Cyrtocapsus Reuter

Henry, Thomas J., 2022, Revision of the New World Plant Bug Genus Cyrtocapsus (Heteroptera: Miridae Bryocorinae: Eccritotarsini), with New and Revised Synonymies, Lectotype Designations, and Descriptions of 12 New Species, Zootaxa 5154 (1), pp. 1-48 : 5-8

Cyrtocapsus Reuter

Cyrtocapsus Reuter 1876: 70 . Type species: Cyrtocapsus caligineus Stål, 1859 , by monotypy.

Pirithous Distant 1884: 302 . Type species: Pirithous pallipes Distant, 1884 , by monotypy. Synonymized by Reuter 1892: 393.

Hemisphaerodella Reuter 1908b: 297 . Type species: Hemisphaerodella mirabilis Reuter, 1908b , by original designation. Synonymized by Henry & Carvalho 1987: 291.

Miccus Bergroth 1910: 65 . Type species: Miccus elutipes Bergroth, 1910 , by original designation. Synonymized by Carvalho 1951: 55.

Lopesiella Wygodzinsky 1946:334 . Type species: Lopesiella mirabilis Wygodzinsky, 1946 , by original designation. Synonymized with Hemisphaerodella by Carvalho 1955: 223.

Diagnosis. Recognized by the relatively small size (2.50–4.90 mm); prominent eyes extending laterally beyond the anterior angles of the pronotum (e.g., Figs. 3 , 10, 17 ); protruding, apically acute clypeus; rugose and transversely striate frons ( Figs. 3, 4 ); distinctly punctate pronotum; prominent calli separated by a wide, deep puncture or pit ( Fig. 3 ); dull, impunctate hemelytron, with a single cell on the membrane; extensive silvery, sericeous setae dorsally and laterally on thorax; and mostly C-shaped left parameres (e.g., Figs. 61, 63 , 73 ) and oval to more elongate ( Figs. 64 , 90 ) and sometimes L-shaped right parameres (e.g., Figs. 68 , 84 ).

Description. Male: Macropterous, small, elongate oval species. Total length of included species from apex of clypeus to apex of hemelytral membrane 2.50–4.32 mm; length from apex of clypeus to cuneal fracture 1.80–2.76 mm; widest point across hemelytra 1.23–1.82 mm. General coloration pale yellowish brown to black, often with paler areas on head, pronotum, and hemelytron. COLORATION. Head: Yellowish brown to uniformly black; frons yellowish brown, sometimes with median line dark brown or black, with inner margin along eyes paler; clypeus usually dark; sides and undersurface often paler yellowish brown to white; labium pale yellowish white to white, sometimes tinged with red; antenna pale yellowish white, white, to black. Pronotum: Yellowish brown, brownish orange, dark brown to uniformly black; basal margins on dark brown to black species sometimes paler brown to brownish orange. Hemelytron: Brownish orange to black; often with the embolium and apical half of corium paler yellowish brown to brownish orange. Legs: Pale yellow, yellowish white, white, or black; coxae pale to dark brown; femora pale to nearly uniformly dark; tibiae pale, dark at only bases, to uniformly black. SURFACE AND VESTITUTRE. Head: Rugose to finely punctate; frons often with rows of transverse striations; usually with rows of silvery sericeous setae along basal margin and median line, inner margins of eyes, and across transverse striations. Pronotum: coarsely punctate on disc, more finely punctate on collar and across calli; silvery sericeous setae dense across calli and collar, less dense and usually finer on disc. Hemelytron: Impunctate, dull; with scattered silvery sericeous setae, often with distinct clusters forming loose patterns and a glabrous spot at middle of clavus. Ventral surface: Pleural areas of thorax and basal areas of abdomen usually with clusters of silver sericeous setae. STRUCTURE. Head: Somewhat triangular, clypeus apically acute; eyes prominent, extending laterally just beyond anterior angles of pronotum. Pronotum: Trapeziform, anterior margin much narrow than posterior margin; laterally straight to concave, with posterior angle flaring, margin with or without a carina lateral to calli; posterior margin nearly straight to weakly convex; calli prominent and separated by a deep, wide puncture or pit. Hemelytron: Macropterous, usually strongly bent downward at cuneal fracture; membrane entire with a single large areole. Legs: Slender, lacking tibial spines.

