Classification of the Derbidae


Muir (1913) divided the Derbidae into 4 groups that were not formally named, based principally on features of the forewing venation (length of the tegmen relative to the body, cubital veins reaching or not reaching the hind margin of the wing, third claval cell open, extending along hind margin to last apical median cell) or closed, cubitus with 0-1 forks vs. 2+), and modifications of the head and prothorax around the antennae.   The four informal groups were referred to as the 1. Cenchrea, 2. Otiocerus, 3. Sikaiana+Zoraida, 4. Derbe+Mysidia groups.  In 1917, Muir formalized these into subfamilies, except groups 3 and 4 were combined except for Rhotana, and the groups were formalized into the Derbinae (Derbini, Sikaianini), Otiocerinae, Cenchreinae, and Rhotaninae, with a key to subfamilies (and genera in the Phillipines) provided. In 1918a, Muir described the Zoraidinae (Zoraidini, Sikaianini), and (1918b) revised the concepts of Derbinae to include tribes Derbini, Cenchreini, Otiocerini, and Rhotanini; with keys to genera.  This classification was reaffirmed in Muir (1923), and modified slightly by Muir (1930), who presented a key to subfamilies and tribes.

The revised classification in Muir (1918a, b; 1930) was adopted (with modifications) in the Metcalf catalogue (1945) is a modification of the classification proposed by Muir in 1918 as modified in 1930, as follows.

Subfamily DERBINAE

Higher taxonomy was entirely revised and reinterpreted by Fennah (1952). Fennah critiqued the principle diagnostic features utilized my Muir (viz. forewings having a reduced or normal cubital area, and “tegmina long and narrow” [in Zoraidinae])  as ignoring other points of similarity that suggested phylogenetic affinities.  Fennah discussed a series of other features of potential phylogenetic significance, presented keys to tribe and genus, and redescribed principle groups.  Fennah asserted that his Cenchreini (which in his definition included Cedusa) was the most basal group of Derbidae.  Fennah’s classification was functionally similar to Muir’s, although genera were rearranged.

Family Derbidae
Subfamily Derbinae
Tribe Zoraidini
Tribe Sikaianini
Tribe Otiocerini
Tribe Rhotanini
Tribe Derbini
Tribe Cenchreini

Subsequently, Broomfield (1985) modified the classification of Derbidae as follows, describing the new tribe Mysidiini.  The Mysidiini were revised with 6 genera and 136 species described, with keys to genera and species and amply illustrated.

Emeljanov (1992, 1996) subsequently revised higher taxa, sinking Mysidiini under Derbini (now a subtribe) and creating a series of new tribes, using a sequence of new features (some elaborate), including the development of a antennal cup (along with associated details), the enlarged antennal pedicle (of males), and the presence and location of sensory pits on the imago.  Emeljanov 1992 described Vinatini, Cedusini, Cedusinae and Nicertini; and Emeljanov (1996) described 9 new tribes, but recognized only 2 subfamilies (Derbinae and Otiocerinae); he also gave a key to tribes, that some potentially challenging diagnostic features such as the presences of stridulatory structures on the hind wing (e.g., on the jugal margin).   Yang and Wu (1994) followed Emeljanov (1992), but Emeljanov (1996) did not place some of Yang & Wu’s (1994) new genera in his subtribes of Zoraidini.

Emeljanov & Fletcher (2004: 40) moved Breddiniolini Fennah, 1950 from the Achilidae to the Derbidae (to Cedusinae).  Emeljanov (1990) also described the fossil tribe Ptychoptilini, moved from the Achilidae by Szwedo (2008). Szwedo (2005, 2006, 2008) presented some modifications to Emeljanov’s (1996) arrangement (esp. to Otiocerines), as did Emeljanov (2008). [I’ll summarize features used and classificatory changes beginning with Emeljanov 1996 in the future]

Cladogram from Emeljanov (1996)

Higher taxonomy of Derbidae (from FLOW) as of Jan 2018

Family Derbidae Spinola 1839: 205
= Derboides Spinola 1839: 205 (original spelling)

Subfamily Cedusinae Emeljanov 1992
= Cedusini Emeljanov 1992; status (subfamily) by Emeljanov 1996
Tribe Breddiniolini Fennah 1950: 36 (Transferred from Achilidae by Emeljanov & Fletcher 2004)

Breddiniola Muir, 1934: 581[Africa]
Breddiniolella Fennah, 1950: 37 (fiji)
Hemielissum Emeljanov & Fletcher, 2004 (Australia)

