Scaphoideini Oman , 1943

Zahniser, James N. & Dietrich, Chris H., 2013, A review of the tribes of Deltocephalinae (Hemiptera: Auchenorrhyncha: Cicadellidae), European Journal of Taxonomy 45, pp. 1-211 : 148-152

Scaphoideini Oman, 1943

Fig. 47

Type genus: Scaphoideus Uhler, 1889 .

Diagnosis

Scaphoideini are medium sized (usually 4-7 mm), often slender, ivory, yellowish, ochraceous, or brown leafhoppers, often marked with orange, brown, or black on head, pronotum, or forewings. None of the following diagnostic characters are present in all taxa, but some combination of these characters is present in all and a few (*) appear to be unique to this tribe: head narrower than pronotum, produced; frontoclypeus long, narrow; antennae long; body slender; head and wings often with brown, orange, ochraceous, or ivory markings; forewing with one or more darkly pigmented reflexed veins in vicinity of outer anteapical cell; profemur row AV setae absent or reduced (without stout setae); metatibia macrosetae in row PD long, as long as or longer than 0.5 x length of protibia*; male or female pygofer with dense tufts of long fine or regular setae*; subgenital plate apex membranous or long, digitate, and somewhat membranous or weakly sclerotized; subgenital plates with long fine setae laterally and/or dorsally (also occurs in many other deltocephaline tribes); basal processes of the aedeagus or connective sometimes present, connected or articulated to base of aedeagus or apex of connective stem.

Description

HEAD. Head often narrower than pronotum, sometimes subequal to or wider than pronotum; anterior margin shagreen; crown rounded to face or angulate. Discal portion of crown glabrous with radial or longitudinal striae. Frontoclypeus not tumid; long, narrow; shagreen. Clypellus widening apically. Lorum subequal to or wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes or near anterodorsal (upper) corners of eyes. Antennae long (longer than width of head). Gena obtusely incised or not incised laterally; with fine erect seta beside laterofrontal suture. Antennal ledge absent. Ocelli close to eyes.

THORAX. Pronotum lateral margin carinate; lateral margin shorter or longer than half basal width of eye.

WINGS. Forewing macropterous; appendix restricted to anal margin; veins sometimes bordered or highlighted at apices with brown coloration; with 3 anteapical cells; often with pigmented reflexed costal veins; usually without A1-A2 crossvein; apical venation not highly reticulate.

LEGS. Profemur row AM with AM1 seta only; intercalary row with one row of five or more fine setae; row AV setae usually reduced or absent, or sometimes short, stout setae. Protibia dorsal surface rounded, convex. Metatibia macrosetae in rows AD and PD often very long, row PD setae often equal to or longer than 0.5 x length of protibia. Metatarsomere I long. Metafemur apical setae 2+2+1.

MALE GENITALIA. Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point. Pygofer with basolateral membranous cleft; often with many long thick or fine setae sometimes arranged in dense tufts or groups. Subgenital plate with macrosetae absent, uniseriate laterally, or uniseriate, distant from lateral margin; often with long fine hairs laterally and/or dorsally; plate apex often long, digitate, and/or membranous or filamentous. Style broadly bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or present, connected to or articulated with connective or near base of aedeagus. Aedeagus with single shaft and gonopore. Connective usually Y -shaped, sometimes linear or with anterior arms closely appressed; articulated or rarely fused ( Sikhamani , Thryaksha , and Univagris ) with aedeagus, or aedeagus dissociated from connective ( Scaphoideus ).

FEMALE GENITALIA. Pygofer with numerous macrosetae, sometimes with dense tufts or groups of fine or thick setae. Ovipositor not protruding far beyond pygofer apex. First valvula convex; dorsal sculpturing pattern often concatenate, sometimes strigate or reticulate; reaching dorsal margin. First valvula ventroapical sculpturing indistinctly delimited or absent. Second valvula abruptly broadened medially or subapically or broad, gradually tapered; with or without dorsal median tooth; dorsoapical teeth large or small, regularly shaped; teeth on apical 1/3 or more.

