Hecalini, Distant, 1908

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: 91-92

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Hecalini   Distant, 1908

Figs 27 View Fig , 28 View Fig

Type genus: Hecalus Stål, 1864   .

= Reuteriellini Evans, 1947. 


Hecalini   are medium sized to large, somewhat to strongly dorsoventrally flattened, stramineous, yellow, green, or brown leafhoppers, sometimes with bright orange or reddish markings. They can be identified by the produced and parabolically shaped head, dorsoventrally flattened body, lateral margin of pronotum as long as or longer than the basal width of the eye, ocelli closer to they eyes than the laterofrontal sutures, apodemes of male sternite I long and relatively narrow, apodemes of male sternite II broad and well-developed, male pygofer often produced or pointed posterodorsally, segment X withdrawn into the pygofer, ventral margins of the male pygofer often lobate, aedeagus often with one or two pairs of apical processes, first valvula dorsal sculpturing granulose to maculose and submarginal, first valvula often with distinctly delimited ventroapical sculpturing, second valvula usually without teeth, humpbacked dorsally, and concave ventrally.


HEAD. Head subequal to or wider than pronotum; often produced anteriorly and/or parabolically shaped. Discal portion of crown glabrous with radial or longitudinal striae, shagreen or granulose. Anterior margin of head shagreen, glabrous, irregularly textured, foliaceous, with a single transverse carina, or with 2 or 3 parallel carinae. Frontoclypeus tumid or flat; texture shagreen. Clypellus parallel-sided, tapering apically, or rarely widening apically; apex following or slightly surpassing normal curve of gena. Lorum narrower than, subequal to, wider than clypellus near base. Antennal bases near middle or posteroventral (lower) corners of eyes. Antennae short, less than 1.5 x width of head. Gena obtusely incised laterally or strongly incised laterally (nearly forming right angle); with or without ( Attenuipyga   , some Glossocratus   ) fine erect seta beside laterofrontal suture. Antennal ledge absent. Ocelli present; close to eyes, closer to eyes than laterofrontal sutures; on anterior margin of head.

THORAX. Pronotum lateral margin carinate; lateral margin as long as or longer than basal width of eye.

WINGS. Forewing brachypterous to macropterous; if macropterous, appendix restricted to anal margin or rarely large, extending around wing apex; with 3 anteapical cells; veins not elevated or rarely elevated ( Attenuipyga   ); without reflexed costal veins; A1-A2 crossvein absent or present; A1 and A2 sometimes confluent ( Attenuipyga   [ Dorycara   ]).

LEGS. Profemur with AM 1 seta only or rarely with AM 1 and with one or more additional proximal setae (some Glossocratus   ) or AM 1 absent ( Attenuipyga   ); intercalary row with one row of five or more fine setae or setae scattered, not in one row, or setae absent or reduced; row AV with short, stout setae or with relatively long macrosetae. Protibia dorsal surface rounded, convex. Metafemur apex macrosetae 2+2+1, 2+2+1 and with additional proximal macrosetae (some Glossocratus   ), or 2+0 ( Attenuipyga   ). Metatarsomere I not expanded apically or expanded apically; plantar setae simple, tapered.

MALE GENITALIA. Apodemes of sternite I usually long, usually as long as those of sternite II, relatively narrow. Apodemes of sternite II large, broad (exception: Glossocratus   ). Valve articulated with pygofer; lateral margin short, articulating with pygofer at a point. Pygofer basolateral membranous cleft present; macrosetae well differentiated into several rows; often produced or pointed posterodorsally, with ‘anal tube’ withdrawn into pygofer; ventral margins often lobate. Subgenital plates free from each other; articulated with valve; macrosetae scattered, irregularly arranged or uniseriate laterally. Style broadly bilobed basally, median anterior lobe pronounced. Basal processes of the aedeagus/connective absent or reduced. Aedeagus with single shaft and gonopore; often with one or two pairs of apical processes, or rarely with preapical processes. Connective anterior arms somewhat divergent, Y - or U -shaped; articulated with aedeagus.

FEMALE GENITALIA. Pygofer with macrosetae absent, reduced, or with numerous macrosetae. Ovipositor usually protruding or sometimes not protruding far beyond pygofer apex. First valvula not strongly convex; dorsal sculpturing pattern granulose or maculose; sculpturing submarginal or rarely reaching dorsal margin; usually with ventroapical sculpturing distinctly delimited. Second valvula gradually broadened medially or subapically; humpbacked dorsally; concave ventrally; usually without dorsal median tooth (except Glossocratus   ); dorsal teeth usually absent but if present (some Glossocratus   ), teeth small, regularly or irregularly shaped, on apical 1/3 or more.

Geography and ecology

Distribution: cosmopolitan. Hecalini   are grass feeders and are sometimes very brightly colored with yellow, green (in live specimens), or orange. Some are extremely elongate and dorsoventrally flattened, apparently an adaptation enhancing crypsis on their grass hosts.


Hecalini   contains 23 genera and 180 species. The three included exemplars ( Hecalus   , Attenuipyga   and Glossocratus   ) were resolved as a monophyletic group here in MP analyses, but not in ML or Bayesian analyses. In the MP analysis, the partitioned Bremer support from the Histone H3 data was -14.4 which apparently strongly conflicted with the signal from the 28S and morphology data sets (DI= 7.7 and 13.2, respectively). This conflict in signal likely influenced the results of the ML and Bayesian analyses which resolved different topologies for the hecalines included here. MP analyses of the combined data suggest that Hecalini   is closely related to Arrugadini, but more data are needed to confirm this.

Hamilton (2000) revised the concept of Hecalini   and excluded several genera that have only a superficial similarity to the tribe. He also pointed out several characters that are shared between Hecalini   and “Dorycephalini”, although of the latter he was apparently referring only to the New World Attenuipyga   and Neoslossonia   , and not to Dorycephalus   which does not share several of the characters he mentioned. The characters listed by Hamilton (2000) and others noted here that are shared between Attenuipyga   , Neoslossonia   , and other Hecalini   and not with Dorycephalus   are listed in the Diagnosis. Based on their shared morphological characters and support from the molecular data, Attenuipyga   and Neoslossonia   are transferred to Hecalini   here. Dorycephalini is maintained as a separate monotypic tribe.

Morrison (1973) revised the Oriental fauna and suggested that Clavena   may belong to Ledrini based on the position of the ocelli stated in the original description of C. sulcata Melichar. He   did not examine the type, so its inclusion in Hecalini   has not been confirmed. Sectoculus   is removed from Hecalini   and placed in Eupelicini: Paradorydiina (see discussion under Paradorydiina).

Selected references

Linnavuori (1957, 1959, 1975), Morrison (1973), Linnavuori & DeLong (1978c), Hamilton (2000), Dmitriev (2002).

Included subtribes:


Australian Museum













Zahniser, James N. & Dietrich, Chris H. 2013


Reuteriellini Evans, 1947.