Phylogenetic Systematics of Odonata

Zygoptera SELYS, 1854

• Included taxa: Caloptera and Euzygoptera, † Steleopteridae and † Eosagrionidae.
• Autapomorphies:
  
  • Wing venation: both pairs of wings are distinctly stalked, with a petiolus that is at least somewhat longer than broad (the total absence of a petiole in Calopterygidae and some Epallagidae clearly is secondary since the closely related Caliphaeinae, Heliocharitidae do possess a distinct petiole, like most other Calopterygoidea, too; even most Epallagidae have a very shortened but still visible petiole); both pairs of wings of identical shape and venation (in the groundplan of Odonata the hindwing was basally somewhat broader than the forewing, even so the greatly broadened hindwings of Anisoptera are a secondary formation; thus neither the identical wingshape of Zygoptera, nor the very different wingshape of Anisoptera represents the ancestral condition, both are alternative autapomorphies); maybe the reduction of the terminal kink of the CP at the nodus is also an autapomorphy of Zygoptera, correlated with a reduction of the nodal furrow since this character is very distinct in the most basal Panodonata († Tarsophlebiidae) and most Epiproctophora, incl. all extant Anisoptera, while it seems to be convergently reduced in Epiophlebia and † Aeschnidiidae; the tubular sclerotised canal of the ScP is obliterated on the venter of the postnodal costal margin.
    
  • Other characters: head capsule transversely very elongated and in the median part dorso-ventrally compressed (hammer-shaped), with very widely separated eyes; cephalic suture between frons and vertex suppressed (the suture between vertex and occiput is retained in basal Lestomorpha); raptorial spines also on femora very long and slender, like on the tibiae (apomorphic contra NEL et al., 1993; and contra LOHMANN, 1995, 1996); thorax and abdomen rather gracile; male paraprocts hypertrophied and functioning as claspers (appendices inferiores); presence of a distinct pattern of movable junctions between crossveins and the concave longitudinal veins, with specialised spines that limit the angle of movability; reduction of the long spines on the dorsal surface of the RP and MP (reversed in some Chlorocyphoidea and Heliocharitidae); a significant increase of spine-density at the apical costal margin, while the Epiophlebiidae and Anisoptera do only have a slight increase or even a decrease in this region; presence of tiny macrotrichia on the ventral surface of the ScP and the dorsal surface of the RA (secondarily absent in higher Coenagrionomorpha); significant lateral compression of the crossveins at least in postero-distal wing area, while these veins always have a more or less round cross section in the other Odonata and also in the Ephemerida (outgroup); on the ventral wing surface the posterior part of the basal brace (ax0) is obliterated or covered by a rather extensive sclerotisation of the wing base; a very extreme obliquity (skewness) of the pterothorax which is much more pronounced than in Epiophlebiidae and Anisoptera (this feature leads to a functional analogy of neoptery in those Zygoptera that fold their wings above the back in rest); a very long and slender abdomen, compared to the relatively small thorax; a more or less triangular cross-section of the abdominal sternites which have a distinct longitudinal keel (convergent to Libelluloidea) while the processus caudalis is totally reduced; presence of an anterior median depression, a true lamina batilliformes, and a "Zipfel" at the male secondary genital apparatus (LINDEBOOM, 1996); enlargement and subdivision of the ligula which serves as "functional penes analogue" in Zygoptera, and which is supplied with sophisticated structures for sperm-removal (CARLE, 1995); formation of an ovipositor-pouch by the enlarged outer valves (valvula 3) of the 9th abdominal sternite (LINDEBOOM, 1996); females do grab their own abdomen instead of the male abdomen with their legs during the formation of the pairing-wheel (maybe the plesiomorphic state, relative to the behaviour in Anisoptera and convergent in Hemiphlebia); fore legs held like the other two leg pairs in flight ("orthoflexate" position sensu LOHMANN, 1995, 1996; contra Lohmann only a difference to Epiproctophora, but a character with uncertain polarity); myofibrils of the flight musculature always occurring in pairs and presence of a second tergopleural muscle that serves for folding the wings into resting position; the side-lobes ("labial palps") of the larval prehensile mask are apically fissured by at least one or two indentions (HEIDEMANN & SEIDENBUSCH, 1993; BECHLY, 1995); side lobes of larval prehensile mask tilted, not in the same plane as the median lobe (HEIDEMANN & SEIDENBUSCH, 1993; LOHMANN, 1996).
• Taxonomy:
  
  • [Agrion sensu FABRICIUS, 1775]
    
  • Agrionida LEACH, 1815
    
  • [Agrionidae sensu STEPHENS, 1836]
    
  • Agrionides WESTWOOD, 1840 (taxon nov.)
    
  • Agrionina SELYS, 1840 (taxon nov.)
    
  • Zygoptera SELYS, 1834 or 1840 ?
    
  • Zygoptera SELYS in SELYS & HAGEN, 1854
    
  • [Agrionidae sensu KIRBY, 1890]
    
  • Calopterygidea KARSCH, 1894 (taxon nov.)
    
  • Zygopterides LUCAS, 1900 (taxon nov.)
    
  • [Agrionidos NAVAS, 1903 (informal name)]
    
  • Agrionodea JACOBSON & BIANCHI, 1905 (taxon nov.)
    
  • Lestomorpha PRITYKINA, 1980 (taxon nov.) (nec Lestomorpha BECHLY, 1996)

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† Steleopteridae HANDLIRSCH, 1906

(Type genus: † Steleopteron HANDLIRSCH, 1906.)

• Included taxa: including the genera † Steleopteron HANDLIRSCH, 1906, † Auliella PRITYKINA, 1968, † Parasteleopteron FLECK & NEL & BECHLY & MARTÍNEZ-DELCLÒS, 2001 and † Euparasteleopteron FLECK & NEL & BECHLY & MARTÍNEZ-DELCLÒS, 2001, but not † Pseudophaea HANDLIRSCH, 1906 which is a nomen dubium in Odonata incertae sedis and † Euphaeopsis HANDLIRSCH, 1906 which belongs to † Isophlebioidea within Epiproctophora.
• Autapomorphies:
  
  • Wing venation: wings very elongate and narrow, with a very long petiolus (convergent to † Triadophlebiomorpha, † Protozygoptera, Zygoptera, and † Parazygoptera); discoidal cell very elongated and basally closed in both pairs of wings (convergent to Caloptera, Lestida, Coenagrionomorpha, † Isophlebiidae, and Euepiproctophora); characteristical pattern of transverse pleats between the longitudinal veins; all longitudinal wing veins are very long, terminating near the apex; MA and CuA are distally zigzagged (convergent to many † Parazygoptera, such as † Asiopteridae, † Mesophlebiinae and † Italophlebia); IR1 very reduced or strongly zigzagged; RP2 not strictly aligned with subnodus (more probably a plesiomorphy); MP distinctly curved after its origin at the distal angle of the discoidal cell (convergent to † Triadophlebiomorpha and Eulestiformia).
    
  • Other characters: abdomen much longer than the wings; strongly hypertrophied female ovipositor.
• Taxonomy:
  
  • Steleopteridae HANDLIRSCH, 1906 (fam. nov.)
    
  • Steleopteridae sensu BECHLY, 1996a (sens. nov.)
  • Steleopteridae sensu FLECK & NEL & BECHLY & MARTÍNEZ-DELCLÒS, 2001 (sens. nov.)

Comment: The distinct petiolation and similar shape of both pairs of wings, the transverse head (only doubtfully known from † Steleopteron), and the increased skewness of the pterothorax all suggest that † Steleopteridae belongs to Zygoptera rather than to Epiproctophora as could be suggested by the robust body, the basal position of the midfork between nodus and arculus, and the presence of transverse furrows on the pterostigmata. The plesiomorphic absence of a longitudinal carina on the abdominal sterna could suggest a position in the stemgroup rather than the crowngroup of Zygoptera, especially because there are no convincing synapomorphies with either Caloptera or Euzygoptera.

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† Eosagrionidae HANDLIRSCH, 1920

(Type genus: Eosagrion HANDLIRSCH, 1920.)

• Included taxa: only including the species † Eosagrion risi HANDLIRSCH, 1920.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Eosagrionidae HANDLIRSCH, 1920 (fam. nov.)

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Caloptera BELYSHEV & HARITONOV, 1983

• Included taxa: † Sieblosiidae and Eucaloptera.
• Autapomorphies:
  
  • Wing venation: the midfork is recessed basally to a position between 12-26 % of wing length (convergent to Lestinoidea and Hypolestinae); pterostigmal brace vein obsolete; basal closure of discoidal cell in forewings including the development of a dorsal arcular bracket (convergent to Lestida, Coenagrionomorpha, Epiophlebiidae, † Isophlebiidae, and Anisopteromorpha).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Normopteroides SELYS, 1840 (taxon nov.)
    
