Phylogenetic Systematics of Odonata

Euzygoptera BECHLY, 1996

• Included taxa: Lestomorpha and Coenagrionomorpha.
• Autapomorphies:
  
  • Wing venation: longitudinal veins rather straight and long (RP3/4 reaching beyond pterostigma, MA reaching pterostigma, and CuA reaching distal of midwing), with the branches of RP distally converging (reversed in Argiolestinae), and only one row of cells between CuA and the hind margin of wings (convergent to Rimanella, Euarchistigmatini, Chlorocyphoidea, Heliocharitidae, Caliphaeidae; reversed in some "Megapodagrionidae"); only the two primary antenodal brackets ax1 and ax2 are retained in the antenodal space (convergent to Pseudolestidae and † Petrolestini); antesubnodal space without any crossveins (convergent to Pseudolestidae, Thaumatoneuridae and Rimanellinae); no antefurcal crossveins present in the space between RP and MA from arculus to midfork (convergent to many Caloptera, in which this character state is caused by a strong shortening of this wing space).
    
  • Other characters: fusion of the meso- and metathoacic ganglia, convergent to Exophytica (CALVERT, 1893; TILLYARD, 1917; contra ASAHINA, 1957); only few macrotrichae on the wing veins.
• Taxonomy:
  
  • Heteropteroides SELYS, 1840 (taxon nov.)
    
  • Rhomboideae BUCHECKER, 1876 (taxon nov.)
    
  • Euzygoptera BECHLY, 1996a (taxon nov.)

Comment: concerning potential conflicting evidence against the monophyly of Euzygoptera see remarks under Caloptera and Hemiphlebiidae.

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Lestomorpha BECHLY, 1996

• Included taxa: Hemiphlebiidae and Lestiformia.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: derived type of "star-shaped" microsculptures (micrasters) on the pterostigmata (BECHLY, 1994); metallic green body colour (even preserved in an undescribed specimen of † Parahemiphlebia); derived colour pattern of the larvae with striped legs (TILLYARD, 1928; secondarily absent in Lestidae); denticles only present on four large proventricular lobes and arranged in a transverse arch (TILLYARD, 1928); characteristical type of the short larval caudal gill lamellae (TILLYARD, 1928; secondarily prolonged in Lestidae), but maybe a symplesiomorphy since already known from Triassic odonatoid larvae (ROZEFELDS, 1985); male ligula with a ventral fossa (no internal fold) on the second segment (KENNEDY, 1919) that is margined by two parallel chitinised bars (secondarily fused in Perilestes and Megalestes; KENNEDY, 1919).
• Taxonomy:
  
  • [partim: Normostigmatina KIRBY, 1890 (taxon nov.)]
    
  • [Archaeoptera BELYSHEV & HARITONOV, 1985 (taxon nov.)]
    
  • Lestomorpha BECHLY, 1996a (taxon nov.) (nec Lestomorpha PRITYKINA, 1980)

Comment: according to TILLYARD (1928) Hemiphlebia shares with all Lestidae a very similar and derived type of elongated larval prehensile mask: the lateral lobes ("palps") with two large marginal horn-like projections (terminal hooks) and a shorter and serrated median projection, and a movable hook that is supplied with setae. But this is more probably a convergence of Hemiphlebia and Lestidae since Chorismagrionidae, Perilestidae and Synlestidae have a very different type of mask with only two or three acute endhooks and no setae at all.

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Hemiphlebiidae TILLYARD, 1926

(Type genus: Hemiphlebia SELYS, 1868.)

• Included taxa: including the extant relic type genus Hemiphlebia SELYS, 1868, and according to BECHLY (1998d) as well the fossil genus † Parahemiphlebia JARZEMBOWSKI et al., 1998, and maybe † Cretarchistigma JARZEMBOWSKI et al., 1998.
• Autapomorphies:
  
  • Wing venation: only 5-7 (rarely 8) postnodal crossveins between nodus and pterostigma (convergent to a few Protoneuridae and some Coenagrionidae); lestine oblique vein secondarily absent (convergent to Caloptera, Coenagrionomorpha and many † Parazygoptera; in Hemiphlebia correlated with the absence of tracheae in convex veins!); wing base with distinctly reduced petiolation; RP1 kinked at the insertion of the pterostigmal brace vein (convergent to Coenagrionida); all intercalary veins (except IR1 and IR2) suppressed (convergent to Chorismagrionidae, Perilestidae, and Coenagrionida).
    
  • Other characters: tiny size; tibial comb of fore legs degenerated (KENNEDY, 1919); legs light brown with characteristical dark brown patches; hamuli anteriores partly membranous (reduction); wing tracheae of all convex wing veins (except RA) suppressed (TILLYARD, 1928); secondary paired projections on the larval prementum that are most likely not homologous with glossae or paraglossae (contra TILLYARD, 1928), but according to TRUEMAN (1999) they could be homologous with a pair of minute teeth on the mask of all other odonate larvae (except Epiophlebia); "Zipfel" of the male secondary genital apparatus is much reduced in size and the ligula is quite derived (LINDEBOOM pers. comm.); male paraprocts hypertrophied and strikingly white (not yet in † Parahemiphlebia which has normal male appendages, but no reduced appendages as maintained in the original description; BECHLY, 1998d), being displayed in a complex courtship behaviour; resting position with wings closely apposed over the dorsum of the body; spermatozoa biflagellate (TRUEMAN, 1999), which is unique within Odonata and even Insecta.
• Taxonomy:
  
  • Hemiphlebiidae TILLYARD, 1926 (fam. nov.)
    
  • Hemiphlebioidea sensu TILLYARD & FRASER, 1938 (stat. nov.)
    
  • Hemiphlebiidae sensu FRASER, 1957
    
  • Hemiphlebioidea sensu FRASER, 1957
    
  • Hemiphleboidea sensu PINHEY in PARKER, 1982 (unjust.emend.)
    
  • Hemiphlebioidea; WHALLEY, 1986
    
  • [partim: Archaeoptera BELYSHEV & HARITONOV, 1985 (taxon nov.)]

Comment: Hemiphlebia shares with Chorismagrion two unique symplesiomorphies within extant Zygoptera: the basally open discoidal cell in forewings (a parallel reversal is very unlikely!), and the presence of a suture between vertex and occiput (also retained in some Synlestidae and "Megapodagrionidae"). Some of the mentioned autapomorphies of Hemiphlebiidae could be rather autapomorphies of the extant genus Hemiphlebia. Many of the alleged plesiomorphies of Hemiphlebia are better explained as autapomorphic reversals or reductions that are correlated with the miniaturisation of the insect. A previously unrecognized, but indeed extremely rare, plesiomorphic behaviour within Zygoptera is the position of the female in the pairing wheel, with the legs grabbing the male abdomen like in Anisoptera, while in all other extant Zygoptera the female legs are usually free in the air or on the ground.
TRUEMAN (1999) argued that the open forewing discoidal cell could be an autapomorphic reversal (convergent to Chorismagrion), because a few specimens have this cell closed, which is interpreted by this author as a putative atavism. However, this variability could as well be interpreted as derived, similar to the fossil † Liassophlebiidae († Heterophlebioptera), confirming the high likelihood of a multiple convergent (or parallel) closure of the forewing discoidal cell, as soon as the hindwing discoidal cell was closed. Consequently, the variability in Hemiphlebia is no evidence for either hypothesis.

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Lestiformia BECHLY, 1996

• Included taxa: † Cretacoenagrionidae and Eulestiformia.
• Autapomorphies:
  
  • Wing venation: postnodal and postsubnodal crossveins aligned (convergent to Coenagrionomorpha ?).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Lestiformia BECHLY, 1996a (taxon nov.)

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† Cretacoenagrionidae BECHLY, 1996

(Type genus: † Cretacoenagrion JARZEMBOWSKI, 1990.)