Hemielissum Emeljanov & Fletcher, 2004 (Australia)

Figures 1-5 from Emeljanov & Fletcher, 2004

Tribe Cedusini Emeljanov 1992
Subtribe Cedusina Emeljanov 1992 (sensu Emeljanov 2008)
Cedusa Fowler, 1904
Emeljanovedusa† Szwedo, 2006
Hauptenia Szwedo, 2006
Malenia Haupt, 1924
Muiredusa Szwedo, 2006
Produsa Szwedo, 2006

Cedusa sp. (all photographs and drawings by Kimberley Shropshire, University of Delaware)

Subtribe Eocenchreina Emeljanov 1992
Anticedusa Emeljanov, 2008 (1 species, Costa Rica)
Eocenchrea Muir, 1913 (=Eocenchres, missp. by Emeljanov 2006: 82)
Melusa Emeljanov, 1995

Anticedusa dominicana

Anticedusa dominicana from Emeljanov 2008

Tribe Goneokarellini Emeljanov 2006: 77
Goneokarella Fennah, 1952

Tribe Ipsnolini Emeljanov 1996: 76
Ipsnola Signoret, 1885

Ipsnola from Emeljanov 1996

Ipsnola from Emeljanov 1996

Ipsnola and Phrygia from Emeljanov 1996

Ipsnola and Phrygia from Emeljanov 1996

Tribe Phrygiini Emeljanov 1996: 76 (genus placed variously in Achilidae and Derbidae)
Phrygia Stål, 1856 (1 species, Brazil)

Tribe Ptychoptilini† Emeljanov, 1990
Ptychogroehnia† Szwedo & Stroinski, 2001
Ptychoptilum† Emeljanov, 1990

Tribe Vinatini Emeljanov 1992
Anerana Emeljanov, 1992
Vinata Distant, 1906 (= Erana Walker 1857, preoccupied, replaced by Vinata Distant, 1906; = Feredayia Kirkaldy, 1910 synonym of Vinata Distant, 1906 by ???)

Family Derbinae Spinola 1839
Tribe Cedochreini Emeljanov, 1996: 82[1]

Cedochrea Emeljanov, 1996 (1 species, Costa Rica)
Cedochrusa Emeljanov 2008

Cedochrusa luculenta

Cedochrusa luculenta from Emeljanov 2008

Cedochrea costariciana from Emeljanov 1995

Cedochrea costariciana from Emeljanov 1995

Tribe Cenchreini Muir, 1917: 54
= Cenchreinae Muir, 1917; status (tribe) by Muir, 1918.
Aethocauda Williams, 1976
Agoo Bahder & Bartlett 2019
Amania Synave, 1973
Anchimothon Fennah, 1952
Basileocephalus Kirkaldy, 1906 (= Aulacocephala Perroud & Montrouzier, 1864 nomen praeoccupatum replaced by Aculocephala Metcalf, 1945; Aculocephala Metcalf, 1945 synonym of Basileocephalus Kirkaldy, 1906 according to Fennah 1969; = Urabunna Distant, 1907 synonym of Basileocephalus Kirkaldy, 1906 by Kirkaldy 1907: 163)
Cenanges Fennah, 1952
Cenchrea Westwood, 1840
Contigucephalus Caldwell, 1944
Copallinges† Szwedo, 2004
Cyclometopum Muir, 1913
Dawnaria Distant, 1911
Fordicidia Distant, 1917
Goneokara Muir, 1913
Herpis Stål, 1862 (= Syntames Fowler 1905: 138)
Lamenia Stål, 1859
Muirileguatia Metcalf, 1945
Neocenchrea Metcalf, 1923
Neolamenia Muir, 1917
Omolicna Fennah, 1945
Oropuna Fennah, 1952
Perandenina Distant, 1911
Persis Stål, 1862
Phaciocephalus Kirkaldy, 1906
Vekunta Distant, 1906

Omolicna cf mcateei (by Kimberley Shropshire, University of Delaware)

Fordicidia robusta

Fordicidia robusta from Locker et al 2009

Vekunta bri

Vekunta bri from Locker et al 2009

Tribe Dawnarioidini Emeljanov 1996: 76; 2006: 84
Dawnarioides Dozier, 1929
Neodawnaria O’Brien, 1982

Tribe Derbini Spinola 1839
= Mysidiini Broomfield 1985: 5; syn by Emeljanov ….
Subtribe Derbina Spinola, 1839
Derbe Fabricius, 1803

