Species of Tytthus are characterized by the small size (lengths ranging from 1.08 mm in brachypterous males of Tytthus wheeleri to more than 3.60 mm in Tytthus mundulus), the relatively broad to nearly round head, usually with a pale yellow spot on the vertex bordering the inner margin of each eye; slightly protruding eyes not touching the anterior margin of the pronotum; smooth, shiny, trapeziform to campanulate pronotum, with lateral margins straight to weakly concave and moderately to strongly flared humeral angles; flat to weakly raised calli; subparallel hemelytra, often brachypterous or abbreviated, with the membrane and cuneus greatly reduced; slender claws with setiform parempodia; slender, tapered abdomen; small genital capsule; simple, C- to weakly S-shaped endosoma, lacking a secondary gonopore; mitt-shaped left paramere; and simple, round to elongate-oval right paramere.

Elongate subparallel to elongate oval species. Head shiny, impunctate, broader than long, sometimes becoming broadly rounded, especially in males, always slightly wider than anterior margin of pronotum; eyes prominent, more so in males, finely granulate, usually with scattered, fine, short setae; in dorsal view, frons and clypeus weakly rounded to prominent and pointed anteriorly; interocular space proportionately narrower in males (because of more prominent eyes) than females, nearly always with a small to large yellow or pale spot adjacent to inner margin of each eye; posterior margin nearly straight, with eyes nearly touching anterior margin of pronotum, to sometimes more narrowed behind eyes forming a necklike area more distinctly separating eyes from pronotum. Labium extending from bases of hind coxae to well onto abdomen near segment III or IV; segment I extending from base of head to bases of forecoxae. Antennal segment I shortest, stoutest; segment II longest; segment III longer than to subequal to segment IV. Pronotum shiny, impunctate, calli usually prominent, often with a glaucous sheen; subrectangular to trapeziform, especially in flightless brachypters, to strongly campanulate or bell-shaped in macropters. Mesoscutum broadly exposed in macropters; concealed by posterior margin of pronotum in brachypters. Scutellum well developed, equilateral. Hemelytra translucent, opaque white to bicolored with dark clouds, transverse bands, or extensive dark areas; macropterous or brachypterous, if only one sex brachypterous, always the female; fully macropterous hemelytra with each cuneus entire and membrane fully developed, extending well beyond apex of abdomen; brachyterous hemelytra (see discussion on brachyptery) abbreviated, ranging from a partially shortened membrane, extending only to apex of abdomen, to a strongly abbreviated membrane represented by only a remnant fringe on posterior edge of coleopteriform corium and clavus, with cuneus absent; in most extreme forms, only short hemelytral pads present, entirely lacking the cuneus and membrane, and extending only to abdominal terga III or IV. Lengths range in macropterous males from 2.14–3.42 mm; brachypterous males 1.08–1.28 mm; macropterous females 1.80–3.52 mm; and brachypterous females 1.44–1.68 mm. Ventral surface shiny, impunctate. Ostiolar evaporative area with a prominent auricle, curving posteriorly, gland opening large and distinct. Legs slender; femora unspotted, sometimes infuscated; tibiae slender, with or without distinct spines; tarsi slender, lengths of segment II and III subequal; claws elongate, slender, parempodia setiform.

Male genitalia: Endosoma relatively simple C-shaped to S-shaped, composed of a single, simple tube, often distally truncate or concave, lacking an apparent secondary gonopore. Left paramere mitt-shaped, with two arms and a narrow basal stem; right arm longest, widest, and most prominent, distally acute to rounded, gradually narrowing from base to apex, often broadened just before apex; left arm much shorter, distally acute. Right paramere elongate oval to nearly round, with a short basal stem. Phallotheca simple, sheathlike, exposed apex gradually narrowing from base to an acute apex.

Members of this genus are so superficially similar to species of the orthotyline genus Cyrtorhinus that Reuter (1875c) placed Tytthus, in synonymy under it, where it remained for the next 80 years. Even H. H. Knight (1923, 1925, 1931), North America’s most knowledgeable and prolific mirid specialist, failed to recognize the misplacement, and R. L. Usinger (1939), who treated several South Pacific species of Tytthus noted “An apparent structural anomaly in Cyrtorhinus which has not been given sufficient attention is the absence, in certain species, of arolia between the claws. The presence or absence and form of the arolia is usually a very reliable guide to relationships in Miridae.” Despite the character differences between these taxa, the species remained together under Cyrtorhinus until Carvalho and Southwood (1955) documented the obvious differences in male genitalia and pretarsal structure.

Another problematic genus, Isoproba Osborn and Drake (1915), has not been mentioned in the primary literature since its original description. Described to accommodate the only included species, Isoproba picea Osborn and Drake from Guatemala, it was said to be “readily separated from the [orthotyline] genus Paraproba Distant and allied genera by the more globose head and the peculiar shape of the thorax (Osborn and Drake 1915).” Carvalho (1952, 1958), however, without explanation, transferred it to the tribe Dicyphini (then placed it in the subfamily Phylinae), whose members also have generally rounded heads, as well as setiform parempodia. Cassis (1984) noted that he was unable to locate the holotype and, therefore, left it in Dicyphini with “considerable reservation.”

I have studied the holotype of Isoproba picea deposited in the Ohio State University collection and, like most species included in the genus Tytthus, it has an overall shiny, fuscous to black head, pronotum, and scutellum, pale translucent hemelytra, and slender legs and antennae. The male genitalia are of the same type as for other species of Tytthus. The left paramere is mitt-shaped, the right paramere is relatively small, elongate oval, and simple, and the endosoma is slender and C-shaped. Isoproba picea differs from other species of Tytthus onlyin having a more distinctly rounded or bulbous head that is narrowed posteriorly into a short neck, especially in males, and the shallowly convex eyes hardly protruding from the side of the head. In addition, I have discovered that Tytthus hondurensis Carvalho (1984) is a junior synonym of Tytthus piceus. As a consequence, Isoproba is placed as a junior synonym of Tytthus.

Wing polymorphism: Slater (1975) separated the various types of wing modifications in the family Lygaeidae (sensu lato) into seven main categories: 1) Aptery (wings entirely absent); 2) Microptery (wings reduced to widely separated pads; 3), Staphylinoidy (wings have the clavus and corium indistinguishably fused into a coriaceous pad, and the wings meet evenly along the midline for their entire length, and usually cover only the first three abdominal segments); 4, Coleoptery (wings may or may not be reduced, but the coriaceous portion is not reduced but lengthened, the clavus and corium are fused, and the wings meet along the midline but do not overlap); 5) Brachyptery (clavus and corium either distinctly separate or fused, but shorter than in macropters, with only the inner portion of the membrane overlapping; 6) Submacroptery (clavus and corium always separate, with membrane slightly shortened, leaving the last abdominal segment exposed); and 7) Macroptery (wings unmodified, fully developed). Of the species of Tytthus exhibiting wing polymorphism, two can be categorized as staphylinoid (Tytthus alboornatus, Tytthus wheeleri), two as brachypterous (Tytthus montanus, Tytthus piceus), and four (Tytthus balli, Tytthus fuscicornis, Tytthus pubescens, and Tytthus uniformis) as submacropterous. The remaining sixteen species are known only from macropterous individuals.

Importance in biological control: It has been documented that most, if not all, species of Tytthus are specialized delphacid and, to a lesser extent, leafhopper egg predators. The best documented species, Tytthus mundulus, provides a good example of successful classical biological control (Hagen and Franz 1973, van den Bosch and Messenger 1973, Rosen 1985, Wheeler 2001). Frederick Muir (1920) discovered while searching for predators of the sugarcane delphacid in Queensland, Australia, that nymphs and adults fed on delphacid eggs. As a consequence, he brought Tytthus mundulus to Hawaii for release into the sugarcane fields. As Usinger (1939) noted, “Muir’s discovery that Tytthus (as Cyrtorhinus) mundulus (Breddin) lives exclusively on the eggs of the sugar-cane leafhopper, Perkinsiella saccaricida Kirkaldy, led to one of the most outstanding successes in the field of biological control of injurious insects.” Zimmerman (1948) summed up the importance of this bug by saying “This one bug has saved the Hawaiian sugar industry and the Territory millions of dollars—its true worth can hardly be estimated.”

Other species also have shown considerable potential in biological control. In South Africa, both Tytthus mundulus and Tytthus parviceps (Reuter) have been investigated for control of a tropiduchid, Numicia viridis Muir, on sugarcane (Carnegie and Harris 1969). Although Tytthus mundulus was the better-known predator, Tytthus parviceps was more easily reared and showed the greatest potential for controlling Numicia viridis. Jhansi et al. (2002) studied the biology and prey preferences of Tytthus parviceps on planthoppers and leafhoppers on rice in India, including the brown planthopper, Nilaparvata lugens (Stål). The Holarctic species Tytthus pubescens (Knight) and Tytthus pygmaeus (Zetterstedt) are known to prey on leafhoppers and delphacids in England (Southwood and Leston 1959, Rothschild 1963, Wheeler and Henry 1992). In coastal eastern North America, Döbel and Denno (1994) considered Tytthus alboornatus (Knight) and Tytthus vagus (Knight) among the major predators of saltmarsh delphacids on two species of Spartina (Poaceae). For additional information on the hosts and habits of these predatory bugs, see the respective species within this revision.

This species is distinguished by the small size, usually brachypterous hemelytra, overall dark brown coloration, with the basal third to half of the corium and clavus, and cuneus (or in brachypters the posterior margin of the corium) pale or white, the pale yellowish brown antennal segment I, the mostly dark brown femora and pale yellowish-brown tibiae and tarsi. Macropterous and brachypterous forms are known for both sexes.

This species is most similar in size and coloration to Tytthus wheeleri, sp. n. In Tytthus alboornatus, antennal segment I is pale yellowish brown and the posterior margin of each hemelytron in brachypters or the cuneus in macropters is pale or white, whereas in Tytthus wheeleri, antennal segment I is dark brown and the posterior margin of each hemelytron in brachypters and or cuneus in macropters is uniformly dark brown.

Macropterous male (n = 1, plus holotype in parentheses) (Fig. 2): Length to apex of hemelytron 2.21 mm (2.24 mm), length to base of cuneus 1.57 mm (1.66 mm), width across hemelytra 0.72 mm (0.77 mm). Head: Length 0.72 mm (0.26 mm), width across eyes 0.51 mm (0.54 mm), interocular width 0.30 mm (0.30 mm). Labium: Length [embedded in glue] (0.94 mm). Antenna: Segment I length 0.29 mm (0.37 mm), II 0.88 mm (1.06 mm), III 0.59 mm (0.72 mm), IV 0.48 mm (missing). Pronotum: Length 0.32 mm (0.35 mm), basal width 0.45 (0.72 mm).

Brachypterous male (n = 5): Length to apex of abdomen 1.34–1.57 mm, length to base of hemelytron (cuneus and membrane absent) 1.15–1.41 mm, width across hemelytra 0.58–0.62 mm. Head: Length 0.22–0.26 mm, width across eyes 0.50–0.59 mm, interocular width 0.29–0.30 mm. Labium: Length 0.83–0.86 mm. Antenna: Segment I length 0.29–0.30 mm, II 0.80–0.86 mm, III 0.53–0.64 mm, IV 0.45–0.61 mm. Pronotum: Length 0.24–0.29 mm, basal width 0.54–0.56 mm.

Coloration: Head: Brown to dark brown, with a small, vague, pale spot near inner margin of each eye; eyes dark brown to reddish brown. Labium: Yellowish brown, apex of segment IV darker brown. Antenna: Segment I pale yellowish brown; segments II–IV yellowish brown, sometimes becoming slightly darker brown. Pronotum: Uniformly shiny brown to very dark brown or fuscous. Mesoscutum: Hidden under base of pronotum in brachypters, narrowly exposed in macropters. Scutellum: Brown to dark brown, with apex pale. Hemelytron: Broadly dark brown, with basal one fourth and narrow apex or cuneus (in macropter) pale or white; membrane on only macropter fully developed, smokey brown. Ventral surface: Thoracic pleural areas brown to dark brown, ventral surface sometime paler yellowish brown; abdomen dark brown to fuscous, especially laterally, ventral area sometimes paler yellowish brown. Ostiolar evaporative area: Dark brown. Legs: Coxae pale yellowish brown to whitish, with bases dark brown; femora dark brown, pale yellowish brown at bases and apices; tibiae, tarsi, and claws pale yellowish brown.

Structure, texture, and vestiture: Head: Shiny, impunctate; broader than long, rounded anteriorly, truncate basally; set with short, recumbent, nearly bristlelike setae on vertex and frons. Labium: Extending beyond metacoxae to second or third abdominal segment. Pronotum: Shiny, impunctate, nearly rectangular, wider than long, anterior angles rounded, base truncate, calli indistinct, not differentiated from discal surface, only slightly raised and rounded; set with scattered, recumbent, brown to nearly black setae. Scutellum: Equilateral, impunctate, with a few scattered, short, recumbent setae. Hemelytron: Macropter with fully developed cuneus and membrane, including two closed cells or areoles; all other specimens (except one macropter) brachypterous (staphylinoid), with clavus fused (and claval suture absent) with corium, cuneal fracture and cuneus absent, and membrane absent or rarely with only a remnant narrow strip along truncate posterior margin, extending from about abdominal tergite IV to nearly to apex of abdomen; set with evenly scattered, short, recumbent brown setae.

Male genitalia: Left paramere (Fig. 109): Mitt-shaped; right arm long, stout; left arm short, apically acute. Right paramere (Fig. 110): Oval. Endosoma (Fig. 111): Slender, S-shaped, apex pointed. Phallotheca (Fig. 112): Relatively slender, apically acute.

Macropterous female (n = 1) (Fig. 3): Length to apex of hemelytron 2.45 mm, length to base of cuneus 1.85 mm, width across hemelytra 0.93 mm. Head: Length 0.32 mm, width across eyes 0.58 mm, interocular width 0.32 mm. Labium: Length 0.99 mm. Antenna: Segment I length 0.32 mm, II 0.91 mm, III 0.69 mm, IV 0.67 mm. Pronotum: Length 0.37 mm, basal width 0.86 mm.

Brachypterous “minor” female [see discussion below] (n = 10): Length to apex of abdomen 1.44–1.79 mm, length to apex of hemelytra 1.34–1.60 mm, width across hemelytra 0.58–0.80 mm. Head: Length 0.26 mm, width across eyes 0.43–0.53 mm, interocular width 0.29–0.30 mm. Labium: Length 0.80–0.91 mm. Antenna: Segment I length 0.26–0.27 mm, II 0.74–0.80 mm, III 0.53–0.59 mm, IV 0.48–0.61 mm. Pronotum: Length 0.27–0.29 mm, basal width 0.50–0.56 mm.

Brachypterous “major” female (n = 2) (Fig. 4): Length to apex of abdomen 2.02–2.30 mm, length to apex of hemelyra 1.60–1.86 mm, width across hemelytra 0.93-0.99 mm. Head: Length 0.27–0.30 mm, width across eyes 0.56–0.61 mm, interocular width 0.34–0.35 mm. Labium: Length 0.91–0.96 mm. Antenna: Segment I 0.30–0.32 length mm, II 0.90–0.96 mm, III 0.64 mm, IV 0.50 mm. Pronotum: Length 0.29–0.34 mm, basal width 0.59–0.64 mm.

Taken by A. G. Wheeler on inland saltgrass, Distichlis spicata (L.) Green [Poeaceae]; saltmeadow cordgrass, Spartina patens (Ait.) Muhl.; and sand cordgrass, Spartina bakeri Merr. [Poeaceae]. Denno (in litt, 2005) informed me that Tytthus alboornatus was extremely abundant only in Spartina patens in Tuckerton, New Jersey, but not in Spartina alterniflora where Tytthus vagus was restricted. Denno (1983) and Hines et al. (2005) reported that Tytthus alboornatus preys on eggs of the delphacids Tumidagena minuta McDermott and possibly Delphacodes detecta (Van Duzee) in Spartina patens.

Previously known only from Florida, New Jersey, New York, and Virginia (Henry and Wheeler 1988, Hoffman 2000). New U. S. state records are Connecticut, Louisiana, South Carolina, and Texas. This distribution indicates that Tytthus alboornatus should occur in all coastal states from at least New England to Texas.

Tytthus alboornatus (Figs 2–4) is one of the smallest species in the genus, second in size only to the similar-appearing Tytthus wheeleri (Figs 50–52). Like Tytthus wheeleri, macropterous forms of this species are extremely uncommon or rare. Of the more than 50 specimens studied, I have found only two macropterous males, including the holotype (from Florida) and three macropterous females (from Connecticut and South Carolina).

In addition, populations include what I call “minor” and “major” females. Along the Connecticut and New Jersey coasts, specimens are much smaller (shorter and more slender), whereas farther south in South Carolina and Florida several specimens are considerably larger, with more well-developed pronota and hemelytra. Two macropterous females from Connecticut also were smaller than the macropter from South Carolina. This size difference may simply be due to more harsh or crowded conditions, with a more limited food supply versus smaller populations with more plentiful prey available, rather than their north/south distributions. More work is needed to better understand the factors that influence size.

Holotype ♂ (00162206) (USNM): UNITED STATES: Florida: Duval Co.: Jacksonville, 30.33194°N, 81.65583°W, 23 Jul 1926, E. D. Ball.

UNITED STATES: Connecticut: New London Co.: Stonington Township, Barn Island, 41.335°N, 71.906°W, 11 Aug 1971, R.T. and J. C. Schuh, 9 ♂♂ (00095349, 00165938 - 00165945), 11 ♀♀ (00095350, 00165946 - 00165955) (AMNH), 1 ♂ ( 00138679) (USNM) (all brachypterous, except two macropterous ♀♀). Tolland Co.: Mansfield Center, 41.76528°N, 72.19861°W, 84 m, 01 Aug 1956, J. A. Slater, 1 ♀ (00166063) (AMNH) (brachypterous). Florida: Brevard Co.: Titusville, 28.61194°N, 80.80778°W, 29 Apr 1952, collector unknown, 1 ♂ (00161887) (USNM) (macropterous). Duval Co.: CR-105, 1 km W. of Dunn Creek, S. of Eastport, 03 Apr 2004, A. G. Wheeler, Jr., ex Spartina patens (Poaceae), 1 ♀ (00161882) (USNM) (brachypterous). Miami-Dade Co.: Biscayne Bay, 25.5747°N, 80.3112°W, 1700, Mrs. A.T. Slosson, 2 ♂♂ (00165956, 00165957) (AMNH) (brachypterous). Louisiana: Tangipahoa Parish: Hammond, 22 June 1948, E. L. Todd, 1 ♂, 1 ♀ (AMNH) (macropterous). New Jersey: Ocean Co.: 2 mi. E. of Manahawkin off Stafford Ave., 20 Jul 1976, collector unknown, Distichlis spicata (Poaceae), 5 ♂♂ (00161870, 00161877 - 00161878, 00161880, 00161886), 2 ♀♀ (00161879, 00161881) (USNM) (all brachypterous). Tuckerton, 39.60305°N, 74.34027°W, 4 m, 01 Aug 2002, R.F. Denno & D. Lewis, Spartina patens (Poaceae), 7 ♂♂ (00161865 - 00161868, 00161872, 00161874 - 00161875), 4 ♀♀ (00161869, 00161871, 00161873, 00161876) (USNM) (all brachypterous). South Carolina: Colleton Co.: Bear Island Wildlife Management Area, near Mary’s House Pond, 03 May 2003, A. G. Wheeler, Jr., ex Spartina patens (Poaceae), 1 ♀ (00161883) (USNM (brachypterous). CR-26, 0.3 km N. of Ashepoo River, N. of Bennetts Pt., 03 May 2003, A. G. Wheeler, Jr., ex Spartina bakeri (Poaceae), 1 ♀ (00161885) (USNM) (brachypterous). CR-26, 1.1 km W. of Bennetts Point, 04 May 2003, A. G. Wheeler, Jr., Spartina bakeri (Poaceae), 1 ♂ (macropterous), 1 ♀ (00161889) (brachypterous) (USNM). Texas: Jefferson Co.: J.D. Murphree Wildlife Management Area, near Lost Lake, 29.78111°N, 93.97333°W, 26 May 1994, R. Vogtsberger, taken by mosquito dipper cup, 1 ♂ (00161888) (USNM) (brachypterous). Virginia: Mecklenburg Co.: Elm Hill WMA, Clyde’s Pond, 11-29 May 1995, VMNH survey, 4 ♂♂ (VMNH). City of Virginia Beach: Dam Neck Naval Base, dune DF site, 25 June 1991, K. A. Buhlmann, 1 ♂ (VMNH).

This species, distinguished by the pale antennae and legs, mostly pale dorsum, with only the frons and basal margin of the head, scutellum, inner margin of the clavus, and the distal third of the corium brown, cannot be easily confused with any other species of the genus. Only macropters are known.

Tytthus amazonicus keys to Tytthus zwaluwenbergi because of the pale tibiae, antennae, and head but its relationship with this central Pacific species almost certainly is only superficial. Tytthus amazonicus is readily separated by the pale head and pronotum invaded with dark brown, the largely pale hemelytra with the inner half of each clavus and the apical half of each corium dark brown, whereas Tytthus zwaluwenbergi is uniformly pale yellowish brown.

Male (n = 5) (Fig. 5): Length to apex of hemelytron 2.83–2.93 mm, length to base of cuneus 2.00–2.10 mm, width across hemelytra 0.86–0.93 mm. Head: Length 0.34–0.37 mm, width across eyes 0.62–0.64 mm, interocular width 0.29–0.30 mm. Labium: Length 1.22–1.28 mm. Antenna: Segment I length 0.37–0.38 mm, II 1.00–1.08 mm, III 0.56–0.66 mm, IV 0.48–0.53 mm. Pronotum: Length 0.35–0.38 mm, basal width 0.83–0.88 mm.

