Monalonion, Herrich-Schaeffer, 1850

Namyatova, Anna A. & Cassis, Gerasimos, 2016, Systematic revision and phylogeny of the plant bug tribe Monaloniini (Insecta: Heteroptera: Miridae: Bryocorinae) of the world, Zoological Journal of the Linnean Society 176 (1), pp. 36-136 : 103-104

MONALONION HERRICH- SCHAEFFER

Figures 9 , 16I–L , 19N, O , 24

Monalonion Herrich-Schaeffer, 1850: 168 (gen. nov.; type species Monalonion parviventre Herrich-Schaeffer, 1859 by monotypy); Fieber, 1858: 300 (key to gen.); Signoret, 1858: 500 (descr.); Walker, 1873: 161 (cat.); Distant, 1883: 246 (descr.); Atkinson, 1890a: 49 (cat.); Kirkaldy, 1906: 134 (list); Kuhlgatz, 1906: 29 (key to gen.); Reuter, 1908: 150 (descr.); Reuter, 1910: 153 (cat.); Bondar, 1939: 1 (note); Knight, 1939: 226 (note); Carvalho, 1952: 59 (cat.): Carvalho, 1955: 39 (key to gen.); Carvalho, 1957: 138 (cat.); Carvalho, 1972: (diag., key to spp.); Lavabre, 1977a: 65 (diag.); de Abreu, 1977: 85 (desc., distrib., ecol.); Schuh, 1995: 517 (cat.); Schuh, 2002 –2013 (cat.); Namyatova et al., in press (phylogeny).

Diagnosis: Among genera of the Monalonion -group, Monalonion can be separated by: ASI very short, c. 1.5– 3× as long as wide; elongate body ( Fig. 9 ), ASII–IV filiform; collar distinctly swollen (as in fig. 7D in Namyatova & Cassis, 2013b); scutellum without spine; forecoxae contiguous (as in fig. 17A in Namyatova et al., in press); foretibia almost straight; secondary gonopore surrounded with small sclerite ( Fig. 16I ); ductus seminis slightly sclerotized basally; and DLP with sclerotized circle ( Fig. 19N ).

