Phylogenetic Systematics of Odonata

Exophytica BECHLY, 1996

• Included taxa: Gomphides and Cavilabiata.
• Autapomorphies:
  
  • Wing venation: no wing venational autapomorphies, except a very homoplastic character (distal accessory oblique vein obsolete in both pairs of wings, convergent to Neoaeshnida and some Petaluridae, etc.).
    
  • Other characters: small spines on the dorsal side of distal side (MAb) of the discoidal triangle secondarily absent; males with apical protibial pads ("callum") in the groundplan (LOHMANN, 1995, 1996; these structures in Gomphides and Cordulegastrida are interpreted by Lohmann and me as plesiomorphic homologues of true tibial keels in Carinitibiata, contra CARLE, 1995); males with peg-like spines ("tumidotrichae" sensu Lohmann) on the meso- and metatibiae (LOHMANN, 1995, 1996; secondarily absent in many Gomphides, some Chlorogomphida and Synthemistidae and all Neolamellida, therefor a rather weak character since it could also be secondarily absent in the non-exophytic Anisoptera, too); meso- and metatibiae of both sexes developed as "plagiotibiae" with an asymmetric cross section (LOHMANN, 1995, 1996; reversed in Neolamellida); the spermal furrow on the third segment of the male vesicula spermalis is completely closed, convergent to some Petaluridae, Austropetaliida and Neoaeshnida (PFAU, 1971; LOHMANN, 1995, 1996); in tandem the male appendices superiores engage only the female occiput (reversal), not parts of the female pronotum like still in Petalurida and Aeshnoptera (WILLIAMSON, 1899; SCHMIDT, 1916; TILLYARD, 1917); fusion of the meso- and metathoacic ganglia, convergent to Euzygoptera (CALVERT, 1893; TILLYARD, 1917; contra ASAHINA, 1957); characteristic reductions in the female ovipositor (pro- and mesogonapophyses at least somewhat reduced and metagonapophyses distinctly reduced; muscles 3, 7 and 8 secondarily absent) which is exophytic, correlated with an oviposition in flight with deposition of a high number of ellipsoidal eggs (ST. QUENTIN, 1962; ASKEW, 1988; PFAU, 1991; LOHMANN, 1995, 1996; contra CARLE, 1995); embryonal germ band is distinctly sigmoidal (ANDO, 1962); presence of an abdominal transverse muscle 4 in larvae (non-phragmatic in groundplan); the sidelobes or lateral spines are at least reduced on the first two abdominal segments of the larvae (LOHMANN, 1996); larvae with a pronounced frontal plate in the groundplan (LOHMANN, 1996), correlated with burrowing life style in the groundplan (maybe a convergence because of different ways of burrowing in Gomphides and Cavilabiata, according to HEIDEMANN & SEIDENBUSCH, 1993; CARLE, 1995).
• Taxonomy:
  
  • Exophytica BECHLY, 1996a (taxon nov.)
    
  • Exophyticata LOHMANN, 1996 (taxon nov.)
    
  • Exophytica sensu NEL et al., 1998

Comment: all previous arguments, e.g. by PFAU (1991) and CARLE (1995), against a sistergroup relationship of Gomphides and Cavilabiata have been demonstrated to be inconclusive by NEL et al., 1998.

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Gomphides BECHLY et al., 1998

• Included taxa: † Araripegomphidae, Progomphidae, and Desmoproctida.
• Autapomorphies:
  
  • Wing venation: pseudo-anal vein PsA and subdiscoidal triangle of hindwings more distinct than in the groundplan of Anisoptera (except in † Araripegomphidae, Lindeniinae and Hagenioidea; at least in the latter two probably a reversal, correlated with the elongated hindwing discoidal triangle); hindwing discoidal triangle in the groundplan with an at least somewhat angulated distal side, caused by a more or less distinct supplementary sector (trigonal planate) in the postdiscoidal area; hypertriangles with distinctly curved anterior margin, convergent to Eurypalpida; membranule more or less reduced; costal margin and RA thickened along the pterostigmata (CARLE, 1986; reversed in Hageniidae); strong tendency towards the development of a straight arculus.
    