Coloration: Head: Pale yellowish brown dorsally, dark brown ventrally, on frons, and narrowly across basal margin; eyes dark brown to reddish brown. Labium: Pale yellowish brown. Antenna: Segments I–III uniformly pale yellow to yellowish brown, segment I sometimes darker brown through middle with apex and base pale, segment IV slightly darker brown. Pronotum: Mostly pale yellowish brown, collar and narrow posterior margin around calli darker brown, entire discal area darker brown on some specimens. Mesoscutum: Pale yellowish brown, tinged with darker brown through middle. Scutellum: Brown. Hemelytron: Predominantly pale or pale yellowish brown, with inner margin of clavus and apical half of corium darker brown; cuneus uniformly pale or pale yellowish brown; membrane translucent brown, veins darker brown. Ostiolar evaporative area: Dark brown. Ventral surface: Thoracic area dark brown; abdomen brown to yellowish brown, darker brown along lateral margins and genital capsule. Legs: Coxae pale yellow, meso- and metacoxae brown at bases; remainder of legs pale yellow.

Structure, texture, and vestiture: Head: Wider than long, impunctate, frons with a glaucus sheen; buccula relatively wide; set with short recumbent setae on frons and a few longer, more erect setae on vertex. Labium: Extending beyond metacoxae to base of abdomen; segment I extending to middle of procoxae. Pronotum: Impunctate, shiny; trapeziform, anterior angles rounded, lateral margins weakly concave, basal angles flared wider, basal margin concave; calli weakly swollen, delimited posteriorly by a shallow impressed line; set with evenly scattered, short, recumbent setae. Mesoscutum: Broadly exposed. Scutellum: Impunctate, equilateral, with scattered, short, recumbent setae. Hemelytron: Macropterous, impunctate, shiny lateral margins subparallel, cuneus longer than wide at base, membrane with two areoles, extending well beyond abdomen.

Male genitalia: Left paramere (Fig. 113): Mitt-shaped, with a long, broad right arm and shorter, more slender left arm. Right paramere (Fig. 114): Elongate oval. Endosoma (Fig. 115): S-shaped, with apex rounded. Phallotheca (Fig. 116): Relatively slender, apically acute.

Female (n = 4) (Fig. 6): Length to apex of hemelytron 2.98–3.14 mm, length to base of cuneus 2.18–2.30 mm, width across hemelytra 0.91–1.07 mm. Head: Length 0.34–0.35 mm, width across eyes 0.58–0.59 mm, interocular width 0.30-0.32 mm. Labium: Length 1.23–1.33 mm. Antenna: Segment I 0.29–0.30 length mm, II 0.85–0.91 mm, III 0.56–0.61 mm, IV mm. Pronotum: Length 0.42–0.43 mm, basal width 0.83–0.96 mm.

Unknown. Most specimens taken at light.

Described and previously known only from Amazonas, Brazil. Peru is a new country record.

BRAZIL: Amazonas: Reserva Ducke, 25 km NNE of Manaus, 120 m, 26 Jul 1973, R.T. Schuh, 2 ♂♂ (00165859, 00165860), 2 ♀♀ (00165857, 00165858) (AMNH), 1 ♂ (00162159), 1 ♀ (00162158) (USNM). On Amazon River, Jul 1900, Boquaert, 1 ♂ (00161884) (USNM). R. Madeira to St. Antonio, May 74, 3 ♀♀ (BNHM). Para: Santarem Co.: Taperinha, 11 Jun 1927 - 20 Jun 1927, Zerny, 1 ♂ (00161615) (USNM). PERU: Junin: Satipo, 11.2667°S, 74.6833°W, Jul 1940 - Aug 1940, P. Paprzycki, 1 ♀ (00161613) (USNM). Loreto: Lake Yarinacocha, 10 km NW of Pucallpa, 150 m, 10 Dec 1971, R. T. Schuh, 1 ♂ (00165855), 1 ♀ (00165856) (USNM).

This species is distinguished by the black head and antennae, orange-brown pronotum often infuscated laterally, orange-brown scutellum, pallid hemelytra infuscated along inner margin of clavus and distal third of corium, orange-brown femora, and dark brown to fuscous tibiae. All known males are macropterous; both macropterous and brachypterous females occur.

Tytthus balli is very similar to Tytthus insperatus and Tytthus uniformis based on the black head and antennal segments I and II, and the orange to brownish-orange pronotum and femora. From Tytthus insperatus, it is distinguished by the uniformly browish-orange pronotum lacking a pale anterior margin and the infuscated apical half of each corium. From Tytthus uniformis it is distinguished by the fuscous tibiae and infuscated hemelytra.

Macropterous male (n = 5, plus holotype in parentheses) (Fig. 7): Length to apex of hemelytron 2.58–2.73 mm (2.68 mm), length to base of cuneus 1.85–1.88 mm (1.96 mm), width across hemelytra 0.70–0.83 mm (0.80 mm). Head: Length 0.34–0.35 mm (0.37 mm), width across eye 0.58–0.59 mm (0.59 mm), interocular width 0.30–0.32 (0.30 mm). Labium: Length 1.05–1.07 mm (1.02 mm). Antenna: Segment I length 0.30–0.32 mm (0.29 mm), II 0.98–1.07 mm (1.02 mm), III 0.69–0.85 mm (0.80 mm), IV 0.48–0.51 mm (missing). Pronotum: Length 0.30-0.32 mm (0.34 mm), basal width 0.72–0.74 mm (0.69 mm).

Coloration: Head: Uniformly shiny black, with a distinct, yellow spot on inner interocular area bordering each eye. Labium: Yellowish brown, apex of segment IV fuscous. Antenna: Uniformly black. Pronotum: Uniformly orange to darker brownish orange, often becoming infuscated laterally and around calli. Mesoscutum: Orange to brownish orange. Scutellum: Dark orange or brownish orange to dark brown. Hemelytron: Pale or whitish on basal half and narrow apical margin of corium, narrow outer margin of clavus, and basal half of cuneus, dark brown on most of clavus, apical half of corium, and apex of cuneus; membrane translucent brown, veins slightly dark brown. Ostiolar evaporative area: Brownish orange to fuscous. Ventral surface: Thoracic area orange to dark brownish orange; abdomen brownish orange to dark brown, often becoming fuscous laterally, genital capsule dark brown to fuscous. Legs: Coxae pale or whitish, orange to dark brownish orange basally; femora orange to brownish orange; tibiae brown on palest specimens, especially protibia, to dark brown or black on darker individuals; tarsi and claws brown to dark brown.

Structure, texture, and vestiture: Head: Broader than long, shiny, impunctate; sparsely set with short, recumbent setae; buccula relative wide, tapering posteriorly. Labium: Extending to apices of metacoxae or base of abdomen; segment I extending only to prosternum. Pronotum: Impunctate, shiny; trapeziform, anterior angles rounded, lateral margins weakly concave, basal angles flared wider, basal margin concave; calli weakly swollen, delimited posteriorly by a shallow impressed line; set with evenly scattered, short, recumbent setae. Mesoscutum: Broadly exposed. Scutellum: Impunctate, equilateral, width a few scattered, recumbent setae. Hemelytron: Macropterous, impunctate, shiny, lateral margins subparallel, cuneus longer than wide at base, membrane with two areoles, extending well beyond abdomen.

Male genitalia: Left paramere (Fig. 117): Mitt-shaped; right arm long, broad; left arm shorter, more slender. Right paramere (Fig. 118): Oval. Endosoma (Fig. 119): C-shaped. Phallotheca (Fig. 120): Slender, relatively straight, apically acute.

Macropterous female (n = 2): Length to apex of hemelytron 2.90–3.07 mm, length to base of cuneus 2.08–2.28 mm, width across hemelytra 0.83–0.93 mm. Head: Length 0.34–0.35 mm, width across eyes 0.58–0.61 mm, interocular width 0.32–0.34 mm. Labium: Length 1.17–1.22 mm. Antenna: Segment I length 0.27–0.32 mm, II 1.02–1.1.18 mm, III 0.77–0.80 mm, IV 0.48–0.54 mm. Pronotum: Length 0.32–0.34 mm, basal width 0.74–0.78 mm. Similar to males in color and shape.

Brachypterous female (n = 4) (Fig. 8): Length to apex of abdomen 2.20–2.38 mm, length to base of cuneus 1.88–1.93 mm, width across hemelytra 0.86–0.90 mm. Head: Length 0.37–0.40 mm, width across eyes 0.58–0.59 mm, interocular width 0.34–0.35 mm. Labium: Length 1.17-1.20 mm. Antenna: Segment I length 0.27–0.29 mm, II 1.01–1.02 mm, III 0.77–0.80 mm, IV 0.48–0.50 mm. Pronotum: Length 0.30–0.32 mm, basal width 0.67–0.69 mm. Similar to males in color, differing in having the cuneus reduced (basal width subequal to length) and the membrane shortened (with the veins absent or indistinct), extending only to the middle of the last abdominal tergite.

Previously recorded only from imported roses (Carvalho and Southwood 1955), which undoubtedly is an incidental record. A. G. Wheeler and I have swept specimens from a salt marsh area containing mixed herbaceous vegetation and Spartina sp. [Poaceae].

This species was described from Jacksonville, Florida (Knight 1931), and later reported from an unspecified Mexican locality based on a specimen intercepted at Brownsville, Texas (Carvalho and Southwood 1955). The specimens listed below from Clay, Hidalgo, and Refugio counties represent the first authentic Texas records.

Holotype ♂ (00162202) (USNM) (macropterous): UNITED STATES: Duval Co.: Jacksonville, 30.33194°N, 81.65583°W, 23 Jul 1926, E. D. Ball. Paratypes: UNITED STATES: Texas: Duval Co.: Jacksonville, 30.33194°N, 81.65583°W, 23 Jul 1926, E. D. Ball, 1 ♂ (00161903) (USNM), 2 ♀♀ (00161912, 00161913), 1 nymph (00161910), 2 ♀♀ (00161914, 00161915) (USNM).Texas: Presidio Co.: Presidio, 29.56056°N, 104.37167°W, 28 Sep 1929, W.L. Owens, 1 ♂ (00167070) (CNC) (macropterous), 1 ♀ (00161911) (USNM) (brachypterous).

UNITED STATES: Florida: Texas: Cameron Co.: Brownsville, 25.90139°N, 97.49722°W, 28 Mar 1945, collector unknown, 1 ♀ (00161917) (USNM) (macropterous). Brownsville, Veteran’s Bridge (Cargo Lot), 27 Jul 2011, S. Guzman, 1 ♂ (USNM). Chambers Co.: Anahuac, 29.77278°N, 94.6825°W, 08 Oct 1918, H. S. Barber, 1 ♂ (00161904) (USNM). Clay Co.: 6 mi NE of Bexar at Red River (at state line), 34.13611°N, 98.13055°W, 20 Sep 2009, G. F. and J. F. Hevel, 1 ♂ (00161909) (USNM) (macropterous). Hidalgo Co.: Hidalgo County, 26.1°N, 98.26278°W, 19 May 1930, J. C. Gaines, Paratype, 1 ♀ (00161916) (USNM). Refugio Co.: 6 mi. N. Bayside, Rt. 136, 20 Apr 1983, T. J. Henry and A. G. Wheeler, Jr, 4 ♂♂ (00161905 - 00161908) (USNM) (macropterous).

This species is very similar to Tytthus parviceps in general size and in sharing the dark base of each tibia, dark antennae with the apex and base of segment I pale, and short, erect, brushlike setae on antennal segment II. Tytthus chinensis almost always has a uniformly black pronotum (Figs 9, 10) or the pronotum with only weak indications of yellow around the calli, and the endosoma (Fig. 123) is C-shaped, whereas Tytthus parviceps has the anterior one third to half of the pronotum around the calli almost always extensively pale yellow (Figs 32, 34) and the endosoma appears more distinctly S-shaped (Fig. 170). All known specimens of both species are macropterous.

Three other exclusively New World species, Tytthus entrerianus, Tytthus femoralis, and Tytthus mexicanus, also have fuscous “knees, ” a character that distinguishes them from all other species of the genus, except. Tytthus chinensis and Tytthus parviceps as noted above and in the key. All three, however, lack the brushlike setae on antennal segment II and the apical one third to two thirds of the hind femora of these species are infuscated.

Male (n = 15) (Fig. 9): Length to apex of hemelytron 2.18–2.60 mm, length to base of cuneus 1.65–1.88 mm, width across hemelytra 0.79–0.93 mm. Head: Length 0.27–0.29 mm, width across eyes 0.56–0.58 mm, interocular width 0.29–0.30 mm. Labium: Length 0.72–0.94 mm. Antenna: Segment I length 0.24–0.26 mm, II 0.78–0.82 mm, III 0.43–0.51 mm, IV 0.30–0.32 mm. Pronotum: Length 0.29–0.32 mm, basal width 0.69–0.80 mm.

Coloration: Uniformly fuscous to black, with a large, yellow, interocular spot touching inner margin of each eye, spots nearly contiguous in some individuals; eyes fuscous to dark reddish brown. Labium: Uniformly pale yellow, except of brown apical half of segment IV. Antenna: Segment I black, with base and apex narrowly pale yellow; segments II–IV uniformly fuscous to black. Pronotum: Usually uniformly black, anterior third around calli frequently weakly tinged with yellow, and less often entire anterior third yellow; posterior half uniformly fuscous to black. Mesoscutum: Uniformly yellowish brown to fuscous. Scutellum: Uniformly fuscous to black. Hemelytron: Uniformly translucent yellow to whitish. Ostiolar evaporative area: Yellowish, with central area of auricle invaded with fuscous, sometimes entirely fuscous. Ventral surface: Anterior half of proacetabula often yellow, propleura, pro- and mesosterna, and metapleura fuscous; abdomen varying from largely yellowish, with only genital capsule fuscous or black to largely fuscous with only ventral area pale. Legs: Uniformly generally yellow; procoxae uniformly yellow, meso-and metacoxae dark brown at bases, yellow beyond; femora yellow, often tinged with orange; tibiae yellow, bases broadly fuscous; tarsi and claws yellowish.

Structure, texture, and vestiture: Head: Weakly shiny, impunctate; buccula slender, extending posteriorly, ending near level with hind margin of eye; thickly set with short to relatively long semierect setae, especially on frons. Labium: Extending to apices of meso- or bases of metacoxae; segment I extending beyond base of head to xyphyus just before procoxae. Antenna: Segment I set with short, recumbent setae and two, long, subapical, bristlelike setae; segment II thickly set with short, recumbent setae, intermixed with row of longer, erect setae (Fig. 59) along ventral surface. Pronotum: Anterior angles rounded; lateral margins weakly concave, gradually widening to rounded posterior angles; posterior margin weakly sinuate. Mesoscutum: Weakly shiny, impunctate; set with a few scattered, semierect setae. Scutellum: Weakly shiny, impunctate; equilateral; set with a few scattered recumbent and semierect setae. Hemelytron: Macropterous, cuneus and membrane fully developed, extending posteriorly well beyond apex of abdomen; evenly set with relatively long, recumbent setae.

Male genitalia: Left paramere (Fig. 121): Mitt-shaped; right arm long, brown; left arm short, spinelike. Right paramere (Fig. 122): Oval. Endosoma (Fig. 123): C-shaped, apex blunt. Phallotheca (Fig. 124): Relatively slender, strongly bent (C-shaped), apically acute.

Female (n = 15) (Fig. 10): Length to apex of hemelytron 2.50–2.78 mm, length to base of cuneus 1.86–2.00 mm, width across hemelytra 0.90–0.96 mm. Head: Length 0.29–0.30 mm, width across eyes 0.60–0.62 mm, interocular width 0.30–0.32 mm. Labium: Length 0.90–0.94 mm. Antenna: Segment I length 0.26–0.27 mm, II 0.45–0.46 mm, III 0.26–0.27 mm, IV 0.29–0.30 mm. Pronotum: Length 0.29–0.30 mm, basal width 0.80–0.85 mm.

Usinger (1939) reported Tytthus chinensis (as Cyrtorhinus riveti) feeding on the eggs of Sogata ochrias (Kirkaldy) [Delphacidae] on Sporobolus virginicus (L.) Kunth [Poaceae] and Nilaparvata lugens (Stål) [Delphacidae] on rice (Oryza sp.) in Guam. He also recorded this species from Bermudagrass, Cynodon dactylon (L.) Pers., in Samoa, and Tradescantia sp. [Commelinaceae] in Tahiti.

This species has been reported from central and southeastern China, Japan, Taiwan, and Australia, India, and the Indo-Pacific Region (Caroline Islands, Childers Island, Cook Islands, Fiji, Gilbert Island, Indonesia, Mariana Islands, Marshall Islands, New Caledonia, New Hebrides, Papua New Guinea, Philippine Islands, Pitcairn Island, Rapa Island, Ryuku Islands, Samoa, Society Islands, Solomon Islands, South Korea, Swain Islands, and Tonga Islands) (Schuh 1984, 1995; Kerzhner and Josifov 1995; Cassis and Gross 1995). Based on specimens in the USNM, most or all records reported by Schuh (1984) from India and Sri Lanka should be applied to Tytthus parviceps; Schuh’s (1984) record from Hawaii is based on two specimens intercepted on international commerce at Honolulu (without origin indicated).

Based on specimens examined, I now also have records of this species from Cambodia, Guadalcanal, Guam, Saipan, and Tinian Island.

Carvalho and Southwood (1955), after consulting with D. R. Malaise at the Swedish Museum of Natural History in Stockholm, reported that the type of Capsus chinensis Stål (1859) must be considered lost. Based on Stål’s original description and study of other types, they concluded that Cyrtorhinus elongatus Poppius and Capsus annulicornis Poppius (both in Deutsches Entomologisches Institut) and Capsus riveti Cheesman (BMNH) are junior synonyms. They characterized Tytthus chinensis as “the smallest species of Tytthus and is distinguished by its black pronotum and scutellum, the dark bases of the tibia and the small size.” I have studied the type of Tytthus riveti and agree that it follows the concept of Tytthus chinensis outlined by Carvalho and Southwood (1955) and Carvalho (1956), including a uniformly dark pronotum.

In addition, Kerzhner and Josifov (1995) suggested that Tytthus koreanus Josifov and Kerzhner (1972) could be a junior synonym of Tytthus chinensis. Although I have not examined type material of Tytthus koreanus, the description and figures presented in the original description indicate that this species is based on pale specimens having the calli or anterior third of the pronotum yellow to brownish yellow. Josifov and Kerzhner (1972) did not indicate if they observed specimens with entirely dark pronota. I also have studied a male and two females from South Korea, lent by Dr. Seunghwan Lee (SNU), that agree with the pale color form of Tytthus chinensis having only the calli and mesoscutum tinged with dull brownish yellow. As a consequence, I consider T. koreanus and the material from Dr. Lee conspecific with that of Tytthus chinensis.

Although most material of what I consider to be Tytthus chinensis has a uniformly fuscous to black pronotum, varying degrees of yellow are present on the calli and anterior third of the pronotum on some specimens from eastern Asia, throughout the Indo-Pacific, and Australia, which may cause confusion with Tytthus parviceps that also has yellow on the anterior area of the pronotum. Nevertheless, I consider all material from eastern Asia (including Korea and Japan), Indonesia, Malaysia, Australia, New Guinea, and the South Pacific Islands Tytthus chinensis based on the dark pronotum in most specimens and C-shaped endosoma. However, because of the strong morphological similarity between T. “chinensis” and T. “parviceps, ” an effort will be made to accumulate fresh material worldwide for conducting a phylogeographic analysis of species limits within this complex using mitochondrial cytochrome oxidase 1 (COI) sequence data in cooperation with colleagues S. Scheffer and M. Lewis (Systematic Entomology Laboratory, ARS, USDA, Beltsville, Maryland, USA).

For the time being, I am restricting the distribution of Tytthus parviceps that has the anterior third of the pronotum broadly yellow (but certain specimens within populations can have a uniformly dark pronotum) to southern Asia (India, Srilanka, Pakistan, Vietnam, Thailand), the Middle East, Afrotropical and Neotropical regions, and subtropical United States. A few specimens from the Indo-Pacific (e.g., Guam), Australia, Cambodia, and other eastern Asian countries are externally indistinguishable from typical specimens of Tytthus parviceps from Africa and the New World. As noted above, the endosoma of typical Tytthus chinensis is C-shaped, whereas this very simple structure in African and New World specimens of Tytthus parviceps is usually S-shaped. Until additional collections can be made and additional DNA sequencing can be conducted on these populations, the strongly yellow phenotypes within populations of Tytthus chinensis must regarded convergent color forms.

To ensure nomenclatural stability, I designate the following female as the lectotype of Cyrtorhinus riveti: Label 1 (circular label with orange ring) “Type”; label 2, “Tahiti: Nr. Papeete[, ] Mar.-Apr.[, ] Miss Cheesman”; label 3 (handwritten), “ Cyrtorhinus riveti Cheesman”; label 4, “St. George Exp.[, ] BM 1925- 235”; label 5 (00085518 (BMNH).”

