Lygaeus compressipes Fabricius, 1803 by original designation [ = A. latipes (Drury, 1782)].

Medium to large size species (body length 20.00–38.00mm), ranging from pale brown, reddish brown to almost black, but never with metallic reflections. Pronotal disc punctate and tuberculated, humeral angles expanded. Apex of scutellum thickened or with an apparent callosity. Metathoracic acetabulum unarmed. Metafemur strongly incrassate, particularly in males. Metatibia conspicuously expanded.

MALE. General: Medium to large size species, body length 20.00–38.00 mm. General color pale brown to black, dull, never with metallic reflections. — Head: Dorsally flat, lateral margins parallel (Fig. 4A). Tylus sharply pointed, projecting beyond the juga as a compressed plate (Fig. 4A–F). Juga globose, not visible from above, flexed below antenniferous tubercles (Fig. 4B–F). Post-tylar depression visible. Preocellar pits deep. Eyes hemispherical, globose, and protuberant; postocular region barely raised, visible in dorsal view. Ocelli rounded, ocellar tubercles present. Bucculae subtriangular, subrectangular, or rounded to oval, extending posteriorly beyond the anterior third of eyes, but never surpassing them (Fig. 4B–F). Antenniferous tubercles broad, oblique at apex (Fig. 4A). Antennae variable in length; scape cylindrical, stout and slightly curved outwards, simple or with inner side longitudinally ridged; pedicel and basiflagellomere cylindrical; distiflagellomere elongate and narrowly fusiform. Antennomeres decreasing in length from scape to basiflagellomere, distiflagellomere always the longest. Labium variable in length, at most surpassing mesocoxae; labial segment I longest, labial segment IV shorter, labial segments II and III subequal. — Thorax: Pronotum trapezoidal, steeply declined anteriorly; with a distinct collar, frontal angles blunt (Fig. 5B, D); anterior lobe smooth, unarmed or with one or two pairs of small central tubercles, calli almost flat, with a conspicuous longitudinal depression between them; anterolateral margins straight, crenulate or tuberculated; posterior lobe punctate and tuberculated; humeral angles slightly to conspicuously expanded, with or without a spine; posterolateral margins tuberculate or spined; triangular processes present or not. Scutellum punctate and transversely striated; basal region elevated, unarmed or with two small medial tubercles; lateral margins thickened; apex thickened or callouse, paler or yellowish. Thoracic pleura smooth or tuberculated, episterna striated, epimera punctate; metathoracic scent gland auricle well developed, lobes of auricle rounded or elongate, posterior lobe reduced; evaporative area well developed, striated or slightly punctate; supracoxal area of metapleura incrassate posteriorly or not. Prosternum with a deep concavity; mesosternum flat, with an anterior projection; metasternum flat. Hemelytra: extending beyond the apex of abdomen; clavus and corium punctate, with decumbent yellowish setae on punctures, costal margin unarmed or with small setiferous tubercles; membrane glabrous or with scattered short decumbent yellowish setae next to veins. Legs: Coxae rounded and setose. Trochanters tuberculate or unarmed ventrally. Pro- and mesofemora cylindrical and slender, always armed ventrally, with one or two rows of spines or setiferous tubercles that become larger towards the apex. Mesofemur with anterior, dorsal and posterior margins unarmed or with spines or setiferous tubercles. Metafemur thickened, laterally compressed. Anterior and posterior margins unarmed or tuberculated. Dorsal margin with two rows of conical setiferous tubercles. Ventral margin with one row of conical or compressed to flat spines that become larger towards the middle and decrease towards the apex, or that become slightly larger towards the apex; and with one row of smaller conical tubercles that become larger towards the apex; both rows ending on an apical flat dentate projection. Pro- and mesotibiae cylindrical with the distal third almost quadrangular, with semierect yellowish setae over the entire surface, thicker and darker on the margins at apical third. Protibiae unarmed or ventrally spined. Mesotibia unarmed. Metatibia with decumbent and semierect yellowish setae over the entire surface, apex with a crown of stiff setae; dorsal and ventral margins expanded. Metatibial dorsal expansion wide along the entire length of the tibia (Fig. 7A–D), or slightly sinuous or lanceolate, narrowing towards the apex (Fig. 7E); unarmed or tuberculated apically; ventral expansion slightly developed to inconspicuous, lanceolate or convex, restricted to basal third, from where it abruptly narrows, with one row of conical irregular setiferous tubercles that become larger to the apex, basally forming two rows of small tubercles. Tarsomere I as long as the length of the tarsomeres II and III combined; tarsomeres II and III sub-equal. — Abdomen: Connexiva smooth. Lateral margin of abdominal sternites III-IV simple or expanded, developed in three different forms: Type I: sternite III fold-like expanded, segment IV similar expanded or not (Fig. 8C–E), Type II: sternite III expanded in a subtriangular projection, posterior segments never expanded (Fig. 8A); Type III: sternites slightly expanded at posterior margin, segment III and IV more developed (Fig. 8B). Sternites with decumbent and semierect yellowish setae; lateral areas with rugosities or punctures around spiracles; spiracles well developed and closer to the anterior margin of abdominal sternites. Male genitalia: Pygophore globose, dorsal aperture rounded to oval, with or without parandria (Fig. 9B, C); dorsal margin straight to convex and declivent posteriorly in lateral view (Fig. 9A); posterodorsal margin straight or slightly sinuous to rounded, simple or with a median depression (Fig. 9B, C); transversal wall of pygophore short. Parameres (Fig. 9D): basal shank wide, narrowing at middle or at the joint with the arm in lateral view, inner margin simple or expanded; arm perpendicular to basal shank, wide and slightly concave, narrowing to apex, ending in a downwardly directed apical dentiform process. Aedeagus (Fig. 9E): Phallosoma membranous and wide, basal region and lateral margins partially or totally sclerotized; conjunctiva membranous, with a pair of small ventral well sclerotized appendages that arise from the apex of two well-developed digitiform membranous sacs, and with two pairs of dorsal appendages: dorsal appendages I well sclerotized, large, broad and flat and lobulated apically; dorsal appendages II small and membranous, posterior to the dorsal sac. Dorsal sac of conjunctiva large and membranous. Vesica well sclerotized, coiled, with almost two coils. Ejaculatory duct basally coiled, apically straight or hook-shaped. FEMALE. Structure and color similar to male, except for: Metafemur flattened and less developed than in males (Fig. 6A–E); ventral margin with one row of flat spines and one row of smaller conical tubercles that become larger towards the apex, ending on an apical flat dentate projection. Expansions of the metatibia more developed than in males (Fig. 7A–E); dorsal expansion widely dilated, or sinuous and foliaceus, more developed than males, unarmed or tuberculated apically; ventral expansion convex on basal third, and abruptly constricted and running parallel to the apex, unarmed or with one row of conical irregular setiferous tubercles that become larger to the apex. Female genitalia: Sternum VII with plica and fisura present, median lobes well developed (Fig. 10A). First gonocoxae large and triangular, distal angle rounded, proximal region of the outer lateral margin with well-developed dorsal apodeme (Fig. 10B). First gonapophysis elongate, digitiform, inner region of the distal third weakly sclerotized and with short and thick peg-like setae (Fig. 10B), first ramus well sclerotized, straight or slightly curved. Second gonocoxa elongate and narrow in the middle (Fig. 10C). Second gonapophysis elongate and well sclerotized, thickened distal region with a wide lateral tooth (Fig. 10B), second ramus well sclerotized, basally curved. Spermatheca of Type III, subtype D (Pluot-Sigwalt and Moulet 2020) (Fig. 10D, E): Seminal receptacle tubular, sinuous and apically oval or globose; intermedian zone straight, distal region with a well sclerotized and pigmented flange (distal flange), median region (flexible zone) straight, unsclerotized and unpigmented, proximal region with a slightly pigmented flange (proximal flange); spermathecal duct short, with a dilation with thickened and strongly pigmented walls.

