Classification of the Cixiidae

Family Cixiidae

Early classifications of Cixiidae, that of Muir (1923, 1925) provided only 1 subfamily (Cixiinae) and 2 tribes (Bothriocerini and Cixiini).  Muir (1925) provided a key to genus.   Interestingly, Muir did not recognize his own tribe Oecleini Muir, 1922. The classification of Muir (1925) was subsequently adopted by the Metcalf catalog (Metcalf 1936).  Metcalf (1938) elevated Bothriocerini to subfamily and defined it to include the new tribe Stenophlepsini (for 4 unspecified genera, but now including only Euryphlepsia and Stenophlepsia, subsequently moved into the Cixiinae); within the Cixiinae, he defined the Bennini, Pintaliini and the subtribe Myndina (for Oecleus and others) in the Cixiini. Little more was done until Emeljanov (1971) separated out the Pentastirini (with 2 subtribes).

Emeljaonv (1989) revised the higher classification, and subsequently described several new tribes in Emeljanov (2002).  Emeljanov (2002)  and Holzinger et al. (2002) provide an overview of the current classification of Cixiidae, which now includes 3 subfamilies (Borystheninae, Bothriocerinae and Cixiinae) and 16 tribes, on which the following classification is based.

The only quantitative phylogenetic work on the family was that by Ceotto and Bourgoin (2008) and Ceotto et al. (2008) (published trees given below).  This work provides and excellent starting point, but does not entirely support or refute the existing classification.  It does point out a variety of issues that remain unresolved (including whether the Delphacidae are derived within the Cixiidae).  It should also be noted that fossils attributed to the Cixiidae (as early as the late Permian) may have a substantive impact on our view of the age and definition of the Cixiidae.

In addition to the work of Ceotto and colleagues, Löcker et al. (2006) has published a morphological phylogeny of the  Gelastocephalini (tree shown below).  Löcker and colleagues have revised the majority of the fauna of Australia and New Zealand, describing many new species.

Tree based on morphology from Ceotto and Bourgoin (2008).

From Ceotto et al 2008 (figure 1):  Majority rule consensus of 18 most-parsimonious trees (length = 5302; CI = 0.39; RI = 0.54) of the Cixiidae family. Bootstrap values (>50%) are above branches and Bremer decay indexes below branches. Node numbers are indicated. The numbers in the table are the contribution of the Cox1, Cytb, 18S rDNA, domain D4–D5 of 28S rDNA, and domain D6–D7 of 28S rDNA partitions to the Bremer support values (results of the Partitioned Bremer Support analysis). The table also indicates the percentage of
negative values for each partition.

From Ceotto et al 2008 (Figure 2). Bayesian inference topology of Cixiidae derived a priori partitioning strategy P3 (Three partitions corresponding to 1st, 2nd and 3rd codon positions of mitochondrial genes, three partitions corresponding to the ribosomal genes: 18S rDNA, regions D4–D5 and D6–D7 of the 28S rDNA. Numbers above branches are posterior probabilities values ( > 50%).

From Ceotto et al. 2008 (Figure 3). Bayesian inference topology of Cixiidae derived with 6Q mixture models. Numbers above branches are posterior probabilities values ( > 50%).

Locker et al. 2006, Fig. 5 (and Figs. 6, 7 below). Subtree a of the phylogram of one of three most parsimonious trees of dataset A obtained under the modified scaled weighting regime. Character numbers of all aut- and synapomorphies with consistency index (CI) ≥ 0.25 are indicated (outgroups and aut- and synapomorphies with CI = 1 in bold).

 

F. Asotocixius diopter (paratype), G. Bothriocera cognita, H. Cixius pini, I. Haplaxius radicis, J. Melanoliarus placitus.

Image Galleries
GBIF
Bugguide
Insects of Australia
iNaturalist
Vectors of Phytoplasma on grepevine.

 

Cixiidae, Peru

Undetermined Cixiid from Tambopata, Madre de Dios, Peru by Arnold Wijker.

