Art.
8 IRZN)]
- Cymatophlebiidae sensu LOHMANN, 1996
(incorrectly indicated as stat. nov.)
- Cymatophlebiinae sensu BECHLY, 1996a
(sens. nov.)
- Cymatophlebiinae sensu BECHLY et
al., 2001 (several new species described)
Valdaeshninae BECHLY et al., 2001
(Type genus: Valdaeshna JARZEMBOWSKI,
1988.)
- Included taxa: Valdaeshna
surreyensis JARZEMBOWSKI, 1988, V. ?
kesslerae BECHLY et al., 2001),
Prohoyaeshna milleri BECHLY et al., 2001, and Hoyaeshna cretacica NEL &
MARTÍNEZ-DELCLÒS, 1993, and maybe
Libellulium agrias WESTWOOD, 1854.
-
Autapomorphies:
- Wing venation: presence of a pseudo-ScP,
derived from postnodal crossveins (only known from
Valdaeshna and
Hoyaeshna, not preserved in
Prohoyaeshna but probably present, since
Prohoyaeshna shares with
Hoyaeshna the basal position of the
pterostigmal brace vein as synapomorphy that is not
shared by Valdaeshna); primary IR1 is
secondarily much elongated and straight (convergent to
Petalurida, Austropetaliida and a few aeshnids, like
Boyeria); RP1 and RP2 secondarily divergent
(convergent Aeschnopsis tischlingeri
and A. jurassica, Archipetaliidae,
Gobiaeshna occulta,
Rudiaeschnidae, and some fossil Gomphaeschninae, e.g.
Alloaeschna and
Plesigomphaeschnaoides,
Progomphaeschnaoides, and
Gomphaeschnaoides); RP3/4 and MA closely
parallel up to the wing margin in both pairs of wings
(convergent to Mesuropetalidae and Euaeshnida);
pseudo-anal vein PsA straight and short, and much less
oblique than in Cymatophlebiinae, correlated
with longitudinal elongate triangles (convergent to
"higher" Aeshnidae; unknown for
Prohoyaeshna; but unknown for
Prohoyaeshna, and transformed in
Valdaeschna ? kesslerae).
- Other characters: not yet known.
-
Taxonomy:
- [Valdaeshninae sensu BECHLY, 1996a (nomen
nudum)]
- Valdaeshninae sensu NEL et al.,
1998 (nomen nudum)
- Valdaeshninae BECHLY et al., 2001
(subfam. nov.)
Comment: Valdaeshna JARZEMBOWSKI, 1988 and
Hoyaeshna NEL &
MARTÍNEZ-DELCLÒS, 1993 have been previously
assigned to Aeshnidae (auct.) (JARZEMBOWSKI, 1988; NEL et
al., 1994), but several character their venation reveals
that they are indeed members of the Cymatophlebiidae
(BECHLY, 1996a; NEL et al., 1998; BECHLY et
al., 2001).
Paneuaeshnida BECHLY
et al., 2001
- Included taxa: Paracymatophlebiidae
and Euaeshnida.
-
Autapomorphies:
- Wing venation: Mspl is rather distinct, long
and rather straight, and more or less parallel to MA in
the groundplan (but its course is still somewhat
irregular in Paracymatophlebiidae and
Eumorbaeschnidae); RP2 and IR2 not parallel (?); RP2
distinctly undulating; RP3/4 and MA closely parallel up
to the wing margin in both pairs of wings (convergent
to Mesuropetaloidea and
Valdaeshninae).
- Other characters: not yet known.
-
Taxonomy:
- Paneuaeshnida BECHLY et al., 2001
(taxon nov.)
Paracymatophlebiidae BECHLY et al., 2001
(Type genus: Paracymatophlebia BECHLY et
al., 2001)
- Included taxa: only including the type
species Paracymatophlebia splendida BECHLY
et al., 2001.
-
Autapomorphies:
- Wing venation: two rows of cells in the
basal area between RP1 and RP2; the distal accessory
oblique vein between RP2 and IR2 is secondarily absent;
RP3/4 and MA more strongly undulating; hypertriangles
unicellular (reversal); secondarily no accessory
cubito-anal crossveins in the submedian space between
CuP-crossing and PsA; anal loop posteriorly poorly
closed.
- Other characters: not yet known.
-
Taxonomy:
- Paracymatophlebiidae BECHLY et al., 2001 (fam. nov.)
Euaeshnida BECHLY, 1996
- Included taxa: Eumorbaeschnidae
and Neoaeshnida.
-
Autapomorphies:
- Wing venation: RP2 and IR2 more distinctly
non-parallel; forewing discoidal triangle more elongate
than that of the hindwing ((CARLE, 1995; convergent to
Valdaeshna; but unknown in
Paracymatophlebiidae); forewing subdiscoidal triangle
unicellular (convergent to Aeschnopsis
tischlingeri ?, Austropetaliida and
Valdaeshna surreyensis; reversed in
Aeshnidae); distal side MAb of the discoidal triangles
at least somewhat bent, or angled, or sigmoidally
curved (BECHLY, 1995); anal loop more or less
transversely enlarged and gaff prolonged.
- Other characters: not yet known.
-
Taxonomy:
- [Aeshninae sensu MUTTKOWSKI, 1910]
- Palanisoptera sensu LOHMANN, 1995 (nec
Palanisoptera PFAU, 1991)
- Euaeshnida BECHLY, 1996a (taxon nov.)
- Palanisoptera sensu LOHMANN, 1996 (nec
Palanisoptera PFAU, 1991)
- Euaeshnida sensu BECHLY et al.,
2000
Comment: the unusual high ranking of the former Aeshnidae
(auct.) was decided to facilitate the subdivision and
phylogenetic classification of this large and diverse
group.
The name Palanisoptera (sensu LOHMANN, 1996) for
this monophylum has been rejected by BECHLY et al.
(2001), because it is not only a younger synonym, but
could lead to considerable confusion because of the previous
use of this name (Palanisoptera PFAU, 1991) for a very
different monophylum (here named Aeshnomorpha).
Eumorbaeschnidae BECHLY et al., 2001
(Type genus: Eumorbaeschna BECHLY et
al., 2001)
- Included taxa: only including the type
species Eumorbaeschna jurassica (CARPENTER,
1932).
-
Autapomorphies:
- Wing venation: in the forewing MA is
distally converging to MP, so that the postdiscoidal
area is distally not widened, but narrowed; in the
forewing a primary antenodal bracket (probably ax2) is
on the level of the basal side of the discoidal
triangle (convergent to Gomphaeschnaoidinae and
Telephlebiidae); RP2 more strongly undulating
(convergent to Cymatophlebiinae and some
Gomphaeschnidae, like Paramorbaeschna
and Linaeschna); RP3/4 and MA are more
strongly undulating (convergent to
Cymatophlebiidae, the petalurid genus
Uropetala, and a few Eurypalpida of the
"corduliid" grade), and strictly parallel up
to the posterior wing margin; hindwing subdiscoidal
triangle two-celled (see comment below).