Acanthocephla arcuata [Uhler?] 1884 NOMEN DUBIUM. The name A. arcuata was published in Kingsley (1884: 290) as a captioned figure without text. P. R. Uhler is inferred to be the author, as he is the only entomologist listed on the title page. This species has not been cited since its first appearance, no type material is known, and the illustration is insufficient for identification; it is therefore considered a nomen dubium.

Acanthocephala hamata Bergroth, 1924 NOMEN DUBIUM. Bergroth (1924) described Acanthocephala hamata from “Brasilia” with only a brief description of one female, which was deposited in the Museo Paulista, Sao Paulo, Brazil. The type material is lost, and the description is too general to diagnose the species, and no specimens can be reliably assigned; besides this name was cited only in one catalog (Packauskas 2010) since its publication, therefore we considered this species as a nomen dubium.

Acanthocephala fulvitarsa (Herrich-Schäffer, 1851) NOMEN DUBIUM. Herrich-Schäffer (1851) published the name Meatapodius fulvitarsus with a brief diagnosis, based in a single specimen that was deposited in the Zoologische Staatssammlung München, Munich, Germany, and since then the species was only cited in few catalogues from French Guiana. Unfortunately, the type material is lost, and the published diagnosis is insufficient to identify the species, hence we considered it as a nomen dubium.

Lygaeus compressipes Fabricius, 1803 by original designation [ = A. latipes (Drury, 1782)] (Fig. 3A).