  • Other characters: prementum without a median cleft (fissum) in larvae and adults (CARLE & LOUTON, 1994; convergent to Eurypalpida); adult labial "palp" with a long spine-like process on the movable end hook which is about as long as the ligula (CARLE, 1995); larval antennae with only 4 segments in the groundplan, with a distinctly enlarged third segment and a vestigial or obsolete fourth segment (CARLE, 1995); complete reduction of the dorsal suture (vestige of the spermal furrow) on the third segment of the male vesicula spermalis (LOHMANN, 1995, 1996; convergent to Eurypalpida); the male hamuli posteriores engaging the female "sternum" 9 with an anterior stroke of the apical hooks (CARLE & LOUTON, 1994; CARLE, 1995); female ovipositor extremely reduced and all its muscles obsolete(progonocoxae and progonapophyses reduced and fused to a vulvar lamina, mesogonapophyses absent and gonostyli obsolete, but metagonapophyses vestigial as small " paired sternal plates 9" that are grasped by the male hamuli posteriores) (CARLE, 1995), convergent to Chlorogomphida and Trichodopalpida (a long plesiomorphic ovipositor is retained in † Araripegomphidae which therefore seem to represent the sistergroup to all other Gomphides; BECHLY, unpubl. pers. observ.); larval mandibles with a movable basal articulation of the second molar segment (LOHMANN, 1995, 1996; here preliminarily interpreted as reversal within Anisoptera; contra CARLE, 1995. Nevertheless this character state could also be interpreted as a plesiomorphy which would imply a convergent fusion in Petalurida, Aeshnoptera and Cavilabiata.); larvae with only two tarsomeres at least on pro- and mesotarsi (CARLE, 1992, 1995); larval abdomen with finger-like projecting antero-lateral sternal apodemes on segments 4-6, instead of only a single ridge-like antero-lateral apodeme on segment 6 in the groundplan of Anisoptera (LOHMANN, 1995, 1996; here regarded as autapomorphy; contra CARLE, 1995); males with numerous irregular short spines on the femora; adult postfrons with a postocellar ridge with two tubercles (CARLE, 1986, 1995; belonging to the groundplan of Gomphides, contra LOHMANN, 1995); epiproct of adult males very strongly bifurcate; costal margin of wings between the two rows of spines rather flat, not developed as a distinctly concave furrow (BECHLY, 1994); number of macrotrichae on the wing veins strongly reduced (convergent to Cordulegastrida); horizontal type of emergence, with the back never pointing downwards; characteristic burrowing and camouflage behaviour, different from Cavilabiata (HEIDEMANN & SEIDENBUSCH, 1993; CARLE, 1995); anterior margin of larval prementum with a dense row of scale-like setae.
• Taxonomy:
  
  • [légion Gomphus SELYS & HAGEN, 1850 (not a valid family-group taxon according Art.11f IRZN)]
    
  • [partim: Integrilabres SELYS, 1854 (taxon nov.)]
    
  • [partim: Dynamophlebiae BUCHECKER, 1876 (taxon nov.)]
    
  • Gomphidae sensu auct. (except BECHLY, 1996a)
    
  • Gomphoidea sensu auct. (except BECHLY, 1996a)
    
  • [Gomphata sensu LOHMANN, 1995 (unpublished manuscript name according to Art. 8 IRZN)]
    
  • Gomphata sensu BECHLY, 1996a (explicitly a citation of Lohmann's unpublished manuscript name)
    
  • Gomphata LOHMANN, 1996 (taxon nov.)
    
  • Gomphides BECHLY et al., 1998 (taxon nov.) (nec Gomphides RAMBUR, 1842)

Comment: since there are no strong autapomorphies in the wing venation of Gomphides, a positive assignment of fossil stemgroup representatives will always remain rather difficult.
Although the name "Gomphata" was already proposed for this clade by LOHMANN (1995, 1996), the new name "Gomphides" should be preferred because of the following two reasons: (1) Lohmann is using the suffix "-ata" explicitly to give equal suffixes to sistergroups, which has to be regarded as a rather scientific and formalistic goal; (2) Since this clade was previously addressed under the family-group name Gomphidae a scientific name which sounds similar and which allows the further use of the vernacular expression "gomphids" would be of significant advantage.

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† Araripegomphidae BECHLY, 1996

(Type genus: † Araripegomphus NEL & PAICHELER, 1994.)

• Included taxa: only including the genus † Araripegomphus NEL & PAICHELER, 1994.
• Autapomorphies:
  
  • Wing venation: discoidal triangles secondarily unicellular (convergent to † Proterogomphidae and Gomphida); anal loop obsolete (reduced and posteriorly open; convergent to Gomphida and Progomphidae ?); presence of a distinct "cordulegastrid gap" (lack of crossveins in the distal antesubnodal space) in both pairs of wings (convergent to † Cordulagomphinae, Gomphaeschnidae, and Cavilabiata); bases of RP and MA (sectors of arculus) with common origin at arculus (convergent to crown-group Eurypalpida).
    
  • Other characters: compound eyes closely approximate (but not touching; convergent to the groundplans of Aeshnomorpha and Cavilabiata).
• Taxonomy:
  
  • Araripegomphidae BECHLY, 1996a (fam. nov.)
    
  • Araripegomphidae LOHMANN, 1996 (fam.nov.) (jun. obj. syn.)

Comment: LOHMANN (1996) mentioned three alleged autapomorphies of † Araripegomphidae which are all invalid: anal loop two-celled (clearly incorrect); gaff secondarily shortened (unwarranted ad hoc assumption; plesiomorphy); secondary antenodal crossveins between ax1 and ax2 aligned (variable).
LOHMANN (1995, 1996) also concluded that † Araripegomphus is a stemgroup representative of Eurypalpida because of several apomorphic similarities. This possibility was already indicated in the original description, but has been rejected by BECHLY (1998d) who discussed the available evidence and arguments. The character pattern rather suggests that the mentioned similarities are convergences and that Araripegomphus is a true Gomphides since † Araripegomphus shares all wing venational apomorphies of Gomphides, and most of the similarities with Eurypalpida cited by Lohmann are also known from some Gomphides. The most important evidence against an eurypalpid relationship is the plesiomorphic state of the gaff (= basal CuA before its branching) and the anal loop which excludes a position within Brachystigmata since a reversal in this character complex can be excluded with great certainty. The closely approximate eyes are unique within Gomphides, but a convergent evolution of this character is quite possible, since the derived state has been evolved at least twice in Aeshnomorpha and Cavilabiata anyway.
The presence of a long ovipositor in female † Araripegomphidae is a unique plesiomorphy that strongly suggests a sistergroup relationship with all other Gomphides (BECHLY, unpubl. pers. observ.).