Male genitalia: Left paramere varying from a stout and strongly C-shaped (e.g., Figs. 63 , 75, 77 ) to a more slender, elongate, open C-shape, often with an apical hook (e.g., Figs. 58 , 71, 82 ). Right paramere varying from oval (e.g., Figs. 64 , 76 , 90), broadly rounded (e.g., Figs. 79 , 98), to elongate, sometimes becoming somewhat L-shaped and apically rounded or truncate (e.g., Figs. 60, 68 , 84 ). The endosoma, as in most Bryocorinae , is comprised of a simple, sclerotized tube-like aedeagus, lacking any accessory spicules or sclerites, encased in membrane (e.g., Figs. 59 , 78, 83 ), varying only slightly at the apex.

The parameres are distinctive for certain species, such as C. atrametatibialis with an open C-shape left paramere ( Fig. 61 ) and an elongate, apically rounded right paramere ( Fig. 62 ); C. haitianus with a mitt-shaped right paramere ( Fig. 79 ); or C. spatulatus with a broad, paddle-shaped right paramere ( Fig. 98 ). Other species have quite similar parameres, such as C. caligineus ( Figs. 63, 64 ), C. grenadaensis ( Figs. 75, 76 ), and C. pallipes ( Figs. 89, 90 ), making it necessary to use external morphology in combination with genitalia to distinguish species.

Female: Similar to male in color, texture, and vestiture. Cyrtocapsus atrametatibialis is the one exception, with sexually dimorphic females having entirely pale yellowish-white hind legs, whereas males have the apical third of the hind femur and the entire hind tibia fuscous to black. Total length of included species from apex of clypeus to apex of hemelytral membrane 2.60–4.90 mm; length from apex of clypeus to cuneal fracture 1.80–2.73 mm; widest point across hemelytra 1.23–1.89 mm.

Female genitalia: Labiate plate ( Figs. 99, 101 ). Posterior wall ( Figs. 100, 102 ). The female genitalia are simple and do not appear to have any characters useful in distinguishing species.

Discussion. The descriptions of the genera Pirithous Distant (1884) and Miccus Bergroth (1910) reflect the confusion previous authors faced in placing species now included in Cyrtocapsus , suggesting they overlooked Reuter’s (1876) original description. Pirithous is not only a junior synonym of Cyrtocapsus , but also a junior homonym of Pirithous Keyserling, 1883 (Arachnida) . Miccus , with M. elutipes as the type species, is a junior synonym. Hemisphaerodella and Lopesiella were based on the nymphal stages of what Henry & Carvalho (1987) considered as C. caligineus and what I now show is C. pallipes , based primarily on distribution.

I have maintained a rather broad definition of the genus for the included species, with the realization that several of them may eventually be placed in new or separate genera following eventual phylogenetic analyses. For the time being, however, I think it is best to maintain a conservative interpretation until more specimens of the more poorly known species are discovered and additional information on hosts and biology are available to help interpret relationships.

The relationship of Cyrtocapsus with other genera within the Eccritotarsini is unclear. Numerous genera have a distinct puncture or pit between the calli. The overall elongate-oval body, extended head with a prominent clypeus, punctate pronotum, and impunctate hemelytra are superficially similar to species of the genera Eurychilella Reuter , Pachymerocerista Carvalho , Pycnoderes Guérin-Menèville , and Sixeonotus Reuter. The silvery sericeous setae on the head, pronotum, and hemelytra, however, are found in only a few genera, including Eurychilella , Pycnoderes , and Sixeonotus . Even so, the setae in these genera are much finer and do not form the distinct patterns or patches found in most species of Cyrtocapsus . A cursory review of the male genitalia of the species in these genera showed that they are quite different from those found in species of Cyrtocapsus , though a much broader survey of the tribe is to needed to better understand the genitalic affinities and resolve the phylogenetic placement of this genus. Ferreira and Henry’s (2011) key to the eccritotarsine genera of Minas Gerais will distinguish Cyrtocapsus from all similar Neotropical genera.

