Subgenus Edessa Fabricius, 1803

Edessa (part.); Amyot & Serville, 1843: 158; Stål, 1872: 49.

Edessa (Edessa); Kirkaldy, 1909: 154; Almeida et al., 2018: 256; Fernandes et al., 2018: 204; Nunes et al., 2020: 402; Fernandes & Silva, 2021: 630.

Type species: Edessa antilope (Fabricius, 1798) (see Silva et al. 2017 for historic notes).

Diagnose. Dorsal surface usually green to brown (Figs. 24 A,E). Ventral surface yellow to reddish-brown (Figs. 24 B,F), but always lighter than dorsal surface. Pronotum with anterolateral margin concave (Figs. 24 C, 25 C). Humeral angles conical (Figs. 24 A, 26 C, 27 C); somewhat developed and projected, with different sizes (Figs. 24 A,B,C, 26 A,C); apex rounded or slightly acuminated (never acute), shiny, smooth or shagreen and with few punctures; concolorous with pronotum (Fig. 24 E), yellow (Figs. 27 A,C), reddish-brown (Fig. 29 A) or black (Figs. 24 A,C, 26 A,C) color extended ventrally (Figs. 24 D,F, 25 B, 26 B,D, 27 D, 28 F).

Description. Head: Triangular, wider than long (Fig. 24 A,C,E). Mandibular plates sulcate, contiguous ahead of tylus and ventrally curved. Bucculae high, slightly setulose. Rostrum setulose, yellow, segment IV distally black (Figs. 24 D, 25 B, 29 F); segments I<II>III<IV. Antennae setulose, usually lighter than body (Figs. 24 A,B, 27 A,C,E, 28 E).

Thorax: Pronotum trapezoidal, wider than long, densely punctured (Figs. 24 A, 26 A, 27 C, 28 C). Scutellum triangular, densely punctured (Figs. 24 A, 25 A, 26 A, 27 C, 28 C, 29 C). Corium densely punctured, normally with at least one vein contrasting with corium color (Figs. 25 A, 26 A, 27 C, 28 C, 29 A). Membrane of the hemelytron brown with metallic shine (Fig. 24 C). Evaporatorium dull, rugose; peritrema ruga-like (Figs. 25 F, 26 A, 27 D, 28 D, 29 B). Metasternal process longer than wide, bifurcated anteriorly; apex of each arm enlarged (Figs. 24 F, 25 B, 26 D, 27 D, 28 F, 29 B). Legs setulose (Fig. 24 B).

Abdomen: Dorsal surface densely and uniformly punctured, small, deep, with concolorous punctures with its surface. Connexivum exposed, punctured, posterolateral angle slightly projected and dark; with (Figs. 24 A,C,E, 27 C, 28 E) or without contrasting spots (Figs. 26 A,C, 29 A). Segment VII posteriorly projected and acute (Figs. 24 A, 25 A, 26 C, 29 C).

Male genitalia: Pygophore trapezoidal and dorsoposteriorly open (Figs. 1 A, 2 A, 5 A, 20 A). Superior process of the genital cup dark, texture different from genital cup (Figs. 1 D,E, 4 D, 12 D). Parameres yellow usually with dark margins (Figs. 1 D,E, 3 D,E, 4 D,E, 12 D,E, 17 D,E); narrow twisted and sulcate at base; apex expanded in different shapes. Proctiger subcylindrical; dorsal surface sulcate posteriorly; lateral face excavated and setose (Figs. 1 D,C, 3 D,E, 5 D,E, 7 D,E, 15 D,E). Ventral rim excavated medially; expansions delimiting excavation somewhat swollen, slightly developed not reaching level of posterolateral angle (Figs. 1 C, 2 C, 3 C, 20 C, 23 C). Posterolateral angles developed, narrow with apex rounded (Figs. 1 C,E, 2 C,E, 3 C,E, 11 C,E, 17 C,E). Ventral surface punctured (Figs. 1 C, 4 C, 8 C, 18 C, 20 C, 23 C).

Female genitalia: Valvifer VIII convex, setulose, punctured, subrectangular. Valvifers IX trapezoidal; presence of a lateral constriction that results in a median keel. Laterotergite VIII distally spiniform. Laterotergite IX triangular; excavated at base. Segment X rectangular and sulcated (Figs. 1 F, 2 F, 7 F, 12 F, 18 F).

