Phylogenetic Systematics of Odonata

Neolamellida BECHLY, 1996

• Included taxa: Gomphomacromiidae and Valvulida.
• Autapomorphies:
  
  • Wing venation: nearly all antenodal crossveins are aligned in both pairs of wings (BECHLY, 1994), and at least all the hindwing antenodal crossveins are more or less enforced and bracket-like (FRASER, 1957; CARLE, 1995); the two primary antenodal brackets ax1 and ax2 are at least somewhat less distinct (FRASER, 1957).
    
  • Other characters: legs with secondarily simplified ortho-tibiae (LOHMANN, 1996); males with ordinary spines instead of peg-like spines on the outer margin of the meso- and metatibiae, and femora of male hindlegs not distinctly thickened as in Chlorogomphoidea and Synthemistidae (LOHMANN, 1996; tumidotrichae reduced, convergent to Neopetaliidae); male abdomen expanded and triquetal at least on the 7. and 8. segment, but not yet supplied with lateral carinae (CARLE & LOUTON, 1994; CARLE, 1995); larval abdomen in the groundplan stout, less than twice as long as wide (CARLE, 1995); larval hind femora in the groundplan more than 1,5 times length of front femora (CARLE, 1995; this character is homoplastic and of doubtful polarity); larval metasternum with anterior and posterior transverse sulci fused medially (CARLE, 1995); posterior margin of larval abdominal segments secondarily without long hair setae (CARLE, 1995; convergent to some Synthemistidae); larval wing pads non-divergent; neo-lamellate sub-type of the larval rectal gills (number of gills increased, placed close together, and with much smaller basal pads) (TILLYARD, 1917).
• Taxonomy:
  
  • Neolamellida BECHLY, 1996a (taxon nov.)
    
  • Firmonervata LOHMANN, 1996 (taxon nov.)

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Gomphomacromiidae TILLYARD & FRASER, 1940

(Type genus: Gomphomacromia BRAUER, 1864.)

• Included taxa: probably only including the two genera Gomphomacromia BRAUER, 1864 and Archaeophya FRASER, 1959.
• Autapomorphies:
  
  • Wing venation: there is no secondary antenodal present between the two primary antenodal brackets ax1 and ax2, therefore the arculus is situated between the first two antenodal crossveins (convergent to Pseudocorduliidae, Austrocorduliidae and Liberaponsida); hindwing discoidal triangle recessed, separated by only about 1/2 length of arculus from the latter (CARLE & LOUTON, 1994; convergent to Pseudocorduliidae and Laterocarinida), correlated with a more basal position of the pseudo-anal crossvein PsA basal of the arculus (CARLE, 1995; conta LOHMANN, 1996); elongate anal loop (CARLE, 1995; convergent to Pseudocorduliidae and Laterocarinida); no cubito-anal crossveins retained, except the CuP-crossing (= anal crossing sensu Fraser) and the pseudo-anal crossvein PsA (convergent to Pseudocorduliidae, Austrocorduliidae and Liberaponsida); hypertriangle without crossveins (convergent to Pseudocorduliidae, Austrocorduliidae and Liberaponsida).
    
  • Other characters: lateral prothoracic lobes shelf-like (CARLE, 1995); lateral abdominal spines absent in larvae (THEISCHINGER & WATSON, 1984); projecting tip of the hyperboloid micropylar cone (MAY, 1995a; LOHMANN, 1996; might rather be a symplesiomorphy); male epiproct trifurcate in the groundplan (LOHMANN, 1995); males without tibial keel on the mesotibiae (LOHMANN, 1996).
• Taxonomy:
  
  • Gomphomacromiinae TILLYARD & FRASER, 1940 (subfam. nov.)
    
  • Gomphomacromiidae; CARLE & LOUTON, 1994 (stat. nov.)
    
  • Gomphomacromiidae sensu MAY, 1995b (sens. nov.)
    
  • [Gomphomacromiida LOHMANN, 1996 (taxon nov.)]

Comment: this taxon might be paraphyletic in its present composition. According to MAY (1995b) and LOHMANN (1996) Pseudocordulia had to be excluded from Gomphomacromiidae.

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Valvulida LOHMANN, 1996

• Included taxa: Pseudocorduliidae and Trichodopalpida.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: ovipositor reduced to a pair of valvula vulvae (LOHMANN, 1996; convergent to Chlorogomphoidea and Gomphides); larval labial palps with medial edge containing small lobes (LOHMANN, 1996); larval prementum secondarily without antero-median protrusion (LOHMANN, 1996); anal pyramid of larvae shortened and the epiproctal process fused with the epiproct (LOHMANN, 1996); male abdominal tergum 1 without ventro-lateral hamule-like projections (CARLE, 1995; LOHMANN, 1996); male epiproct unifurcate (LOHMANN, 1996; contra LOHMANN, 1995).
• Taxonomy:
  
  • Valvulida LOHMANN, 1996 (taxon nov.)

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Pseudocorduliidae LOHMANN, 1996

(Type genus: Pseudocordulia TILLYARD, 1909.)

• Included taxa: only including the type genus Pseudocordulia TILLYARD, 1909.
• Autapomorphies:
  
  • Wing venation: there is no secondary antenodal present between the two primary antenodal brackets ax1 and ax2 (convergent to Gomphomacromiidae, Austrocorduliidae and Liberaponsida), and the arculus is aligned with ax2 (FRASER, 1960); hindwing discoidal triangle recessed, separated by only about 1/2 length of arculus from the latter (CARLE & LOUTON, 1994; convergent to Gomphomacromiidae and Laterocarinida); elongate anal loop (CARLE, 1995; convergent to Gomphomacromiidae and Laterocarinida); no cubito-anal crossveins retained (convergent to Gomphomacromiidae, Austrocorduliidae and Liberaponsida), except the CuP-crossing (= anal crossing sensu Fraser) and the pseudo-anal crossvein PsA; hypertriangle without crossveins (convergent to Gomphomacromiidae, Austrocorduliidae and Liberaponsida).
    