• Included taxa: only including the type species † Cretacoenagrion alleni JARZEMBOWSKI, 1990.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Cretacoenagrionidae BECHLY, 1996a (fam. nov.)

Comment: the attribution of † Cretacoenagrion to Lestomorpha rather than Coenagrionidae is based on the most parsimonious interpretation of the character pattern. Even though there are no strong synapomorphies with Lestomorpha and Lestiformia known at all, the suggested position requires the fewest number of character gains or reversals. For example the plesiomorphic presence of an open discoidal cell and a lestine oblique vein would contradict any position within Coenagrionomorpha. Phenetically the wing venation of this fossil taxon is nearly identical to the extant genus Chorismagrion, only differing in the plesiomorphic straight course of MP.

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Eulestiformia BECHLY, 1996

• Included taxa: Chorismagrionidae and Lestida.
• Autapomorphies:
  
  • Wing venation: MP distinctly curved after its origin at the distal angle of the discoidal cell (convergent to † Triadophlebiomorpha and † Steleopteridae).
    
  • Other characters: male secondary genitalia with elongated, leaf-shaped hamuli anteriores (FRASER, 1957), and suppressed terminal projections of ligula (KENNEDY, 1919; maybe an autapomorphy of Lestiniformia since † Cretacoenagrion is only known as a single wing); larval gizzard (proventriculus) with only eight folds.
• Taxonomy:
  
  • [légion Lestes SELYS, 1840 (not available as family-group taxon according Art.11f IRZN)]
    
  • Lestinoidea sensu auct. (except BECHLY, 1996a)
    
  • Eulestiformia BECHLY, 1996a (taxon nov.)

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Chorismagrionidae TILLYARD & FRASER, 1938

(Type genus: Chorismagrion MORTON, 1914.)

• Included taxa: only including the extant relic type species Chorismagrion risi MORTON, 1914.
• Autapomorphies:
  
  • Wing venation: the origin of IR2 is shifted several cells distal of the midfork (this character could well be regarded as a strong potential synapomorphy with Perilestidae since it is a quite unique derived state within Odonata, only present in these two groups and the "megapodagrionid" genus Arrhenocnemis); the posterior margin of the subdiscoidal cell [CuP & AA'] is mostly fused to the hind margin (a potential synapomorphy with Perilestidae; convergent to Synlestidae); all intercalary veins (except IR1 and IR2) suppressed (a potential synapomorphy with Perilestidae; convergent to Hemiphlebiidae and Coenagrionida).
    
  • Other characters: secondarily non-metallic dull body coloration (also a potential synapomorphy with Perilestidae).
• Taxonomy:
  
  • Chorismagriinae TILLYARD & FRASER, 1938 (subfam. nov.) (nom. imperf.)
    
  • Chorismagriinae sensu FRASER, 1957
    
  • Chorismagrioninae sensu ST. QUENTIN & BEIER, 1968 (nom. correct.)
    
  • Chorismagrioninae sensu O’FARREL, 1970
    
  • Chorismagrioninae sensu DAVIES, 1981 (sens. nov.)
    
  • [partim: Archaeoptera BELYSHEV & HARITONOV, 1985 (taxon nov.)]
    
  • Chorismagrionidae sensu BRIDGES, 1993 (stat. nov.)

Comment: it cannot be excluded that Chorismagrionidae is rather the sistergroup of Perilestidae because of the mentioned potential synapomorphies. This was already suggested by FRASER (1957), but would of course imply that the closed discoidal cell was independantly acquired in Perilestidae which is by no means unlikely since this character has evolved several times within Zygoptera and Epiproctophora anyway.

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Lestida BECHLY, 1996

• Included taxa: Perilestidae and Lestodea.
• Autapomorphies:
  
  • Wing venation: basal closure of discoidal cell in forewings including the development of a dorsal arcular bracket (convergent to Caloptera, Coenagrionomorpha, Epiophlebiidae, † Asiopteridae, † Isophlebiidae, and Anisopteromorpha).
    
  • Other characters: adult males with very long forcep-like cerci; adult males without hairs or spines on the ligula shaft (KENNEDY, 1919; convergent to Chlorocyphoidea and Platycnemididae).
• Taxonomy:
  
  • Lestida BECHLY, 1996a (taxon nov.)

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Perilestidae TILLYARD & FRASER, 1938

(Type genus: Perilestes HAGEN in SELYS, 1862.)

• Included taxa: Nubiolestinae and Perilestinae.
• Autapomorphies:
  
  • Wing venation: wings very slender and extremely petiolated; nodus in a very basal position (about 26-28 % of wing length; convergent to Platystictidae and Pseudostigmatoidea); basal half of RA and [RA & RP] greatly thickened (FRASER, 1957); dorsal arcular bracket reduced (convergent to Lestidae, Polythoridae and Eurypalpida); apex of discoidal cell is close to the hind margin of the wing; IR2 distinctly shortened (convergent to Chorismagrionidae ?)and arising close to the origin of RP2 or even on RP2; the posterior margin of the subdiscoidal cell [CuP & AA'] is mostly fused to the hind margin (convergent to Synlestidae and Chorismagrionidae ?); all intercalary veins (except IR1 and IR2) suppressed (convergent to Hemiphlebiidae, Chorismagrionidae and Coenagrionida).
    
  • Other characters: tarsal claws secondarily without ventro-apical hook (convergent to Leptobasinae and Teinobasini, and Onychothemistina); abdomen extremely long and slender; adult males with vestigial paraprocts; secondary non-metallic body coloration (convergent to Chorismagrionidae ?).
• Taxonomy:
  
  • Perilestidae; FRASER, 1957 (stat. nov.) (nom. transl. ex Perilestinae TILLYARD & FRASER, 1938)
    
  • Perilestinae sensu FRASER, 1957

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Nubiolestinae BECHLY, 1996

(Type genus: Nubiolestes FRASER, 1945 nom. subst. pro Eolestes FRASER, 1944 which is an jun. obj. syn. and homonoym of Eolestes SCHMIDT, 1943 and a homonym of Eolestes COCKERELL, 1940; = Camerunolestes SCHMIDT, 1958 nom. subst. pro Eolestes SCHMIDT, 1943, nec Eolestes COCKERELL, 1940.)

• Included taxa: only including the type genus Nubiolestes FRASER, 1945.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Nubiolestinae BECHLY, 1996a (subfam. nov.)

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Perilestinae TILLYARD & FRASER, 1938

(Type genus: Perilestes HAGEN in SELYS, 1862.)

• Included taxa: only including the two neotropical genera Perilestes HAGEN in SELYS, 1862 and Perissolestes KENNEDY, 1941.
• Autapomorphies:
  
  • Wing venation: wing venation less reticulate; midfork shifted distinctly distal of the nodus; lestine oblique vein reduced; together with the reduction of the subdiscoidal vein (basal CuA), the discoidal bracket (dorsal sclerotisation on MAb and the subdiscoidal vein) is strongly reduced (convergent to Protoneuridae and Anisoptera); apex of discoidal cell touching the hind margin of the wing; IR1 strongly shortened, and IR2 further shortened, too; postnodal crossveins aligned with the rows of crossveins below, forming several pseudo-transverse-veins (convergent to some "Megapodagrioninae" and all Coenagrioniformia); anal vein [CuP & AA'] is completely fused with the hind margin.
    
  • Other characters: larvae with a dense medio-longitudinal row of small spines on the abdominal terga.
• Taxonomy:
  
  • Perilestinae TILLYARD & FRASER, 1938 (subfam. nov.)
    
  • Perilestinae sensu BECHLY, 1996a (sens. nov.)

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Lestodea BECHLY, 1996

• Included taxa: Synlestidae and Lestinoidea.
• Autapomorphies:
  
  • Wing venation: arculus shifted basally beneath the ax2.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Lestodea BECHLY, 1996a (taxon nov.)