Subtribe Mysidiina Broomfield, 1985
Amysidiella Broomfield, 1985
Dysimia Muir, 1924
Dysimiella Broomfield, 1985
Ipsemysidia Broomfield, 1985
Mysidaloides Broomfield, 1985
Mysidia Westwood, 1840
Neomysidia Broomfield, 1985
Paramysidia Broomfield, 1985
Pseudomysidia Metcalf, 1938
Symidia Muir, 1918


Tribe Nicertini Emeljanov 1992
Nicerta Walker, 1857

Subfamily Otiocerinae Muir, 1917
= Subfamily Zoraidinae Muir 1918b: 173 (as subfamily and tribe)[2]
Tribe Aquaeliciini Banaszkiewicz & Szwedo, 2005
Aquaelicium Distant, 1917
Ileifea Banaszkiewicz & Szwedo, 2005
Muiravea Banaszkiewicz & Szwedo, 2005
Ravola Banaszkiewicz & Szwedo, 2005
Razanus Banaszkiewicz & Szwedo, 2005
Synavea Emeljanov, 1995
Vizimbum Banaszkiewicz & Szwedo, 2005

Aquaelicium typicum (lectotype) from Locker et al 2009

Aquaelicium typicum (lectotype) from Locker et al 2009; scalebar applies to figures A–B.

Aquaelicium brunnescens (paralectotype)

Aquaelicium brunnescens (paralectotype) from Locker et al 2009


Tribe Kamendakini Emeljanov 2006: 87
Kamendaka Distant, 1906 (= Nicertoides Matsummura 1910: 14 (subgenus), = Eosaccharissa Kirkaldy 1907: 125 (subgenus))

Tribe Neocyclokarini Emeljanov 2006: 89
Neocyclokara Muir, 1917

Tribe Otiocerini Muir, 1917: 54
= Otiocerinae Muir, 1917: 54; status (tribe) by Muir 1918: 229; returned to subfamily by Broomfield 1985 (also subfamily in Em., 2006)
Anomaladerbe Muir, 1922
Anotia Kirby, 1821 (= Amalopota Van Duzee 1889: 176)
Apache Kirkaldy, 1901
Archara Metcalf, 1945
Banksiella Muir, 1917
Cobacella Fennah, 1952
Dendrokara Melichar, 1914
Deribia Westwood, 1841
Eusyphax Fennah, 1956
Flaccia Stål, 1866
Harpanor Fennah, 1950
Heronax Kirkaldy, 1906
Kaha Kirkaldy, 1906
Kampulokara Muir, 1913
Kubilaya Koçak & Kemal, 2010 (=Iquitosa Fennah, 1945 nomen praeoccupatum replaced by Kubilaya Koçak & Kemal, 2010)
Kuranda Distant, 1907
Labicerus Erichson, 1848
Lugeilangor† Szwedo, 2005
Makula Distant, 1907
Mysidioides Matsumura, 1905
Neodendrokara Muir, 1917
Nesocore Kirkaldy, 1907
Nesokaha Muir, 1913
Nesoneura Kirkaldy, 1907
Nesoniphas Kirkaldy, 1907
Niphadodite Kirkaldy, 1907
Paralyricen Muir, 1913
Pepleuca Emeljanov, 1999 (= Platocera Muir, 1913 synonym of Pepleuca Emeljanov, 1999 by ??)
Phra Distant, 1911
Platocerella Fennah, 1952
Platonax Metcalf, 1938
Positrona† Emeljanov, 1994
Pyrrhoneura Kirkaldy, 1906
Pyrrhonice Emeljanov, 1994
Sayiana Ball, 1928
Shellenius Ball, 1928
Swezeyia Kirkaldy, 1906
Tempora Matsumura, 1914

Anotia westwoodi

Subtribe Nicertina Emeljanov, 1992
Epotiocerus Matsumura, 1914
Interamma Walker, 1870
Leptaleocera Muir, 1913
Megatropis Muir, 1913
Nicerta Walker, 1857
Robigus Distant, 1911
Vivaha Distant, 1906

Subtribe Otiocerina Muir, 1917
Otiocerus Kirby, 1821

Otiocerus coquebertii Kirby, 1821

Otiocerus coquebertii

Tribe Patarini Emeljanov 2006: 89 (redefined Banaszkiewicz & Szwedo, 2005)
Anapatara Emeljanov, 1995
Patara Westwood, 1841