- Other characters: not yet known.
-
Taxonomy:
- [Eumorbaeschnidae sensu BECHLY, 1996a
(nomen nudum)]
- Eumorbaeschnidae BECHLY et al., 2001
(fam. nov.)
Comment: the genus Morbaeschna NEEDHAM,
1907 is a junior subjective synonym of
Mesuropetala HANDLIRSCH, 1906 since Needham
erroneously believed that the specimen figured by him is
conspecific with "Aeschna"
muensteri GERMAR, 1839, of which the true holotype is in
Munich and represents a very badly preserved specimen of
Mesuropetala koehleri (HAGEN, 1848), so that
Mesuropetala muensteri (GERMAR) has to be
regarded as the valid name of the latter species. This
complex taxonomic problem was solved by BECHLY et
al. (2001) by the description of a new genus
Eumorbaeschna. The valid specific name is
E. jurassica (CARPENTER, 1932) since the paratype of
Cymatophlebia jurassica CARPENTER, 1932 is
conspecific with the aeshnid described by NEEDHAM (1907)
under the name "Morbaeschna
muensteri". This is also confirmed by the original
description of the holotype of C. jurassica by
CARPENTER (1932).
As somewhat less parsimonious alternative the strong
undulation of RP2 could also be regarded as a derived
groundplan character of Aeshnida that has been preserved in
Cymatophlebiinae, Eumorbaeschnidae and some
Gomphaeschnidae, while it was more or less reduced in
Progobiaeshnidae, Valdaeshninae,
Rudiaeschnidae, Paracymatophlebiidae, some
Gomphaeschnidae, and all Aeshnodea. An unicellular
hypertriangle could be a further autapomorphy of
Eumorbaeschnidae, although in one specimen the hypertriangle
is apparently divided by several crossveins, so that this
character is ambiguous. Furthermore, the hypertriangle is
also unicellular in some Neoaeshnida. The two-celled
subdiscoidal triangle of the hindwings cannot be regarded as
a very convincing autapomorphy of this taxon, since it is
somewhat variable in Eumorbaeschna jurassica
and also very homoplastic (convergently present in
Cymatophlebiella euryptera, Progobiaeshna
liaoningensis, Hypopetalia pestilens, and
Cymatophlebioidea, except Valdaeshna
surreyensis).
Neoaeshnida BECHLY,
1996
- Included taxa: Gomphaeschnidae
and Aeshnodea.
-
Autapomorphies:
- Wing venation: Mspl better defined
(non-zigzagged) and strictly parallel to MA in the
groundplan (a distinct curvature of Mspl is a
convergence of Aeshnidae to Aeschnidiidae, some
species of Cymatophlebia, and some
Libellulidae); both pairs of wings with a strong convex
secondary longitudinal vein (trigonal planate) in the
postdiscoidal area, correlated with a distinctly angled
or sigmoidally curved distal side MAb of the discoidal
triangles (BECHLY, 1995, 1996; CARLE, 1995; convergent
to Valdaeshna); in both pairs of wings
MP and CuA are closely parallel with only one row of
cells between them up to the wing margin (MP and CuA
not diverging near the posterior wing margin); the
distal accessory oblique vein between RP2 and IR2 is
secondarily absent (two oblique veins are retained in
Progobiaeshnidae, Cymatophlebioidea, and
in most known wings of Eumorbaeschnidae), so
that only the basal lestine oblique vein is retained
(contrary to Austropetaliida) and shifted basally,
close to the subnodus (this basal position is constant
in Neoaeshnida, but rather variable in the other
Aeshnida); the distal primary antenodal bracket ax2 is
shifted distinctly basal of the level of distal angle
of discoidal triangle in the forewings (convergent to
Cymatophlebiidae, Gomphaeschnaoidinae,
Telephlebiidae, and probably also
Eumorbaeschnidae); in the groundplan only two rows of
cells in the basal part of the postdiscoidal area
between the level of the distal angle of the discoidal
triangle and the level of the midfork (convergent to
Exophytica, Aeschnopsis perkinsi and
A. jurassica, Austropetaliida; retained
in Gomphaeschnidae, Brachytronidae and Telephlebiidae,
but again reversed in Aeshnidae); tendency towards the
formation of a longitudinal accessory anal loop between
CuAb and the most basal posterior branch of CuAa
(homology and polarity somewhat unsafe, since within
Gomphaeschnidae only known from the two fossil species
Paramorbaeschna araripensis and
Alloaeschna paskapooensis, while it is present
in most Aeshnodea, except a few taxa, e.g.
Brachytron).
- Other characters: larval rectal gills of the
derived implicate or foliate duplex type (TILLYARD,
1917); larval anal pyramid hyperattenuate, with a long
epiproctal process and long paraprocts (CARLE, 1995,
1996; LOHMANN, 1996); larval prementum more widened
distally (CARLE, 1996); intraocellar lobe more strongly
developed; terminal segment of male vesicula spermalis
swab-like with the "penile prepuce" being
obsolete (CARLE, 1995, 1996); abdominal tergites with
distinct lateral carinae (CARLE, 1995, 1996).
-
Taxonomy:
- Neoaeshnida BECHLY, 1996a (taxon nov.)
- Neoaeshnida sensu BECHLY et al.,
2001
Comment: the non wing venational characters are mostly
unknown in all fossil Panaeshnida, and might therefore
represent autapomorphies for more inclusive monophyla.
Gomphaeschnidae
TILLYARD & FRASER, 1940
(Type genus: Gomphaeschna SELYS, 1871; =
Gomphaeshna TILLYARD & FRASER, 1940, unjust.
emend. and jun. obj. syn.)
- Included taxa: "Gomphaeschninae"
and Gomphaeschnaoidinae.
-
Autapomorphies:
- Wing venation: the most distal part of the
antesubnodal area between RA and RP is free of
antesubnodal crossveins (LOHMANN, 1996) (convergent to
Araripegomphidae, Cordulagomphinae, and
Cavilabiata; however, such a "cordulegastrid
gap" is not present in Alloaeschna
quadrata); no accessory cubito-anal crossveins in
the submedian space between CuP-crossing and PsA
(reversal, convergent to Brachytron);
discoidal triangles only divided into two cells by a
single crossvein (reversal; not yet present in the most
basal genus Oligoaeschna and some specimens of
Gomphaeschna furcillata); hypertriangles
secondarily unicellular (apparently also present in
"Oligoaeschna"
oligocenica, but not in the extant species of
the most basal genus Oligoaeschna).