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Progomphidae CARLE, 1986

(Type genus: Progomphus SELYS, 1854.)

• Included taxa: including the genus Progomphus SELYS, 1854.
• Autapomorphies:
  
  • Wing venation: anal loop obsolete (convergent to Gomphida and † Araripegomphidae ?); hindwing subdiscoidal triangle divided into two cells by a transverse crossvein.
    
  • Other characters: adult males with a transverse row of denticles (convergent to Lestinogomphini) or a chitinous ridge on the anterior face of the basally broadly expanded hamuli posteriores (CARLE, 1986); prementum of larval prehensile mask secondarily elongated (here not regarded as plesiomorphy, contra LOHMANN, 1995, 1996); adult males with a uniquely shaped trifurcate epiproct (the bifurcate epiproct of all Desmoproctida is here regarded as plesiomorphic, contra LOHMANN, 1995, 1996).
• Taxonomy:
  
  • Progomphini CARLE, 1986 (trib. nov.)
    
  • Progomphini sensu BELLE, 1996
    
  • Progomphidae sensu BECHLY, 1996a (stat. nov.)

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Desmoproctida BECHLY, 1996

• Included taxa: Lindeniidae and Oligophlebiata.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larval epiproct process situated on the same level as the epiproct and fused with the latter (convergent to Trichodopalpida), and larval anal pyramid shortened (convergent to Trichodopalpida; but still rather long in Zonophoridae and Gomphoidini) (DE MARMELS, 1990; LOHMANN, 1995, 1996).
• Taxonomy:
  
  • Desmoproctida BECHLY, 1996a (taxon nov.)

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Lindeniidae JACOBSON & BIANCHI, 1905

(Type genus: Lindenia DE HAAN, 1826.)

• Included taxa: Gomphoidinae and Lindeniinae.
• Autapomorphies:
  
  • Wing venation: hypertriangle divided by crossveins into at least 2 cells in both pairs of wings; IR2 and MA with a more or less distinct secondary branch.
    
  • Other characters: at least males with large lateral leaf-like expansions on abdominal segment 8; adult males with a distinctly shortened epiproct.
• Taxonomy:
  
  • Lindeniidae sensu BECHLY, 1996a (stat. et sens. nov.) (nom. transl. ex Lindeniinae JACOBSON & BIANCHI, 1905)

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Gomphoidinae JACOBSON & BIANCHI, 1905

(Type genus: Gomphoides SELYS, 1854, nec Gomphoides SELYS, 1850; the latter is maybe an unavailable name that should otherwise be surpressed by the ICZN.)

• Included taxa: Aphyllini, Diaphlebiini, Gomphoidini, and Peruviogomphini.
• Autapomorphies:
  
  • Wing venation: anal triangle extended to tornus (CARLE, 1986; convergent to Idiogomphoides) and separated into three cells by two parallel crossveins instead of the usual Y-shaped vein; membranule reduced or absent.
    
  • Other characters: male prepuce vestigial (CARLE, 1986); larval prehensile mask with the endhook of the lateral lobes ("palps") strongly prolonged and rectangular inclined medially, in the groundplan with a serrated medio-distal margin (GEIJSKES, 1970); 10th abdominal segment of larvae more or less elongated.
• Taxonomy:
  
  • Gomphoidinae JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • Gomphoidiinae sensu TILLYARD & FRASER, 1940 (unjust. emend.)
    
  • Gomphoidinae sensu FRASER, 1957
    
  • Gomphoidini sensu CARLE, 1986 (stat. nov.)
    
  • Gomphoidinae; BELLE, 1996 (sens. nov.)

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Aphyllini BELLE, 1996

(Type genus: Aphylla SELYS, 1854.)

• Included taxa: including the genera listed in BELLE (1996).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Aphyllini BELLE, 1966 (trib. nov.)

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Diaphlebiini BELLE, 1996

(Type genus: Diaphlebia SELYS, 1854.)

• Included taxa: including the genera listed in BELLE (1996).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Diaphlebiini BELLE, 1966 (trib. nov.)

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Gomphoidini JACOBSON & BIANCHI, 1905

(Type genus: Gomphoides SELYS, 1854, nec Gomphoides SELYS, 1850; the latter is maybe an unavailable name that should otherwise be surpressed by the ICZN.)

• Included taxa: including the genera listed in BELLE (1996).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Gomphoidinae JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • Gomphoidini; CARLE, 1986 (stat. nov.)
    
  • Gomphoidini sensu BELLE, 1996 (sens. nov.)

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Peruviogomphini BELLE, 1996

(Type genus: Peruviogomphus KLOTS, 1944.)