  • Other characters: terrestrial larvae (WATSON, 1982); male hamuli posteriores with posterior lobe erect-chisel-like (CARLE, 1995); female abdominal sternite 9 with medial carina and sternite 10 with lateral carinae (CARLE, 1995).
• Taxonomy:
  
  • Pseudocorduliidae LOHMANN, 1996 (fam. nov.)

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Trichodopalpida BECHLY, 1996

• Included taxa: Macromiidae and Laterocarinida.
• Autapomorphies:
  
  • Wing venation: presence of at least an indistinct Rspl that is more or less parallel to IR2 in the groundplan (convergent to † Aeschnidiidae, Aeshnoptera and † Hemeroscopus baissicus; the alleged Rspl within Lindeniidae and in Cordulegastrida is indeed a secondary branch of IR2).
    
  • Other characters: males with well-developed genital lobes at the ventral margin of the second abdominal segment (SCHMIDT, 1916), correlated opposable hamuli posteriores, and basal asymmetry of penile flagellae (SCHMIDT, 1916; CARLE & LOUTON, 1994; CARLE, 1995, MAY, 1995b); females with further reduced ovipositor (the progonapophyses are shorter than sternum 9 and reduced to a valvula vulvae, the mesogonapophyses are strongly or completely reduced, and the metagonapophyses are peg-like or obsolete) (CARLE & LOUTON, 1994; CARLE, 1995; LOHMANN, 1995); frontal plate of larval head reduced (LOHMANN, 1995); larval prehensile mask with small tufts of raptorial setae ("palpotrichae") on the inner margin of the lateral lobes ("palps") (THEISCHINGER & WATSON, 1984; CARLE, 1995; LOHMANN, 1995); ventral lobes of larval gizzard less than half as wide as the dorsal lobes, with teeth separated by less than the width of the dorsal lobes (CARLE, 1995); larval gizzard with medial chute-like sclerotisation between the dorsal lobes (CARLE, 1995); larval epiproct fused with process and situated on the same level (LOHMANN, 1995; convergent to Gomphides-Desmoproctida); eggs with a concave micropylar cone and subapical micropyle openings opposite to each other like an eye of a needle (MAY, 1995a; LOHMANN, 1995); larvae with dorsal abdominal spines (CARLE, 1995; a somewhat doubtful character since such spines are also present within Neoaeshnida and Gomphides); second segment of male vesicula spermalis less curved in the groundplan (SCHMIDT, 1916).
• Taxonomy:
  
  • [MCL-Complex sensu CARLE, 1995 (informal name)]
    
  • Trichodopalpida BECHLY, 1996a (taxon nov.)
    
  • Trichopalpida LOHMANN, 1996 (taxon nov., based on a suggestion of Bechly)

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Macromiidae NEEDHAM, 1903

(Type genus: Macromia RAMBUR, 1842.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: RP3/4) and MA distinctly undulate in both pairs of wings (NEEDHAM & WESTFALL, 1955); the two primary antenodal brackets ax1 and ax2 are indistinct or even indistinguishable from the secondaries (FRASER, 1937, 1957); sectors of arculus stalked (convergent to Synthemistidae, Idionychidae, Neophyinae and Libellulidae); the areas without crossveins basal and distal of the subnodus ("cordulegastrid gap" and "libellulid gap") are secondarily shortened; cubital cell (between CuP-crossing and pseudo-anal crossvein PsA) divided by accessory cubito-anal crossveins (convergent to Chlorogomphoidea, Synthemistidae, Idomacromiidae and some Idionychidae).
    
  • Other characters: tarsal claws with ventral tooth ("claw hook") distinctly prolonged, about as long as claw tip (GLOYD, 1959; CARLE & LOUTON, 1994; convergent to Zygonychina and some species of Macrothemis); unique armature of the larval gizzard (GLOYD, 1959); top of larval head with nipple-like projection or low rounded tubercle near the postero-lateral angle (GLOYD, 1959); larvae with very long spidery legs and rather small head even in the ultimate instar; larval frontal plate developed as upcurving pyramidal horn (TILLYARD, 1911, 1917; NEEDHAM & WESTFALL, 1955); incisor 2 of larval mandible is smaller than all other incisors (WATSON, 1956); specialised spines that accompany the movable crossvein junctions are obsolete.
• Taxonomy:
  
  • Macromiinae NEEDHAM, 1903 (subfam. nov.)
    
  • [group Macromina TILLYARD, 1911]
    
  • Marcomiini sensu TILLYARD, 1917 (stat. nov.)
    
  • Epophthalmiinae TILLYARD & FRASER, 1940 (subfam. nov. with type genus Epophthalmia BURMEISTER, 1839) (jun. subj. syn.)
    
  • Macrominae sensu NEEDHAM & WESTFALL, 1955 (incorr. subseq. spell.)
    
  • Macromiidae sensu DAVIES, 1981 (stat. nov.)

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Laterocarinida BECHLY, 1996

• Included taxa: Austrocorduliidae and Mediocostida.
• Autapomorphies:
  
  • Wing venation: basal side of hindwing discoidal triangle near arculus, separated by less than 1/2 length of arculus (GLOYD, 1959; CARLE & LOUTON, 1994; convergent to Gomphomacromiidae and Pseudocorduliidae); anal loop more or less elongate (convergent to Gomphomacromiidae and Pseudocorduliidae).
    
  • Other characters: male with elongated middorsal carina (convergent to † Tarsophlebiidae, Aeshnida and Synthemistidae) and lateral abdominal carinae on the terga of at least two segments, convergent to Aeshnodea sens. nov. (CALVERT, 1893; GLOYD, 1959; CARLE & LOUTON, 1994; CARLE, 1995).
• Taxonomy:
  
  • Laterocarinida BECHLY, 1996a (taxon nov.)

