Classification of the Nogodinidae

The Nogodinidae were first recognized as a tribe of Ricaniidae by Melichar (1898) and subsequently recognized as either tribe or subfamily (in Melichar 1906) within Ricaniidae.  Kirkaldy (1907: 9, 93) established Bladinini as a group within Ricaniidae, subsequently included in the Nogodinidae.  Muir (1930) first recognized Nogodinidae as a family. Muir segregated Nogodinidae (from Ricaniidae) on the basis of having two spines on the second hind tarsomere, front longer than wide, and having lateral carinae on the clypeus. Melichar (1923) gives keys and descriptions of 35 genera, the arrangement of which may have formed the basis for subsequent groupings, and Metcalf (1938) gives a key to 6 Neotropical genera.  Metcalf (1954) – the Metcalf catalog – recognized no subfamily groupings (including Bladinini, despite recognizing Bladina as a genus of Nogodinidae).

 

Key to New World genera of Nogodinidae from Metcalf (1938)

Key to New World genera of Nogodinidae from Metcalf (1938)

Fennah (1956) provided a key to planthopper families, that (in effect) redefined Nogodinidae.  Fennah (1967) erected Mithymnini for the South African genera Xosias Kirkaldy, Mithymna Stal, Colmadona Kirkaldy and Monteira Melichar (transferred from lssidae). The first formal classification of Nogodinidae was presented by Fennah (1978), revised by Fennah (1984, 1987), who included Elicaini  Melichar (1915: 379) corrected to Elicini by Fennah 1978: 118) in Nogodinidae.

Fennah’s (1978) arrangement was:

Subfamily Nogodininae
Tribe Nogodinini
subtribe Nogodinina – Nogodina, Varciopsis, Biolleyana, Orthothyreus and Neovarcia
subtribe Vutinina – Vutina.
Tribe Epacriini Gerstaecker, 1895 – Philbyella, Privesomorphus, Epacria, Colpocara, Criopaca, Diazanus, Morsina and Psiadiicola
Tribe Bladinini
subtribe Gaetuliina – Gaetulia, Nurunderia, Indogaetulia, Salona, Pucina, Laberia, Paralasonia and provisionally Riancia, Exphora and Siopaphora.
subtribe Elicina – Elica and Canna
subtribe Bladinina – Bladina
Tribe Pisachini – Pisacha, Soaemis and Goneopsara
Tribe Varciini
subtribe Varciina – Convarcia, Detya, Miriza, Paravarcia, Probletomus, Varcia and Varciella.
subtribe Sassulina – Aphanophrys, Decoomana, Mindura, Paradetya and Sassula.

Fennah (1984) redescribed Nogodinidae, redefined Gaetuliina to include a list of genera transferred from Issidae: Acrisius, Numithia, Dictidea, Dictyobia, Dictyonia, Osbornia, Neaethis, Dictyonyssus, Misodema, Dictyssa, Danepteryx (from the Nearctic) and Gamergus, Gamergomorphus, Paragamergomormorphus, Johannesburgia and Mangola (from Africa); Alleloplasis and Bilbilis (from Australia). He also described tribe Lipocalliini (for Lipocallia and Bilbilicallia). redefined Mithymnini (to include Mithymna, Xosias, Telmessodes, Telmosias, Colmadona and tentatively Stilpnochlaena), and Monteira, Afronias, Kiomonia, Gamergomimus (nov) to Epacriini.  Fennah (1987) described the subfamily Gastriniinia for Gastrinia transferred from Issidae.

Petrulevicius, (2005) described the extinct tribe Celinapterixini for the new genus Celinapterix Petrulevicius, 2005. Gnezdilov (2007) moved Tonginae from Acanaloniidae (as placed by Emeljanov 1999) into Nogodinidae and Gnezdilov (2012) moved Colpopterini Gnezdilov 2003 into Nogodinidae as a subfamily.

Gnezdilov (2017a) revised the higher classification of Nogoninidae and presented a key to higher groupings (presented below).  He described the new tribe Bilbiliini (for Bilbilis) In the Nogodininae, raised Bladinini to subfamily (Bladinini) and subsumed Tonginae as a tribe of Nogodininae.

Quantitative phylogenetic systematics effort that have included Nogodinidae have been limited.  There are three studies of family-level phylogenetic studies that have included multiple taxa of Nogodinidae (Urban & Cryan 2007, Song & Liang 2013, Gnezdilov et al. 2015).  All of these have suggested that Nogodinidae are para- or polyphyletic, but none of these were undertaken to examine the monophyly of Nogodinidae and the taxon sampling, while indicative, is too sparse to draw conclusions.

