Makilingia Baker, 1904

Dietrich, Christopher H. & Zahniser, James N., 2019, Review of the leafhopper genus Makilingia Baker (Hemiptera: Cicadellidae: Mileewinae), Zootaxa 4559 (3), pp. 473-500 : 474-478

Makilingia Baker

Type species: Makilingia nigra Baker 1914

Diagnosis. Head with crown depressed, anterior margin sharply carinate and encroaching onto eye laterally; lateral frontal sutures not extended onto crown; gena broad, without distinct emargination below eye; pronotum and base of forewing distinctly punctate; front femur with seta AM1 small and inconspicuous.

Description. Small to medium-sized, cylindrical leafhoppers (5–10 mm). Coloration usually mostly dark brown or black with legs yellow, usually with symmetrical yellow or orange markings dorsally on head and forewing, rarely mostly pale yellow ( Figs. 1 , 2 ). Dorsum of head, pronotum and mesonotum, gena, lorum and base of forewing distinctly punctate. Head produced, ocelli on crown well separated from anterior margin and eyes, crown with anterior margin parabolic, sharply carinate with carina elevated and encroaching onto eyes laterally; posterior margin of eye not extended to posterior margin of crown medially; face convex; frontoclypeus shagreen with muscle scars visible, with or without punctations medially; lateral frontal sutures weakly divergent, extended to dorsal margin of face; clypeal suture poorly delimited or absent; anteclypeus nearly as long as frontoclypeus, weakly to strongly convex, tapered distally, apical margin extended beyond lower margin of gena; lorum small, well separated from genal margin, not extended dorsad of anteclypeus; gena broad, lateral margin convex, not emarginate below eye, completely concealing proepisternum; antennal ledge absent; antenna shorter than width of head; rostrum extended to middle coxae.

Pronotum wider than head, convex, lateral margins as long as or longer than eyes, divergent posteriorly in dorsal view, carinate, carina distinctly below posterior corner of eye. Exposed part of mesonotum and scutellum variable in size interspecifically, scutellar suture deeply impressed. Forewing with varying amounts of opaque sclerotization, venation somewhat poorly delimited, especially near base; R three branched, s crossvein absent (present in M. pruinosa ); r-m1 present or absent; m-cu2 present; supernumerary r-m and m-cu crossveins sometimes present; CuA reaching wing margin at claval apex; inner apical cell nearly parallel-sided. Hind wing with submarginal vein usually complete, R and M separate distally. Prothoracic femur slender, AM1 small, inconspicuous; intercalary row extended from basal third to apex, with numerous setae becoming shorter and finer distally; rows AV and PV absent; tibia cylindrical or very weakly bicarinate dorsally, dorsal rows with fine setae, row AV with several macrosetae becoming longer distally. Mesothoracic legs approximately same size as prothoracic legs, without conspicuous macrosetae. Hind femur with macrosetal formula 2+1+1, antepenultimate seta often much smaller than others; tibia slender, PD with more macrosetae than AD, intercalary setae absent; tarsomere I with pair of dorsoapical macrosetae, pecten usually with 2 platellae.

Male abdomen with 2S apodemes weakly developed. Pygofer ( Fig. 3 ) with base usually broadly sclerotized dorsally, lobe with longitudinal row of enlarged setae near base, ventral appendage present or absent. Segment X well sclerotized dorsoapically and laterally with distinct articulation to dorsal margin of pygofer adjacent to dorsal connective. Valve large, convex, posterior margin rounded, joined basally to pygofer by narrow sclerotized ridge but with extensive membranous cleft more distad. Subgenital plates movably articulated to valve, fused to each other near base but separated and divergent for most of length, each plate broad and somewhat depressed basally, narrowing distally, with darkly pigmented dorsal denticulate callosity opposite style apex, distal lobe weakly sclerotized compressed, macrosetae absent but scattered short, stout setae present. Style apodeme elongate, straight; apophysis short with large, rounded basal lobe ventrally, apex falcate, with few widely spaced ventral preapical setae. Connective elongate, anterior arms very short and bowed laterad, strongly compressed median anterior ventral keel usually present, stem arcuate in lateral view, widened posteriorly in ventral view, articulated to aedeagus. Aedeagus short, preatrium developed, shaft weakly sclerotized posteriorly, atrium usually with prominent lateral lobes or flanges. Well-sclerotized V-shaped dorsal connective present and articulated between atrium and basal ventrolateral lobes of segment X; gonopore poorly delimited and situated in large, membranous area on posterior surface of shaft.

Female sternite VII covering base of ovipositor, sternite VIII variably sclerotized and folded dorsad of sternite VII in repose. First valvulae slender, dorsal sculpture concatenate, ventroapical sculpture irregularly granulose. Second valvulae with distal toothed blades occupying approximately two thirds total length, slightly sinuate to straight, dorsal margin arcuate with several large teeth distributed over most of blade and small serrations between teeth. Third valvulae (gonoplacs) with sparse small setae ventrally.