- Other characters: derived type of male
vesicula spermalis (LOHMANN, 1996) with a pair of long
processes on the incus of the first segment,
secondarily absent ligula hooks on the second segment,
and a pair of posteriorly directed style-like processes
and a pair of lateral incisions on the third segment
which is strongly curved, and characteristically curved
ram-horn-like flagellae on the terminal segment (maybe
only an autapomorphy of Gomphaeschna); median
lobes of the terminal segment of the male vesicula
spermalis are secondarily unfused (a reversal that
might be explained with an ontogenetic phenomenon, e.g.
paedomorphosis, contra LOHMANN, 1996).
-
Taxonomy:
- [partim: Brachytridi TILLYARD &
FRASER, 1940 (taxon nov.)]
- [partim: Brachytrini FRASER, 1957 (taxon
nov., division not tribe)]
- [Gomphaeschnoidea sensu LOHMANN, 1995: 54
(invalid stat. nov., since unpublished according to
Art. 8 IRZN)]
- Gomphaeschnidae sensu BECHLY, 1996a: 384
(stat. et sens. nov.) (nom. transl. ex Gomphaeshninae
TILLYARD & FRASER, 1940)
- [Gomphaeschnata LOHMANN, 1996: 224 (taxon nov.)
(redundant taxon)]
- Gomphaeschnidae sensu BECHLY et
al., 2001 (several new fossil genera and
species described)
Comment: LOHMANN (1996) suggested that the absence of a
dorso-longitudinal abdominal carina is a unique
symplesiomorphy of his Gomphaeschnata. Since such a carina is
indicated in Mesuropetalidae and known to be
well-developed in many Austropetaliida, all
Cymatophlebiidae, some Gomphaeschnidae (e.g.
Oligoaeschna and Sinojagoria) and
all Aeshnodea, BECHLY (1996a) regarded its absence as an
autapomorphic reversal in Gomphaeschna. Furthermore
such a carina is even present in many specimens of type
species Gomphaeschna furcillata (SAY, 1839), so that
its suppression might only represent a derived trend within
the genus Gomphaeschna. Since several of the
mentioned putative autapomorphies of Gomphaeschnidae seem to
be rather homoplastic or insufficiently known (body
characters), only the reduction in the number of antesubnodal
crossveins between RA and RP (distal of the arculus and basal
of the subnodus) could be regarded as relatively
"good" autapomorphy of paleontological relevance,
although it evolved at least one further time by convergence
as an autapomorphy of Cavilabiata, too.
The two-celled male anal triangle of Gomphaeschna
seems to be an autapomorphy of this genus (convergent to some
Aeshnodea, e.g. Brachytron, Basiaeschna,
and some Aeshnini, and all Eurypalpida) since the anal
triangle is three-celled in Linaeschna,
Oligoaeschna and Gomphaeschnaoides
(still unknown in the other fossil taxa).
"Gomphaeschninae"
TILLYARD & FRASER, 1940
(Type genus: Gomphaeschna SELYS, 1871; =
Gomphaeshna TILLYARD & FRASER, 1940, unjust.
emend. and jun. obj. syn.)
- Included taxa: including the type
genus Gomphaeschna SELYS, 1871, and maybe the
extant genera Linaeschna MARTIN, 1909 and
Oligoaeschna SELYS, 1889, and the fossil genera
Alloaeschna WIGHTON & WILSON, 1986 and
Cretalloaeschna JARZEMBOWSKI & NEL,
1996. According to PETERS (pers. comm. 1999) there are no
synapomorphies supporting an inclusion of
Linaeschna into Gomphaeschninae, and the
plesiomorphic(?) absence of a trigonal supplement might
even indicate a more basal phylogenetic position within
Euaeshnida!
-
Autapomorphies:
- Wing venation: not yet known.
- Other characters: not yet known.
-
Taxonomy:
- [Jagorinae sensu FRASER, 1936: 55 (nomen
nudum)]
- Gomphaeshninae TILLYARD & FRASER, 1940: 376
(nom. imperf.)
- Gomphaeshninae; FRASER, 1957: 97
- Gomphaeschnini; DAVIES, 1981: 28 (stat. nov.)
(incorr. subseq. spell. according to Art. 33(b)(i)
IRZN, since not demonstrably intentional)
- Gomphaeschninae; WIGHTON & WILSON, 1986:
505-506
- Gomphaeschninae; MARTÍNEZ-DELCLÒS
& NEL, 1993
- Gomphaeschninae sensu BECHLY et
al., 2001 (sens. nov.)
Comment: no autapomorphies of this taxon are yet known,
therefore it might be paraphyletic in its present generic
composition.
Gomphaeschnaoidinae BECHLY et al., 2001
(Type genus: Gomphaeschnaoides CARLE &
WIGHTON, 1990.)
- Included taxa: Sinojagorini and
Gomphaeschnaoidini.
-
Autapomorphies:
- Wing venation: in the forewing there is only
a single secondary antenodal crossvein between ax1 and
ax2 which is aligned like a primary antenodal bracket
(unknown in Plesigomphaeschnaoides); in
the forewing ax2 is shifted basally on the level of the
basal angle of the discoidal triangle (convergent to
Telephlebiidae and probably also
Eumorbaeschnidae) (unknown in
Plesigomphaeschnaoides); pterostigmal brace
vein very oblique.
- Other characters: not yet known.
-
Taxonomy:
- Gomphaeschnaoidinae BECHLY, 1998d (nomen
nudum)
- Gomphaeschnaoidinae BECHLY 2001, 2001 (subfam. nov.)
Sinojagorini BECHLY et al., 2001
(Type genus: Sinojagoria BECHLY et
al., 2001.)
- Included taxa: only including the type
species Sinojagoria imperfecta BECHLY
et al., 2001.
-
Autapomorphies:
- Wing venation: anal loop large, six- or
seven-celled, nearly as wide as long (this character is
very weak, but the taxon is monophyletic by monotypy
anyway).
- Other characters: not yet known.
-
Taxonomy:
- Sinojagorini BECHLY et al., 2001
(trib. nov.)
Comment: unfortunately, two of the known autapomorphies of
Gomphaeschnaoidini (the basally widened cell below the
pterostigma, and the weakly defined posterior branches of
hindwing CuAa) are currently unknown in
Sinojagoria. Nevertheless, the hypothesis of a
sistergroup relationship between Sinojagoria
and the remaining Gomphaeschnaoidinae is reasonably
supported, and there is also no conflicting evidence against
this hypothesis.
Gomphaeschnaoidini BECHLY et al., 2001
(Type genus: Gomphaeschnaoides CARLE &
WIGHTON, 1990.)
- Included taxa: only including the type
genus Gomphaeschnaoides CARLE &
WIGHTON, 1990, and the new fossil genera
Paramorbaeschna,
Progomphaeschnaoides, and
Plesigomphaeschnaoides that are all described in
BECHLY et al. (2001).