• Included taxa: including the genera listed in BELLE (1996).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Peruviogomphini BELLE, 1966 (trib. nov.)

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Lindeniinae JACOBSON & BIANCHI, 1905

(Type genus: Lindenia DE HAAN, 1826.)

• Included taxa: including the genera listed in BRIDGES (1994) under Lindeniini.
• Autapomorphies:
  
  • Wing venation: secondary branch of IR2 very distinct and more parallel to it, therefore IR2 appears to be dichotomously forked distal of the lestine oblique vein, unlike Cordulegastrida (BELLE, 1979; BECHLY, 1994); hindwing discoidal triangle longitudinal elongate with a strongly sigmoidal and angulated distal side, caused by the development of a more distinct supplementary sector (trigonal planate) in the postdiscoidal area (convergent to Hagenioidea); discoidal triangles divided in more than two cells in both pairs of wings (convergent to a few Progomphidae and some Gomphoidini); at least in forewings the basal part of the subdiscoidal cell (between CuP-crossing and pseudo-anal vein PsA) is traversed by supplementary cubito-anal-crossveins (CARLE, 1986); pseudo-anal vein PsA is secondarily less distinct in the hindwing, correlated with the elongated hindwing discoidal triangle (convergent to Hagenioidea); forewing subdiscoidal triangle divided into three cells (not yet in the most basal genera (Lindenia, Ictinogomphus, and Sinictinogomphus); hypertriangle divided by at least two or more crossveins (except in the very basal genus Sinictinogomphus; convergent to the gomphoidine genus Phyllogomphoides).
    
  • Other characters: all larval tarsi, incl. metatarsi, with only two tarsomeres (CHAO, 1990; CARLE, 1992; convergent to Archaeogomphus); male auricles dorso-ventrally compressed and produced posteriorly (CARLE, 1986); larval anal pyramid further shortened (convergent to Gomphida).
• Taxonomy:
  
  • [légion Lindenia SELYS, 1854 (not a valid family-group taxon according Art.11f IRZN)]
    
  • Lindeniinae JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • [Ictinini TILLYARD, 1917 (objectively invalid trib. nov. that is based on a homonym type genus)]
    
  • [Ictininae FRASER, 1934 (stat. nov.) (objectively invalid name that is based on a homonym type genus)]
    
  • Ictinogomphinae TILLYARD & FRASER, 1940 (subfam. nov. with the type genus Ictinogomphus COWLEY, 1934 nom. subst. pro Ictinus RAMBUR, 1842, which is a junior homonym) (jun. subj. syn.)
    
  • Lindeniinae; DAVIES, 1981
    
  • [Division Lindenia CARLE, 1986]
    
  • Lindeniinae; CARLE, 1986 (sens. nov.)
    
  • Lindeniini; CARLE, 1986 (stat. nov.)
    
  • Ictinogomphinae; BELLE, 1996 (sens. nov.)
    
  • Ictinogomphini; BELLE, 1996 (stat. nov.)

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Oligophlebiata BECHLY, 2003e

• Included taxa: Hagenioidea and Brevicubitalia.
• Autapomorphies:
  
  • Wing venation: in hindwings less than 5 antefurcal (= postmedian) crossveins are retained between the sectors of arculus (BECHLY, 1996a).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Oligophlebiata BECHLY, 2003e (taxon nov.)

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Hagenioidea TILLYARD & FRASER, 1940

(Type genus: Hagenius SELYS, 1854.)

• Included taxa: † Proterogomphidae and Hageniidae.
• Autapomorphies:
  
  • Wing venation: branching of RP at midfork symmetrical (FRASER, 1957; convergent to Eugomphida); distal side (MAb) of triangle strongly angulate, correlated with the development of a more distinct supplementary sector (trigonal planate) in the postdiscoidal area (convergent to the hindwing of Lindeniinae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • [partim: Eugomphida sensu BECHLY, 1996a (taxon nov.)]
    
  • Hagenioidea sensu BECHLY et al., 1998 (stat. nov.) (nom. transl. ex Hageniinae TILLYARD & FRASER, 1940)

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† Proterogomphidae BECHLY et al., 1998

(Type genus: † Proterogomphus BECHLY et al., 1998)

• Included taxa: † Cordulagomphinae and † Proterogomphinae.
• Autapomorphies:
  
  • Wing venation: discoidal triangles secondarily unicellular (convergent to † Araripegomphidae and Gomphida); only two cells beneath the pterostigmata; vein pseudo-IR1 very distinct and originating on RP1 beneath the distal side of the pterostigma; anal loop longer than wide and with only one or two cells (convergent to many Gomphida); forewings with a large elongated cell beneath the submedian area.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • [Proterogomphidae sensu BECHLY, 1996a (nomen nudum)]
    
  • Proterogomphidae BECHLY et al., 1998 (fam. nov.)

