Phylogenetic Systematics of Odonata

© Günter Bechly, Böblingen, 2007

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Phylogenetic Systematics of basal Pterygota and Stem-Group Odonates


Neoptera MARTYNOV, 1923 (= Neopterygota CRAMPTON, 1924)

Palaeoptera MARTYNOV, 1923

Discussion: Some of the characters that have been considered as putative autapomorphies of Palaeoptera have meanwhile been demonstrated to represent symplesiomorphies or convergences: short and bristle-like antennae in adults (invalid character which does neither belong to the groundplan of Odonatoptera, nor Ephemeroptera, since a relatively long antenna is still present in † Protereisma and † Namurotypus, so that it is irrelevant that the alleged long antenna of † Lithoneura lameerei turned out to be a plant remain according to WILLMANN, pers. comm.); enlarged complex eyes (apparently neither belonging to the groundplan of Odonatoptera, nor Ephemeroptera, as indicate by the character state in † Protereismatidae and † Eugeropteridae); aquatic larvae (most likely a symplesiomorphy since the neopterous † Lemmatophoridae had ephemerid-like larvae with abdominal gills).
There is also considerable conflicting evidence in favour of a monophyly of Metapterygota BÖRNER, 1909 (= Odonatoptera + Neoptera): derived type of anterior mandibular articulation (STANICZEK, 1996; regarded as convergence by KUKALOVÁ-PECK, 1991); loss of ventral abductor muscles of mandible (not more than one tentorio-mandibular-muscle is retained); the tentorio-lacinial-muscle is completely reduced (could be a convergence, since this muscle is reduced within Zygentoma, too); superlinguae reduced or suppressed, and the hypopharynx therefore not significantly three-lobed (convergent to Zygentoma, thus a weak character; a somewhat three-lobed hypopharynx is retained or regained in Hemiphlebia and quite indistinctly also in Epiophlebia); reduction of several pterothoracic muscles (phragma II - tergum II, profurcasternum - mesobasalare, furca - 1. axillary); muscular closing apparatus for stigmal openings, with a hairy "weir-plate" and muscles that have their insertion directly on the sclerotised stigmal lip (could be a plesiomorphy that is secondarily absent in Ephemeroptera since this character shall not be present in all Metapterygota); reduction or suppression of paracercus (terminal filum) at least in adults (convergent to † Palaeodictyopteroida); complete suppression of subimaginal stages (at least doubtful, because of palaeozoic "protodonate" larvae with large and expanded, but curved, wing sheaths); presence of a nerve connection between corpora allata and corpora cardiaca (a weak character, since also present by convergence in some "apterygotes" and the derived ephemerid Prosopistoma); two derived changes in the ribosomal DNA, including one restriction site change and one 18S insertion (WHEELER in FERNHOLM, 1989).
In spite of this conflicting evidence, I preliminarily prefer the Palaeoptera hypothesis, since it seems to be supported by the better evidence, especially by strong wing venational autapomorphies for Eupalaeoptera, while no wing venational autapomorphies at all are known for Metapterygota. The wing venation of the most basal representatives of Ephemeroptera († Bojophlebiidae and † Protereismatidae) and Odonatoptera († Eugeropteridae and † Erasipteridae) is strikingly similar in several derived features that can certainly not be postulated to belong to the groundplan of † Palaeodictyopteroida or Neoptera, and thus strongly suggest a unique common groundplan of Ephemeroptera and Odonatoptera.
Finally, there is some new evidence (WILLKOMMEN, pers. comm.) that the palaeopterous wing folding could indeed be apomorphic rather the plesiomorphic, and thus independently derived within † Palaeodictyopteroida and in Eupalaeoptera.

