Phylogenetic Systematics of
Odonata
© Günter Bechly, Böblingen, 2007
Phylogenetic classification of fossil and
extant odonates
(please click on the taxa names for a link to
the referring characters!)
Pterygota
Odonatoclada
Stigmoptera
Epiproctophora
Anisoptera
Exophytica
Cavilabiata
Haplohamulida
Zygoptera
Euzygoptera
Fossil Odonatoptera incertae
sedis:
- Camptotaxineuridae TILLYARD,
1937
(Type genus: Camptotaxineura TILLYARD,
1937)
Maybe belonging to Protanisoptera.
- Kaltanoneuridae ROHDENDORF,
1961
(Type genus: Kaltanoneura ROHDENDORF,
1961)
Probably belonging to Protozygoptera.
- Liadotypidae MARTYNOV, 1937
(Type genus: Liadotypus MARTYNOV, 1937)
Still classified as Meganisoptera by ZESSIN (1983),
but excluded from Odonatoptera by BRAUCKMANN & ZESSIN
(1989). According to NEL (pers. comm., 1998), who re-examined
the holotype, it is most likely an Isophlebioptera, thus not
a "protodonate" at all, but a true crown-group
Odonata.
- Malmagrion
eichstaettense (HAGEN, 1962)
This tiny fossil damselfly from the Solnhofen limestone was
generally regarded as a Protomyrmeleontidae (e.g.
BRIDGES, 1994), but was transferred to Odonatoptera
incertae sedis by NEL (1992). However a rexamination
of the holotype (BECHLY, unpubl.) suggests that it could
indeed be a protomyrmeleontid and even a senior synonym of
Protomyrmeleon jurassicus NEL, 1992.
- Palaeophlebiinae
(Type genus: Palaeophlebia BRAUER, 1889, nec
Palaeophlebia SELYS, 1889)
Belonging to Stigmoptera incertae sedis.
- Perissophlebia
multiseriata TILLYARD, 1918
This species was placed in Odonata incertae sedis by
TILLYARD (1918, 1923), PRITYKINA (1981), and ROZEFELDS
(1985). It was even regarded as a Palaeoptera incertae
sedis by CARPENTER (1992). BECHLY (1997i) regarded it as
an Odonatoptera - Stigmoptera incertae sedis,
because of the odonatoid-like pattern of the main
longitudinal veins, the polyneural cross-venation, and the
presence of a pterostigma. Quite unique autapomorphies of
this genus are the very oblique distal margin of the
pterostigma, and the presence of more than one row of cells
between the costal margin and RA and between RA and RP distal
of the pterostigma. The dense venation might be correlated
with the probable very large total size of the wing.
- {Protokennedya TILLYARD, 1928
(nomen nudum)}
An invalid name coined by Tillyard for the hypothetical
ancestor of Odonata sensu Tillyard.
- Samarurinae TILLYARD, 1917
(Type genus: Samarura BRAUER, REDTENBACHER,
& GANGLBAUER, 1889)
This taxon traditionally incorporated a probably polyphyletic
assemblage of Mesozoic fossil odonatoid larvae with
synlestine habitus. The material of the type species
Samarura gigantea at PIN in Moscow was recently
re-examined by NEL, FLECK and MARTÍNEZ-DELCLÒS
(in prep.), who could definitely confirm its attribution to
Isophlebioidea, since the typical derived wing
venation is visible on the larval wing sheaths of one
specimen. This attribution is also confirmed by the extremely
large size of the larvae (length about 12 cm!) and the
ocurence of adult isophlebiids as only fossil odonates in the
same layers.
- Triassophlebia
stigmatica TILLYARD, 1922
Classified in Mesophlebiidae by TILLYARD (1922, 1923)
(followed by FRASER, 1957), but later regarded by CARPENTER
(1992) and BRIDGES (1994) as an "Anisozygoptera"
incertae sedis, possibly related to
Triassolestes, while ROZEFELDS (1985) and NEL et
al.(1993) regarded Triassophlebia as an
Odonatoptera incertae sedis. BECHLY (1997i) regarded
this species as a Stigmoptera incertae sedis,
although it is quite possible that it belongs to
Parazygoptera.
- Schlechtendaliola nympha
HANDLIRSCH, 1919
(= Schlechtandaliana nympha sensu BRIDGES,
1994, incorr. subseq. spell.)