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Austrocorduliidae BECHLY, 1996

(Type genus: Austrocordulia TILLYARD, 1909)

• Included taxa: including the extant genera Lauromacromia GEIJSKES, 1970 and Syncordulia SELYS, 1882, as well as all the remaining Australian "gomphomacromiines" listed in BRIDGES (1994), except Hesperocordulia TILLYARD, 1911.
• Autapomorphies:
  
  • Wing venation: there is no secondary antenodal present between the two primary antenodal brackets ax1 and ax2, therefore the arculus is situated between the first two antenodal crossveins (convergent to Gomphomacromiidae, Pseudocorduliidae and Liberaponsida); reduction of all cubito-anal crossveins, except the CuP-crossing (= anal crossing sensu Fraser) and the pseudo-anal crossvein PsA (convergent to Gomphomacromiidae, Pseudocorduliidae and Liberaponsida); hypertriangle without crossveins (convergent to Gomphomacromiidae, Pseudocorduliidae and Liberaponsida).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Austrocorduliidae BECHLY, 1996a (fam. nov.)

Comment: this taxon might be paraphyletic in its present composition.

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Mediocostida BECHLY, 1996

• Included taxa: Idionychidae and Longiansida.
• Autapomorphies:
  
  • Wing venation: anal loop with two parallel cell rows that are separated by a concave bisector Cuspl ("midrib") which is still zigzagged in the groundplan (GLOYD, 1959; BECHLY, 1994); distinct Rspl present in both pairs of wings and more or less parallel to IR2 (convergent to Aeshnida).
    
  • Other characters: dark coloured areas of the body with distinct metallic-greenish gloss in the groundplan.
• Taxonomy:
  
  • Mediocostida BECHLY, 1996a (taxon nov.)

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Idionychidae TILLYARD & FRASER, 1940

(Type genus: Idionyx HAGEN, 1867.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: the two primary antenodal brackets ax1 and ax2 very indistinct or even indistinguishable from the secondaries (FRASER, 1937, 1957); sectors of arculus stalked (convergent to Synthemistidae, Macromiidae, Neophyinae and Libellulidae), with a long fusion (FRASER, 1957) ); postdiscoidal field of forewing commencing with a single row of cells.
    
  • Other characters: thorax rather small and weak (FRASER, 1957); mesotibiae with a strongly shortened tibial keel (LOHMANN, 1995).
• Taxonomy:
  
  • Idionychinae TILLYARD & FRASER, 1940 (subfam. nov.)
    
  • Idionychidae sensu BECHLY, 1996a (stat. nov.)

Comment: the inclusion of the genus Macromidia is doubtful, and this genus could at best represent the sistergroup of all other Idionychidae. It cannot be excluded that Macromidia might rather be a basal Macromiidae.

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Longiansida BECHLY, 1996

• Included taxa: Idomacromiidae and Liberaponsida.
• Autapomorphies:
  
  • Wing venation: anal loop and midrib distinctly elongated (with the tip pointing to the wing apex in the groundplan), correlated with an strongly elongated and sigmoidal gaff (= basal CuA before its branching).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Longiansida BECHLY, 1996a (taxon nov.)

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Idomacromiidae TILLYARD & FRASER, 1940

(Type genus: Idomacromia KARSCH, 1896.)

• Included taxa: Nesocorduliinae and Idomacromiinae.
• Autapomorphies:
  
  • Wing venation: RP3/4) and MA undulate in both pairs of wings (convergent to Macromiidae); unique shape of the elongated anal loop that has secondarily two rows of cells in the basal part anterior of the midrib; at least the hindwing cubital cell (between CuP-crossing and pseudo-anal crossvein PsA) is divided by several accessory cubito-anal crossveins (convergent to Chlorogomphoidea, Synthemistidae, Macromiidae and some Idionychidae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Idomacromiidae sensu BECHLY, 1996a (stat. et sens. nov.) (nom. transl. ex Idomacromiinae TILLYARD & FRASER, 1940)

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Nesocorduliinae BECHLY, 1996

(Type genus: Nesocordulia MCLACHLAN, 1882)

• Included taxa: only including the type genus Nesocordulia MCLACHLAN, 1882.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Nesocorduliinae BECHLY, 1996a (subfam. nov.)

Comment: although autapomorphies are not yet known this taxon is most probably monophyletic since it is mono-generic and endemic to Madagascar.

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Idomacromiinae TILLYARD & FRASER, 1940

(Type genus: Idomacromia KARSCH, 1896.)

• Included taxa: only including the type genus Idomacromia KARSCH, 1896.
• Autapomorphies:
  
  • Wing venation: very distinct Rspl and Mspl in both pairs of wings (convergent to † Aeschnidiidae, Euaeshnida, and Anauriculida); venation very dense (FRASER, 1957); the two primary antenodal brackets ax1 and ax2 are completely indistinguishable from the secondaries (FRASER, 1957); cubital cell (between CuP-crossing and pseudo-anal crossvein PsA) divided by accessory cubito-anal crossveins in both pairs of wings (convergent to Chlorogomphoidea, Synthemistidae, Macromiidae and some Idionychidae); anal loop very elongated with more than 8 cells in the row posterior of the midrib which is very straight (FRASER, 1957); hypertriangles traversed by several crossveins in both pairs of wings.
    
  • Other characters: females with a large pseudo-ovipositor (FRASER, 1957).
• Taxonomy:
  
  • Idomacromiinae TILLYARD & FRASER, 1940 (subfam. nov.)
    
  • [group Idocordulina TILLYARD, 1911 (taxon nov.)]
    
  • {Idocorduliini TILLYARD, 1917 (trib. nov.) (according to Art. 11 IRZN unavailable family-group name since not based on a type genus)}

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Liberaponsida BECHLY, 1996

• Included taxa: Cordulephyidae and Haplohamulida.
• Autapomorphies:
  
  • Wing venation: there is no secondary antenodal present between the two primary antenodal brackets ax1 and ax2, therefore the arculus is situated between the first two antenodal crossveins (convergent to Gomphomacromiidae, Pseudocorduliidae and Austrocorduliidae); bridge-space free (TILLYARD, 1911), rarely one crossvein present (reversed in some Tetrathemistinae); no cubito-anal crossveins retained, except the CuP-crossing (= anal crossing sensu Fraser) and the pseudo-anal crossvein PsA (convergent to Gomphomacromiidae, Pseudocorduliidae and Austrocorduliidae ?); hypertriangle without crossveins (convergent to Gomphomacromiidae, Pseudocorduliidae and Austrocorduliidae; reversed in some Libellulinae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Liberaponsida BECHLY, 1996a (taxon nov.)

