Phylogenetic Systematics of Odonata


© Günter Bechly, Böblingen, 2007


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Phylogenetic Systematics of Zygoptera / Caloptera




Zygoptera SELYS, 1854



Steleopteridae HANDLIRSCH, 1906

(Type genus: † Steleopteron HANDLIRSCH, 1906.)

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.



Eosagrionidae HANDLIRSCH, 1920

(Type genus: Eosagrion HANDLIRSCH, 1920.)



Caloptera BELYSHEV & HARITONOV, 1983

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.



Sieblosiidae HANDLIRSCH, 1907

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



Eucaloptera BECHLY, 1996



Amphipterygida BECHLY, 1996



Diphlebiidae HEYMER, 1975

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



Philoganginae KENNEDY, 1920

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



Diphlebiinae HEYMER, 1975

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



Amphipterygoidea TILLYARD, 1917

(Type genus: Amphipteryx SELYS, 1853.)



Pseudolestidae FRASER, 1957

(Type genus: Pseudolestes KIRBY, 1900.)



Thaumatoneuridae TILLYARD & FRASER, 1938

(Type genus: Thaumatoneura MCLACHLAN, 1897.)



Dysagrioninae COCKERELL, 1908

(Type genus: † Dysagrion SCUDDER, 1878.)



Petrolestini COCKERELL, 1927

(Type genus: † Petrolestes COCKERELL, 1927.)



Dysagrionini COCKERELL, 1908

(Type genus: † Dysagrion SCUDDER, 1878.)



Thaumatoneurinae TILLYARD & FRASER, 1938

(Type genus: Thaumatoneura MCLACHLAN, 1897.)



Euarchistigmatini CARLE & WIGHTON, 1990

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



Thaumatoneurini TILLYARD & FRASER, 1938

(Type genus: Thaumatoneura MCLACHLAN, 1897.)



Amphipterygidae TILLYARD, 1917

(Type genus: Amphipteryx SELYS, 1853.)



Rimanellinae DAVIES & TOBIN, 1984

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



Amphipteryginae TILLYARD, 1917

(Type genus: Amphipteryx SELYS, 1853.)



Calopterygomorpha BECHLY, 1996



Chlorocyphoidea COWLEY, 1937

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



Libellaginidae JACOBSON & BIANCHI, 1905

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



Libellagininae JACOBSON & BIANCHI, 1905

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



Sclerocyphini BECHLY, 1996

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



Libellaginini JACOBSON & BIANCHI, 1905

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



Melanocyphina BECHLY, 1996

(Type genus: Melanocypha FRASER, 1949)



Libellaginina JACOBSON & BIANCHI, 1905

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



Rhinocyphinae BECHLY, 1996

(Type genus: Rhinocypha RAMBUR, 1842)



Disparocyphini MUNZ, 1919

(Type genus: Disparocypha RIS, 1916.)



Rhinocyphini BECHLY, 1996

(Type genus: Rhinocypha RAMBUR, 1842)

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.)



Calopterygiformia BECHLY, 1996



Euphaeida BECHLY, 1996



Epallagoidea NEEDHAM, 1903

(Type genus: Epallage CHARPENTIER, 1840.)



Zacallitidae COCKERELL, 1928

(Type genus: † Zacallites COCKERELL, 1928.)



Epallagidae NEEDHAM, 1903

(Type genus: Epallage CHARPENTIER, 1840.)

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.)



Eodichromini COCKERELL, 1923

(Type genus: † Eodichroma 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.)

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.)



Polythoridae MUNZ, 1919

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



Euthorinae FRASER, 1957

(Type genus: Euthore SELYS, 1869.)



Polythorinae MUNZ, 1919

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



Miocorini FRASER, 1957

(Type genus: Miocora CALVERT, 1917.)



Polythorini MUNZ, 1919

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



Calopterygida BECHLY, 1996



Heliocharitidae TILLYARD & FRASER, 1939

(Type genus: Heliocharis SELYS, 1853.)

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.



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.)



Caliphaeidae FRASER, 1929

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



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.)

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).



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.)



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.)





Last Update: 10th August, 2007

© Günter Bechly, Böblingen, 2007