Comments: The nominal subgenus is the largest of Edessa with 79 species (Table 1) that can be separated in several groups of species. Among the species mentioned by Stål (1872) as part of the species group Edessa, only Edessa cervus (Fabricius, 1787), E. antilope (Fabricius, 1798), E. taurina Stål, 1872, E. arabs (Linnaeus, 1758) and E. cylindricornis Stål, 1872 were considered here as part of the subgenus Edessa . Three species groups described to Edessa belong to the nominal subgenus: cervus (Fernandes & van Doesburg 2000b), metallica (Fernandes & Campos 2011) and ovina (Fernandes & Silva 2021) . The nominal subgenus can be separated from most genera of Edessinae and subgenera of Edessa by the shape and color of the humeral angle. A comprehensive diagnose separating them can be found in Almeida et al. (2018) and Nunes et al. (2020). Four genera— Anisoedessa Nunes & Fernandes, 2019, Calcatedessa Silva & Fernandes, 2021, Graziaedessa Eger, 2021, and Mediocampus Thomas, 1994 —were never included in any diagnostic discussion between the genera of Edessinae . The subgenus Edessa can be separated from these four genera by the shape of the humeral angle described above. In all these four genera the humeral angles are dentiform or rounded, slightly developed and flat.

E. sexdens group

(Figs. 1–32)

Diagnosis. Specimens large (13.2–25.2 mm). Dorsal surface green to orange (Figs. 24 C, 27 E). Underside of body dark yellow to orange with transversal brown, green or black lines on thorax and abdomen (Figs. 9 B, 24 B,D, 26 D). Humeral angles as long as wide, with apices black (Fig. 26 E), brown (Fig. 24 A) or concolorous with surface (Fig. 29 E) and slightly curved backwards (Fig. 24 A) or, more rarely, straight (Fig. 24 E). Corium with all veins concolorous (Fig. 24 C) or yellow (Fig. 25 A). Connexival segments with (Fig. 24 C) or without (Fig. 26 A) spots, these spots may (Fig. 24 D) or not (Fig. 26 B) extend to ventral side; with or without small concavities. Legs reddish brown to brown (Fig. 24 B,D). Posterolateral angles of pygophore projected, tumid, medially slightly excavated (Figs. 2 A,B, 11 A,B). Superior process of the genital cup laminar, rectangular, usually continuing ventrally in a carina (Figs. 1 B,E, 6 B); oblique to paramere (Figs. 1 B,D,E, 3 E, 5 E, 20 D). Parameres club-like with three lobes (Figs. 1 D,E, 2 D,E, 4 D,E, 6 D,E, 11 D, E, 13 D, E).

Description. Head: Punctured, punctures small, deep, sparsely distributed and concolorous with surface, brown or black. Mandibular plates deeply sulcate medially (Figs. 24 A,C, 25 A,C, 26 C,E). Bucculae tall, receiving nearly all first rostral segment (Figs. 24 B, 25 D, 26 F, 27 F, 29 F); margin rounded (Figs. 24 B, 25 B, 26 D). Rostrum brown (Fig. 24 B). Antennomers, II+III shorter than IV; IV subequal in length to V (Fig. 24 D).

Thorax: Pronotum densely and uniformly punctured (Figs. 24 A, 25 C, 26 A, 27 C, 28 C, 29 A, 30 A); punctures small, deep, and brown, black or concolorous with surface; cicatrices and apices of humeral angles not punctured. Anterolateral angle of pronotum with a small tooth concolorous with surface (Figs. 24 A,B, 26 A,B, 28 E,F, 29 C). Anterolateral margin grooved, sulci extending ventrally (Figs. 24 F, 25 E,F, 26 A,B, 29 E,F). Scutellum with punctures deep, brown, black or concolorous with surface and evenly distributed; apex acute (Figs. 24 C, 25 C, 26 C, 27 A, 29 A, 30 E). Corium with punctures deep, brown or concolorous with surface and uniformly distributed (Figs. 24 A, 25 A, 27 E, 28 C). Membrane brown (Fig. 24 E). Ventral surface of thorax with brown or black stripes (Figs. 24 B, 25 B); propleuron, mesopleuron and metapleuron with brown, green, black or concolorous punctures. Evaporatorium rugulose, concolorous or with contrasting stripes with pleura, opaque (Figs. 24 D, 25 B). Peritreme reaching 3/4 of the distance from ostiole of scent gland to lateral margin of metapleuron (Fig. 25 B). Metasternal process as long as wide, raised anteriorly; anterior apex of each arm truncated and laterally expanded (Figs. 1 G, 24 B,F, 25 F, 26 B,F, 27 D, 28 F); anterior face of each arm rounded, flat or excavated.