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Synlestidae TILLYARD, 1917

(Type genus: Synlestes SELYS, 1868.)

• Included taxa: including the genera listed in BRIDGES (1994) for Synlestinae.
• Autapomorphies:
  
  • Wing venation: the posterior margin of the subdiscoidal cell [CuP & AA'] is mostly fused to the hind margin (convergent to Chorismagrionidae and Perilestidae).
    
  • Other characters: pedicel segment of larval antenna strongly prolonged (maybe not a groundplan character since only pronounced in the genus Synlestes; convergent to Chlorocyphidae and Calopterygida); interpleural suture of the pterothorax secondarily well-developed (reversal that might be easily explained with paedogenesis; convergent to Calopterygiformia).
• Taxonomy:
  
  • Synlestinae TILLYARD, 1917 (subfam. nov.)
    
  • Synlestidae; nom. transl. TILLYARD, 1926 (stat. nov.)
    
  • Chlorolestidae FRASER, 1951 (fam. nov. with the type genus Chlorolestes SELYS, 1862) (jun. subj. syn.)
    
  • Chlorolestinae FRASER, 1951 (subfam. nov. with the type genus Chlorolestes SELYS, 1862) (jun. subj. syn.)
    
  • Synlestidae sensu BECHLY, 1996a (sens. nov.)

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Lestinoidea CALVERT, 1901

(Type genus: Lestes [LEACH] [1815]; = Puella BRULLÉ, 1832, jun. subj. syn.; = Anapates CHARPENTIER, 1840, jun. subj. syn.; = Anapetes PINHEY, 1980, incorr. subseq. spell.)

• Included taxa: Megalestidae and Lestidae.
• Autapomorphies:
  
  • Wing venation: the midfork is recessed basally to a position of 20-26 % of wing length (KENNEDY, 1919; convergent to Caloptera and Hypolestinae), therefore the subnodus is located between the bases of RP2 and IR2 that are widely separated (convergent to Hypolestinae); MA at least distally zigzagged.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Lestini FRASER, 1951 (stat. nov., superfamily not tribe) (nom. imperf.)
    
  • Lestoidea; FRASER, 1954 (nom. correct.) (nec Lestoidea TILLYARD, 1913)
    
  • Lestinoidea; FRASER, 1957 (unjustified emendation of Lestoidea FRASER, 1954, nec Lestoidea TILLYARD, 1913, since family-group taxa do not compete in homonymy with genus-group taxa according to IRZN)
    
  • Lestidoidea; DAVIES, 1981 (incorr. subseq. spell.)
    
  • Lestionidea sensu BECHLY, 1996a (sens. nov.)

Comment: the suprafamily name Lestoidea has an identical spelling as the older generic name Lestoidea TILLYARD, 1913. Therefore, Fraser (1957) introduced the replacement name Lestinoidea which is not in accordance with the present rules of zoological nomenclature since genus-group names do not compete with family-group names for synonymy or homonymy. Nevertheless I preliminarily followed the suggestion of Fraser to avoid misunderstandings.

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Megalestidae TILLYARD & FRASER, 1938

(Type genus: Megalestes SELYS, 1862.)

• Included taxa: including the type genus Megalestes SELYS, 1862.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Megalestinae TILLYARD & FRASER, 1938 (subfam. nov.)
    
  • Megalestidae sensu BECHLY, 1996a (stat. nov.)

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Lestidae CALVERT, 1901

(Type genus: Lestes [LEACH] [1815]; = Puella BRULLÉ, 1832, jun. subj. syn.; = Anapates CHARPENTIER, 1840, jun. subj. syn.; = Anapetes PINHEY, 1980, incorr. subseq. spell.)

• Included taxa: Sympecmatinae and Lestinae.
• Autapomorphies:
  
  • Wing venation: the distal discoidal vein MAb is very oblique, so that the distal angle of the discoidal cell is very acute (not in the genus Archilestes; convergent to Coenagrionidae); nodal veinlet less oblique and anterior part of the ventral subnodal bracket strongly thickened (convergent to Heliocharitidae); dorsal arcular bracket reduced (convergent to Perilestidae ?, Polythoridae and Eurypalpida); MA more strongly zigzagged; area between IR2 and RP3/4 distally strongly widened with three rows of cells between these two veins instead of only one (also in Orolestes).
    
  • Other characters: larval prehensile mask with a triangularly projecting anterior margin, a less distinct median cleft, and with numerous dorsal setae on the prementum and lateral lobes (convergent to Coenagrionodea and Cavilabiata); side lobe of larval mask with 4 processes, of which 3 are sharp hooks and one is developed as a serrated protuberance (Archilestes and Orolestes have a plesiomorphic mask with only three sharp processes, while Austrolestes annulosus has four sharp processes, contrary to the other Austrolestes spp. which have a "normal" lestid mask); larval caudal gill lamellae secondarily elongated; larval caudal gills with the tracheal branches running at right angle to the main stem; oviposition in non-aquatic plants (convergent to Palaemnematinae).
• Taxonomy:
  
  • Lestidae; JACOBSON & BIANCHI, 1905 (stat. nov.) (nom. transl. ex Lestinae CALVERT, 1901)
    
  • Lestidae; TILLYARD, 1917

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Sympecmatinae FRASER, 1951

(Type genus: Sympecma SELYS, 1840, nec BURMEISTER, 1839; = Sympycna CHARPENTIER, 1840, jun. obj. syn.; = Sympegma ST. QUENTIN, 1970, incorr. subseq. spell.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: discoidal cell in forewings narrower and shorter than in hindwings (ASKEW, 1988).
    
  • Other characters: resting position with wings closely apposed over the dorsum of the body; secondarily non-metallic body coloration.
• Taxonomy:
  
  • Sympecmatinae FRASER, 1951 (subfam. nov.)
    
  • Sympecmatinae sensu FRASER, 1957
    
  • Sympycnatinae BELYSHEV & HARITONOV, 1975 (jun. obj. syn.)

Comment: the genus Ceylonolestes is certainly not a synonym of Indolestes since it shares with Austrolestes the derived Coenagrion-like imaginal body coloration which is unique within Lestomorpha. Indolestes and Sympecma could be sistergroups, too, as is indicated by the dull brown body coloration of adults.

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Lestinae CALVERT, 1901

(Type genus: Lestes [LEACH] [1815]; = Puella BRULLÉ, 1832, jun. subj. syn.; = Anapates CHARPENTIER, 1840, jun. subj. syn.; = Anapetes PINHEY, 1980, incorr. subseq. spell.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: arculus mainly formed by the basal discoidal crossvein, so that the discoidal cell is nearly touching the RA, convergent to Polythoridae (also in Archilestes, Orolestes, and Chalcolestes viridis, but not in "Lestes" macrostigma and only indistinct in Platylestes).
    
  • Other characters: prementum of larval prehensile mask strongly elongated (also in Archilestes and Orolestes, but not in Chalcolestes viridis, and only slightly so in "Lestes" macrostigma).
• Taxonomy:
  
  • Lestinae CALVERT, 1901 (subfam. nov.)
    
  • Lestinae; NEEDHAM, 1903
    
  • Lestinae; TILLYARD, 1917 (sens. nov.)
    
  • Lestinae; FRASER, 1957 (sens. nov.)

Comment: the position of Archilestes and Orolestes is somewhat enigmatic, since they are in two characters more plesiomorphic than all other Lestidae, while they agree in two derived characters with Lestinae. This homoplasy suggests that the referring characters have to be regarded as rather weak evidence. Nevertheless "Lestes" macrostigma and Chalcolestes viridis indeed seem to be very basal representatives of Lestinae, while Archilestes and Orolestes are either highly derived Lestinae, or the sistergroup of all other Lestidae. In the latter case it could be necessary to establish a separate subfamily Archilestinae or Orolestinae.