Patara vanduzei (photographs by Kimberley Shropshire, University of Delaware)

Tribe Phenicini Emeljanov 2006: 92
Equirria Distant, 1917
Fescennia Stål, 1866
Metaphenice Emeljanov, 1995
Paraphenice Muir, 1924 (=Imbalara Hesse, 1925 synonym of Paraphenice Muir, 1924 by Muir 1928: 502)
Phenice Westwood, 1840

Paraphenice aurea

Paraphenice aurea from Locker et al 2009

Tribe Rhotanini Muir, 1918: 228-229
= Rhotaninae Muir, 1918; status (tribe) by Muir, 1918; RESTORED to subfamily be Broomfield 1985; reduced to tribe in Emeljanov 1996.. see Zelazny & Webb (2011)
Alara Distant, 1911
Dichotropis Muir, 1913
Levu Kirkaldy, 1906
Muiralevu Zelazny, 1981 (=Formolevu Yang & Wu, 1993 synonym of Muiralevu Zelazny, 1981 according to Zelazny & Webb 2011: 55
Rhotana Walker, 1857 (= Decora Bierman, 1910, = Genestia Stål, 1858)
Rhotanella Fennah, 1970
Saccharodite Kirkaldy, 1907
Sumangala Distant, 1911

Alara hyalina

Alara hyalina from Zelazny 1981

Derbidae forewing venation from Zelazny 1981

Derbidae forewing venation from Zelazny 1981

Rhotana species from Zelazny & Webb-2011

Figures 74-77 from Zelazny and Webb 2011; top row Rhotana pellax, bottom Rhtana semiopalina

Tribe Sikaianini Muir 1917: 56
Ceropupa Emeljanov, 1995
Distantinia Muir, 1917
Leomelicharia Muir, 1913
Muiria Kirkaldy, 1907
Mula Ball, 1928 but placed in Patarini elsewhere
Sikaiana Distant, 1907 (= Euklastus Metcalf 1923)

Sikaiana albomaculata

Sikaiana albomaculata from Locker et al 2009

Sikaiana harti

Sikaiana harti

Tribe Zoraidini Muir 1918b: 173 (
= Subfamily Zoraidinae Muir 1918: 173; status (tribe) by Fennah 1952 (subfamily not indicated); status (subfamily) by Broomfield (1984: 4); status (tribe of Otiocerinae) by Emeljanov (2006: 92)/

Subtribe Lyddina Emeljanov 2006: 92
Acanthocerana Metcalf, 1945 (doubtfully included Em. 2006)
Diostrombus Uhler, 1896
Formodanga Yang & Wu, 1993
Helcita Stål, 1856
Lydda Westwood, 1840
Lyddastrombus Van Stalle, 1992
Neoproutista Yang & Wu, 1993
Pamendanga Distant, 1906
Proutista Kirkaldy, 1904
Shizuka Matsumura, 1914


Proutista moesta from Locker et al 2009

 Proutista moesta (From Locker et al 2009): A–D habitus; E head; F body lateral. Scalebar 1 mm applies to figures B–D.

Subtribe Zeugmatina Emeljanov 2006: 92 (Emeljanov, 1995)
Zeugma Westwood, 1840

subtribe Zoraidina Muir 1918: 173 (status by Emeljanov 2006: 93; constituent genera from Metcalf less genera specified in Emeljanov 2006)
Losbanosia Muir, 1917
Monochorhynchus Muir, 1917
Neocamma Melichar, 1915
Neodiostrombus Muir, 1918
Neozoraida Muir, 1918
Parapamendanga Yang & Wu, 1993
Parapeggia Yang & Wu, 1993
Peggia Kirkaldy, 1901
Peggiopsis Muir, 1913
Pseudohelcita Muir, 1918
Raizoda Muir, 1924
Shirakiana Metcalf, 1945
Stenopeggia Fennah, 1952
Teutberga Jacobi, 1917
Zorabana Van Stalle, 1984
Zoraida Kirkaldy, 1900
Zoraidoides Distant, 1914

Diprora Williams, 1976
Nesorhamma Fennah, 1956
Stenogeggia Fennah, 1952 (may just be a misspelling of Stenopeggia)

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Original publication not found

Cedusa ussurica (Anufriev, 1968)

Malenia turkestanica Dubovsky, 1965


[1] Pagination based on English version of text.

[2] As a subfamily, includes Zoraidini and Sikaianini.

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