 

Family Cixiidae Spinola 1839

Subfamily Borystheninae Emeljanov, 1989
Tribe Borysthenini Emeljanov, 1989
Borysthenes Stal 1866

Borysthenes

I suspect this is Borysthenes (Wiki commons, L. Shyamal)

Subfamily Bothriocerinae Muir 1923
Tribe Bothriocerini Muir 1923
Bothriocera Burmeister 1835
Bothrioceretta Caldwell 1950
Bothriobaltia† Szwedo, 2002

Bothriocera from Texas

Bothriocera from Texas (from iNaturalist)

 Subfamily Cixiinae Spinola 1839
Tribe Andini Emeljanov, 2002
Andes Stal 1866
Andixius Emeljanov & Hayashi, 2007
Parandes Muir, 1925

Tribe Bennarellini Emeljanov, 1989
Amazobenna Penny 1980
Bennarella Muir 1930
Loisirella Holzinger, Holzinger & Egger, 2013
Noabennarella Holzinger & Kunz, 2006

Tribe Bennini Metcalf 1938
Benna Walker 1857 (Type genus; = Kotonisia Matsumura, 1938 synonym of Benna Walker, 1857 according to Tsaur 2009: 67)
Bennaria Melichar, 1914 synonym of Benna Walker, 1857 according to Fennah 1970: 46.
Canobenna Hoch, 2013
Celebenna Hoch & Wessel, 2011
Clarabenna Hoch, 2013
Cordobenna Hoch, 2013
Cyclobenna Hoch, 2013
Dentobenna Hoch, 2013
Fuscobenna Hoch, 2013
Goniobenna Emeljanov, 1989
Jonabenna Hoch, 2013
Kinabenna Hoch, 2013
Kotonisia Matsumura, 1938
Madangabenna Hoch, 2013
Malukubenna Hoch, 2013
Minabenna Hoch, 2013
Negrobenna Hoch, 2013
Papuabenna Hoch, 2013
Rhyobenna Hoch, 2013
Romabenna Hoch, 2013
Sanghabenna Hoch, 2013
Silangobenna Hoch, 2013
Singabenna Hoch, 2013
Sinubenna Hoch, 2013
Striabenna Hoch, 2013
Thaumatobenna Hoch, 2013
Torrebenna Hoch, 2013

Tribe Brixidiini Emeljanov 2002
Brixidia Haglund 1929

Tribe Brixiini Emeljanov, 2002
Brixia Stal 1856
Caffrocixius Fennah 1967
Cibrica Emeljanov, 2007
Innobindus Jacobi 1928
Ithma Fennah 1969
Melandeva Distant 1906
Solonaima Kirkaldy 1906
Typhlobrixia Synave 1953
Undarana Hoch & Howarth 1989

Brixia albomaculata, Wynaad, India (L. Shyamal, Wiki commons)

Tribe Cajetini Emeljanov, 2002
Cajeta Stal, 1866

Tribe Cixiini Spinola 1839
Achaemenes Stål, 1866
Aka White, 1879
Anilus Distant, 1906 [Anila Distant, 1906 missp.]
Ankistrus Tsaur & Hsu, 1991
Apartus Holzinger, 2002
Aselgeoides Distant 1917
Asotocixius Kramer, 1983
Calamister Kirkaldy, 1906
Caravella Emelyanov 2006
Cermada Emeljanov, 2000
Chathamaka Larivière, 1999
Chidaea Emeljanov, 2000 (or Chidea)**
Cixiosoma Berg, 1879
Cixius Latreille, 1804 (Type genus)
Ferricixius Hoch & Ferreira, 2012
Flachaemus Van Stalle, 1988
Glisachaemus† Szwedo, 2007
Gonophallus Tsaur & Hsu, 1991
Ibleocixius D’Urso & Grasso, 2009
Iolania Kirkaldy, 1902
Koroana Myers, 1924
Leades Jacobi, 1928
Leptolamia Metcalf, 1936
Macrocixius Matsumura, 1914
Malpha Myers, 1924
Microledrida Fowler, 1904
Monomalpha Emeljanov, 2000
Nanocixius Wagner, 1939
Neocixius Wagner, 1939
Pachyntheisa Fowler, 1904
Platycixius Van Duzee, 1914
Sardocixius Holzinger, 2002
Semicixius Tsaur & Hsu, 1991
Simplicixius Holzinger, 2002
Sphaerocixius Wagner, 1939
Stegocixius Kramer, 1983
Tachycixius Wagner, 1939
Taomma Emeljanov, 2007
Trirhacus Fieber, 1875
Tsauria Koçak & Kemal, 2009 (= Discophorellus Tsaur & Hsu, 1991 nomen praeoccupatum replaced by Tsauria Koçak & Kemal, 2009 nomen novum, according to Xing & Chen 2014: 149)
Yanganaka Löcker, 2015

Tachycixius pilosus, borrowed from British bugs

Tribe Duiliini Emeljanov, 2002
Duilius Stal, 1858 (Duiliopsis De Bergevin, 1933 synonym of Duilius Stål, 1858 according to Emeljanov 2007: 126)