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Coenagrionomorpha BECHLY, 1996

• Included taxa: Hypolestidae, Megapodagrionidae, and Coenagrioniformia.
• Autapomorphies:
  
  • Wing venation: pterostigma shortened; postnodal and postsubnodal crossveins aligned (convergent to Lestiformia ?); lestine oblique vein secondarily absent (convergent to Hemiphlebiidae, Caloptera and many † Parazygoptera); basal closure of discoidal cell in forewings including the development of a dorsal arcular bracket (convergent to Caloptera, Lestida, Epiophlebiidae, † Asiopteridae, † Isophlebiidae, and Anisopteromorpha); strong tendency towards the formation of pseudo-transverse veins in the distal part of the wing, caused by an alignment of the postnodal rows of crossveins between the costal margin and the hind margin.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Coenagrionoidea sensu auct. (except. BECHLY, 1996a)
    
  • Coenagrionomorpha BECHLY, 1996a (taxon nov.)

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Hypolestidae TILLYARD & FRASER, 1938

(Type genus: Hypolestes GUNDLACH, 1888, nec Hypolestes HAGEN, 1868, nomen nudum; = Ortholestes CALVERT, 1891, jun. subj. syn.)

• Included taxa: Heteragrioninae, Philogeniinae and Hypolestinae.
• Autapomorphies:
  
  • Wing venation: amphipterygid type of pterostigmata (basal margin strongly slanting, often with hyperstigmal crossveins between itself and the costal margin).
    
  • Other characters: larvae with saccoid caudal gills and a filamentous apex (convergent to Eucaloptera and Palaemenatinae).
• Taxonomy:
  
  • [partim: Normostigmatina KIRBY, 1890 (taxon nov.)
    
  • Hypolestidae; ALAYO SOTO, 1985 (stat. nov.) (nom. transl. ex Hypolestinae TILLYARD & FRASER, 1938)
    
  • Hypolestidae sensu BECHLY, 1996a (sens. nov.)

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Heteragrioninae RACENIS, 1959

(Type genus: Heteragrion SELYS, 1862.)

• Included taxa: at least including the type genus Heteragrion SELYS, 1862, but the remaining generic composition still uncertain.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Heteragrionini RACENIS, 1959 (trib. nov.)
    
  • Heteragrioninae sensu BECHLY, 1996a (stat. nov.)

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Philogeniinae RACENIS, 1959

(Type genus: Philogenia SELYS, 1862.)

• Included taxa: at least including the type genus Philogenia SELYS, 1862. Maybe also including the genus Paraphlebia HAGEN, 1861.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Philogenini RACENIS, 1959 (trib. nov.)
    
  • Philogeninae sensu BECHLY, 1996a (stat. nov.)
    
  • Philogeniinae sensu BECHLY, 2003e (nom. correct.)

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Hypolestinae TILLYARD & FRASER, 1938

(Type genus: Hypolestes GUNDLACH, 1888, nec Hypolestes HAGEN, 1868, nomen nudum; = Ortholestes CALVERT, 1891, jun. subj. syn.)

• Included taxa: Philosinini, Hypolestini, and Lestoideini.
• Autapomorphies:
  
  • Wing venation: the midfork is at least somewhat recessed basally (convergent to Caloptera and Lestinoidea), therefore the subnodus is located between the bases of RP2 and IR2 that are widely separated (convergent to Lestinoidea).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Hypolestinae TILLYARD & FRASER, 1938 (subfam. nov.)
    
  • Hypolestinae; DAVIES, 1981 (sens. nov.)
    
  • Hypolestinae; NEL & PAICHELER, 1994 (sens. nov.)
    
  • Hypolestinae sensu BECHLY, 1996a (sens. nov.)

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Philosinini KENNEDY, 1925

(Type genus: Philosina RIS, 1917.)

• Included taxa: including the type genus Philosina RIS, 1917, and the sister-genera Lestomima MAY, 1933 and Rhipidolestes RIS, 1912 that have a very elongate discoidal cell. Maybe also including the fossil taxa † Eolestes synthetica COCKERELL, 1940 and † Prohypolestes dauphinensis NEL & PAICHELER, 1994.
• Autapomorphies:
  
  • Wing venation: longitudinal wing veins curved (reversal), thus somewhat shortened (especially MA, MP and CuA).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Philosininae KIRBY, 1925 (subfam. nov.)
    
  • Philosinini sensu BECHLY, 1996a (stat. et sens. nov.)

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Hypolestini TILLYARD & FRASER, 1938

(Type genus: Hypolestes GUNDLACH, 1888, nec Hypolestes HAGEN, 1868, nomen nudum; = Ortholestes CALVERT, 1891, jun. subj. syn.)

• Included taxa: only including the type genus Hypolestes GUNDLACH, 1888.
• Autapomorphies:
  
  • Wing venation: characteristical distal curvature of the veins IR1, RP2 and IR2.
    
  • Other characters: adult males with four lobes on the terminal ligula segment (KENNEDY, 1919); body pruinose blue.
• Taxonomy:
  
  • Hypolestinae sensu DAVIES, 1981 (sens. nov.)
    
  • Hypolestidae sensu ALAYO SOTO, 1985 (stat. nov.)
    
  • Hypolestini sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Hypolestinae TILLYARD & FRASER, 1938)

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Lestoideini MUNZ, 1919

(Type genus: Lestoidea TILLYARD, 1913.)

• Included taxa: only including the type genus Lestoidea TILLYARD, 1913.
• Autapomorphies:
  
  • Wing venation: discoidal cell rectangular (convergent to Platystictidae, Protoneuridae and most Caloptera); MP strongly reduced and CuA completely fused to the hind margin (convergent to † Batkeniidae, Disparocypha, Platystictinae, and Protoneuridae, incl. Isostictinae); two characteristical straight intercalary veins between RP1 and IR1 (FRASER, 1957); MA very zigzagged.
    
  • Other characters: resting position with wings closely apposed over the dorsum of the body.
• Taxonomy:
  
  • [légion Lestoidea TILLYARD, 1913 (not available as family-group taxon according to Art. 11f IRZN)]
    
  • Lestoidinae MUNZ, 1919 (subfam. nov.)
    
  • Lestoideidae; TILLYARD & FRASER, 1938 (stat. nov.)
    
  • Lestoideinae sensu FRASER, 1957 (stat. rest. et sens. nov.)
    
  • Lestoideinae; NOVELO-GUTIERREZ, 1995 (sens. nov.)
    
  • Lestoideidae; VAN TOL, 1995 (sens. nov.)
    
  • Lestoideini sensu BECHLY, 1996a (stat. nov.)

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Megapodagrionidae CALVERT, 1913

(Type genus: Megapodagrion SELYS, 1885.)

• Included taxa: Argiolestinae and Megapodagrioninae.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: the three larval caudal gill lamellae held in a horizontal plane.
• Taxonomy:
  
  • [légion Podagrion SELYS, 18?? (not available as family-group taxon according Art.11f IRZN)]
    
  • [partim: Normostigmatina KIRBY, 1890 (taxon nov.)]
    
  • Megapodagriidae; TILLYARD, 1926 (stat. nov.) (nom. transl. ex Megapodagrioninae CALVERT, 1913) (incorr. subseq. spell.)
    
  • Megapodagrionidae; LIPPIT WILLEY, 1955
    
  • Megapodogrionidae; ROSS & JARZEMBOWSKI in BENTON, 1993 (incorr. subseq. spell.)
    
  • Megapodagrionidae sensu BECHLY, 1996a (sens. nov.)

Comment: this group could well be a paraphyletic in the present composition since the characteristical gills are only known from very few species and might thus represent a synapomorphy for a more subordinate group.

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Argiolestinae FRASER, 1957

(Type genus: Argiolestes SELYS, 1862.)