AUSTRALIA: New South Wales: Deewhy Beach dunes, 28 Jan 1957, W. W. Wirth, 1 ♂ (00162347), 1 ♀ (00167399) (USNM). Narrabri, 25 Jan 1960, M. Nikitin, by light, 1 ♀ (BMNH). Northern Territory: Katherina River, 14.5°S, 132.25°E, 12 Nov 1979, Zaytsev, 1 ♂ (00235059) (ZISP). Katherine, 14.467°S, 132.267°E, 17 Nov 1990, W. F. Chamberlain, Light Trap, 1 ♂ (00167391) (USNM). Queensland: Cape York Islands Co.: Banaga, N. Cape York, gum forest, Jan 1958, P.F. Darlington, 1 ♂ (00409566) (KU). Gracemere, 05 Nov 1990, W. F. Chamberlain, Light Trap, 1 ♂ (00167392), 1 ♀ (00167398) (USNM). Mossman, 16.47°S, 145.37172°E, 36 m, 11 Nov 1990, W. F. Chamberlain, 1♀ (00167400), Light Trap, 2 ♂♂ (00167389, 00167393) (USNM). Mt. Berryman Rd., Laidley, 05 Dec 1990, W. F. Chamberlain, Light Trap, 2 ♂♂ (00167395, 00167396) (USNM); 06 Dec 1990, W. F. Chamberlain, Light Trap, 1 ♂ (00167397) (USNM); 07 Dec 1990, W. F. Chamberlain, Light Trap, 1 ♂ (00167394) (USNM). Mt. Isa, 15 Nov 1990, W. F. Chamberlain, Light Trap, 1 ♂ (00167390), 1 ♀ (00167402) (USNM). Tozer Range, Cape York Peninsula, 12.7833°S, 143.2167°E, 122 m, 01 Jul 1948 - 05 Jul 1948, G. M. Tate, 1♀ (00167403) (USNM). BISMARCK ARCHIPELIGO: New Britain, Keravat, 17 Nov 1957, J. Smart, 1 ♂ (BMNH). CAMBODIA: Kampong Seila Dist. National Road, Pk 135, Boeung Trach Village, Picnic Resort, 11–12 Nov 2010, L. T. & R. K. Duval, S. H. Lee, W. Lee, & S. Kim, 1 ♂, 1 ♀ (SNU). FIJI: Viti Levu: Suva, 02 Mar 1985, Koebele Collection, 1 ♂ (00162346) (USNM). none, 13 Apr 1959, Haw, 2 ♀♀ (00162356, 00162357) (USNM). GUAM: Mariana Is.: Agana, port, 15 Aug 1945, H. S. Dybas, 1 ♀ (00162351) (USNM). Asan Village, Asan River at Rt. 1, 13.47278°N, 144.7135°E, 08 Jun 2008, R. S. Zack, 1 ♂ (00410393) (WSU). Asan Village, Rt. 1, Asan River outlet at Nino Perdito church, 13.47286°N, 144.71655°E, 13 Mar 2011, R. S. Zack, 38 ♂♂ (00410329 - 00410366), 26 ♀♀ (00410367 - 00410392) (WSU). Mangilao Village, University of Guam campus, Marine Biology Lab area, 13.42856°N, 144.79855°E, 03 Aug 2005, R. S. Zack, 1 ♂ (00410396) (WSU). Mangilao Village, University of Guam campus near Marine Lab dormitory field, 13.42888°N, 144.80083°E, 30 May 2008, R. S. Zack, 1 ♂ (00410395) (WSU). Mongmong-Toto-Maite Village, Pipeline Rd. off of Rt. 33, 13.45678°N, 144.76826°E, 06 Jun 2008, R. S. Zack, 1 ♂ (00410394) (WSU). Pt. Oca, 13.503°N, 144.771°E, 23 Jun 1945, G.E. Bohart and J.L. Gressit, 1 ♀ (00162354) (USNM); 16 Jul 1945, G.E. Bohart and J.L. Gressit, 1 ♂ (00162342), 1 ♀ (00162350) (USNM); 20 Dec 1945, J. L. Gressitt, Light Trap, 1 ♂ (00095351) (AMNH), 1 ♂ (00162341), 2 ♀♀ (00162352, 00162353), Light Trap, 1 ♀ (00162355) (USNM), 2 ♂♂ (00410279, 00410280), 2 ♀♀ (00410277, 00410278) (WSU). INDONESIA: Sulawesi: Utara, Dumoga-Bone N. P., 2 ♂♂ (BMNH). MICRONESIA: Ngulu Atoll: Ngulu Island, 03 Oct 1952, N. L. H. Krauss, 2 ♀♀ (00162358, 00162359) (USNM). NEW HEBRIDES: Erromana, Jul 1930, L. E. Cheesman, B. M. 1930–477, 2 ♂♂ (BMNH). Malekula, Malua Bay, May 1929, Miss L. E. Cheesman, B. M. 1929–410, 1 ♀ (BMNH). NORTHERN MARIANA ISLANDS: Rota Island: (Luta), “Josen Cristina Country” Park, 14.11836°N, 145.1857°E, 14 May 2004, R. S. Zack, 2 ♂♂ (00410401, 00410402) (WSU). (Luta), farm plots/stream area, 14.11851°N, 145.17846°E, 14 May 2004, R. S. Zack, 1 ♂ (00410403) (WSU). Saipan: 1 - 2 mi. E. of Tanapag, 10 Jan 1945, H. S. Dybas, 1 ♀ (00162349) (USNM). Saipan: Garapan, sweeping beach, 15.19458°N, 145.71678°E, 20 Jun 2006, R. S. Zack, 3;m (00410398 - 00410400) (WSU). Saipan; Kagman, Kagman Exp. Farm, Northern Marianas College, 15.17503°N, 145.77166°E, 19 Jun 2006, R. S. Zack, 1 ♂ (00410397) (WSU). Tinian: Tinian Island, 15.02333°N, 145.63305°E, 11 Jun 1946, H. K. Townes, 3 ♂♂ (00162343 - 00162345) (USNM). SAMOA: Tutuila Island: Pago Pago, 04 Sep 1923, Swezey and Wilder, 1 ♂ (00162340) (USNM); 20 Sep 1923, Swezey and Wilder, 1 ♀ (00162348) (USNM). Tutuila, 1930, Swezey & Wilder, 1 ♂, 1 ♀ (BMNH). Upolu, Apia, 9 Dec 1923, Swezey & Wilder, on Bermuda grass, 2 ♂♂ (BMNH). SOLOMON ISLANDS: Guadalcanal: Guadalcanal, 29 Mar 1944, L. J. Lipovsky, 1 ♀ (00409562) (KU); 04 Apr 1944, L. J. Lipovsky, 1 ♀ (00409563) (KU); 19 Apr 1944, L. J. Lipovsky, 1 ♂ (00409564) (KU). Ilu, 8 Apr 1963, M. Mquillan, 2 ♂♂ (BMNH). Kukun, 17 Dec 1962, P. Greenslade, 1 ♂ (BMNH). Nr. Honiara, 25027 Jun 1965, Roy. Soc. Exped. B.M. 1966-1, at light and sweeping around pond, 2 ♀♀ (BMNH). SOUTH KOREA: Gyeongsangbuk-do, Is. Ulleung, 30–31 Aug. 2010, L. T. & R. K. Duwal, 1 ♂, 2 ♀♀ (SNU). THAILAND: Bangkok: Bangkapi at light, 15 Dec 1951, M. E. Griffith, 1 ♂ (00409565) (KU).

This distinct species is recognized by the overall shiny fuscous to black body, antennae, and femora, contrasting with the pale or white cuneus, basal area of corium and clavus, antennal segments III and IV, and tibiae.

It is similar to the Nearctic Tytthus montanus in the overall dark brown to fuscous dorsum and femora and the basally pale corium, but differs in having most of the cuneus, antennal segments III and IV pale (segments I and II uniformly black) and most of the tibiae (except bases) pale or whitish, whereas in Tytthus montanus the cuneus and antennal segment III and IV are uniformly black (segments I pale and II pale on basal third to half) and the tibiae are pale only on the distal halves.

Holotype male (Fig. 11): Length to apex of hemelytron ca 3.40 mm (wing separated from body), length to base of cuneus 2.33 mm (wing separated from body), width across hemelytra 1.12 mm. Head: Length 0.43 mm, width across eyes 0.64 mm, interocular width 0.32 mm. Labium: Length 1.12 mm. Antenna: Segment I length 0.32 mm, II 1.04 mm, III 0.64 mm, IV ca 0.51 mm (curled and in glue). Pronotum: Length 0.42 mm, basal width 0.82 mm.

Coloration: Head: Uniformly black, with a vague pale spot on interocular space adjacent to each eye; eyes dark reddish brown. Labium: Segments I, II, and apex of IV dark brown; segment III, apex of II, and basal two thirds of IV pale or whitish. Antenna: Segments I and II fuscous to black; segment III and IV pale or whitish. Pronotum: Uniformly black. Hemelytron: Fuscous to black, with basal one fourth of corium and clavus and cuneus, except for apex, pale or white; membrane translucent brown. Ostiolar evaporative area: Fuscous to black. Ventral surface: Propleura black, ventral areas of thorax dark to reddish brown; abdomen dark reddish brown. Legs: Procoxae reddish brown, mesocoxa reddish brown with only apex pale, metacoxa uniformly pale to whitish; femora uniformly black; tibiae pale yellow to whitish, with only basal one fourth of each fuscous to black; tarsi and claws pale yellow to whitish.

Structure, texture, and vestiture: Head: Shiny, impunctate, width subequal to length, shiny; buccula slender (less than half the width of labial segment I), tapering posteriorly; sparsely set with scattered, short, semierect, dark brown setae on vertex and frons and a few longer, erect setae along posterior margin. Labium: Extending to bases of mesocoxae; segment I extending only to base of head. Pronotum: Shiny, impunctate; anterior angles rounded, lateral margins weakly concave, posterior anterior angles strongly flared, posterior margin concave; sparsely set with scattered, short, erect and semierect, dark brown setae. Mesoscutum: Broadly exposed, impunctate, sparsely set with short, erect, dark brown setae. Scutellum: Equilateral, impunctate, sparsely set with short, erect, dark brown setae. Hemelytron: Macropterous, impunctate, basal width of cuneus about two thirds the length, membrane fully developed with two areoles; evenly set with short, semierect dark brown setae (pale setae on pale or white areas).

Male genitalia: The unique holotype was not dissected.

Female: Unknown.

The holotype was taken on Oryza sp. (Poaceae).

Described and known only from the holotype collected in Colombia.

Holotype ♂ (00162207) (USNM): COLOMBIA: Valle del Cauca: Palmira, 25 Oct. 1958, G. Bravo, Arroz, Oryza sp. (Poaceae).

The left hemelytron of the holotype is missing, and the right one is glued to the point next to the specimen. Otherwise, the specimen is in reasonably good condition. Carvalho (1984) illustrated the holotype, apparently before the specimen was damaged. Figure 11 depicts the original condition, using Adobe Photoshop to reconstruct the position of both hemelytra.

This species is recognized by the dark brown head and antennal segments I and II, the pale brown pronotum with the anterior half darker brown or reddish brown, the translucent, smoky-brown hemelytra, and the mostly pale legs with only the distal thirds of the femora fuscous. All known specimens of this species are macropterous.

This species keys to Tytthus femoralis based on the pale tibiae with fuscous knee spots and the apically fuscous hind femora. Tytthus entrerianus can be distinguished from Tytthus femoralis by the dark brown or fuscous antennal segment II and having only apical third of the hind femur infuscated.

Male (n = 1, holotype in parentheses) (composite description based on Carvalho and Carpintero, 1986, and one USNM paratype) (Fig. 12): Length to apex of hemelytron 2.65 mm (3.00 mm), length to base of cuneus 1.80 mm (not given), width across hemelytra 0.74 mm (0.90 mm). Head: Length 0.40 mm (0.30 mm), width across eyes 0.54 mm (0.50 mm), interocular width 0.29 mm (0.30 mm). Labium: Not visible, imbedded in glue (not given). Antenna: Segment I length 0.34 mm (0.30 mm), II 0.94 mm (1.00 mm), III 0.74 mm (0.70 mm), IV 0.56 mm (0.50 mm). Pronotum: Length 0.27 mm (0.20 mm), basal width 0.67 mm (0.70 mm).

Coloration: Head: Uniformly dark brown; pale spot near inner margin of eye absent. Labium: Not visible. Antenna: Segments I and II dark brown; segment III and IV brownish yellow. Pronotum: Anterior half dark brown; posterior half yellowish brown. Mesoscutum: Reddish brown. Scutellum: Reddish brown. Hemelytron: Brown, with basal one fourth and cuneus paler yellowish brown; membrane pale translucent brown. Ostiolar evaporative area: Brown. Ventral surface: Thorax and abdomen brown to reddish brown, genital capsule dark brown. Legs: Pale brownish yellow, apical third of each femur and bases of tibiae (knees) dark brown.

Structure, texture, and vestiture: Shiny, impunctate (buccula imbedded in glue); sparsely set with short, semierect setae. Labium: Not visible. Antenna: Segment I with a few scattered, short, recumbent setae and two long, erect, bristlelike setae at apex; segment II with relatively few, short, recumbent setae. Pronotum: Shiny, impunctate; anterior angles rounded; lateral margins straight, gradually widening to posterior angles; posterior margin weaky concave. Mesoscutum: Shiny, impunctate; set with only a few semierect setae. Scutellum: Shiny, impunctate; set with a few semierect setae. Hemelytron: Macropterous, impunctate, subparallel; length of cuneus two times length; membrane entire with two areoles; set with sparsely scattered, recumbent, brown setae.

Male genitalia (based on Carvalho and Carpintero’s 1986, figures): Left paramere (Fig. 125): Mitt-shaped; right arm long, slender; left arm shorter, spinelike. Right paramere (Fig. 126): Oval. Endosoma (Fig. 127): C-shaped, with apparent secondary gonopore at middle. Phallotheca: Not figured.

Female: None examined; described from 5 paratype ♀♀. Carvalho and Carpintero (1986) indicated that the general aspect and dimensions of females were similar to males.

Unknown.

Described and known only from Entre Rios Province, Argentina.

Paratype: ARGENTINA: Entre Rios: Concordia, 31.4°S, 58.033°W, 21 m, Apr 1939 [L. Carpintero], ♂ (00138710) (USNM).

ARGENTINA: Entre Rios: Concordia, 31.4°S, 58.033°W, Jan 1979, collector unknown, 1 ♂ (00138710) (USNM).

This species is distinguished by the combination of a black head, pronotum, and scutellum; translucent hemelytra usually marked with dark brown through middle of the corium; the fuscous antennal segment I, with only the apex pale; the pale antennal segment II having a fuscous basal band; the fuscous hind femur contrasting with the pale yellow pro- and mesofemora; and the pale tibiae with the bases or “knees” fuscous. All known specimens of this species are macropterous.

Tytthus femoralis keys to Tytthus entrerianus based on the pale tibiae with knee spots and the apically infuscated hind femora. It can be distinguished from Tytthus entrerianus by the pale antennal segment II and the more extensively infuscated hind femur.

Male (n =10, plus holotype in parentheses) (Fig. 13): Length to apex of hemelytron 2.56–3.42 mm (2.83 mm), length to base of cuneus 1.90–2.25 mm (2.08 mm), width across hemelytra 0.80–0.99 mm (0.88 mm). Head: Length 0.30–0.34 mm (0.35 mm), width across eyes 0.61–0.67 mm (0.63 mm), interocular width 0.27–0.29 mm (0.29 mm). Labium: Length 1.20–1.38 mm (1.28 mm). Antenna: Segment I length 0.38–0.50 mm (0.38 mm), II 0.94–1.21 mm (1.10 mm), III 0.58–0.72 mm (0.66 mm), IV 0.40–0.59 mm (0.58 mm). Pronotum: Length 0.34–0.38 mm (0.35 mm), basal width 0.75–0.90 mm (0.78 mm).

Coloration: Head: Uniformly fuscous to black, with a large, yellow, interocular spot near inner margin of each eye, spots nearly converging on some specimens; eyes fuscous to dark reddish brown. Labium: Pale yellow, with apical half of segment IV brown. Antenna: Segment I fuscous to black, with apical one quarter pale or yellowish; segment II pale yellowish brown, with a distinct fuscous band at base and sometimes with apex infuscated; segments III and IV black. Pronotum: Uniformly black. Mesoscutum: Uniformly black. Scutellum: Uniformly black. Hemelytron: Translucent, with a dark brown cloud or patch on apical half of cor- ium and inner apical half of clavus ranging from somewhat indistinct to a definite dark pattern (Figs 13, 14); membrane pale translucent brown. Ostiolar evaporative area: Dark brown to fuscous, with a yellow patch on posterior edge. Ventral surface: Thorax and abdomen uniformly fuscous to black. Legs: Coxae pale yellow, with basal thirds to halves dark brown; pro- and meso femora pale yellow, metafemur dark brown to fuscous, with basal third and apex pale yellow; tibiae pale, each with a fuscous base or “knee”; tarsi, and claws pale yellow.

Structure, texture, and vestiture: Head: Shiny, impunctate; buccula relatively broad, tapering posteriorly, ending near level with middle of eye; set with scattered, semierect setae. Labium: Extending to metacoxe or base of abdomen; segment I extending beyond base of head onto xyphus to bases of procoxae. Antenna: Segment I sparsely set with recumbent setae and two erect, subapical bristlelike setae; segment II set with only very short, recumbent setae. Pronotum: Shiny, impunctate; anterior angles rounded; lateral margins weakly concave, widening at posterior angles; posterior margin weakly sinuate; calli weakly swollen; set with relatively long, recumbent setae. Mesoscutum: Weakly shining, impunctate, broadly exposed; set with a few scattered semierect setae. Scutellum: Shiny, impunctate; equilateral; set with scattered erect and semierect setae. Hemelytron: Macropterous, with cuneus and membrane fully developed, extending well beyond apex of abdomen; evenly set with relatively long, recumbent setae.

Male genitalia: Left paramere (Fig. 128): Mitt-shaped; right arm long, broad, acute apically; left arm short, apically acute. Right paramere (Fig. 129): Endosoma (Fig. 130): S-shaped. Phallotheca (Fig. 131): Slender, apically acute.

Female (n =5) (Fig. 14): Length to apex of hemelytron 2.82–3.52 mm mm, length to base of cuneus 2.05–2.50 mm, width across hemelytra 1.00–1.14 mm. Head: Length 0.30–0.35 mm, width across eyes 0.62–0.69 mm, interocular width 0.30–0.32 mm. Labium: Length 1.12–1.50 mm. Antenna: Segment I length 0.32–0.50 mm, II 0.85–1.17 mm, III 0.53–0.75 mm, IV 0.54–0.59 mm.

Pronotum: Length 0.40–0.41 mm, basal width 0.91–0.99 mm.

Similar to males in general appearance and coloration, differing primarily in the overall broader form. The one color exception is that the ventral area of the abdomen in many females is pale yellow, whereas in males the abdomen is always uniformly fuscous to black.

The specific epithet A "femoralis" denotes the dark brown to fuscous hind femur in contrast to the uniformly pale pro- and mesofemora.

Four specimens from San Carlos, Ecuador, with the label a "Host-Eggs of Perkinsiella spp. " Three specimens intercepted on Musa sp. certainly represent incidental or sitting records.

So far recorded from Bolivia, Brazil, Colombia, Costa Rica, Cuba, Ecuador, Guatemala, Honduras, Jamaica, Mexico, Panama, and Peru.

Holotype ♂ (00161890) (USNM): ECUADOR: Manabi: Bahia de Caraquez (35 kms SE), 10 May 1975, Ashley B. Gurney. Paratypes: BOLIVIA: El Beni: Capivara on Rio Itenez, approx. 20 km E. of Versalles, 22 Jul 1964, J. K. Bouseman, J. Lussenhop, 1 ♂ (00165937) (AMNH). BRAZIL: Amazonas: Reserva Ducke, 25 km NNE Manaus, 2.9136°S, 59.9464°W, 120 m, 26 Jul 1973, R. T. Schuh, 1 ♀ (00165831) (AMNH). COLOMBIA: Valle del Cauca: Jamundi, 3.25833°N, 76.54°W, 13 Jul 1985, F. Garcia, on Jameo arroz (Poaceae), 6 ♂♂ (00161854 - 00161859), 2 ♀♀ (00161860, 00161861) (USNM). COSTA RICA: San Jose: San Jose, 9.9333°N, 84.0833°W, 1147 m, 1932, H. Schmidt, 1 ♂ (00165935), 1 ♀ (00165936) (AMNH). CUBA: Origin unknown, intercepted at Baltimore, Maryland by APHIS/PPQ, 26 Nov 1925, in “ship’s light socket”), 1 ♀ (00161862) (USNM). ECUADOR: Chimborazo: Huigra, 15 Jun 1914, H. S. Parish, 1 ♂ (00161899) (USNM). El Oro: Victoria-Arenillas, 150 m, 18 Aug 1977 - 19 Aug 1977, L. Pena G., 1 ♀ (00161931) (USNM). Guayas: Duran, 91 m, 23 Jun 1914, H. S. Parish, 2 ♂♂ (00161900, 00161901) (USNM), 2 ♂♂ (CU). Ingenio San Carlos, 07 May 1982, Robert Morey, host–eggs of Perkinsiella spp., 1 ♂ (00161898), 3 ♀♀ (00161930, 00161936 - 00161937) (USNM). Los Rios: Babahoyo, 21 Jun 1975, Cohen, Langley & Monnig, at blacklight, 1 ♂ (00161934) (USNM). Chone, 0.6833°S, 80.1°W, 110 m, 09 May 1975, Ashley B. Gurney, 2 ♂ (00161929, 00161935) (USNM). Manabi: Chone, 0.6833°S, 80.1°W, 110 m, 09 May 1975, Ashley B. Gurney, 2 ♂♂ (00161929, 00161935) (USNM). Napo: Baeza (72 Km E), 1280 m, 14 Apr 1977, Elaine R. Hodges, near cut trees and bamboo, 1 ♂ (00161933) (USNM). Origin unknown, intercepted at Long Beach, California, by APHIS/PPQ, 12 May 1997, on Musa sp., 1 ♂ (00161891), 1 ♀ (00161932) (USNM). Origin unknown, intercepted at San Diego, California by APHIS/PPQ, 32.71528°N, 117.15639°W, 20 May 2008, on Musa sp. (fruit), 1 ♂ (00161902) (USNM). GUATEMALA: Locality unknown, 13 Sep 2007, P. Perez, Musa sp. (Musaceae), 1 ♀ (00161864) (USNM). HONDURAS: Atlantida: Lancetilla, 1900 - 1900, M. Bates, Paratype, 2 ♂♂ (00165933, 00165934) (AMNH). JAMAICA: St. Ann Parish: 5 mi S of St Anns Bay, 05 Jul 1971, J. A. Slater & R. M. Baranowski & J. E. Harrington, 1 ♂ (00165830) (AMNH). MEXICO: Origin unknown, 03 Nov 1988, intercepted at Elpaso, Texas, by APHIS/PPQ, on Musa sp. (Musaceae), 1 ♀ (00161863) (USNM). PANAMA: Pinogana: El Real, 08 Aug 1952, F. S. Blanton, 4 ♀♀ (00161850 - 00161853), 1 ♀ (00161849) (USNM). PERU: Huanuco: Leoncio Prado Co.: Tingo Maria, 671 m, 19 Apr 1969–24 Apr 1969, P. & P. Spangler, 1 ♂ (00161938), 1 ♀ (00161939) (USNM).

