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 the 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 (and Rhotana) groups.  In 1917, Muir formalized these into subfamilies, except groups 3 and 4 were combined but excluded Rhotana, and the groups were formalized into the 1. Cenchreinae, 2. Otiocerinae, 3. Derbinae (Derbini including Zoraida, Sikaianini), and 4. Rhotaninae, and a key to subfamilies (and genera in the Philippines) 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.

Family DERBIDAE
Subfamily ZORAIDINAE
Tribe ZORAIDINI
Tribe SIKAIANINI
Subfamily DERBINAE
Tribe DERBINI
Tribe RHOTANINI
Tribe CENCHREINI
Tribe OTIOCERINI

Higher taxonomy was entirely revised and reinterpreted by Fennah (1952) (using 6 tribes, but not specifying subfamilies). 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 (mostly using sensory pits on the body and forewing venation).  He utilized 3 subfamilies – Derbinae, Otiocerinae (= Zoraidinae) and Cedusinae.  Emeljanov sunk Mysidiini under Derbini (now a subtribe) and creating a series of new tribes (viz. Ipsnolini, Goneokarellini, Phrygiini, Vinatini, Cedusini (including subtribe Eocenchreina), Cedochreini, Dawnarioidini, Nicertini, Kamendakini, Patarini, Neocyclocarini and Phenicini), using a sequence of new features (some elaborate), including the development of an 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.

Cladogram from Emeljanov (1996)

Emeljanov & Fletcher (2004: 40) moved Breddiniolini Fennah, 1950 from the Achilidae to the Derbidae (to Cedusinae).  Emeljanov (1990) also described the Eocene 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).

Emeljanov & Shcherbakov (2020) described the new tribe Derbachilini Emeljanov & Shcherbakov 2020† including 2 species in 2 genera from mid-Cretaceous Burmese amber.  They revised the status of Cedusinae Emeljanov, 1992, observing that Breddiniolini Fennah, 1950 should have priority as Breddiniolinae Fennah, 1950. Because this work is recent, I am still working to understand the implications of these changes.

A commentary on higher-level features is adopted (abridged and lightly edited) below directly from Emeljanov & Shcherbakov (2020)

The Derbinae was separated from Cedusinae by the presence of sensory pits in the adult (Emeljanov 1996). The presence of sensory pits on the head, as well as the costa and clavus in the forewing, was the only criterion for distinguishing Cedochreini and Cenchreini (Derbinae) from Cedusini (Cedusinae): the pits are absent in Cedusini as opposed to present on the basal part of the costa in Cedochreini, and on the head, the costa and Pcu in Cenchreini (Emeljanov, 1992, 1996). However, there are exceptions: in some Cedochreini the pits are developed also at the scutellar margin of clavus (Cedochrusa Emeljanov, 2008; Emeljanov, 2008a, fig. 12), and vice versa, the pits on tegmen are restricted to the basal part of the costa also in some Cenchreini (Dawnaria girdlestoni (Muir, 1913)), Nicertini (Leptaleocera coccinea Muir, 1913) and Otiocerini (Paralyricen jepsoni Muir, 1913; Platocera annulipes Muir, 1913, Pyrrhoneura immaculata Muir, 1913, etc.) [Muir, 1913, pl. 3].

Another important set of character changes at the transition from the basal derbid tribes to Derbinae is associated with the apex of clavus, postclaval area and postclaval crossvein. Achilidae and the most primitive, still achilid-like derbids (grade A) have tegmina not as long as the body, their postnodal parts overlapping in repose (often short) and sometimes with extra folding, the apex of the clavus transversely or obliquely truncate, CuP sharply bent posteriorly near the apex, Pcu+1A joining CuP there, CuA2 reaching apical margin, claval furrow continued into long and wide postclaval area, and postclaval crossvein long and transverse. Somewhat more derived derbids that had lost the achilid habitus (grade B), have the tegmina longer than the body length, steeply tectiform in repose, CuP nearly straight apically, and Pcu+1A joining either CuP near the apex (Cedusini Eocenchreina, Cedochreini, some Cenchreini—e.g. Goneokara Muir, 1913, Agoo Bahder et Bartlett, 2019), or 2A or their junction (cixiid condition restored: Cedusina, Dawnarioidini, other Cenchreini—e.g. Basileocephalus Kirkaldy, 1906, Dawnaria Distant, 1911). Further transformations of the claval region occur in still more derived derbids: CuP after the junction with Pcu+1A running along posterior margin up to postclaval crossvein, clavus open, postclaval area short and narrow, CuA2 joining posterior margin, postclaval crossvein short and oblique. This condition recorded already in a few Cenchreini (some Phaciocephalus Kirkaldy, 1906, Herpis Stål, 1862), is further developed in higher Derbidae (Fennah, 1952). The characters that unite all Derbidae are the fusion of tergite IX with the anal tube in males, and probably also peculiar wax pores on the abdomen. In the evolution of Derbidae, there was a transformation of the basal, achilid-like forms into higher derbids through the accumulation of derived characters. Accordingly, the tribes of Derbidae were arranged in a phylogenetic sequence [Emeljanov, 1995]. The character changes that have occurred from primitive tribes to typical derbids are as follows (described as ‘derbization of achilids’):