  • Equinervulées SELYS, 1854 (taxon nov.)
    
  • Dyorthophlebiae BUCHECKER, 1876 (taxon nov.)
    
  • Calopterycoptera ZALESSKY, 1934 (taxon nov.)
    
  • [Agrionidea sensu ??]
    
  • [Calopterygoidea sensu FRASER, 1957]
    
  • Caloptera BELYSHEV & HARITONOV, 1983 (taxon nov.)

Comment: the reduction of the posterior metathoracic tergal sclerite (PFAU., 1986, 1991), the enlargement of the anterior metathoracic tergal sclerite (PFAU, 1986, 1991), the suppression of the lestine oblique vein, and the suppression of the cephalic suture between vertex and occiput could be potential synapomorphies of Eucaloptera and Coenagrionomorpha which could then be united in a new taxon Agrionoptera, with the Lestoptera (= Lestomorpha) as sistergroup.

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† Sieblosiidae HANDLIRSCH, 1907

(Type genus: † Sieblosia HANDLIRSCH, 1907; = † Stenolestes SCUDDER, 1895, sen. subj. syn.)

• Included taxa: including the genera † Stenolestes SCUDDER, † Oligolestes SCHMIDT, 1958, † Parastenolestes NEL & PAICHELER, 1994, and † Paraoligolestes NEL & ESCUILLIÉ, 1993.
• Autapomorphies:
  
  • Wing venation: nodal furrow further reduced; nodal and subnodal veinlets less oblique, transverse or even with reversed obliquity; pterostigmata very elongated (but unknown in † Petrolestes); highly specialised nodus (plesiomorphic absent in the most basal genus † Oligolestes) which seems to be traversed by the ScP because the terminal kink of CP is shifted basally together with the nodal and subnodal veinlets and the nodal membrane sclerotisation is reduced.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Sieblosiidae HANDLIRSCH, 1907 (fam. nov.)
    
  • Sieblosiidae; FISCHER, 1974 (sens. nov.)
    
  • Sieblosiidae; NEL, 1986 (sens. nov.)
    
  • Sieblosiidae; NEL & PAICHELER, 1994 (stat. rest. et sens. nov.)

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Eucaloptera BECHLY, 1996

• Included taxa: Amphipterygida and Calopterygomorpha.
• Autapomorphies:
  
  • Wing venation: tendency towards a more or less rectangular discoidal cell (reversed in Amphipterygidae and many Chlorocyphoidea ?); very oblique basal margin of the pterostigma which is developed as an apparent secondary branching of RA; lestine oblique vein reduced (convergent to Hemiphlebiidae, Coenagrionomorpha and many † Parazygoptera); strong tendency towards coloured wings (convergent to Eurypalpida).
    
  • Other characters: larval caudal gills are always saccoid or triquetral (convergent to Hypolestidae and Palaemnematinae); lateral lobes ("palps") of larval prehensile mask with three distinct endhooks (polarity unsafe, more probably a symplesiomorphy with Hemiphlebiidae, Perilestidae, Lestidae, and "Megapodagrionidae").
• Taxonomy:
  
  • Eucaloptera BECHLY, 1996a (taxon nov.)

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Amphipterygida BECHLY, 1996

• Included taxa: Diphlebiidae and Amphipterygoidea.
• Autapomorphies:
  
  • Wing venation: all secondary antenodal crossveins between ScP and RA distal of ax2 are suppressed (reversed in Philoganginae); antesubnodal space at least without crossveins in its basal half (the distal crossveins only retained in Amphipteryginae; convergent to Chlorocyphoidea).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Amphipterygida BECHLY, 1996a (taxon nov.)

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Diphlebiidae HEYMER, 1975

(Type genus: Diphlebia SELYS, 1869 nom. subst. pro Dineura SELYS, 1859 (= Dinevra incorr. orig. spell.), which is a junior homonym.)

• Included taxa: Philoganginae and Diphlebiinae.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: paraprocts of adult males more or less reduced, only the cerci functioning as male claspers; unique type of larvae (FRASER, 1938; NOVELO-GUTIERREZ, 1995; body and legs strongly flattened; ventral margin of eyes with rows of strong spines; mandibles flattened ventrally with strong, sharp spines; occiput strongly concave; prementum with strong marginal spines); adults large and robust, with a very strong thorax (FRASER, 1938); resting position with widespread wings that are strictly horizontally outstretched (FRASER, 1938; BECHLY, 1994; NOVELO-GUTIERREZ, 1995; convergent to Anisoptera); larvae adapted to a live in torrential mountain streams, clinging to the underside of stones (FRASER, 1938).
• Taxonomy:
  
  • Diphlebiidae HEYMER, 1975 (fam. nov.)
    
  • Diphlebiidae DAVIES & TOBIN, 1984 (jun. obj. syn. and homonym)
    
  • Diphlebiidae; NOVELO-GUTIERREZ, 1995 (sens. nov.)
    
  • Diphlebiidae sensu BECHLY, 1996a (sens. nov.)

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Philoganginae KENNEDY, 1920

(Type genus: Philoganga KIRBY, 1890 nom. subst. pro Anisoneura SELYS, 1859, which is a junior homonym.)

• Included taxa: only including the type genus Philoganga KIRBY, 1890.
• Autapomorphies:
  
  • Wing venation: wings very slender with parallel costal and hind margins; discoidal cell short; basal accessory antenodal crossveins present in the subcostal space between ax0 and ax1; base of IR2 widely separated from the midfork (convergent to † Zacallitidae and Epallaginae); presence of a characteristical oblique vein between RP1 and IR1.
    
  • Other characters: very large size; adult males with vestigial paraprocts.
• Taxonomy:
  
  • Philoganginae KENNEDY, 1920 (subfam. nov.)
    
  • Philoganginae sensu KENNEDY, 1925
    
  • Philoganginae sensu FRASER, 1957
    
  • Philoganginae sensu BECHLY, 1996a

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Diphlebiinae HEYMER, 1975

(Type genus: Diphlebia SELYS, 1869 nom. subst. pro Dineura SELYS, 1859 (= Dinevra incorr. orig. spell.), which is a junior homonym.)

• Included taxa: only including the type genus Diphlebia SELYS, 1869.
• Autapomorphies:
  
  • Wing venation: antesubnodal space without crossveins (rarely one basal crossvein present).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Diphlebiinae sensu BRIDGES, 1993 (stat. nov.) (nom. transl. ex Diphlebiidae HEYMER, 1975)
    
  • Diphlebiinae; NOVELO-GUTIERREZ, 1995 (sens. nov.)
    
  • Diplebiinae sensu BECHLY, 1996a

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Amphipterygoidea TILLYARD, 1917

(Type genus: Amphipteryx SELYS, 1853.)

• Included taxa: Thaumatoneuridae and Amphipterygidae, and probably also Pseudolestidae as taxon incertae sedis.
• Autapomorphies:
  
  • Wing venation: nodus in a very basal position (at distinctly less than 40 % of wing length); midfork further recessed, too.
    
  • Other characters: resting position with wings closely apposed over the dorsum of the body.
• Taxonomy:
  
  • Amphipterygoidea sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Amphipterygini TILLYARD, 1917)

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Pseudolestidae FRASER, 1957

(Type genus: Pseudolestes KIRBY, 1900.)

• Included taxa: only including the extant type species Pseudolestes mirabilis KIRBY, 1900 from the island Hainan, which could very well turn out to be a genuine Amphipterygidae, when the larva will be discovered.
• Autapomorphies:
  
  • Wing venation: male hindwings opaque metallic coloured and much shorter and broader than the totally differently shaped forewings (FRASER, 1957); the hindwing discoidal cell is strongly elongated; hindwing pterostigma recessed; longitudinal wing veins more strongly curved in the hindwing; only the two primary antenodal brackets ax1 and ax2 are retained in the antenodal space (convergent to Euzygoptera; maybe a synapomorphy with † Petrolestini or Rimanella); antesubnodal space without any crossveins (maybe a synapomorphy with Rimanellinae or Thaumatoneuridae); midfork further recessed, being very close to the arculus (maybe a synapomorphy with Amphipterygidae; convergent to many other Caloptera, Lestidae and Hypolestidae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Pseudolestinae FRASER, 1956 ? (subfam. nov.)
    
  • Pseudolestidae FRASER, 1957 (stat. nov.)
    
  • Pseudolestidae sensu BECHLY, 1996a (sens. nov.)