A. (Acanthocephala) carioca Olivera et al., 2024

A. (Acanthocephala) equalis (Westwood, 1842)

A. (Acanthocephala) flaviantennata Olivera et al., 2024

A. (Acanthocephala) gamboensis Olivera et al., 2024

A. (Acanthocephala) guatemalena Distant, 1881

A. (Acanthocephala) harryi Olivera et al., 2024

A. (Acanthocephala) latipes (Drury, 1782)

A. (Acanthocephala) nigra Olivera et al., 2024

A. (Acanthocephala) rufa Olivera et al., 2024

A. (Acanthocephala) pleuritica (Costa, 1863)

A. (Acanthocephala) scutellata (Signoret, 1862)

A. (Acanthocephala) spinosa Olivera et al., 2024

Medium size species (body length 20.00–29.00 mm). Species with large eyes in relation to the head height. Antennae slender, as long or longer than the body length; scape with inner side longitudinally ridged. Anterior lobe of pronotum unarmed, posterior lobe with small shiny tubercles on humeral angles, partially anastomosed on posterior region. Humeral angles weakly expanded, acute and spined. Dorsal expansion of metatibia wide and well-developed, ending rounded in both sexes. Abdominal expansion of type II, present only in males.

MALE. General: Slender and medium size species, body length 20.00–29.00 mm. General color brown to black. — Head: Eyes relatively large, almost occupying the entire head height in lateral view (Fig. 4B). Buccula reaching the anterior third of eye. Antennae slender, as long as or longer than the body length. Scape, pedicel, and basiflagellomere with semierect dark stiff setae; distiflagellomere with abundant decumbent yellowish setae and semierect sparse stiff darker setae. Scape longitudinally ridged on inner side; distiflagellomere conspicuously longer than rest of antennal segments, longer than the length of the pedicel and basiflagellomere combined. Labium variable in length, extending to the anterior region of mesosternum to surpassing the mesocoxae. — Thorax: Pronotum (Fig. 5B) with long and erect dark setae, and short decumbent yellowish setae, scattered or forming a conspicuous patch on anterior lobe; anterior lobe and calli unarmed; posterior lobe with small rounded tubercles on humeral angles, partially anastomosed on posterior region (Fig. 11E, F); anterolateral margins slightly crenulate; humeral angles spined and weakly expanded, elevated or not over the pronotal disc; posterolateral margins with small tubercles; triangular processes present. Base of scutellum unarmed or with two small central tubercles. Thoracic pleura never tuberculated; supracoxal area of metapleura incrassate posteriorly. Hemelytra: clavus and corium with punctures along and next to veins and in the center of the cells, veins concolorous or contrasting reddish, costal margin unarmed; membrane glabrous. Legs: Femora with semierect dark setae all over the surface, and erect setae ventrally; with decumbent yellowish setae in some species. Mesofemur with dorsal, anterior and posterior margins unarmed. Ventral margin of metafemur with conical or compressed to flat spines (Fig. 6A). Metatibial dorsal expansion unarmed, widely dilated, wider at basal third, simple or with a notch from where it slightly narrows to expand again to the apex, ending rounded (Fig. 7A); ventral expansion convex, unarmed or tuberculated; anterior and posterior margins always unarmed. Metatarsus contrasting yellowish. — Abdomen: Terga entirely black. Abdominal sternites with expansion of type II: sternite III laterally with a subtriangular expansion (Fig. 8A). FEMALE. Structure and color similar to male, except for: Supracoxal area of metapleural not incrassate. Abdominal sternite expansion absent.

Widely distributed in the New World, known from Central America to southern South America (Argentina, Brazil, Colombia, Costa Rica, Guyana, Jamaica, Surinam, Panama, Peru, and Venezuela (Olivera et al. 2024a, 2024b; Coreoidea Species File 2025).

Acanthocephala guatemalena Distant, 1881 REVALIDATED NAME. Originally described as Acanthocephala declivis var. guatemalena, this species was synonymized under A. declivis by Lethierry and Severin (1894). Posteriorly was regarded as junior synonym of A. alata by Brailovsky (2006), a status retained to the present day. Distant (1881) described and distinguished this variety by the rounded apex of the dorsal expansion of the metatibia, unlike A. declivis (Say) in which is truncated. After examining the type material of these species, we can distinguish A. guatemalena from A. alata and A. declivis by the posterior lobe of pronotum with short, smooth and shiny tubercles partially anastomosed on the posterior region; the humeral angles ending in a subacute spine directed backwards; the unarmed dorsal margin of mesofemur; and the unarmed dorsal expansion of the metatibia with the apex rounded. On the other hand, A. alata and A. declivis present the posterior lobe of pronotum with sparse and small, rounded and not anastomosed tubercles; the humeral angles conspicuously wide and unspined; the dorsal margin of the mesofemur armed with a row of conical setiferous tubercles; and the dorsal expansion of the metatibia tuberculated on the distal third and with the apex straight. Therefore, we remove A. declivis var. guatemalena from synonymy and establish it as a valid species (A. guatemalenastat. nov.) in the subgenus A. (Acanthocephala).