The following key is completely revised from previous versions provided by Carvalho (1954, 1985) and Henry & Carvalho (1987). I note that Henry & Carvalho (1987) erred in couplet 9 of their key. The first rung (running to C. andinus Carvalho [now C. caligineus in this work]) should read “Second antennal segment 0.70 mm long...,” not “ 2.2 mm.” The second rung (running to C. grenadensis Carvalho ) should have read “Second antennal segment less than 0.50 mm long...,” not “2.00 mm.” That key, however, is now obsolete because of the new species and the numerous nomenclatural changes presented in this paper and should not be followed.

Key to the species of Cyrtocapsus Reuter

1. Hind femur brown to black, at least on apical third (and sometimes paler ventrally); tibiaefully or in part pale or all dark brown to black............................................................................................. 2

– Hind femur pale yellow to yellowish white with, at most, a short fuscous streak on either side near apex; tibiae pale....... 9

2. Only hind femur with brown or black areas................................................................. 3

– All femora with brown or black areas..................................................................... 5

3. Dorsal apical half of hind femur and entire hind tibia fuscous or black (male); distribution: Bolivia.................................................................................... C. atrametatibalis Henry , n. sp. ( Figs. 10, 11 )

– Hind femur partially to entirely black; hind tibia uniformly pale or fuscous only at base …............................4

4. Body uniformly black; hind femur uniformly fuscous or black; hind tibia pale; distribution: Colombia...................................................................................... C. columbiensis Carvalho ( Figs. 15, 16 )

– Body dark brown, with inner margins of eyes, basal margin of pronotum, embolium, and apex of corium yellowish brown; apical third of hind femur and base of hind tibia fuscous or black; distribution: Costa Rica and Mexico............................................................................... … C. metafemoratus Henry , n. sp. ( Figs. 40, 41 )

5. Middle and hind coxae dark brown; calli shiny, impunctate, glabrous............................................ 6

– Middle and hind coxae pale yellow to yellowish white; calli punctate, covered with silvery sericeous setae.............. 7

6. Large species, male length 3.72–4.08 mm; females, 3.80–4.16 mm; hind femur fuscous mostly on sides of apical two thirds, with dorsal and ventral surfaces frequently paler; distribution: Panama........... C. schaffneri Henry , n. sp. ( Figs. 52–54 ) Small species, male lengths 2.85–2.94 mm (females unknown); hind femur entirely dark brown to fuscous; distribution: Costa Rica................................................................ C. glabratus Henry , n. sp. ( Figs. 27, 28 )

7. Antennal segment II dark brown or black; hind tibia uniformly fuscous, sometimes paler at apex; distribution: Bolivia and Pará, Brazil................................................................. C. paraensis Carvalho ( Figs. 46, 47 )

– Antennal segment II uniformly pale; apical halves of all tibiae pale or white....................................... 8

8. Hemelytron black, with embolium only slightly paler; distribution: Tabasco, Mexico...................................................................................................... C. marginatus (Distant) ( Figs. 36, 38 )

– Hemelytron dark brown, with embolium contrasting yellowish brown to brownish orange; distribution: Mexico (Veracruz) and Venezuela................................................................ C. femoralis Reuter ( Figs. 23, 24 )

9. Pronotum and hemelytron predominately dark brown, fuscous, or black......................................... 10

– Pronotum and hemelytron yellowish brown or brownish orange, if dark brown then embolium contrastingly paler yellowish brown or brownish orange............................................................................. 22

10. All coxae pale yellowish white, or white, with bases of middle and hind coxa sometimes dark brown.................. 11

– Only front coxa at least partially yellowish white to white, middle and hind coxae uniformly dark brown............... 20