Palaeodictyopteroida (sensu BECHLY, 1996a) (= Palaeodictyopteroidea sensu auct.; = † Protorrhynchota ROHDENDORF, 1968)

Comment: the supra-ordinal name Palaeodictyopteroidea has to be rejected since the suffix "-oidea" is reserved for superfamilies.
The branched CuP in the groundplan of † Palaeodictyopteroida is a remarkable plesiomorphy that might even indicate a sistergroup relationship with all other Pterygota, since it otherwise only occurs in some † Protorthoptera and † Titanoptera, most likely as a reversal.
If the ability of wing flexing should be a symplesiomorphy of Neoptera and † Diaphanopterodea, the latter group must be the most basal clade within † Palaeodictyopteroida, while the other three groups would form a monophylum with † "Palaeodictyoptera" as most basal grade, and † Megasecoptera and † Permothemistida as sister-groups. However, if the ability of wing flexing should be a convergence of Neoptera and † Diaphanopterodea, the phylogenetic relationships within † Palaeodictyopteroida could be as follows: the paraphyletic † "Palaeodictyoptera" are the most basal group of † Palaeodictyopteroida. The † Megasecoptera, † Diaphanopterodea, and † Permothemistida (= † Archodonata) together form a monophyletic group, with † Megasecoptera as basal clade and † Diaphanopterodea and † Permothemistida being sistergroups (BECHLY, unpubl.).
The † Synthonopterodea are certainly no † Palaeodictyopteroida (contra BECHLY, 1996a), but most likely belong to the basal stemgroup of Ephemeroptera, and include the † Bojophlebiidae. The alleged "beak", long antennae, and "prothoracic paranotal lobes" of † Lithoneura lameerei were based on misinterpretations according to WILLMANN (1997).
An alleged protodonate from the Lower Triassic of Germany was described by KUHN (1937) as † Thuringopteryx gimmi from the Chirotherium-sandstone (Middle "Buntsandstein") of Saalfeld in Thuringia / eastern Germany. This species was also regarded as protodonate (probably new family) by MÜLLER (1965), although it had already been transferred to Ensifera incertae sedis (possibly belonging to Haglidae) by ZEUNER (1939), followed by CARPENTER (1992). According to the better illustrations in the redescription by MÜLLER (1965) the attribution to Ensifera has to be rejected, because of the long unbranched veins ScP, RA and RP1. It is also certainly neither an Odonatoptera, nor an Ephemeroptera, since the intercalary veins IR1 and IR2 are absent. According to BECHLY (1997i) the wing venation identifies † Thuringopteryx as a member of † Palaeodictyoptera, most likely related to Spilapteridae. This is quite remarkable, since previously there were no Triassic Palaeodictyopteroida known at all, and the whole group was believed to have suffered extinction in the Permian.

Eupalaeoptera Bechly, 2003e (= Hydropalaeoptera sensu KUKALOVÁ-PECK)

Ephemeroptera (= Panephemeroptera CRAMPTON, 1928)

Comment: the † Triplosobidae are here regarded as Palaeoptera incertae sedis, and the † Syntonopterodea, including † Bojophlebiidae, are preliminarily regarded as most basal stemgroup representatives of Ephemeroptera, while the † Permoplectoptera, e.g. † Protereismatidae, certainly belong to the stemgroup of Ephemeroptera (STANICZEK pers. comm.).

Odonatoptera MARTYNOV, 1932

Geroptera BRODSKY, 1994

Comment: the two monospecific genera † Eugeropteron RIEK & KUKALOVÁ-PECK, 1984 and † Geropteron RIEK & KUKALOVÁ-PECK, 1984 could represent a single species since the described differences could rather be due to intraspecific variability, as indicated by a recently discovered and still undescribed specimen (KUKALOVÁ-PECK pers. comm., and pers. observ. on a cast of the referring specimen).

Neodonatoptera BECHLY, 1996

Comment: Many of the characters that are not referring to the wings might also represent synapomorphies of the Odonatoptera since they are not known in † "Eugeropteridae", or of a more subordinate group since the body characters of † "Erasipteridae" are incompletely known either.

"Eomeganisoptera" ROHDENDORF, 1962

Comment: maybe a paraphyletic group which represents the most basal grade of Neodonatoptera, as indicated by the uniquely retained archaedictyon of † Erasipteron, the still retained prothoracic winglets in † Erasipteroides, and the still relatively long median stem.


Meganisoptera MARTYNOV, 1932

Comment: TILLYARD & FRASER (1938: 140-142) and FRASER (1957: 21) caused considerable confusion (e.g. see TRUEMAN, 1991) through their most unconventional redefinition of the order Protodonata (CARPENTER, 1960, 1992). They did not synonymise the latter group with Meganisoptera, but instead restricted Protodonata to include just a few non-odonatoid (mostly palaeodictyopteroid) families, while they included Meganisoptera as suborder of Odonata.