Abdomen: Dorsally brown with punctures small, deep and evenly distributed. Connexivum (Figs. 25 C,E) densely and uniformly punctured only on concavities; or densely and uniformly punctured except the outer margin; punctuation black or concolorous with connexivum. Abdominal segment VII with posterolateral angles developed, with apices acute and black (Fig. 24 A), brown (Fig. 25 A), green (Fig. 9 A) or concolorous with surface (Fig. 26 A). Ventrally, abdomen medially raised sometimes with brown to black spots forming or not a complete band (Fig. 26 B,D,F). Intersegmental areas of abdomen black (Fig. 24 D) or brown (Fig. 25 D). Pseudosutures black (Fig. 24 B), brown (Fig. 26 B) or concolorous with surface (Fig. 24 C). Rounded shallow excavation near each spiracle. Spiracles elliptical (Fig. 24 B).

Male genitalia: Pygophore trapezoidal (Figs. 1 A, 2 A, 3 A). Dorsal rim sinuous, wrinkled, ending on each side near posterolateral angles (Figs. 1 A,B, 14 A,E, 15 A,E). Posterolateral angles developed, inner part brown, strongly punctured (Figs. 1 A,B, 2 D,E, 6 D,E); apices rounded (Figs. 1 A,B, 7 D, 12 D). Proctiger laterally constricted with dense band of short setae along the edge (Figs. 1 D,E, 12 D,E, 15 D,E, 20 D,E) of the subtriangular (Figs. 1 D,E) or subelliptical (Figs. 2 D,E) posterior face. Ventral rim setulose (except Edessa (E.) bella) with median excavation U- or V-shaped (Figs. 1 C, 7 C, 11 C, 12 C). Ventral surface (Fig. 1 C) punctured; punctures large, shallow, sparse, dark (Fig. 1 C) or concolorous with surface (Fig. 2 C).

Female genitalia: Valvifers VIII setulose and punctured (Fig. 1 F); punctures dark (Fig. 1 F) or concolorous with surface (Fig. 7 F); convex, medial part with a conspicuous protuberance (Fig. 1 F); distal margin straight (Fig. 1 F). Valvifers IX (Fig. 1 F) trapezoidal, large, carinated (except Edessa (E.) miniata; Edessa (E.) electa (Fig. 5 F); Edessa (E.) nigricornis; Edessa (E.) nigroangulata, Edessa (E.) schirmeri). Laterotergites VIII uniformly punctured; punctures small, shallow, concolorous with surface; distal margin acutely projected posteriorly, apices black (Figs. 1 F, 6 F, 8 F). Laterotergites IX excavated at the base; apices black surpassing sclerite uniting laterotergites VIII, in ventral view (Figs. 1 F, 5 F).

Comments: The species belonging to the E. sexdens group have a large morphological variety, but also groups of very similar species. Despite this, the group has a set of characteristics that are not found in other species of the subgenus Edessa: connexival segments with a pair of dark spots (Fig. 24 A); superior process of the genital cup laminar dorsally and continued ventrally by a carina (Fig. 1 B,E); and oblique to the parameres (Fig. 1 B,D,E); parameres with three lobes (Fig. 1 D,E) (Silva 2017). Other characteristics are also related to the group, such as: anterior arms of the metasternal process with excavated apex (Fig. 1 G); ventral side of the abdomen raised medially (Fig. 24 B); rounded shallow excavation near each spiracle (Fig. 24 B); union of dorsal rim with posterolateral angles grooved (Fig. 2 A,B). However, within the E. sexdens group there are species that do not have the entire set of characteristics mentioned above. A few species do not have the spots on connexival segments (Fig. 26 A); and others do not have the ventral carina of the superior process of the genital cup (Figs. 4 B,E, 7 B, 8 B).

