Phylogenetic Systematics of Odonata

© Günter Bechly, Böblingen, 2007

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Phylogenetic Systematics of "Anisozygopteres"

Epiproctophora BECHLY, 1996

Isophlebioptera BECHLY, 1996

Euthemistidae PRITYKINA, 1968

(Type genus: † Euthemis PRITYKINA, 1968.)

Comment: Both mentioned putative autapomorphies are also present in † Mesoepiophlebia veronicae, either by convergence, or as synapomorphy. Furthermore the characteristic pattern of intercalary veins is convergently also present in † Erichschmidtiidae and † Stenophlebiidae, but † Euthemistidae are differing from the former by the presence of secondary antenodal crossveins between costal margin and ScP, and from both groups by the lack of any intercalaries in the extremely narrow postdiscoidal area, and by the presence of pektinate convex "secondary branches" of RP3/4 and MP. The presence of secondary antenodal crossveins between the costal margin and ScP could indicate a sistergroup relationship of † Euthemistidae and all other † Isophlebioptera, but on the other hand such crossveins are also present in a few subgroups of † Isophlebioptera as obvious reversals, so that an autapomorphic reversal cannot be totally excluded for † Euthemistidae either.

Parazygoptera BECHLY, 1997

Comment: the petiolation of the hindwing most probably represents a plesiomorphy, since this state is also present in † Campyloptera, † Triadophlebiomorpha, † Protozygoptera, Zygoptera, † Tarsophlebiidae, Epiophlebiidae, † Heterophlebioptera, and † Stenophlebioptera. The secondary loss of the lestine oblique vein in many † Parazygoptera is clearly due to a multiple convergence (since this veinlet is retained in † Mesoepiophlebia, † Ensphingophlebia, † Cyclothemis and † Shurabiola, and maybe also in † Pseudotriassothemis and † Mesophlebia), probably caused by the shifting of the origin of RP2 distal of the subnodus, as a pre-disposition.

Sphenophlebiidae BECHLY, 1997

(Type genus: † Sphenophlebia BODE, 1953.)

Euparazygoptera BECHLY, 1997

Asiopteridae PRITYKINA, 1968

(Type genus: † Asiopteron PRITYKINA, 1968.)

Comment: † "Sphenophlebia" pommerana ANSORGE, 1996 was transferred to the genus † Turanopteron PRITYKINA, 1968 by BECHLY (1997i) because of the nearly identical venation that was already recognized by ANSORGE (1996). † Oreopteron asiaticum PRITYKINA, 1968 and † Oreopterella paula PRITYKINA, 1968 seem to be closely related, since they uniquely share a zigzagged RP1, while † "Oreopteron" simile PRITYKINA, 1968 has to be transferred to the genus † Asiopteron PRITYKINA, 1968. Since † Oreopteridae and the genus † Oreopteron are not monophyletic, and † Asiopteridae lacks any autapomorphies if † Oreopteridae are excluded, † Oreopteridae is here regarded as a synonym of † Asiopteridae. A detailed phylogenetic system of † Asiopteridae and several new genera and species will be introduced in NEL et al. (in prep.). CARLE & WIGHTON (1990: 58) suggested that † Oreopteridae and † Asiopteridae are synonyms of † Progonophlebiidae, although they did not justify this hypothesis, which has to be rejected according to BECHLY (1997i) (also see NEL et al., 1993).
Sogjutella is certainly no member of † Asiopteridae (= † Oreopteridae) since it shares the unique kink in RA between nodal and subnodal veinlet as strong synapomorphy with † Triassolestoidea. Likewise † "Sogdopteron" legibile shares a very complex synapomorphy in the anal area with † Triassolestini.

Triassolestoidea TILLYARD, 1918

(Type genus: † Triassolestes TILLYARD, 1918.)

Cyclothemistidae BECHLY, 1996

(Type genus: † Cyclothemis PRITYKINA, 1980.)

Cyclothemistinae BECHLY, 1996

(Type genus: † Cyclothemis PRITYKINA, 1980.)

Comment: the sister-genera † Shurabiola and † Sogjutella share the complete suppression of all secondary antenodal and all antesubnodal crossveins as putative synapomorphies. Autapomorphies of † Sogjutella are the oblique distal side (MAb) of the discoidal cell (reversal), the suppression of the lestine oblique vein between RP2 and IR2, and the strongly reduced anal and cubito-anal area in the hindwing, with only one row of cells, which is correlate with the generally reduced venation and the slender shape of the wing.

