Neella Reuter, 1908

Mingheti, Eugenia, Montemayor, Sara Itzel & Dellapé, Pablo Matías, 2025, Phylogenetic revision of the Neella-Neoneella complex (Hemiptera: Heteroptera: Miridae: Bryocorinae), with description of five new genera, Zoological Journal of the Linnean Society 204 (2) : -

Neella Reuter, 1908

( Figs 1C, 2F, 3D, E, K, 5C, K, O, 8)

Tope species: Eccritotarsus eucosmus Stål, 1862 . By original designation.

Neella Reuter, 1908: 152–154 (original description); Reuter, 1910: 153 (catalogue); Bergroth, 1922: 17 (species transfer); Costa Lima, 1942: 103–104 (citation); Carvalho, 1945: 161– 172 (description of new species; redescription of known species; key); Hsiao, 1946: 385–387 (description of a new species; species transfer; key); Carvalho, 1948: 100, 103–104 (description of a new species); Carvalho, 1952: 56; Carvalho, 1954a: 1–19 (description of new species; redescription of known species; key); Carvalho, 1955: 32 (key); Carvalho, 1960: 47, 49 (key); Carvalho and Gomes, 1971: 466–468 (description of new species); Carvalho, 1974b: 324–325 (description of a new species); Carvalho and Schaffner, 1985: 7–14 (description of new species); Carvalho and Ferreira, 1995: 477 (key); Carpintero, 1998: 145 (citation); Kerzhner and Konstantinov, 1999: 124; Ferreira et al., 2001: 162 (distribution); Schuh, 2002–13 (online catalogue); Ferreira et al., 2006: 4, 10 (distribution); Coelho, 2008: 18, 25,104, 116, 149 (key; distribution); Ferreira and Henry, 2011: 2, 13, 19–20, 34 (key; distribution); Namyatova et al., 2016: 14 (citation); Etl et al., 2016: 300, 302–303 (host plants); Barbosa et al., 2019: 158 [ Neela misspelling] (host plant); Álvarez-Zapata et al., 2022: 101 , 105, 107, 109–110, 114, 116, 129–130, 136, 144 (generic and species key; figs; distribution); Ferreira et al., 2024: 207, 214, 215, 222, 242 (diagnosis; generic key; figs).

Included species: N. bicolor Hsiao, 1946 N. carvalhoi Hsiao, 1946

N. eucosma ( Stål, 1862)

N. fasciata Hsiao, 1946

N. floridula (Distant, 1883) N. mantiqueirae Carvalho, 1954a N. peruana Carvalho, 1974b N. similaris Carvalho and Gomes, 1971

Diagnosis: Orange to reddish, with brown to black areas in pronotum and hemelytra; eyes large, more than half of head height, posterior margin concave; vertex as maximum 2.5× as wide as one eye; posterior margin of head with a group of recumbent setae behind eyes; labrum flat in lateral view; labium at least extending to metacoxae; antennal segment II swollen subapically; anterior margin of collar concave; posterior pronotal margin straight, pronotal width across calli half as wide as posterior margin width; inner margin of cuneus concave; cell angled before apex of cuneus; tibiae with only one type of setae; lef paramere with two or three curvatures, tapering towards apex; right paramere different from lef, C-shaped, tapering towards apex.