-
Autapomorphies:
- Wing venation: presence of a characteristic
elongate distal paranal cell in the hindwing, directly
basal of the anal loop (convergent to
Cordulagomphinae contra CARLE & WIGHTON,
1990); pterostigmal brace vein somewhat undulating;
basally widened cell bellow the pterostigma, caused by
a curvature of RP1 at the pterostigmal brace (but
unknown in Sinojagoria); posterior
branches of CuAa are relatively weakly defined in the
hindwing (but unknown in
Sinojagoria).
- Other characters: not yet known.
-
Taxonomy:
- Gomphaeschnaoidini BECHLY et al., 2001
(trib. nov.)
Comment: except the character of the paranal cell all
mentioned synapomorphies are not yet known for
Plesigomphaeschnaoides, but the latter genus is
phenetically very similar to the remaining gomphaeschnaoidine
genera.
Aeshnodea BECHLY, 1996
- Included taxa: Allopetaliidae and
Euaeshnodea.
-
Autapomorphies:
- Wing venation: undulation of RP2 modified to
a characteristical curvature beneath the pterostigma;
undulation of RP3/4 and MA completely absent
(reversal); discoidal triangles more strongly
longitudinal elongated, at least in the forewings (a
very homoplastic and dubious character which is also
present in some Gomphaeschnidae and even in some
Cymatophlebiidae); PsA reduced to an oblique
cubito-anal crossvein (reversal)
- Other characters: adult postclypeus more or
less transversely enlarged; imaginal labium without
median cleft (LOHMANN, 1996; a dubious groundplan
character since a fissum is retained in some basal
aeshnids like Nasiaeschna and
Austroaeschna as indicated by this author in
the same publication!); epiproct of adult males
apically only slightly bifid or even not at all bifid
(a bifid epiproct is here regarded as plesiomorphic;
contra LOHMANN, 1996).
-
Taxonomy:
- [Aeshnoidea sensu LOHMANN, 1995
(unpublished according to Art. 8 IRZN)]
- Aeshnodea BECHLY, 1996a (taxon nov.)
- Aeshnata LOHMANN, 1996 (taxon nov.) (nec Aeshnata
BECHLY, 1996a)
- Aeshnodea sensu BECHLY et al., 2001
Allopetaliidae
COCKERELL, 1913
(Type genus: Allopetalia SELYS, 1873.)
- Included taxa: at least including the
extant genus Allopetalia SELYS, 1873, and
preliminarily also the fossil genus
Baissaeshna PRITYKINA, 1977. Maybe the genus
Basiaeschna SELYS, 1883 also belongs to this
group, but according to PETERS (pers. comm. 1998) it rather
seems to be the sistergroup of Oplonaeschna within
Aeshnidae (see below).
-
Autapomorphies:
- Wing venation: two rows of cells between
Rspl and IR2, and between Mspl and MA; two rows of
cells between RP1 and RP2 basal of the pterostigma
(reversal; but not present in
Basiaeschna).
- Other characters: not yet known.
-
Taxonomy:
- Allopetaliini COCKERELL, 1913 (trib. nov.)
- Allopetaliidae sensu BECHLY et
al., 2001 (stat. et sens. nov.)
Discussion: PRITYKINA (1977) regarded the extant genera
Oligoaeschna and Oplonaeschna as closest
relatives of Baissaeshna. This hypothesis is
based on symplesiomorphic and convergent similarities, and
rather improbable, since Baissaeshna does not
seem to belong to Gomphaeschnidae (like
Oligoaeschna) and certainly not to Aeshnidae (like
Oplonaeschna, as demonstrated by BECHLY, 1996,
1997), but most likely belongs to a basal grade within
Aeshnodea, as is indicated by the apomorphic presence of a
seven-celled anal loop and an accessory anal loop, the
plesiomorphic presence of several crossveins that divide the
discoidal triangle and hypertriangle, and the plesiomorphic
absence of the oblique RP3/4-MA brace, as well as the
plesiomorphic absence of the other autapomorphies of either
Aeshnidae, or Telephlebiidae stat. nov. Among extant aeshnids
Baissaeshna is most similar and probably most
closely related to Basiaeschna and/or
Allopetalia, because of the mentioned putative
synapomorphies, and numerous symplesiomorphies (e.g. Rspl and
Mspl not curved, but parallel to IR2 and MA). The unforked
IR2 is a potential symplesiomorphy, too, although it cannot
be totally excluded that this might be a reversal (as
probably the case in Boyeria, and almost certainly
in Oplonaeschna). As demonstrated by BECHLY et
al. (2001) the previous hypothesis of WIGHTON &
WILSON (1986) that Baissaeshna and
Gobiaeshna shall be sister-genera or even synonyms,
cannot be upheld, since both species belong to different,
only remotely related, monophyla within Aeshnoptera. The most
parsimonious interpretation is a close relationship of
Baissaeshna with Allopetalia (and
Basiaeschna?) in a separate family Allopetaliidae. A
few plesiomorphic characters of the latter taxon (e.g. the
unforked IR2 and the smaller anal loop) suggest a sistergroup
relationship with all remaining Aeshnodea (Euaeshnodea).
Eueshnodea BECHLY et
al., 2001
- Included taxa: Brachytronidae and
Aeshnoidea.
-
Autapomorphies:
- Wing venation: IR2 with a distal dichotomic
furcation (secondarily rather indistinctly developed as
secondary branch in Aeshna juncea, A.
caerulea, and Oreaeschna, while
completely suppressed in Oplonaeschna and most
species of Boyeria, except B. vinosa
and B. grafiana; in Anax the IR2-fork
is present, but much shorter, narrower and shifted
distally; but probably plesiomorphic absent in
Allopetaliidae); anal loop enlarged (generally more
than 5-celled, convergent to Baissaeshna
prisca)..
- Other characters: not yet known.
-
Taxonomy:
- Euaeshnodea BECHLY et al., 2001 (taxon
nov.)
Brachytronidae
COCKERELL, 1913
(Type genus: Brachytron EVANS, 1845.)
- Included taxa: at least including the
sister-genera Brachytron EVANS, 1845 and
Aeschnophlebia SELYS, 1883. According to PETERS
(pers. comm. 1998) there is strong morphological evidence
that the two sistergenera Nasiaeschna and
Epiaeschna also belong to this group.
-
Autapomorphies:
- Wing venation: pterostigmal brace vein
reduced (the well-defined pterostigmal brace vein of
Aeshnoidea has to be regarded as symplesiomorphy;
contra PETERS, 1987); MP and CuAa distally
more divergent in the forewing (reversal); pterostigma
basally prolonged, except in Nasiaeschna
(PETERS, pers. comm. 1998).
- Other characters: males secondarily without
constriction of the basal abdomen (PETERS, pers. comm.
1998); keel on abdominal tergites VIII and IX (PETERS,
pers. comm. 1998); occiput with tubercle (PETERS, pers.
comm. 1998).