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Thaumatoneuridae TILLYARD & FRASER, 1938

(Type genus: Thaumatoneura MCLACHLAN, 1897.)

• Included taxa: † Dysagrioninae and Thaumatoneurinae.
• Autapomorphies:
  
  • Wing venation: the basal costal margin between wing base and nodus is distinctly convex curved; antesubnodal space without any crossveins (convergent to Diphlebiinae, Pseudolestidae, Rimanellinae and Euzygoptera).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Thaumatoneuridae sensu BECHLY, 1995 (stat. nov.) (nom. transl. ex Thaumatoneurinae TILLYARD & FRASER, 1938)
    
  • Thaumatoneuridae sensu BECHLY, 1996a (sens. nov.)

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† Dysagrioninae COCKERELL, 1908

(Type genus: † Dysagrion SCUDDER, 1878.)

• Included taxa: † Petrolestini and † Dysagrionini.
• Autapomorphies:
  
  • Wing venation: rather unique shape of the discoidal cell (only similarly developed in some † Sieblosiidae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Dysagrioninae COCKERELL, 1908 (subfam. nov.)
    
  • Dysagrioninae; NEL & PAICHELER, 1994
    
  • Dysagrioninae sensu BECHLY, 1996a (sens. nov.)

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† Petrolestini COCKERELL, 1927

(Type genus: † Petrolestes COCKERELL, 1927.)

• Included taxa: including the two genera † Petrolestes COCKERELL, 1927 and † Congqingia ZHANG, 1992 (by the way: the holotype of † Conqingia rhora ZHANG, 1992 was offered for sale by a Chinese trader at a fossil fair at Stuttgart and is now deposited at the GPIT in Tübingen, Germany).
• Autapomorphies:
  
  • Wing venation: only the two primary antenodal brackets ax1 and ax2 are retained in the antenodal space (convergent to Pseudolestidae and Euzygoptera); midfork further recessed, being situated midway between nodus and arculus; IR1 shortened and strongly zigzagged (unknown in † Petrolestes).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Petrolestinae COCKERELL, 1927 (subfam. nov.)
    
  • Petrolestinae; NEL, 1986 (sens. nov.)
    
  • Congqingiidae ZHANG, 1992 (fam. nov. with the type genus Congqingia ZHANG, 1992) (jun. subj. syn. in BECHLY, 2003e)
    
  • Petrolestinae; NEL & PAICHELER, 1994 (sens. nov.)
    
  • Petrolestini sensu BECHLY, 1996a (stat. et sens. nov.)
    
  • Congqingiini sensu BECHLY, 1996a (stat. et sens. nov.) (jun. subj. syn. in BECHLY, 2003e)

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† Dysagrionini COCKERELL, 1908

(Type genus: † Dysagrion SCUDDER, 1878.)

• Included taxa: including the two genera † Dysagrion SCUDDER, 1878 and † Phenacolestes COCKERELL, 1908.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Dysagrionini sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Dysagrioninae COCKERELL, 1908)

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Thaumatoneurinae TILLYARD & FRASER, 1938

(Type genus: Thaumatoneura MCLACHLAN, 1897.)

• Included taxa: † Euarchistigmatini and Thaumatoneurini.
• Autapomorphies:
  
  • Wing venation: nodus in an extremely basal position (subnodus between the bases of IR2 and RP3/4 that are strongly shifted basally, too), correlated with a large number of postnodal crossveins; IR2 apparently arising on RP3/4 (but still variable in † Euarchistigma); very dense wing venation with a high number of cells; longitudinal wing veins distally distinctly curved to the posterior wing margin; discoidal cell perfectly rectangular.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Thaumatoneurinae TILLYARD & FRASER, 1938 (subfam. nov.)
    
  • Thaumatoneurinae; FRASER, 1957 (sens. nov.)
    
  • Thaumatoneurinae; NEL & PAICHELER, 1994 (sens. nov.)
    
  • Thaumatoneurinae sensu BECHLY, 1996a (sens. nov.)

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† Euarchistigmatini CARLE & WIGHTON, 1990

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

• Included taxa: probably only including † Euarchistigma atrophium CARLE & WIGHTON, 1990, but certainly not † Cretarchistigma JARZEMBOWSKI, et al., 1998, which has to be regarded as Zygoptera incertae sedis (maybe Hemiphlebiidae according to BECHLY, 1998d).
• Autapomorphies:
  
  • Wing venation: small size of the wings (wing length less than 3 cm); all secondary antenodal crossveins suppressed (convergent to Pseudolestidae, † Petrolestini and Euzygoptera); only one row of cells between CuA and the wing margin; all intercalary veins (except IR1 and IR2) suppressed (maybe correlated with the small size); curious shape of pterostigmata.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Euarchistigmatinae CARLE & WIGHTON, 1990 (subfam. nov.)
    
  • Euarchistigmatini sensu BECHLY, 1996a (stat. nov.)

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Thaumatoneurini TILLYARD & FRASER, 1938

(Type genus: Thaumatoneura MCLACHLAN, 1897.)

• Included taxa: only including the type species Thaumatoneura inopinata MCLACHLAN, 1897.
• Autapomorphies:
  
  • Wing venation: large wing span; colour pattern of wings; characteristical triadic branching pattern of CuA (convergent to Megaloprepus); subdiscoidal cell traversed by a crossvein (FRASER, 1957); basal discoidal veinlet (= posterior part of arculus) is arising from [M & Cu] with a primary insertion; several rows of cells between RA and RP1 distal of the pterostigmata.
    
  • Other characters: adult males with reduced paraprocts; large size; larvae adapted to rapids and waterfalls.
• Taxonomy:
  
  • Thaumatoneurini sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Thaumatoneurinae TILLYARD & FRASER, 1938)

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Amphipterygidae TILLYARD, 1917

(Type genus: Amphipteryx SELYS, 1853.)

• Included taxa: Rimanellinae and Amphipteryginae.
• Autapomorphies:
  
  • Wing venation: secondarily (?) more oblique distal discoidal vein MAb, causing a more acute apex of the discoidal cell; midfork further recessed, being very close to the arculus (maybe a synapomorphy with Amphipterygidae; convergent to many other Caloptera, Lestidae and Hypolestidae).
    
  • Other characters: the plesiomorphic honeycomb microsculpture of the pterostigmata is obliterated and transformed into a crumpled microsculpture (further derived in one species, Devadatta podolestoides that has many small tubercles); larval gizzard with 32 proventricular folds, each carrying a row of minute denticles; unique type of larval caudal gills and accessory with a basal pair of retractible gill tufts (external rectal gills) that are developed from the laminae sub-anales (WATSON, 1966; LIEFTINCK, 1971; NOVELO-GUTIERREZ, 1995); adult males with four lobes on the terminal segment of the ligula (KENNEDY, 1919).
• Taxonomy:
  
  • [légion Amphipteryx SELYS, 1853 (not a valid family-group taxon according Art.11f IRZN)]
    
  • Amphipterygini TILLYARD, 1917 (trib. nov.)
    
  • Amphipteryginae; KENNEDY, 1920 (stat. nov.)
    
  • Amphipterygidae; TILLYARD, 1926 (stat. nov.)
    
  • Amphipterygidae; NOVELO-GUTIERREZ, 1995 (sens. nov.)
    
  • Amphipterygidae sensu BECHLY, 1996a (sens. nov.)

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Rimanellinae DAVIES & TOBIN, 1984

(Type genus: Rimanella NEEDHAM, 1934 nom. subst. pro Rima NEEDHAM, 1933, which is ajunior homonym.)

• Included taxa: including the two sister-genera Rimanella NEEDHAM, 1934 and Pentaphlebia FÖRSTER, 1909.
• Autapomorphies:
  
  • Wing venation: antesubnodal space without any crossveins (maybe synapomorphic with Pseudolestidae; convergent to Euzygoptera).
    
  • Other characters: larval head with sharp cephalic lobes (NOVELO-GUTIERREZ, 1995); larval prehensile mask with a subquadrate or subrectangular (NOVELO-GUTIERREZ, 1995); larval lateral caudal gills (paraprocts) long and triquetral with a trumpet-shaped base, and the median caudal gill (epiproct) very short and trifid (Fraser, 1955; NOVELO-GUTIERREZ, 1995).
• Taxonomy:
  
  • Rimanellidae DAVIES & TOBIN, 1984 (fam. nov.)
    
  • Pentaphlebiinae NOVELO-GUTIERREZ, 1995 (subfam. nov.) (jun. subj. syn.)
    
  • Rimanellinae sensu VAN TOL, 1995 (stat. nov.)
    