• Included taxa: including the remaining genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: longitudinal wing veins curved, thus somewhat shortened (especially MA, MP and CuA) and the branches of RP distally diverging (reversal); more than one row of cells between CuA and the hind margin (reversal).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Argiolestinae FRASER, 1957 (subfam. nov.)
    
  • Argiolestini; RACENIS, 1959 (stat. nov.)
    
  • incl. Austroargiolestini RACENIS, 1959 (trib. nov. with the type genus: Austroargiolestes KENNEDY, 1925) (jun. subj. syn.)
    
  • Argiolestinae sensu BECHLY, 1996a (sens. nov.)

Comment: maybe paraphyletic, since some genera have derived similarities with Platystictidae, e.g. Burmargiolestes and especially Protolestes.

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Megapodagrioninae CALVERT, 1913

(Type genus: Megapodagrion SELYS, 1885.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: very long spidery legs.
• Taxonomy:
  
  • {Podagrioninae JACOBSON & BIANCHI, 1905 (according to Art. 39 IRZN objectively invalid family-group name, since based on a homonym type genus)}
    
  • Megapodagrioninae CALVERT, 1913 (subfam. nov.)
    
  • Megapodagrioninae; TILLYARD, 1917
    
  • Megapodagriinae; TILLYARD & FRASER, 1938 (incorr. subseq. spell.)
    
  • Megapodagriinae; FRASER, 1957 (sens. nov.)
    
  • incl. Tatocnemidinae RACENIS, 1959 (subfam. nov.) (jun. subj. syn.)
    
  • Megapodagrioninae sensu BECHLY, 1996a (sens. nov.)

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Coenagrioniformia BECHLY, 1996

• Included taxa: Platystictidae and Coenagrionida.
• Autapomorphies:
  
  • Wing venation: pterostigma rather short with only 2 (rarely 3) crossveins beneath it; pterostigmal brace vein ventrally serrated like RP1; subnodus aligned with the base of IR2 (reversed in several subordinate taxa) that is at least somewhat strengthened and dorsally united with the subnodus by a common sclerotisation (interradial bracket) which is usually serrated; number of hexagonal and pentagonal cells strongly reduced (maybe synapomorphic with some "Megapodagrionidae"); in the total wing space between RP1 and RP2 there are only two rows of cells present that are separated by the IR1 (convergent to a few Megapodagrioninae; reversed in Pseudostigmatidae and the coenagrionid genus Boninagrion); postnodal crossveins aligned with the rows of crossveins below, forming several pseudo-transverse-veins (convergent to Perilestinae and some "Megapodagrioninae"; reversed within Platystictinae); all intercalary veins (except IR1 and IR2) suppressed (convergent to Hemiphlebiidae, Chorismagrionidae, and Perilestidae; reversed in Megaloprepini).
    
  • Other characters: resting position with wings closely apposed over the dorsum of the body; the specialised spines at the movable crossvein-junctions are only present along the ventral sides of RP2, RP3/4 and MP (reduced on the dorsal side, where they are still present in Lestomorpha, Caloptera and many Anisoptera); lateral lobes ("palps") of the larval prehensile mask with only two endhooks.
• Taxonomy:
  
  • Coenagrioniformia BECHLY, 1996a (taxon nov.)

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Platystictidae LAIDLAW, 1924

(Type genus: Platysticta SELYS, 1860.)

• Included taxa: Palaemnematinae, Sinostictinae, and Platystictinae.
• Autapomorphies:
  
  • Wing venation: wings with falcate apex (convergent to some Protoneuridae, e.g. Phylloneura); longitudinal veins RA, RP1, IR1, RP2 and IR2 strongly converging to the apex (convergent to some Protoneuridae, e.g. Phylloneura); costal edge of pterostigma much shorter than the radial edge (KENNEDY, 1919); nodus in a very basal position (about 22 % of wing length; convergent to Perilestidae, Pseudostigmatoidea, and some Protoneuridae, e.g. Phylloneura); discoidal cell rectangular (convergent to Lestoideini, Protoneuridae and most Caloptera); CuP-crossing (= anal crossing sensu Fraser) recessed in a very basal position (mistaken as pcv by FRASER, 1957) (convergent to some Protoneuridae, e.g. Nososticta and Isostictinae); the real subdiscoidal cell is traversed by three crossveins that delimit a pseudo-subdiscoidal-cell (reduced in Protosticta and Drepanosticta carmichaeli); MA very long (in the groundplan also MP), at least reaching up to 90 % of wing length (convergent to Pseudostigmatoidea); CuA completely fused with the hind margin, thus only retained as subdiscoidal vein (convergent to † Batkeniidae, Disparocypha, Lestoideini, and Protoneuridae, incl. Isostictinae).
    
  • Other characters: microsculpture of the pterostigmata consisting of tiny pyramid-like structures (convergent to Chlorocyphoidea and Epallagidae).
• Taxonomy:
  
  • [partim: Normostigmatina KIRBY, 1890 (taxon nov.)]
    
  • [légion Platysticta LAIDLAW, 1917 (not available as family-group taxon according to Art. 11f IRZN)]
    
  • Platystictidae TILLYARD & FRASER, 1938 (stat. nov.) (nom. transl. ex Platystictinae LAIDLAW, 1924)

Comment: KENNEDY (1919) mentions some similarities in the wing venation and the male genitalia between Platystictidae and basal Pseudostigmatidae which he interpreted as evidence of a common ancestry, but which are of uncertain polarity and uncertain homology. The striking similarities with certain Protoneuridae are without doubt convergences as is clearly demonstrated by the plesiomorphic type of the larval mask.

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Palaemneminae TILLYARD & FRASER, 1938

(Type genus: Palaemnema SELYS, 1860.)

• Included taxa: only including the type genus Palaemnema SELYS, 1860.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larvae with saccoid caudal gills (convergent to most basal Eucaloptera and Hypolestidae; but polarity within Platystictidae unclear); larval prehensile mask with a medially widened prementum which as two parallel medio-dorso-longitudinal patches of minute striations (DE MARMELS, 1990); oviposition in non-aquatic plants (convergent to Lestidae, but maybe rather an autapomorphy of Platystictidae).
• Taxonomy:
  
  • Palaemnematinae TILLYARD & FRASER, 1938 (subfam. nov.) (nom. imperf.)
    
  • [Palaemnematinae FRASER, 1957 (jun. obj. syn. and homonym)]
    
  • Palaemneminae sensu DAVIES & TOBIN, 1984 (nom. correct.)

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Sinostictinae WILSON, 1997

(Type genus: Sinosticta WILSON, 1997.)

• Included taxa: only including the single species Sinosticta ogatai WILSON, 1997.
• Autapomorphies:
  
  • Wing venation: presence of one to three supplementary cubito-anal crossvein between CuP-crossing (= pcv sensu Fraser) and the pseudo-subdiscoidal cell (basally limited by "anal crossing" sensu Fraser) (WILSON, 1997).
    
  • Other characters: not yet known (probably the genital organs and anal appendages have autapomorphic chraracters).
• Taxonomy:
  
  • Sinostictinae WILSON, 1997 (subfam. nov.)

Comment: this new subfamily was erected by WILSON (1997) for Sinosticta ogatai from Hong Kong that was previously classified in the genus Drepanosticta. This species is the only "Old World" Platystictidae that retained a long vein MP just like the Palaemnematinae in the "New World".

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Platystictinae LAIDLAW, 1924

(Type genus: Platysticta SELYS, 1860.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: MP much shortened; origin of RP2 shifted basally nearer to the nodus, correlated with an elongation of IR1 (FRASER, 1957; only distinct in the genus Platysticta); many pentagonal cells because of more or less zigzagged convex longitudinal veins IR1, IR2 and MA (FRASER, 1957; only distinct in the genus Platysticta).
    
  • Other characters: larvae with triquetral caudal gills (convergent to Rimanellinae and Calopterygida; but polarity within Platystictidae unclear).
• Taxonomy:
  
  • Platystictinae LAIDLAW, 1924 (subfam. nov.)
    