(1) shortening of the apical segment of rostrum—at least thrice as long as wide in Breddiniolini, Ipsnolini, Goneokarellini, Vinatini, Ptychoptilini, Derbachilini; at most twice in other derbids (except for Archara Metcalf, 1945 of Otiocerini) [Fennah, 1952];

(2) reduction of lateral teeth on hind tibia—four or three teeth in Breddiniolini; three or two in Derbachile gen.n.; two in Vinatini; one tooth in Ipsnolini, Goneokarellini, Phrygiini and Eocenchrea Muir, 1913 (Cedusini Eocenchreina); no teeth in other derbids, except Zoraidini and Sikaianini;

(3) reduction of subapical setae on apical pectens of hind tarsomeres — setae present in Breddiniolini and Ipsnolini; absent in Derbachile and other derbids;

(4) medial shortening of the pronotum and reduction  of its disc—pronotal disc elevated, tricarinate, triangular, nearly as long as wide in Ipsnolini, Vinatini, Ptychoptilini, Derbachile; much shorter in Goneokarellini, Phrygiini, some Cenchreini; not elevated, ecarinate in other derbids;

(5) reduction of M fork in the hind wing — M forked before r-m in Breddiniolini, Derbachilini; beyond r-m in Ipsnolini, Phrygiini, Cedochreini; simple in other primitive derbids;

(6) reduction of CuA1 fork in the hind wing — CuA1 forked before m-cu in Breddiniolini; beyond m-cu in Ipsnolini, Goneokarellini, Phrygiini, Phrygiini, Derbachilini; simple in other derbids;

(7) modification of antenna — enlarged and somewhat flattened in Derbachilini trib.n.; enlarged in Vinatini, Derbini, and flattened in some Otiocerinae;

(8) development of subantennal lobe — ridge at lorogenal boundary in Phrygiini [Emeljanov, 1995]; low ridge in Derbachile hochae; lobe in Cedusini, Cedochreini, Cenchreini, most Otiocerinae;

(9) shortening of ScRA in the hind wing—in Cedusini (except Emeljanovedusa Szwedo, 2006 from Baltic amber) and higher tribes;

(10) increase of wing/body size ratio — tegmen length greater than body length in Breddiniolini, Derbachilini, and in Cedusini and higher tribes;

(11) loss of tegminal overlap and truncate apex of clavus—tegmina overlapping, clavus truncate in Breddiniolini, Ipsnolini, Goneokarellini, Phrygiini, Vinatini, Ptychoptilini and Derbachilini; tegmina not overlapping, clavus more acute or open in Cedusini and higher tribes;

(12) development of sensory pits in adults—on costa in tegmen (sometimes on its base only) in Derbachilini, Dawnarioidini, and some Cedochreini, Cenchreini, Nicertini and Otiocerini; on C and/or other veins and/or head in higher tribes.

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

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

Subfamily Breddiniolinae Fennah, 1950 (status revised by Emeljanov & Shcherbakov 2020)
= Cedusinae Emeljanov 1992, syn by Emeljanov & Shcherbakov 2020: 238. 
= 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)

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

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

Tribe Derbachilini Emeljanov & Shcherbakov 2020†
Derbachile Emeljanov & Shcherbakov 2020†
Achiderbe Emeljanov & Shcherbakov 2020†

Tribe Goneokarellini Emeljanov 2006: 77
Goneokarella Fennah, 1952

Tribe Ipsnolini Emeljanov 1996: 76
Ipsnola Signoret, 1885

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 ???)

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

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

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

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; scalebar applies to figures A–B.

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

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
Otiocerus Kirby, 1821
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

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

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

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

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)

Tribe Zoraidini Muir 1918b: 173 (http://biodiversitylibrary.org/page/34176021)
= 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

 

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

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

References [Incomplete, borrowed freely from FLOW]

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The 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.