This new species is distinguished by the small size, the uniformly dark brown head and pronotum, fuscous to black antennae with segment II thickened and subequal to apical diameter of segment I, the pale or whitish hemelytra, and the brownish-yellow legs. The only known male (holotype) of this species is macropterous and the only known female is brachypterous with an abbreviated membrane.

It is superficically similar to several species, such as Tytthus mexicanus and Tytthus panamensis, based on the dark head, pronotum and scutellum and pale hemelytra. It is distinguished from these and all other species by the pale tibiae lacking knee spots, uniformly fuscous antennae, the thickened antennal segment II, pale femora, and relatively small size.

Holotype male (Fig. 15): Length to apex of hemelytron 2.14 mm, length to base of cuneus 1.54 mm, width across hemelytra 0.69 mm. Head: Length 0.29 mm, width across eyes 0.51 mm, interocular width 0.29 mm. Labium: Length 0.69 mm. Antenna: Segment I length 0.22 mm, II 0.75 mm, III 0.43 mm, IV 0.29 mm. Pronotum: Length 0.27 mm, basal width 0.67 mm.

Coloration: Head: Uniformly dark brown, with a somewhat indistinct, small, yellow interocular spot near inner margin of eye; eyes dark brown to reddish brown, especially around margins. Labium: Uniformly brownish yellow, with only apex of segment IV dark brown or fuscous. Antenna: Uniformly fuscous to black. Pronotum: Uniformly dark brown. Mesoscutum: Dark brown. Scutellum: Dark brown. Hemelytron: Pale or whitish, evenly tinged with pale brown; membrane pale translucent brown. Ostiolar evaporative area: Pale brownish yellow. Ventral surface: Propleura dark brown; ventral areas of thorax and abdomen reddish brown, genital capsule darker brown to nearly black. Legs: Coxae pale to pale brownish yellow, with bases reddish brown; femora brownish yellow, tinged with orange or brownish orange; tibiae, tarsi, and claws pale browish yellow.

Structure, texture, and vestiture: Head: Shiny, impunctate; buccula slender, ending posteriorly about level with middle of eye; sparsely set with very short, recumbent and semierect, setae. Labium: Extending to apices of meso- or bases of metacoxae; segment I short, extending only slightly beyond base of head. Antenna: Segment I with only a few short, recumbent setae and two erect, subapical, bristlelike setae; segment I densely set with short, recumbent setae; segment II gradually thickened to apex, apical width equal to or greater than diameter of segment I. Pronotum: Shiny, impunctate; calli weakly swollen; anterior angles rounded; lateral margins nearly straight, gradually widening to posterior angles; basal margin weakly concave; sparsely set with only short, recumbent setae. Mesoscutum: Narrowly exposed; sparsely set with short, recumbent setae. Scutellum: Impunctate, weakly shiny; sparsely set with short, recumbent setae. Hemelytron: Entire, suparallel; length of cuneus about two times basal width; membrane with two areoles, extending well beyond apex of abdomen.

Male genitalia: Left paramere (Fig. 132): Mitt-shaped; right arm long, evenly slender, and rounded apically; left arm short, apically acute. Right paramere (Fig. 133): elongate oval. Endosoma (Fig. 134): C-shaped or weakly S-shaped. Phallotheca (Fig. 135): Relatively slender, apically acute.

Brachypterous female (n = 1; somewhat teneral) (Fig. 16): Length to apex of hemelytron 1.57 mm, length to base of cuneus 1.36 mm, width across hemelytra 0.70 mm. Head: Length 0.27 mm, width across eyes 0.53 mm, interocular width 0.32 mm. Labium: Length 0.64 mm. Antenna: Segment I length 0.19 mm, II 0.61 mm, III 0.40 mm, IV 0.27 mm. Pronotum: Length 0.27 mm, basal width 0.67 mm.

Similar to male in general coloration, but differing in the broader, more oval form, the slightly more slender antennal segment II (but with apex still nearly equal to apical diameter of segment I), and abbreviated hemelytron, with the length of the cuneus subequal to the basal width and the membrane shortened and extending only to the apex of the abdomen.

Host. Beaten by A. G. Wheeler from crowns of Muhlenbergia rigens (Benth) Hitchc. [Poaceae].

Known only from New Mexico, USA.

Holotype ♂ (00161896) (USNM): UNITED STATES: New Mexico: Grant Co.: Little Walnut Road, Gila National Forest, North of Silver City, 12 May 2008, A. G. Wheeler, Jr. Paratype: Same data as for holotype, 1 ♀ (00161895) (USNM).

This species is distinguished by the black head and antennae, strongly infuscated or brown pronotum with the anterior collarlike margin narrowly whitish, the translucent smoky-brown hemelytra, and the orange-brown legs with a slender, dorsal and anterior red line on each femur, and a posterior red line on the metafemur. All known specimens of this species are fully macropterous.

Tytthus insperatus is similar to Tytthus balli and Tytthus uniformis in overall size, body shape, and general coloration. It differs from both species in having an imbrowned pronotum with a pale or white collar and distinct red lines on the femora.

Holotype male (Fig. 17): Length to apex of hemelytron 2.98 mm, length to base of cuneus 2.06 mm, width across hemelytra 0.77 mm. Head: Length 0.45 mm, width across eyes 0.59 mm, interocular width 0.27 mm. Labium: Length 1.09 mm. Antenna: Segment I length 0.35 mm, II 1.44 mm, III 0.91 mm, IV 0.54 mm. Pronotum: Length 0.34 mm, basal width 0.70 mm.

Coloration: Head: Uniformly fuscous to nearly black, with only a small, indistinct pale, interocular spot near inner margin of each eye; eyes fuscous. Labium: Pale brownish yellow, apical half of segment IV darker brown. Antenna: Uniformly black. Pronotum: Dark brown, fading to paler orange brown posteriorly, narrow anterior collar white. Mesoscutum: Pale brownish orange. Scutellum: Pale brownish orange. Hemelytron: Uniformly pale translucent brown, with pale orange highlights; membrane clear or translucent. Ostiolar evaporative area: Pale brownish orange. Ventral surface: Thoracic area pale brownish orange; abdomen pale with green and orange highlights; genital capsule fuscous. Legs: Coxae pale brownish orange; femora pale brownish orange with a slender dorsal and anterior red stripe on pro- and mesofemora and a dorsal, anterior, and posterior red stripe on metafemur; tibiae, tarsi, and claws fuscous to black.

Structure, texture, and vestiture: Head: Shiny, impunctate, width subequal to length; nearly glabrous with only a few scattered erect and semierect setae; buccula narrow, tapering posteriorly ending near level with middle of eye. Labium: Extending to mesocoxae; segment I not extending beyond base of head. Antenna: Segment I only slightly thicker than segment II, sparsely set with short, recumbent setae and two erect, subapical, bristlelike setae; segment II densely set with short, recumbent setae. Pronotum: Shiny, impunctate; calli weakly swollen; anterior angles angulate; lateral margins convex, flaring at posterior angles; posterior margin weakly concave; nearly glabrous, with only a few scattered, recumbent setae. Mesoscutum: Shiny, impunctate, with a few scattered, recumbent setae. Scutellum: Shiny, impunctate, with a few scattered, recumbent setae. Hemelytron: Macropterous, subparallel, cuneus and membrane fully developed, extending well beyond apex of abdomen; evenly set with short, recumbent setae.

Male genitalia: Left paramere (Fig. 136): Mitt-shaped; right arm, long, stout, bluntly rounded apically; left arm short, apically blunt. Right paramere (Fig. 137): Elongate oval. Endosoma (Fig. 138): Strongly S-shaped. Phallotheca (Fig. 139): Slender, apically acute.

Female (n = 3) (Fig. 18): Length to apex of hemelytron 3.14–3.33 mm, length to base of cuneus 2.33–2.43 mm, width across hemelytra 0.96–1.06 mm. Head: Length 0.40–0.43 mm, width across eyes 0.61–0.66 mm, interocular width 0.32–0.34 mm. Labium: Length 1.12–1.15 mm. Antenna: Segment I length 0.32–0.34 mm, II 1.12–1.47 mm, III 0.96–1.01 mm, IV 0.53–0.54 mm. Pronotum: Length 0.34–0.37 mm, basal width 0.77–0.82 mm.

Similar to male in overall coloration and body form.

Unknown. The Maricopa specimen below from “on cotton” almost certainly represents an incidental or sitting record.

Described from Tucson (Pima County), Arizona, and later reported from Buckeye (Maricopa County), Arizona, and Calexico (Imperial County), California (Carvalho and Southwood 1955).

Holotype ♂ (00162203) (USNM): UNITED STATES: Arizona: Pima Co.: Tucson, 32.22167°N, 110.92583°W, 07 Jun 1924, A. A. Nichol, . Paratypes: Same data as for holotype, 1 ♀ (allotype) (00161894) (USNM), 1 ♀ (00167071) (CNC).

UNITED STATES: Arizona: Cochise Co.: W Hereford, Rio San Pedro, 4150’, 22 July 1974, E. R. Hoebeke, 1 ♂ (CU). Maricopa Co.: Buckeye, 33.37028°N, 112.58306°W, 06 Jun 1935, H. G. Johnston, on cotton, 1 ♂ (00138719) (USNM).

Based on the original description and adult habitus illustration, this species can be distinguished by the overall dark color with the cuneus, embolium, and the narrow outer edge of the corium pale yellow, the medially fuscous membrane, the dark antennal segment I with only the apex pale, and the pale antennal segment II with the apex fuscous. This combination of characters places Tytthus juturnaiba near the North American Tytthus vagus, but differs in having an apparent pale yellow antennal segment II, with only the apex black, and uniformly pale yellow legs, whereas Tytthus vagus has a uniformly black antennal segment II, and the hind femur is fuscous on the apical two thirds.

I have, however, studied a photograph of the holotype of Tytthus juturnaiba stored in the PBI Heteroptera Species Database (http://research.amnh.org/pbi/heteropteraspeciespage/popupimage.php?imagename=AMNH_PBI 00174988.jpg) that appears to indicate that antennal segment I actually is pale at the base and apex (not just pale at the apex) and segment II is uniformly dark (not yellow, with the apex fuscous). Based on this photograph, which contradicts the antennal characters described in the original description, Tytthus juturnaiba almost certainly is conspecific with Tytthus neotropicalis and would run to this species in my key. Nevertheless, I refrain from making a formal synonymy until I can examine the holotype (which was unvailable at the time of this study) deposited in the Museu Nacional, Rio de Janeiro, Brazil.

Holotype male (based Carvalho and Wallerstein 1978): Length to apex of hemelytron 2.90 mm, width across hemelytra 0.90 mm. Head: Length 0.20 mm, width across eyes 0.60 mm, interocular width 0.28 mm. Antenna: Segment I length 0.30 mm, II 0.80 mm, III 0.50 mm, IV missing. Pronotum: Length 0.40 mm, basal width 0.80 mm. Cuneus: Length 0.40 mm, basal width 0.28 mm.

Coloration (based on Carvalho and Wallerstein 1978): General coloration black, outer margin of exocorium, embolium, and cuneus pale yellow; middle of membrane fuscous; antenna pale yellow, segment I (except extreme apex) and apex of segment II black. Undersurface black. Legs pale yellow.

Structure, texture, and vestiture (based on Carvalho and Wallerstein 1978): Information not given.

Male genitalia (based on Carvalho and Wallerstein 1978): Left paramere (Fig. 140): Mitt-shaped; right arm longest; left arm shorter, spinelike. Right paramere (Fig. 141): Elongate oval. Endosoma (Fig. 142): Strongly C-shaped, bent at middle. Phallotheca (Fig. 143): Broad, apically acute.

Female (based on Carvalho and Wallerstein 1978): Length to apex of hemelytron 3.20 mm, width across hemelytra 1.00 mm. Head: Length 0.20 mm, width across eyes 0.50 mm, interocular width 0.28 mm. Antenna: Segment I length 0.20 mm, II 0.80 mm, III and IV missing. Pronotum: Length 0.40 mm, basal width 0.90 mm. Cuneus: Length 0.44 mm, basal width 0.28 mm.

Unknown.

Described and known only from the holotype and allotype taken at Logoa Juturnaiba, Araruama, Estado do Rio de Janeiro, Brazil, Nov. 1976 (Carvalho and Wallerstein 1978).

None.

This species is distinguished by the black antennae, except for the pale apex of segment I; the black pronotum; the brown mesoscutum and scutellum; the smoky brown to dirty white hemelytra; and the pale yellowish legs, with distinct black spines and knee spots at the bases of the tibiae.

Tytthus mexicanus is most similar to Tytthus femoralis and Tytthus entrerianus based on the pale tibiae with knee spots and antennal segment II lacking erect, bristlelike setae. It can be distinguished from both species by the uniformly pale hind femora and dark brown to fuscous antennal segment II.

Male (n = 4; plus holotype in parentheses) (Fig. 19): Length to apex of hemelytron 2.75–2.58 mm (2.55 mm), length to base of cuneus 2.90–2.05 mm, width across hemelytra 0.88-0.93 mm (0.86 mm). Head: Length 0.32–0.34 mm (0.30 mm), width across eyes 0.59–0.61 mm (0.59 mm), interocular width 00.27–0.29 mm (0.27 mm). Labium: Length 1.12–1.14 mm (1.12 mm). Antenna: Segment I length 0.30–0.34 mm (0.32 mm), II 0.91–1.07 (1.02 mm), III 0.54–0.64 mm (0.58 mm), IV 0.51–0.64 mm (missing). Pronotum: Length 0.34–0.35 mm (0.32 mm), basal width 0.78–0.82 mm (0.77 mm).

Coloration: Head: Uniformly fuscous to black, with a large, distinct, pale yellow, interocular spot near inner margin of each eye, spots nearly contiguous in some specimens; eyes reddish brown, often fading to silver with a reddish tinge. Labium: Uniformly pale yellow; apex of segment IV dark brown to fuscous. Antenna: Segment I fuscous to black, with apical one fourth yellow (length of yellow area equal to diameter of segment) and narrowed basal one fourth shiny; segment II dark brown to black (basal area darker on “paler” dark brown segments); segments III and IV dark brown to fuscous. Pronotum: Uniformly shiny fuscous, weakly swollen calli sometimes very slightly paler brown. Mesoscutum: Dark brown to fuscous, lateral angles pale yellow on some specimens. Scutellum: Dark brown to fuscous, slightly paler apically. Hemelytron: Pale translucent smoky brown to dirty white; veins brown and often narrow inner margin of clavus brown. Ostiolar evaporative area: Dark brown to fuscous, often invaded with pale areas posteriorly. Ventral surface: Thorax uniformly dark brown to fuscous; abdomen dark brown to fuscous laterally, pale ventrally, genital capsule black. Legs: Coxae pale brownish yellow, with bases brown; femora uniformly pale yellow to whitish; tibia pale yellow to whitish with spines and bases, or knees, dark brown to fuscous; tarsi and claws pale yellow.

Structure, texture, and vestiture: Head: Shiny, impunctate, wider than long; buccula relatively narrow, ending posteriorly near level with middle of eye; sparsely set with long, erect and suberect, pale brown to brown setae. Labium: Extending to bases of hind coxae or just onto abdominal segment II; segment I extending to bases of fore coxae. Antenna: Segment I with short, recument setae and two long, black, bristlelike setae near apex before pale area; segment II thickly set with short, recumbent pale brown setae much shorter than diameter of segment. Pronotum: Shiny, impunctate; calli weakly swollen but distinct; anterior rounded; lateral margins concave, strongly flaring at posterior angles; posterior margin weakly sinuate; set with short, erect to semierect setae. Mesoscutum: Shiny, impunctate, broadly exposed; with numerous semierect and recumbent, pale brown setae. Scutellum: Shiny, impunctate; thickly set with semierect and recumbent pale brown setae. Hemelytron: Macropterous, subparallel when paired, with fully developed cuneus and membrane, extending well beyond apex of abdomen; evenly scattered with short, recumbent, pale brown setae.

Male genitalia: (Fig. 144): Mitt-shaped; right arm longest, stout, slightly constricted basally, weakly pointed apically; left arm short, apically acute. Right paramere (Fig. 145): Elongate oval, strongly rounded apically. Endosoma (Fig. 146): C-shaped, nearly S-shaped with apical quarter curving slightly upward. Phallotheca (Fig. 147): moderately stout, apically acute.

Female:(n = 2) (Fig. 20): Length to apex of hemelytron 2.88–2.93 mm, length to base of cuneus 2.10–2.23 mm, width across hemelytra 0.99–1.04 mm. Head: Length 0.32–0.37 mm, width across eyes 0.61–0.62 mm, interocular width 0.29–0.30 mm. Labium: Length 1.20–1.22 mm. Antenna: Segment I length 0.29–0.30 mm, II 0.88–0.91 mm, III 0.61–0.64 mm, IV 0.61–0.62 mm. Pronotum: Length 0.35–0.37 mm, basal width 0.86–0.88 mm.

Similar to male in general color and pubescence, differing primarily in the slightly broader form.

Named for the country in which it was collected, Mexico.

Unknown.

Known only from Baja California Sur, Nayarit, and Sinaloa, Mexico.

Holotype ♂ (00166142) (USNM): MEXICO: Sinaloa: Culiacan Co.: Camino Real Tres Rios, 3 km N Mex. 15 at toll bridge gate, 08 Aug 1981, S. Nichols. Paratypes: MEXICO: Nayarit: San Blas, 16 Oct 1973, S.C. Williams, K.B. Blair, & C.L. Mullinex, 1 ♂ (00409874), 1♀ (00409875) (CAS). Sinaloa: Culiacan Co.: Camino Real Tres Rios, 3 km N Mex. 15 at toll bridge gate, 08 Aug 1981, S. Nichols, 2 ♂ (00163423, 00166141) (USNM). Other Paratypes: MEXICO: Baja Calif. Sur: San Javier 11 Oct 1981, D. Faulkner & F. Andrews, at blacklight, 1 ♂ (CDFA). 112.2 mi SE San Perdito near Rancho Saucito, 8 Oct 1981, F. Andrews & D. Faulkner, 1 ♀ (SDNH). Nayarit: Choix, 5 Aug 1968, A. Sears, R. C. Gardner, & C. S. Glaser, 1 ♂ (UCD). 5.5 mi NW Choix, 14 Jul 1968, A. Sears, R. C. Gardner, & C. S. Glaser, 1 ♀ (UCD). Mazatlan, 27 Mar 1979, L. D. French, 1 ♂ (UCD).

This species is distinguished by the overall dark brown to nearly black coloration, with the basal thirds of the corium and clavus white, the pale yellow antennal segment I and black segment II, the dark brown metafemur, and pale yellow tibiae. It is known from only two macropterous males (Fig. 21), one macropterous female (Fig. 22), and numerous brachypterous females (Fig. 23).

Tytthus montanus is similar to Tytthus wheeleri in having mostly dark brown hemelytra with only the base pale or white but is distinguished from that species by the larger size (2.70 mm vs. less than 2.00 mm in Tytthus wheeleri), pale yellow antennal segment I, red-streaked pro- and mesofemora, and dark brown metafemur (versus entirely pale yellow legs in Tytthus wheeleri). It is also similar to Tytthus alboornatus but is distinguished by the distally dark hemelytron (versus distally white or pale cuneus or pale area across posterior margin of hemelytra in brachypters).

Macropterous male (n=1; holotype in parentheses) (Fig. 21): Length to apex of hemelytron 2.83 mm (2.75 mm), length to base of cuneus 1.98 mm (1.75 mm), width across hemelytra mm. Head: Length 0.38 mm (0.37 mm), width across eyes 0.64 mm (0.64 mm), interocular width 0.34 mm (0.32 mm). Labium: Length 0.98 mm (0.93 mm). Antenna: Segment I length 0.27 mm (0.30 mm), II 0.93 mm (0.93 mm), III 0.56 mm (missing), IV mm (missing). Pronotum: Length 0.45 mm (0.43 mm), basal width 0.72 mm (0.74 mm).

Coloration: Head: Dark reddish brown (holotype) to black; pale yellow interocular spot near inner margin of eye indistinct; eyes dark brown to nearly black. Labium: Mostly pale yellow, with segment I and apex of segment IV dark brown. Antenna: Segment I pale yellow, narrowly fuscous at base; segments II mostly fuscous to black, with only base pale yellow; segments III–IV nearly black (segments II–IV yellowish brown in holotype). Pronotum: Uniformly dark brown (holotype) to black. Mesoscutum: Uniformly dark brown (holotype) to black. Scutellum: Uniformly dark brown (holotype) to black. Hemelytron: Largely dark brown (holotype) to black, with only basal third of corium and clavus white; membrane translucent brown to smoky black. Ostiolar evaporative area: Dark reddish brown (holotype) to black. Ventral surface: Uniformly dark reddish brown (holotype) to black. Legs: Coxa pale yellow, reddish brown or black at bases; pro- and mesofemora pale yellow, lightly tinged with pale orange to more reddish brown with bases pale (holotype), metafemur dark reddish brown (holotype) to dark brown, with basal one third and apex pale yellow; pro- and mesotibiae pale yellow to more reddish brown with bases pale, metatibia yellow (holotype) to dark brown; tarsi and claws pale yellow.

Structure, texture, and vestiture: Head: Shiny, impunctate, width subequal to length; buccula very narrow, tapering posteriorly, ending at level before middle of eye. Labium: Extending to bases of mesocoxae; segment I not extending to base of head. Antenna: Segment I with only very short, fine, recumbent setae, with two erect, subapical, bristlelike setae; segment II thickly set with short, recumbent setae, intermixed with a few more erect, short setae on distal half. Pronotum: Shiny, with a glaucous sheen over weakly defined calli, impunctate; anterior angles rounded, lateral margins concave, strongly flaring at posterior angles; posterior margin weakly sinuate; sparsely set with short, fine, recumbent setae. Mesoscutum: Shiny, impunctate, with a few scattered, recumbent setae. Scutellum: Shiny, impunctate, with a few short, fine, recumbent setae. Hemelytron: Macropterous, cuneus and membrane fully developed, extending well beyond apex of abdomen; evenly set with scattered, short, fine, recumbent setae.