Stål (1870) originally included three species in the subgenus A. (Acanthocephala): A. latipes, A. alata, and A. declivis. However, our cladistic analysis does not support the inclusion of the latter two species, which are herein transferred to A. (Pronoptera). In accordance to our results, we include in the nominotypical subgenus A. (Acanthocephala) the species forming the clade A. Additionally, although A. equalis and A. guatemalena were not included in the phylogenetic analysis, they are assigned to A. (Acanthocephala) based on the presence of the subgeneric autapomorphies, which were assessed from photographs of the type material of these species. This subgenus is easily distinguished as the only one that exhibits a type II abdominal expansion and a wide, well developed dorsal expansion of the metatibia along its entire length in both sexes, with a conspicuously rounded apex.

Acanthocephala heissi Brailovsky, 2006 by present designation (Fig. 3B).

A. (Spinipedia) heissi Brailovsky, 2006

Medium size species (body length 25.00–27.00 mm). Eyes small in relation to the head height. Antennae stout, as long as the body length; scape simple, never with an inner longitudinal ridge. Anterior lobe of pronotum with one pair of small central tubercles, posterior lobe with short and small rounded tubercles all over its surface. Humeral angles expanded, acuminate and spined. Protibia spined. Dorsal expansion of metatibia in both sexes, wide and well-developed, ending in acute angle. Abdominal expansion type III, present only in males.

MALE. General: General color dark brown to black. Medium size specimens, body length 25.00–27.00 mm. — Head: Eyes relatively small, never occupying the entire head height in lateral view (Fig. 4C). Antennae stout, as long as the body length. Scape, pedicel, and basiflagellomere with semierect stiff dark setae; distiflagellomere with abundant decumbent yellowish setae and semierect sparse stiff darker setae. Scape simple, not longitudinally ridged on inner side; distiflagellomere shorter than or equal to the combined length of the pedicel and basiflagellomere. Labium short, not reaching the mesocoxae. — Thorax: Pronotum with long and erect dark setae and short decumbent yellowish setae, never forming a conspicuous patch on anterior lobe; anterior lobe with one pair of small central tubercles, posterior lobe with short and small rounded tubercles; anterolateral margins with well-developed conical tubercles; humeral angles spined and expanded, acuminate, ending in a sharp elongate broad spine, elevated over the pronotal disc; posterolateral margins with well-developed tubercles; triangular processes present. Scutellum with decumbent yellowish setae and erect stiff darker setae; base unarmed. Propleura always tuberculated, meso- and metapleurae unarmed; supracoxal area of metapleura incrassate posteriorly; anterior and posterior lobes of metathoracic scent gland auricle elongate, evaporatorium finely punctate. Thoracic sterna with abundant erect dark setae and semierect and decumbent yellowish setae. Hemelytra: clavus and corium with punctures along and next to veins and in the center of the cells, veins concolorous, costal margin unarmed; membrane glabrous. Legs: Femora with decumbent yellowish setae and semierect dark setae all over the surface, and erect setae ventrally. Mesofemur with anterior, posterior and dorsal margins unarmed. Ventral margin of metafemur with conical and slightly compressed spines, but never flat, that becomes larger towards the middle; anterior and posterior margins with small rounded tubercles of irregular size (Fig. 6B). Protibia ventrally spined. Metatibial dorsal expansion widely dilated, wider at basal third, with a notch from where it slightly narrows to the apex, ending in acute angle, with one or two conspicuous teeth apically; ventral expansion inconspicuous and tuberculated (Fig. 7B); anterior and posterior margins always unarmed. Tarsi brown. — Abdomen: Terga with yellowish markings, never entirely black. Abdominal sternites with expansion of type III: sternites III, IV, V and VI slightly laterally expanded at posterior margin (Fig. 8B). FEMALE. Structure and color similar to male, except for: Supracoxal area of metapleura not incrassate. Anterior margin of metafemur unarmed, posterior margin with small apical tubercles. Ventral expansion of metatibia lanceolate and unarmed (Fig. 7B). Abdominal sternites without expansions.

The subgeneric name refers to the ventrally spined protibia of the type species. The gender is feminine.

Known from Central America (Guatemala) and Mexico (Brailovsky 2006; Olivera et al. 2024b).

This subgenus is easily distinguished as the only one that exhibits a type III abdominal expansion and a tuberculate protibia. Consequently, we establish A. (Spinipedia) (clade B) to accommodate A. heissi.

