Phylogenetic Systematics of Odonata


© Günter Bechly, Böblingen, 2007


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Phylogenetic Systematics of Euanisoptera / Cavilabiata




Cavilabiata BECHLY, 1996


Cordulegastrida BECHLY, 1996


Zoraenidae LOHMANN, 1992

(Type genus: Zoraena KIRBY, 1890.)


Cordulegastridae HAGEN, 1875

(Type genus: Cordulegaster [LEACH], [1815].)


Cordulegastrinae HAGEN, 1875

(Type genus: Cordulegaster [LEACH], [1815].)


Taeniogastrini LOHMANN, 1992

(Type genus: Taeniogaster SELYS, 1854.)


Cordulegastrini HAGEN, 1875

(Type genus: Cordulegaster [LEACH], [1815].)


Thecagastrinae LOHMANN, 1992

(Type genus: Thecagaster SELYS, 1854; = Kuldanagaster YOUSUF & YUNUS, 1974, jun. subj. syn.)


Pangaeagastrini BECHLY, 1996

(Type genus: Pangaeagaster LOHMANN, 1992)


Thecagastrini LOHMANN, 1992

(Type genus: Thecagaster SELYS, 1854; = Kuldanagaster YOUSUF & YUNUS, 1974, jun. subj. syn.)


Lauragastrina BECHLY, 1996

(Type genus: Lauragaster LOHMANN, 1992)


Thecagastrina LOHMANN, 1992

(Type genus: Thecagaster SELYS, 1854; = Kuldanagaster YOUSUF & YUNUS, 1974, jun. subj. syn.)


Cristotibiata BECHLY, 2003e


Neopetaliidae TILLYARD & FRASER, 1940

(Type genus: Neopetalia COWLEY, 1934 nom. subst. pro Petalia HAGEN in SELYS, 1854, nec Petalia XXX.)


Brachystigmata BECHLY, 1996


† Nannogomphidae BECHLY, 1996

(Type genus: † Nannogomphus HANDLIRSCH, 1906-1908)

Comment: there are no strong synapomorphies known for † Nannogomphus bavaricus, † Prohemeroscopus jurassicus, and † Prohemeroscopus ? kuehnapfeli, but all three species belong to the same grade of Brachystigmata (basal of Eubrachystigmata), and † Nannogomphus exactly corresponds to the wing venation that would have to be expected from a miniaturized relative of † Prohemeroscopus.
The following characters are autapomorphies of † Prohemeroscopus jurassicus: the forewing subdiscoidal triangle is widened with a curved or angled posterior margin (convergent to † Juracorduliidae, Paneurypalpida, Petalurida, and some Gomphides).
The following characters are autapomorphies of † Nannogomphus bavaricus: very small body size; hindwing CuAa secondarily without posterior branches; there are no antesubnodal crossveins present in the hindwing, between RA and RP, distal of the arculus and basal of the subnodus; anal loop slightly reduced in size and with only four cells; pterostigma further shortened, only 1-2 cells long (convergent to † Juracorduliidae and Eurypalpidiformia); discoidal triangles unicellular in both pairs of wings (convergent to † Hemeroscopidae and Paneurypalpidomorpha). For a redescription of the holotype of † Nannogomphus bavaricus and discussion of its phylogenetic position see BECHLY & NEL & MARTÍNEZ-DELCLÒS, 1996. Although † Nannogomphus without doubt belongs to Brachystigmata, its few derived similarities with Eurypalpida (e.g. unbranched CuAa) rather seem to be convergences due to the small size of † Nannogomphus.


Eubrachystigmata BECHLY, 2003e


† Hemeroscopidae PRITYKINA, 1977

(Type genus: † Hemeroscopus PRITYKINA, 1977.)

Comment: BECHLY et al. (1998) and BECHLY (1998d) suspected that the alleged larvae of † Hemeroscopidae most probably represent larval † Aeschnidiidae, just like † Sonidae (larvae) with which they should be most closely related. The alleged libelluloid-like larval mask and gizzard described by PRITYKINA (1977) was considered by these authors as fragments of another type of larvae. However, HUANG & NEL (2001) could recently describe fossil dragonfly larvae that clearly belong to the alleged hemeroscopid larval type, and which indeed do possess a spoon shaped mask. Since these larvae are found together with numerous adult † Hemeroscopidae, it is well possible that they represent hemeroscopid larvae indeed. Anyway, these larvae do neither seem to be related to the sonid larvae, nor to the true aeschnidiid larvae.


Neobrachystigmata BECHLY, 2003e


Chlorogomphida BECHLY, 1996


† Araripechlorogomphidae BECHLY & UEDA, 2002

(Type genus: † Araripechlorogomphus BECHLY & UEDA, 2002

Comment: the only known specimen is also discussed and figured in BECHLY (1998d), and represents the first fossil record of chlorogomphids, and the first New World record, too.


Chlorogomphoidea NEEDHAM, 1903

(Type genus: Chlorogomphus SELYS, 1854.)


Chloropetaliidae CARLE, 1995

(Type genus: Chloropetalia CARLE, 1995.)


Chlorogomphidae NEEDHAM, 1903

(Type genus: Chlorogomphus SELYS, 1854.)


Eorogomphinae CARLE, 1995

(Type genus: Eorogomphus CARLE, 1995.)