Comment: the monophyly of Liberaponsida is still somewhat unsafe, and it can not be excluded that Cordulephyidae might eventually represent the sistergroup of Idomacromiidae.

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Cordulephyidae TILLYARD, 1917

(Type genus: Cordulephya SELYS, 1870.)

• Included taxa: Neophyinae and Cordulephyinae.
• Autapomorphies:
  
  • Wing venation: the two primary antenodal brackets ax1 and ax2 are very indistinct or even indistinguishable from the secondaries (FRASER, 1937, 1957); discoidal triangles secondarily quadrangular in both pairs of wings (TILLYARD, 1911; FRASER, 1957; convergent to some Tetrathemistinae); postdiscoidal field of forewing commencing with a single row of cells; Rspl strongly reduced or secondarily absent; CuAa parallel to hind margin of forewing.
    
  • Other characters: derived "similarities" of larvae (FRASER, 1957).
• Taxonomy:
  
  • Cordulephyidae sensu BECHLY, 1996a (stat. et sens. nov.) (nom. transl. ex Cordulephyini TILLYARD, 1917)

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Neophyinae TILLYARD & FRASER, 1940

(Type genus: Neophya SELYS, 1881.)

• Included taxa: only including the type species Neophya rutherfordi SELYS, 1881.
• Autapomorphies:
  
  • Wing venation: sectors of arculus stalked (convergent to Synthemistidae, Macromiidae, Idionychidae and Libellulidae) and strongly curved (costal side of hypertriangle very convex); trigonal vein that is separating the hypertriangle from the discoidal triangle, is distinctly curved; cubito-anal field of hindwings enormously dilated (FRASER, 1957); Rspl secondarily absent; RP2 and IR2, as well as RP3/4) and MA distally converging; besides the anal crossing (= CuP) no cubito-anal-crossveins present in hindwing (convergent to Pseudocordulia, Austrocordulia, Hesperocordulia, some Corduliidae and all Anauriculida) because of reduction of the basal side of the hindwing "subdiscoidal triangle".
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Neophyinae TILLYARD & FRASER, 1940 (subfam. nov.)

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Cordulephyinae TILLYARD, 1917

(Type genus: Cordulephya SELYS, 1870.)

• Included taxa: only including the type genus Cordulephya SELYS, 1870 (contra BRIDGES, 1994).
• Autapomorphies:
  
  • Wing venation: hindwing discoidal triangle secondarily shifted far distad of the arculus, correlated with an "asthenogenetic" hindwing like Tetrathemis; cubito-anal-field of hindwing strongly reduced, thus anal loop strongly reduced, too (only 2-4 cells), and anal angle, anal triangle and membranule vestigial (TILLYARD, 1911; FRASER, 1957); RP3/4 and MA distally diverging in forewings (reversal).
    
  • Other characters: wings are closely apposed over the dorsum at rest (TILLYARD, 1911; FRASER, 1957); unique shape of the larval prehensile mask, with several extremely deep incisions in the medio-apical margin of the lateral lobes ("palps") (TILLYARD, 1911); unique armature of larval gizzard, with only 3 minor denticles on the outer lateral edge of the tooth on the two outer lobes (TILLYARD, 1911); mesotibiae secondarily without tibial keel (LOHMANN, 1995; convergent to Anauriculida).
• Taxonomy:
  
  • Cordulephyini TILLYARD, 1917 (trib. nov.)
    
  • Cordulephyinae sensu TILLYARD & FRASER, 1940 (stat. nov.)

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Haplohamulida BECHLY, 1996

• Included taxa: Oxygastridae and Italoansida.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: male hamuli anteriores strongly reduced (SCHMIDT, 1916); tip of horn-like median lobe at the terminal segment of the male vesicula spermalis is prolonged to an unpaired "flagellum" (MAY, 1995b); male auricles without teeth (LOHMANN, 1996); body colour pattern without the plesiomorphic yellow stripes on the pterothorax.
• Taxonomy:
  
  • Haplohamulida BECHLY, 1996a (taxon nov.)

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Oxygastridae BECHLY, 1996

(Type genus: Oxygastra SELYS, 1870)

• Included taxa: probably only including the extant genera Hesperocordulia TILLYARD, 1911, Neocordulia SELYS, 1882, and Oxygastra SELYS, 1870.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larval abdomen with medio-dorsal tufts of setae instead of the dorsal spines that are secondarily absent (unknown for Hesperocordulia).
• Taxonomy:
  
  • Oxygastridae BECHLY, 1996a (fam. nov.)

Comment: this taxon might be paraphyletic in its present composition.

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Italoansida BECHLY, 1996

• Included taxa: Corduliidae and Anauriculida.
• Autapomorphies:
  
  • Wing venation: hindwing discoidal triangle recessed to the level of the arculus (FRASER, 1957; reversed in some Tetrathemistinae); forewing discoidal triangle divided in the groundplan into two cells by a longitudinal crossvein (FRASER, 1957); forewing subdiscoidal triangle divided into 3 cells in the groundplan (FRASER, 1957); wings with a Mspl, that is still somewhat indistinct and more or less parallel to MA in the groundplan (the alleged Mspl within Lindeniidae is indeed a secondary branch of MA); hindwing with elongated and boot-shaped anal loop ("italian loop" sensu TILLYARD, 1917) that nearly reaches the hind margin of the wing with its "toe", and with a straight (non-zigzagged) and forked midrib (Cuspl).
    
  • Other characters: body colour pattern without the plesiomorphic yellow markings on the abdomen; male secondary genitalia with muscle M6 (as antagonist to muscle M8III) attached near the basal joint of the vesicula spermalis, correlated with a derived "ejaculation chamber" with a reversed mechanism in which compression causes a sucking and decompression a spurting of sperm (PFAU, 1971; maybe an autapomorphy for a more inclusive group within the Eurypalpida since not yet analysed in all non-italoansid Eurypalpida; MAY & COOK, 1993 however mention derived similarities in the male ligula of all Italoansida, too); male hamuli anteriores obsolete, only the basal piece remains as vestige and fuses with the lamina anterior (SCHMIDT, 1916; GLOYD, 1959; CARLE & LOUTON, 1994); larval prehensile mask with the raptorial setae on the inner margin of the lateral lobes ("palps") shortened and its serrations less deep but more numerous.
• Taxonomy:
  
  • Italoansida BECHLY, 1996a (taxon nov.)

