Phylogenetic Systematics of Odonata


© Günter Bechly, Böblingen, 2007


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Phylogenetic Systematics of basal Pananisoptera and Anisoptera / Petalurida




Pananisoptera BECHLY, 1996


† Liassogomphidae TILLYARD, 1935

(Type genus: † Liassogomphus COWLEY, 1934 nom. subst. pro † Gomphites TILLYARD, 1925.)


† Juragomphidae NEL & BECHLY & MARTÍNEZ-DELCLÒS & FLECK, 2001

(Type genus: † Juragomphus NEL & BECHLY & MARTÍNEZ-DELCLÒS & FLECK, 2001.)

Comment: the broad and long membranule which is present in † Juragomphidae, but absent in † Liassogomphidae, could represent a synapomorphy of † Juragomphidae and Neoanisoptera.


Neoanisoptera BECHLY, 1998


† Aeschnidiidae NEEDHAM, 1903

(Type genus: † Aeschnidium WESTWOOD, 1854; = Estemoa GIEBEL, 1856, jun. obj. syn.)

Comment: ZHANG (1999) and FLECK et al. (2002) recently described the first genuine aeschnidiid larvae from the Lower Cretaceous of China. These larvae are very large, have an hypertrophied ovipositor and forcep-like paraprocts, spidery legs, as well as a strange mask with very long and narrow prementum and concave mentum and palps. The larval wing sheaths clearly show the typical aeschnidiid wing venation (transverse discoidal triangles, Rspl and Mspl, etc.). I here consider the following fossil dragonfly larvae from China as also belonging to the same type of aeschnidid larvae: † Dissurus HONG, 1982, † Neimengogomphus HONG, 1985, and † Yixiangomphus LIN, 1986. The † Sonidae sensu PRITYKINA, 1986, have been restricted by BECHLY et al. (1998) to the holotypical larva of † Sona nectes, and have been attributed to † Aeschnidiidae by BECHLY (1998d). The alleged adults of † Sonidae are unrelated to these larvae and have been classified by BECHLY et al. (1998) in a new family † Proterogomphidae within Gomphides. The apparently vestigial ovipositor-Anlagen in the alleged female ultimate instar larvae of † Sona nectes seem to contradict an attribution to † Aeschnidiidae, but could be explained if these larvae are no ultimate instar larvae at all. Contrary to earlier beliefs of mine the larvae of † Hemeroscopidae do not seem to be related with † Sona, but indeed seem to belong to Cavilabiata (the dense fringe of swimming hairs on the larval tibiae and tarsi therefore must be considered as a strange convergence). The Brazilian genus † Nothomacromia CARLE, 1995 († Nothomacromiidae), can also be attributed to † Aeschnidiidae because of the large size, the hypertrophied paraprocts, and the spidery legs that all are derived similarities with the undisputed Chinese aeschnidiid larvae. According to BECHLY (1998d) the giant larvae from Lower Cretaceous of Santana in Brazil simply represent late stages of † Nothomacromia. Contrary to FLECK et al. (2002), I do not consider the flat gomphid-like mask of † Sona and † Nothomacromia as compelling evidence against an attribution to † Aeschnidiidae, because if the peculiar mask of the Chinese aeschnidiid larvae is considered as a convergence to Cavilabiata, it could as well be a convergence WITHIN † Aeschnidiidae rather than a convergence OF † Aeschnidiidae.


Anisoptera SELYS in SELYS & HAGEN, 1854

Comment: most of the non wing venational characters are unknown from fossil stemgroup representatives of Anisoptera and could therefore be autapomorphies for more inclusive monophyla.


Petalurida BECHLY, 1996

Comment: the dissimilar discoidal triangles in fore and hindwings, with the forewing discoidal triangle being at least somewhat more transverse, seem to be a symplesiomorphy of Petaluridae, Gomphidae and Eurypalpida, contra NEL et al. (1998), and the broad female abdomen seems to be a symplesiomorphy with † Isophlebioptera († Isophlebia) and † Aeschnidiidae.


† Protolindeniidae HANDLIRSCH, 1906

(Type genus: Protolindenia DEICHMÜLLER, 1886.)


Petalurodea BECHLY, 1996


† Cretapetaluridae NEL et al., 1998

(Type genus: † Cretapetalura NEL et al., 1998)

Comment: the alleged larval autapomorphies mentioned by BECHLY (1996a) are obsolete since they exclusively refer to † Nothomacromia sensibilis (CARLE & WIGHTON, 1990), which indeed seems to be the larva of an † Aeschnidiidae (= † Sonidae s.str.) according to BECHLY et al., (1998), BECHLY (1998d), and FLECK et al. (2002).


Petaluroidea NEEDHAM, 1903

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


† Aktassiidae PRITYKINA, 1968

(Type genus: † Aktassia PRITYKINA, 1968.)


† Pseudocymatophlebiinae NEL et al., 1998

(Type genus: † Pseudocymatophlebia NEL et al., 1998.)


† Aktassiinae PRITYKINA, 1968

(Type genus: † Aktassia PRITYKINA, 1968.)

Comment: A severe problem is the recent discovery (NEL and BECHLY, unpubl.) that † Aktassia indeed does have a strong and oblique pterostigmal brace vein that is more or less aligned with the basal margin of the pterostigma. This either must be a strange reversal, or it is a unique plesiomorphy within Petalurida that questions the here proposed phylogenetic position of † Aktassia. The pterostigmal brace vein is completely suppressed in † Aeschnogomphus (convergent to Cordulegastrida and Chlorogomphida). † Aktassia has several distinct autapomorphies that are all due to its extremely dense wing venation: postsubnodal space divided into two rows of cells; basal part of the subdiscoidal cell (between CuP-crossing and pseudo-anal vein PsA) traversed by crossveins (convergent to † Aeschnidiidae).


Petaluridae NEEDHAM, 1903

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

Comment: the mentioned non-wing venational characters are mostly unknown in the fossil petalurids and therefore could represent autapomorphies for more inclusive monophyla within Petalurida. Nevertheless, they are derived groundplan characters of all extant Petaluridae compared to any other group of extant dragonflies.


Tachopteryginae FRASER, 1933

(Type genus: Tachopteryx UHLER in SELYS, 1859.)


Tanypterygini TILLYARD & FRASER, 1940

(Type genus: Tanypteryx KENNEDY, 1917.)


Tachopterygini FRASER, 1933

(Type genus: Tachopteryx UHLER in SELYS, 1859.)


Tachopteryx UHLER in SELYS, 1859

(Type species: Tachopteryx thoreyi (HAGEN, 1857).)


Phenes RAMBUR, 1842

(Type species: Phenes raptor RAMBUR, 1842.)


Petalurinae NEEDHAM, 1903

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


Uropetala SELYS, 1857

(Type species: Uropetala carovei (WHITE, 1846).)

Comment: although there are no autapomorphies known yet (but these probably exist), this genus is certainly monophyletic, as is strongly indicated by the phenetic similarity of the two species that are both endemic to New Zealand.


Petalura LEACH, 1815

(Type species: Petalura gigantea LEACH, 1815, 1814-1817.)


Euanisoptera BECHLY, 1996




Last Update: 10th August, 2007

© Günter Bechly, Böblingen, 2007