11. Antennal segments I and II uniformly or in large part fuscous or black; hind femur sometimes with a fuscous streak on either side near apex....................................................................................... 12

– Antennal segment I white; segment II white, sometimes with apex fuscous; hind femur without a fuscous streak near apex... .................................................................................................. 14

12. Antennal segments I–III uniformly black or reddish brown; underside of head white, contrasting with fuscous clypeus; veins on hemelytral membrane with silvery sericeous setae; hind femur with a fuscous streak on inner and outer surface of apical half; right paramere mitt-shaped ( Fig. 79 ); distribution: Dominican Republic and Haiti......................................................................................................... C. haitianus Carvalho ( Figs. 32, 33 )

– Antennal segment I pale, with apical third dark brown or fuscous; segment II largely fuscous, with only base pale; underside of head fuscous to brownish orange; veins on hemelytral membrane lacking silvery sericeous setae; femora with or without a fuscous streak........................................................................................13

13. Pronotum uniformly black; hind femur pale yellowish white, without fuscous markings; right paramere ( Fig. 98 ) broadly rounded; distribution: Bolivia and Brazil..................................... C. spatulatus Henry , n. sp. ( Fig. 55 )

– Pronotum dark brown through middle, pale yellowish brown laterally; hind femur with a brown streak on either side of apical fourth; right paramere elongate and slender; distribution: Goiás, Brazil............. C. discalis Henry , n. sp. ( Figs 18, 19 )

14. Basal halves of middle and hind coxae brown; apical third of antennal segment II fuscous; distribution: Peru.................................................................................... C. apicalis Henry , n. sp. ( Figs. 8, 9 )

– All coxae uniformly pale or white, with only bases of middle and hind coxae sometimes narrowly brown; antennal segment II uniformly pale or white............................................................................... 15

15. Hemelytron fuscous or black, with contrastingly pale brownish-orange embolium; apical two thirds of hind tibia brown; distribution: Peru.......................................................... C. costatus Henry , n. sp. ( Fig. 17 )

– Hemelytron uniformly fuscous or black; hind tibia pale...................................................... 16

16. Lower half of head, apical third of cuneus, and usually last female genital segment and ovipositor white; distribution: British Virgin Islands, Puerto Rico, U.S. Virgin Islands................................ C. lazelli Henry , n. sp. ( Figs. 35, 36 )

– Lower half of head pale brown or brownish orange but never white; cuneus, female genital segment, and ovipositor fuscous or black............................................................................................. 17

17. Lateral margin of pronotum lacking a distinct carina adjacent to calli; distribution: Bolivia (female)...................................................................................... C. atrametatibialis Henry , n. sp. ( Fig. 12 )

– Lateral margin of pronotum with a distinct carina lateral to calli............................................... 18

18. Apex of corium with a triangular yellowish-white area bordering cuneus; basal margin of pronotum usually paler dark yellowish brown or brownish orange; distribution: Argentina, Brazil, Chile, and Paraguay........ … C. rostratus Reuter ( Figs. 49–51 )

– Paler area at apex of corium narrow, not well-defined or absent; pronotum uniformly fuscous to black................. 19

19. Small species, male lengths 3.04–3.48 mm, bases of middle and hind coxae often narrowly brown; left paramere ( Fig. 67 ) broad through middle and apically hooked; right paramere ( Fig. 68 ) L-shaped, elongate, and apically rounded or truncate; distribution: northern South America, Central America, Mexico, and Jamaica.......... C. elutipes (Bergroth) ( Figs. 20–22 )

– Larger species, male lengths 3.49–4.12 mm, coxae uniformly white; left paramere ( Fig. 85 ) stout, C-shaped; right paramere ( Fig. 86 ) elongate, thickened through main body, narrowing and pointed apically; distribution: Costa Rica, Mexico, Panama.................................................................. C. mesoamericanus Henry , n. sp. ( Fig. 39 )