Geography and ecology

Distribution: cosmopolitan. Scaphoideini are among the most commonly encountered forest-dwelling deltocephalines throughout the tropical and warm-temperate regions of the world. Nymphs and adults are often collected on understory grasses or herbaceous vegetation but most species appear to require woody hosts for oviposition. Scaphoideus titanus Ball, 1932 is a serious pest and vector of grape flavescence dorée in Europe and of chrysanthemum yellows. S. luteolus Van Duzee, 1894 is a vector of elm phloem necrosis in the United States. Osbornellus borealis DeLong & Mohr, 1936 is a vector of western X disease.

Remarks

Scaphoideini contains 61 genera and 631 species. Oman et al. (1990) included 12 genera, significantly fewer than the circumscription here. Several genera included previously ( Acunasus DeLong, 1945 , Danbara Oman, 1949 , Sobara Oman, 1949 and Twiningia Ball, 1931 ) are not included here. Twiningia was included in the phylogenetic analyses here and was found to be closely related to Platymetopius Burmeister, 1838 (Athysanini) , and the others are excluded because they do not possess characters considered here to be diagnostic for the tribe.

The phylogenetic analyses presented here consistently resolved a clade including Scaphoideini sensu novo plus Drabescini. This clade was in turn resolved as sister to Phlepsiini. Scaphoideini was resolved as paraphyletic with respect to Drabescini and three undescribed genera, but with little or no branch support for the basal branches of the group. Genera of Scaphoideini included in phylogenetic analyses here that were not included previously are Loipothea Linnavuori, 1969 , Omanana DeLong, 1942 , Scaphoidophyes Kirkaldy, 1906 , Anoplotettix Ribaut, 1942 and Mimotettix Matsumura, 1914 . Further taxon sampling and molecular data may help to resolve the relationships within this large tribe and its relationship to the other related tribes, but the available evidence suggests that this is a lineage distinct from other Athysanini , the tribe in which many of the genera were previously placed. The circumscription of the tribe here is based on the phylogenetic analyses and the morphological characters identified here as diagnostic for most members of the group. Representatives of most genera of Athysanini were examined or were reviewed from the literature for their potential placement in Scaphoideini. Further adjustments to the circumscription may be needed after more detailed study of the group.

The Perugrampta group (including Perugramptella , Penthigrampta , and Stenogrampta ) is placed here pending further study. These genera possess the following characters that are diagnostic for Scaphoideini: profemur row AV setae absent, darkly pigmented reflexed branches of R vein, male or female pygofer or subgenital plates with tufts of fine or thick setae, subgenital plates with long fine setae laterally, and basal processes of the aedeagus or connective present and connected to base of aedeagus.

Selected references

Barnett (1977, 1979, 1980), Martinson (1977), Knight & Fletcher (2007), Viraktamath & Mohan (1994, 2004), Stiller (2001), Zhang & Dai (2006), Freytag (2008), Dominguez & Godoy (2010), Dai & Dietrich (2011).

Included genera

Acastroma Linnavuori, 1969 placement nov. (transferred from Athysanini )

Afroideus Linnavuori, 1961 placement nov. (previously unplaced in Deltocephalinae )

Alemaia Heller & Linnavuori, 1968 placement nov. (previously unplaced in Deltocephalinae )

Amimenus Ishihara, 1953 placement nov. (previously unplaced in Deltocephalinae )

Anoplotettix Ribaut, 1942 placement nov. (previously placed in Platymetopiina)

Bampurius Dlabola, 1977 placement nov. (transferred from Athysanini )

Banus Distant, 1908 placement nov. (transferred from Athysanini )

Burakia Kocak, 1981 placement nov. (previously placed in Platymetopiina)

Cantura Oman, 1949 placement nov. (previously unplaced in Deltocephalinae )

Caphodus Oman, 1938 placement nov. (previously unplaced in Deltocephalinae )

Capoideus Theron, 1974 placement nov. (previously unplaced in Deltocephalinae )

Coroticus Distant, 1918 placement nov. (transferred from Scaphytopiini )

Dariena Linnavuori & DeLong, 1977 placement nov. (transferred from Athysanini )

Delospilopterus Stiller, 2001 placement nov. (transferred from Athysanini )

Euscelidella Evans, 1954 placement nov. (previously unplaced in Deltocephalinae )

Ferganotettix Dubovsky, 1966 placement nov. (transferred from Athysanini )

Grammacephalus Haupt, 1929 placement nov. (transferred from Scaphytopiini )