Pseudotriassothemistinae BECHLY, 1997

(Type genus: † Pseudotriassothemis BECHLY, 1997)

Comment: † P. okafujii and † P. nipponensis at least are sister-species (synapomorphy: unique crossvein-brace between RP and IR2 below the subnodus) or might even represent fore and hindwing of the same species, but this could only be positively demonstrated if a specimen with both pairs of wings preserved together would be found. An autapomorphy of † P. minensis seems to be the secondary presence of secondary antenodal crossveins between costal margin and ScP distal of ax2 (convergent to † Mesophlebiinae and † Italophlebia).

Triassolestidae TILLYARD, 1918

(Type genus: † Triassolestes TILLYARD, 1918.)

Mesophlebiinae TILLYARD, 1916

(Type genus: † Mesophlebia TILLYARD in TILLYARD & DUNSTAN, 1916.)

Comment: † Mesophlebia tillyardi HANDLIRSCH, 1939 is a junior subjective synonym of † Mesophlebia antinodalis TILLYARD, 1916 according to COWLEY (1942). † Progonophlebia cromptoni ZEUNER, 1958 (nec "cramptoni": NEL et al., 1993) is an junior objective synonym of † Progonophlebia woodwardi TILLYARD, 1925 according to NEL & JARZEMBOWSKI (1997), since the two referring holotypes are indeed part and counterpart of the same specimen.
Unique autapomorphies of † Mesophlebia are an oblique vein that is obliquely slanted towards the apex between RP and IR2 below the transverse subnodus, the reversed obliquity of the nodal crossvein (convergent to † Triassolestodes), and a more distinct constriction of the space between RA and RP1 beneath the pterostigma.

Triassolestinae TILLYARD, 1918

(Type genus: † Triassolestes TILLYARD, 1918.)

Triassothemistini FUJIYAMA, 1991

(Type genus: † Triassothemis CARPENTER, 1961.)

Comment: distinct autapomorphies of † Italophlebia are the distally strongly converging veins RP1 and RP2, the distally zigzagged vein MA, and the shortened and zigzagged IR1. Autapomorphies of the new subgenus † Afrotriassothemis BECHLY, 1997 are the numerous cells below the elongated pterostigma, very long and straight IR1, RP2 and IR2 basally very close together fore a longer distance, and two rows of cells between the distal parts of IR2 and RP3/4.

Triassolestini TILLYARD, 1918

(Type genus: † Triassolestes TILLYARD, 1918.)

Discussion: a strong synapomorphy of all † Triassolestini, except the most basal † "Sogdopteron" legibile, is a unique veinal pattern in the forewing base, in which [AA & CuP] is basally fused with the hind margin, and distally fused with [MP & CuA], only two short portions remained free, the basal one looking like the CuP-crossing, and the distal one as oblique "pseudo-subdiscoidal veinlet" beneath the tip of the discoidal cell (unique within Odonatoptera).
A distinct synapomorphy of † Triassolestodes and † Triassoneura is the short basal fusion of MP and CuA distal of the forewing discoidal cell (unique within Odonatoptera, but similar to the hindwing of † Tarsophlebiidae). A further synapomorphy of these two genera might be the circumstance that ax2 is secondarily shifted basal of the arculus (reversal), but this character is unknown in † Triassolestes.
The holotypical wing of † Triassolestes looks very similar to the forewing of † Triassoneura and † Triassolestodes. The only conflicting character would be the closed discoidal cell, but according to NEL (pers. comm.) this character state is based on an error of Tillyard. An unusual feature (autapomorphy) of † Triassolestes is the distally widened postdiscoidal area with more than one row of cells in the widened part. However, such an autapomorphic reversal happened in † Cyclothemistidae, too.
Autapomorphies of † Triassolestodes are the reversed obliquity of the nodal crossvein (convergent to † Mesophlebia), a unique type of hindwing arculus (NEL pers. comm.), and the reduced wing venation with only one row of cells between all the longitudinal veins, giving the wing a zygopteroid appearance. Distinct autapomorphies of † "Sogdopteron" legibile PRITYKINA, 1980 are the basally zigzagged RP2 and the distal obliteration of MA. The distally converging veins MA and MP may be a derived similarity with † Triassothemistini, but the position of the arculus closer to ax2 than to ax1 has to be regarded as stronger evidence for a relationship with † Triassolestini, since this character state is a very rare reversal within Epiproctophora.