20. Anterior coxa entirely pale or white; embolium black; distribution: North, Central, and South America, and West Indies............................................................................. C. pallipes (Distant) ( Figs. 44, 45 )

– Anterior coxa brown, except narrowly at apex; embolium paler brown, at least basally............................. 21

21. Larger species, length 3.90 mm or more; antennal segment II long, length 0.66–73 mm; embolium and apical half of corium yellowish brown; distribution: Argentina, Ecuador, Paraguay, and Peru................. C. caligineus (Stål) ( Figs. 13, 14 )

– Smaller species, length 3.50 mm or less, antennal segment II shorter, length 0.56–0.59 mm; hemelytra, including corium, uniformly dark brown, embolium entirely to pale on basal half only; distribution: Costa Rica, Dominica, Grenada, and Mexico............................................................. .. C. grenadensis Carvalho ( Figs. 29, 30 )

22. Pronotum yellowish brown to brownish orange, calli sometimes fuscous........................................ 23

– Pronotum, including calli, reddish brown to dark brown, disc sometimes with basal margin paler yellowish brown....... 26

23. Area across collar and calli fuscous to black, contrasting with brown to brownish-orange disc........................ 24

– Area across collar and calli yellowish brown or brown orange, concolorous with disc, at most with median line fuscous... 24

24. General color brownish orange; corium dark brown on basal two thirds, yellowish brown apically; distribution: Bolivia and Paraguay............................................................... C. paraguaiensi s Carvalho ( Fig. 48 )

– General color brown; hemelytron uniformly brown, accented with costal vein, claval suture, and, inner margin of clavus fuscous; distribution: Honduras........................................... C. suturalis Henry , n. sp. ( Figs. 56, 57 )

25. Pronotum yellowish brown to brownish orange, with median line between calli black; scutellum, clavus and inner margin of corium dark brown or fuscous; distribution: Guyana.................................. C. guianus Carvalho ( Fig. 31 )

– Pronotum, scutellum, and hemelytra uniformly yellowish brown, with only claval suture accented with dark brown or fuscous; distribution: Bahia , Brazil............................................... C. nordestinus Carvalho ( Figs. 42, 43 )

26. Second antennal segment red or reddish brown at apex; right paramere elongate ( Fig. 81 ); distribution: Brazil, Suriname, Trinidad, and Venezuela....................................................... C. intermedius Reuter ( Fig. 34 )

– Second antennal segment uniformly pale yellowish white; right paramere broadly rounded, with apex somewhat truncate ( Fig. 72 ); distribution: Minas Gerais, Brazil........................................ C. fiuzai Henry , n. sp. ( Figs. 25, 26 )

Figures

FIGURES 2–7. Scanning electron micrographs of Cyrtocapsus lazelli. 2, head and pronotum, lateral aspect. 3, head and pronotum, dorsal aspect. 4, head, frontal aspect. 5, ostiolar evaporative area and auricle. 6, parameres, lateral aspect. 7, pretarsal claws.

FIGURES 8–17. Habitus images of Cyrtocapsus spp. 8, 9,Cyrtocapsus apicalis, holotype male.8, dorsal aspect. 9, lateral aspect. 10–12, C. atrametatibialis. 10, male, dorsal aspect. 11, male, lateral aspect. 12, female, dorsal aspect. 13, 14, C. caligineus. 13, male, dorsal aspect. 14, female, dorsal aspect. 15, 16, C. columbiensis. 15, male, dorsal aspect. 16, female, dorsal aspect. 17, C. costatus, holotype male, dorsal aspect.

FIGURES 58–70. Male genitalia of Cyrtocapsus spp. 58–60, C. apicalis. 58, left paramere. 59, endosoma. 60, right paramere. 61, 62, C. atrametatibialis. 61, left paramere. 62, right paramere. 63, 64, C. caligineus. 63, left paramere. 64, right paramere. 65, 66, C. columbiensis. 65, left paramere. 66, right paramere. 67, 68, C. elutipes. 67, left paramere. 68, right paramere. 69, 70, C. femoralis. 69, left paramere. 70, right paramere.