Tribe Eucarpiini Emelyanov 2002
Adolendana Distant 1917
Bajauana Distant 1907
Caneirona Distant 1916
Dilacreon Fennah, 1980
Dysthaetias Kirkaldy 1907
Eucarpia Walker 1857
Kirbyana Distant 1906
Neocarpia Tsaur & Hsu, 2003
Nesochlamys Kirkaldy 1907
Nothocharis Muir 1925
Phytocentor Fennah 1980
Pterolophus Emeljanov, 2013

Tribe Gelastocephalini Emeljanov, 2000
Subtribe Gelastocephalina Emeljanov, 2000
Aubirestus Löcker & Larivière, 2006
Balyadimetopia Löcker & Larivière, 2006
Barbonia Löcker & Larivière, 2006
Carolus Kirkaldy 1906
Candicarina Löcker & Larivière, 2006
Corylonga Löcker & Fletcher, 2006
Fletcherolus Löcker & Larivière, 2006
Gelastocaledonia Löcker & Larivière, 2006
Gelastocephalus Kirkaldy 1906 (=Tarberus Jacobi 1928, synonym of Gelastocephalus Kirkaldy, 1906 according to Fennah 1969)
Gurrundus Löcker & Larivière, 2006
Guttala Löcker & Larivière, 2006
Hartliebia Löcker & Larivière, 2006
Holgus Löcker & Larivière, 2006
Larivierea Löcker & Fletcher, 2006
Lipsia Löcker & Fletcher, 2006
Metaplacha Emeljanov 2000
Melanoclypeus Löcker & Fletcher, 2006
Novotarberus Löcker & Fletcher, 2006
Payastylus Löcker & Fletcher, 2006
Rokebia Löcker & Fletcher, 2006
Ronaldia Emeljanov, 2000
Schuerrera Löcker & Fletcher, 2006
Wernindia Löcker & Fletcher, 2006
Yarnikada Löcker & Fletcher, 2006

Subtribe Rhigedanina Emeljanov, 2000
Dysoliarus Fennah 1949
Orphninus Emeljanov, 2000
Rhigedanus Emeljanov, 2000

Incertae subtribe
Cicerama Emeljanov, 2007

Tribe Mnemosynini Emeljanov, 1992
Autrimpus† Szwedo, 2004
Mnaomaia† Szwedo, Bourgoin & Lefebvre, 2006
Mnasthaia† Szwedo, Bourgoin & Lefebvre, 2006
Mnemosyne Stal, 1866  (type genus)
Stalisyne† Szwedo, Bourgoin & Lefebvre, 2006

Tribe Oecleini Muir 1922 (= Myndini Muir 1923)
Achaebana Attié, Bourgoin & Bonfils, 2002
Antillixius Myers, 1928
Borbonomyndus Attié, Bourgoin & Bonfils, 2002
Colvanalia Muir, 1925
Confuga Fennah, 1975
Eumyndus Synave, 1956
Fipsianus Holzinger, 2009
Haplaxius Fowler, 1904
Mundopa Distant, 1906
Myndodus Emeljanov, 1992
Myndus Stål, 1862
Nesomyndus Jacobi, 1917
Notolathrus Remes Lenicov, 1992
Nymphocixia Van Duzee, 1923
Nymphomyndus Emelyanov 2006
Oecleus Stål, 1862
Perindus Emeljanov, 1989
Pinacites Emeljanov, 1972
Proclytus Emelyanov 2007
Rhamphixius Fowler, 1904
Tiriteana Myers, 1924
Trigonocranus Fieber, 1875
Volcanalia Distant, 1917

Oecleus sp. (San Diego, CA)

Tribe Pentastirini Emeljanov, 1971
Subtribe Pentastirina Em., 1971 (= Oliarina Emeljanov, 1971)
Afroreptalus Van Stalle, 1986
Anoculiarus Dlabola, 1985
Arosinus Emeljanov, 2007
Atonurus Emeljanov, 1992
Atretus Emeljanov, 2007
Cordoliarus Löcker, 2006
Cyclopoliarus Fennah, 1971
Dorialus Van Stalle, 1986
Epoliarus Matsumura, 1910
Eumecurus Emeljanov, 1971
Helenolius Van Stalle, 1986
Hyalesthes Signoret, 1865
Indolipa Emeljanov, 2001
Kibofascius Van Stalle, 1986
Lalobidius Van Stalle, 1985
Manurevana Hoch, 2006
Melanoliarus Fennah, 1945
Mesoliarus Matsumura, 1910
Miclucha Emeljanov, 2001
Narravertus Van Stalle, 1986
Nesoliarus Kirkaldy, 1909
Nivcentia Holzinger, 2004 (Vincentia Uhler, 1895 nomen praeoccupatum replaced by Nivcentia Holzinger, 2004: 951)
Norialsus Van Stalle, 1986
Oecleopsis Emeljanov, 1971
Oliarellus Emeljanov, 1971
Oliarus Stål, 1862
Oliarissa Fennah, 1945
Oliaronus Ball, 1934
Olipa Emeljanov, 2001
Oliparisca Emeljanov, 2001
Oteana Hoch, 2006
Ozoliarus Löcker, 2006
Peartolus Van Stalle, 1986
Pentastira Kirschbaum 1868
Pentastiridius Kirschbaum, 1868
Prosops Buckton, 1893
Pseudolarius Haupt, 1927
Reptalus Emeljanov, 1971
Setapius Dlabola, 1988
Siniarus Emeljanov, 2007
Suriola Emeljanov, 1992
Trirhacus Fieber, 1875
Urvillea Kirkaldy, 1907
Zeoliarus Larivière & Fletcher, 2008