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Corduliidae SELYS in SELYS & HAGEN, 1850

(Type genus: Cordulia [LEACH] [1815].)

• Included taxa: Corduliinae and Neurocorduliinae.
• Autapomorphies:
  
  • Wing venation: not yet known, except the presence of only one crossvein beneath the pterostigma (convergent to some Anauriculida).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Cordulina SELYS, 1850 (nom. imperf.)
    
  • Cordulines; SELYS & HAGEN, 1850
    
  • [partim: Dynamiophlebiae BUCHECKER, 1876 (taxon nov.)]
    
  • Cordulina; CABOT, 1890
    
  • Cordulidae; BANKS, 1892 (stat. nov.)
    
  • Corduliidae; KARSCH, 1894
    
  • Corduliidae TILLYARD, 1926
    
  • Corduliidae sensu BECHLY, 1996a (sens. nov.)

Comment: since some Corduliidae share with Anauriculida the absence of cubito-anal crossveins and absence of mesotibial keels, this taxon might be paraphyletic in its present composition.

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Corduliinae SELYS in SELYS & HAGEN, 1850

(Type genus: Cordulia [LEACH] [1815].)

• Included taxa: including the remaining corduliine genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Cordulinae; BUCHECKER, 1876 (stat. nov. ?) (nom. transl. ex Cordulina SELYS, 1850)
    
  • incl. Epithecinae BUCHECKER, 1876 (subfam. nov. with type genus Epitheca) (jun. subj. syn.)
    
  • Corduliinae; KIRBY, 1890 (nom. correct.)
    
  • Cordulinae; NEEDHAM, 1903
    
  • Cordulinae; CALVERT, 1909
    
  • [partim: group Eucordulina TILLYARD, 1911 (taxon nov.)]
    
  • Corduliinae; TILLYARD, 1917
    
  • {partim: Eucorduliini TILLYARD, 1917 (trib. nov.) (according to Art. 11 IRZN unavailable family-group name since not based on a type genus)}
    
  • Corduliinae TILLYARD, 1926 (sens. nov.)
    
  • Corduliinae; TILLYARD, 1929 (sens. nov.)
    
  • Corduliinae; TILLYARD & FRASER, 1940 (sens. nov.)
    
  • Corduliinae sensu BECHLY, 1996a (sens. nov.)

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Neurocorduliinae BECHLY, 1996

(Type genus: Neurocordulia Selys, 1871)

• Included taxa: including the extant genera Heteronaias NEEDHAM & GYGER, 1937, Libellulosoma MARTIN, [1907], Aeschnosoma SELYS, 1870, and Neurocordulia SELYS, 1871.
• Autapomorphies:
  
  • Wing venation: forewing discoidal triangle divided into three cells (unique within Cavilabiata).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Neurocorduliinae BECHLY, 1996a (subfam. nov.)

Comment: Aeschnosoma and Libellulosoma are sister-genera with the quadri-cellular hypertriangle of the forewings as synapomorphy.

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Anauriculida BECHLY, 1996

• Included taxa: Hemicorduliidae and Libellulida.
• Autapomorphies:
  
  • Wing venation: male hindwing anal angle reduced (FRASER, 1937, 1957; BECHLY, 1994; convergent to Anacina and † Aeschnidiidae, and correlated with the reduction of auricles); the two primary antenodal brackets ax1 and ax2 are less distinct (FRASER, 1937); besides the anal crossing (= CuP) no cubito-anal-crossveins present in hindwing (BECHLY, 1994; convergent to Pseudocordulia, Austrocordulia, Neophya, Hesperocordulia; maybe synapomorphic with some Corduliidae; reversed in some derived libellulids) because of reduction of the basal side of the hindwing "subdiscoidal triangle"; characteristical curved shape of Rspl that distally "rejoining" the IR2 (NEL & MARTÍNEZ-DELCLÒS, 1993; convergent to † Aeschnidiidae, † Cymatophlebiidae and higher Aeshnidae); Mspl very distinct and curved, distally "rejoining" the MA (NEL & MARTÍNEZ-DELCLÒS, 1993; convergent to † Aeschnidiidae and Aeshnidae; reduced in Tetrathemistinae).
    
  • Other characters: male auricles reduced (FRASER, 1937, 1957; BECHLY, 1994; convergent to Anacina); at least mesotibiae without tibial keel.
• Taxonomy:
  
  • Anauriculida BECHLY, 1996a (taxon nov.)

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Hemicorduliidae BECHLY, 1996

(Type genus: Hemicordulia SELYS, 1870)

• Included taxa: only including the genus Hemicordulia SELYS, 1870 and maybe certain species of Procordulia MARTIN, 1907.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Hemicorduliidae BECHLY, 1996a (fam. nov.)

Comment: although no autapomorphies are not yet known the genus Hemicordulia most probably is monophyletic.

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Libellulida BECHLY, 1996

• Included taxa: Urothemistidae and Eulibellulida (= Libellulidae).
• Autapomorphies:
  
  • Wing venation: the two primary antenodal brackets ax1 and ax2 are very indistinct in the forewings and indistinguishable from the secondaries in hindwings (FRASER, 1957); the second crossvein between RP1 and RP2 is developed as "libellulid oblique vein"; the anal triangle of the male hindwing is secondarily absent; anal loop reaches the hind margin of the hindwing (well-developed "toe"); the area between RA and RP1 stays only unicellular for 2-5 cells (with rare exceptions that do not represent the groundplan condition) because RA and RP1 are distinctly diverging (convergent to Austropetaliida and Neopetaliidae, where this character state is caused by an elongated IR1).
    