-
Taxonomy:
- Brachytronini COCKERELL, 1913 (trib. nov.)
- Brachytronini; TILLYARD, 1917
- Brachytroninae; FRASER, 1936
- Brachyirinae; TILLYARD & FRASER, 1940
(incorrect original spelling)
- Brachytrinae; TILLYARD & FRASER, 1940
- [partim: Brachytridi TILLYARD &
FRASER, 1940 (taxon nov.)]
- [partim: Brachytrini FRASER, 1957 (taxon
nov., division not tribe)]
- Brachytrinae; FRASER, 1957
- Brachytroninae; DAVIES, 1981
- Brachytronini; DAVIES, 1981
- Brachytronidae sensu BECHLY, 1996a (stat.
nov.)
Aeshnoidea LEACH,
1815
(Type genus: Aeshna FABRICIUS, 1775; =
Aeschna ANONYMUS, 1801.)
- Included taxa: Telephlebiidae and
Aeshnidae. Maybe also
including the following genera as rather basal Aeshnoidea
incertae sedis: Nasiaeschna, and the
sister-genera Allopetalia and
Basiaeschna. The latter two genera have been
classified in separate tribes Allopetaliini and
Basiaeschnini by COCKERELL (1913) who included
Gomphaeschna in Allopetaliini and
Oplonaeschna in Basiaeschnini.
-
Autapomorphies:
- Wing venation: strong tendency towards the
development of an accessory anal loop between the two
main branches of CuA in the hindwing; presence of
several accessory cubito-anal crossveins in both pairs
of wings.
- Other characters: third abdominal segment
distinctly constricted in adult males; epiproct of
adult males triangularly elongated and apically not
bifid; larvae with distinctly larger compound eyes
(PETERS, 1987); larval paraprocts elongated.
-
Taxonomy:
- Aeshnoidea; TILLYARD, 1926 (stat. nov.) (nom.
transl. ex Aeshnidae LEACH, 1815)
- Aeshnidea; PRITYKINA, 1968 (incorr. subseq.
spell.)
- Aeshniidea; HENNIG, 1969, 1981 (incorr. subseq.
spell.)
- Aeshnoidea; CARLE, 1986 (sens. nov.)
- Aeshnoidea sensu BECHLY, 1996a (sens.
nov.)
Telephlebiidae
COCKERELL, 1913
(Type genus: Telephlebia SELYS, 1883.)
- Included taxa: Austroaeschninae
and Telephlebiinae.
-
Autapomorphies:
- Wing venation: ax2 recessed to the basal
angle of the discoidal triangle (close to, at, or even
basal of the arculus).
- Other characters: not yet known.
-
Taxonomy:
- Telephlebiini COCKERELL, 1913 (trib. nov.)
- [partim: Brachytridi TILLYARD &
FRASER, 1940 (taxon nov.)]
- [partim: Brachytrini FRASER, 1957 (taxon
nov., division not tribe)]
- [partim: Brachytrinae; FRASER, 1957]
- Caliaeschnidae BECHLY, 1996a (fam. nov. with the
type genus: Caliaeschna SELYS, 1883) (jun.
subj. syn.)
- Telephlebiidae sensu BECHLY et
al., 2001 (stat. nov.)
Austroaeschninae
BECHLY, 1996
(Type genus: Austroaeschna SELYS, 1883.)
- Included taxa: including the genera
Spinaeschna THEISCHINGER, 1982,
Notoaeschna TILLYARD, 1916, Austroaeschna
SELYS, 1883 (= Dromaeschna FÖRSTER, 1908),
Acanthaeschna SELYS, 1883, and
Planaeschna MCLACHLAN, 1896.
-
Autapomorphies:
- Wing venation: not yet known.
- Other characters: not yet known.
-
Taxonomy:
- Austroaeschninae BECHLY, 1996a (subfam. nov.)
Comment: the sister-genera Spinaeschna and
Notoaeschna share spiny larval paraprocts as unique
synapomorphy.
Telephlebiinae
COCKERELL, 1913
(Type genus: Telephlebia SELYS, 1883.)
- Included taxa: including the genera
Boyeria MCLACHLAN, 1896, Caliaeschna
SELYS, 1883, Gynacanthaeschna FRASER, 1921,
Cephalaeschna SELYS, 1883 (= Indophlebia
FRASER, 1935), Periaeschna MARTIN, 1908,
Petaliaeschna FRASER, 1927, and
Telephlebia SELYS, 1883. The unbranched IR2 of
Boyeria (in some species, e.g. B. vinosa
and B. grafiana, an indistinctly branched IR2 has
been retained!) has to be regarded as reversal, convergent
to Oplonaeschna.
-
Autapomorphies:
- Wing venation: median space traversed by
crossveins; hypertriangles traversed by at least three
crossveins in both pairs of wings (convergent to
Aeshnidae).
- Other characters: not yet known.
-
Taxonomy:
- Telephlebiini COCKERELL, 1913 (trib. nov.)
- Boyeriini COCKERELL, 1913 (trib. nov. with the type
genus Boyeria MCLACHLAN, 1896) (jun. subj.
syn.)
- Caliaeschninae BECHLY, 1996a (subfam. nov. with the
type genus: Caliaeschna SELYS, 1883) (jun.
subj. syn.)
- Telephlebiinae sensu BECHLY et
al., 2001 (stat. nov.)
Aeshnidae LEACH, 1815
(Type genus: Aeshna FABRICIUS, 1775; =
Aeschna ANONYMUS, 1801.)
- Included taxa: Epiaeschninae, Oplonaeschninae,
Gynacanthinae, and
Aeshninae.
-
Autapomorphies:
- Wing venation: characteristical bulge in the
distal part of the MA in both pairs of wings
("aeshnid bulla"); anal loop further enlarged
(about 10 cells or more); subdiscoidal triangle of both
wings traversed by one crossvein (convergent to
Hypopetalia, Cymatophlebiidae, and
Eumorbaeschnidae); Rspl and Mspl distinctly
curved, with more than one row of cells (mostly even
more than two!) between them and IR2 or MA (convergent
to Oligoaeschna venusta, Basiaeschna
janata and Aeschnophlebia kolthoffi that
have two rows of cells between the mentioned veins, but
always straight radial and median supplements); the
widened wing space between IR2 and Rspl and MA and Mspl
is at least basally divided by oblique intercalary
veinlets; in both pairs of wings more than two rows of
cells in the basal part of the postdiscoidal field
between the level of the distal angle of the discoidal
triangle and the level of the midfork (convergent to
Aeschnidioptera, most Petalurida,
Mesuropetalidae, Cymatophlebioidea,
Eumorbaeschnidae, Alloaeschna,
Linaeschna, Petaliaeschna, few
Lindeniidae like Cacoides and
Melanocacus, and many Libellulidae, except
Tetrathemistinae; reversed in Subaeschna and
Coryphaeschna); hypertriangles traversed by at
least three crossveins in forewings and more than three
crossveins in hindwings (convergent to Telephlebiinae;
reversed in Oplonaeschninae and many Aeshnini and
Anacini).