  • Platystictinae sensu TILLYARD & FRASER, 1938 (sens. nov.)

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Coenagrionida BECHLY, 1996

• Included taxa: Pseudostigmatoidea and Coenagrionodea.
• Autapomorphies:
  
  • Wing venation: RP1 distinctly kinked at the insertion of the pterostigmal brace vein (convergent to Hemiphlebiidae; reduced in Pseudostigmatidae).
    
  • Other characters: larval prehensile mask with the anterior margin of the prementum triangularly projecting (convergent to Calopterygida), and with strongly reduced or completely suppressed median cleft; larval prehensile mask with two apical endhooks, and a row of dorsal setae on the lateral lobes ("palps") (convergent to Lestidae and Cavilabiata).
• Taxonomy:
  
  • Coenagrionida BECHLY, 1996a (taxon nov.)

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Pseudostigmatoidea KIRBY, 1890

(Type genus: Pseudostigma SELYS, 1860.)

• Included taxa: Coryphagrionidae and Pseudostigmatidae.
• Autapomorphies:
  
  • Wing venation: wings very long and slender (thus all longitudinal veins strongly elongated and very straight in the groundplan); nodus in a extremely basal position (16-23 % of wing length; convergent to Perilestidae and Platystictidae), correlated with a basal recession of the midfork (at about 24 % of wing length) and a very high number of postnodal crossveins; the basal wing space between RP3/4 and IR2 is very narrowed for a considerable distance; the discoidal cell is at least somewhat elongated; the longitudinal veins converge near the wing apex; RA and RP1 are sigmoidally curved at the apex; MA, MP and CuA are very long, ending near the wing apex; more than one row of cells in the poststigmal area between the costal margin and RA; [M & Cu] or MP rather straight at the arculus, being only smoothly curved or indistinctly kinked.
    
  • Other characters: giant size; very elongated and slender abdomen; exophytic oviposition in phytotelmata (oviposition by "bombing").
• Taxonomy:
  
  • Pseudostigmatoidea sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Pseudostigmatina KIRBY, 1890)

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Coryphagrionidae PINHEY, 1962

(Type genus: Coryphagrion MORTON, 1924.)

• Included taxa: on including the species Coryphagrion grandis MORTON, 1924.
• Autapomorphies:
  
  • Wing venation: the single row of cells between CuA and the hind margin of the wing contains numerous cells with a more or less trigonal or trapezoidal shape; kink of RP1 at the pterostigmal brace vein hypertrophied.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Coryphagriinae PINHEY, 1962 (subfam. nov.) (nom. imperf.)
    
  • Coryphagrioninae ST. QUENTIN & BEIER, 1968 (nom. correct.)
    
  • Coryphagrionidae sensu BECHLY, 1995 (stat. nov.)

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Pseudostigmatidae KIRBY, 1890

(Type genus: Pseudostigma SELYS, 1860.)

• Included taxa: Mecistogastrinae and Pseudostigmatinae.
• Autapomorphies:
  
  • Wing venation: pterostigmata reduced (incl. the brace) and replaced by characteristical pseudo-pterostigmata; more than two rows of cells between RP1 and RP2 (reversal, correlated with the large size); RA, RP1 and IR1 apically strongly curved towards the hind margin and converging to the same point at the hind margin (reversed in Anomisma).
    
  • Other characters: unique shape of larval caudal gill lamellae which are stalked at the base (convergent to some Calicnemidinae like Copera) and spatulate distally; larvae with branched spines on the distal parts of the tibiae and the sides of the tarsi (PINHEY in PARKER, 1982); adult males with reduced paraprocts; terminal segment of male ligula with an apical filamentous projection (SCHMIDT, 1915; KENNEDY, 1919); feeding on prey in spider webs.
• Taxonomy:
  
  • [légion Pseudostigma SELYS (not available as family-group taxon according Art.11f IRZN)]
    
  • Pseudostigmatina KIRBY, 1890 (described as new "division")
    
  • Pseudostigmatinae sensu MUTTKOWSKI, 1910 (stat. nov. ?)
    
  • Pseudostigmatinae sensu TILLYARD, 1917
    
  • Pseudostigmatinae sensu MUNZ, 1919
    
  • Pseudostigmatidae sensu TILLYARD & FRASER, 1938 (stat. nov.)
    
  • [Anormostigmatini NEEDHAM, 1903 (taxon nov.; according to Art. 11 IRZN not available as family-group name, since not derived from a generic name)]

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Mecistogastrinae BECHLY, 1996

(Type genus: Mecistogaster RAMBUR, 1842.)

• Included taxa: only including the type genus Mecistogaster RAMBUR, 1842.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larval gills more strongly pedunculate.
• Taxonomy:
  
  • Mecistogastrinae BECHLY, 1996a (subfam. nov.)

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Pseudostigmatinae KIRBY, 1890

(Type genus: Pseudostigma SELYS, 1860.)

• Included taxa: Pseudostigmatini and Megaloprepini.
• Autapomorphies:
  
  • Wing venation: field between CuA and the hind margin of the wing secondarily with more than one row of cells.
    
  • Other characters: adult males with more or less reduced cerci; derived type of male ligula (KENNEDY, 1919); larval caudal gills with an apical pale area or white spot (RAMIREZ, 1997).
• Taxonomy:
  
  • Pseudostigmatinae; MUTTKOWSKI, 1910 (stat. nov. ?) (nom. transl. ex Pseudostigmatina KIRBY, 1890)
    
  • Pseudostigmatinae; TILLYARD, 1917
    
  • Pseudostigmatinae; MUNZ, 1919
    
  • Pseudostigmatinae sensu BECHLY, 1996a (sens. nov.)

Comment: the larvae of Mecistogaster are lacking the tarsal pulvilli that are present in the larvae of Pseudostigma and Megaloprepus, but the polarity of this character is not yet known.

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Pseudostigmatini KIRBY, 1890

(Type genus: Pseudostigma SELYS, 1860.)

• Included taxa: only including the type genus Pseudostigma SELYS, 1860.
• Autapomorphies:
  
  • Wing venation: accessory anterior IR1 more distinct.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Pseudostigmatini sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Pseudostigmatina KIRBY, 1890)

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Megaloprepini BECHLY, 1996

(Type genus: Megaloprepus RAMBUR, 1842.)

• Included taxa: Anomismina and Megaloprepina.
• Autapomorphies:
  
  • Wing venation: wings less narrow and slender; triadic branching of veins CuA and MA; field between RP1/2 and RP3/4 widened by a distal shifting of the furcation into RP1 and RP2, and with at least a very short part with more than one row of cells; RP3/4 apically forked.
    
  • Other characters: further derived type of male ligula, with the base of the apical segment crooked or offset and an erect internal fold (KENNEDY, 1919; not yet analysed in Anomisma).
• Taxonomy:
  
  • Megaloprepini BECHLY, 1996a (trib. nov.)

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Anomismina BECHLY, 1996

(Type genus: Anomisma MCLACHLAN, 1877.)

• Included taxa: only including the type genus Anomisma MCLACHLAN, 1877.
• Autapomorphies:
  
  • Wing venation: a single accessory antenodal present between costal margin and ScP distal of ax2; no pterostigma present at all; discoidal cell extremely elongated; discoidal and subdiscoidal cell traversed by 2-3 crossveins; two accessory cubito-anal crossveins present; accessory anterior IR1 obliterated.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Anomismina BECHLY, 1996a (subtrib. nov.)

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Megaloprepina BECHLY, 1996

(Type genus: Megaloprepus RAMBUR, 1842.)

• Included taxa: including the two genera Megaloprepus RAMBUR, 1842 and Microstigma RAMBUR, 1842.
• Autapomorphies:
  
  • Wing venation: large part of the postsubnodal space divided into two rows of cells; field between RP1/2 and RP3/4 with a longer area with more than one row of cells.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Megaloprepina BECHLY, 1996a (subtrib. nov.)