Notes. Makilingia , the only genus included in tribe Makilingiini, has an unusual combination of morphological traits that suggest mixed affinities. The structure of the head, with a moderately to strongly convex face and dorsal ocelli, and ovipositor with the toothed blades of the second valvulae occupying more than half the total length of the valvulae, clearly places it in a lineage with Cicadellinae , Mileewinae (where it is currently placed), and Signoretiinae . The presence of distinct punctations on the dorsum and somewhat enlarged pronotum of Makilingia suggests a relationship to Signoretiinae but, unlike members of that subfamily, the pronotum does not extend all the way to the scutellum and the hind leg chaetotaxy is not reduced. The male genitalia are similar to those of Cicadellinae and Signoretiinae (and different from Mileewini and Tinteromini) in having the subgenital plates triangular. Like other Mileewinae , Makilingia has the submarginal vein of the hind wing very close to the wing margin and this was the main basis for Dietrich's (2011) placement of the group as a tribe of Mileewinae . A recent phylogenomic analysis of Membracoidea ( Dietrich et al., 2017) did not recover Mileewinae sensu Dietrich (2011) as monophyletic but provided support for a relationship between Makilinigiini, Tungurahualini, and Cicadellinae . Additional analyses will be needed to resolve the relationship of Makilingiini to other leafhoppers. Some undescribed leafhopper genera and species from Melanesia (New Guinea, Solomon Islands, New Caledonia) resemble Makilingia to various degrees, including the presence of distinct pits on the head and pronotum, basally fused male subgenital plates lacking macrosetae, and presence of a sclerotized dorsal connective articulated between the aedagal atrium and base of anal tube (Viraktamath & Webb, ms. in prep.).

Comparative study of the male genitalia of available specimens revealed considerable variation in several structures that appears to be diagnostic at the species level. The overall structure of the genital capsule is remarkably uniform but variation occurs in the degree of sclerotization of the pygofer base and the length, vestiture and orientation of the ventral pygofer appendage, which is absent in a few species. Species may be divided into two groups based on the structure of the subgenital plate. One group has the plates normally developed with the distal, unpigmented section extended posterad well beyond the basal darkly sclerotized section. The other group has the distal section strongly reduced, membranous and extended dorsad. The structure of the styles and connective are relatively constant among species with only minor variation in relative proportions and in the shape and vestiture of the style apophysis. The aedeagus is also conservative in structure, the main variation occurring in the relative lengths and orientations of the preatrium and shaft, and the shape of the lateral lobes of the atrium, which bears posteriorly directed processes in a few species.

Variation also occurs in the shape and dentition of the female second valvulae, but not to the extent found in the related tribe Mileewini ( Krishnankutty & Dietrich 2011, Yang et al. 2014).

Figures

FIGURE 1. Makilingia spp., dorsal and lateral habitus: A–B, Makilingia banahaoensis, male; C, M. banahaoensis, female; D– E, M. colorata lectotype male; F–G, M. costalis, holotype male; H–I, M. davaoensis, holotype male; J–K, M. flavifrons, male; L–M, M. frontalis, lectotype male; N–O, M. haightiana lectotype male; P–Q, M. intermedia (lectotype male of M. variabilis); R–S, M. lineata holotype female; T–U, M. lobata, holotype male; V–W, M. maculamima, holotype male; X–Y, M. maculata, lectotype male; Z–AA, M. nigra lectotype male; BB–CC, M. nigramima holotype male; DD–EE, M. pallida, lectotype male; FF, M. pallida (cotype of M. pallida benguetensis Baker), female; GG–HH, M. panayensis, holotype female; II–JJ, M. panayensis, male; Scale bars: 1 mm.

FIGURE 2. Makilingia spp., dorsal and lateral habitus: A–B, M. paranigra, holotype male; C–D, M. pruinosa, male; E–F, M. siamensis, holotype male; G–H, M. sibuyanensis lectotype male. I–J, M. similima, lectotype male; K–L, M. speciosa, paralectotype female; M–N, M. surigaoensis, lectotype male; O–P, M. suturalis, lectotype male; Q–R, M. tenebrifrons, holotype male. S–T, M. tettigonoides, lectotype male. U–V, M. uncinata, holotype male; W–X, M. viraktamathi, holotype male; Y–Z, M. woodworthi, holotype male. AA–BB, M. xanthopicta, holotype male.

FIGURE 3. A, Makilingia xanthopicta n. sp., male genital capsule, lateral view, with parts labeled. B–F, Makilingia suturalis Melichar, holotype (drawings by D. A. Young): B, head and pronotum, dorsal view, left side; C, male genital capsule, lateral view; D, left male subgenital plate, ventral view; F, aedeagus, lateral view; F, style and connective, ventral view.