Figures 71– 84. Male genitalia of Cyrtocapsus spp. 71, 72, C. fiuzai. 71, left paramere. 72, right paramere. 73, 74, C. glabratus. 73, left paramere. 74, right paramere. 75, 76, C. grenadensis. 75, left paramere. 76, right paramere. 77–79, C. haitianus. 77, left paramere. 78, endosoma. 79, right paramere. 80, 81, C. intermedius. 80, left paramere. 81, right paramere. 82–84, C. lazelli. 82, left paramere. 83, endosoma. 84, right paramere.

FIGURES 85–98. Male genitalia of Cyrtocapsus spp. 85, 86, C. mesoamericanus. 85, left paramere. 86, right paramere. 87, 88, C. metafemoratus. 87, left paramere. 88, right paramere. 89, 90, C. pallipes. 89, left paramere. 90, right paramere. 91, 92, C. paraguaiensis. 91 left paramere. 92, right paramere. 93, 94, C. rostratus. 93, left paramere. 94, right paramere. 95, 96, C. schaffneri. 95, left paramere. 96 right paramere. 97, 98, C. spatulatus. 97, left paramere. 98, right paramere.

FIGURES 99–102. Female genitalia Cyrtocapsus spp. 99, 100, C. lazelli. 99, Ventral aspect of ovipositor and labiate plate. 100, Dorsal aspect of ovipositor, posterior wall, and sclerotized rings. 101, 102, C. pallipes. 101, Ventral aspect of ovipositor and labiate plate. 102, Dorsal aspect of ovipositor, posterior wall, and sclerotized rings.

FIGURES 40–48. Habitus images of Cyrtocapsus spp. 40, 41, C. metafemoratus. 40, male, dorsal aspect. 41, holotype female, dorsal aspect. 42, 43, C. nordestinus. 42, male, dorsal aspect. 43, male, lateral aspect. 44, 45, C. pallipes. 44, male, dorsal aspect. 45, female, dorsal aspect. 46, 47, C. paraensis, holotype female (of C. bolivianus). 46, dorsal aspect. 47, lateral aspect. 48, C. paraguaiensis, holotype female, dorsal aspect.

FIGURES 49–57. Habitus images of Cyrtocapsus spp. 49–51, C. rostratus. 49, male dorsal aspect. 50, female lectotype, dorsal aspect. 51, female lectotype, lateral aspect. 52–54, C. schaffneri. 52, male, dorsal aspect. 53, female, dorsal aspect. 54, C. female, lateral aspect. 55, C. spatulatus, holotype male, dorsal aspect. 56, 57. C. suturalis. 56, male, dorsal aspect. 57, male, lateral aspect.

FIGURES 18–28. Habitus images of Cyrtocapsus spp. 18, 19, C. discalis, holotype male. 18, dorsal aspect. 19, lateral aspect. 20–22, C. elutipes. 20, male, dorsal aspect. 21, female, dorsal aspect. 22, female, lateral aspect. 23, 24, C. femoralis, lectotype male. 23, dorsal aspect. 24, lateral aspect. 25, 26, C. fiuzai. 25, male, dorsal aspect. 26, female, dorsal aspect. 27, 28, C. glabratus, holotype male. 27, dorsal aspect. 28, lateral aspect.

FIGURES 29–39. Habitus images of Cyrtocapsus spp. 29, 30, C. grenadensis. 29, male, dorsal aspect. 30, female, dorsal aspect. 31, C. guianus, holotype female, dorsal aspect. 32, 33, C. haitianus. 32, male dorsal aspect. 33, female, dorsal aspect. 34, C. intermedius, female, dorsal aspect. 35, 36, C. lazelli. 35, male, dorsal aspect. 36, female, dorsal aspect. 37, 38, C. marginatus. 37, lectotype male, dorsal aspect. 38, paralectotype female, dorsal aspect. 39, C. mesoamericanus, holotype male, dorsal aspect.