Selenothemistidae HANDLIRSCH, 1939

(Type genus: † Selenothemis HANDLIRSCH, 1920.)

Isophlebiida BECHLY, 1996

Archithemistidae TILLYARD, 1917

(Type genus: † Archithemis HANDLIRSCH, 1906; = † Diastatommites TILLYARD, 1925 nom. subst. pro. † Diastatomma GIEBEL, 1856, nec Diastatomma BURMEISTER, 1839; = † Diastatommites HANDLIRSCH, 1920, nomen nudum.)

Comment: † Petrophlebia anglicana was already correctly placed by FRASER (1957), and later transferred to † Campterophlebiidae by NEL et al. (1993), but in both cases without explication. Thus † Petrophlebiidae is not a synonym of † Campterophlebiidae (contra NEL et al., 1993). Although there are only few putative synapomorphies known for † Archithemistidae, this taxon very probably is monophyletic since the three included species are so similar that they could easily be classified in the same genus.

Isophlebioidea HANDLIRSCH, 1906

(Type genus: † Isophlebia HAGEN, 1866.)

Campterophlebiidae HANDLIRSCH, 1920

(Type genus: † Campterophlebia BODE, 1905.)

Comment: the only Triassic (Raetian) species † Sogdophlebia singularis PRITYKINA, 1970 was originally described in † Heterophlebiidea - † Liassophlebiidae (sensu Pritykina), but transferred to † Isophlebioidea - † Campterophlebiidae by NEL et al. (1993). This placement is also endorsed by BECHLY (1997i).

Isophlebiidae HANDLIRSCH, 1906

(Type genus: † Isophlebia HAGEN, 1866.)

Euepiproctophora BECHLY, 2003e

Comment: the hypothesis that Epiophlebia is closer related with Anisoptera than † Isophlebioptera is convincingly demonstrated by the mentioned larval synapomorphies. The presence of a synlestine type of larva (with three caudal gills of characteristical shape including a median gill that is formed by a long epiproct; no anal pyramid; abdomen very elongate and slender) in basal extant Zygoptera (Hemiphlebiidae, Synlestidae) and fossil isophlebioid larvae, as well as the oldest known odonate larva from the Triassic of Australia, is such a distinct and complex similarity that it cannot be reasonably regarded as convergence, but only as a symplesiomorphy. Therefore the strongly Anisoptera-like-larvae (with a shortened and broadened abdomen with well-developed rectal gills and with suppressed caudal gills) represent a very strong synapomorphy for Epiophlebiidae and Anisoptera (but not † Isophlebioptera). The attribution of the referring fossil larvae (e.g. † "Samarura" gigantea) from the Lower Cretaceous of Sibiria to † Isophlebioptera is absolutely certain, since FLECK and NEL (pers. comm. 1998) could recently identify the synapomorphic characters of the isophlebioid wing venation on the wing sheath of one of the referring fossil larvae at PIN in Moscow. Furthermore, the only fossil adult odonates in the same layers are large isophlebioids, while the only occuring fossil odonate larvae are these very large † "Samarura" larvae (length about 12.5 cm!).
FRASER (1957) proposed that a tendency towards a division of the hindwing discoidal cell into an anterior hypertriangle and a posterior discoidal triangle might indicate a close relationship of Epiophlebia and Anisoptera, too. However, the division of the hindwing discoidal cell is very variable Epiophlebia, and the formation of a true hypertriangle is even still variable in † Liassophlebia. Furthermore I regard the alleged division of the hindwing discoidal cell in some specimens of Epiophlebia as based on an erroneous interpretation anyway, since the referring crossvein is clearly developed in the median space, forming a second "arculus", while the discoidal cell is never divided at all.

Epiophlebiidae MUTTKOWSKI, 1910

(Type genus: Epiophlebia CALVERT, 1903 nom. subst. pro Palaeophlebia SELYS, 1889, nec † Palaeophlebia BRAUER, 1889; = Neopalaeophlebia HANDLIRSCH, 1906, jun. obj. syn.)