  • Rimanellinae sensu BECHLY, 1996a

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Amphipteryginae TILLYARD, 1917

(Type genus: Amphipteryx SELYS, 1853.)

• Included taxa: including the two sister-genera Amphipteryx SELYS, 1853 and Devadatta KIRBY, 1890.
• Autapomorphies:
  
  • Wing venation: basal side of pterostigmata extremely slanting, with several hyperstigmal crossveins between the oblique basal side of the pterostigma and the costal margin.
    
  • Other characters: surface of the pterostigmata with a unique longitudinal crumpling; larval cerci plate-like, closely apposed in the midline (NOVELO-GUTIERREZ, 1995); larvae without lamina supra-analis (NOVELO-GUTIERREZ, 1995); stem of the larval gill tufts issuing from a strap-like process and ending in a sclerotised plate (NOVELO-GUTIERREZ, 1995); larval sternite 10 subdivided (NOVELO-GUTIERREZ, 1995).
• Taxonomy:
  
  • [légion Amphipteryx SELYS, 1853 (not a valid family-group taxon according Art.11f IRZN)]
    
  • Amphipterygini TILLYARD, 1917 (trib. nov.)
    
  • Amphipteryginae; KENNEDY, 1920 (stat. nov.)
    
  • Amphipteryginae sensu NOVELO-GUTIERREZ, 1995 (sens. nov.)
    
  • Amphipteryginae sensu BECHLY, 1996a

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Calopterygomorpha BECHLY, 1996

• Included taxa: Chlorocyphoidea and Calopterygiformia.
• Autapomorphies:
  
  • Wing venation: antenodal field with very numerous antenodal crossveins that are very close together (distance less than 1,0 mm); kink of ScP at nodus very abrupt and Z-like (convergent to most Anisoptera); strong tendency towards a basal curving of RP1/2 which is arising on RP with a secondary insertion (only absent in some basal Chlorocyphoidea, the † Zacallitidae and all Polythoridae, but at least in the latter maybe secondarily so); strong tendency towards an elongation of the discoidal cell and its subdivision by crossveins (convergent to Devadatta and Anomisma; not present in † Zacallitidae, † Eodichrominae and some genera of extant Epallagidae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Calopterygomorpha BECHLY, 1996a (taxon nov.)

---

Chlorocyphoidea COWLEY, 1937

(Type genus: Chlorocypha FRASER, 1928 nom. subst. pro Libellago SELYS, 1953, nec Libellago SELYS, 1840.)

• Included taxa: Libellaginidae and Chlorocyphidae.
• Autapomorphies:
  
  • Wing venation: discoidal cell extremely elongated and always traversed by at least one crossvein; only one row of cells between MP and CuA (postsubdiscoidal space not distally widened) and between CuA and the hind margin (convergent to Rimanella, Heliocharitidae, Caliphaeidae and most Euzygoptera); basal part of the antesubnodal space free of crossveins ("chlorocyphoid gap"); RP3/4 waving and MA distinctly upward curved immediately after the discoidal cell (reversed in Disparocypha); CuA zigzagged (reversed in Aristocypha).
    
  • Other characters: compound eyes very large and placed more closely together than in other Zygoptera (FRASER, 1957), only slightly more separated than their width; postclypeus (epistoma) inflated and projected forward as a prominent snout (FRASER, 1957); thorax rather robust; abdomen distinctly shorter than wings (FRASER, 1957; except in Rhinoneura); microsculpture of the pterostigmata consisting of tiny pyramid-like structures (convergent to Epallagidae and Platystictidae); crossveins often supplied with macrotrichae; resting position with wings closely apposed over the dorsum of the body; all three larval caudal gills spike-like, the median "spike" (appendix dorsalis of the epiproct) distinctly shorter than the lateral "spikes" (paraprocts) (FRASER, 1957); adult males without hairs or spines on the ligula shaft (KENNEDY, 1919; convergent to Lestida and Platycnemididae).
• Taxonomy:
  
  • [Productinases SELYS, 1854 (taxon nov.)]
    
  • Chlorocyphoidea sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Chlorocyphidae COWLEY, 1937)

---

Libellaginidae JACOBSON & BIANCHI, 1905

(Type genus: Libellago SELYS, 1840, nec Libellago SELYS, 1953.)

• Included taxa: Libellagininae and Rhinocyphinae.
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: larval prehensile mask extremely elongate, with a deeply fissured prementum (median cleft hypertrophied; convergent to Calopterygidae), the edges of the fissures overlapping (FRASER, 1957; convergent to Calopterygidae); larval antenna eight-segmented with the penultimate segment strongly prolonged (FRASER, 1957); lateral lobes of the larval prehensile mask furnished with basal setae and distal hooks (FRASER, 1957).
• Taxonomy:
  
  • [légion Libellago SELYS, 1853 (not available as family-group taxon according Art.11f IRZN)]
    
  • Libellaginidae; LAIDLAW, 1934 (stat. nov. ?) (nom. transl. ex Libellagininae JACOBSON & BIANCHI, 1905)
    
  • Libellaginidae; LIEFTINCK, 1934 (stat. nov. ?) (nom. transl. ex Libellagininae JACOBSON & BIANCHI, 1905)
    
  • Libellaginidae sensu BECHLY, 1996a (sens. nov.)

---

Libellagininae JACOBSON & BIANCHI, 1905

(Type genus: Libellago SELYS, 1840, nec Libellago SELYS, 1953.)

• Included taxa: Sclerocyphini and Libellaginini.
• Autapomorphies:
  
  • Wing venation: sectors of arculus arising from the same point (FRASER, 1957); number of antenodal crossveins reduced to 5-9 (FRASER, 1957).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Libellagininae JACOBSON & BIANCHI, 1905 (subfam. nov.)
    
  • Libellaginae; LAIDLAW, 1917 (nom. correct.)
    
  • Libellaginae; LAIDLAW, 1920
    
  • Libellagininae sensu BECHLY, 1996a (sens. nov.)

---

Sclerocyphini BECHLY, 1996

(Type genus: Sclerocypha FRASER, 1949, stat. rest.)

• Included taxa: only including the type genus Sclerocypha FRASER, 1949.
• Autapomorphies:
  
  • Wing venation: costal margin conspicuously thickened at the nodus (FRASER, 1957).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Sclerocyphini BECHLY, 1996a (trib. nov., and stat. rest. for Sclerocypha FRASER, 1949)

---

Libellaginini JACOBSON & BIANCHI, 1905

(Type genus: Libellago SELYS, 1840, nec Libellago SELYS, 1953.)

• Included taxa: Melanocyphina and Libellaginina.
• Autapomorphies:
  
  • Wing venation: no antenodal crossveins between the primary antenodal brackets ax1 and ax2.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Libellagini; TILLYARD, 1917 (stat. nov.) (nom. transl. ex Libellagininae JACOBSON & BIANCHI, 1905)
    
  • Libellaginini sensu BECHLY, 1996a (sens. nov.)

---

Melanocyphina BECHLY, 1996

(Type genus: Melanocypha FRASER, 1949)

• Included taxa: only including the type genus Melanocypha FRASER, 1949.
• Autapomorphies:
  
  • Wing venation: two rows of cells between the CuA and the hind margin (FRASER, 1957; convergent to Paracypha).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Melanocyphina BECHLY, 1996a (subtrib. nov.)

---

Libellaginina JACOBSON & BIANCHI, 1905

(Type genus: Libellago SELYS, 1840, nec Libellago SELYS, 1953.)

• Included taxa: only including the two genera Libellago SELYS, 1840 and Pachycypha LIEFTINCK, 1950.
• Autapomorphies:
  
  • Wing venation: number of antenodal crossveins reduced to 5-6, rarely 7 (FRASER, 1957); MA strongly zigzagged (convergent to Chlorocyphidae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Libellaginina sensu BECHLY, 1996a (stat. et sens. nov.) (nom. transl. ex Libellagininae JACOBSON & BIANCHI, 1905)

---

Rhinocyphinae BECHLY, 1996

(Type genus: Rhinocypha RAMBUR, 1842)

• Included taxa: Disparocyphini and Rhinocyphini, also including the African genus Platycypha FRASER, 1949.
• Autapomorphies:
  
  • Wing venation: RP1/2 basally at least somewhat upward curved and arising on RP with a secondary insertion (convergent to Chlorocyphidae, Epallagidae and Calopterygida); unique enforced oblique vein between RP1/2 and IR2.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Rhinocyphinae BECHLY, 1996a (subfam. nov.)

---

Disparocyphini MUNZ, 1919

(Type genus: Disparocypha RIS, 1916.)