This fossil was regard as nymphal wing of a
"protodonate" larva by HANDLIRSCH (1919).
BRAUCKMANN & ZESSIN (1989) and BRIDGES (1994) regarded
its position as uncertain, and indeed its odonatoid
affinities are at least doubtful.
Palaeodictyopteroida that had once been
classified as fossil Odonatoptera:
- Calvertiellidae MARTYNOV,
1931
(Type genus: Calvertiella TILLYARD,
1925)
- Permothemistidae sensu
ROHDENDORF, 1962 nom. correct. pro Permothemidae MARTYNOV,
1938 ( nom. subst. pro Palaeothemidae MARTYNOV, 1932)
(Type genus: Permothemis MARTYNOV, 1934, nom.
subst. pro Palaeothemis MARTYNOV, 1932, nec
Palaeothemis FRASER, 1923)
Classified within Protodonata in a separate suborder
Archodonata by MARTYNOV (1932), followed by TILLYARD
& FRASER (1938) and FRASER (1957). Transferred by
SINITSHENKOVA (1980) to a separate suborder
Permothemistida within Palaeodictyopteroida. CARPENTER
(1992) synonymised Archodonata with
Palaeodictyopteroida.
- Protagrionidae HANDLIRSCH,
1906
(Type genus: Protagrion BRONGNIART,
1893)
This family was classified within Protodonata in a
separate suborder Palaeodonata by TILLYARD &
FRASER (1938) and FRASER (1957).
- Spilapteridae HANDLIRSCH, 1906 (=
Dunbariidae HANDLIRSCH, 1937; = Doropteridae
ZALESSKY, 1946)
(Type genus: Spilaptera BRONGNIART,
1885)
The genus Dunbaria (=
Doropteron) was classified by ZALESSKY (1946) in a
separate suborder Hemiodonata, that was synonymised by
CARPENTER (1992) with Palaeodictyoptera.
- Thuringopteryx gimmi
KUHN, 1937
This taxon was classified by KUHN (1937) and MÜLLER
(1965) as Protodonata, but was regarded by ZEUNER
(1939) and CARPENTER (1992) as Saltatoria - Ensifera. Finally
it was transferred to Spilapteridae by BECHLY (1997i),
as first known Triassic representative of
Palaeodictyoptera.
Fossil Neoptera (and other taxa) that
had once been classified as Odonatoptera:
- Aeroplana mirabilis
TILLYARD, 1918
This species was originally described in a new family and
suborder of "Protodonata". TILLYARD (1923) elevated
Aeroplanoptera to a separate order, but
Aeroplana was later generally considered to be a
Phasmatodea (MARTYNOV, 1928; RIEK, 1956; CARPENTER,
1992).
- Antitaxineura anomala
TILLYARD, 1935
This fossil species has erroneously been indicated to be of
Triassic age in CARPENTER (1992) although it was described
from the Upper Permian of New South Wales. It was regarded as
Odonata incertae sedis by CARPENTER (1992) and
BRIDGES (1994), but shall represent a Plecoptera according to
Riek in ROZEFELDS (1985).
- Austrolestidion
duaringae TILLYARD, 1916
This fossil was originally described as "two damselfly
larvae", but turned out to be a single crustacean
(ROZEFELDS, 1985).
-
"Cordulagomphus"
santanensis CARLE & WIGHTON,
1990
The holotype was originally described as a dragonfly larva,
but was recently demonstrated to represent an adult earwig by
BECHLY (in press a) who therefore transferred this species to
Dermaptera incertae sedis.
- Tcholmanvissiidae ZALESSKY, 1934
(= Tillyardiellidae HANDLIRSCH, 1937)
(Type genus: Pinegia MARTYNOV, 1928; =
Thnetodes MARTYNOV, 1928; =
Tcholmanvissia ZALESSKY, 1929; =
Tillyardiella MARTYNOV, 1930)
Tillyardiella was erroneously classified
within Protodonata / Palaeodonata by TILLYARD & FRASER
(1938), but was transferred to Saltatoria:
"Ensifera" by SHAROV (1968) and CARPENTER
(1992).
Last Update: 27th November, 2007
© Günter Bechly, Böblingen, 2007