Male genitalia: Left paramere (Fig. 148): Mitt-shaped; right arm longest, stout, distally blunt; left arm short, blunt. Right paramere (Fig. 149): Oval. Endosoma (Fig. 150): C-shaped to weakly S-shaped. Phallotheca (Fig. 151): Slender, apically acute.

Macropterous female (n = 1) (Fig. 22): Length to apex of hemelytron 2.65 mm, length to base of cuneus 1.87 mm, width across hemelytra 0.94 mm. Head: Length 0.37 mm, width across eyes mm, interocular width 0.32 mm. Labium: Length 0.91 mm. Antenna: Segment I length 0.22 mm, II 0.59 mm, III 0.40 mm, IV 0.40 mm. Pronotum: Length 0.42 mm, basal width 0.72 mm.

Very similar to the two macropterous males in overall shape and structure, differing in the significantly shorter antennal segments and in having all femora and the basal halves of the tibiae dark brown to fuscous.

Brachypterous female (n = 5) (Fig. 23): Length to apex of hemelytron 1.85–2.20 mm, length to base of cuneus 1.53–1.75 mm, width across hemelytra 0.77-0.86 mm. Head: Length 0.38–0.42 mm, width across eyes 0.61–0.62 mm, interocular width 0.32–0.34 mm. Labium: Length 0.88-0.91 mm. Antenna: Segment I length 0.22-0.24 mm, II 0.64–0.78 mm, III 0.35–0.42 mm, IV 0.34–0.35 mm. Pronotum: Length 0.37–0.40 mm, basal width 0.56–0.58 mm. Similar to macropters in overall color pattern, but like the macropterous female, differing from males in the shorter antennal segment and darkened femora.

Similar in color to the macropterous males and the one macropterous female, but differing especially in the nearly quadrate pronotum that has only weakly flared posterior margins and the brachypterous hemelytron with a greatly shortened cuneus (fracture still visible on most specimens) and a much abbreviated membrane lacking any trace of areoles (Fig. 23).

No specific host. Specimens from Arizona were swept from a dry grass and flower meadow in eastern Arizona; the one male from Utah was beaten from the bases of bunch grasses.

Described and previously known only from Montana. Arizona and Utah represent new state records.

Although I have considered the series of 18 females collected in Arizona by Leonard Kelton (CNC) conspecific with the holotype male and one other male taken in northern Utah, I note that all femora of this southern population are uniformly dark except at the bases, whereas the Montana and Utah males have pale front and middle femora and only the hind femora dark. In addition, the actual measurements of the antennal segments are considerably smaller in these females than males, a variation not observed in other species.

Holotype ♂ (macropterous) (00162198) (USNM): UNITED STATES: Montana: Granite Co.: Drummond, July 10, 1935, Oman.

UNITED STATES: Arizona: Apache Co.: Big Lake, Apache National Forest, 33.88667°N, 109.41667°W, 2774 m, 12 Aug 1967 - 14 Aug 1967, L. A. Kelton, 18 ♀♀ (00112047 - 00112061, 00112063, 00161925 - 00161926) (CNC). Utah: Cache Co.: 0.4 mi off Rt 89 along Franklin Basin Rd, 41.93555°N, 111.57222°W, 2004 m, 17 Jul 2001 - 18 Jul 2001, T. J. Henry and A. G. Wheeler, Jr., 1 ♂ (00161897) (USNM).

Tytthus mundulus is distinguished by the fuscous to black head, pronotum, and scutellum; the translucent hemelytra with the clavus, most of the corium, and membrane tinged with brown; and the yellow legs and antennal segment I (apex sometimes narrowly infuscated), with contrasting fuscous to black antennal segments II–IV. Antennal segment II has short erect setae along the entire dorsal and ventral surface (most evident ventrally), similar to those found on in Tytthus chinensis and Tytthus parviceps; Tytthus mundulus, however, lacks the fuscous knee spots found in these two species. All known specimens of this species are macropterous.

Male (n =5) (Fig. 24): Length to apex of hemelytron 3.07–3.26 mm, length to base of cuneus 2.21–2.30 mm, width across hemelytra 0.93–1.01 mm. Head: Length 0.34–0.35 mm, width across eyes 0.70–0.72 mm, interocular width 0.34–0.35 mm. Labium: Length 1.04–1.15 mm. Antenna: Segment I length 0.38–0.40 mm, II 1.23–1.26 mm, III 0.66–0.67 mm, IV 0.54–0.56 mm. Pronotum: Length 0.38–0.40 mm, basal width 0.83–0.86 mm.

Coloration: Head: Uniformly black, with an indistinct, pale yellow, interocular spot near inner margin of each eye; eyes dark reddish brown. Labium: Pale yellow, with only apical half of segment IV brown. Antenna: Segment I pale yellow, sometimes tinged with pale brownish orange, apex sometimes narrowly dark brown; segments II–IV dark brown to fuscous. Pronotum: Uniformly shiny, black. Mesoscutum: Shiny black. Scutellum: Shiny black. Hemelytron: Clavus, inner two thirds of corium and most of membrane tinged with brown to dark brown, leaving only outer margin of corium clear. Ostiolar evaporative area: Dark brown to fuscous. Ventral surface: Uniformly dark brown to fuscous. Legs: Coxae yellow, with bases dark brown to fuscous; femora, tibiae, tarsi, and claws uniformly yellow.

Structure, texture, and vestiture: Head: Shiny, impunctate, with a glaucous patch along inner margin of each eye; wider than long; buccula relatively broad, ending near level with posterior margin of eye; sparsely set with short, recumbent setae, more so on glaucous patches, and with a few longer, erect setae along posterior margin. Labium: Extending to bases of mesocoxae; segment I extending beyond base of head to anterior edge of prosternum. Antennae: Segment I sparsely set with short, fine, recumbent setae and two erect, subapical, bristlelike setae; segment II thickly set dorsally and ventrally with short, erect and semierect, somewhat bristlelike setae, forming a “bottlebrush” appearance (similar to Tytthus chinensis). Pronotum: Shiny, impunctate; calli weakly swollen; anterior angles rounded; lateral margins weakly concave, moderately flaring at posterior angles; posterior margin sinuate; thickly set with semierect and recumbent setae. Mesoscutum: Shiny, impunctate, broadly exposed; with semierect and recumbent setae. Scutellum: Shiny, impunctate, equilateral; thickly set with semierect and recumbent setae. Hemelytron: Macropterous, subparallel when paired, with fully developed cuneus and membrane, extending well beyond apex of abdomen.

Male genitalia: Left paramere (Fig. 152): Mitt-shaped; right arm longest, gradually tapering to a point; left arm short, apically acute. Right paramere (Fig. 153): Elongate oval. Endosoma (Fig. 154): Strongly C-shaped. Phallotheca (Fig. 155): Relatively slender, apically acute.

Female (n = 5) (Fig. 25): Length to apex of hemelytron 3.07–3.64 mm, length to base of cuneus 2.27–2.66 mm, width across hemelytra 1.04–1.22 mm. Head: Length 0.35–0.37 mm, width across eyes 0.74–0.77 mm, interocular width 0.35–0.37 mm. Labium: Length 1.06–1.22 mm. Antenna: Segment I length 0.34–0.38 mm, II 0.98–1.10 mm, III 0.58–0.59 mm, IV 0.50–0.58 mm. Pronotum: Length 0.40–0.43 mm, basal width 0.91–1.01 mm.

This species is an egg predator of the sugarcane delphacid, Perkinsiella sacharicida Kirkaldy (Zimmerman1948, Wheeler 2001). It has also been taken on corn and taro in Hawaii.

This Indo-Pacific species has been reported from Fiji, Hawaii, New Caledonia, Papua New Guinea, the Philippine Islands, and Queensland, Australia (Schuh 1984). It has been successfully introduced into Hawaii to control the sugarcane delphacid (Zimmerman 1948, Carvalho and Southwood 1955), representing one of the best examples of successful classical biological control (Wheeler 2001). It was released in Florida, according to Nguyen et al. (1984), but apparently it has not become established (Wheeler 2001). It was also introduced into South Africa to control a tropiduchid, Numicia viridis Muir, on sugarcane but without success (Carnegie and Harris 1969). See Wheeler (2001) for additional information about this beneficial species.

AUSTRALIA: Queensland: North Queensland Co.: Halifax Apr. 1920, F. Muir, 1 ♀ (BMNH). Upper Mulgrave River, 8 miles from Goldsborough Road, 09 May 1967, D. H. Colless, 1 nymph (00161946) (USNM). FIJI: Natova, Apr. 1919, R. Veitch, 1 ♂, 5♀♀ (BMNH). MAURITIUS: Locality unknown, 20.2°S, 57.5°E, 02 May 1965, J.R. Williams, 2 ♂♂ (00138689, 00138743) (AMNH). PAPUA NEW GUINEA: Eastern Highlands: No. 11, Arau, Kratke Mountains, Valley of the upper Wanton River, 12 Oct 1959, L. J. Brass, 1 ♂ (00138756) (AMNH). Morobe Province: Lae at head of Huon Gulf near mouth of Markham River, camp #1, 6.723°S, 146.991°E, 2 m, 07 Oct 1959 - 19 Oct 1959, L. J. Brass, 1 ♀ (00095355) (AMNH). South High Province: Upper Mendi, Hoai Village, 1970 m, 4-5 Nov. 1981, B. M. Thistleton, from sugarcane, 2 ♂♂, 1 ♀ (BMNH). PHILIPPINES: Luzon: Los Banos, 21 Nov 1921, F. X. Williams, sugarcane, 1 ♀ (00161942) (USNM). UNITED STATES: Hawaii: Honolulu Co.: Honolulu, Mar 1930, F.C. Hadden, 2 ♂♂ (00167072, 00167073) (CNC). Intercepted on ship at Honolulu port which originated from Treasure Island, California, 08 Mar 1939, R. G. Oakley, 1 ♂ (00162001) (USNM). Waialua, 22 Apr 1925, F. Muir, 1 nymph (00161928) (USNM); 22 Jan 1930, O. H. Swezey, 1 ♀ (00161919) (USNM). Waipio, 02 May 1923, O. H. Swezey, 1♀ (00161918) (USNM). Maui Co.: 2191 S. Kihei Road, 29 Feb 1984, G. M. Stonedahl, at mercury vapor light, 30 ♂♂ (00138660, 00138662, 00138666, 00138668, 00138672 - 00138673, 00138682, 00138693, 00138711, 00138745, 00138749, 00138754, 00138757, 00138760, 00138787, 00165832 - 00165846), 8 ♀♀ (00165847 - 00165854) (AMNH), 2 ♂♂ (00161944, 00161945) (USNM). Oahu Co.: Barber’s Point, Mar 1960, E. J. Ford, Jr., at light, 1 ♂ (00161947), 4 ♂ (00161952 - 00161954, 00161958) (USNM). Ewa Beach, 22 Nov 1982, collector unknown, 2 ♂♂ (00161949, 00161957), 2 ♀♀ (00161955, 00161956) (USNM). Honolulu, 21.3069°N, 157.8583°W, 35 m, 24 Feb 1943, N. L. H. Krauss, on corn plants, 1 ♀ (00161940) (USNM). John Rodgers [airport], May 1958, E. J. Ford, Jr., corn, 1 ♂ (00161941) (USNM). Kailua, 09 Oct 1930, O. H. Swezey, 1 ♂ (00161920) (USNM); 15 Sep 1950, T.H., feeding on corn leafhopper eggs, 1 ♂ (00161943) (USNM). Mokuleia, 05 Feb 1940, T. H., on green corn, 2 ♂♂ (00161950, 00161951), 1 ♀ (00161948) (USNM). Punloa, 21 Dec 1926, O. H. Swezey, 1 ♂ (00161922) (USNM). Waianae, 13 Jan 1931, O. H. Swezey, Taro, 1 ♀ (00161921) (USNM).

This species is readily distinguished from all other species of Tytthus by the dark brown to fuscous head, pronotum, and scutellum; pale translucent hemelytra; pale yellow legs; and especially the pale first antennal segment having a broad, dark band through the middle. No other species has a broad band on antennal segment I with the base and apex pale. Males and females of this species are always macropterous.

As noted in the diagnosis of Tytthus juturnaiba, a photograph of the holotype stored in the PBI Heteroptera Species Database contradicts the color of antennal segments I and II described in the original description. If my interpretation of the banded antennal segment I in the photograph is correct, the two species probably are conspecific. A final decision, however, must await examination of the holotype of Tytthus juturnaiba.

Male (n = 10) (26, 63, 64): Length to apex of hemelytron 2.40–2.60 mm, length to base of cuneus 1.68–1.80 mm, width across hemelytra 0.82–0.83 mm. Head: Length 0.27–0.29 mm, width across eyes 0.54–0.59 mm, interocular width 0.27–0.29 mm. Labium: Length 0.94–1.07 mm. Antenna: Segment I length 0.24–0.29 mm, II 0.75–0.96 mm, III 0.37–0.51 mm, IV 0.32–0.34 mm. Pronotum: Length 0.32–0.34 mm, basal width 0.74–0.77 mm.

Coloration: Head (Figs 65–67: Uniformly black, with an indistinct, pale yellow, interocular spot near inner margin of each eye; eyes reddish brown. Labium: Pale yellow, with apical half of segment IV brown. Antenna: Segment I pale or whitish on apical and basal fourth, with a broad, uniformly dark brown to fuscous band through middle and a very narrow dark brown ring at base; segments II–IV uniformly dark brown to fuscous. Pronotum: Uniformly dark brown to fuscous. Mesoscutum: Uniformly dark brown to fuscous. Scutellum: Uniformly dark brown to fuscous. Hemelytron: Translucent, highlighted or tinged with pale brown on clavus and inner half of corium, inner half of clavus along claval commissure accented with darker brown; translucent dusky brown. Ostiolar evaporative area (Fig. 68): Dark reddish brown. Ventral surface: Thorax and abdomen uniformly dark reddish brown to fuscous. Legs: Coxae pale yellow, with bases reddish brown; femora pale yellowish, with metafemur sometimes accented with pale orange; tibiae, tarsi, and claws (Fig. 70) pale yellow.

Structure, texture, and vestiture: Head: Shiny, impunctate, wider than long; buccula narrow, tapering posteriorly, ending near hind margin of eye; set with scattered, relatively long, semierect setae. Labium: Extending just beyond metacoxae to base of abdomen; segment I extending past base of head to middle of xyphyus before procoxae. Antenna: Segment I with short, relatively sparse, recumbent setae and two or three erect, subapical, bristlelike setae; segment II evenly set with short, recumbent setae. Pronotum: Shiny, impunctate; calli weakly swollen, entire area covered with a glaucous sheen; anterior angles rounded; lateral margins concave, strongly flaring at posterior angles; posterior margin weakly sinuate; evenly set with recumbent and semierect setae, especially on disc. Mesoscutum: Broadly exposed; set with scattered semierect setae. Scutellum: Impunctate, equilateral; set with scattered, relatively long, semierect setae. Hemelytron: Macropterous, cuneus and membrane fully developed, extending well beyond apex of abdomen; evenly set with recumbent setae.

Male genitalia (Fig. 69): Left paramere (Fig. 156): Mitt-shaped; right arm long and broad, tapering to a point apically; left arm shorter, pointed. Right paramere (Fig. 157): Elongate oval. Endosoma (Fig. 158): Strongly C-shaped. Phallotheca (Fig. 159): Relatively slender, apically acute.

Female (n = 10) (Fig. 27): Length to apex of hemelytron 2.66–3.01 mm, length to base of cuneus 2.02–2.18 mm, width across hemelytra 0.96–1.04 mm. Head: Length 0.29–0.32 mm, width across eyes 0.56–0.61 mm, interocular width 0.27–0.32 mm. Labium: Length 1.06–1.17 mm. Antenna: Segment I length 0.26–0.27 mm, II 0.70–0.86 mm, III 0.45–0.54 mm, IV 0.43–0.45 mm. Pronotum: Length 0.32–0.35 mm, basal width 0.80–0.86 mm.

Similar to males in overall appearance, differing primarily in the slightly broader body form.

No specific host known. Associated with grass savannas and marshes and pine-oak sand scrub.

This species was described from Brazil (Carvalho 1954) and later reported from Ecuador, Peru, Puerto Rico, Surinam (Schuh 1995), and Cuba (Hernández and Henry 2010). New country records are Argentina, Belize, Bolivia, Brazil, Guyana, Haiti, Mexico (Veracruz), Nicaragua, Panama, Paraguay, the United States (Florida), and Venezuela.