- Other characters: adult postclypeus very
much transversely enlarged; larval paraprocts further
elongated.
-
Taxonomy:
- Aeshnidae LEACH, 1815
- Aeschnides LEACH, 1815 (incorr. subseq. spell.)
?
- Aeschnidae; BURMEISTER, 1839
- Aeschnoides; SELYS, 1840 (incorr. subseq.
spell.)
- Aeschnides; RAMBUR, 1842 (incorr. subseq.
spell.)
- Aeschnines; SELYS & HAGEN, 1850 (incorr.
subseq. spell.)
- Aeschnina; SELYS, 1850 (incorr. subseq.
spell.)
- [Aeschnidees; SELYS, 1854 (trivial name?)]
- Aeschnina; HAGEN, 1861 (incorr. subseq.
spell.)
- Aeschnidae; BUCHECKER, 1876
- Aeshnina; CABOT, 1881 (incorr. subseq.
spell.)
- Aeschnidae; KIRBY, 1890
- Aeschninae; KIRBY, 1890 (stat. nov. ?)
- Aeschnii; ACLOQUE, 1897 (incorr. subseq.
spell.)
- [Esnidos NAVAS, 1903 (trivial name?)]
- Aeschninae; NEEDHAM, 1903
- Aeschnidae; NEEDHAM, 1903
- Aeschnodea; JACOBSON & BIANCHI, 1905 (stat.
nov. ?)
- Aeschnidae; JACOBSON & BIANCHI, 1905
- Aeshninae; CALVERT, 1909
- Aeschninae; RIS, 109
- Aeshninae; MUTTKOWSKI, 1910
- Aeshninae; FRASER, 1936
- Aeshninae; TILLYARD & FRASER, 1940 (sens.
nov.)
- Aeshnidae; TILLYARD & FRASER, 1940
- Aeshnina; NEL & MARTÍNEZ-DELCLÒS
& ESCUILLIÉ & BRISAC, 1994 (incorr.
subseq. spell.)
- Aeshnidae sensu BECHLY, 1996a (sens.
nov.)
Epiaeschninae BECHLY,
1996
(Type genus: Epiaeschna HAGEN, 1875.)
- Included taxa: only including the
genus Epiaeschna HAGEN, 1875.
-
Autapomorphies:
- Wing venation: not yet known.
- Other characters: not yet known.
-
Taxonomy:
- [partim: Brachytridi TILLYARD &
FRASER, 1940 (taxon nov., section)]
- [partim: Brachytrini FRASER, 1957 (taxon
nov., division not tribe)]
- [partim: Brachytrinae; FRASER, 1957]
- Epiaeschninae BECHLY, 1996a (subfam. nov.)
Comment: the genus Epiaeschna that was previously
classified as "Brachytronini", could belong to
Aeshnidae because of several derived similarities, but it
does not share any known synapomorphies with one of the other
subfamilies of Aeshnidae. The rather small anal loop and the
less curved Rspl and Mspl would indicate that Epiaeschninae
might be the sistergroup of all other Aeshnidae. On the other
hand, the stronger curved RP1 with only one row of cells
between RP1 and the pterostigma, and the presence of oblique
veinlets that divide the distal area between IR2 and Rspl and
between MA and Mspl, could be putative synapomorphies with
Aeshninae. However, according to PETERS (pers. com. 1998)
there is strong morphological evidence supporting a
sistergroup relationship of Epiaeschna and
Nasiaeschna within Brachytronidae (see above).
Oplonaeschninae
BECHLY, 1996
(Type genus: Oplonaeschna SELYS, 1883)
- Included taxa: only including the
genus Oplonaeschna SELYS, 1883.
-
Autapomorphies:
- Wing venation: IR2 secondarily unbranched
(reversal; convergent to Boyeria); hindwing
discoidal triangle much less narrow than forewing
discoidal triangle (WIGHTON & WILSON, 1986;
reversal, probably convergent to
Alloaeschna and Allopetalia); both
triangles only traversed by two crossveins
(reversal).
- Other characters: not yet known.
-
Taxonomy:
- [partim: Brachytridi TILLYARD &
FRASER, 1940 (taxon nov., section)]
- [partim: Brachytrini FRASER, 1957 (taxon
nov., division not tribe)]
- [partim: Gomphaeschninae; FRASER,
1957]
- Oplonaeschninae BECHLY, 1996a (subfam. nov.)
Comment: the genus Oplonaeschna that was
previously classified as "Gomphaeschnini", also
belongs to the Aeshnidae, but it does not share any known
synapomorphies with either Epiaeschninae, or Gynacanthinae,
or Aeshninae. The less elongate and less narrow triangles
could indicate a sistergroup relationship of Oplonaeschninae
with all remaining Aeshnidae. According to PETERS (pers.
comm. 1998) there is strong morphological evidence for an
inclusion of Basiaeschna SELYS, 1883 within
Aeshnidae - Oplonaeschninae as sistergenus to
Oplonaeschna (e.g. structure of the male hamuli,
shape of anal triangle and membranule, curved Rspl and Mspl,
trigonal supplement fusing with Mspl, reduced IR2 fork,
etc.).
Gynacanthinae COCKERELL,
1913
(Type genus: Gynacantha RAMBUR, 1842; =
Triacanthagyna SELYS, 1883, jun. obj. syn; =
Acanthagyna KIRBY, 1890, jun. subj. syn..)
- Included taxa: including the genera
listed in BRIDGES (1994) under Gynacanthini.
-
Autapomorphies:
- Wing venation: membranule strongly reduced;
characteristically shaped (symmetrical and hexagonal)
anal loop.
- Other characters: compound eyes very large
and very broadly confluent, correlated with crepuscular
habits (FRASER, 1957); anal appendages long and thin in
both sexes (FRASER, 1957); female genital region with
the dentigerous plate produced into a robust
two-pronged fork (FRASER, 1957; convergent to
Periaeschna, Gynacanthaeschna,
"Aeshna" ornithocephala and
Polycanthagynini. According to PETERS (pers. comm.)
"Aeshna" ornithocephala probably
represents a member of Polycanthagynini!).
-
Taxonomy:
- Gynacanthini COCKERELL, 1913 (trib. nov.)
- Neuraeschnini COCKERELL, 1913 (jun. subj.
syn.)
- Gynacanthaginae sensu TILLYARD &
FRASER, 1940 (stat. nov.) (nom. imperf.)
- [partim: Aeshnidi TILLYARD & FRASER,
1940 (taxon nov., section)]
- [partim: Aeshnini FRASER, 1957 (taxon
nov., division not tribe)]
- Gynacanthinae sensu LIEFTINCK, 1954 (nom.
correct.)