• Included taxa: including the two genera Disparocypha RIS, 1916 and Rhinoneura LAIDLAW, 1915.
• Autapomorphies:
  
  • Wing venation: very elongate and narrow wings; characteristically shaped pterostigmata (FRASER, 1957).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Disparocyphinae MUNZ., 1919 (subfam. nov.)
    
  • Disparocyphinae; DAVIES, 1981 (sens. nov.)
    
  • Disparocyphini sensu BECHLY, 1996a (stat. et sens. nov.)

---

Rhinocyphini BECHLY, 1996

(Type genus: Rhinocypha RAMBUR, 1842)

• Included taxa: including the genera Rhinocypha RAMBUR, 1842, Aristocypha LAIDLAW, 1950, Calocypha FRASER, 1928, Cyrano NEEDHAM & GYGER, 1939, Heliocypha FRASER, 1928, Heterocypha LAIDLAW, 1950, Indocypha FRASER, 1949, Paracypha FRASER, 1949, Platycypha FRASER, 1949, and Sundacypha LAIDLAW, 1950.
• Autapomorphies:
  
  • Wing venation: RP1/2 strongly upward curved at its base and arising on RP with a secondary insertion (convergent to Chlorocyphidae, Epallagidae and Calopterygida).
    
  • Other characters: adult males with four lobes on the terminal ligula segment (KENNEDY, 1919; these four lobes seem to be non-homologous to the lobes of the other quadrilobate ligulae within Caloptera).
• Taxonomy:
  
  • Rhynocyphini BECHLY, 1996a (trib. nov.)

Comment: Cyrano and Indocypha are sister-genera, as well as Rhinocypha and Aristocypha.

---

Chlorocyphidae COWLEY, 1937

(Type genus: Chlorocypha FRASER, 1928 nom. subst. pro Libellago SELYS, 1953, nec Libellago SELYS, 1840.)

• Included taxa: only including the African genus Chlorocypha FRASER, 1928, and maybe also Africocypha PINHEY, 1961.
• Autapomorphies:
  
  • Wing venation: MA strongly zigzagged (convergent to Libellaginina); RP1/2 strongly upward curved at its base and arising on RP with a secondary insertion (convergent to Rhinocyphinae, Epallagidae and Calopterygida).
    
  • Other characters: larval antenna shortened, with only six segments, but the pedicel segment is strongly prolonged (half of the total antennal length; convergent to Synlestidae and Calopterygida) (PINHEY in PARKER, 1982); the median caudal gill of the larvae is vestigial (PINHEY in PARKER, 1982); larval prehensile mask secondarily without a median cleft in the prementum (PINHEY in PARKER, 1982).
• Taxonomy:
  
  • Chlorocyphidae COWLEY, 1937 (fam. nov.)
    
  • Chlorocyphinae; DAVIES, 1981 (stat. nov.)
    
  • Chlorocyphidae sensu BECHLY, 1996a (sens. nov.)

---

Calopterygiformia BECHLY, 1996

• Included taxa: Euphaeida and Calopterygida.
• Autapomorphies:
  
  • Wing venation: presence of basal accessory antenodal crossveins between ax0 and ax1 (dubious since apparently absent in † Zacallitidae, and certainly absent in † Eodichrominae).
    
  • Other characters: interpleural suture of the pterothorax secondarily well-developed (FRASER, 1954; ASAHINA, 1957; reversal, convergent to some Synlestidae that might be explained with paedogenesis; again reduced in many Epallagidae); the ventral spines on the cross-venation are distinctly longer and more numerous in the hindwings.
• Taxonomy:
  
  • Calopterygiformia BECHLY, 1996a (taxon nov.)

---

Euphaeida BECHLY, 1996

• Included taxa: Epallagoidea and Polythoridae.
• Autapomorphies:
  
  • Wing venation: more strongly developed dorsal discoidal bracket on the distal side MAb of the discoidal cell and on the subdiscoidal veinlet (basal CuA).
    
  • Other characters: larvae with 6-7 pairs of ventro-lateral abdominal gills (6 pairs on segments 2-7 in Polythoridae and 7 pairs on segments 2-8 in Epallagidae, but the polarity of this difference is unclear; the abdominal gills are secondarily absent in some genera of Epallagidae, e.g. Anisopleura).
• Taxonomy:
  
  • [Euphaeoidea sensu HEYMER, 1975]
  • Euphaeida BECHLY, 1996a (taxon nov.)

---

Epallagoidea NEEDHAM, 1903

(Type genus: Epallage CHARPENTIER, 1840.)

• Included taxa: † Zacallitidae and Epallagidae.
• Autapomorphies:
  
  • Wing venation: wing venation more densely reticulated (convergent to Polythorinae and Calopterygidae); petiolus shortened (FRASER, 1940; convergent to Calopterygidae); discoidal cell very short (FRASER, 1940); distal discoidal vein MAb and subdiscoidal vein with reversed obliquity (convergent to Calopterygidae); CuA sigmoidally and smoothly curved (FRASER, 1940); curved intercalary veins between CuA and hind margin (FRASER, 1940); [M & Cu] or MP not kinked or bent at arculus (convergent to Calopterygoidea).
    
  • Other characters: thorax and abdomen relatively strong and stout, rather anisopterid-like (not yet known from † Eodichrominae).
• Taxonomy:
  
  • Euphaeoidea; HEYMER, 1975 (stat. et sens. nov.) (nom. transl. ex Euphaeinae JACOBSON & BIANCHI, 1905) (jun. subj. syn.)
    
  • Euphaeoidea sensu BECHLY, 1996a (sens. nov.)
    
  • Epallagoidea sensu BECHLY, 1998e (stat. nov.) (nom. transl. ex Epallaginae NEEDHAM, 1903)

---

† Zacallitidae COCKERELL, 1928

(Type genus: † Zacallites COCKERELL, 1928.)

• Included taxa: only including the type species † Zacallites balli COCKERELL, 1928.
• Autapomorphies:
  
  • Wing venation: apex of both wings dark coloured; base of IR2 widely separated from midfork (convergent to Philoganginae and Epallaginae); cubito-anal area secondarily expanded (convergent to † Eodichrominae, or a symplesiomorphy ?); hindwing with accessory convex intercalary "anal" vein (certainly not homologous with the concave intercalary "anal" vein in † Eodichrominae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Zacallitidae COCKERELL, 1928 (fam. nov.)
    
  • Zacallitidae sensu FRASER, 1940 (pos. nov.)
    
  • Zacallitidae; CARPENTER, 1992 (treated as junior subjective synonym of Epallagidae)
    
  • Zacallitidae; BRIDGES, 1994 (treated as junior subjective synonym of Epallagidae)
    
  • Zacallitidae sensu BECHLY, 1996a (stat. rest. et sens. nov.)

---

Epallagidae NEEDHAM, 1903

(Type genus: Epallage CHARPENTIER, 1840.)

• Included taxa: † Eodichrominae and Epallaginae.
• Autapomorphies:
  
  • Wing venation: antenodal crossveins more numerous, and both rows of antenodal crossveins strictly aligned (convergent to Calopterygoidea and Eurypalpida); at least a smooth basal curving of RP1/2 towards RA (convergent to many Chlorocyphoidea and all Calopterygida); arculus shifted basally between ax2 and ax1 (maybe rather a convergence of † Eodichrominae and Heliocharitidae, since this state cannot be determined in Epallaginae and Calopterygoidea).
    
  • Other characters: microsculpture of pterostigmata consisting of tiny pyramid-like structures (convergent to Devadatta, Chlorocyphoidea and Platystictidae; not yet known in Zacallitidae).
• Taxonomy:
  
  • [légion Euphaea SELYS, 1853 (not available as family-group taxon according to Art. 11f IRZN)]
    
  • [partim: Planinases SELYS, 1854 (taxon nov.)]
    
  • Epallaginae NEEDHAM, 1903 (subfam. nov.)
    
  • Euphaeinae JACOBSON & BIANCHI, 1905 (subfam. nov. with the type genus Euphaea SELYS, 1840; = Euphaea RAMBUR, 1842, jun. obj. syn. and homonym; = Pseudophaea KIRBY, 1890, erroneous nom. subst. pro Euphaea RAMBUR, 1842, nec Euphaea SELYS, 1840, jun. obj. syn.) (jun. subj. syn.)
    
  • Epallagidae sensu HANDLIRSCH, 1906 (stat. nov.) (nom. transl. ex Epallaginae NEEDHAM, 1903)
    
  • Epallaginae TILLYARD, 1917 (subfam. nov.) (jun. obj. syn. and homonym)
    
  • Epallagini TILLYARD, 1917 (trib. nov.) (jun. obj. syn. and homonym)
    
  • Epallagidae sensu TILLYARD & FRASER, 1939 (stat. et sens. nov.) (nom. transl. ex Epallaginae TILLYARD, 1917) (jun. obj. syn. and homonym)
    
  • Euphaeidae; MONTGOMERY, 1959 (stat. nov.)
    