Redescription: Male: Body length 7–12 mm. COLORA- TION ( Fig. 9 ). Variable, mostly brown to dark brown or yellow, sometimes with reddish areas; antennae and legs usually dark colored, legs sometimes with yellow or pale brown band on femur or mostly yellow. TEXTURE. Body mostly smooth, without punctures, wrinkles and tubercles; vertex without flattened areas; semicircular depression between scutellum and mesoscutum absent; striations on scutellum laterally absent; only small depression on anterior angle of pronotum present (as in fig. 9H in Namyatova & Cassis, 2013b). VESTITURE. Setae on dorsum and thoracic pleura absent; only short simple suberect setae on ASII–IV, legs and abdomen present; setae on ASII sometimes slightly longer than width of hind tibia; setae on tibia and hind femur sometimes very dense, shorter or longer than width of hind tibia; setae on abdomen pale, short and adpressed and very rare; black spinules on femora and tibiae absent. STRUCTURE. Head. Distance between eye and pronotum subequal to half of eye diameter (as in Fig. 10J ), sometimes subequal to eye diameter; occipital region delimited with distinct depression; longitudinal depression on vertex as long or slightly longer than eye diameter; eyes not stylate, in line with contour of head, c. 0.2–0.3× as long as head width; distance between antennal fossae oval as long as or slightly longer than antennal fossa diameter; frons straight or slightly convex, without ridges, outgrowths or longitudinal depression; anterior view of head c. 1.4– 1.7× as wide as high; eye c. 1.7–2.8× as long as distance between eye and apex of clypeus; antennal fossa oval c. 0.5–0.7× as long as eye height, not raised (as in fig. 6D–F in Namyatova & Cassis, 2013b), inferior margin placed distinctly above inferior margin of eye; position of base of clypeus varying from distinctly below inferior margin of antennal fossa to near halfway of antennal fossa height, not delimited with depression; head swollen in lateral view; length of gula varies from slightly longer to almost twice as long as buccula. Labium. Length varying from reaching middle of mesosternum to slightly surpassing posterior margin metasternum; c. 2–3× as long as wide; LSIII c. 3–4× as long as wide, varying from slightly shorter to distinctly longer than LSI; LSIII c. 2–5× as long as wide, varying from slightly shorter to distinctly longer than LSII; LSV c. 4–8× as long as wide, from slightly longer almost twice as long as long as LSIII. Antenna. Slightly shorter or longer than body; ASI c. 2–3× as long as wide, c. 0.3–0.7× as long as head width, widened basally; ASII c. 4–6× as long as ASI, c. 1.4–2× as long as head and pronotum combined; ASIII c. 0.7–0.9× as long as ASII; ASIV c. 0.2– 0.3× as long as ASIII; ASII–IV filiform. Thorax. Collar distinctly delimited laterally and posteriorly, swollen; calli separated, flat almost indistinct (as in Fig. 10J ); depression delimiting calli posteriorly absent (as in Fig. 10J ); humeral angles of pronotum rounded, not dilated (as in Fig. 10J ); posterior margin of pronotum slightly sinuate (as in Fig. 10J ); scutellum flat, rounded apically; without outgrowth or medial depression; metepimeron c. 3–4× as high as long, rounded (as in fig. 9C in Namyatova & Cassis, 2013b); metasternum rounded posteriorly, without medial projection on to abdominal segment II (as in fig. 17B in Namyatova et al., in press). Hemelytron. Costal margin concave or almost straight, hemelytron almost not tapering posteriorly; claval commissure c. 2–5× as long as scutellum, concave (as in fig. 11G in Namyatova et al., in press); R + M distinct, reaching posterior margin of corium; medial fracture subparallel to R + M; corium without swelling posteriorly; cuneus c. 3–4× as long as wide, c. 0.7–1.2× as long as pronotum, medial margin distinctly concave (as in fig. 13C in Namyatova et al., in press); membrane cell c. 1.4–2.2× as long as pronotum, rounded or acute apically; auxiliary vein absent; distance between cell and apex of membrane c. 0.2–0.4× as long as cell. Legs. Forecoxae contiguous (as in fig. 17A in Namyatova et al., in press); femora more or less swollen apically (as in fig. 18A in Namyatova et al., in press); forefemur almost straight, middle and hind femora distinctly curved, hind tibia sometimes with medial part widened; foretibia slightly shorter, as long as or slightly longer than head and pronotum combined; tibiae without swellings; segment I of hind tarsus distinctly longer than segment II and subequal to or slightly longer than segment III (as in fig. 19A in Namyatova et al., in press); claw broadly rounded (fig. 10F in Namyatova & Cassis, 2013b); basal tooth on claw triangular, short (as in fig. 10B in Namyatova & Cassis, 2013b). Genitalia ( Fig. 16I–L ). Genital capsule slightly longer than wide, without outgrowth(s); ventral wall shortened anteriorly; left paramere r-shaped, twice as long as right paramere; phallobase sclerite of primary gonopore suboval, without outgrowths supporting ductus seminis; ductus seminis longer than phallotheca, with coils forming wide tube, sclerotized basally and with narrow sclerite around secondary gonopore, attached to phallobase medially; sclerotized part of phallotheca occupying entire dorsal side, rounded apically, without ridge or outgrowth(s); endosoma with small sclerites or sclerotized areas.

Female: Body length 8–12.5 mm. Coloration, surface, vestiture and structure as in male, generally larger than males and eye usually smaller than in male ( Fig. 9 ). Genitalia ( Fig. 19N, O ). DLP with sclerotized ring, sometimes indistinct, mostly without striations or with some medially, especially at base of lateral oviducts; lateral oviducts placed at halfway of or in posterior half of DLP, removed from each other and from lateral margins of DLP; spermathecal gland placed in posterior half, medially; posterior wall of bursa copulatrix with tubercles, without outgrowths or sclerotizations; base of second valvula with bifurcate outgrowth; ventral wall of bursa copulatrix membranous.

Distribution: Widely distributed in Latin America, including Mexico ( Fig. 24 ).

Host plants: Many species of Monalonion are known to damage cocoa pods ( Distant, 1917; Costa Lima, 1938; Bondar, 1939; Carvalho, 1972; de Abreu, 1977; Entwistle, 1977). Monalonion schaefferi was also recorded from cashew ( Piart, 1977), and M. velenzagi is known from avocado ( Carvalho & Costa, 1988). Species are also known from Ficus sp. , Cercopia adenopsis ( Moraceae ), Hamelia patens (Rubiaceae) and Begonia spp. (Begoniaceae) ( Costa Lima, 1938; Bondar, 1939; Carvalho, 1972; Piart, 1977).