- Gynacanthagini sensu DAVIES, 1981 (stat.
rest.)
- Gynacanthinae sensu BECHLY, 1996a (stat.
rest.)
Comment: the monophyly of Gynacanthinae is not very
well-supported and the very different state of the
"aeshnid bulla" and the Rp3/4-MA-brace within this
group could represent evidence for paraphyly. According to
PETERS (pers. comm. 1998) the gynacanthines could represent
the most basal group of Aeshnidae.
BRIDGES (1994) discussed the probable synonymies of the
genera Triacanthagyna, Acanthagyna, and the
type genus Gynacantha, but did not make the
appropriate decisions because he believed these would
"greatly upset nomenclature". Contrary to BRIDGES
(1994) the case seems to be rather simple: obviously neither
RAMBUR (1842), nor SELYS (1857) designated a type species for
Gynacantha. The first subsequent designation of a
type species (G. trifida) for Gynacantha
was made by KIRBY (1890) and is the only valid one. According
to Art. 67k IRZN it is irrelevant that G. trifida
was already the type species of Triacanthagyna. The
neglectance of the possible restriction of
Gynacantha by SELYS (1857) is irrelevant, too. The
only important point is, that the subsequent type species
belongs to the originally included species by RAMBUR (1842)
(Art. 69a(i) IRZN). Therefore the arguments by KIMMINS (1936)
are invalid, and the subsequent type species designations by
CALVERT (1905) and TILLYARD (1916) are invalid, too. The
subsequent designation of a type species (G.
nervosa) for Acanthagyna by COWLEY (1934) is
perfectly valid. Thus, the situation is absolutely clear:
Triacanthagyna definitely is a junior objective
synonym of Gynacantha, and Acanthagyna is a
valid generic name. Any confusion can be easily avoided if
Acanthagyna is regarded as a new junior subjective
synonym of Gynacantha. There is no taxonomic
confusion resulting from a classification of all the
referring species in the genus Gynacantha, because
of the following four reasons: (1) all referring species
belong to the same subfamily; (2) neither
Gynacantha, nor Acanthagyna hav been
demonstrated to be either monophyletic or paraphyletic, thus
a synonymisation does not conflict with any phylogeny; (3)
there exist no objective criteria anyway for the delimitation
of genera (of course genera should be monophyletic, but
whether a given monophylum is treated as genus, or as tribe,
or as suborder, is completely arbitrary); (4)
Triacanthagyna and Acanthagyna were treated
by various authors in the past as synonyms of
Gynacantha, and the referring type species were all
among the originally included species of Gynacantha
sensu Rambur.
Aeshninae LEACH, 1815
(Type genus: Aeshna FABRICIUS, 1775; =
Aeschna ANONYMUS, 1801.)
- Included taxa: Polycanthagynini,
Aeshnini, and Anacini.
-
Autapomorphies:
- Wing venation: at the "aeshnid
bulla" a characteristical oblique crossvein is
developed as convex brace between RP3/4 and MA,
appearing like a branch of MA, while the main branch of
MA becomes concave for a short distance (1-2 cells) and
is distally less distinct (FRASER, 1957; BECHLY, 1994);
the widened wing space between IR2 and Rspl and MA and
Mspl is also distally divided by oblique intercalary
veinlets (PETERS, 1987); branching point of IR2 shifted
distally beneath the pterostigma, at least in
forewings; in both pairs of wings the vein RP1 is more
strongly curved, with only one row of cells between RP1
and the pterostigma (maybe a synapomorphy with
Epiaeschninae, and a convergence with
Brachytron); two rows of cells between the
basal parts of MP and CuAa in hindwings (reversed in
Polycanthagyna, Coryphaeschna and
Remartinia.
- Other characters: larval gills of foliate
duplex type (TILLYARD, 1917; only known from Aeshnini
and Anacini, but maybe more generally
distributed).
-
Taxonomy:
- Aeshnidae LEACH, 1815
- Aeschnides LEACH, 1815 (incorr. subseq. spell.)
?
- Aeschnidae; BURMEISTER, 1839
- Aeschnoides; SELYS, 1840 (incorr. subseq.
spell.)
- Aeschnides; RAMBUR, 1842 (incorr. subseq.
spell.)
- Aeschnines; SELYS & HAGEN, 1850 (incorr.
subseq. spell.)
- Aeschnina; SELYS, 1850 (incorr. subseq.
spell.)
- [Aeschnidees; SELYS, 1854 (trivial name?)]
- Aeschnina; HAGEN, 1861 (incorr. subseq.
spell.)
- Aeschnidae; BUCHECKER, 1876
- [partim: Dynamophlebiae BUCHECKER, 1876 (taxon
nov.)]
- Aeshnina; CABOT, 1881 (incorr. subseq.
spell.)
- Aeschnidae; KIRBY, 1890
- Aeschninae; KIRBY, 1890 (stat. nov. ?)
- Aeschnii; ACLOQUE, 1897 (incorr. subseq.
spell.)
- [Esnidos NAVAS, 1903 (trivial name?)]
- Aeschninae; NEEDHAM, 1903
- Aeschnidae; NEEDHAM, 1903
- Aeschnodea; JACOBSON & BIANCHI, 1905 (stat.
nov. ?)
- Aeschnidae; JACOBSON & BIANCHI, 1905
- Aeshninae; CALVERT, 1909
- Aeschninae; RIS, 109
- Aeshninae; MUTTKOWSKI, 1910
- Aeshninae; FRASER, 1936
- Aeshninae; TILLYARD & FRASER, 1940 (sens.
nov.)
- Aeshnidae; TILLYARD & FRASER, 1940
- [partim: Aeshnidi TILLYARD & FRASER,
1940 (taxon nov., section)]
- [partim: Aeshnini FRASER, 1957 (taxon
nov., division not tribe)]
- Aeshnina; NEL & MARTÍNEZ-DELCLÒS
& ESCUILLIÉ & BRISAC, 1994 (incorr.
subseq. spell.)
- Aeshninae sensu BECHLY, 1996a (sens.
nov.)
Polycanthagynini
TILLYARD & FRASER, 1940
(Type genus: Polycanthagyna FRASER, 1933.)
- Included taxa: only including the
species P. erythromelas (MCLACHLAN, 1896), P.
melanictera (SELYS, 1883), P. ornithocephala
(MCLACHLAN, 1896), and according to PETERS (pers. comm.)
probably also "Aeshna" petalura
(MARTIN, 1908).
-
Autapomorphies:
- Wing venation: not yet known.
- Other characters: female genital region with
the dentigerous plate prolonged backwards and ending in
several robust spines (FRASER, 1957); females with
bright heliochromatic colouring (FRASER, 1957).