Distribution (Figs. 30, 31, 32): MEXICO: Chihuahua, Jalisco, Veracruz, Morelos, Guerrero, Quintana Roo, Chiapas, Mexico City; JAMAICA; GUATEMALA: El Petén, Izabal; HONDURAS; NICARAGUA: Léon; COSTA RICA: Guanacaste, Alajuela, Heredia, Limón Puntarenas, San Jose; PANAMA: Bocas Del Toro, Colón, Panama, West Panama; COLOMBIA: Bogotá, Cauca; VENEZUELA: Zulia, Carabobo, Aragua, Caracas, Monagas, Bolívar, Amazonas; TRINIDAD AND TOBAGO; GUYANA; SURINAME: Marowijne, Paramaribo; FRENCH GUYANA: Cayenne; BRAZIL: Pará, Amazonas, Maranh„o, Mato Grosso, Acre, Rondônia, Góias; ECUADOR: Napo; PERU: Satipo; BOLIVIA: La Paz, Santa Cruz, Cochabamba, ARGENTINA: Formosa.

Identification key to the species of the E. sexdens group

1- Body predominantly orange to red with large black areas on pronotum, corium and ventral surface of abdomen (Fig. 27 E,F)...................................................................... Edessa (E.) miniata Westwood, 1837

- Body predominantly green to brown with dark marks usually present only on humeral angles, connexivum, and stripes on ventral surface of abdomen (Figs. 24 C,D; 9 A,B; 29 E,F)........................................................ 2

2- Ventral surface of the body with alternated yellow and green stripes; humeral angles narrow and laterally directed (Fig. 9 A,B)................................................................... Edessa (E.) imbecilla Breddin, 1903

- Ventral surface of the body yellow to orange usually with brown lines, sometimes without lines; humeral angles different from above (Figs. 25 C,D; 29 A,B)........................................................................... 3

3- Humeral angles long and sloping forward (Figs. 25 A; 26 C,E)................................................. 4

- Humeral angles usually short and slightly sloping backward or directed sideward (Figs. 26 A; 28 E)................... 8

4- Humeral angles straight, posterior margin slightly crenulated (Fig. 25 A,B).......... Edessa (E.) cylindricornis Stål, 1872

- Humeral angles distally curved backward, posterior margin smooth (Fig. 26 C,E).................................. 5

5- Humeral angles narrow, about 4 x longer than wide (Fig. 26 C,D); ventral surface with contrastingly black wide stripes (Fig. 26 D)................................................................... Edessa (E.) helicoceras Breddin, 1905

- Humeral angles wide, about twice longer than wide; ventral surface usually with narrow brown to black stripes, if broad, stripes are brown (Figs. 24 C,D; 26 E,F; 27 C,D)................................................................. 6

6- Humeral angles dorsally mostly green to olive green with more than distal third black (Fig. 26 E,F)............................................................................................. Edessa (E.) helix Erichson, 1848

- Humeral angles dorsally mostly yellow or with apices black (Figs. 24 C,D; 27 C,D)................................ 7

7- Humeral angles yellow without black apices (Fig. 27 C,D); connexival segments without black marks (Fig. 27 C); ventral stripes on abdomen broad (Fig. 27 D)................................... Edessa (E.) lewisi Fernandes & Silva, 2015

- Humeral angles with apices black (Fig. 24 C,D); connexival segments with black spots (Fig. 24 C); ventral stripes on abdomen narrow (Fig. 24 D)................................................... Edessa (E.) bella Fernandes & Silva, 2015

8- Connexival segments without dark spots (Figs. 26 A; 28 A)................................................... 9

- Connexival segments with dark spots (Figs. 24 A,E; 29 C,E)................................................. 13

9- Humeral angle with black apices (Figs. 26 A; 28 A)......................................................... 10

- Humeral angles with yellow to light brown apices (Figs. 25 E; 29 A)........................................... 11

10- Humeral angles barely developed, black part longer than wide (Fig. 26 A,B); evaporatorium darker than thorax (Fig. 26 B)......................................................................... Edessa (E.) gentilitia Distant, 1890

- Humeral angles little developed, black part as long as wide in dorsal view (Fig. 28 A,B); evaporatorium concolorous with thorax (Fig. 28 B)........................................................... Edessa (E.) nigriclava Walker, 1868