Figures

Figure 9. Habitus photographs. Monalonion dissimulatum ♂ AMNH_PBI 19564, ♀ AMNH_PBI 5258; Physophoropterella bondroiti ♂ AMNH_PBI 400348, ♀ AMNH_PBI 5037; Physophoroptera mirabilis ♂ AMNH_PBI 5070, ♀ AMNH_PBI 5083; Arthriticus eugeniae ♂ AMNH_BPI 19573, ♀ AMNH_PBI 19575; Ragwelellus suspectus ♂ AMNH_PBI 34153, ♀ AMNH_PBI 34154; Ragwelellus vittatus ♂ AMNH_PBI 34152, ♀ AMNH_PBI 45821; Schuhirandella fulva ♂ PT AMNH_PBI 400345, ♀ PT AMNH_PBI 19576; Helopeltis clavifer ♂ AMNH_PBI 40416, ♀ AMNH_PBI 202016; Afropeltis corbisieri ♂ AMNH_PBI 5133, ♀ AMNH_PBI 5216.

Figure 10. Scanning electron micrographs. Head and pronotum, dorsal view. A, Mansoniella nitida ♀ AMNH_PBI 46067; B, Poppiusia leroyi ♀ AMNH_PBI 5838; C, Chamus bellus ♀ AMNH_PBI 5223; D, Lycidocoris mimeticus ♀ AMNH_PBI 5043; E, Physophoropterella bondroiti ♂ AMNH_PBI 19110; G, Odoniellia reuteri ♂ AMNH_PBI 19090; H, Volkelius carvalhoi ♀ PT AMNH_PBI 19630; I, Yangambia vesiculata ♀ AMNH_PBI 19084; J, Ragwelellus suspectus ♂ AMNH_PBI 19629. Head, dorsal view. F, Sahlbergella tai ♀ AMNH_PBI 5106. Head, anterior view. K, Parachamus bellus ♀ AMNH_ PBI 5837. Head, lateral view. L, Poppiusia leroyi ♀ AMNH_PBI 5838. Frons, dorsal view. M, Poppiusia leroyi ♀ AMNH_PBI 5838.

Figure 16. Male genitalia, dorsal view. Mansoniella flava AMNH_PBI 19270. A, aedeagus; B, right paramere; C, left paramere; D, genital capsule. Mansoniella sassafti AMNH_PBI 19276. E, aedeagus; F, right paramere; G, left paramere; H, genital capsule. Monalonion dissimulatum no USI. I, aedeagus; J, right paramere; K, left paramere; L, genital capsule. Odoniellia rubra AMNH_PBI 18951. M, aedeagus. AMNH_PBI 18950. N, right paramere; O, left paramere; P, genital capsule. Odoniellia similis AMNH_PBI 5036. Q, aedeagus; R, left paramere; S, genital capsule. Pachypeltis chinensis no USI. T, aedeagus; U, right paramere; V, left paramere; W, genital capsule. Pachypeltis sp. (Bouganville) AMNH_PBI 46046. P, aedeagus; Q, right paramere; R, left paramere; S, genital capsule. Pararculanus madagascariensis no USI. X, aedeagus; Y, right paramere; Z, left paramere; AA, genital capsule. Pararculanus piperis AMNH_PBI 18979. AB, aedeagus; AC, right paramere; AD, left paramere; AMNH_PBI 271336. AE, genital capsule. The smaller scale is for genital capsule, the larger scale is for aedeagi and parameres.

Figure 19. Female genitalia, bursa copulatrix. Eupachypletis pilosus AMNH_PBI 46068. A, dorsal labiate plate; B, posterior wall. Helopletis clavifer AMNH_PBI 74192. C, dorsal labiate; D, posterior wall. Lycidocoris modestus AMNH_PBI 5099. E, dorsal labiate plate. Lycidocoris mimeticus AMNH_PBI 5086. F, dorsal labiate plate; G, posterior wall. Mansoniella nitida AMNH_PBI 45970. H, dorsal labiate plate; I, posterior wall. Lycidocoris thoracicus AMNH_PBI 5096. G, dorsal labiate plate; K, posterior wall. Mansoniella sassafri AMNH_PBI 19067. L, dorsal labiate plate; M, posterior wall. Monalonion sp. AMNH_PBI 19632. N, dorsal labiate plate. Monaloion dissimulatum no USI. O, posterior wall. Odoniella rubra AMNH_PBI 18952. P, dorsal labiate plate; Q, posterior wall. Rhopaliceschatus quadrimaculatus AMNH_PBI 20523. R, dorsal labiate plate.

Figure 24. Distribution maps of Monalonion, Odoniella, Pararculanus, Physophoroptera, Poppiusia, Ragwelellus, Rhopaliceschatus, Volkelius and Yangambia.