  • Other characters: male tibial keels secondarily absent on all legs (FRASER, 1957); posterior tubercles of compound eyes secondarily absent; body colour secondarily non-metallic (FRASER, 1957); the "Anlagen" of the vulvar scale (ovipositor vestige) are not externally visible in female larvae (HEIDEMANN & SEIDENBUSCH, 1993); male secondary genitalia with strongly reduced ligula and without suture in the lamina anterior, thus apparent absence of the basal pieces of the vestigial hamuli anteriores (SCHMIDT, 1916); males guard the ovipositing females or oviposition is performed in tandem (convergent to some Aeshnidae); serrations of the inner margin of the lateral lobes ("palps") of the larval prehensile mask much flattened and indistinct (small undulations); resting behaviour usually with a horizontal or even upturned position of the abdomen (vertical position in most other Anisoptera, except many gomphids, and secondarily so in Zygonyx) (ASKEW, 1988).
• Taxonomy:
  
  • Libellulida BECHLY, 1996a (taxon nov.) (nec Libellulida LEACH, 1815)

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Urothemistidae LIEFTINCK, 1954

(Type genus: Urothemis BRAUER, 1868.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: very oblique and distinct "libellulid oblique vein" (BECHLY, 1994); venation quite open, with few antenodal crossveins (about 5) and only very few (3) antesubnodal crossveins at least in hindwings (FRASER, 1957; BECHLY, 1994).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Urothemistinae LIEFTINCK, 1954 (subfam. nov.)
    
  • Macrodiplactidae FRASER, 1957 (fam. nov. with type genus Macrodiplax BRAUER, 1868) (nom. imperf.) (jun. subj. syn.)
    
  • Macrodiplacinae sensu DAVIES, 1981 (nom. correct. et stat. nov.) (jun. subj. syn.)
    
  • Urotheminae sensu PINHEY, 1962 (incorr. subseq. spell.)
    
  • Macrodiplacidae sensu BECHLY, 1996a (jun. subj. syn.)
    
  • Urothemistidae sensu BECHLY, 2003e (stat. nov.)

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Eulibellulida BECHLY, 1996

• Included taxa: preliminarily only including the family Libellulidae LEACH, 1815, with the type genus Libellula LINNAEUS, 1758.
• Autapomorphies:
  
  • Wing venation: sectors of arculus stalked (convergent to Synthemistidae, Macromiidae, Idionychidae and Neophyinae); all antenodal crossveins are developed as antenodal brackets, therefore the two primaries ax1 and ax2 are completely indistinguishable from the secondaries in both pairs of wings (FRASER, 1937, 1957; BECHLY, 1994); a strong concave supplementary sector developed between RP1 and RP2 (apical supplement); the most distal antenodal between costal margin and ScP is developed as antenodal oblique vein, with a strong tendency towards suppression of the corresponding antenodal between ScP and RA.
    
  • Other characters: temples of head secondarily not swollen (FRASER, 1957); male secondary genitalia with a derived mechanism for loading the sperm-reservoir, correlated with a reduction of the dorso-ventral muscle M9b (PFAU, 1971; but this could also be an autapomorphy for Libellulida or even Anauriculida since not yet analysed in the respective groups).
  • Taxonomy (of Libellulidae):
    
  • Libellulida LEACH, 1815 (nec Libellulides LEACH, 1815)
    
  • Libellulidae BURMEISTER, 1839 (nec Libellulidae STEPH., 1836, nec Libellulidae WESTWOOD, 1840)
    
  • Libelluloides SELYS, 1840 (nec Libelluloides LAICHARTING, 1781)
    
  • Libellulides RAMBUR, 1842 (nec Libellulides LEACH, 1815, nec Libellulides WESTWOOD, 1840)
    
  • Libellulines SELYS & HAGEN, 1850
    
  • Lbellulidae SELYS, 1850
    
  • Libellulina SELYS, 1850 (nec Libellulina SELYS, 1840, nec Libellulina NEWMAN, 1834)
    
  • Libellulina; HAGEN, 1861
    
  • [incl. Monotoxophlebiae & Dytoxophlebiae BUCHECKER, 1876]
    
  • Libellulinae; KIRBY, 1890 (nec Libellulinae LATREILLE, 1802, nec Libellulinae SELYS, 1840, nec Libellulinae SWAINSON, 1840)
    
  • Libellulida; HAECKEL, 1896
    
  • Libellulii; ACLOQUE, 1897
    
  • [Libellulidos NAVAS, 1903 (trivial name?)]
    
  • Libellulinae; NEEDHAM, 1903
    
  • Libellulidae; NEEDHAM, 1903
    
  • [Libellulodea JACOBSON & BIANCHI, 1905 (taxon nov. ?)]
    
  • Libellulinae; TILLYARD, 1926 (sens. nov.)
    
  • Libelluloidea; TILLYARD, 1926 (stat. nov.) (nec Libelluloidea KARSCH, 1896, nec Libelluloidea HANDLIRSCH, 1906)
    
  • Libelluloidea; TILLYARD & FRASER, 1940
    
  • Libelluloidea; PRITYKINA, 1980
    
  • [Eulibellulida BECHLY, 1996a (taxon nov.)]

Comment: the new taxon Eulibellulida which is presently still coextensive with Libelluloidea (auct.) and Libellulidae (auct.), shall facilitate a future subdivision and phylogenetic systematisation of this diverse group into several superfamilies and families.

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"Tetrathemistinae" TILLYARD, 1917

(Type genus: Tetrathemis BRAUER, 1868.)

• Included taxa: Brachydiplacini and Tetrathemistini.
• Autapomorphies:
  
  • Wing venation: anal loop more or less reduced or even obsolete (FRASER, 1957); cubito-anal area with accessory crossveins (FRASER, 1957); bridge-space secondarily divided by crossveins (FRASER, 1957); at least the forewing discoidal triangle secondarily quadrilateral (FRASER, 1957); Rspl secondarily more or less parallel to IR2 (FRASER, 1957); libellulid oblique vein secondarily absent.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Tetratheminae; TILLYARD, 1926 (stat. nov.) (nom. transl. ex Tetrathemini TILLYARD, 1917)
    
  • Tetratheminae; TILLYARD & FRASER, 1940
    
  • Tetrathemistinae; DAVIES, 1981 (nom. correct.)
    