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† Cordulagomphinae CARLE & WIGHTON, 1990

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

• Included taxa: including the genus † Cordulagomphus CARLE &WIGHTON, 1990 (incl. † Procordulagomphus NEL & ESCUILLIÉ, 1994) and a new undescribed genus (BECHLY, 1998d).
• Autapomorphies:
  
  • Wing venation: the most distal postmedian (antefurcal) crossvein and the most basal postnodal crossvein are very oblique ("postnodal oblique vein" sensu BECHLY, 1994; convergent to Eurypalpida, and reversed within † Procordulagomphus); only two antefurcal crossveins in both pairs of wings (convergent to Gomphidae); hindwing CuAa shortened and with strongly reduced branching (convergent to † Nannogomphus, except in the most basal (still undescribed) genus (BECHLY, 1998d); anal loop more distinctly longitudinal elongate; presence of a "cordulegastrid gap" (sensu BECHLY, 1996) of crossveins in the distal part of the antesubnodal area (convergent to † Araripegomphidae, Gomphaeschnidae, and Cavilabiata), except in the most basal (still undescribed) genus (BECHLY, 1998d); small size with a wing length of less than 25 mm (correlated with a rather open wing venation), except in the most basal (still undescribed) genus (BECHLY, 1998d); postdiscoidal intercalary vein secondarily less well defined, except in the most basal (still undescribed) genus (BECHLY, 1998d).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • [partim: Neogomphida sensu BECHLY, 1996a (taxon nov.)]
    
  • Cordulagomphinae CARLE & WIGHTON, 1990 (subfam. nov.)
    
  • Cordulagomphidae sensu BECHLY, 1996a (stat. et sens. nov.)
    
  • Cordulagomphinae sensu BECHLY, 1998d (stat. rest.)

Comment: the close relationship of † Cordulagomphinae with † Proterogomphus was only recently revealed by the discovery of a still undescribed new genus of † Cordulagomphinae from the Santana Formation. † Cordulagomphus santanensis CARLE & WIGHTON, 1990 obviously is not a fossil gomphid larva at all but an adult earwig (!) and therefore has to be excluded from Odonata.

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† Proterogomphinae BECHLY et al., 1998

(Type genus: † Proterogomphus BECHLY et al., 1998.)

• Included taxa: only including the closely related species † Proterogomphus krauseorum BECHLY et al., 1998 and † Proterogomphus renateae BECHLY et al., 1998. The former was previously regarded as the adult of † Sona nectes PRITYKINA, 1986, which is based on a holotypical larva that is probably unrelated and of uncertain relationship. An attribution of the latter larva to † Aeschnidiidae was suspected by BECHLY (1996, 1997) and BECHLY et al. (1998), but conflicting evidence was recently presented by FLECK & NEL & BECHLY & ESCUILLIÉ (2002).
• Autapomorphies:
  
  • Wing venation: anal loop only unicellular (convergent to many other Gomphides); hindwing discoidal triangles distinctly longitudinal elongate (convergent to Hageniidae and Lindeniinae), correlated with a somewhat less distinct pseudo-anal vein PsA.
    
  • Other characters: adult males with unifurcate epiproct (very uncertain character since based on an isolated abdomen fragment which was assigned to the referring fossil wings by PRITYKINA (1986) without any evidence.).
• Taxonomy:
  
  • Proterogomphidae BECHLY et al., 1998 (fam. nov.)
    
  • Proterogomphinae sensu BECHLY, 1998d (stat. nov.)

Comment: new family-group taxon, and a new genus and species, for the adult specimens that have been formerly classified in † Sonidae sensu PRITYKINA, 1986. Also including a new species from the Upper Jurassic Solnhofen Limestone of Germany that has been described by BECHLY et al. (1998).

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Hageniidae TILLYARD & FRASER, 1940

(Type genus: Hagenius SELYS, 1854.)

• Included taxa: according to BECHLY et al. (1998), this family includes the genera Hagenius SELYS, 1854, Sieboldius SELYS, 1854, and † Mesoictinus NEL et al., in prep.
• Autapomorphies:
  
  • Wing venation: discoidal triangles distinctly longitudinal elongate in both pairs of wings, correlated with secondarily less distinct pseudo-anal veins PsA and subdiscoidal triangles; distal side (MAb) of triangle strongly sigmoidal, correlated with the further strengthening of the supplementary sector (trigonal planate) in the postdiscoidal area (CARLE, 1986; convergent to the hind of Lindeniinae); MP and CuAa strongly divergent towards the wing margin in hindwings (CARLE, 1986; convergent to Octogomphinae); IR1 is secondarily elongated in both wings (convergent to Petaluridae, Austropetaliida, Petaliaeschna and Neopetaliidae); costal margin and RA not widened along pterostigmata (CARLE, 1986; reversal).
    
  • Other characters: occiput secondarily not ridged (convergent to Epigomphidae, Octogomphinae, and Austrogomphinae); labrum about 3 times as wide as long (CARLE, 1986); basi-lateral location of male cercal spine (CARLE, 1986; convergent to Hemigomphini); very large body size; basi-ventral surface of hind femora with overlapping flat-appressed spines (CARLE, 1986); anterior lamina raised with medial seam (CARLE, 1986); female sternum 9 membranous (convergent to Zonophoridae and Octogomphinae); length of middle maxillary tooth of posterior tooth row 60-70 % of proximal tooth (convergent to Octogomphinae, Anisogomphini and some Gomphini).
• Taxonomy:
  
  • [Hagenia series LAIDLAW, 1930 (not available as family-group taxon according to Art. 11f IRZN)]
    
  • Hageninae TILLYARD & FRASER, 1940 (subfam. nov.) (nom. imperf.)
    