Metapodius thomasii Uhler, 1872 by present designation (Fig. 3C).

A. (Contrastata) thomasii (Uhler, 1872)

Stout species, medium to large size (body length 27.00–32.00 mm). Eyes small in relation to the head height. Antennae stout, shorter than the body length; scape simple, not longitudinally ridged. Anterior lobe of pronotum with one pair of small central tubercles, posterior lobe with small, rounded tubercles all over its surface. Humeral angles rounded, weakly expanded, ending acutely but spineless. Dorsal expansion of metatibia wide and well-developed in both sexes, ending acute in males and rounded in females. Male with expansion of abdominal sternites type I, developed only anteriorly in sternite III, sternite IV never expanded.

MALE. General: General color dark brown to black, with light brown to yellowish contrasting structures. Medium to large size species, body length 27.00–32.00 mm. — Head: Eyes relatively small, never occupying the entire head height in lateral view (Fig. 4D). Buccula reaching the middle third of the eye. Antennae stout, shorter than the body length. Scape, pedicel, and basiflagellomere with semierect stiff dark setae; distiflagellomere with abundant decumbent yellowish setae and semierect sparse stiff darker setae. Scape simple, not longitudinally ridged on inner side; distiflagellomere shorter than the length of the pedicel and basiflagellomere combined. Labium extending to mesocoxae. — Thorax: Pronotum with scattered semierect yellowish setae and short decumbent yellowish setae, never forming a conspicuous patch; anterior lobe with one pair of small central tubercles between calli; posterior lobe with dark and abundant small rounded tubercles; anterolateral margins with well-developed conical tubercles; humeral angles rounded, ending acutely but spineless, weakly expanded, not elevated over the pronotal disc; posterolateral margins with small tubercles; triangular processes present. Scutellum with decumbent yellowish setae; base with two small central tubercles. Propleura tuberculated, meso- and metapleurae unarmed; supracoxal area of metapleura incrassate posteriorly; anterior lobe of metathoracic scent gland auricle rounded, posterior lobe elongate, evaporatorium with conspicuous rugosities. Thoracic sterna with abundant decumbent and semierect yellowish setae. Hemelytra: clavus and corium entirely punctate, veins concolorous, costal margin unarmed; membrane glabrous. Legs: Femora with semierect dark setae all over the surface, and erect setae ventrally. Mesofemur with dorsal, anterior and posterior margins unarmed. Ventral margin of metafemur with conical and compressed spines, but never flat, that becomes larger towards the middle (Fig. 6C); anterior and posterior margins with small rounded tubercles of irregular size. Protibia unarmed. Metatibial dorsal expansion widely dilated, wider at basal third, from where it narrows to the apex, ending in acute angle, with small tubercles apically; ventral expansion slightly lanceolate and tuberculated; anterior margin unarmed; posterior margin with small conical tubercles (Fig. 7C). Pro-, meso- and metatarsi yellowish to orange. — Abdomen: Terga with a central longitudinal yellowish fringe, never entirely black. Abdominal sternites with expansion of type I: sternite III expanded only anteriorly; sternite IV not expanded (Fig. 8C). FEMALE. Structure and color similar to male, except for: Supracoxal area of metapleura not incrassate. Metatibial dorsal expansion wider than in males, with a notch at basal third, ending rounded (Fig. 7C); ventral expansion slightly convex; posterior margin unarmed. Abdominal sternites expansions absent.

The name Contrastata comes from Latin and means contrast, referring to the marked color contrast between the yellow distiflagellomere, tibiae, and tarsi, and the dark body of the type species Metapodius thomasii Uhler, 1872. The gender is feminine.

Known from North America (Mexico and United States (Brailovsky 2006; McPherson et al. 2011)).

Acanthocephala thomasii was included in A. (Metapodiessa) by Van Duzee (1916). Nevertheless, the results of our analysis do not support this hypothesis, and we transfer A. thomasii to A. (Contrastata) n. subgen. (clade C). This new subgenus can be distinguished by the dorsal expansion of the metatibia, which in both sexes is wide and well developed along its entire length. In contrast, in A. (Metapodiessa), the dorsal expansion of the metatibia is narrow and lanceolate in males, and sinuous or triangular in females, being wide only at the basal third and narrowing toward the apex.

Diactor alatus Burmeister, 1835 by present designation (Fig. 3D).