Phylogeny of Fulgoroidea from Urban and Cryan (2007); note polyphyly of Nogodinidae.

Maximum-likelihood tree Song & Liang

Figure 1. Maximum-likelihood tree estimated from the combined datatset of 65 taxa of Fulgoroidea. Values above branches denote ML bootstrap support ($70). The red numbers in circles correspond to ten major clades discussed in the text. doi:10.1371/journal.pone.0058400.g001

The 50% majority-rule consensus tree, Bayesian, from Song and Liang 2013

Figure 2. The 50% majority-rule consensus tree estimated from the Bayesian analysis based on the combined dataset. Values above branches denote posterior probabilities ($0.9). doi:10.1371/journal.pone.0058400.g002

 

Phylogeny of Issidae from Gnezdilov et al. (2015), note paraphyly of Nogodinidae.

Key to subfamilies and tribes of Nogodinidae (modified from Gnezdilov 2017a)
1(2) Third segment of rostrum twice as short as second one, not narrowing apically (Fig. 2). First metatarsomere with peculiar oval area of short dense setae on its ventral surface (Fig. 1). Anterior connective lamina of gonapophyses VIII wide, with 5 large flat teeth (Fig. 9) … Subfamily Bladininae Kirkaldy, 1907 (tribe Bladinini)

2(1) Third segment of rostrum as long as second one or slightly shorter, conically narrowing apically (Fig. 4). First metatarsomere with long, usually hair-shaped setae ventrally (Fig. 3). Anterior connective lamina of gonapophyses VIII narrow (Fig. 7), if wide (Fig. 13) than with another shape and arrangement of teeth … 3

3(4) Paradiscal field / paranotal lobe border of pronotum in shape of distinct carina and each paranotal lobe with another distinct transverse carina below this border. Fore wing without precostal area (Fig. 50). Gonoplacs long and narrow, large anal tube clasping the gonoplacs dorsally and laterally (Fig. 51) … Subfamily Gastriniinae Fennah, 1987 (tribe Gastriniini)

4(3) Paradiscal field / paranotal lobe border of pronotum and paranotal lobes without transverse carinae. Fore wing with precostal area. Gonoplacs wide or beak-shaped and curved … 5

5(6) Fore wing with rich transverse venation usually only in it s distal half (Gnezdilov,
2012, figs 4, 35). Hind wing well developed, with simple venation (R and CuA with two branches, M simple or with 2 branches and Pcu simple) (Gnezdilov, 2012, figs 15, 36) … Subfamily Colpopterinae Gnezdilov, 2003 (tribe Colpopterini)

6(5) Fore wing usually with rich transverse venation on whole area (Fig. 23). Hind wing rudimentary or well developed, with R, M, and CuA with two and more branches and Pcu simple (Figs 33, 34) or if M simple than Pcu with two branches (Gnezdilov & Constant, 2014, figs 3, 18) … Subfamily Nogodininae Melichar, 1898

7(8) Forewing with precostal margin hemicircularly protruding below the eye (in  lateral view) (Fig. 38, hp). First metatarsomere with only 2 latero-apical spines and without intermediate spines … Tribe Lipocalliini Fennah, 1984

8(7) Fore wing with precostal margin not protruding below the eye (in lateral view). First metatarsomere with 2 latero-apical spines and several intermediate spines.

9(14) First metatarsomere with straight row of intermediate spines.

10(11) Macropterous, fore wing with wide precostal area. Hind wing well developed, with first and second anal veins not fused medially and second anal vein with 2 branches (A2 2) (Fig. 33). Gonoplacs flattened laterally … Tribe Nogodinini Melichar, 1898

11(10). Subbrachypterous, fore wing with more narrow precostal area. Hind wings rudimentary or reduced. Gonoplacs convex … 12

12(13) Pronotum with anterior margin strongly protruding and reaching anterior margin of eyes (in dorsal view) (Fig. 25). Gonoplacs hemisphaerical (Fig. 10, 48) … Tribe Bilbiliini Gnezdilov

13(12) Pronotum with anterior margin less protruding, not reaching anterior margin of eyes (in dorsal view) (Fennah, 1967, fig. 24B). Gonoplacs convex or cylindrical, each with peculiar invagination (Gnezdilov, 2017a, fig. 8) … Tribe Mithymnini Fennah, 1967

14(9) First metatarsomere with intermediate spines arranged in shape of “V” letter (Fig. 3).

15(20) Gonoplacs convex or nearly cylindrical, with denticles or teeth. Hind wing with first anal vein 3-branched (A1 3) with its posterior branch (A1.3) fused on a long distance with second anal vein which is simple (A2 1) (Fig. 32).