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Coenagrionodea BECHLY, 1996

• Included taxa: Coenagrionidae and Platycnemidoidea.
• Autapomorphies:
  
  • Wing venation: terminal kink of CP at nodus and nodal membrane sclerotisation completely suppressed; basal and distal margin of pterostigma thickened and fused with the strongly thickened pterostigmal part of RA (convergent to † Tarsophlebiidae and many Gomphides) to a U-shaped structure; all intercalary veins (except IR1 and IR2) suppressed (convergent to Hemiphlebiidae, Chorismagrionidae and Perilestidae).
    
  • Other characters: macrotrichae on wing veins completely suppressed, except along the margin and the ventral side of ScP; larval prehensile mask with dorsal setae on the prementum (except in Argiinae; convergent to Lestidae and Cavilabiata); proventricular folds of larval gizzard divided into an membranous oral part with numerous smaller denticles and an anal cuticular plate with fewer and larger denticles (BULLENS, 1966; not yet analysed in Protoneuridae, and also not yet known from other Coenagrionomorpha, thus maybe an autapomorphy for a more inclusive monophylum); unique oviposition behaviour with the male remaining attached to the female neck in a peculiar upright "watch-tower-position" during oviposition.
• Taxonomy:
  
  • [partim: Normostigmatina KIRBY, 1890 (taxon nov.)]
    
  • Coenagrionodea BECHLY, 1996a (taxon nov.)

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Coenagrionidae KIRBY, 1890

(Type genus: Coenagrion KIRBY, 1890 nom. subst. pro Agrion [LEACH] [1815], nec Agrion FABRICIUS, 1775.)

• Included taxa: Argiinae, Coenagrioninae, Agriocnemidinae, Ischnurinae, Leptobasine, and Pseudagrioninae, which are all still of doubtful phylogenetic status and position.
• Autapomorphies:
  
  • Wing venation: the distal discoidal vein MAb is very oblique, so that the anterior side of the discoidal cell is much shorter than the posterior side (convergent to Lestinae).
    
  • Other characters: anal appendages of adult males reduced, especially cerci that are never forcipate (convergent to Parahemiphlebia and some Protoneuridae; reversed in a few species like Coenagrion armatum).
• Taxonomy:
  
  • [légion Agrion SELYS, 18?? (not available as family-group taxon according Art.11f IRZN)]
    
  • {Agrionidae STEPHENS, 1836 (according to Art. 39 IRZN objectively invalid new family-group name, since based on a homonym type genus)}
    
  • {Agrionines; SELYS in SELYS & HAGEN, 1850 (objectively invalid name)}
    
  • Coenagrionidae; KARSCH, 1894 (stat. nov.) (nom. transl. ex Coenagrioninae KIRBY, 1890)
    
  • {Agrionii; ACLOQUE, 1897 (incorr. subseq. spell.) (objectively invalid name)}
    
  • {Agrioninae; LUCAS, 1900 ? (objectively invalid name)}
    
  • {Agrioninae; NEEDHAM, 1903 (objectively invalid name)}
    
  • {Agrionidae; NEEDHAM, 1903 (objectively invalid name)}
    
  • {Agrioninae; JACOBSON & BIANCHI, 1905 (objectively invalid name)}
    
  • {Agrioninae; RIS, 1909 (objectively invalid name)}
    
  • Coenagrioninae; MUTTKOWSKI, 1910
    
  • {Agrionidae; TILLYARD, 1917 (objectively invalid name)}
    
  • {Agrioninae; TILLYARD, 1917 (objectively invalid name)}
    
  • Coenagrionidae; MUNZ, 1919
    
  • Coenagrioidea; TILLYARD, 1926 (stat. nov.) (nom. imperf.)
    
  • Coenagriidae; TILLYARD, 1926 (nom. imperf.)
    
  • Coenagrioidea; TILLYARD & FRASER, 1938 (nom. imperf.)
    
  • {Puellidae AKRAMOWSKI, 1948 (probably an invalid taxon since the family-group name was erroneously based on the type genus Puella BRULLÉ, 1832, which is a jun. subj. syn. of Lestes [LEACH] [1815] according to the subsequent designation of a type species by COWLEY (1934))}
    
  • Coenagrionoidea; MACNEILL, 1960
    
  • Coenagrionidea; HENNIG, 1969 (nom. imperf.)
    
  • Coenagrionoidea; PRITYKINA, 1980

Comment: although probably monophyletic, there is some weak conflicting evidence from the morphology of the larval mask, suggesting that Argiinae might be the sistergroup of all remaining Coenagrionodea, and genera like Acanthagrion, Aeolagrion and Ceriagrion could be closer related to Platycnemidoidea because of the pattern of premental setae. If this taxon should indeed be monophyletic it could be retransfered to superfamily rank (Coenagrionoidea) to allow a future phylogenetic systematisation of the current subfamilies.

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Argiinae TILLYARD, 1917

(Type genus: Argia RAMBUR, 1842.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: wings only very shortly petiolated.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Argiini TILLYARD, 1917 (trib. nov.)
    
  • Argiinae sensu TILLYARD & FRASER, 1938 (stat. nov.)

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Coenagrioninae KIRBY, 1890

(Type genus: Coenagrion KIRBY, 1890 nom. subst. pro Agrion [LEACH] [1815], nec Agrion FABRICIUS, 1775.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Coenagrioninae KIRBY, 1890
    
  • Coenagrioninae; MUTTKOWSKI, 1910
    
  • {Agrionini sensu TILLYARD, 1917 (stat. nov.) (objectively invalid name)}
    
  • Coenagriinae sensu TILLYARD & FRASER, 1938 (nom. imperf.)
    
  • Coenagriinae sensu FRASER, 1957
    

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Agriocnemidinae FRASER, 1957

(Type genus: Agriocnemis SELYS, 1869 or 1877, a case for ICZN according to BRIDGES, 1994.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: arculus shifted distal of ax2 (reversal, convergent to Ceratura).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Agriocneminae FRASER, 1957 (subfam. nov.) (nom. imperf.)
    
  • Agriocnemidinae sensu DAVIES, 1981 (nom. correct.)

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Ischnurinae FRASER, 1957

(Type genus: Ischnura CHARPENTIER, 1840.)

• Included taxa: including the genera listed in BRIDGES (1994), and also including the genus Ceratura SELYS, 1876 (stat. rest.).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: adult females with a prominent ventral spine on the apical margin of the 8th abdominal segment (vulvar spine).
• Taxonomy:
  
  • Ischnurinae FRASER, 1957 (subfam. nov.)

Comment: to remove the homonymy with the scorpion family Ischnuridae SIMON, 1879 a conservation as the correct spelling Ischnurainae FRASER, 1957 was proposed as CASE 3120 to the ICZN by FET & BECHLY, 2000.

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Leptobasinae DAVIES & TOBIN, 1984

(Type genus: Leptobasis SELYS, 1877.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: tarsal claw hooks obsolete or absent (convergent to Perilestidae and Onychothemistina; maybe synapomorphic with "Teinobasini").
• Taxonomy:
  
  • [partim: Amphicneminae sensu FRASER, 1957]
    
  • Leptobasinae DAVIES & TOBIN, 1984 (subfam. nov.)

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Pseudagrioninae TILLYARD, 1917

(Type genus: Pseudagrion SELYS, 1876.)

• Included taxa: including the genera listed in BRIDGES (1994), and thus also including the former tribus Teinobasini.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Pseudagrionini TILLYARD, 1917 (trib. nov.)
    
  • incl. Teinobasini TILLYARD, 1917 (trib. nov. with type genus Teinobasis KIRBY, 1890) (jun. subj. syn.)
    