A. (Pronoptera) alata (Burmeister, 1835)

A. (Pronoptera) brunnea Olivera et al., 2024

A. (Pronoptera) declivis (Say, 1832)

A. (Pronoptera) macrotuberculata Olivera et al., 2024

A. (Pronoptera) maculata Olivera et al., 2024

A. (Pronoptera) mercur (Mayr, 1865)

A. (Pronoptera) panamensis Distant, 1881

A. (Pronoptera) pilosa Olivera et al., 2024

A. (Pronoptera) pittieri Montandon, 1895

A. (Pronoptera) thoracica (Dallas, 1852)

Medium to large size species (body length 27.00–38.00 mm). Eyes small in relation to the head height. Antennae stout, as long as or shorter than the body length; simple, never with inner side longitudinally ridged. Anterior lobe of pronotum unarmed or with one or two pairs of small central tubercles, posterior lobe with rounded tubercles all over its surface. Humeral angles conspicuously expanded, wide, spineless or sharply acute, elevated over the pronotal disc. Dorsal expansion of metatibia wide and well-developed, ending in a straight angle in both sexes. Abdominal expansion of type I; always present in males, sternite III expanded only anteriorly, sternite IV not expanded; absent or present in females.

MALE. General: Medium to large size species, body length 27.00–38.00 mm. General color brown to black. — Head: Eyes relatively small, never occupying the entire head height in lateral view (Fig. 4E). Buccula extended to the posterior third of the eye. Antennae stout, as long as or shorter than the body length. Scape, pedicel, and basiflagellomere with semierect dark stiff setae always present and with or without decumbent yellowish setae; distiflagellomere with abundant decumbent yellowish setae and semierect sparse darker stiff setae. Scape simple, not longitudinally ridged on inner side; distiflagellomere shorter than or equal to the combined length of the pedicel and basiflagellomere. Labium extending to mesocoxae. — Thorax: Pronotum with long and erect stiff setae and short decumbent yellowish setae, more abundant on anterior lobe, but never forming a conspicuous patch; anterior lobe and calli unarmed, or with one or two pairs of small rounded central tubercles next to collar and between calli; posterior lobe with small to large and well-developed rounded tubercles on all its surface; anterolateral margins with well-developed conical tubercles (Fig. 5C); humeral angles conspicuously expanded, wide and spineless or sharply acute, elevated over the pronotal disc (Fig. 5C, E, F); posterolateral margins with small to well-developed tubercles; triangular processes present. Propleura always tuberculated, meso- and metapleurae unarmed; supracoxal area of metapleura incrassate posteriorly. Hemelytra: clavus and corium entirely punctate, veins concolorous, costal margin unarmed or with one or more small setiferous tubercles (Fig. 5F); membrane glabrous or with scattered short decumbent yellowish setae. Legs: Femora with decumbent yellowish setae and semierect dark setae all over the surface, in some with erect setae ventrally. Mesofemur with anterior margin unarmed, and dorsal and posterior margins unarmed or with small tubercles. Ventral margin of metafemur with conical and slightly compressed laterally spines, but never flat, that becomes larger towards the middle (Fig. 6D); anterior and posterior margins unarmed or with small rounded tubercles of irregular size. Protibia unarmed. Metatibial dorsal expansion well-developed, wider at basal third, from where it slightly narrows to the apex, ending in a straight angle, distal third unarmed or with small setiferous tubercles; ventral expansion inconspicuous and tuberculated (Fig. 7D); anterior and posterior margins always unarmed. Tarsi dark brown or yellowish to orange. — Abdomen: Terga with rounded yellowish macula around scent glands scars, never entirely black. Abdominal sternites with expansion of type I: sternite III expanded only anteriorly; sternite IV not expanded (Type I) (Fig. 8D). FEMALE. Structure and color similar to male, except for: Supracoxal area of metapleura not incrassate. Ventral expansion of metatibia slightly convex on the basal third and abruptly constricted and running parallel to the apex (Fig. 7D). Abdominal sternite expansion absent or present, if it is present less developed than in males.

The suffix -ptera meaning wings or wing-like parts, refers to the wide and wing-like pronotal humeral angles of the species included in this subgenus. The gender is neuter.

Known from North America to northern South America (Brazil, Colombia, Costa Rica, Guatemala, Honduras, Nicaragua, Mexico, Panama, and United States (Brailovsky 2006; McPherson et al. 2011; Olivera et al. 2024a, 2024b; Coreoidea Species File 2025)).

Acanthocephala panamensis Distant, 1881 REVALIDATED NAME. Originally described as Acanthocephala declivis var. panamensis, this species was synonymized under A. declivis by Lethierry and Severin (1894). Posteriorly was regarded as junior synonym of A. alata by Brailovsky (2006), a status retained to the present day. After a comprehensive examination of the type specimens of these names, we recognize A. panamensis as clearly distinguishable from A. declivis and A. alata by the presence of abundant rounded and well developed shiny tubercles irregularly arranged on the posterior lobe of pronotum, the conspicuously acute humeral angles, the presence of two medial rounded tubercles at the base of the scutellum, and the presence of small setiferous tubercles on the basal region of the costal margin of hemelytra. Therefore, we remove A. declivis var. panamensis from synonymy and establish it as a valid species (A. panamensisstat. nov.) in the subgenus A. (Pronoptera).