  • Epallaginidae; DAVIES, 1981 (emend.)
    
  • Euphaeidae; CARPENTER, 1992 (sens. nov.)
    
  • Euphaeidae; BECHLY, 1996a (sens. nov.)

Comment: † Epallagites avus COCKERELL, 1924 from the Eocene of Colorado (Green River) is only known by a poorly preserved small fragment and thus should be preliminarily regarded as a Zygoptera - Caloptera incertae sedis (CARPENTER, 1992; NEL & PAICHELER, 1992) until better preserved material becomes available.

---

† Eodichrominae COCKERELL, 1923

(Type genus: † Eodichroma COCKERELL, 1923.)

• Included taxa: † Eodichromini and † Litheuphaeini.
• Autapomorphies:
  
  • Wing venation: cubito-anal area secondarily expanded (convergent to † Zacallitidae ?), with an unique accessory concave "anal" vein (intercalary) between CuA and the hind margin; ax1 and ax2 closely approximated; no antefurcal crossveins between basal parts of RP and MA.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Eodichrominae COCKERELL, 1923 (subfam. nov.) (nom. imperf.)
    
  • Parazacallitinae NEL, 1988 (subfam. nov. with type genus Parazacallites NEL, 1988) (subj. jun. syn.)
    
  • Parazacallitinae; BRIDGES, 1994 (treated as jun. sub. syn. of Chlorocyphidae)
    
  • Eodichromatinae sensu BECHLY, 1998e (unjust. emend. et sens. nov.)

---

† Eodichromini COCKERELL, 1923

(Type genus: † Eodichroma COCKERELL, 1923.)

• Included taxa: only including the species † Eodichroma mirifica COCKERELL, 1923 and † Parazacallites NEL, 1988.
• Autapomorphies:
  
  • Wing venation: anal area with a unique recurrent intercalary vein parallel to anal vein beneath subdiscoidal cell; small size (wing length only 20-21.5 mm).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Eodichromatini sensu BECHLY, 1998e (stat. nov.) (unjust. emend. et nom. transl. ex Eodichrominae COCKERELL, 1923)

Comment: the type genus † Eodichroma has the following two autapomorphies: characteristical pattern of intercalary veins in the cubito-anal area that form a asymmetrical triadic fork with CuA (quite different from the symmetrical triadic fork in Polythoridae, thus probably no putative synapomorphy); basal half of wing dark coloured. The second genus † Parazacallites has the following two autapomorphies: arculus shifted basal to ax1; pattern of secondary veins in the cubito-anal area.

---

† Litheuphaeini BECHLY, 1996

(Type genus: † Litheuphaea FRASER, 1955.)

• Included taxa: only including the species † Litheuphaea carpenteri FRASER, 1955 and a new species (L. ludwigi BECHLY, 1998e).
• Autapomorphies:
  
  • Wing venation: all secondary antenodal crossveins between ScP and RA suppressed (convergent to Amphipterygida sensu BECHLY, 1996a).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Litheuphaeini BECHLY, 1996 (trib. nov.)
    
  • Litheuphaeini sensu BECHLY, 1998e (pos. nov.)

Comment: the present conclusions are based on recent studies by the author (BECHLY, 1998e) of the holotype of † Litheuphaea carpenteri at MCZ, and a new specimen from Baltic amber in coll. Ludwig, Berlin. The drawings of the wing venation of † Litheuphaea carpenteri in the original description and later publications are definitely incorrect!

---

Epallaginae NEEDHAM, 1903

(Type genus: Epallage CHARPENTIER, 1840.)

• Included taxa: including all extant genera of Epallagidae listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: base of IR2 widely separated from the midfork (convergent to Philoganginae and † Zacallitidae); base of RP1/2 strongly curved towards RA, arising on RP with a secondary insertion (convergent to many Chlorocyphoidea and all Calopterygida); all antenodal crossveins developed as enforced brackets, so that the two primary antenodal brackets ax1 and ax2 can no longer be distinguished from the secondary antenodal crossveins (convergent to Calopterygoidea and many Eurypalpida).
    
  • Other characters: numerous macrotrichae on the dorsal surface of the wing vein ScP (unknown in the fossil Euphaeida and therefore maybe an autapomorphy for a more inclusive monophylum); lateral lobes of larval prehensile mask with only two endhooks (unknown in the fossil Euphaeida and therefore maybe an autapomorphy for a more inclusive monophylum).
• Taxonomy:
  
  • [partim: Regulières SELYS, 1854 (taxon nov.)]
    
  • Epallaginae NEEDHAM, 1903 (subfam. nov.)
    
  • Euphaeinae JACOBSON & BIANCHI, 1905 (subfam. nov. with the type genus Euphaea SELYS, 1840; = Euphaea RAMBUR, 1842, jun. obj. syn. and homonym; = Pseudophaea KIRBY, 1890, erroneous nom. subst. pro Euphaea RAMBUR, 1842, nec Euphaea SELYS, 1840, jun. obj. syn.) (jun. subj. syn.)
    
  • Epallaginae TILLYARD, 1917 (subfam. nov.) (jun. obj. syn. and homonym)
    
  • Euphaeinae; BECHLY, 1996a (sens. nov.)
    
  • Euphaeini sensu BECHLY, 1996a (sens. nov.)
    
  • Epallaginae sensu BECHLY, 1998e (sens. nov.)

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Polythoridae MUNZ, 1919

(Type genus: Polythore CALVERT, 1917 nom. subst. pro Thore SELYS, 1853, nec Thore KOCK, 1850.)

• Included taxa: Euthorinae and Polythorinae.
• Autapomorphies:
  
  • Wing venation: distal half of wings distinctly broadened; discoidal cell touches RA (convergent to Lestinae), because the arculus is only formed by the basal discoidal crossvein (= posterior arculus) that is developed as an apparent branch of [M & Cu] (unique type of arculus within Odonatoptera); insertion of arcular crossvein on [RP & MA] of secondary type (convergent to Eulestiformia sensu BECHLY, 1996a); dorsal arcular bracket reduced (convergent to Lestidae and Eurypalpida); median space traversed by numerous crossveins (convergent to most Calopterygidae); submedian space (cubital cell and subdiscoidal cell) traversed by numerous crossveins (convergent to most Calopterygoidea), so that the CuP-crossing (= anal crossing sensu FRASER, 1995) is unidentifiable; discoidal cell traversed by several crossveins (convergent to Calopterygidae); anterior side of discoidal cell concave (MA basally curved), and basal side at least twice as long as distal side; CuA secondarily forked into CuAa and CuAb (convergent to Calopterygidae), with a concave intercalary vein between these branches (triadic branching; reversed in Miocorini).
    
  • Other characters: adult males with four lobes of unique shape (the lateral ones having a filamentous apex) on the terminal segment of the ligula (KENNEDY, 1919); adult males with reduced paraprocts; resting position with wings closely apposed over the dorsum of the body (also occurring within Epallaginae); larvae with several angular projections on the saccoid caudal gills.
• Taxonomy:
  
  • [légion Thore SELYS, 1853 (not available as family-group taxon according to Art. 11f IRZN)]
    
  • [partim: Planinases SELYS, 1854 (taxon nov.)]
    
  • [Irregulières SELYS, 1854 (taxon nov.)]
    
  • {Thorinae sensu NEEDHAM, 1903 (according to Art. 39 IRZN objectively invalid name since based on a homonym type genus)}
    
  • Polythorinae sensu MUNZ, 1919 (subfam. nov.)
    
  • Polythoridae sensu TILLYARD & FRASER, 1939 (stat. nov.)

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Euthorinae FRASER, 1957

(Type genus: Euthore SELYS, 1869.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Euthorinae FRASER, 1957 (subfam. nov.)

---

Polythorinae MUNZ, 1919

(Type genus: Polythore CALVERT, 1917 nom. subst. pro Thore SELYS, 1853, nec Thore KOCK, 1850.)

• Included taxa: Miocorini and Polythorini.
• Autapomorphies:
  
  • Wing venation: the distal primary antenodal ax2 is obsolete (with exception of Chalcopteryx scintillans and some species of Cora, e.g. C. terminalis which have retained or regained the presence of ax1 and ax2.); wing venation more densely reticulated (convergent to Epallagoidea and Calopterygidae).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Polythorinae MUNZ, 1919
    
  • Polythorinae; FRASER, 1957 (sens. nov.)
    
