Phylogenetic Systematics of Odonata


© Günter Bechly, Böblingen, 2007


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Phylogenetic Systematics of Stem-Group Odonates / Discoidalia




Discoidalia BECHLY, 1996


† Triadophlebioptera BECHLY, 1996


† Triadotypomorpha BECHLY, 1996


† Triadotypidae GRAUVOGEL & LAURENTIAUX, 1952

(Type genus: † Reisia HANDLIRSCH, 1912; = † Triadotypus GRAUVOGEL & LAURENTIAUX, 1952, jun. subj. syn.)

Comment: REIS (1909) described † Handlirschia gelasii of which the genus name is a junior homonym of † Handlirschia KOHL, 1896. HANDLIRSCH (1912) therefore erected the substitute genus † Reisia. LAURENTIAUX-VIEIRA & RICOUR & LAURENTIAUX (1952) synonymised † Reisia gelasii (REIS, 1909) with † Triadotypus guillaumei GRAUVOGEL & LAURENTIAUX, 1952, but did not consider the correct taxonomical consequences, therefore BECHLY (1997i) recognized the senior synonym † Reisia gelasii (REIS, 1909) as valid name for † Triadotypus guillaumei GRAUVOGEL & LAURENTIAUX, 1952. According to Art. 40(a) IRZN the family-group name † Triadotypidae does not have to be changed in this case, although the name of the type genus changed.
Reisia was placed by REIS (1909) and HANDLIRSCH (1912, 1920) in "Protodonata", and by CARPENTER (1931, 1992) in Palaeoptera incertae sedis, while BRIDGES (1994) did not mention this genus at all.


† Piroutetiidae NEL, 1989

(Type genus: † Piroutetia MEUNIER, 1907.)


† Triadophlebiomorpha PRITYKINA, 1981


† Zygophlebioidea (sens. nov. in BECHLY, 1996a)

(Type genus: † Zygophlebia PRITYKINA, 1981.)


† Zygophlebiidae PRITYKINA, 1981

(Type genus: † Zygophlebia PRITYKINA, 1981.)


† Xamenophlebiidae PRITYKINA, 1981

(Type genus: † Xamenophlebia PRITYKINA, 1981.)


† Triadophlebiida BECHLY, 1996


† Mitophlebiidae PRITYKINA, 1981

(Type genus: † Mitophlebia PRITYKINA, 1981.)


† Triadophlebioidea PRITYKINA, 1981

(Type genus: † Triadophlebia PRITYKINA, 1981.)


† Triadophlebiidae PRITYKINA, 1981

(Type genus: † Triadophlebia PRITYKINA, 1981.)


† Paurophlebiidae BECHLY, 1996

(Type genus: † Paurophlebia PRITYKINA, 1981.)


† Neritophlebiinae BECHLY, 1996

(Type genus: † Neritophlebia PRITYKINA, 1981.)


† Paurophlebiinae BECHLY, 1996

(Type genus: † Paurophlebia PRITYKINA, 1981.)


† Nonymophlebiini BECHLY, 1996

(Type genus: † Nonymophlebia PRITYKINA, 1981.)


† Paurophlebiini BECHLY, 1996

(Type genus: † Paurophlebia PRITYKINA, 1981.)


Stigmoptera BECHLY, 1996

Comment: According to NEL (unpubl., pers. comm. 1998) the odonatoid pterostigma could rather be an autapomorphy of Discoidalia, and being secondarily absent in † Triadophlebioptera, since he recently discovered an undescribed triadophlebiomorph odonate with pterostigma in the collection of PIN in Moscow. The non-wing venational characters are mostly unknown from † Protanisoptera and † Triadophlebioptera and might therefore represent autapomorphies for more inclusive monophyla.


† Protozygoptera TILLYARD, 1925


† Permagrionoidea TILLYARD, 1928

(Type genus: † Permagrion TILLYARD, 1928.)


† Permagrionidae TILLYARD, 1928

(Type genus: † Permagrion TILLYARD, 1928.)


† Permolestidae MARTYNOV, 1932

(Type genus: † Permolestes MARTYNOV, 1932.)


† Archizygoptera HANDLIRSCH, 1906


† Permepallagidae MARTYNOV, 1938

(Type genus: † Permepallage MARTYNOV, 1938.)


† Kennedyidae TILLYARD, 1925

(Type genus: † Kennedya TILLYARD, 1925.)