16(19) Metope often with sublateral carinae converging above metopoclypeal suture in shape of “V” letter (Meng et al., 2014, fig. 31). Gonoplacs convex or nearly cylindrical, with thicken distal parts.

17(18) Gonoplacs with dense rows of teeth on its thicken distal parts which are situated nearly perpendicular to lateral sides of gonoplacs (Meng et al., 2014, figs 32, 33). Apices of anterior connective laminae of gonapophyses VIII are well visible above the gonoplacs … Tribe Pisachini Fennah, 1978

18(17) Gonoplacs with rudimentary areas of denticles on its thicken distal parts which are mainly smooth. Apices of anterior connective laminae of gonapophyses VIII not visible above the gonoplacs … Tribe Varciini Fennah, 1978

19(16) Metope with sublateral carinae converging on upper margin of metope in shape of inverted “V” letter (Chan & Yang, 1994, figs 31B, 32B) or horse-shoe-shaped (Fennah, 1954, fig. 7A) or without carinae (Gnezdilov & Constant, 2014, figs 4, 13). Gonoplacs flat … Tribe Tongini Kirkaldy, 1907

20(15) Gonoplacs flattened laterally, nearly triangular, without denticles or teeth (Gnezdilov, 2017b, fig. 15). Hind wing (if not reduced) with first and second anal veins separated and second anal vein simple (A2 1) (Gnezdilov, 2017a, fig. 13; Gnezdilov, 2017b, figs 4, 18) … Tribe Epacriini Fennah, 1978

Family Nogodinidae Melichar, 1898
Subfamily Nogodininae Bladininae Kirkaldy 1907: 9, 93
Tribe Bladinini Kirkaldy 1907: 9, 93
Subtribe Bladinina Kirkaldy 1907 (sensu Fennah 1978: 117, 118)
Bladina Stål, 1859

Bladina fowleri from Belize (Photograph by Kimberley Shropshire, University of Delaware)

Subfamily Colpopterinae Gnezdilov, 2003
= Colpopterini Gnezdilov 2003; status (subfamily) and transfer to Nogodinidae (from Issidae) by Gnezdilov (2012: 757).
Tribe Colpopterini Gnezdilov 2003.
Bumerangum Gnezdilov, 2012 (Africa)
Caudibeccus Gnezdilov & O’Brien, 2008 (Neotropical)
Colpoptera Burmeister, 1835 (Neotropical) (= Leptophara Stål, 1869 nomen praeoccupatum replaced by Hesperophara Kirkaldy, 1904: 279; = Hesperophara Kirkaldy, 1904 synonym of Colpoptera Burmeister, 1835 by Gnezdilov & O’Brien 2008: 21).
Dozierana Gnezdilov, 2018 (Type species Thionia gibbicollis Dozier, 1931, 1 species, Haiti)
Jamaha Gnezdilov & O’Brien, 2008 (Neotropical)
Neocolpoptera Dozier, 1931 (Neotropical)
Ugoa Fennah, 1945 (Neotropical)

Colpoptera sp. adventive to Florida.

Subfamily Gastriniinae Fennah 1987: 363
Tribe Gastriniini Fennah 1987 (tribe by implication)
Gastrinia Stål, 1859 (= Gastrinia Stål, 1859 nomen praeoccupatum replaced by Amfortas Kirkaldy, 1904: 280; = Amfortas Kirkaldy, 1904 nomen nudum synonym of Gastrinia Stål, 1859 nomen revivisco by Metcalf 1938: 385)

Gastrinia vaginata Stal

Gastrinia vaginata Stal from Gnezdilov (2017a)

Subfamily Nogodininae Melichar, 1898
Tribe Bilbiliini Gnezdilov, 2017
Bilbilis Stål, 1861 (from Mithymnini by Gnezdilov 2007: 296)

 

Bilbiliini

Nogodinidae. 35-37-Bilbilis modestus (Stål), holotype: 35-dorsal view; 36-lateral view; 37-type labels; 38-Bilbilicallia truncatella (Walker), female, holotype, “New Holl.” (BMNH), fore wing. Abbreviation: hp-hemicircularly protruding costal margin of forewing (Gnezdilov 2017).