  • Hageninae sensu FRASER, 1957
    
  • Hageninae sensu BELLE, 1979
    
  • Hageniinae sensu DAVIES, 1981 (incorr. subseq. spell., since not demonstrably intentional according to Art. 33(b)(i) IRZN)
    
  • [Division Hagenius CARLE, 1986 (informal name)]
    
  • Hageniinae sensu CARLE, 1986
    
  • Hageniini sensu CARLE, 1986 (stat. nov.)
    
  • Hageniidae sensu BECHLY, 1996a (stat. nov.)
    
  • Hageniidae sensu BECHLY et al., 1988 (nom. correct.)

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Brevicubitalia BECHLY, 2003e

• Included taxa: Zonophoridae and Gomphida.
• Autapomorphies:
  
  • Wing venation: CuAa distinctly shortened and with a distinct course (distally converging with the wing margin, instead of running parallel to it), and with a reduced pektinate branching.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Pangomphida BECHLY, 1996 Internet website
    
  • Brevicubitalia BECHLY, 2003e (taxon nov.)

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Zonophoridae BELLE, 1983

(Type genus: Zonophora SELYS, 1854.)

• Included taxa: including the genera listed in BRIDGES (1994) under Zonophorini.
• Autapomorphies:
  
  • Wing venation: hindwing discoidal triangle somewhat longitudinal elongate (?).
    
  • Other characters: female sternum 9 membranous (CARLE, 1986; convergent to some Gomphida); male tibial laminae (?) vestigial, at most 1/10 length of tibia (CARLE, 1986; convergent to several Gomphida); lateral lobes ("palps") of larval prehensile mask with a strong somewhat elongated and curved endhook and a serrated inner margin (dissimilar and probably convergent to Gomphoidini); unique larval gizzard with two dissimilar pairs of lobes with large and minute teeth (BELLE, 1983).
• Taxonomy:
  
  • Zonophorinae BELLE, 1983 (subfam. nov.)
    
  • Zonophorini sensu CARLE, 1986 (stat. nov.)
    
  • Zonophorinae sensu BELLE, 1996 (sens. nov.)
    
  • Zonophorini sensu BELLE, 1996 (sens. nov.)
    
  • Zonophoridae sensu BECHLY, 1996a (stat. nov.)

---

Gomphida BECHLY, 1996

• Included taxa: Epigomphidae and Gomphidae.
• Autapomorphies:
  
  • Wing venation: discoidal triangles secondarily unicellular in both pairs of wings (convergent to † Araripegomphidae and † Proterogomphidae; reversed in a few species); anal loop reduced to only one ot two cells or even completely suppressed (convergent to † Araripegomphidae, Progomphidae, and † Proterogomphidae; reversed in Onychogomphini).
    
  • Other characters: location of the male cercal spine not basi-ventral (CARLE, 1986; reversed in Octogomphini); larval anal pyramid further shortened (convergent to Lindeniinae); female auricles well-developed.
• Taxonomy:
  
  • Gomphida BECHLY, 1996a (taxon nov.)

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Epigomphidae FRASER, 1934

(Type genus: Epigomphus SELYS, 1854.)

• Included taxa: Epigomphinae and Archaeogomphinae.
• Autapomorphies:
  
  • Wing venation: pterostigmal brace reduced or obsolete (CARLE, 1986; convergent to Hagenoides).
    
  • Other characters: occiput secondarily not ridged (convergent to Hageniidae, Octogomphinae, and Austrogomphinae).
• Taxonomy:
  
  • Epigomphidae sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Epigomphinae FRASER, 1934)

Comment: this proposed preliminary classification of Epigomphidae is mainly based on the work of CARLE (1986), but probably does not yet represent a truely phylogenetic systematisation.

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Epigomphinae FRASER, 1934

(Type genus: Epigomphus SELYS, 1854.)

• Included taxa: Epigomphini, Macrogomphini, and Microgomphini.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • [partim: Gomphus series LAIDLAW, 1922]
    
  • Epigomphinae FRASER, 1934 (subfam. nov.)
    
  • Epigomphinae; BELLE, 1996 (sens. nov.)
    
  • Epigomphinae sensu BECHLY, 1996a (sens. nov.)

---

Epigomphini FRASER, 1934

(Type genus: Epigomphus SELYS, 1854.)

• Included taxa: Epigomphina and Leptogomphina.
• Autapomorphies:
  
  • Wing venation: pterostigmal brace obsolete (convergent to some Macrogomphini ?).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Epigomphini; CARLE, 1986 (stat. nov.) (nom. transl. ex Epigomphinae FRASER, 1934)
    
  • Epigomphini; BELLE, 1996 (sens. nov.)
    
  • Epigomphini sensu BECHLY, 1996a (sens. nov.)

---

Epigomphina FRASER, 1934

(Type genus: Epigomphus SELYS, 1854.)