† Protomyrmeleontoidea HANDLIRSCH, 1906

(Type genus: † Protomyrmeleon GEINITZ, 1887.)


† Batkeniidae PRITYKINA, 1981

(Type genus: † Batkenia PRITYKINA, 1981.)

Comment: † "Triassoneura" primitiva PRITYKINA, 1981 is phenetically very similar to the genus † Batkenia but has to be preliminarily regarded as † Batkeniidae incertae sedis. It was transferred to a new genus † Paratriassoneura in BECHLY (1997i). A putative autapomorphy of this new genus is the separation of the origins of RP2 and IR2 by six cells.


† Batkeniinae PRITYKINA, 1981

(Type genus: † Batkenia PRITYKINA, 1981.)


† Batkenia PRITYKINA, 1981

(Type species: † Batkenia pusilla PRITYKINA, 1981.)


† Terskeja PRITYKINA, 1981

(Type species: † Terskeja paula PRITYKINA, 1981.)


† Voltzialestinae BECHLY, 2003e

(Type genus: † Voltzialestes gen. nov. in NEL et al., 1996)


† Protomyrmeleontidae HANDLIRSCH, 1906

(Type genus: † Protomyrmeleon GEINITZ, 1887.)


† Triassagrioninae TILLYARD, 1922

(Type genus: † Triassagrion TILLYARD, 1922.)


† Protomyrmeleontinae HANDLIRSCH, 1906

(Type genus: † Protomyrmeleon GEINITZ, 1887.)


Panodonata BECHLY, 1996

Comment: several of the mentioned non-wing autapomorphies of Panodonata are unknown from most fossils and therefore might rather represent autapomorphies for more inclusive groups within Odonatoptera.


† Tarsophlebiidae HANDLIRSCH, 1906

(Type genus: † Tarsophlebia HAGEN, 1866.)

Comment: the sistergroup relationship of † Tarsophlebia with all crowngroup Odonata is indicated by several plesiomorphic characters of † Tarsophlebia (viz ligula orimentary; vesicula spermalis still very short and flat with a very wide porus; hindwing discoidal cell basaly still open, thus arculus incomplete; legs still with four tarsomeres of equal length), that are developed in the derived state in all extant odonates (BECHLY & BRAUCKMANN & ZESSIN & GRÖNING, 2001). The body characters are presently only known from † Tarsophlebia eximia. The alleged male auricles of † Tarsophlebia eximia are based on a misinterpretation of hamuli posteriores by NEL et al. (1993) (BECHLY & BRAUCKMANN & ZESSIN & GRÖNING, 2001). Likewise the interpretation of the tarsus was erroneous since † Tarsophlebia indeed retained four tarsomeres of equal length, just like † Meganisoptera and † Protozygoptera. The further characters cited by NEL et al. (1993) are clearly symplesiomorphies, especially the acute distal angles of the forewing discoidal and subdiscoidal cell and the aligned [RP & MA]-MAb-CuA that are also present in Epiophlebiidae, † Isophlebioptera and † Heterophlebioptera. The similarities of † Tarsophlebiidae and Epiproctophora (NEL et al., 1993), e.g. the less separated and relatively large eyes, the presence of two cephalic sutures, and the small leg spines, are either symplesiomorphies, or at least characters of uncertain polarity that are here regarded as symplesiomorphies, too.
Recent studies of NEL (pers. comm.) at PIN in Moscow, and of BECHLY (unpubl.) at MCZ, revealed that the alleged calopterygoid-like anal appendages of † Tarsophlebia, with apparently two pairs of claspers, are clearly based on misinterpretations due to artifacts of preservation (FLECK et al., 2004): Indeed † Tarsophlebia definitely does not possess zygopteroid but anisopteroid appendages. There are no visible paraprocts, the epiproct is small, and the cerci are very long, with a double-barreled basal petiole and a distal plate-like expansion. The broken double-parreled petioles of the two cerci have been commonly misinterpreted as two pairs of claspers, while the distal plates have been overlooked or regarded as artifacts.


Odonata FABRICIUS, 1793

Comment: several of the mentioned non-wing autapomorphies of Odonata are unknown from most fossils and therefore might rather represent autapomorphies for more inclusive groups within Odonatoptera.




Last Update: 10th August, 2007

© Günter Bechly, Böblingen, 2007