-
Taxonomy:
- Polycanthaginae TILLYARD & FRASER, 1940
(subfam. nov.) (nom. imperf.)
- Polycanthagyni sensu DAVIES, 1981 (stat.
nov.) (emend.; nom. imperf.)
- Polycanthagini sensu DAVIES & TOBIN,
1985 (unjustified emendation; nom. imperf.)
- Polycanthaginini sensu NEL &
MARTÍNEZ-DELCLÒS & ESCUILLIÉ
& BRISAC, 1994 (emend; nom. imperf.)
- Polycanthagynini sensu BECHLY, 1996a (nom.
correct.)
Comment: maybe the sistergroup of Anacini, as indicated by
the states of the ax1, the oblique RP3/4-MA brace, the
presence of one row of cells between the distal RP3/4 and MA,
and the more strongly curved Rspl. The female genital plate
developed as digging organ for oviposition in dry soil is a
convergence with Periaeschna,
Gynacanthaeschna, and Gynacanthinae.
Aeshnini LEACH, 1815
(Type genus: Aeshna FABRICIUS, 1775; =
Aeschna ANONYMUS, 1801.)
- Included taxa: including the genera
listed in BRIDGES (1994) under Aeshnini.
-
Autapomorphies:
- Wing venation: anal triangle very narrow;
anal triangle only two-celled (convergent to
Gomphaeschnidae, Coryphaeschna,
Castoraeschna and Remartinia), except
in the most basal species like Aeshna cyanea
(PETERS, 1987).
- Other characters: not yet known.
-
Taxonomy:
- Aeschnini; COCKERELL, 1913 (stat. nov. ) (nom.
transl. ex Aeshnidae LEACH, 1815)
- Aeschnini; TILLYARD, 1917
- Aeshnini; DAVIES, 1981 (sens. nov.)
- Aeshnini sensu BECHLY, 1996a (sens.
nov.)
Comment: both mentioned autapomorphies are only present in
some of the species of the genus Aeshna which was
already demonstrated to be paraphyletic by PETERS, 1987.
Anacini COCKERELL, 1913
(Type genus: Anax LEACH, 1815 1814-1817.)
- Included taxa: Amphiaeschnina and
Anacina.
-
Autapomorphies:
- Wing venation: the basal primary antenodal
bracket ax1 is at least somewhat more oblique in the
forewing than in the hindwing; sectors of arculus
arising from its anterior half (ASKEW, 1988), so that
the posterior part of the arculus is prolonged; Rspl
more strongly curved; only one row of cells between the
distal part of RP3/4 and MA (PETERS, 1987), and a more
developed oblique brace between these two veins,
correlated with a distally rather indistinct MA.
- Other characters: not yet known.
-
Taxonomy:
- Anactinini COCKERELL, 1913
- Anactini; DAVIES, 1981 (stat. nov.)
- Anactini sensu BECHLY, 1996a (sens.
nov.)
- Anacini sensu BECHLY, 2003e (nom.
correct.)
Amphiaeschnina
COCKERELL, 1913
(Type genus: Amphiaeschna SELYS, 1871.)
- Included taxa: including
"Aeshna" mixta LATREILLE, 1805,
"Aeshna" affinis VAN DER
LINDEN, 1820, Hesperaeschna californica CALVERT,
1895, Hesperaeschna biliosa KENNEDY, 1938, and
Neureclipa NAVÁS, 1911, as well as the
genera Amphiaeschna SELYS, 1871,
Castoraeschna CALVERT, 1952,
Coryphaeschna WILLIAMSON, 1903,
Oreaeschna LIEFTINCK, 1937, and
Remartinia NAVÁS, 1911. According to PETERS
(pers. comm.) there are also African species which may
rather belong to the genus Hesperaeschna
COCKERELL, 1913.
-
Autapomorphies:
- Wing venation: anal loop only with two
transverse rows of cells (only present in
"Aeshna" mixta,
"Aeshna" affinis,
Hesperaeschna californica, Hesperaeschna
biliosa, and Neureclipa).
- Other characters: not yet known.
-
Taxonomy:
- Amphiaeschnini COCKERELL, 1913 (trib. nov.)
- Hesperaeschnina BECHLY, 1996a (subtrib. nov. with
the type genus: Hesperaeschna COCKERELL, 1913)
(jun. subj. syn.)
- Amphiaeschnina sensu BECHLY, 2003e (stat.
nov.)
Anacina COCKERELL, 1913
(Type genus: Anax LEACH, 1815 1814-1817.)
- Included taxa: including the
sister-genera Anaciaeschna SELYS, 1878 and
Anax LEACH, 1815 (incl. HemianaxSELYS,
1883). According to PETERS (pers. comm. 1998)
Anaciaeschna shall not be the sistergenus to
Anax.
-
Autapomorphies:
- Wing venation: the basal primary antenodal
bracket ax1 is much more strongly oblique in the
forewing; pterostigmata elongated and shifted basally
(not yet in Anaciaeschna; convergent to
Isophlebiidae and Aeschnidiidae), therefore the
pterostigmal brace vein is situated midway between
nodus and apex (not yet in Anaciaeschna;
convergent to Petalurida); the oblique median brace at
the aeshnid bulla is more strongly developed; discoidal
triangles very narrow and elongate (not yet in
Anaciaeschna); male anal angle rounded
(Anaciaeschna) or completely reduced
(Anax); male triangle reduced (not yet in
Anaciaeschna); membranule prolonged up to
tornus; gaff (= basal CuA before its branching)
strongly elongated and very straight in hindwings; RP2
sharply curved towards RP1 beneath the distal part of
the pterostigma (FRASER, 1957; not yet in
Anaciaeschna); Mspl and Rspl more strongly
curved, with a broader area between Mspl and MA and
Rspl and IR2; distal part of Rspl obliterated, not
reaching the hind margin of the wing (maybe a
synapomorphy with some Amphiaeschnina, like A.
mixta, A. affinis, Neureclipa
and Remartinia); bifurcation of IR2 strongly
reduced (not yet in Anaciaeschna).
- Other characters: males without auricles on
the second abdominal segment (except in
Anaciaeschna); male abdomen supplied with
supplementary ridges on segments 4-8 (FRASER, 1957;
rudimentary or vestigial in Anaciaeschna and
Hemianax; maybe synapomorphic with
Oreaeschna); adult males with a short and
stout epiproct (except in Anaciaeschna).
-
Taxonomy:
- Anactinini sensu COCKERELL, 1913
- Anaxinae sensu FRASER, 1936 (stat. nov.)
(unjust. emend.)
- Anactinae sensu TILLYARD & FRASER,
1940
- Anactini sensu DAVIES, 1981 (stat.
nov.)
- Anactina sensu BECHLY, 1996a (stat.
nov.)
- Anacina sensu BECHLY, 2003e (nom.
correct.)