Redescription: Male. TBL 4.00–5.89; PBL 3.38–5.32; BW 1.51–2.59. Female. TBL 3.47–5.70; PBL 3.19–5.22; BW 1.44–2.66. Coloration: Orange to reddish, with brown to black areas in pronotum and hemelytra. Head: pale yellow to reddish, sometimes with brown to black areas; eyes black, golden or silver, or black with golden or silver areas; antennal segments I–II yellow, orange, reddish or brown reddish; III, pale yellow to reddish or brown; IV, pale yellow to reddish, sometimes distal half brown. Torax: collar pale yellow to dark red or brown; calli pale yellow to reddish; posterior pronotal lobe pale yellow to dark red or brown, in some species with dark transverse fascia or semicircular area; mesoscutum and scutellum yellow to black; pleura pale yellow to reddish, in some species propleura bicoloured. Hemelytron: pale yellow to reddish, brown or black, unicolorous, or with longitudinal brown fascia extending to membrane; membrane translucent to black; veins yellow to black. Legs: pale yellow to dark brown; usually femora and tibiae bicoloured; tarsi and claws pale yellow to black. Abdomen: pale yellow to dark red. Surface: Shiny; posterior pronotal lobe evenly punctate; head, collar, pronotum, and hemelytra with short, recumbent or semierect setae; labium with semierect setae; antennal segments I–II with short and recumbent or semierect setae, sometimes with some erect setae; III–IV, with abundant recumbent to semierect setae; veins with very short and recumbent setae; pleura and abdomen with semierect setae, longer than dorsum; coxae, trochanters, femora, and tarsi with short and semierect setae; femora with dorsal setae as long as ventral setae; tibiae with thicker setae. Structure: Head: at least 2× as wide as long; clypeus not visible from above, flat or rounded in lateral view; frons rounded to globose in lateral view; vertex flat, as maximum 2.5× as wide as one eye; eyes large, stylate, more than half of head height, upper margin of eyes overcoming dorsal margin of head; posterior margin concave; gula not visible to short; labrum flat in lateral view; labium at least extending to metacoxae; antennal segment I straight; II more than 2× as long as I, swollen subapically; III, tapering towards apex; IV, straight and thinner than III; posterior margin of head with a group of recumbent setae behind eyes. Torax: anterior margin of collar concave and posterior margin straight or convex, thinner than antennal segment I; calli separated in central area, ataining or not lateral margins of pronotum; pronotal width across calli half as wide as posterior margin width, posterior margin straight; mesoscutum usually exposed; scutellum with basal depression. Hemelytron: lateral margins slightly convex to convex; embolium thin, bent upwards; claval commissure longer than scutellum and half of pronotum length combined; anterior half of medial fracture adjacent to R + M vein, posterior half separated from R + M vein; cuneus longer than wide, inner margin concave; cell angled before or in middle of membrane, before apex of cuneus, inner margin straight or convex and posterior margin straight or concave; veins angled before apex of cuneus. Legs: profemora widened basally; meso- and metafemora straight; metafemora slightly curved; protibia slightly flatened distally. Male genitalia: Genital capsule wider than long; genital opening broad, facing dorsally; right wall equal to or shorter than lef; dorsal wall well developed or reduced, in some species with projections, posterior margin concave; ventral wall more developed than dorsal, posterior margin convex, with one or two projections; subgenital plate with or without projections, ventral to parameres insertion. Lef paramere smaller than right, curved to sinuate; body with two or three curvatures, tapering towards apex; apical process rounded or pointed, in some species bifurcate. Right paramere different from lef, C-shaped; basal process narrow; apical process simple or bifurcate. Aedeagus small and simple; phallotheca membranous with some sclerotized areas; seminal duct folded initially, membranous, sclerotized posterior to its distal curvature or completely sclerotized, at least extending to half of phallotheca; endosoma membranous, in some species apex sclerotized.

Distribution: Argentina, Brazil, Colombia, Costa Rica, Ecuador, El Salvador, French Guiana, Guatemala, Honduras, Mexico, Panama, Paraguay, and Peru ( Fig. 8).

Discussion: In our analysis Neella is highly polyphyletic, indicating several species require transfer to other known or to new genera. Te phylogeny recovers seven species in the Neella clade (plus N. similaris ) and as the sister-group of the clade Egerocoris , Laterocavocoris gen. nov., Lelena gen. nov., Neoneella , and Puncticollus gen. nov. Te genus is supported by five synapomorphies, one continuous character: relation of posterior pronotal margin width/anterior pronotal margin width (11: 0.193 –0.198), three discrete contradictory synapomorphic characters: the cylindrical antennal segment II (38:0), tibiae with one type of setae (89:0) and the right paramere C-shaped in dorsal view (122:1), and a discrete synapomorphic character: lef paramere with a basal and distal curvatures in straight angle (137:1). Te GC frequencies indicate a low support due to high contradictory information ( Fig. 7).

Troughout the years different authors have related Neella with other genera of Eccritotarsini. Reuter (1908) differentiated it from Sosinas Distant by the vertical position of the head, the flat vertex, the flatened clypeus in lateral view, the more rounded inner margin of the eyes, the long labium, the large and transverse calli, and the scutellum with a triangular depression at the anterior margin. Also, when Costa Lima (1942) described Neoneella he recognized its affinity with Neella , but distinguished them by the absence of sexual dimorphism in the development of the cuneus in Neella . Hsiao (1946) considered Neella closely related to Tenthecoris Scot , but pointed out that in Neella the body is more elongated, the lateral margins of the hemelytra are less convex, the vertex is flat, and the labium long, with segments III–IV short and thin. Finally, Carvalho (1960) highlighted that Neella is related to Adneella , Neoneella , Paraneella , and Proneella , transferring one species from Neella to Adneella and suggesting that four other species should also be transferred. In our hypothesis, Sosinas and Tenthecoris are located basally together with Paraneella , Adneella, Pocnoderes , Parafurius , and Globicephalomiris gen. nov., whereas Neella is related to Proneella and other genera (see ‘Phylogenetic results’ for a more detailed analysis of characters).