  • Tetrathemistinae sensu BECHLY, 1996a (sens. nov.)

Comment: all mentioned characters do not belong to the groundplan of "Tetrathemistinae", but those genera of Brachydiplacinae and Tetrathemistinae that share these derived characters states might form a monophylum, while those genera that do not are of uncertain position.

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Brachydiplacini TILLYARD, 1917

(Type genus: Brachydiplax BRAUER, 1868.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Brachydiplacini TILLYARD, 1917 (trib. nov.)
    
  • Brachydiplacinae sensu TILLYARD & FRASER, 1940 (stat. nov.)
    
  • Brachydiplactinae FRASER, 1957 (incorr. subseq. spell.)

Comment: probably non-monophyletic in the present extent.

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Tetrathemistini TILLYARD, 1917

(Type genus: Tetrathemis BRAUER, 1868.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: arculus situated beneath the second antenodal or even distal of it; sectors of arculus with long stalk; anal loop obsolete (FRASER, 1957; not in the groundplan); postdiscoidal area with only one row of cells (FRASER, 1957; not in the groundplan).
    
  • Other characters: body colouring steely black with bright citron yellow markings (FRASER, 1957).
• Taxonomy:
  
  • Tetrathemini TILLYARD, 1917 (trib. nov.) (nom. imperf.)
    
  • Tetratheminae sensu TILLYARD, 1926
    
  • Tetratheminae sensu TILLYARD & FRASER, 1940
    
  • Tetrathemistinae sensu DAVIES, 1981 (nom. correct.)

Comment: probably non-monophyletic in the present extent; including several genera that have previously been classified in Brachydiplactinae.

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"Libellulinae" LEACH, 1815

(Type genus: Libellula LINNAEUS, 1758.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larval occiput with parallel lateral borders, like Macromiidae (HEIDEMANN & SEIDENBUSCH, 1993); larval legs stronger (profemur max. 4 times longer than broad) and more densely haired (HEIDEMANN & SEIDENBUSCH, 1993); mature males with bluish pruinescent body colour in the groundplan (rather doubtful character).
• Taxonomy:
  
  • Libellulinae KIRBY, 1890 (stat. nov. ?)
    
  • Libellulinae; NEEDHAM, 1903
    
  • Libellulinae; JACOBSON & BIANCHI, 1905 (sens. nov.)
    
  • Libellulini sensu TILLYARD, 1917 (stat. nov.)
    
  • Libellulinae; TILLYARD, 1926 (sens. nov.)

Comment: the mentioned characters are mainly synapomorphies on Libellula and Orthetrum, and may not belong to the groundplan of Libellulinae which therefore might be paraphyletic in the present extent. A distinct monophylum within Libellulinae includes all species with an reduced cubito-anal field in the hindwing and a position of the arculus distal of the second antenodal which might even be synapomorphies with Tetrathemistini.

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Trithemistinae TILLYARD, 1917

(Type genus: Trithemis BRAUER, 1868.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: nodus shifted distad, thus apical part of wing shortened (FRASER, 1957; somewhat doubtful character).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Trithemini TILLYARD, 1917 (trib. nov.) (nom. imperf.)
    
  • Tritheminae sensu TILLYARD & FRASER, 1940 (stat. nov.) (nom. imperf.)
    
  • Trithemistinae sensu DAVIES, 1981 (nom. correct.)

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Sympetrinae TILLYARD, 1917

(Type genus: Sympetrum NEWMAN, 1833.)

• Included taxa: Sympetrini and Leucorrhiniini.
• Autapomorphies:
  
  • Wing venation: supplementary crossveins in the cubito-anal space and the bridge-space present in the groundplan.
    
  • Other characters: larvae without a fringe of setae along the suture between vertex and occiput which belongs to the groundplan of Haplohamulida (HEIDEMANN & SEIDENBUSCH, 1993; but not yet examined in all genera and the possibly related Trithemistinae).
• Taxonomy:
  
  • Sympetrinae; TILLYARD & FRASER, 1940 (stat. nov.) (nom. transl. ex Sympetrini TILLYARD, 1917)
    
  • Sympetrinae sensu BECHLY, 1996a (sens. nov.)

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Sympetrini TILLYARD, 1917

(Type genus: Sympetrum NEWMAN, 1833.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: mature males with bright red body colour in the groundplan (rather doubtful character).
• Taxonomy:
  
  • Sympetrini TILLYARD, 1917 (trib. nov.)
    
  • Sympetrinae sensu TILLYARD & FRASER, 1940

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Leucorrhiniini TILLYARD, 1917

(Type genus: Leucorrhinia BRITTINGER, 1850.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: sectors of arculus at base secondarily unfused (arculus not stalked) (FRASER, 1957).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Leucorrhiniini TILLYARD, 1917 (trib. nov.)
    
  • Leucorrhininae sensu TILLYARD & FRASER, 1940 (stat. nov. and incorr. subseq. spell.)
    
  • Leucorrhiniinae; NEL & PAICHELER, 1993

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Palpopleurinae JACOBSON & BIANCHI, 1905

(Type genus: Palpopleura RAMBUR, 1842.)

• Included taxa: Palpopleurini and Diastatopidini.
• Autapomorphies:
  
  • Wing venation: basal costal field of forewings expanded and with a sinuous hump in the strongly convex costal margin (secondarily absent in Perithemis) (FRASER, 1957); wings relatively broad and short (FRASER, 1957).
    
  • Other characters: territorial and courtship behaviour (?).
• Taxonomy:
  
  • Palpopleurinae JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • Palpopleurini sensu TILLYARD, 1917 (stat. nov.)
    
  • Palpopleurinae sensu DAVIES, 1981

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Palpopleurini JACOBSON & BIANCHI, 1905

(Type genus: Palpopleura RAMBUR, 1842.)

• Included taxa: only including the two genera Palpopleura RAMBUR, 1842 and Perithemis HAGEN, 1861.
• Autapomorphies:
  
  • Wing venation: not yet known, except the small wing span and typical wing colour pattern.
    