  • Pseudagriinae; TILLYARD, 1926 (stat. nov.) (incorr. subseq. spell.)
    
  • partim: Amphicneminae FRASER, 1957 (subfam. nov. with type genus Amphicnemis SELYS, 1863) (jun. subj. syn.)
    
  • Pseudagrioninae; St.Quentin & BEIER, 1968

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Platycnemidoidea JACOBSON & BIANCHI, 1905

(Type genus: Platycnemis BURMEISTER, 1839, nec CHARPENTIER, 1840).)

• Included taxa: "Platycnemididae" and Protoneuridae.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larval prehensile mask with a transverse row of 4 setae (only 2 in some Protoneurinae) on the dorsal side of the prementum.
• Taxonomy:
  
  • Platycnemidoidea sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Platycnemididinae JACOBSON & BIANCHI, 1905)

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"Platycnemididae" JACOBSON & BIANCHI, 1905

(Type genus: Platycnemis BURMEISTER, 1839, nec CHARPENTIER, 1840.)

• Included taxa: "Calicnemidinae" and Platycnemidinae.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: adult males without hairs or spines on the ligula shaft (KENNEDY, 1919; convergent to Chlorocyphoidea and Lestida).
• Taxonomy:
  
  • [légion Platycnemis SELYS, 1863 (not available as family-group taxon according Art.11f IRZN)]
    
  • Platycnemididinae sensu JACOBSON & BIANCHI, 1905 (subfam. nov.) (nom.imperf.)
    
  • Platycneminae sensu TILLYARD, 1917
    
  • Platycnemini sensu HANDLIRSCH, 1926-1930 (stat. nov.)
    
  • Platycnemininae sensu FRASER, 1929
    
  • Platycnemididae sensu TILLYARD & FRASER, 1938 (stat. nov. et nom. correct.)
    
  • Platycnemididae sensu FRASER, 1957

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"Calicnemidinae" FRASER, 1957

(Type genus: Calicnemia STRAND, 1928; = Calicnemis SELYS, 1863, jun. obj. syn.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Calicneminae FRASER, 1957 (subfam. nov.) (nom. imperf.)
    
  • Calicnemidinae sensu DAVIES, 1981 (nom. correct.)

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Platycnemidinae JACOBSON & BIANCHI, 1905

(Type genus: Platycnemis BURMEISTER, 1839, nec CHARPENTIER, 1840.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: discoidal cell more or less rectangular (maybe rather a synapomorphy with Protoneuridae).
    
  • Other characters: male tibiae dilated.
• Taxonomy:
  
  • Platycnemididinae JACOBSON & BIANCHI, 1905 (subfam. nov.) (nom. imperf.)
    
  • Platycneminae; TILLYARD, 1917 (nom. imperf.)
    
  • Platycnemininae; FRASER, 1929 (nom. imperf.)
    
  • Platycnemininae sensu FRASER, 1957 (sens. nov.)
    
  • Platycnemidinae sensu DAVIES, 1981 (nom. correct.)

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Protoneuridae JACOBSON & BIANCHI, 1905

(Type genus: Protoneura SELYS, 1857.)

• Included taxa: "Protoneurinae" and Isostictinae.
• Autapomorphies:
  
  • Wing venation: discoidal cell rectangular (convergent to Lestoideini, Platystictidae and most Caloptera; maybe synapomorphic with Platycnemidinae); discoidal bracket (dorsal sclerotisation on MAb and the subdiscoidal vein) strongly reduced (convergent to Perilestidae and Anisoptera); CuA completely fused with the hind margin, thus only retained as subdiscoidal vein (convergent to † Batkeniidae, Disparocypha, Lestoideini, and Platystictidae).
    
  • Other characters: larval caudal gills are bipartite; adult males with four lobes at the terminal segment of the ligula (KENNEDY, 1919; maybe a synapomorphy with Platycnemidinae; reversed in most Isostictini and some Protoneurinae, but preserved in the former Caconeurinae and Disparoneurinae).
• Taxonomy:
  
  • Protoneurinae sensu JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • Protoneurinae sensu TILLYARD, 1917
    
  • Protoneuridae sensu TILLYARD, 1926 (stat. nov.)
    
  • Protoneurini sensu HANDLIRSCH, 1926-1930 (stat. nov.)
    
  • Protonevridae sensu RACENIS, 1959 (incorr. subseq. spell.)
    
  • Protoneuridae; LIEFTINCK, 1975 (sens. nov.)
    
  • Protoneuridae; WATSON, 1992
    
  • Protoneuridae sensu BECHLY, 1996a

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"Protoneurinae" JACOBSON & BIANCHI, 1905

(Type genus: Protoneura SELYS, 1857.)

• Included taxa: including the genera listed in BRIDGES (1994) under Protoneuridae (thus including the former separate subfamilies Caconeurinae and Disparoneurinae according to WATSON, 1992).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Protoneurinae JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • Protoneurinae; FRASER, 1957 (sens. nov.)
    
  • incl. "Caconeurinae" FRASER, 1957 (subfam. nov. with type genus Caconeura KIRBY, 1890) (jun. subj. syn.)
    
  • incl. "Disparoneurinae" FRASER, 1957 (subfam. nov. with type genus Disparoneura SELYS, 1860) (jun. subj. syn.)
    
  • Protoneurinae; WATSON & THEISCHINGER, 1984 (sens. nov., synonymised with Disparoneurinae)
    
  • Protoneurinae; WATSON, 1992 (sens. nov., synonymised with Disparoneurinae and Caconeurinae)
    
  • Protoneurinae sensu BECHLY, 1996a (sens. nov.)

Comment: the genera Chlorocnemis and Proneura have uniquely retained a short free portion of CuA and therefore could represent the sistergroup of all remaining Protoneuridae. A partly free AA is only retained in all Caconeurinae, many Disparoneurinae and a few Protoneurinae, while in the other genera and in all Isostictinae there are no vestiges of AA and CuA at all.

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Isostictinae FRASER, 1955

(Type genus: Isosticta SELYS, 1885.)

• Included taxa: † Eoprotoneurini and Isostictini.
• Autapomorphies:
  
  • Wing venation: arculus secondarily shifted distal of ax2 (maybe a synapomorphy with certain Protoneurinae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Isostictinae FRASER, 1955 (subfam. nov.)
    
  • Isostictinae sensu FRASER, 1957
    
  • Isostictinae sensu WATSON, 1969
    
  • Isostictidae sensu LIEFTINCK, 1975 (stat. nov.)
    
  • Isostictidae sensu WATSON, 1992
    
  • Isostictinae sensu BECHLY, 1996a (stat. rest. et sens. nov.)

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† Eoprotoneurini CARLE & WIGHTON, 1990

(Type genus: † Eoprotoneura CARLE & WIGHTON, 1990.)

• Included taxa: only including the species † Eoprotoneura hyperstigma CARLE & WIGHTON, 1990.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Eoprotoneurinae CARLE & WIGHTON, 1990 (subfam. nov.)
    
  • Eoprotoneurini sensu BECHLY, 1996a (stat. nov.)

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Isostictini FRASER, 1955

(Type genus: Isosticta SELYS, 1885.)

• Included taxa: including the genera listed in BRIDGES (1994) under Isostictidae.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larvae with unique type of caudal gills that are nodate with a strong constriction at nodus, and basal of the nodus somewhat saccoid (they are not basally saccoid and distally filiform as maintained by FRASER, 1957).
• Taxonomy:
  
  • Isostictinae sensu FRASER, 1955 (subfam. nov.)
    
  • Isostictinae sensu FRASER, 1957
    
  • Isostictidae sensu LIEFTINCK, 1975 (stat. nov.)
    
  • Isostictidae sensu WATSON
    
  • Isostictinae sensu BECHLY, 1996a (stat. rest. et sens. nov.)
    
  • Isostictini sensu BECHLY, 1996a (stat. nov.)