  • {Thorinae NEEDHAM, 1903 (subfam. nov.) (according to Art. 39 IRZN objectively invalid name since based on a homonym type genus)}
    
  • Polythorinae sensu BECHLY, 1996a (sens. nov.)

---

Miocorini FRASER, 1957

(Type genus: Miocora CALVERT, 1917.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: CuA secondarily unbranched (reversal).
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Miocorinae FRASER, 1957 (subfam. nov.)
    
  • Miocorini sensu BECHLY, 1996a (stat. nov.)

---

Polythorini MUNZ, 1919

(Type genus: Polythore CALVERT, 1917 nom. subst. pro Thore SELYS, 1853, nec Thore KOCK, 1850.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: not yet known.
    
  • Other characters: not yet known.
• Taxonomy:
  
  • Polythorinae sensu FRASER, 1957 (sens. nov.)
    
  • Polythorini sensu BECHLY, 1996a (stat. nov.) (nom. transl. ex Polythorinae MUNZ, 1919)

---

Calopterygida BECHLY, 1996

• Included taxa: Heliocharitidae and Calopterygoidea.
• Autapomorphies:
  
  • Wing venation: RP1/2 strongly curved after its base, arising on RP with a secondary insertion (convergent to Rhinocyphini, Chlorocyphidae and Epallagidae), and apparently fused to RA for a short distance (convergent to some Epallaginae); discoidal cell rather elongate and narrow, and traversed by at least one crossvein; arculus shifted basally near ax1 (maybe rather a convergence of † Eodichrominae and Heliocharitidae, since this state cannot be determined in Epallaginae and Calopterygoidea).
    
  • Other characters: larvae with very long and slender legs; pedicel segment of larval antenna distinctly elongated to nearly half of the length of the complete antenna (convergent to Chlorocyphidae and Synlestidae); larvae with strongly elongated and triquetral lateral caudal gills and a shorter lamellate median caudal gill; larval prehensile mask with the anterior margin of the prementum triangularly projecting (convergent to Coenagrionodea) and an elongated and widened median cleft (more than a simple notch); adult males with four lobes of typical calopterygid shape on the terminal segment of the ligula (KENNEDY, 1919; LINDEBOOM, 1996, and pers. comm.); larval prehensile mask with the three endhooks of the lateral lobes ("palps") being strongly elongated and very acute.
• Taxonomy:
  
  • [partim: Planinases SELYS, 1854 (taxon nov.)]
    
  • Calopterygida BECHLY, 1996a (taxon nov.)

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Heliocharitidae TILLYARD & FRASER, 1939

(Type genus: Heliocharis SELYS, 1853.)

• Included taxa: only including the two species Heliocharis amazona SELYS, 1853 and Dicterias atrosanguinea SELYS, 1853 according to the revision of DUNKLE (1991), but for a different opinion see COSTA & SANTOS (1991).
• Autapomorphies:
  
  • Wing venation: arculus secondarily shifted between ax1 and ax2 or even to ax1 (convergent to the coenagrionid genus Metathemis and all Epiproctophora); nodal veinlet less oblique and anterior part of the subnodal veinlet strongly thickened (convergent to Lestidae); only one row of cells between MP and CuA (postsubdiscoidal space not distally widened) and between CuA and the hind margin (convergent to Rimanella, Chlorocyphoidea, Caliphaeidae and most Euzygoptera).
    
  • Other characters: dorsal surface of wing vein ScP with extremely long macrotrichae (max. 0,3 mm); legs very long and slender (in adults and larvae), and secondarily without distinct raptorial spines (DUNKLE, 1991; unique within Odonata); adult labial palps with a very long and sharp movable hook (DUNKLE, 1991).
• Taxonomy:
  
  • [légion Dicterias SELYS, 1853 (not available as family-group taxon according to Art. 11f IRZN)]
    
  • Heliocharitidae TILLYARD & FRASER, 1939 (fam. nov.)
    
  • Heliocharitidae sensu FRASER, 1957
    
  • Dicteriastidae MONTGOMERY, 1959 (fam. nov. with the type genus Dicterias SELYS, 1853) (nom. imperf.)
    
  • Dicteriidae sensu RACENIS, 1968 (emend., nom. imperf.)
    
  • Dicteriastatidae sensu BELYSHEV & HARITONOV, 1983 (emend., nom. imperf.)
    
  • Dicteriadidae sensu DUNKLE, 1991 (emend., nom. correct.)
    
  • Dicteryastidae sensu NEL & PAICHELER, 1993 (incorr. subseq. spell.)

Comment: contrary to the opinion of DUNKLE (1991) the "légions" of Selys clearly have to be rejected as available family-group taxa according to Art. 11(f)(i)(1) IRZN.

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Calopterygoidea SELYS, 1850

(Type genus: Calopteryx [LEACH] [1815], emended by BURMEISTER, 1839 pro Calepteryx [LEACH] [1815], incorr. orig. spell.; = Agrion FABRICIUS, 1775, nec Agrion [LEACH] [1815], senior objective synonym but a nomen ambiquum. For a complete synonymy see remark under Calopterygidae.)

• Included taxa: Caliphaeidae and Calopterygidae.
• Autapomorphies:
  
  • Wing venation: both rows of antenodal crossveins are strictly aligned (convergent to Epallagidae and Eurypalpida) and developed as brackets so that the two primary antenodal brackets ax1 and ax2 can not be identified; submedian space (cubital cell and subdiscoidal cell) traversed by numerous crossveins (convergent to Polythoridae), so that the CuP-crossing (= anal crossing sensu Fraser) is unidentifiable; [M & Cu] or MP not kinked or bent at the arculus (convergent to Epallagoidea).
    
  • Other characters: adult males with hypertrophied paraprocts; resting position with wings closely apposed over the dorsum of the body; body with metallic colouring; larval prehensile mask elongate, with a deeply fissured prementum (median cleft hypertrophied; convergent to Libellaginidae), the edges of the fissures overlapping (FRASER, 1957; convergent to Libellaginidae; still not certainly known for Caliphaeidae).
• Taxonomy:
  
  • [partim: Regulières SELYS, 1854 (taxon nov.)]
    
  • Calopterygoidea; MONTGOMERY, 1959 (stat. nov.) (nom. transl ex Calopterygina SELYS, 1850)
    
  • Calopterygoidea; MACNEILL, 1960
    
  • Agrioidea; TILLYARD, 1926 (stat. nov.) (nom. transl. ex Agrioninae KIRBY, 1890) (incorr. subseq. spell.) (jun. obj. syn.)
    
  • Agrionoidea; RACENIS, 1968
    
  • Agrionidea; HENNIG, 1969 (incorr. subseq. spell.)
    
  • Agrionidea; PRITYKINA, 1968 (incorr. subseq. spell.)
    
  • Calopterygoidea sensu BECHLY, 1996a (sens. nov.)

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Caliphaeidae FRASER, 1929

(Type genus: Caliphaea HAGEN in SELYS, 1859; = Notholestes MCLACHLAN, 1887, jun. subj. syn.)

• Included taxa: only including the two genera Caliphaea HAGEN in SELYS, 1859 and Noguchiphaea ASAHINA, 1976.
• Autapomorphies:
  
  • Wing venation: only one row of cells between MP and CuA (postsubdiscoidal space not distally widened) and between CuA and the hind margin (convergent to Rimanella, Chlorocyphoidea, Heliocharitidae and most Euzygoptera); at least CuA and RP3/4 are forked.
    
  • Other characters: allegedly breeding in stagnant waters (FRASER, 1957; would be quite unique in Eucaloptera, and rather seems to be a doubtful myth).
• Taxonomy:
  
  • Caliphinae FRASER, 1929: 595 (subfam. nov.) (nom.imperf.)
    
  • Caliphaeinae; FRASER, 1934: 148 (nom. correct.)
    
  • Caliphaeinae; TILLYARD & FRASER, 1939
    
  • Caliphaeidae HEYMER, 1975 (stat. nov.)

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Calopterygidae SELYS, 1850

(Type genus: Calopteryx [LEACH] [1815], emended by BURMEISTER, 1839 pro Calepteryx [LEACH] [1815], incorr. orig. spell.; = Agrion FABRICIUS, 1775, nec Agrion [LEACH] [1815], senior objective synonym but a nomen ambiquum. Type species Libellula virgo LINNAEUS, 1758 by subsequent type designation by WESTWOOD, 1840. Complete synonymy of the type genus see remark below.)

