Guapinannus Wygodzinsky, 1951

Frankenberg, Sarah, Knyshov, Alexander, Hoey-Chamberlain, Rochelle & Weirauch, Christiane, 2021, Taxonomic revision of Guapinannus Wygodzinsky, 1951 (Hemiptera: Schizopteridae), with description of 19 new species, Zootaxa 4958 (1), pp. 261-286 : 263-265

Guapinannus Wygodzinsky, 1951

( Figs 1–9)

Type species: Guapinannus bierigi Wygodzinsky, 1951

Diagnosis: Recognized among Schizopteridae by the relatively small eyes ( Fig. 5C, D), four-segmented labium ( Fig. 1A), flat and wide forewings ( Figs 1–4), unique forewing venation with long fracture in the costal margin, long rc1, rc, and tc cells ( Figs 1B, 2, 6A), C+S, with An1/2 claval process except G. dispar sp. n. ( Figs 2, 6A, D), better developed in males than in females ( Fig 1B), males with wing organ associated with An1 on corium adjacent to the claval process ( Figs 2–4, 6A, D), 2-2-3 tarsal formula in both sexes, male genitalia simple ( Fig. 7) and female with well-developed ovipositor ( Figs 1C, 8).

Re-description: Male: Ovoid (greatest length to width ratio 1.57–1.85) to elongate ovoid (length to width ratio 3.4), total body length 0.66–2.06 mm; length from posterior margin of pronotum to posterior wing margin ( 0.51–1.49 mm). General coloration: uniformly pale or a mix of pale and darker brown or reddish brown. Vestiture: Dorsum with dense setae, setae long or short, setae on posterior margin of forewing long or extremely long. Structure: Head: Eye ranging from small (less than one quarter head height) to relatively large (almost half as high as head) ( Fig. 5C, D); ocellus adjacent to eye ( Fig. 5D), at 11 o’clock position seen in lateral view and about twice the size of one ommatidium; four-segmented labium with segments 1, 2 and 3 of similar length and 4 about 2.5–3 times as long as any other segment ( Fig. 1A); muscle scars present on the frons and vertex present ( Figs 1A, 5C). Thorax: Collar in dorsal view narrow ( Fig. 5C); proepisternal lobe inflated ( Fig. 5D); separation of anterior and posterior lobes (dorsal view) marked by weak depression ( Fig. 5C); anterior pronotal lobe with muscle scars, margin concave or straight; posterior margin slightly concave to almost straight; pits on pronotum ranging from small to large, evenly distributed or grouped ( Fig. 1A); scutellum in dorsal view abruptly or gradually narrowed to tip, tip pointed, rounded, or slightly inflated ( Figs 2–4). Forewing: With outline elongate oval or broad anteriorly; with ( G. dispar , sp. n., Fig. 2) or without (all other species) distinct honey-comb pattern; membrane (cub, dc1, rc2–3 and areas distal to these cells) contributing about one third or less to forewing length; posterior distal margin rounded or square; clavus twice as long as wide; vein-tracing areoles along margins of a combination of scc, rc1, rc1, rc2, and tc on proximal part of wing absent or present ( Fig. 6A, C); C+Sc narrow ( Fig. 2, G. anaticulus , sp. n.), fairly broad ( Fig. 2, G. clava, sp. n.), broad ( Fig. 2, G. falcis , sp. n.), or very broad ( Fig. 3, G. lutuosus , sp. n.), either of uniform width ( Fig. 2, G. anaticulus , sp. n.) or wider at widest part of wing ( Fig. 2, G. auriculu s, sp. n.); R flattened in distal part (black arrow in Fig. 1B); distal process of R pronounced (white arrow in Fig. 1B) or weakly developed; An1/2 claval process well developed in males (except G. tenuis , sp. n.), overlapping base of An1 wing organ, apex acute and in lateral view pointing dorsad ( Figs 2–4, 6A, D); An1 wing organ on corium strongly inflated in all males, with or without (e.g., Fig. 4, G. tergus , sp. n.) notch articulating with claval process, wing organ rounded, elongate, or acute, with or without median notch; scc about 1.5 times longer than wide ( Fig. 6A) (except ~2 times longer than wide in G. castigatus , sp. n.); scc undivided except in G. dispar , sp. n. ( Fig. 2); rc1 about 3 times as long as wide except in G. dispar , sp. n., ranging from parallel-sided to slightly or much wider anteriorly; rc about 4 or 5 times as long as wide; tc 3 times or more than 3 times as long as wide; part of An1 and Cu forming posterodistal margin of tc of similar width as other veins (e.g., Fig. 2, G. artus , sp. n.) or thicker (e.g., Fig. 2, G. falcis , sp. n.); distal margin of rc1, rc, and tc cells double s-shaped ( Fig. 2, G. anaticulus , sp. n.), s-shaped ( Fig. 1, G. bierigi ), or angular s-shaped ( Fig. 2, G. dispar , sp. n.); rc2–3 almost triangular (e.g., Fig. 2, G. anaticulus , sp. n.), trapezoidal (e.g., Fig. 2, G. castigatus , sp. n.), or rectangular ( Fig. 2, G. dispar , sp. n.); cub oval, almost reaching wing margin or distant form wing margin (except elongate in G. dispar ; Fig. 2); dc1 with basal portion fairly narrow and elongate ( Fig. 2, G. anaticulus , sp. n.) or relatively wide and short ( Fig. 2, G. auriculu s, sp. n.); M beyond Cu short. Legs: males and females with tarsal formula 2-2-3, with claws long except short in G dispar , sp. n. Genitalia ( Fig. 7): Pygophore without lobe on right side ( Fig. 7, G. anaticulus , sp. n.), with small ( Fig. 7, G. artus , sp. n.) or large lobe ( Fig. 7, G. auriculus , sp. n.), or with slightly extended right margin ( Fig. 7, G. castigatus , sp. n.). Vesica about one half loop (e.g., Fig. 7, G. castigatus , sp. n.), between half and one loop (e.g., Fig. 7, G. anaticulus , sp. n.), or one loop (e.g., Fig. 7, G. plurilobus , sp. n.), either not reaching right pygophore margin or surpassing margin, at midpoint thick (e.g., Fig. 7, G. graziae , sp. n.), very thick (e.g., Fig. 7, G. robustus , sp. n.), or more slender (e.g., Fig. 7, G. falcis , sp. n.), tapering to blunt or acute tip, tip itself blunt, bottle-opener shaped (e.g., Fig. 7, G. uncus , sp. n.), slightly s-shaped (e.g., Fig. 7, G. falcis , sp. n.), rounded (e.g., Fig. 7, G. artus , sp. n.), or fairly straight (e.g., Fig. 7, G. tatumbia , sp. n.). Anophoric ridge with process present (e.g., Fig. 7, G. artus , sp. n.) or absent ( Fig. 7, G. castigatus , sp. n.), process short or long, with single lobe curved or relatively straight, or with multiple lobes, tip pointed, blunt, or combination of pointed and blunt. Right paramere curved or straight, tapering towards apex or broad throughout; basal process of paramere broad, narrow, thumb-shaped, or duck head-shaped. Left paramere straight or curved, tapering towards apex or broad throughout, basal process broad or narrow.