Namurotypidae BECHLY, 1996

(Type genus: † Namurotypus BRAUCKMANN & ZESSIN, 1989)

Meganeuromorpha PRITYKINA, 1980

Paralogidae HANDLIRSCH, 1906

(Type genus: † Paralogus SCUDDER, 1893.)

Kargalotypidae ZESSIN, 1983

(Type genus: † Kargalotypus ROHDENDORF, 1962.)

Kohlwaldiidae GUTHÖRL, 1962

(Type genus: † Kohlwaldia GUTHÖRL, 1962.)

Meganeuridae HANDLIRSCH, 1906

(Type genus: † Meganeura BRONGNIART, 1885.)

Carpentertypinae ZESSIN, 1983

(Type genus: † Carpentertypus ZESSIN, 1983.)

Tupinae HANDLIRSCH, 1919

(Type genus: † Tupus SELLARDS, 1906; = † Typus SELLARDS, 1909, unjustified emendation, rejected by ICZN Opinion 1317, 1984, objectively invalid name no. 2161 on the Official Index.)

Comment: most likely even in this very restricted composition still a paraphyletic group.

Meganeurinae HANDLIRSCH, 1906

(Type genus: † Meganeura BRONGNIART, 1885.)

Odonatoclada BECHLY, 2003e

Lapeyriidae NEL & GAND & GARRIC, 1999

(Type genus: † Lapeyria NEL & GAND & GARRIC, 1999.)

Campylopterodea ROHDENDORF, 1962

Comment: unfortunately † Campyloptera is not well-known and many important character states (e.g. discoidal area) are unknown. However, the shape of the costal margin strongly suggests that a true nodus was present in basal position (putative synapomorphy with Nodialata), vein MA is unbranched (putative synapomorphy with Nodialata), and even an apical pterostigma seems to have been present according to CARPENTER (1943), which is a derived similarity with Stigmoptera (putative synapomorphy?). Also the petiolated wing could represent a putative synapomorphy with Discoidalia, and the complete suppression of a free AA2 could represent a further putative synapomorphy with Stigmoptera. On the other hand, the plesiomorphic presence of a long free vein CuP excludes a position within crowngroup Stigmoptera. Therefore, most likely † Campylopterodea is the sistertaxon to Stigmoptera (together constituting the new taxon Panstigmoptera), but I here preliminarily retain † Campylopterodea as a taxon incertae sedis within Odonatoclada, until a redescription of the holotype supports the evidence for the mentioned phylogenetic hypothesis.

Nodialata BECHLY, 1996

Protanisoptera CARPENTER, 1931

Polytaxineuridae TILLYARD, 1935

(Type genus: † Polytaxineura TILLYARD, 1935.)

Ditaxineuromorpha PRITYKINA, 1980

Permaeschnidae MARTYNOV, 1931 (sens. nov.)

(Type genus: † Permaeschna MARTYNOV, 1931.)

Comment: the attribution of † Gondvanoptilon to † "Erasipteridae" by MARTINS-NETO (1996) is only based on symplesiomorphic similarities and ignores the very strong synapomorphies with † Protanisoptera and † Ditaxineuromorpha. † Gondvanoptilon brasiliense (the spelling change to brasiliensis by Martins-Neto was not justified, since ptlion is neutrum!) is so similar to † Permaeschna dolloi that even its status as a distinct genus must be regarded as very doubtful.

Ditaxineurida BECHLY, 2003e

Callimokaltaniidae ZALESSKY, 1955 (stat. rest.)

(Type genus: † Callimokaltania ZALESSKY, 1955.)

Ditaxineuroidea TILLYARD, 1926

(Type genus: † Ditaxineura TILLYARD, 1926.)

Hemizygopteridae ZALESSKY, 1955

(Type genus: † Hemizygopteron ZALESSKY, 1955.)

Ditaxineuridae TILLYARD, 1926

(Type genus: † Ditaxineura TILLYARD, 1926.)

Last Update: 16th August, 2007

© Günter Bechly, Böblingen, 2007