ARGENTINA: Misiones: Parque Nacional de Iguazú, 25.61666°S, 54.33333°W, Dec 1975, collector unknown, 1 ♀ (00095357) (AMNH). BELIZE: Toledo: Punta Gorda, 16.1°N, 88.8°W, Mar 1931, J.J. White, 1 ♀ (00161324) (USNM). BOLIVIA: El Beni: 40 km E. San Borja, Estacion Biologica Beni, Estancia El Porvenir, 06 Sep 1987 - 08 Sep 1987, W. E. Steiner, at black light trap in open grass savanna and marsh, 4 ♂♂ (00161989 - 00161990, 00161994, 00161997) (USNM). Capivara on Rio Itenez, approx. 20 km E. of Versalles, 22 Jul 1964, J. K. Bouseman, J. Lussenhop, 1 ♂ (00165932) (AMNH). Sara: Locality unknown, 1700, Steinbach, 1 ♂ (00138669) (USNM). BRAZIL: Amazonas: Estirao do Equador, AM, 4.52378°S, 71.56085°W, Oct 1979, Alvarenga, 1 ♂ (00161385) (USNM). Ceara: Barbalha, 7.3167°S, 39.2833°W, May 1969, M. Alvarenga, 1 ♂ (00161384) (USNM). Minas Gerais: Vicosa, 20.75°S, 42.8833°W, 653 m, 13 Oct 1985 - 01 Nov 1985, T. J. Henry and P. S. F. Ferreira, 15 ♂♂ (00161893, 00161959 - 00161965, 00161968 - 00161972, 00161978, 00162006), 18 ♀♀ (00161892, 00161966 - 00161967, 00161973 - 00161977, 00161979 - 00161988) (USNM). Para: Belem Co.: 8 km E Belem, Ananindeua, 12 Jun 1973, R. T. Schuh, 1 ♀ (00138665) (USNM). Rio Guama, 28 Aug 1973, R. T. Schuh, 2 ♂♂ (00138658, 00138734) (USNM). Jacareacanga, 6.2667°S, 57.65°W, 88 m, Jun 1970, F. R. Barbosa, 1 ♂ (00161386) (USNM). Parana: Antonia, Reserva Sapitanduva, 05 Oct 1986, Lev. Ent. ProfauPar. lampada, 1 ♀ (00161380) (USNM). Rio de Janeiro: Conceicao de Macabu, Aug 1977, M. Alvarenga, 1 ♀ (00138738) (USNM); Sep 1978, Alvarenga, 2 ♂♂ (00161322, 00161323) (USNM). Est do Rio, 1947, J.C.M. Carvalho, 3 ♀♀ (00161991 - 00161993) (USNM). Estrada Rio-Sao Paulo, Km 47, 22.8669°S, 43.7789°W, 12 Nov 1943, O. Braga, 1 ♂ (00174909) (MNRJ). Guaratiba, 22.9667°S, 42.8°W, Nov 1941, J. C. M. Carvalho, 1 ♂ (00161379) (USNM). Rondonia: 62 km SW of Ariquemes, near Fzda. Rancho Grande, 10.32921°S, 63.46881°W, 30 Mar 1992 - 10 Apr 1992, J. E. Eger, 1 ♂ (00161338), 2 ♀♀ (00161339, 00161340) (USNM). Santa Catarina: Nova Teutonia, 27.18333°S, 52.38333°W, 642 m, Dec 1944, F. Plaumann, 3 ♀♀ (00161381 - 00161383) (USNM); 03 Aug 1951, F. Plaumann, 1 ♀ (00161336) (USNM). São Paulo: São Paulo Co.: km 47, Estr. Rio, 1950, J. Maldonado C., 1 ♂ (00161390) (USNM). CUBA: Havana: Habana-Alamar-Cojimer, 210 m, 10 Aug 1966 - 24 Aug 1966, Jar. Prokop, 1 ♂ (00161320) (USNM). Mantanzas: Varadero, 23.1536°N, 81.2514°W, 31 Jan 1965, Jar. Prokop, 1 ♀ (00161321) (USNM). HAITI: Ouest: Port-au-Prince, 18.5392°N, 72.335°W, 99 m, Jul 1961, J. Maldonado C., 1 ♂ (00161998) (USNM). MEXICO: Veracruz: Los Tuxtlas area, Rio Maquinas, “Los Tuxtlas” Biological Station, 31 km NE of Catemaco, 04 May 1981 - 14 May 1981, C. M. & O. S. Flint, Jr., 1 ♀ (00161389) (USNM). Los Tuxtlas area, Rio Palma, below La Palma, “Los Tuxtlas” Biological Station, 31 km NE of Catemaco, 05 May 1981, C. M. & O. S. Flint, Jr., 1 ♀ (00161388) (USNM). Veracruz, 19.18333°N, 96.11666°W, 1 m, Dec 1961, N. L. H. Krauss, 1 ♀ (00161343) (USNM). NICARAGUA: Managua: Managua, 12.15083°N, 86.26833°W, Baker, 1 ♂ (00161378), 1 ♀ (00161595) (USNM). PANAMA: Canal Zone: Playa Venado, 30 Jul 1975 - 31 Jul 1975, E. M. & J. L. Fisher, 1 ♂ (00138763) (UCR). Cocle: Agua Dulce, 1951, Blanton, 2 ♂♂ (00161328, 00161329) (USNM); 07 Aug 1951, Blanton, 2 ♂♂ (00161326, 00161327), 1 ♀ (00161325) (USNM). Aguadulce, 07 Aug 1951, Blanton, 4 ♂♂ (00161353 - 00161356) (USNM); 25 Sep 1951, F. S. Blanton, 1 ♀ ( 00161344) (USNM). Puerto Obaldia, 04 Nov 1952, collector unknown, 1 ♀ (00161609) (USNM). Rio Hato, 8.3833°N, 80.1667°W, 03 Aug 1953, F. S. Blanton, 1 ♀ (00161319) (USNM). Darien: El Real, 8.1333°N, 77.7167°W, 17 m, 19 Mar 1953, F. S. Blanton, 2 ♀♀ (00161995, 00161996) (USNM). Garachine, 8.06472°N, 78.36277°W, 18 Feb 1953, F. S. Blanton, 1 ♀ (00161999) (USNM). Panama: Las Cumbres, 9.06°N, 79.32°W, 113 m, 28 Jul 1971, M. Daykin, 1 ♂ (00138698) (UCD). Tocumen, 9.0833°N, 79.3833°W, 25 Mar 1952, F. S. Blanton, 2 ♀♀ (00161341, 00162000) (USNM); 17 Nov 1952, F. S. Blanton, 1 ♀ (00161924) (USNM); 04 Dec 1952, F. S. Blanton, light trap, 1 ♀ (00161923) (USNM); 05 Oct 1953, F. S. Blanton, 1 ♀ (00161342) (USNM). PARAGUAY: Alto Parana: Reserva Biol. Tati Yupi, 14 Nov 1990, G. Arriagada, 2 ♂♂ (00161345, 00161346), 6 ♀♀ (00161347 - 00161352) (USNM). Tarija: Gran Chaco Co.: 260 kilometers west of the Paraguay River, 13 Jul 1935, Alberto Schulze, 1 ♂ (00161927) (USNM). PERU: La Libertad: Trujillo, near mouth of Rio Moche, 02 Jul 1972, R.T. and J.C. Schuh, 1 ♀ (00138706) (USNM). Lima: Callao, 12.03333°S, 77.13333°W, 3 m, 17 Nov 1950, Michelbacher and Ross, 1 ♀ (00161611) (USNM). Loreto: Km 3 Tournavista Rd., 34 km W Pucallpa, 8.48333°S, 74.8°W, 300 m, 13 Dec 1971 - 31 Dec 1971, R. T. & J. C. Schuh, light trap, 63 ♂♂ (00138655 - 00138657, 00138659, 00138671, 00138674 - 00138675, 00138678, 00138680, 00138683, 00138687, 00138691, 00138694, 00138696 - 00138697, 00138699, 00138702, 00138705, 00138712, 00138718, 00138721 - 00138728, 00138730 - 00138733, 00138737, 00138739 - 00138740, 00138744, 00138746 - 00138747, 00138750, 00138752 - 00138753, 00138758 - 00138759, 00138761 - 00138762, 00138764 - 00138767, 00138770 - 00138775, 00138778 - 00138779, 00138782 - 00138786, 00138788), 26 ♀♀ (00138661, 00138663, 00138667, 00138670, 00138676, 00138681, 00138685 - 00138686, 00138688, 00138695, 00138700 - 00138701, 00138704, 00138707 - 00138709, 00138714 - 00138715, 00138717, 00138720, 00138735, 00138751, 00138769, 00138777, 00138780, 00161318) (USNM). Lake Yarinacocha, 10 km NW of Pucallpa, 150 m, 08 Dec 1971 - 10 Dec 1971, R. T. Schuh, light trap, 2 ♀♀ (00138736, 00138741) (USNM); 08 Dec 1971, R. T. Schuh, light trap, 1 ♂ (00138692) (USNM). Madre de Dios: Tambopata Co.: Rio Tambopata Reserve, 30 air km SW Pto. Maldonado, 290 m, 26 Nov 1979 - 30 Nov 1979, J. B. Heppner, subtropical moist forest, 1 ♀ (00161337) (USNM). PUERTO RICO: Caguas: Caguas, May 1965, Ricardo Jorge, 2 ♂♂ (00161361, 00161362) (USNM). El Verde, Jun 1967, J. Maldonado C., 1 ♂ (00161360), 1 ♀ (00161359) (USNM). Cayey: Cayey, Jul 1961, J. Maldonado C., 1 ♂ (00161363), 3 ♀♀ (00138713, 00161364 - 00161365), 1 nymph (00161366) (USNM). Comerio: Comerio, 05 Jun 1961, J. Maldonado C., 1 ♂ (00161376) (USNM). Guanica: Locality unknown Co.: Lake Guanica, 23 Jul 1936, H. L. Dozier, 1 ♀ (00161335) (USNM). Guayanilla: Guayanilla, Sep 1960 - Nov 1960, E. Murphy, 2 ♀♀ (00161370, 00161371) (USNM). Maricao: Maricao, Jul 1960, J. Maldonado C., 1 ♂ (00161375) (USNM). Mayaguez: Sabanetas Co.: Mani Beach, 04 Aug 1935, H. L. Dozier, 1 ♀ (00161334) (USNM). Mayaguez, Mar 1960, J. Maldonado C., 1 ♀ (00161372) (USNM); Jun 1962, J. Maldonado C., 3 ♂♂ (00161367 - 00161369) (USNM); Jul 1975, J. Maldonado C., 1 ♀ (00161373) (USNM). Mayaguez, Dec 1964, R. Ricardo, 1 ♂ (00161357), 1 ♀ (00161358) (USNM). Rio Grande: El Yunque, Apr 1967 - Jun 1967, J. Maldonado C., 1 ♀ (00161374) (USNM). San Juan: San Juan, 18.46633°N, 66.10573°W, 02 Aug 1914 - 03 Aug 1914, collector unknown, 1 ♀ (00161377) (USNM). Cayey, 18.113°N, 66.166°W, 431 m, 1961, Julio, 1 ♂ (00095356) (AMNH). UNITED STATES: Florida: Highlands Co.: Archbold Biological Station, S of Lake Placid, hill area E. of Station, 27.18333°N, 81.34166°W, 01 Jan 2007, W.E. Steiner and J.M. Swearingen, at black light in pine-oak sand scrub, 1 ♀ (00161391) (USNM). VENEZUELA: Aragua: Maracay Co.: Maracay, 10 Jul 1968, J. Maldonado C., 1 ♀ (00161387) (USNM). Maracay, 10.24694°N, 67.59583°W, 548 m, 10 Jul 1968, J. Maldonado C., 1 ♂ (00161433) (USNM). Guanare: Estado Portuguesa, 10 Sep 1957 - 13 Sep 1957, Borys Malkin, 2 ♂♂ (00161330, 00161331), 2 ♀♀ (00161332, 00161333) (USNM).

This species is recognized by the combination of the dark brown head, pronotum, and scutellum; the pale antennal segment I, with only the base fuscous, the brown antennal segment II; and the uniformly pale yellow legs. All known specimens are macropterous.

Tytthus pallidus keys out with Tytthus piceus based on the pale antennal segment I and the pale hemelytra with smoky-brown shading. It can be distinguished from Tytthus piceus by the broader head, the longer antennal segment I that is longer than the interocular width, and the less prominent calli lacking a glaucous sheen.

Holotype male (Fig. 28): Length to apex of hemelytron 2.88 mm, length to base of cuneus 2.21 mm, width across hemelytra 0.88 mm. Head: Length 0.35 mm, width across eyes 0.61 mm, interocular width 0.29 mm. Labium: Length 1.31 mm. Antenna: Segment I length 0.35 mm, II 1.22 mm, III and IV missing. Pronotum: Length 0.35 mm, basal width 0.75 mm.

Coloration: Head: Uniformly dark brown; pale interocular spot found in all other species apparently absent; fuscous to dark reddish brown. Labium: Pale brownish yellow. Antenna: Segment I uniformly pale yellow, with a dark brown or fuscous ring at base; segments II–IV dark brown. Pronotum: Uniformly dark brown. Mesoscutum: Uniformly dark brown. Scutellum: Uniformly dark brown. Hemelyton: Uniformly pale translucent brown. Ostiolar evaporative area: Dark brown to fuscous. Ventral surface: Thorax dark brown to dark reddish brown; abdomen dark reddish brown on segment II, III, and genital capsule, slightly paler in between. Legs: Coxa pale yellow, reddish brown at bases; femora, tibiae, tarsi, and claws uniformly pale yellow.

Structure, texture, and vestiture: Antenna: Segment I set with a few, scattered, recumbent setae and two erect, subapical, bristlelike setae. Labium: Extending beyond metacoxae to abdominal segment II or III. Pronotum: Shiny, impunctate; anterior angles rounded; lateral margins weakly concave, flaring at posterior angles; posterior margin distinctly sinuate; calli weakly swollen; set with relatively long, semierect setae. Mesoscutum: Impunctate, broadly exposed; with a few scattered semierect setae. Scutellum: Weakly shining, impunctate; equilateral; set with scattered, semierect setae. Hemelytra: Macropterous, cuneus and membrane fully developed, extending well beyond apex of abdomen.

Male genitalia: Left paramere (Fig. 160): Mitt-shaped; right arm long, broad, apically blunt; left arm short, apically acute. Right paramere (Fig. 161): Round. Endosoma: Teneral and damaged; not drawn. Phallotheca (Fig. 162): slender, apically acute.

Female (n = 4) (Fig. 29): Length to apex of hemelytron 2.69–3.33 mm, length to base of cuneus 2.05–2.40 mm, width across hemelytra 0.82–1.02 mm. Head: Length 0.32–0.38 mm, width across eyes 0.56–0.64 mm, interocular width 0.29–0.30 mm. Labium: Length 1.12–1.44 mm. Antenna: Segment I length 0.26–0.34 mm, II 0.85–1.23 mm, III 0.61–0.78 mm, IV 0.51–0.56 mm. Pronotum: Length 0.29–0.37 mm, basal width 0.69–0.85 mm.

The specific epithet “pallidus” refers to the characteristic pale yellow first antennal segment.

Unknown.

Known from Brazil and Panama.

Holotype ♂ (00162199) (USNM): PANAMA: Darien: [Pinogana District], El Real, 19 Mar. 1953, F. S. Blanton, 1 ♂ (USNM). El Real, 08 Aug 1952, F. S. Blanton, 2 ♀♀ (00161392, 00161393) (USNM). Paratypes: BRAZIL: Amazonas: Reserva Ducke, 25 km NNE of Manaus, 120 m, 26 Jul 1973, R.T. Schuh, 2 ♀♀ (00165829) (AMNH).

Tytthus panamensis is a small species distinguished by the combination of the fuscous head, pronotum, and scutellum; the fuscous antennal segment I, with only the apex pale; the pale yellowish brown antennal segment II; and the uniformly pale legs. All known specimens are macropterous.

It is most similar to Tytthus pallidus, sp. n. in overall size and color, including the reduced or apparent lack of an interocular spot on the head and the uniformly pale legs. It differs in having a dark brown antennal segment I, with only the apex pale, and a pale brown antennal segment II, whereas Tytthus pallidus has a pale antennal segment I, with only a narrow dark ring at the base, and segment II is dark brown. Tytthus panamensis keys near Tytthus vagus based on the dark antennal segment I with only the apex pale, but can be distinguished by the uniformly pale yellowish brown antennal segment II and hind femur.

Male (n =1, plus holotype in parentheses) (Fig. 30): Length to apex of hemelytron 2.21 mm (2.37 mm), length to base of cuneus 1.57 mm (1.63 mm), width across hemelytra 2.21 mm (2.37 mm). Head: Length 0.26 mm (0.29 mm), width across eyes 0.48 mm (0.51 mm), interocular width 0.24 mm (0.26 mm). Labium: Length 0.85 mm (0.93 mm). Antenna: Segment I length 0.30 mm (0.35 mm), II 0.88 mm (0.99 mm), III missing (0.56 mm), IV missing (missing). Pronotum: Length 0.22 mm (0.22 mm), basal width 0.58 mm (0.66 mm).

Coloration: Head: Uniformly dark brown, pale interocular spot found in all other species (except Tytthus pallidus) apparently absent; eyes dark brown to fuscous. Labium: Uniformly pale yellow. Antenna: Segment I dark brown, narrowly pale at apex; segment II pale yellowish brown; segments III and IV brown. Pronotum: Uniformly dark brown. Mesoscutum: Uniformly dark brown. Scutellum: Uniformly dark brown. Hemelytron: Uniformly pale translucent brown. Ostiolar evaporative area: Reddish brown. Ventral surface: Thorax and abdomen uniformly dark reddish brown. Legs: Procoxae pale yellow; meso- and metacoxae pale yellow, with bases reddish brown; femora, tibiae, tarsi, and claws uniformly pale yellow.

Structure, texture, and vestiture: Head: Shiny, impunctate; buccula slender, tapering posteriorly, ending near level with middle of eye. Labium: Extending to apices of meso- or bases of metacoxae; segment I extending beyond base of head to anterior edge of xyphus before procoxae. Antenna: Segment I with relatively sparse, short, recumbent setae and two, long, subpical, bristlelike setae; segment II thickly set with short, recumbent setae, intermixed with a few more scattered, semierect setae. Pronotum: Shiny, impunctate; anterior angles weakly rounded; lateral margins straight, gradually widening to posterior angles; basal margin weakly sinuate; set with scattered recumbent and semierect setae. Mesoscutum: Shiny, impunctate, broadly exposed; set with a few short, semierect setae. Scutellum: Shiny, impunctate, equilateral; set with a few scattered, short, recumbent and semierect setae. Hemelytron: Macropterous, cuneus and membrane fully developed, extending posteriorly well beyond apex of abdomen; evenly set with relatively long, recumbent setae.

Male genitalia: Not examined. See note below.

Female (n = 1) (Fig. 31): Length to apex of hemelytron 2.91mm, length to base of cuneus 1.98 mm, width across hemelytra 0.80 mm. Head: Length 0.34 mm, width across eyes 0.59 mm, interocular width 0.29 mm. Labium: Length 1.20 mm. Antenna: Missing. Pronotum: Length 0.27 mm, basal width 0.74 mm.

Similar to male in overall appearance, differing primarily in the broader body form.

Unknown.

Described and known only from Panama.

The one male paratype previously had been dissected and the genitalia apparently were placed in a glass genitalia vial sealed with a cork stopper. Since that dissection, the cork has dried and crumbled and the genitalia are missing from the vial. I now have placed the glass vial inside a polyethylene vial, sealed it with a neoprene stopper, and reattached it to the specimen pin.

Holotype ♂ (00162200) (USNM): PANAMA: Canal Zone: Corozal, 8.98844°N, 79.5719°W, 14 Apr 1912, A. Busck, at light. Paratypes: Same data as for holotype, 1 ♂ (00161395) (USNM), 1 ♀ (00161394) (USNM).

PANAMA: Canal Zone:. Fort Gulick, 9.31667°N, 79.86667°W, 21 Aug 1952, F. S. Blanton, 1 ♀ (00162201) (USNM). Darien: Garachine, 8.06472°N, 78.36277°W, 17 Feb 1953, F. S. Blanton, 1 ♀ (00161396) (USNM).

This widespread circumtropical species is best distinguished by the pale yellow anterior area of the pronotum on almost all specimens, a characteristic found only in some specimens of Tytthus chinensis and certain color forms of Tytthus pygmaeus. It is also recognized by the combination of the black head, with large, yellow interocular spots; black antennal segment I, with the apex and base of segment I narrowly pale, yellowish-brown to brown segment II with the base black (but often uniformly fuscous or black); black scutellum, pale translucent hemelytra; and uniformly pale yellow legs, with a fuscous knee spot on each tibia. Tytthus parviceps is readily distinguished from Tytthus pymaeus by the knee spots at the bases of the tibiae. From Tytthus chinensis it is distinguished by the usually extensively yellow anterior third to one half of the pronotum and the more strongly C-shaped (Fig. 165) endosoma. Only macropterous individuals of this species are known.

Male (n = 10) (Fig. 32): Length to apex of hemelytron 2.11–2.46 mm, length to base of cuneus 1.50–1.70 mm, width across hemelytra 0.74–0.82 mm. Head: Length 0.26–0.27 mm, width across eyes 0.53–0.56 mm, interocular width 0.27–0.29 mm. Labium: Length 0.82–0.86 mm. Antenna: Segment I length 0.24–0.26 mm, II 0.72–0.80 mm, III 0.42–0.48 mm, IV 0.37–0.40 mm. Pronotum: Length 0.27–0.30 mm, basal width 0.67–0.74 mm.

Coloration: Head: Uniformly fuscous to black, with a large, yellow, interocular spot touching inner margin of each eye, spots nearly contiguous in some individuals; eyes fuscous to dark reddish brown. Labium: Uniformly pale yellow, except brown apical half of segment IV. Antenna: Segment I black, with the apex pale yellow; segment II yellowish brown to brown, with base black but sometimes uniformly black; segments III–IV uniformly fuscous to black. Pronotum: Anterior half typically broadly yellow from margin to margin, sometimes yellow area reduced so lateral margins and area between calli become invaded with fuscous, to the extreme with yellow greatly reduced or absent; posterior half uniformly fuscous to black. Mesoscutum: Uniformly yellowish brown to fuscous. Scutellum: Uniformly fuscous to black. Hemelytron: Uniformly translucent yellow. Ostiolar evaporative area: Uniformly yellow to yellow with central area of auricle invaded by fuscous. Ventral surface: Anterior half of proacetabula yellow, propleura, pro- and mesosterna black; metapleura yellowish, invaded by fuscous; abdomen largely yellowish, with only genital capsule fuscous or black. Legs: Uniformly yellow, with only bases of tibiae fuscous.

Structure, texture, and vestiture: Head: Weakly shiny, impunctate; buccula slender, extending posteriorly, ending near level with hind margin of eye; thickly set with short to relatively long semierect setae, especially on frons. Labium: Extending to apices of meso- or bases of metacoxae; segment I extending beyond base of head to xyphyus just before procoxae. Antenna: Segment I set with short, recumbent setae and two, long, subapical, bristlelike setae; segment II thickly set with short, recumbent setae, intermixed with row of longer, erect setae (similar to Tytthus chinensis) along ventral surface. Pronotum: Anterior angles rounded; lateral margins weakly concave, gradually widening to rounded posterior angles; posterior margin weakly sinuate. Mesoscutum: Weakly shiny, impunctate; set with a few scattered, semierect setae. Scutellum: Weakly shiny, impunctate; equilateral; set with a few scattered recumbent and semierect setae. Hemelytron: Macropterous, cuneus and membrane fully developed, extending posteriorly well beyond apex of abdomen; evenly set with relatively long, recumbent setae.

Male genitalia: Left paramere (Fig. 163): Mitt-shaped; right arm long, broad, almost triangular, bluntly pointed apically; left arm relatively long, apically pointed. Right paramere (Fig. 164): Oval. Endosoma (Fig. 165): Strongly C-shaped, blunt apically. Phallotheca (Fig. 166): Slender, apically acute.

Female (n = 10) (Figs 33, 34): Length to apex of hemelytron 2.40–3.04 mm, length to base of cuneus 1.79–2.21 mm, width across hemelytra 1.02–1.09 mm. Head: Length 0.29–0.30 mm, width across eyes 0.59–0.64 mm, interocular width 0.30–0.32 mm. Labium: Length 0.88–1.04 mm. Antenna: Segment I length 0.26–0.29 mm, II 0.69–0.86 mm, III 0.43–0.56 mm, IV 0.43–0.50 mm. Pronotum: Length 0.34–0.38 mm, basal width 0.86–0.98 mm.

Similar to male, differing primarily in the larger size and broader body form.

In the New World, Tytthus parviceps has been associated with the delphacid Saccarosydne saccharivora (Westwood) (Carvalho and Southwood 1955) and the sugarcane delphacid, Perkinsiella saccaricida Kirkaldy (Sosa 1985), on sugarcane in Florida and Ecuador. It also has been taken on Spartina alterniflora Loisel [Poaceae] in North Carolina (Wheeler and Henry 1992). In the Old World, it is known to prey on eggs of the brown plant hopper, Nilaparvata lugens (Stål), in India and sugarcane delphacid in Australia (Bull 1981).

In the New World, this widespread circumtropical and subtropical species is known from Central and South America (see specimen data below), the West Indies, Bermuda, and Florida and North Carolina in the United States (Henry and Hilburn 1990, Wheeler and Henry 1992). In the Old World, it has been reported from southern Europe, the Middle East, Africa, eastern (Far-Eastern Russia, Korea) and southeastern Asia, Australia, India, and islands in the Indian, Pacific, and southern Atlantic oceans (Wheeler and Henry 1992, Kerzhner and Josifov 1999). Cassis and Gross (1995) omitted this species from the Australian list, suggesting that reports in the literature should be referred to the similar-appearing Tytthus chinensis.

Tytthus parviceps, described from Egypt, is similar to Tytthus chinensis in nearly all external characters, except for the strongly pale yellow anterior area around the calli of the pronotum, which may cause considerable confusion between these species when individuals of Tytthus parviceps with greatly reduced yellow markings or an entirely dark pronotum are encountered, or when specimens of Tytthus chinensis with more extensive yellow markings are found. Dissection of representative males of Tytthus parviceps from different regions (list localities) shows that the endosoma is consistently S-shaped, whereas specimens of Tytthus chinensis from (list localities) have a C-shaped endosoma.

China (1924) synonymized Cylloceps pellicia Uhler with Cyrtorhinus parviceps after examining the “type-specimen” deposited in the Natural History Museum (BMNH). I have examined the female syntype of Cyrtorhinus parviceps from Egypt and a female syntype of Cylloceps pellicia from St. Vincent in the Natural History Museum, as well as a male syntype of Capsus pellicia from Cuba (labeled in Uhler’s hand) in the USNM collection, and agree that Uhler’s species is a junior synonym of Tytthus parviceps as now defined.