• Included taxa: including the genera EpigomphusSELYS, 1854 and Eugomphus KENNEDY, 1947.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Epigomphini sensu CARLE, 1986 (stat. nov.)
    
  • Epigomphina sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Epigomphinae FRASER, 1934)

---

Leptogomphina CARLE, 1986

(Type genus: Leptogomphus SELYS, 1878.)

• Included taxa: including the genera Heliogomphus LAIDLAW, 1922 and Leptogomphus SELYS, 1878.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Leptogomphini CARLE, 1986 (trib. nov.)
    
  • Leptogomphina sensu BECHLY, 1996a (stat. nov.)

---

Macrogomphini CARLE, 1986

(Type genus: Macrogomphus SELYS, 1857.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Macrogomphini CARLE, 1986 (trib. nov.)

---

Microgomphini CARLE, 1986

(Type genus: Microgomphus SELYS, 1857.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: medial ocellus smaller than lateral ocelli.
• Taxonomy:
  
  • Microgomphini CARLE, 1986 (trib. nov.)

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Archaeogomphinae CARLE, 1986

(Type genus: Archaeogomphus WILLIAMSON, 1919, nec Archaeogomphus LIN, 1976.)

• Included taxa: Archaeogomphini, Agriogomphini, Cyanogomphini, and Lestinogomphini.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: middle maxillary tooth of posterior tooth row absent.
• Taxonomy:
  
  • Archaeogomphinae sensu BECHLY, 1996a (stat. nov.) (nom.transl. ex Archaeogomphini CARLE, 1986)

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Archaeogomphini CARLE, 1986

(Type genus: Archaeogomphus WILLIAMSON, 1919, nec Archaeogomphus LIN, 1976.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Archaeogomphini CARLE, 1986 (trib. nov.)
    
  • Archaeogomphini; BELLE, 1996 (sens. nov.)

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Agriogomphini BELLE, 1996

(Type genus: Agriogomphus SELYS, 1869.)

• Included taxa: including the genera listed in BELLE (1996).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Agriogomphini BELLE, 1966 (trib. nov.)

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Cyanogomphini CARLE, 1986

(Type genus: Cyanogomphus SELYS, 1873.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Cyanogomphini CARLE, 1986 (trib. nov.)
    
  • Cyanogomphini; BELLE, 1996 (sens. nov.)

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Lestinogomphini CARLE, 1986

(Type genus: Lestinogomphus MARTIN, 1911.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Lestinogomphini CARLE, 1986 (trib. nov.)

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Gomphidae RAMBUR, 1842

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

• Included taxa: Austrogomphinae, Phyllogomphinae, Onychogomphinae, Gomphinae, and Octogomphinae.
• Autapomorphies:
  
  • Wing venation: branching of RP at midfork symmetrical (FRASER, 1957; convergent to Hagenioidea); lestine oblique vein less than three cells distal of the subnodus also in the forewings (convergent to † Proterogomphidae and a few Lindeniidae and Epigomphidae; reversed in a few species); in hindwings only 1 or 2 and in forewings only 1-4 antefurcal (= postmedian) crossveins retained between the sectors of arculus (BELLE, 1979) (convergent to † Cordulagomphinae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Gomphides RAMBUR, 1842
    
  • Gomphines SELYS & HAGEN, 1850
    
  • Gomphina; BRAUER, 1856
    
  • Gomphinae; SELYS, 1858
    
  • Gomphina; HAGEN, 1861
    
  • Gomphina; CABOT, 1872
    
  • Gomphidae; BANKS, 1892 (stat. nov.)
    
  • Gomphinae; NEEDHAM, 1897, 1903
    
  • Gomphinae; CALVERT, 1903
    
  • Gomphinae; JACOBSON & BIANCHI, 1905
    
  • Gomphina; HANDLIRSCH, 1906
    
  • Gomphinae; RIS, 1909
    
  • Gomphoidea; CARLE, 1986 (stat. nov.)
    
  • Gomphinidae ROSS & JARZEMBOWSKI in BENTON, 1993 (incorr. subseq. spell.)
    
  • [partim: Eugomphida sensu BECHLY, 1996a (taxon nov.)]
    
  • [partim: Neogomphida sensu BECHLY, 1996a (taxon nov.)]
    
  • Gomphoidea sensu BECHLY, 1996a
    
  • Gomphidae sensu BECHLY, 2003e (sens. nov.)

Comment: the new higher taxa Eugomphida and Neogomphida, suggested by BECHLY (1996), are here regarded as non-monophyletic, but the names might be considered as available for a future phylogenetic systematisation within the present group, just like Gomphoidea, while the name Gomphidae could then be used in a more restrictive sense.

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Austrogomphinae CARLE, 1986

(Type genus: Austrogomphus SELYS, 1854.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Austrogomphinae CARLE, 1986 (subfam. nov.)
    
  • Austrogomphini CARLE, 1986 (trib. nov.)
    
  • Austrogomphidae sensu BECHLY, 1996a (stat. nov.)
    
  • Austrogomphinae sensu BECHLY, 2003e (stat. rest.)

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Phyllogomphinae CARLE, 1986

(Type genus: Phylogomphus SELYS, 1854.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Phyllogomphinae CARLE, 1986 (subfam. nov.)
    