11- Evaporatorium dark brown (Fig. 29 B); ventral surface of abdomen with wide transversal and longitudinal dark brown to black stripes (Fig. 29 B)...................................................... Edessa (E.) phoenicopus Dallas, 1851

- Evaporatorium concolorous with thorax (Figs. 25 F; 27 B); ventral surface of abdomen with narrow transversal black stripes (Figs. 25 F; 27 B).................................................................................... 12

12- Paramere anterior process truncated (Fig. 6 D,E); superior process of genital cup narrow, short and continuing ventrally in high crenulate carina (Fig. 6 B)....................................................... Edessa (E.) fallenii Stål, 1872

- Paramere anterior process developed and triangular (Fig. 12 D,E); superior process of genital cup large, continuing ventrally in a short carina crenulate (Fig. 12 B)........................................... Edessa (E.) leucocera Dallas, 1851

13- Apices of humeral angles concolorous with dorsal surface, yellow or light brown (Figs. 24 A,E; 29 C,E)............... 14

- Apices of humeral angles black or blackish (Figs. 25 C; 28 C,E)............................................... 17

14- Humeral angles directed sideward (Fig. 24 E,F); spots on connexivum small, rounded and not reaching any of the margins separating segment (Fig. 24 E)............................................... Edessa (E.) congrua Walker, 1868

- Humeral angles directed backwards (Figs. 24 A; 29 C,E); spots on connexivum large, subrectangular and usually reaching at least one of the margins separating segments (Figs. 24 A; 29 C,E)............................................. 15

15- Apices of humeral angles not clearly globose, acuminated and short (Fig. 29 C,E)................................. 16

- Apices of humeral angles clearly globose (Fig. 24 A,B)............................. Edessa (E.) alces Erichson, 1848

16- Specimens large (about 20 mm) (Fig. 29 E,F); venter uniformly dark yellow or orange (Fig. 29 F); apices of anterior bifurcation of metasternal process rounded (Fig. 23 G)....................................... Edessa (E.) urus Erichson, 1848

- Specimens small (around 15 mm) (Fig. 29 C,D); venter uniformly greenish yellow or ochraceous (Fig. 29 D); apices of anterior bifurcation of metasternal process laterally expanded (Fig. 20 G)................... Edessa (E.) sexdens Fabricius, 1803

17- Body olive brown (Figs. 28 C,D); humeral angles robust, globose, almost as long as wide at base (Fig. 28 C,D); venter reddish brown (Fig. 28 C)......................................................... Edessa (E.) nigricornis Stål, 1862

- Body green to olive green (Figs. 25 C; 28 E); humeral angles not as robust or globose as above, wider at base than long (Figs. 25 C; 28 E); venter greenish yellow or orange (Figs. 25 D; 28 F; 10 F,G)........................................ 18

18- Black spots on connexival segments large (Fig. 28 E); stripes on intersegmental areas solid black (Fig. 28 F)........... 19

- Black spots on connexival segments smaller than above (Fig. 25 C); stripes on intersegmental areas faded brown (Fig. 25 D) .................................................................................................. 21

19- Intersegmental stripes of abdomen complete (Fig. 22 B)......................... Edessa (E.) subandina Breddin, 1904

- Intersegmental stripes of abdomen incomplete (Figs. 19 B, 28 F).............................................. 20

20- Connexival spots large and expanded ventrally (Fig. 28 E,F); pseudosutures faded brown (Fig. 28 F); intersegmental stripes of abdomen narrow (Fig. 28 F)................................... Edessa (E.) nigroangulata Fernandes & Silva, 2015

- Connexival spots small and not expanded ventrally (Fig. 19 A,B); pseudosutures solid black (Fig. 19 B); intersegmental stripes of abdomen wide (Fig. 19 B)............................................... Edessa (E.) schirmeri Breddin, 1904

21- Connexivum greenish-yellow, dark spots triangular (Fig. 25 C); apex of anterior bifurcation of metasternal process straight (Fig. 5 G); propleuron with long dark stripe (Fig. 25 D)............................. Edessa (E.) electa Walker, 1868

- Connexivum orange, dark spots rectangular (Fig. 10 F); apex of anterior bifurcation of metasternal process rounded (Fig. 11 G); propleuron with short dark stripe (Fig. 10 B,G)................................... Edessa (E.) lavata Breddin, 1903

Species description