Also, Josifov and Kerzhner (1972), in describing Tytthus koreanus, initially indicated that it was similar to Tytthus parviceps (Reuter) based on the extensive pale areas on the pronotum. However, they also added that the new species might be conspecific with Tytthus chinensis described from Hong Kong, which according to the original description (Stål 1860) and to Reuter (1903) is a larger species (3.00 mm) than Tytthus parviceps. Unfortunately, the type of Tytthus chinensis is lost and the current concept of this species was established by Carvalho and Southwood (1955) and Carvalho (1956) as smaller, with the pronotum entirely black. I have examined a large amount of material from Australia and the Indo-Pacific Region that contains a few individuals among mostly black specimens exhibiting a yellow pronotal color not too unlike that described by Josifov and Kerzhner (1972) for Tytthus koreanus. In addition, I have studied three specimens from Korea that also have more yellow on the anterior area of the pronotum but less than for typical specimens of Tytthus parviceps. As a consequence, I am treating Tytthus koreanus as a junior synonym of Tytthus chinensis (which see) until additional studies can show otherwise.

To ensure nomenclatural stability, I am selecting the following syntypes as lectotypes:

Cyrtorhinus parviceps ♀ (BMNH): label 1 (circular label with orange ring) “Type”; label 2, “Le Cairo, V. 86, E. Antian, Schweinfor”; label 3 “Cyrtorhinus parviceps Reut. type; label 4 (here added, red printed label), “LECTOTYPE: ♀, Cyrtorhinus parviceps Uhler, by T. J. Henry”; label 4 “00085519 (BMNH).

Cylloceps pellicia: ♂ (USNM): label 1, “Cuba”; label 2 (handwritten), “152”; label 3, “PR Uhler Collection”; label 4 (handwritten), “Cylloceps pellicia Cuba: Uhler”; label 5 (handwritten, except “Det. Uhler”), “Cylloceps pellicia Uhler Cuba [printed] Det Uhler”; label 6 (name handwritten; remainder printed), “Tytthus parviceps (Rt.) det JCMCarvalho)”; label 7 (here added, printed red label), “LECTOTYPE: ♂, Cylloceps pellicia Uhler, by T. J. Henry.” A female from “St. Vincent, May, H. H. Smith” in the BMNH with 00085520 (BMNH) is here designated as a paralectotype.

ANTIGUA AND BARBUDA: Antigua: Galleon Beach, 03 Aug 1992, H.V. & R.M. Baranowski, 2 ♀♀ (00161495, 00161496) (USNM). BAHAMAS: Andros Island: Fresh Creek, 24.7°N, 77.76666°W, 23 Apr 1953, E.B. Hayden, 1 ♂ (00165901) (plus 10 ♂♂, 3 ♀♀) (AMNH). Mangrove Cay, 24.28333°N, 77.66666°W, 26 Apr 1953, E.B. Hayden, 12 ♂♂ (00165889 - 00165900), 6 ♀♀ (00165861 - 00165866) (AMNH). Bimini: South Bimini Island, 25.70623°N, 79.28204°W, 6 m, Jun 1951, Cazier and C. & P. Vaurie, 1 ♀ (00165913) (AMNH). Exuma Cays: Staniard Cay, 24.18333°N, 76.41666°W, 13 Jan 1953, E. B. Hayden & L. Giovannoli, 9 ♂♂ (00165876 - 00165884), 7 ♀♀ (00165867 - 00165873) (AMNH). Nassau: New Providence Co.: New Providence Island, 03 Jan 1953, E.B. Hayden, 1 ♀ (00165874) (AMNH). San Salvador Island: nr. Cockburn Town, 24.0333°N, 74.5167°W, 18 Mar 1953, L. Giovannoli & G.B. Rabb, 4 ♂♂ (00165885 - 00165888), 1 ♂ (00165875) (AMNH). BARBADOS: H. H. Smith, 1 ♂ (BMNH). BELIZE: Belize: Big Falls Ranch, 23 Dec 1975, R. Akers, Oryza sativa (Poaceae), 1 ♂ (00161546) (USNM). BERMUDA: Devonshire Parish: Devonshire Marsh, 13 Jan 1988, T. J. Henry, 1 ♂ ( 00161503) (USNM); 14 Jul 1988 - 22 Jul 1988, M.R. Wilson & D.J. Hilburn, 1 ♂ (00161502) (USNM). Paget Parish: Paget Marsh, 23 Jul 1971, N. L. H. Krauss, 1 ♂ (00161505) (USNM); 13 Jan 1988, T. J. Henry, 1 ♀ (00161504) (USNM); 14 Jul 1988 - 22 Jul 1998, M.R. Wilson & D.J. Hilburn, 15 ♂♂ (00161468 - 00161482), 3 ♀♀ (00161483 - 00161485) (USNM) (an additional 10 ♂♂, 5 ♀♀ in BMNH). BRAZIL: Para: Santarem Co.: Taperinha, 11 Jun 1927 - 20 Jun 1927, Zerny, 1 ♀ (00161614) (USNM). BURKINA FASO: Kadiogo: Upper Volta, Ouagadougou, 03 Nov 1973 - 05 Nov 1973, R. Linnavuori, 1 ♂ (00166054), 3 ♀♀ (00166055 - 00166057) (AMNH). CAMEROON: Locality unknown, 1983, Irct A. Renou, 3 ♂♂ (00161707 - 00161709), 4 ♀♀ (00161705 - 00161706, 00161767 - 00161768), 2 nymphs (00161710, 00161711) (USNM). CAYMAN ISLANDS: Grand Cayman: Grand Cayman, 05 Aug 1988, P. Fitzgerald, 1 nymph (00161587) (USNM). CHAD: Farcha, 20 May 1973 - 22 May 1973, R. Linnavuori, 1 ♀ (00166052) (AMNH). CHILE: Vicuna: El Pangue, Nov 1961, L.E. Peña, 2 ♂♂ (00161464, 00161465), 1 ♀ (00161466) (USNM). COSTA RICA: Guanacaste: 10 mi NW of Liberia, 10.723°N, 85.344°W, 25 Jul 1965, Paul J. Spangler, 1 ♀ (00161586) (USNM). CUBA: Cienfuegos: La Jiquima, 29 Jul 1956, J. M. Osorio, 2 ♂♂ (00161519, 00161520), 6 ♀♀ (00161521 - 00161526) (USNM). La Habana: Alamar-Cojimar, 10 m, 04 May 1966 - 05 May 1966, Jar. Prokop, 1 ♂ (00161463) (USNM). Habana-Marianao, 15 m, 15 Jun 1966 - 20 Jul 1966, F. Gregor, 1 ♂ (00161462) (USNM). Pinar del Rio: La Jiquima, 27 Jul 1956, J. Acuna, 5 ♂♂ (00161729 - 00161730, 00161539, 00161729 - 00161730), 6 ♀♀ (00161540 - 00161544, 00161728) (USNM). DEMOCRATIC REPUBLIC OF THE CONGO: Haut-Katanga: Lubumbashi, 10 Apr 1971 - 11 Apr 1971, R. Linnavuori, 1 ♂ (00166024) (AMNH); 25 Apr 1971 - 26 Apr 1971, R. Linnavuori, 1 ♂ (00166023) (AMNH). Nord-Kivu: Lac Edouard, 15 Jan 1936, Dr. H. Damas, 1 ♂ (00161766) (USNM). DOMINICAN REPUBLIC: La Altagracia: Nisibon, 03 May 1978, Woodruff & Fairchild, 1 ♀ (00161584) (USNM). Nisibon, Finca Papagayo, 46 m, 04 Apr 2000 - 07 Apr 2000, T. J. Henry and R. E. Woodruff, 5 ♂♂ (00161486 - 00161490), 3 ♀♀ (00161491 - 00161493) (USNM). Pendernales: Cabo Rojo, Alcoa Headquarters, 13 Jun 1988, R. E. Woodruff, 1 ♀ (00161583), 1 ♀ (00161582) (USNM). Puerto Plata: Rio Camu, 19 km NE of Jarabacoa, 12 Jun 1969, Flint & Gomez, 1 ♀ (00161591) (USNM). San Juan: Locality unknown, 18.88°N, 71.27°W, 19 Oct 1967, collector unknown, 1 ♀ (00161550) (USNM). San Pedro de Macoris: Santo Domingo, 8 mi up Macoris river, 16 Jul 1917, Harold Morrison, 3 ♂♂ (00161536 - 00161538) (USNM). ECUADOR: El Oro: Victoria-Arenillas, 150 m, 18 Aug 1977 - 19 Aug 1977, L. Pena G., 1 ♂ (00161580) (USNM). Guayas: Ingenio San Carlos, 29 Nov 1995, Walter Gordillo, 1 ♂ (00161547), 2 ♀♀ (00161548, 00161549) (USNM). Imbabura: Otavalo, 01 Jan 1970, R. Levi-Castillo, 1 ♂ (00161531), 2 ♀♀ (00161530, 00161610) (USNM). Pichincha: Ingenio San Carlos, 07 May 1982, Robert Morey, 1 ♂ (00161721), 1 ♀ (00161722) (USNM). EGYPT: 6th of October Governorate: W. Desert, Bahariya Oasis, 12 May 1995, Dr. Ullrich, 1 ♂ (00161648) (USNM); 30 Apr 1996, Dr. Ullrich, 2 ♂♂ (00161651, 00161652), 2 ♀♀ (00161649, 00161650) (USNM). Aswan: Aswan, 24.0875°N, 32.8989°E, 25 Dec 1993, Dr. Ullrich, 2 ♂♂ (00161572, 00161573), 1 ♀ (00161571) (USNM). Cairo Governorate: 25 km SW of Cairo, Wadi Digla, 23 Jan 1998, Dr. Ullrich, 1 ♂ (00161575), 1 ♀ (00161576) (USNM). Cairo, Digla, Road to Wadi Digla, 29.95861°N, 31.32041°E, 01 Feb 1998, Dr. Ullrich, 4 ♂♂ (00161657 - 00161660), 3 ♀♀ (00161661 - 00161663) (USNM); 11 Feb 1998, Dr. Ullrich, 1 ♂ (00161577), 1 ♀ (00161616) (USNM). Dahshur, ca. 23 km S. Cairo, 29.79908°N, 31.24625°E, 29 May 1996, Dr. Ullrich, 12 ♂♂ (00161617 - 00161624, 00161632 - 00161633, 00161645 - 00161646), 17 ♀♀ (00161625 - 00161630, 00161634 - 00161644), 1 nymph (00161631) (USNM). El-Marg, 02 Apr 1912, Alfieri, 1 ♂ (00161654) (USNM). Maadi, 05 May 1931, Alfieri, 1 ♂ (00161655) (USNM); 29 Aug 1992, Dr. Ullrich, 1 ♀ (00161647) (USNM); 28 May 1996, Dr. Ullrich, 1 ♀ (00161653) (USNM). Meadi, 12 June 1961, R. Linnavuori, 1 ♂ (BMNH). Umgeb, Cairo, 1-19 June 1961, R. Linnavuori, 1 ♀ (BMNH). Fayum Governorate: Fayum, 13-14 June 1961, R. Linnavuori, 2 ♀♀ (BMNH). Qahirah: Cairo (‘Le Caire, ’ ‘Al Qahirah’), 30.05°N, 31.25°E, May 1888, Schweinfurth, syntype, 1 ♀ (00085519) (BMNH). Meadi, 29.9667°N, 31.25°E, 20 m, 06 May 1931, H. Priesner, 1 ♂ (00235057) (ZISP). EL SALVADOR: La Libertad: La Libertad, 10 m, 29 Oct 1965, N. L. H. Krauss, 1 ♀ (00161589) (USNM). GHANA: Accra, 27 Sep 1943 - 29 Sep 1943, M.A. Locke, 1 ♂ (00161780), 3 ♀♀ (00161781 - 00161783) (USNM). GRENADA: Balthazar, Windward side, 07 Aug 1963, O. S. Flint, 1 ♀ (00161588) (USNM). GUATEMALA: San Marcos: 17.3 km SE of Talisman, Rio Cabuz at Hwy. CA2, 14.85°N, 92.06666°W, 200 m, 23 May 1973, G. F. Hevel, 1 ♀ (00161585) (USNM). GUYANA: Mibicury, 26 Feb 1998, Srinivasan, on “paddy plant hopper, ” 8 ♂♂, 18 ♀♀ (BMNH). Demerara (?), Sahr, C. B. Williams, 1910, 1 ♀ (BMNH). HAITI: Gonave Id., 21 Jul 1931, B. M. 1931-448, 2 ♂♂, 1 ♀ (BMNH). INDIA: Manipur: Delhi, 01 Jan 1950, P.K. Pathek, 2 ♂♂ (00161701, 00161702), 2 ♀♀ (00161703, 00161704) (USNM). Orrisa: Cuttack, CRRI, 1980, “predatory on brown planthopper, ” 3 ♂♂, 1 ♀ (BMNH). IRAN (ISLAMIC REPUBLIC OF): Hormozgan: Rodan, 27 Mar 2001 - 29 Mar 2001, R. & S. Linnavuori, 3 ♂♂ (00161675 - 00161677), 1 ♀ (00161678) (USNM). Khuzestan: Dezful Research Station, garden, 30 Oct 1971 - 10 Nov 1971, L. Knutson, 1 ♂ (00161656) (USNM). Hormozgan Garband, 23-25 Feb. 2001, 26 Mar., 4 Apr. 2001, R. Linnavuori, 13 ♂♂, 11 ♀♀ (NMW). Harmozagan Oeshm Ramkan, 14-15 May 2002, R. Linnavuori, 1 ♂ (NMW). ISRAEL: Mehoz HaZafon (Northern region): Dan, 33.21666°N, 35.65°E, 07 Jul 1958, R. Linnavuori, 1 ♂ (00235058) (ZISP). Deganya, 32.7°N, 35.56666°E, 23 Jul 1958, R. Linnavuori, 1 ♀ (00235173) (ZISP).N. Distr., Bet-Savoa Res., 8-9 Aug. 1986, R. Linnavuori, 1 ♂, 3 ♀♀ (NMW). N. Distr., Dan, 11 June 1986, R. Linnavuori, 1 ♀ (NMW). S. Distr. Melot hakikkar, 16-20 Jul. 1986, R. Linnavuori, 1 ♂, 1 ♀ (NMW). S. Distr., Nahal ‘Aruoot, 22 Jul. 1986, R. Linnavuori, 9 ♂♂, 6 ♀♀ (NMW). Yad Hashmone, 10 June 1986, Linnavuori, 1 ♀; S. Distr., Be’er Shev’, June 1986, R. Linnavuori, 2 ♀♀ (NMW). S. Distr., Revivim, 4 June 1986, R. Linnavuori, 1 ♀, (NMW). S. Distr., ‘En Gedi, 28 Apr. 1986, R. Linnavuori, 3 ♂♂, 7 ♀♀ (NMW). S. Distr., ‘Ein Avdat, 10 Aug. 1986, R. Linnavuori, 1 ♀ (NMW). S. Distr., ‘En Besor, 6 Jul. 1986, R. Linnavuori, 8 ♂♂, 3 ♀♀ (NMW). S. Distr., Be’er Sheva’, May 1986, R. Linnavuori, 1 ♂, 2 ♀♀ (NMW). S. Distr., Na’ot hakikkar, 16-20 Jul. 1986, R. Linnavuori, 1 ♂ (NMW). S. Distr., Eliias nr Imma, 23 June 1986, R. Linnavuori, 1 ♂, 1 ♀ (NMW). IVORY COAST: Gouméré, 19 Sept. 1973, R. Linnavuori, 3 ♂♂, 6 ♀♀ (NMW). Foro Foro, 25-28 Sept. 1973, R. Linnavuori, 6 ♂♂, 8 ♀♀ (NMW). Lamto, 8-9 Oct. 1973, R. Linnavuori, 1♀ (NMW); Adiopodoumé, 29 Sept.-7 Oct. 1973, R. Linnavuori, 1 ♀ (NMW). Man, 14-21 Oct. 1973, R. Linnavuori, 2 ♂♂, 2 ♀♀ (NMW). JAMAICA: Clarendon Parish: Monymusk Estate, Jul 1959, F. D. Bennett, (Poaceae), 1 ♀ (00161532) (USNM); 10 Dec 1959, F. D. Bennett, (Poaceae), 1 ♂ (00161535), 2 ♀♀ (00161533, 00161534) (USNM). J. Faradane, B. M. 1970-455, 1 ♂, 3 ♀♀ (BMNH). MAURITIUS: Bel Etan, 3 Oct 1957, J. R. Williams, on sugarcane, 3 ♂♂, 1 ♀ (BMNH). MEXICO: Tamaulipas: Tampico, 22.2167°N, 97.85°W, 2 m, 29 Dec 1908, collector unknown, 1 ♂ (00161402) (USNM). Yucatan: Progreso, 21.28°N, 89.67°W, 01 Apr 1992, J. R. Vockeroth, 1 ♂ (00112062), 1;f (00166921) (CNC). NICARAGUA: Granada: Santa Lastenia, Malacatoya, 08 Nov 1971, Ev. Vogel, 4 ♀♀ (00161513 - 00161516) (USNM). Locality unknown, Nov 1989, J. M. Maes, 10 ♂♂ (00161596 - 00161605), 2 ♀♀ (00161606, 00161607) (USNM). NIGERIA: Lagos: Ikoyi, 25 Oct 1975, J. Riley, 2 ♀♀ (BMNH). NE St. Yola, 25 Aug. 1973, R. Linnavuori, 2 ♂♂, 4 ♀♀ (NMW). Tahoua- In Waggeur, 12-13 Nov. 1973, R. Linnavuori, 4 ♂♂, 2 ♀♀ (NMW). Kw. St. nr. Sinau, 4 Sept. 1975, R. Linnavuori, 1 ♂, 2 ♀♀ (NMW). N. C. St., Malumfashi, 26-30 Jul. 1973, R. Linnavuori, 3 ♂♂, 9 ♀♀ (NMW). W. St. Ife, 7-8 Jul., 14 Aug. 1973, R. Linnavuori, 1 ♀. NW St. Badeggi, 8-9 Aug. 1973, R. Linnavuori, 1 ♂ (NMW). Oyo State, Ibadan, INTA 1994, C. T. Williams & S. Okhidie, from filtering irrigated lowland rice, 1 ♂ (NMW). Niamey, 9 Nov. 1973, R. Linnavuori (NMW), 1 ♂, 5 ♀♀. Ile-Ife, 5 Aug. 1969, J. T. Medler, 2 ♂♂, 1 ♀ (NMW). N. Bussa K state, 12 Jan. 1970, J. T. Medler, 1 ♀ (NMW). B. Pl. St., nr. Makurdi, 30 Aug. 1973, R. Linnavuori, 3 ♂♂, 3 ♀♀ (NMW). Kwara: Shaganu Biological Station, 20 Jul 1973 - 22 Jul 1973, R. Linnavuori, 1 ♂ (00166053) (AMNH). NE State: Serti, 20 Aug 1975, R. Linnavuori, 1 ♂ (00161733), 1 nymph (00161734) (USNM). Yola, 25 Aug 1973, R. Linnavuori, 1 ♂ (00161731), 1 ♀ (00161732) (USNM). Oyo: Ibadan, Oct 1982, C.I.E., on rice, 1 ♂, 1 ♀ (BMNH). OMAN: Wadi Ghul, 1450 m, 23E14'N, 57E09'E, 1 Nov. 1990, M. D. Gallagher & J. C. Deeming, 1 ♂ (NMW). DhoPar, Ain Hamran, streamside vegetation, 10 Oct. 1990, J. C. Deeming, 2 ♂♂ (NMW). PALESTINE TERRITORY[?]: West Bank: Beit-el-Ghofr, N of Haz, about 16 mi NW of San’a, ca 9, 300 ft, 4 Feb. 1938, from a lucerne field, 1 ♂ (BMNH). PANAMA: Barro Colorado Island: Canal Zone, 19 Oct 1971, D. Engleman, 1 ♀ (00165989) (AMNH). Canal Zone: Barro Colorado Island, 9.16667°N, 79.85°W, 08 Aug 1972, D. Engleman, light trap, 1 ♂ (00165958) (AMNH). Fort Amador, Feb 1964, Ch. Keenan, 1 ♀ (00161467) (USNM). Playa Venado, 30 Jul 1975 - 31 Jul 1975, E. M. ♀ J. L. Fisher, 1 ♂ (00161497) (USNM). Cocle: Agua Dulce, 07 Aug 1951, Blanton, 1 ♀ (00161581) (USNM). El Chiru, 10 Nov 1952, F. S. Blanton, 1 ♂ (00161593) (USNM). Herrera: Port Chitre, 24 Oct 1952, F. S. Blanton, 1 ♂ (00161719), 1 ♀ (00161720) (USNM). PERU: Lima: Barranca, 01 Jan 1900, Sauer, 1 ♂ (00161507) (USNM); 16 Sep 1940, collector unknown, 2 ♀♀ (00161608, 00161612) (USNM). PHILIPPINES: Bulacan: Malolos, 29 May 1972, A. D. Pawar, 1 ♀ (00166922) (CNC). Laguna: Los Banos, International Rice Research Institute, 10 Jul 1972, A. D. Pawar, Oryza sp. (Poaceae), 1 ♂ (00112064) (CNC). Mountain: Sagada, 09 Apr 1972, A. D. Pawar, 1 ♀ (00166923) (CNC). PUERTO RICO: Arecibo: Arecibo, 30 Jul 1914 - 01 Aug 1914, collector unknown, 1 ♀ (00165914) (AMNH). Caguas: Caguas, May 1965, Ricardo Jorge, 2 ♂♂ (00161528, 00161712), 3 ♀♀ (00161529, 00161713 - 00161714) (USNM). Cayey: Cayey, Sep 1960 - Nov 1960, M. M. Beauchamp, 1 ♂ (00161718), 1 nymph (00161717) (USNM); Sep 1960, M. Santiago, 2 ♂♂ (00161725, 00161726) (USNM). Comerio: Comerio, 05 Jun 1961, J. Maldonado C., 1 ♂ (00161727) (USNM). Guanica: State Forest, 30 Jun 1955, J. A. Ramos, J. Maldonado, 1 nymph (00161461) (USNM). Guayanilla: Guayanilla, Sep 1960 - Nov 1960, E. Murphy, 2 ♀♀ (00161715, 00161716) (USNM). Lajas: Lajas, 18.0519°N, 67.0597°W, 47 m, 01 May 1954 - 20 May 1954, J. Maldonado Capriles, 1 ♂ (00161592) (USNM); Sep 1960, J. Maldonado C., 1 ♂ (00161724) (USNM). Mayaguez: Guanajibo, 03 Aug 1935, H. L. Dozier, 1 ♀ (00161527) (USNM). Mayaguez, Mar 1960, J. Maldonado C., 1 ♂ (00161723) (USNM); Jul 1975, J. Maldonado C., 2 ♀♀ (00161451, 00161452) (USNM). San Juan: Rio Piedras, 18.3994°N, 66.0503°W, 29 m, 01 Jan 1972, G. F. and S. Hevel, 1 ♂ (00161590) (USNM). San Juan, 18.46633°N, 66.10573°W, 18 Feb 1964, L.T. Sanders, 1 ♀ (00161545) (USNM). Manati, 18.429°N, 66.492°W, 53 m, 27 Jun 1915 - 29 Jun 1915, collector unknown, 1 ♂ (00095358) (AMNH). REUNION: Savannah, 17 Jan 1990, B. Vercambre, 1 ♀ (00161553) (USNM); 09 Feb 1990, B. Vercambre, 1 ♂ (00161551), 1 ♀ (00161552) (USNM). RODRIGUES ISLAND: Aug-Nov 1918, H. J. Snell and H. P. Thomasset, 1 ♂, 1 ♀ (BMNH). SAINT LUCIA: Castries, 0-210 meters, Aug 1976, N. L. H. Krauss, 1 ♂ (00165986) (AMNH). ST. HELENA: Diana’s Peak, 27 June 1959, . R. Wallace, swept from grass, 1 ♀. Picquet Post. 27 Feb 1936, H. F. D. Barlett, 1 ♂ (BMNH). SAUDI ARABIA: Ash Sharqiyah: Al Hasa, Oct 1977, R. Linnavuori, 3 ♂♂ (00161697 - 00161699), 1 ♀ (00161700) (USNM). Al Hasa, 10 Nov 1917, R. Linnavuori, 4 ♂♂ (00165919 - 00165922), 2 ♀♀ (00165923, 00165924) (AMNH); Oct 1977, R. Linnavuori, 1 ♂ (00165925), 3 ♀♀ (`00165926 - 00165928) (AMNH); 05 Dec 1977, R. Linnavuori, 1 ♂ (00165931), 2 ♀♀ (00165929, 00165930) (AMNH); 02 Oct 1978, R. Linnavuori, 4 ♂♂ (00165915 - 00165918) (AMNH). El Riyadh, 24.63333°N, 46.71666°E, 31 Mar 1959 - 05 Apr 1959, E. W. Diehl, 1 ♀ (00235174) (ZISP). SENEGAL: Dakar: Dakar, 25 Oct 1943, J. Phillips, 2 ♂♂ (00161777, 00161778), 1♀ (00161779) (USNM). SIERRA LEON: Njala, 21-23 Nov. 1981, L. H. Rolston, 2 ♂♂, 3 ♀♀ (USNM). SOMALIA: Sar Uanle, 28 May 1973, S.B.S., 1 ♂ (00165999) (AMNH); 08 Jun 1973, S.B.S., 1 ♂ (00166022), 1 ♀ (00166040) (AMNH); 11 Jun 1973, S.B.S., 1 ♀ (00166025) (AMNH); 11 Jun 1973, R. Linnavuori, 1 ♀ (00166041) (AMNH). SOUTH AFRICA: Cape Province: Rondvlei near Knysna, 08 Feb 1968, T. Schuh, J. A. & S. Slater, M. Sweet, 1 ♂ (00165906) (AMNH). Wilderness, 12 Mar 1968 - 13 Mar 1968, P. J. Spangler, 1 ♂ (00161771), 2 ♀♀ (00161769, 00161770) (USNM). Gauteng: Lyttelton, 25.833°S, 28.216°E, 1492 m, 12 Jan 1968, J. A. & S. Slater, 1 ♀ (00165912) (AMNH); 29 Feb 1968, J. and S. Slater, 4 ♂♂ (00165902 - 00165905), 1 ♀ (00095359) (AMNH). KwaZulu-Natal: Port-Shepstone, 5.97, 1926, 1 ♀ (BMNH). Northern Cape: McDougall Bay, Port Nolloth, 29.28796°S, 16.87945°E, 2 m, 06 Sep 2004, Schuh, Schwartz, Henry, Wyniger, Forero, Sporobolus virginicus (L.) Kunth (Poaceae), det. J Manning VOUCHER-NYBG, 17 ♂♂ (00161735 - 00161751), 19 ♀♀ (00161752 - 00161765, 00161772 - 00161776) (USNM). SPAIN: Canary Islands: Grand Canary Island, Maspalomas, 14 Aug 1966, K. M. Guichad & P. H. Ward, 9 ♂♂, 6 ♀♀ (BMNH). SUDAN: Bahr El-Jabal: Equatoria, Juba, 27 Feb 1963 - 02 Mar 1963, R. Linnavuori, 5 ♂♂ (00166002 - 00166005, 00166013), 1 ♀ (00166042) (AMNH). Equatoria, Juba-Nimule, 10 Mar 1963 - 11 Mar 1963, R. Linnavuori, 1 ♂ (00166016) (AMNH). Equatoria, Lalyo-Juba, 26 Feb 1963 - 27 Feb 1963, R. Linnavuori, 1 ♀ (00166049) (AMNH). Equatoria, Loka Forest, 08 Apr 1963 - 10 Apr 1963, R. Linnavuori, 1 ♀ (00166050) (AMNH). Equatoria, Nimule, 11 Mar 1963 - 13 Mar 1963, R. Linnavuori, 1 ♂ (00166015) (AMNH). Central Equatoria, Juba, 27 Feb-2 Mar 1963, R. Linnavuori, 2 ♂♂, 2 ♀♀ (BMNH). Blue Nile: Singa-Damazin, 15 Nov 1962 - 17 Nov 1962, R. Linnavuori, 9 ♂♂ (00165975 - 00165983), 5 ♀♀ (00166030 - 00166034) (AMNH). Umm Banein, 14 Nov 1962, R. Linnavuori, 5 ♂♂ (00166007 - 00166011), 1 ♀ (00166035) (AMNH). Wad Madani [Medani], 14.4°N, 33.5°E, 11 Nov 1962 - 12 Nov 1962, R. Linnavuori, 1 ♂ (00166012), 2 ♀♀ (00166044, 00166045) (AMNH). Wad es Zaki, 10 May 1963, R. Linnavuori, 1 ♂ (00166014) (AMNH). Kassala: Kassala, 30 Nov 1962, R. Linnavuori, 1 ♀ (00166043) (AMNH). Khartoum: Khartoum, 13 Aug 1966, Stys, 1 ♂ (00161569), 1 ♀ (00161570) (USNM). Nile: Ed Damer, 05 Jul 1961 - 10 Jul 1961, R. Linnavuori, 2 ♂♂ (00166019, 00166020), 1 ♀ (00166047) (AMNH). Shendi, 02 Nov 1962 - 05 Nov 1962, R. Linnavuori, 1 ♀ (00166048) (AMNH). Southern Kordofan: Lake Keilak, 08 Feb 1963 - 11 Feb 1963, R. Linnavuori, 1 ♂ (00166018) (AMNH). Upper Nile, Malakal, 05 Jan 1963 - 20 Jan 1963, R. Linnavuori, 2 ♂♂ (00165984, 00166006) (AMNH). Upper Nile: Malakal, 05 Jan 1963 - 20 Jan 1963, R. Linnavuori, 3 ♀♀ (00166026 - 00166028) (AMNH). Warab: Bahr el Ghazal, 19 Feb 1963, R. Linnavuori, 1 ♂ (00166021) (AMNH). White Nile: Blue Nile, Kosti, 22 Jan 1963, R. Linnavuori, 2 ♂♂ (00166000, 00166001) (AMNH). Kordofan, Kadugli, 02 Feb 1963 - 14 Feb 1963, R. Linnavuori, 1 ♂ (00166017), 1 ♀ (00166029) (AMNH). APO 625, AMM #A857, 30 Oct 1943, collector unknown, 1 ♂ (00161784) (USNM). TAIWAN: Nantou: Ren-ai Township Co.: Huei-Sun For. Rec. Area, 5km NE of Meiyuan, 24.0667°N, 120.9833°E, 733 m, 10 Jul 1992 - 11 Jul 1992, T. J. Henry and A. G Wheeler, Jr., 1 ♀ (00161420) (USNM). TANZANIA: Ukiriguru, 18 May 1961, I. A. D. Robertson, ex light trap, 1 ♂, 5 ♀♀ (AMNH). TRINIDAD AND TOBAGO: Arima: Curepa, 11 Aug 1975, F. D. Bennett, 1 ♀ (00165988) (AMNH). Port-of-Spain: Port-of-Spain, 10.65°N, 61.5167°W, 09 Sep 1969, H.A. Wright, 1 ♀ (00161494) (USNM). UNITED STATES: Florida: Alachua Co.: Gainesville, 29.63527°N, 82.37111°W, 24 m, 29 Dec 1964, F.W. Mead, Medicago sativa (Fabaceae), 3 ♂♂ (00161670 - 00161672) (USNM); 25 Oct 1982, R. Nguyen, Saccharum officinarum (Poaceae), 1 ♀ (00161666) (USNM). Brevard Co.: Orsino, 13 Aug 1951, collector unknown, 1 ♂♂ (00161594) (USNM). Broward Co.: Deerfield Beach, 26 Jul 1948, R. H. Beamer, 1 ♀ (00165991) (AMNH). W. Park, 29.66806°N, 82.40194°W, 18 Aug 1939, collector unknown, 2 ♀♀ (00161400, 00161401) (USNM). Hendry Co.: Locality unknown, 31 Aug 1982, D. G. Hall, 1 ♀ (00161421) (USNM). Highlands Co.: Lake Placid, 27.29278°N, 81.36306°W, 13 Jul 1948, R. H. Beamer, 1 ♀ (00165992) (AMNH), 1 ♂ (00161673) (USNM). Sebring, 27.49555°N, 81.44083°W, 41 m, 24 Jul 1950 - 31 Jul 1950, C. T. Parsons, 2 ♀♀ (00165966, 00165967) (AMNH); 06 Aug 1950 - 12 Aug 1950, C. T. Parsons, 9 ♂♂ (00165960 - 00165965, 00165971 - 00165973), 1 ♀ (00165974) (AMNH); 10 Aug 1950 - 30 Aug 1950, C. T. Parsons, 2 ♀♀ (00165969, 00165970) (AMNH); 01 Sep 1950 - 16 Sep 1950, C. T. Parsons, 1 ♀ (00165968) (AMNH). Indian River Co.: Fellsmere, 04 Sep 1937, H.A.J., 1 ♀ (00161664) (USNM). Sebastian, 27.81642°N, 80.47061°W, Apr 1700, G. Nelson, 1 ♂ (00165985) (AMNH). Jefferson Co.: Monticello, 30.545°N, 83.87°W, 08 Oct 1965, W.H. Whitcomb, 1 ♀ (00161665) (USNM). Miami-Dade Co.: Airport Fumigation Site, 25.79966°N, 80.30733°W, 02 Jun 2008, T. Dobbs, light trap, 1 ♀ (00161412) (USNM). Biscayne Bay, 25.5747°N, 80.3112°W, 1700, Mrs. A.T. Slosson, 2 ♂♂ (00165907, 00165908) (AMNH), 1 nymph (00161404) (USNM); 01 Jan 1900, collector unknown, 2 ♀♀ (00165994, 00165995) (AMNH), 1 ♂ (00161397) (USNM). Coconut Grove, Mar 1900, G. Fairchild, 1 ♀ (00165993) (AMNH). Everglades National Park, 25.3125°N, 80.9375°W, 19 Jul 1973, C. W. O’Brien, 1 ♂ (00165959) (AMNH). Homestead, 25.46833°N, 80.47778°W, 26 Aug 1968, R. M. Baranowski, 1 ♂ (00161417) (USNM); 08 Dec 1974, J. A. Slater, 1 ♀ (00165990) (AMNH). Miami, 25.77389°N, 80.19389°W, 1 m, 01 May 2005, APHIS port inspector, 2 ♀♀ (00161498, 00161499) (USNM); 20 Dec 2009, APHIS port inspector, 1 ♂ (00161501) (USNM). Monroe Co.: Big Pine Key, Watsons Hammock, 24.68643°N, 81.36715°W, 14 Apr 1981, T. J. Henry and A. G. Wheeler, Jr., 3 ♀♀ (00161409 - 00161411) (USNM). Orange Co.: Orlando, 28.53805°N, 81.37944°W, 34 m, 02 Jul 1918, Geo. G. Ainslie, rye, 1 ♀ (00161399) (USNM). Winter Park, 28.59972°N, 81.33944°W, 16 Aug 1944, H. T. Fernald, 1 ♂ (00165910) (AMNH). Osceola Co.: CR 532, 1 km E. of CR 545, 3.5 km NNW of Loughman, 05 Apr 2003, A. G. Wheeler, Jr., 1 ♀ (00161413) (USNM). Palm Beach Co.: 2 miles south of co. line on Rt. 27, 20 Apr 1981, T. J. Henry, 1 ♂ (00161405) (USNM). Belle Glade, 26.68417°N, 80.66778°W, 10 Jul 1963, Tullosand Brandt, 1 ♀ (00161674) (USNM). Canal Point, 07 Feb 1983, O. Sosa Jr., 2 ♀♀ (00161418, 00161419) (USNM). Lake Worth, 26.58°N, 80.04°W, 01 Jan 1750, Mrs. A.T. Slosson, 1 ♀ (00165996) (AMNH); 01 Jan 1900, collector unknown, 1 nymph (00161398) (USNM). Palm Beach, 26.70528°N, 80.03667°W, 1 m, 27 Jul 1948, R. H. Beamer, 1 ♀ (00161500) (USNM); 19 Nov 1982, D. G. Hall, 3 ♂♂ (00161667 - 00161669) (USNM); 24 Nov 1982 - 26 Nov 1982, D. G. Hall, sugarcane, 1 ♂ (00161406), 2 ♀♀ (00161407, 00161408) (USNM). Pinellas Co.: Saint Petersburg, 27.7709°N, 82.6793°W, 12 Aug 1910, J. C. Bradley, 1 ♂ (00165987) (AMNH). Polk Co.: Lakeland, 28.0392°N, 81.95°W, 02 Oct 1948, R. F. Hussey, 1 ♂ (00165909) (AMNH); 14 Nov 1948, R. F. Hussey, 3 ♂ (00161554 - 00161556) (USNM); 24 Sep 1952, R. F. Hussey, 1 ♀ (00165911) (AMNH). North Carolina: Camden Co.: North River, 17 Oct 1959, L. Davis, 1 ♂ (00161414), 2 ♀♀ (00161415, 00161416) (USNM). Virginia: Louisa Co.: 4 mi. S. of Cuckoo, 01 Jun 1985, J. Kloke & D.R. Smith, 1 ♀ (00161828) (USNM). VENEZUELA: Aragua: Maracay Co.: Maracay, 10 Jul 1968, J. Maldonado C., 7 ♂♂ (00161557 - 00161563), 5 ♀♀ (00161564 - 00161568) (USNM). Maracay, 10.24694°N, 67.59583°W, 548 m, 10 Jul 1968, J. Maldonado C., 11 ♂♂ (00161422 - 00161432), 6 ♀♀ (00161434 - 00161439) (USNM). Carabobo: Mariara, Las Vueltas, 10.2983°N, 67.7161°W, 450 m, Aug 1970, L. J. Joly, 1 ♂ (00161460) (USNM). Saman Mocho, 10.1206°N, 67.8914°W, 595 m, Aug 1970, L. J. Joly, 1 ♂ (00161440), 5 ♀♀ (00161441 - 00161445) (USNM). Saman Mocho, 10.1206°N, 67.8914°W, 450 m, Aug 1970, L. J. Joly, 1 ♂ (00161453), 7 ♀♀ (00161454 - 00161459, 00161578) (USNM). Tocuyo, Aug 1970, L. J. Joly, 2 ♂♂ (00161447, 00161448), 1 ♀ (00161446) (USNM). VIRGIN ISLANDS (U.S.): St. Thomas: 3 miles from Charlotte Amalie, 31 May 1917, Harold Morrison, 1 ♂ (00161512) (USNM). Charlotte Amalie, 18.3439°N, 64.9311°W, 02 Jun 1917, Harold Morrison, 2 ♀♀ (00161517, 00161579), 1 nymph (00161518), 1 ♂ (00161403) (USNM). YEMEN: Abyan: Al Kowd, 13.08333°N, 45.36666°E, Jan 2000, van Harten Awad, 4 ♂♂ (00161691 - 00161694), 2 ♀ (00161695, 00161696) (USNM). Ta’izz: Ta’izz, Aug 1999 - Nov 1999, van Harten Awad, 7 ♂♂ (00161679 - 00161685), 5 ♀♀ (00161686 - 00161690) (USNM). Ta’izz, Jan., Jul., Aug., Sept., & Nov. 1999, van Harten & Awad, 30 ♂♂, 18 ♀♀ (NMW). Ta’izz, May, June, Aug., & Sept. 2000, van Hartenn & Al Yarimi, 8 ♂♂, 12 ♀♀ (NMW). Ta’izz, Abkhawkhah, 4 May 1992, R. Linnavuori, 1 ♂ (NMW). Abyan, A Kowd, Jul., Sept., & Dec. 1999, van Harten, Al Harun, & Sallam, 8 ♂♂, 10 ♀♀. Abyan, A Kowd, 4 May & 8-12 July 2001, van Harten & Asi Harun, 5 ♂♂, 1 ♀ (NMW). Abyan, A Kowd, 7 Sept. 2002, van Harten & Alö Harun, 3 ♀♀ (NMW). Abyan, A Kowd, Sept. 2003, van Harten & Al Haurn, 1 ♂. A Kowd, Jan., Feb., May, June, Jul., Oct. 2000, van Harten & Sallam, 22 ♂♂, 19 ♀♀ (NMW). Hudaydah, Zabid, 19-20 Mar. 1992, R. Linnavuori, 3 ♂♂, 1 ♀ (NMW). Sana’a Sana’a, Feb.-Mar. 1992, R. Linnavuori, 3 ♂♂, 1 ♀ (NMW). Sana’a, May & 1-10 Jul. 1999, van Harten, 3 ♂, 4 ♀♀ (NMW). Lahj: Lahi, 3-5 2002, van Harten & Sallam, 1 ♂, 3 ♀♀ (NMW). Al Kadan, Oct. 2001, van Harten & Abdul-Haq, 8 ♂♂, 2 ♀♀ (NMW).