  • Phyllogomphini CARLE, 1986 (trib. nov.)

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Onychogomphinae CHAO, 1984

(Type genus: Onychogomphus SELYS, 1854.)

• Included taxa: Crenigomphini and Onychogomphini.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: adult males with the two branches of the epiproct less diverging or even closely parallel (convergent to Desmogomphus).
• Taxonomy:
  
  • Onychogomphinae CHAO, 1984 (subfam. nov.)
    
  • Onychogomphinae; DAVIES & TOBIN, 1985
    
  • [Ophiogomphinae CARLE, 1986 (nomen nudum)]
    
  • Onychogomphinae; BELLE, 1996 (sens. nov.)

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Crenigomphini CARLE, 1986

(Type genus: Crenigomphus SELYS, 1892.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Crenigomphini CARLE, 1986 (trib. nov.)

---

Onychogomphini CHAO, 1984

(Type genus: Onychogomphus SELYS, 1854.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: secondarily enlarged anal loop with 1-3 cells along the anal vein; male hindwings with the anal triangle divided into 4 cells.
    
  • Other characters: male epiproct with arms being close and parallel (CHAO, 1984).
• Taxonomy:
  
  • Onychogomphini CARLE, 1986 (stat. nov.) (nom. transl. ex Onychogomphinae CHAO, 1984)
    
  • Onychogomphini; BELLE, 1996 (sens. nov.)

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Gomphinae RAMBUR, 1842

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

• Included taxa: Anisogomphini, Cyclogomphini, Gomphini, and Neurogomphini.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Gomphides RAMBUR, 1842
    
  • Gomphines SELYS & HAGEN, 1850
    
  • Gomphina; BRAUER, 1856
    
  • Gomphinae; SELYS, 1858
    
  • Gomphina; HAGEN, 1861
    
  • Gomphina; CABOT, 1872
    
  • Gomphinae; NEEDHAM, 1897, 1903
    
  • Gomphinae; CALVERT, 1903
    
  • Gomphinae; JACOBSON & BIANCHI, 1905
    
  • Gomphina; HANDLIRSCH, 1906
    
  • Gomphinae; RIS, 1909
    
  • Gomphinae; FRASER, 1934
    
  • Gomphinae; DAVIES & TOBIN, 1985 (sens. nov.)

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Anisogomphini CARLE, 1986

(Type genus: Anisogomphus SELYS, 1857.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Anisogomphini CARLE, 1986

---

Cyclogomphini CARLE, 1986

(Type genus: Cyclogomphus SELYS, 1854.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
• Wing venation: not yet known.
  
• Other characters: not yet known.

• Taxonomy:
  
  • Cyclogomphini CARLE, 1986 (trib.nov.)

---

Gomphini RAMBUR, 1842

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

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Gomphini; TILLYARD, 1917 (stat. nov.) (nom. transl. ex Gomphides RAMBUR, 1842)
    
  • Gomphini; DAVIES & TOBIN, 1985
    
  • Gomphini; CARLE, 1986 (sens. nov.)

---

Neurogomphini CARLE, 1986

(Type genus: Neurogomphus KARSCH, 1890.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Neurogomphini CARLE, 1986 (trib. nov.)

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Octogomphinae CARLE & COOK, 1984

(Type genus: Octogomphus SELYS, 1873.)

• Included taxa: Hemigomphini, Anomalophlebiini, Octogomphini, and Trigomphini.
• Autapomorphies:
  
  • Wing venation: distal primary antenodal ax2 of hindwing midway between basal primary antenodal ax1 and nodus (except in Hemigomphus); MP and CuAa strongly divergent towards wing margin in hindwings (convergent to Hageniidae).
    
  • Other characters: 2-5 ventral teeth on the larval ligula (CARLE & COOK, 1984); female sternum 9 membranous (convergent to Zonophoridae and Hageniidae); length of middle maxillary tooth of posterior tooth row 60-70 % of proximal tooth (convergent to Hageniidae, Anisogomphini and some Gomphini).
• Taxonomy:
  
  • Octogomphinae; DAVIES & TOBIN, 1985 (stat. nov.) (nom. transl. ex Octogomphini CARLE & COOK, 1984)
    
  • Octogomphinae; BELLE, 1996 (sens. nov.)

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Hemigomphini CARLE, 1986

(Type genus: Hemigomphus SELYS, 1854.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Hemigomphini CARLE, 1986 (trib. nov.)
    
  • Hemigomphini; BELLE, 1996 (sens. nov.)

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Anomalophlebiini BELLE, 1996

(Type genus: Anomalophlebia BELLE, 1995.)

• Included taxa: including the genera listed in BELLE (1996).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Anomalophlebiini BELLE, 1966 (trib. nov.)

---

Octogomphini CARLE & COOK, 1984

(Type genus: Octogomphus SELYS, 1873.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Octogomphini CARLE & COOK, 1984 (trib. nov.)
    
  • Octogomphini; DAVIES & TOBIN, 1985

---

Trigomphini CARLE, 1986

(Type genus: Trigomphus BARTENEV, 1911.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Trigomphini CARLE, 1986 (trib. nov.)