Comment: according to LINDEBOOM (pers. comm.) the secondary male genital apparatus of Epiophlebia has several apparently plesiomorphic features that could be regarded as evidence for a sistergroup relationship of Epiophlebia with the remaining extant Odonata, since Zygoptera and Anisoptera share the putative apomorphic states (ligula detached and lifted from sternum, plate-like hamuli anteriores, and a lamina anterior with a median depression). However, the polarity of the latter two character states (which are correlated anyway) is doubtful, since † Tarsophlebia, which certainly has a more basal position than Epiophlebia, also has plate-like hamuli anteriores and a median depression in the lamina anterior, which is strongly suggesting a symplesiomorphy of † Tarsophlebiidae with Zygoptera and Anisoptera. Therefore, the hook-like hamuli anteriores and the entire lamina anterior rather have to be regarded as autapomorphic reversals of Epiophlebia. A sistergroup relationship of † Tarsophlebia with all extant Odonata (incl. Epiophlebia) is well-supported by the following unique plesiomorphies of † Tarsophlebia: ligula orimentary (same shape as in the early ontogenesis in Zygoptera); vesicula spermalis still very short and flat with a very wide porus; hindwing discoidal cell basaly still open (arculus incomplete); legs clearly still with four tarsomeres of equal length. All extant Odonata share the referring apomorphic states as synapomorphies. Consequently, only the "primitive" ligula (and maybe the absence of a discal node) remains as potential evidence for a most basal position of Epiophlebia within extant Odonata. The Tarsophlebia-like ligula in the ontogenesis of extant Zygoptera (e.g. Calopteryx) indicates that the apparently plesiomorphic ligula of Epiophlebia could as well be explained by paedomorphosis, correlated with the secondary loss of function of the ligula by the autapomorphic enlargement of the hamuli posteriores, whichtake over the function as sperm intromittent organ. The undisputet autapomorphic enlargement of the hamuli posteriores also implies an increased need for mechanical stability, which might also explain the secondary loss of the median depression of the lamina anterior.

Anisopteromorpha BECHLY, 1996

Erichschmidtiidae BECHLY, 1996

(Type genus: † Erichschmidtia PRITYKINA, 1968.)

Comment: † Trigonophlebia shares with † Erichschmidtia a rather unique cell below the subdiscoidal cell, convergent to † Tarsophlebiopsis mayi, but does have secondary antenodal crossveins between costal margin and ScP that would conflict a sistergroup relationship of these two genera. Most probably the † Erichschmidtiidae represent the sistergroup of all other Anisopteromorpha, although it cannot be totally excluded that they might either belong to † Heterophlebioptera, or even to † Isophlebioptera, e.g. as possible sistergroup of † Euthemistidae.

Heterophlebioptera BECHLY, 1996

Myopophlebiidae BODE, 1953

(Type genus: † Myopophlebia BODE, 1953.)

Heterophlebioidea NEEDHAM, 1903

(Type genus: † Heterophlebia BRODIE, 1849.)

Liassophlebiidae TILLYARD, 1925

(Type genus: † Liassophlebia TILLYARD, 1925.)

Comment: the genus † Grimmenopteron which is only known by a single forewing, shares the two respective autapomorphies of † Liassophlebiidae, but on the other hand has retained two unique plesiomorphies within † Heterophlebioidea. The very large size of most † Liassophlebiidae might represent an ingroup synapomorphy for the referring species. Therefore the smaller size of † Grimmenopteron could rather be a plesiomorphy than an autapomorphy (contra ANSORGE, 1996).

Heterophlebiidae NEEDHAM, 1903

(Type genus: † Heterophlebia BRODIE, 1849.)

Trigonoptera BECHLY, 1996

Stenophlebioptera BECHLY, 1996

Gondvanogomphidae BECHLY, 1996

(Type genus: † Gondvanogomphus SCHLÜTER & HARTUNG, 1982.)

Stenophlebiidae NEEDHAM, 1903

(Type genus: † Stenophlebia HAGEN, 1866.)

Comment: not including the genus † Prostenophlebia that is most probably closely related to † Erichschmidtia, contra NEL et al. (1993).

Last Update: 10th August, 2007

© Günter Bechly, Böblingen, 2007