• Included taxa: Hetaerininae and Calopteryginae.
• Autapomorphies:
  
  • Wing venation: wings secondarily not petiolated (convergent to most Epallagidae) and more densely reticulated (convergent to Polythorinae and Epallagoidea), with much more antenodal crossveins (convergent to Epallagidae); median space traversed by numerous crossveins (reversed in several genera of Calopteryginae; convergent to Polythoridae); discoidal cell very elongate and narrow, and traversed by numerous crossveins; distal discoidal vein MAb and subdiscoidal vein with reversed obliquity (convergent to Epallagoidea).
    
  • Other characters: derived type of microsculpture of the pterostigmata (HEYMER, 1975; BECHLY, 1994).
• Taxonomy:
  
  • Calopterygina SELYS, 1850
    
  • Calopterygines; SELYS & HAGEN, 1850
    
  • Calopterygidae; BUCHECKER, 1876 (stat. nov.)
    
  • Calopterygii; ACLOQUE, 1897 (incorr. subseq. spell.)
    
  • Calopterygidae; NEEDHAM, 1903
    
  • Agrionidae KIRBY, 1890 (fam. nov.) (jun. obj. syn.)
    
  • Agrionidae; MUTTKOWSKI, 1910
    
  • Agriidae; TILLYARD, 1926 (sens. nov.) (incorr. subseq. spell.)
    
  • Agriuntidae; FORBES, 1940 (incorr. subseq. spell.)
    
  • Agrioidae; WHATMOUGH in MONTGOMERY, 1954 (incorr. subseq. spell.)
    
  • Agriadidae; WHATMOUGH in MONTGOMERY, 1967 (incorr. subseq. spell.)

Comment: the type genus Calopteryx [LEACH] [1815] involves one of the most difficile taxonomic problems, known as the "Agrion versus Calopteryx problem". Calopteryx [LEACH] [1815] is an junior objective synonym but in very common use since many years (Art. 79c IRZN), while the previous use of the senior synonym Agrion FABRICIUS, 1775 has caused a significant confusion in the past (Art. 79a IRZN), also because the objectively invalid homonym Agrion [LEACH] [1815] was often used instead of the valid replacement name Coenagrion KIRBY, 1890. This ambiquity also led to confusion of the referring family-group taxa (Agrionidae sensu Calopterygidae versus Agrionidae sensu Coenagrionidae). NEEDHAM (1903) and TILLYARD (1917) for example used Agrionidae in the sense of Coenagrionidae, while KIRBY (1890) and TILLYARD & FRASER (1938) used Agrionidae or Agriidae in the sense of Calopterygidae. Furthermore there was great confusion about the subsequent type species designation of Libellula virgo LINNAEUS, 1758 for Agrion FABRICIUS, 1775 by LATREILLE (1810). Therefore, Agrion FABRICIUS, 1775 has to be regarded as a nomen ambiquum. Only a partially suppression (Art. 79bii IRZN) of the valid senior objective synonym Agrion FABRICIUS, 1775, only for the purpose of the Principle of Priority but not for the pupose of the Principle of Homonymy, could solve this complex nomenclatorial problem (also see BRIDGES, 1994). This case definitely should be referred soon to the Commision (ICZN) for a ruling according to Art. 79 IRZN.
Synonymy of Calopteryx: Agrion FABRICIUS, 1775 (available and valid senior objective synonym), nec Agrion [LEACH], [1815] (available and valid senior objective synonym); Agrionus RAFINESQUE, 1815 (available but invalid junior objective synonym through unjustified emendation); Calepteryx LEACH, 1815 (unavailable incorrect original spelling); Agrio SELYS, 1831 (available but invalid junior objective synonym through unjustified emendation); Calopteryx BURMEISTER, 1839 (justified emendation pro Calepteryx LEACH, 1815); Callepteryx HAGEN, 1840 (unavailable incorrect subsequent spelling); Callipteryx AGASSIZ (1847) 1842-1847 (available but invalid junior objective synonym through unjustified emendation); Agrium AGASSIZ, 1847 (available but invalid junior objective synonym through unjustified emendation); Sylphis HAGEN (?) in SELYS, 1853 (available and valid junior subjective synonym); Calopterix BENTIVOGLIO, 1897 (unavailable incorrect subsequent spelling); Calapteryx BRAUNER, 1902 (available but invalid objective junior synonym through unjustified emendation); Anaciagrion KENNEDY, 1920 (available and valid junior subjective synonym); Pseudomatrona FRASER, 1935 (unavailable name, since published in synonymy).
As explained by MONTGOMERY (1954) there are several severe errors in the paper of TILLYARD (1926), e.g. the stem of the genus Agrion is agrio- not agri-, and thus the family-group names must end on -agrionidae not -agriidae. The derivation of the name Agrion is somewhat disputed since the are several alternatives (see MONTGOMERY 1954, 1967).

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Hetaerininae TILLYARD & FRASER, 1939

(Type genus: Hetaerina HAGEN in SELYS, 1853; = Heterina MUNZ, 1919, incorr. subseq. spell.; = Mnesarete COWLEY, 1934 nom. subst. pro Lais HAGEN (?) in SELYS, 1853, which is a junior homonym; new synonymy.)

• Included taxa: only including the genus Hetaerina HAGEN in SELYS, 1853. The genus Mnesarete COWLEY, 1934 should be synonymised with Hetaerina since the latter otherwise would be paraphyletic.
• Autapomorphies:
  
  • Wing venation: arculus obliquely slanted, straight and posterior part of arculus strongly reduced (convergent to Chlorogomphoidea and Eurypalpida); sectors of arculus diverging from one point; discoidal cell strongly curved (FRASER, 1957, the anterior side being convex; red colour pattern of the wings (FRASER, 1957); MP of both pairs of wings with a unique kink shortly distal of the discoidal cell (MP sigmoidally curved; FRASER, 1957), apparently correlated with a unique curvature of CuA beneath the discoidal cell; the CuA of forewings is basally much stronger curved than in hindwings, and the forewing cubito-anal space more reticulated than that of the hindwing (contrary to Calopteryginae); subdiscoidal vein (basal CuA) distinctly shortened.
    
  • Other characters: lateral gills of larvae of unique shape (strongly broadened and more distinctly triquetral); male ligula secondarily with only two terminal lobes since the apical pair is completely reduced (KENNEDY, 1919; a vestige of the second pair of ligula lobes is uniquely retained in the species Mnesarete (= Lais) pudica which therefore could be the sistergroup of all other Hetaerininae which implies the paraphyly of the genus Mnesarete).
• Taxonomy:
  
  • [légion Hetaerina SELYS, 1853 (not available as family-group taxon according to Art. 11f IRZN)]
    
  • Hetaerininae TILLYARD & FRASER, 1939 (subfam. nov.)
    
  • Hetaerinidae sensu FRASER, 1954 (stat. nov.)
    
  • Hetaerinidae sensu FRASER, 1957
    
  • Hetaerininae sensu FRASER, 1957
    
  • Hetaerinidae sensu HEYMER, 1975
    
  • Hetaerinidae sensu PFAU, 1991
    
  • Hetaerininae sensu BECHLY, 1996a

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Calopteryginae SELYS, 1850

(Type genus: Calopteryx [LEACH] [1815], emended by BURMEISTER, 1839 pro Calepteryx [LEACH] [1815], incorr. orig. spell.; = Agrion FABRICIUS, 1775, nec Agrion [LEACH] [1815], senior objective synonym but a nomen ambiquum. For a complete synonymy see remark under Calopterygidae.)

• Included taxa: including the genera listed in BRIDGES (1994).
• Autapomorphies:
  
  • Wing venation: wings more or less broadened (FRASER, 1957); the cubito-anal field of the hindwing more is expanded than that of the forewing (both pairs of wings distinctly different); all longitudinal veins are distally strongly curved towards the wing margin; discoidal cell extremely elongate and narrow (FRASER, 1957); CuA secondarily forked into two strong branches CuAa and CuAb, especially in the hindwing.
    
  • Other characters: median cleft of larval mask enlarged, reaching nearly half of the length of the prementum.
• Taxonomy:
  
  • Calopterygina SELYS, 1850
    
  • Calopterygines; SELYS & HAGEN, 1850
    
  • [légion Calopteryx SELYS, 1853 (not available as family-group taxon according Art.11f IRZN)]
    
  • Calopteryginae; WALKER, 1853
    
  • Calopteryginae; CALVERT, 1893
    
  • Vestalinae NEEDHAM, 1903 (subfam. nov.) (jun. subj. syn.)
    
  • Agrioninae KIRBY, 1890 (subfam. nov.) (jun. obj. syn.)
    
  • Agriinae; TILLYARD & FRASER, 1938 (incorr. subseq. spell.)