Female: As male, but with less well developed An1/2 claval process and without An1 wing organ on corium ( Fig. 1B). Genitalia: with well-developed ovipositor ( Figs 1C, 8). Reservoir of spermatheca kidney-shaped with short spermathecal duct ( Fig. 8).

Distribution. Species of Guapinannus are documented from Central and South America, ranging from Veracruz in Mexico to the southern part of Brazil. The greatest species diversity is found in Central America and the northern areas of South America. Specimens have been collected in tropical wet or moist forest, including cloud forest and mixed hardwood forest, from elevation ranging from 5 meters above sea level in Panama to more than 200 meters in Chiapas, Mexico.

Collecting method. The majority of specimens for which collection method was recorded were collected using leaf litter sifting and Berlese and Winkler extraction methods. A small number of specimens are derived from Malaise and flight intercept traps.

Discussion. The forewing shape, structure with distinct claval suture, and venation set apart Guapinannus from all other genera of Schizopteridae . While Humpatanannus and related genera have not been included in phylogenetic analyses, genitalic features are strikingly different in the two groups, and they are unlikely to be close relatives. The non-genitalic morphology of species of Guapinannus is fairly uniform. Most species are mid-sized and ovoid with some variation, with only few species being distinctly smaller or bigger or markedly elongated ovoid. Some forewing venation features are fairly distinctive and useful as diagnostic characters at the species level, among them the width of the C+Sc and the relative proportion of certain cells. As in many other genera of Schizopteridae , the bulk of diagnostic features is derived from male genitalic structures, in particular the length, curvature, and tip shape of the vesica, shape of the anophoric process, where present, and shape of the pygophore that is expanded into a distinct lobe on the right side in some species. We described as new two species for which only female specimens are known ( G. dispar , sp. n. and G. minutus , sp. n.). Both are distinctive based on non-genitalic morphology and G. dispar , sp. n. in addition is geographically isolated from all other known species of Guapinannus .

As part of this study we discovered a so far undocumented structure on the forewings that is restricted to males that we interpret as a novel male-specific wing organ. This wing organ is located on the posterior margin of the corium, associated with An1, consist of a strongly inflated semicircular or elongate area and in most species features a median notch. SEM observation of this wing organ in one species ( Fig. 6D) shows a tuft of hairlike structures that may represent a trichome. Based on our observations, males of all species of Guapinannus possess this wing organ. Among closely related taxa (based on Knyshov et al., 2020), Caucanannus also have a male-specific wing organ, although in a different position on the forewing ( Weirauch et al., 2020), while wing organs have so far not been documented in Luachimonannus and Kokeshia .