Chlorogomphinae NEEDHAM, 1903

(Type genus: Chlorogomphus SELYS, 1854.)


Sinorogomphini CARLE, 1995

(Type genus: Sinorogomphus CARLE, 1995.)


Chlorogomphini NEEDHAM, 1903

(Type genus: Chlorogomphus SELYS, 1854.)


Paneurypalpidomorpha BECHLY, 2003e


† Juracorduliidae BECHLY, 2003e

(Type genus: † Juracordulia BECHLY, 1998g)


Eurypalpidomorpha BECHLY, 2003e


† Valdicordulioidea BECHLY, 1996

(Type genus: † Valdicordulia JARZEMBOWSKI & NEL, 1996)

Comment: besides the two mentioned weak synapomorphies, there is a very large phenetic similarity between † Valdicorduliidae and † Araripephlebiidae, which of course includes numerous symplesiomorphies, but probably also some more putative synapomorphies.


† Valdicorduliidae BECHLY, 1996

(Type genus: † Valdicordulia JARZEMBOWSKI & NEL, 1996)

Comment: contrary to the original description certainly not a member of crown-group Eurypalpida as is clearly demonstrated by the following plesiomorphies: a distal furcation of CuAa is retained; all secondary antenodal crossveins non-aligned; pseudo-anal vein PsA still distinctly developed in the hindwing; area between RP2 and IR2 distally still widened.


† Araripephlebiidae BECHLY, 1998d

(Type genus: † Araripephlebia BECHLY, 1998d.)

Comment: BECHLY (1998d) regarded † Araripephlebiidae as more closely related to Chlorogomphoidea mainly because of the hindwing discoidal triangle that is more transverse than the forewing discoidal triangle. However, the latter character is hardly developed in Chloropetaliidae and basal genera of Chlorogomphidae (see CARLE, 1995) and thus may not belong to the groundplan of Chlorogomphoidea.


Eurypalpidiformia BECHLY, 2003e


† Eocorduliidae BECHLY, 1996

(Type genus: † Eocordulia PRITYKINA, 1986)


Paneurypalpida BECHLY, 1996


† Araripelibellulidae BECHLY, 1996

(Type genus: † Araripelibellula NEL & PAICHELER, 1994)

Comment: LOHMANN (1996) included the † Condaliidae, † Eocorduliidae, and † Araripelibellulidae (all sensu BECHLY, 1996a) in a single family † Condaliidae sensu Lohmann, and furthermore regards the referring three family group taxa of BECHLY (1996a) as nomina nuda. Since these taxa are all monotypic and the original publication includes a statement "fam. nov.", a designation of a type genus and a reference to the description of the type genus, all three taxa are valid according to Art. 13 IRZN. LOHMANN (1996) regarded the elongated anal loop as a putative synapomorphy of his family † Condaliidae with his higher taxon Firmonervata (= Neolamellida), but this character is unknown for † Condalia, and furthermore represents a very homoplastic character anyway. The only other alleged synapomorphy with Firmonervata (antenodal crossveins aligned) is absent in † Eocordulia, and is very homoplastic, too. The retention of several plesiomorphies (forewing arculus angled; bases of RP and MA separated at arculus; a more distal position of ax2 in the forewing; anal loop small with only two cells; pterostigmal brace aligned with basal side of pterostigma) excludes a position of † Araripelibellula within crown-group Eurypalpida. The fact that † Cratocordulia shows the derived states of these characters has to be regarded as convergence to crown-group Eurypalpida, since this genus shares all listed synapomorphies with the other † Araripelibellulidae. † Eocordulia still has a three-celled male anal triangle, while † Araripelibellula shares with crown-group Eurypalpida the derived two-celled anal triangle as putative synapomorphy. Consequently, † Eocordulia does not belong to † Araripelibellulidae and is therefore here retained as separate family with a more basal position in the stemgroup of Eurypalpida.


† Mesocorduliinae BECHLY, 2003e

(Type genus: † Mesocordulia DONG & ZIGUANG, 1996)


† Araripelibellulinae BECHLY, 1996

(Type genus: † Araripelibellula NEL & PAICHELER, 1994)


Eurypalpida BECHLY, 1996

Comment: LOHMANN (1995, 1996) cites the character "females in the groundplan with only a single spermatheca" as potential autapomorphy of Eurypalpida, but since Trichodopalpida have paired spermathecae (compare MILLER, 1990) this derived state most probably represents a convergence of Synthemistidae and Gomphomacromia.


Synthemistidae TILLYARD, 1911

(Type genus: Synthemis SELYS, 1870.)


Synthemiopsinae CARLE, 1995

(Type genus: Synthemiopsis TILLYARD, 1917.)


Synthemistinae TILLYARD, 1911

(Type genus: Synthemis SELYS, 1870.)


Palaeosynthemistini CARLE, 1995

(Type genus: Palaeosynthemis FÖRSTER, 1903.)


Synthemistini TILLYARD, 1911

(Type genus: Synthemis SELYS, 1870.)


Eusynthemistina CARLE, 1995

(Type genus: Eusynthemis FÖRSTER, 1903.)


Synthemistina TILLYARD, 1911

(Type genus: Synthemis SELYS, 1870.)




Last Update: 10th August, 2007

© Günter Bechly, Böblingen, 2007