  • Other characters: tiny size and very characteristical body shape (e.g. short and broad abdomen), with yellowish colouring, correlated with a wasp-mimicry.
• Taxonomy:
  
  • Palpopleurini; TILLYARD, 1917 (stat. nov.) (nom. transl. ex Palpopleurinae JACOBSON & BIANCHI, 1905)
    
  • [Diastatopidinae sensu TILLYARD & FRASER, 1940 (subfam. nov.)]
    
  • Palpopleurini sensu BECHLY, 1996a (sens. nov.)

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Diastatopidini TILLYARD & FRASER, 1940

(Type genus: Diastatops RAMBUR, 1842.)

• Included taxa: only including the two genera Diastatops RAMBUR, 1842 and Zenithoptera SELYS, 1877.
• Autapomorphies:
  
  • Wing venation: wings coloured dark with blue-metallic gloss.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Diastatopidinae TILLYARD & FRASER, 1940 (subfam. nov.)
    
  • Diastatopidini sensu BECHLY, 1996a (stat. et sens. nov.)

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Onychothemistinae TILLYARD & FRASER, 1940

(Type genus: Onychothemis BRAUER, 1878.)

• Included taxa: Rhyothemistini and Onychothemistini.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: at least slightly metallic body colour (reversal) (FRASER, 1957).
• Taxonomy:
  
  • Onychotheminae TILLYARD & FRASER, 1940 (subfam. nov.) (nom. imperf.)
    
  • Onychothemistinae; DAVIES, 1981 (nom. correct.)
    
  • Onychothemistinae sensu BECHLY, 1996a (sens. nov.)

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Rhyothemistini TILLYARD & FRASER, 1940

(Type genus: Rhyothemis HAGEN, 1867, nec Rhysthemis GRÜNBERG, 1911, incorr. subseq. spell, nec Rhyiothemis OGUMA, 1922, incorr. subseq. spell.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: abdomen very short, only slightly longer than half wing length (FRASER, 1957); body never fully chitinised (FRASER, 1957), correlated with a weak flight (fluttering and soaring); mimicry of Rhopalocera and large Hymenoptera (FRASER, 1957).
• Taxonomy:
  
  • Rhyotheminae TILLYARD & FRASER, 1940 (subfam. nov.) (nom. imperf.)
    
  • Rhyothemistini; DAVIES, 1981 (nom. correct. et transl.)

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Onychothemistini TILLYARD & FRASER, 1940

(Type genus: Onychothemis BRAUER, 1878.)

• Included taxa: Zygonychina and Onychothemistina.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Onychothemistini sensu BECHLY, 1996a (stat. et sens. nov.) (nom. transl. ex Onychotheminae TILLYARD & FRASER, 1940)

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Zygonychina FRASER, 1957

(Type genus: Zygonyx SELYS, 1867.)

• Included taxa: including the genera and species listed in BRIDGES (1994), except the type species Olpogastra lugubris KARSCH, 1895.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: the hook of the tarsal claws is strongly prolonged (convergent to Macromiidae and some species of Macrothemis) (FRASER, 1957); derived "corduliine-like" behaviour, with a resting position in a more or less vertical attitude (FRASER, 1957), the abdomen hanging down.
• Taxonomy:
  
  • Zygonictinae FRASER, 1957 (subfam. nov.) (nom. imperf.)
    
  • Zygonychinae; DAVIES, 1981 (nom. correct.)
    
  • Zygonichinae; ASKEW, 1988
    
  • Zygonychina sensu BECHLY, 1996a (stat. et sens. nov.)

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Onychothemistina TILLYARD & FRASER, 1940

(Type genus: Onychothemis BRAUER, 1878.)

• Included taxa: this taxon does not only include the genus Onychothemis BRAUER, 1868, but also Olpogastra lugubris KARSCH, 1895 that was previously classified as Zygonictinae by FRASER, 1957.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: hook of tarsal claws completely reduced (convergent to Perilestidae, and Leptobasinae and Teinobasini) (FRASER, 1957).
• Taxonomy:
  
  • Onychothemistina sensu BECHLY, 1996a (stat. et sens. nov.) (nom. transl. ex Onychotheminae TILLYARD & FRASER, 1940)

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Zyxommatinae JACOBSON & BIANCHI, 1905

(Type genus: Zyxomma RAMBUR, 1842.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: anal loop secondarily posteriorly open because AA1b and CuAa apically do not fuse (FRASER, 1957; but not yet present in the most basal genus Parazyxomma).
    
  • Other characters: very large and very broadly confluent eyes (like gynacanthine Aeshnidae), correlated with crepuscular habits (FRASER, 1957).
• Taxonomy:
  
  • Zyxomminae JACOBSON & BIANCHI, 1905 (subfam. nov.) (nom. imperf.)
    
  • {Zyxommatinae TILLYARD & FRASER, 1940 (subfam. nov.) (jun. obj. syn. & homonym)}
    
  • Zyxommatini; DAVIES, 1981 (stat. nov.)

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Pantalinae JACOBSON & BIANCHI, 1905

(Type genus: Pantala HAGEN, 1861.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: Rspl and Mspl distinct like a primary longitudinal vein and with 2-3 cell rows between (FRASER, 1957), in forewings and/or hindwings; hindwing anal field strongly broadened, with numerous small cells basal of the anal loop, correlated with a characteristical triangular shape of hindwings (FRASER, 1957).
    
  • Other characters: relatively small thorax and abdomen shorter than wing span; eyes large and broadly confluent (probably convergent to Zyxommatinae).
• Taxonomy:
  
  • Pantalinae JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • Trameini TILLYARD, 1917 (trib. nov. with type genus Tramea HAGEN, 1861, jun. obj. syn. of Trapezostigma HAGEN, 1849)
    
  • Trameinae; TILLYARD & FRASER, 1940 (stat. nov.)
    
  • Tramiini; WESERBERG-LUND, 1943 (incorr. subseq. spell.)
    
  • Trameini; DAVIES, 1981 (sens. nov.)
    
  • {Pantaliinae FRASER, 1957 (subfam. nov.) (jun. obj. syn. & homonym)}
    
  • Pantalinae; BRIDGES, 1994 (incorrectly attributed to FRASER, 1957)