Phylogenetic Systematics of Odonata

© Günter Bechly, Böblingen, 2007

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Phylogenetic Systematics of Euanisoptera / Aeshnoptera

Aeshnoptera BECHLY, 1996

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.)

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.)

Mesuropetalidae BECHLY, 1996

(Type genus: † Mesuropetala HANDLIRSCH, 1906.)

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.)

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

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 Lohmann’s 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

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.)

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.)

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.)

Phyllopetaliinae BECHLY, 1996

(Type genus: Phyllopetalia SELYS, 1857)

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.)

Phyllopetaliini BECHLY, 1996

(Type genus: Phyllopetalia SELYS, 1857.)

Rheopetaliina CARLE, 1996

(Type genus: Rheopetalia CARLE, 1996.)

Phyllopetaliina BECHLY, 1996

(Type genus: Phyllopetalia SELYS, 1857.)

Panaeshnida BECHLY et al., 2001

Progobiaeshnidae BECHLY et al., 2001

(Type genus: † Progobiaeshna BECHLY et al., 2001)

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

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.)

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)

Cymatophlebiidae HANDLIRSCH, 1906

(Type genus: † Cymatophlebia DEICHMÜLLER, 1886.)

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.)

Valdaeshninae BECHLY et al., 2001

(Type genus: † Valdaeshna JARZEMBOWSKI, 1988.)

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

Paracymatophlebiidae BECHLY et al., 2001

(Type genus: † Paracymatophlebia BECHLY et al., 2001)

Euaeshnida BECHLY, 1996

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)

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

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.)

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.)

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.)

Sinojagorini BECHLY et al., 2001

(Type genus: † Sinojagoria BECHLY et al., 2001.)

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.)

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

Allopetaliidae COCKERELL, 1913

(Type genus: Allopetalia SELYS, 1873.)

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

Brachytronidae COCKERELL, 1913

(Type genus: Brachytron EVANS, 1845.)

Aeshnoidea LEACH, 1815

(Type genus: Aeshna FABRICIUS, 1775; = Aeschna ANONYMUS, 1801.)

Telephlebiidae COCKERELL, 1913

(Type genus: Telephlebia SELYS, 1883.)

Austroaeschninae BECHLY, 1996

(Type genus: Austroaeschna SELYS, 1883.)

Comment: the sister-genera Spinaeschna and Notoaeschna share spiny larval paraprocts as unique synapomorphy.

Telephlebiinae COCKERELL, 1913

(Type genus: Telephlebia SELYS, 1883.)

Aeshnidae LEACH, 1815

(Type genus: Aeshna FABRICIUS, 1775; = Aeschna ANONYMUS, 1801.)

Epiaeschninae BECHLY, 1996

(Type genus: Epiaeschna HAGEN, 1875.)

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)

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..)

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.)

Polycanthagynini TILLYARD & FRASER, 1940

(Type genus: Polycanthagyna FRASER, 1933.)

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.)

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.)

Amphiaeschnina COCKERELL, 1913

(Type genus: Amphiaeschna SELYS, 1871.)

Anacina COCKERELL, 1913

(Type genus: Anax LEACH, 1815 1814-1817.)

Last Update: 10th August, 2007

© Günter Bechly, Böblingen, 2007