 

Bilbilini

Nogodinidae. 45-47-Bilbilis elongatus Gnezdilov, 2017, holotype: 45dorsal view; 46-frontal view; 47-lateral view; 48-B. dorsalis (Walker), holotype, ovipositor (From Gnezdilov 2017)

Tribe Celinapterixini† Petrulevicius, 2005
Celinapterix† Petrulevicius, 2005

Tribe Epacriini Fennah 1978: 114
Afronias Fennah, 1967
Colpocara Bergroth, 1920
Criopaca Schmidt, 1918
Diazanus Hesse, 1925
Epacria Gerstaecker, 1895
Gamergomimus Fennah, 1984
Kiomonia Schmidt, 1911
Mangola Melichar, 1906
Monteira Melichar, 1906 (= Monteirina Lallemand, 1931 synonym by Synave 1957: 66)
Morsina Melichar, 1902
Paramangola Synave, 1956
Philbyella China, 1938
Privesomorphus Schmidt, 1912
Psiadiicola Fennah, 1978
Rileyopsis De Bergevin, 1917

Tribe Lipocalliini Fennah 1984: 84
Bilbilicallia Jacobi, 1928 (Australia, &c)
Lipocallia Kirkaldy, 1906 (Australia, &c)

 

Lipocallia australensis

Lipocallia australensis Kirkaldy male syntype from New South Wales Government, Department of Primary Industries Biosecurity Collections.

Tribe Mithymnini Fennah 1967: 697
Bowesdorpia Synave, 1956 (to Mithymnini by Gnezdilov 2007: 296)
Colmadona Kirkaldy, 1901 (= Telmessus Stål, 1866 nomen praeoccupatum replaced by Colmadona Kirkaldy, 1901: 6)
Fovealvus Gnezdilov & Wilson, 2007
Mithymna Stål, 1862
Paralusanda Synave, 1956
Stilpnochlaena Fennah, 1967
Telmessodes Fennah, 1967
Telmosias Fennah, 1967
Xosias Kirkaldy, 1904 (= Eteocles Stål, 1866 nomen praeoccupatum replaced by Xosias Kirkaldy, 1904: 280)

Paralusanda sinuatipennis

Paralusanda sinuatipennis Synave from Gnezdilov 2017b

Tribe Nogodinini Melichar, 1898 see Schmidt 1912: 83
Subtribe Nogodinina Melichar, 1898 see Schmidt 1912: 83 (see Fennah 1978: 114)
Biolleyana Distant, 1909
Neovarcia Schmidt, 1919
Nogodina Stål, 1859
Orthothyreus Schmidt, 1919
Tainosia† Szwedo & Stroinski, 2001
Tonacatecutlius† Stroinski & Szwedo, 2000
Varciopsis Jacobi, 1916

Nogodinia reticulata lectotype

Type specimen of Nogodinia reticulata (Fabricius 1803) from the Fabricius collection.  Photo courtesy Niels Kristensen,Natural History Museum of Denmark, University of Copenhagen.

Nogodinidae from Tiskita Jungle Lodge in Costa Rica (Biolleyana sp. poss. costalis); photo by Brian Cutting, University of Delaware, Department of Entomology)

Subtribe Vutinina  Fennah 1978: 114
Vutina Stål, 1862

Tribe Pisachini Fennah 1978: 118
Goniopsara Metcalf, 1952 (= Goniopsis Melichar, 1899 nomen praeoccupatum replaced by Goniopsara Metcalf, 1952: 226)
Goniopsarites Meng, Wang & Wang 2014
Pisacha Distant, 1906 (= Soaemis Jacobi, 1916 by Ishihara 1965) ( = Soaemis Jacobi, 1916 synonym of Pisacha Distant, 1906 by Ishihara 1965: 207; Soaemis Jacobi, 1916 status revivisco by Fennah 1978: 118; Soaemis Jacobi, 1916 synonym of Pisacha Distant, 1906 by Meng et al. 2014: 87)

Tribe Varciini Fennah 1978: 118
Subtribe Varciina Fennah 1978: 118
Convarcia Schmidt, 1919
Detya Distant, 1906
Miriza Stål, 1862
Paravarcia Schmidt, 1919
Probletomus Gerstaecker, 1895
Varcia Stål, 1870 (= Aphanophrys Melichar, 1898, subgenus according to Melichar 1923; full genus by Fennah 1978)
Varciella Melichar, 1923

Subtribe Sassulina Fennah 1978: 118
Aphanophrys Melichar, 1898
Decoomana Lallemand, 1942
Mindura Stål, 1862
Paradetya Schmidt, 1919
Sassula Stål, 1870