Reptalus panzeri

Reptalus panzeri from here.

Tribe Pintaliini Metc. 1938
Aulocoryoha Berg, 1879
Cubana Uhler 1895
Cubanella Fennah 1948
Diastrocixius Caldwell, 1944
Monorachis Uhler 1901
Muirolonia Metcalf 1936
Notocixius Fennah, 1965
Pintalia stal, 1862

Pintalia delicata

Pintalia delicata

Tribe Semonini Emeljanov, 2002
Betacixius Matsumura 1914
Huttia Myers 1924
Kuvera Distant 1906
Parasemo Lariviere 1999
Semo White 1879

Tribe Stenophlepsiini Metcalf, 1938
Euryphlepsia Muir 1922
Stenophlepsia Muir 1922

Incertae tribe/subfamily
Meenocixius Attie, Bourgoin & Bonfils 2002 (Cixiinae, incertae sedis)
Bangoliarus Shakila, 1984
Balticixius† Lefebvre, Bourgoin & Nel, 2007
Cretofennahia† Martins-Neto & Szwedo, 2007
Issomimus Jacobi, 1910
Karebodopoides† Szwedo, 2001
Kulickamia† Gebicki & Szwedo, 2000
Mundopoides† Cockerell, 1925
Oeclixius† Fennah, 1963
Oligocixia† Gebicki & Wegierek, 1993
Perkunus† Szwedo & Stroinski, 2007
Chlorodus Johnson & Ledig, 1918

Huanh and Holzinger 2010, Figures 1–13. Habitus photographs, dorsolateral view. 1. Duilius tenuis, female; 2. Pentastiridius moestus, male; 3. P. sudanicus, male; 4. P. limbifer, male; 5. Eumecurus incompletus, male; 6 E. kibuyanus, male; 7. E. decempunctatus, male; 8. E. mashonanus, male; 9. E. brachycephalus, male; 10. Atonurus meridianus, male; 11. E. hottentottus, female; 12. A. namibianus, male; 13. E. eryx, female. Scale bar A = 1 mm for figs 1–6, scale bar B = 1 mm for figs 7–13.

Muirolonia metallica

Undetermined Cixiid from Ecuador, from INaturalist jmtobin6975. I suspect Muirolonia metallica (Cixiinae: Pintaliini)

(the following table is out of date, but indicative)

Species count
PA NA ET OR AU  NT total spp
Borystheninae Emeljanov, 1989 0 0 0 25 0 0 25
Bothriocerinae Muir 1923 0 9 0 0 0 37 46
Cixiinae Spinola 1839 184 168 217 205 315 249 1338
Andini Emeljanov, 2002 0 0 0 1 5 0 6
Bennarellini Emeljanov, 1989 0 0 0 0 0 4 4
Bennini Metcalf 1938 0 0 0 23 0 0 23
Brixidiini Emeljanov, 2002 0 0 0 0 0 0 0
Brixiini Emeljanov, 2002 0 0 7 1 31 0 39
Cajetini Emeljanov, 2002 0 0 0 0 1 0 1
Cixiini Spinola 1839 39 34 51 16 37 32 209
Duiliini Emeljanov, 2002 0 0 0 0 0 0 0
Eucarpiini Emeljanov, 2002 0 0 0 79 52 0 131
Gelastocephalini Emeljanov, 2000 0 0 0 0 64 0 64
Oecleini Muir 1922 (= Myndini Muir 1923) 6 78 21 22 2 58 187
Pentastirini Emeljanov, 1971 139 52 127 49 116 70 553
Pintaliini Metcalf 1938 0 4 0 0 0 85 89
Semonini Emeljanov, 2002 0 0 0 0 7 0 7
Stenophlepsiini Metcalf 1938 0 0 0 14 0 0 14
unplaced 1 0 2 0 0 0 3
Totals 369 345 425 435 630 535 2739

 

References (higher classification only)

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