Gunghuyana Distant, 1910 placement nov. (previously unplaced in Deltocephalinae )

Loipothea Linnavuori, 1969 placement nov. (previously unplaced in Deltocephalinae )

Longicornis Li & Song, 2008 placement nov. (transferred from Athysanini )

Madagella Evans, 1954 placement nov. (previously placed in Platymetopiina)

Melanetettix Knight & Fletcher, 2007 placement nov. (transferred from Athysanini )

Mesotettix Matsumura, 1914 placement nov. (previously unplaced in Deltocephalinae )

Mimotettix Matsumura, 1914 placement nov. (transferred from Athysanini )

Mirzayansus Dlabola, 1979 placement nov. (transferred from Athysanini )

Monobazus Distant, 1908 placement nov. (transferred from Athysanini )

Moorada Ghauri, 1975 placement nov. (transferred from Athysanini )

Nataretus Theron, 1980 placement nov. (transferred from Athysanini )

Neolimnus Linnavuori, 1953 placement nov. (transferred from Athysanini )

Neophlepsius Linnavuori, 1955 placement nov. (transferred from Athysanini )

Omanana DeLong, 1942 placement nov. (transferred from Athysanini )

Osbornellus Ball, 1932 placement nov. (transferred from Athysanini )

Pachodus Linnavuori, 1961 placement nov. (transferred from Athysanini )

Parallygus Melichar, 1903 placement nov. (transferred from Athysanini )

Paramelia Evans, 1954 placement nov. (previously placed in Platymetopiina)

Penthigrampta Dietrich & Rakitov, 2002 placement nov. (transferred from Penthimiini)

Perugrampta Kramer, 1965 placement nov. (transferred from Penthimiini)

Perugramptella Dietrich & Rakitov, 2002 placement nov. (transferred from Penthimiini)

Phlogotettix Ribaut, 1942 placement nov. (previously unplaced in Deltocephalinae )

Phlogothamnus Ishihara, 1961 placement nov. (previously unplaced in Deltocephalinae )

Platyretus Melichar, 1903 placement nov. (transferred from Athysanini )

Premanus DeLong, 1944 placement nov. (transferred from Athysanini )

Prescottia Ball, 1932 placement nov. (previously unplaced in Deltocephalinae )

Scaphodhara Viraktamath & Mohan, 1994 placement nov. (previously unplaced in Deltocephalinae )

Scaphoidella Vilbaste, 1968 placement nov. (transferred from Athysanini )

Scaphoideus Uhler, 1889 placement nov. (transferred from Athysanini )

Scaphoidophyes Kirkaldy, 1906 placement nov. (previously unplaced in Deltocephalinae )

Scaphoidula Osborn, 1923 placement nov. (previously unplaced in Deltocephalinae )

Scaphomonus Viraktamath, 2009 placement nov. (transferred from Athysanini )

Sikhamani Viraktamath & Webb, 2006 placement nov. (transferred from Scaphytopiini )

Soleatus DeLong, 1971 placement nov. (previously unplaced in Deltocephalinae )

Spathanus DeLong, 1945 placement nov. (transferred from Athysanini )

Stenogrampta Dietrich & Rakitov, 2002 placement nov. (transferred from Penthimiini)

Sudhamruta Viraktamath & Anantha Murthy, 1999 placement nov. (transferred from Scaphytopiini )

Tbilisica Dlabola, 1958 placement nov. (previously placed in Platymetopiina)

Thamnophryne Kirkaldy, 1907 placement nov. (previously unplaced in Deltocephalinae )

Thryaksha Viraktamath & Anantha Murthy, 1999 placement nov. (transferred from Scaphytopiini )

Tongdotettix Kwon, 1980 placement nov. (previously placed in Platymetopiina)

Tuakamara Webb, 1980 placement nov. (previously placed in Platymetopiina)

Univagris Viraktamath & Anantha Murthy, 1999 placement nov. (transferred from Scaphytopiini )

Wanritettix Vilbaste, 1969 placement nov. (transferred from Athysanini )

Figures

Fig. 47. Scaphoideini. Scaphoideus omani sp. nov. A–F, H–I. Standard views (see Material & Methods). J. Lateral view of connective and connective processes. K. ♀ sternite VII. L. Style, ventrally. M. Ventral view of connective and connective processes.