Subfamily Tonginae Kirkaldy, 1907: 9 (see Baker 1927; to Nogodinidae from Issidae by Gnezdilov 2007 as comprised by Fennah 1954)
Tribe Tongini Kirkaldy, 1907 (by implication)
Andrewsiella Izzard, 1936 (from Acanaloniidae)
Atylana Melichar, 1906
Capelopterum Melichar, 1906
Chondroptera Bergroth, 1910
Cotylana Fennah, 1954
Distiana Metcalf, 1952 (= Moniana Distant, 1909 synonymby Metcalf 1952)
Epitonga Fennah, 1954
Forculus Distant, 1912
Forculusoides Distant, 1916
Glyphotonga Schmidt, 1910 ( = Neodelia Distant, 1914 synonym of Glyphotonga Schmidt, 1910 by Baker 1927; = Oxychara Melichar, 1923 synonym of Glyphotonga Schmidt, 1910 by Baker 1927)
Hemitonga Schmidt, 1911
[Issarius Metcalf, 1950 to incertae sedis]
Lollius Stål, 1866 ( = Okissus Matsumura, 1916 synonym by Gnezdilov 2009: 87; = Ecapelopterum Chan & Yang, 1994 synonym by Gnezdilov 2009: 87)
Orthophana Melichar, 1923
subgenus Eupharos Gnezdilov & Constant, 2014
subgenus Orthophana Melichar, 1923
Oryxana Distant, 1910 (= Issella Metcalf, 1952, = Buehleria Lallemand & Synave, 1953, = Issina Melichar 1906, preoccupied)
Paratonga Schmidt, 1910
Paratylana Melichar, 1906
Pseudotylana Melichar, 1906
Scalabis Stål, 1870
Tonga Kirkaldy, 1900 (= Cyrene Westwood, 1845, preoccupied, = Sutteria Lallemand & Synave, 1953)
Tylana Stål, 1861

Image of Andrewsiella oceanica Izzard 1936 from Fletcher, 2009

Cotylana acutipennis (Kirkaldy) (Male syntype, Cairns, Qld); New South Wales Government, Department of Primary Industries, Biosecurity Collections .

Distiana andrewsi (Kirby) New South Wales Government, Department of Primary Industries, Biosecurity Collections.

Orthophana (Eupharos) tamdaoina Gnezdilov and Constant 2014 from original publication

Tonga sp.

Atylana rarotongae Eyles & Linnavuori, 1974 (Image from the B. P. B. Museum, Cook Islands Biodiversity Website

Lollius kuroiwae (Matsumura, 1916) from Gnezdilov 2009 (Iriomote Island)

Nogodinidae incertae sedis
Aphaena rotundipennis† Scudder, 1878
Cotradechites† Fennah, 1968
Detyopsis† Cockerell, 1920
Dilaropsis† Cockerell, 1920
Eoricania† Henriksen, 1922
Ficarasites† Scudder, 1890
Giselia† Haupt, 1956
Hammapteryx† Scudder, 1890
Iranissus Dlabola, 1980 moved to Nogodinidae by Mozaffarian & Wilson 2011, not placed to subfamily or tribe.
Issarius Metcalf, 1950
Issidius Puton, 1898
Lithopsis† Scudder, 1878
Neoricania† Carpenter, 1990
Paralatindia saussurei† Scudder, 1890
Thaumastocladius† Cockerell & Sandhouse, 1921
Henriksenopterix† Petrulevicius, 2005
Tritophania† Jacobi, 1938

Transfers

Gamergus Stal, 1859 to Gaetuliini (Tropiduchinae) by Gnezdilov 2007
Agenia Descourtilz, 1825 see Issidae transferred from Tonginae by …? Agenia Descourtilz, 1825 (Fulgoroidea incertae sedis) according to Dlabola (1987): 62
Chimetopon Schmidt, 1910 see Issidae transferred from Tonginae by …? Issini Spinola, 1839
Chondroptera Bergroth, 1910 see Issidae transferred from Tonginae by …?
Mushya Kato, 1933 syn of Aphelonema Uhler, 1876 by Chan & Yang (1994) see Peltonotus Mulsant & Rey, 1855 (Caliscelidae)
Mijas Rambur, 1840 see Issidae transferred from Tonginae by …? Mijas Rambur, 1840 (Fulgoroidea incertae sedis) according to Dlabola (1987): 62

Tropiduchidae Stål, 1866 – Elicinae Melichar, 1915
Hemithiscia Schmidt, 1912 (2 species, Dutch east Africa – http://biodiversitylibrary.org/page/8824629)
Paraphilatis Melichar, 1912 (1 species, Congo) (http://biodiversitylibrary.org/page/5889788)
Parathiscia Melichar, 1901 (2 species, Africa; Cameroon)
Pseudothiscia Schmidt, 1912 (1 species, South Africa; http://biodiversitylibrary.org/page/8824628)

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