Acanthocephala thoracica (Dallas, 1852) REVALIDATED NAME. Originally described as Metapodius thoracicus, this species was synonymized under A. alata by Stål (1870). Subsequently, A. alata was synonymized under A. declivis by Distant (1881). Finally, Brailovsky (2006) revalidated A. alata as a valid name, leaving M. thoracicus as its junior synonym. After a detailed examination of the type specimens of the aforementioned names, we observed that the specimen described by Dallas can be distinguished from A. declivis and A. alata specimens by the largest body size; the more acute and elongate humeral angles, ending conspicuously acute and directed backwards; and by the absence of the characteristic central macula in the posterior lobe of the pronotum of A. alata. We remove M. thoracicus from synonymy and re-established it as a valid species in the subgenus A. (Pronoptera).

In accordance with our results, we assign the species in the clade D to the new subgenus A. (Pronoptera). We transfer A. alata and A. declivis from A. (Acanthocephala), and include within it the species A. brunnea, A. macrotuberculata, A. maculata, A. panamensis, A. pittieri, and A. pilosa. In addition, the species A. mercur and A. thoracica, although not included in the phylogenetic analysis, are assigned to A. (Pronoptera) based on the presence of the subgeneric autapomorphies. These autapomorphies were observed from the original description and from photographs of the type material. This new subgenus can be distinguished by the conspicuous expansion of the humeral angles and the dorsal expansion of the metatibia. Although the metatibia is wide and well developed along its entire length in both sexes, similar to A. (Acanthocephala), it differs by having a straight apex. In contrast, the metatibia apex in A. (Acanthocephala) is distinctly rounded.

Cimex femoratus Fabricius, 1775 by present designation (Fig. 3E).

A. (Metapodiessa) affinis (Walker, 1871)

A. (Metapodiessa) angusta Olivera et al., 2024

A. (Metapodiessa) angustipes (Westwood, 1842)

A. (Metapodiessa) apicalis (Westwood, 1842)

A. (Metapodiessa) bicoloripes (Stål, 1855)

A. (Metapodiessa) concolor (Herrich-Schäffer, 1841)

A. (Metapodiessa) consobrina (Westwood, 1842)

A. (Metapodiessa) dallasi Lethierry and Severin, 1894

A. (Metapodiessa) distanti Olivera et al., 2024

A. (Metapodiessa) femorata (Fabricius, 1775)

A. (Metapodiessa) granulosa (Dallas, 1852)

A. (Metapodiessa) latiantennata Olivera et al., 2024

A. (Metapodiessa) luctuosa (Stål, 1855)

A. (Metapodiessa) surata (Burmeister, 1835)

A. (Metapodiessa) terminalis (Dallas, 1852)

A. (Metapodiessa) unicolor (Westwood, 1842)

Medium size species (body length 23.00–30.00). Eyes small in relation to the head height. Antennae stout, shorter than the body length; simple, never with inner side longitudinally ridged. Anterior lobe of pronotum unarmed or with one or two pairs of small central tubercles, posterior lobe with rounded tubercles all over its surface. Humeral angles weakly expanded and nearly rounded, obtuse or slightly acute, spineless, not elevated over pronotal disc. Dorsal expansion of metatibia lanceolate or wider and slightly sinuous in males, sinuous or triangular in females, wider to the apex at basal third and narrowing to the apex, ending in acute angle on both sexes. Abdominal expansion of type I, always present in males; absent or present in females, if it is present, only the sternite III is anteriorly expanded.

MALE. General: Medium size species, body length 23.00–30.00. General color pale brown to dark brown. — Head: Eyes relatively small, never occupying the entire head height in lateral view (Fig. 4F). Buccula usually extended to the posterior third of eye. Antennae stout, shorter than the body length. Scape, pedicel, basi- and distiflagellomere with semierect stiff dark setae and with decumbent yellowish setae. Scape simple, not longitudinally ridged on inner side; distiflagellomere shorter than or equal to the combined length of the pedicel and basiflagellomere. Labium extending to mesocoxae. — Thorax: Pronotum (Fig. 5D) with long and erect stiff dark setae and short decumbent yellowish setae, never forming a conspicuous patch; anterior lobe and calli unarmed, or with one or two pair of small rounded central tubercles next to collar and between calli; posterior lobe with rounded tubercles on all its surface; anterolateral margins with conspicuous conical tubercles; humeral angles spineless, obtuse or acute, nearly rounded and weakly expanded, not elevated over the pronotal disc; posterolateral margins with small tubercles; triangular processes short or absent. Scutellum with abundant decumbent yellowish setae and erect stiff darker setae; base unarmed or with two small central tubercles. Thoracic pleura unarmed or tuberculated; supracoxal area of metapleura incrassate posteriorly; anterior lobe of metathoracic scent gland auricle rounded, posterior lobe elongate, evaporatorium with conspicuous rugosities. Thoracic sterna with abundant, erect, dark setae, and decumbent, yellowish setae. Hemelytra: clavus and corium entirely punctate, veins concolorous, costal margin unarmed; membrane glabrous or with scattered short decumbent yellowish setae. Legs: Femora with decumbent yellowish setae and semierect dark setae all over the surface, and erect setae ventrally. Mesofemur with dorsal margin unarmed, and with anterior and posterior margins unarmed or with small tubercles. Ventral margin of metafemur with conical spines that become larger towards the middle and with a median spine conspicuously larger, or with compressed spines that become slightly larger towards the apex (Fig. 6E); anterior and posterior margins with abundant rounded tubercles of irregular size. Protibia unarmed. Metatibia dorsal expansion slightly developed, lanceolate or slightly sinuous, ending in acute angle, distal third always armed with small setiferous tubercles; ventral expansion lanceolate, slightly developed on basal third to inconspicuous, with one row of conical irregular setiferous tubercles that become larger to the apex (Fig. 7E). Pro-, meso- and metatarsi pale brown to yellowish. — Abdomen: Terga with a longitudinal yellowish fringe or rounded maculae around scent glands scars, never entirely black. Abdominal sternites with expansion of type I: sternite III expanded across the entire margin or expanded only anteriorly; sternite IV expanded only anteriorly, or not expanded (Fig. 8E). FEMALE. Structure and color similar to male, except for: Supracoxal area of metapleura simple or incrassate posteriorly, but less developed than in males. Dorsal expansion of metatibia triangular and developed on basal third, or foliaceus and sinuous, wider to the apex at basal third, simple or with a notch from where it narrows to the apex, ending in acute angle, distal third unarmed or with small setiferous tubercles; ventral expansion convex on basal third, and abruptly constricted and running parallel to the apex (Fig. 7E). Abdominal sternites expansion absent or present, if it is present only the sternite III is expanded anteriorly.

Widely distributed in the New World, known from Central America to southern South America (Argentina, Barbados, Brazil, Canada, Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Panama, United States, and Venezuela (Brailovsky 2006; McPherson et al. 2011; Olivera et al. 2024a, 2024b; Coreoidea Species File 2025)).

Acanthocephala granulosa (Dallas, 1852) REVALIDATED NAME. This taxon was originally described as Metapodius granulosus and was synonymized under A. femorata by Barber (1926). After a detailed examination of the type material of both names, we observed that the specimens described by Dallas can be distinguished from A. femorata by having the posterior lobe of the pronotum with abundant and well-developed rounded tubercles, antennae paler than the rest of the body, and the unarmed scutellum, metapleurae, and posterior margin of the mesofemora. In contrast, A. femorata presents the posterior lobe of the pronotum with smaller and sparser tubercles, reddish to brown antennae, and the scutellum, metapleurae, and posterior margin of the mesofemora tuberculated. According to these, we remove from synonymy A. granulosa and re-established it as a valid species in the subgenus A. (Metapodiessa).

Acanthocephala luctuosa (Stål, 1855) REVALIDATED NAME. Originally described as Metapodius luctuosus, this species, was also synonymized under A. femorata by Barber (1926). We examined the type material of these species and we observed that it can be easily distinguished from A. femorata and A. granulosa by the structure of the scape, which is conspicuously broader than the other antennal segments; the coloration pattern of the antennae, which is brown with the distiflagellomere contrastingly yellowish; and by the ornamentation of the pronotum, with the posterior lobe slightly punctate and with small and rounded scattered tubercles. We remove from synonymy A. luctuosa and re-established it as a valid species in the subgenus A. (Metapodiessa).

Stål (1870) included six species in A. (Metapodiessa): A. bicoloripes, A. femorata, A. granulosa, A. luctuosa, A. surata, and A. terminalis. Later, Van Duzee (1916) included A. thomasii (which is transferred to A. (Contrastata) in the present work), and Blöte (1938) included A. consobrina. According to our results, we also include in this subgenus the remaining species forming clade E. In addition, the species A. unicolor, although not included in the phylogenetic analysis, is assigned to A. (Metapodiessa) based on the presence of the subgeneric autapomorphies, which were assessed from photographs of the type specimens. This subgenus can be distinguished by the dorsal expansion of the metatibia, which exhibits distinct sexual dimorphism: in males, the expansion is narrow (lanceolate or slightly sinuous) along its entire length, whereas in females it is wide (sinuous or triangular) only at the basal third, narrowing toward the apex.