This species, known from macropterous males and females (Figs 35, 36) and brachypterous females (Fig. 37), is distinguished by the bulbous black head, the black pronotum and scutellum having a distinct glaucous sheen, the raised finely punctate pronotal calli, the pale translucent-brown hemelytra, and the pale yellow to white first antennal segment with a narrow black ring at the base.

Tytthus piceus keys out with Tytthus pallidus, n. sp but can be distinguished by the bulbous head, especially in males, the shorter antennal segment I that is only subequal to the interocular width, and the distinct calli that are covered in a glaucous sheen.

Male (n = 10; holotype in parentheses) (Figs 35, 71, 72): Length to apex of hemelytron 2.35–2.50 mm (2.70 mm), length to base of cuneus 1.70–1.75 mm (1.89 mm), width across hemelytra 0.62–0.66 mm (wings folded). Head: Length 0.34–0.37 mm (0.35 mm), width across eyes 0.51–0.54 mm (0.50 mm), interocular width 0.24–0.26 mm (0.26 mm). Labium: Length 1.07–1.14 mm (imbedded in glue). Antenna: Segment I, length 0.26–0.27 mm (antennae missing); II, 0.98–1.02 mm; III, 0.50–0.56 mm; IV, 0.40–0.50 mm. Pronotum: Length 0.32–0.34 mm (0.32 mm), basal width 0.59–0.61 mm (0.59 mm).

Coloration: Head (Figs 73–75): Shiny black; eyes black. Labium: Segment I fuscous to black, apex paler yellowish brown, tinged with red or reddish brown in some specimens; segments II–IV pale yellowish brown, apex of segment IV fuscous. Antenna: Segment I pale yellowish brown to white, with a narrow black ring at base; segments II–IV uniformly black. Pronotum, mesoscutum, and scutellum: Shiny black, with a distinct glaucous sheen. Hemelytron: Uniformly smoky brown, slightly darker brown on clavus. Ostiolar evaporative area (Fig. 76): Dark reddish brown to fuscous. Ventral surface: Thorax dark brown or fuscous; abdomen in males dark brown, sometimes paler ventrally, genital capsule fuscous to black; abdomen in females yellowish green to pale brown, with broad lateral margins and ovipositor fuscous to black. Legs: Uniformly yellowish brown, inner face of pro- and mesofemora and outer face of metafemur usually with a narrow reddish line; claw (Fig. 78).

Structure, texture, and vestiture: Head: Impunctate, round or bulbous in both sexes, slightly wider than long; with scattered, long, erect setae on frons and vertex and a few short, erect setae on eyes. Labium: Extending to abdominal segment II or III. Pronotum: Trapeziform, narrowest anteriorly, lateral margins weakly concave, flaring to humeral angles; impunctate, except for a few scattered punctures on distinctly swollen calli. Mesoscutum: Broadly exposed. Scutellum: Equilateral, weakly convex, rising just above level of hemelytra. Hemelytron: Translucent, subparallel; in macropterous males and females (Figs 35, 36) cuneus longer than wide at base, membrane fully developed, extending well beyond apex of abdomen; in brachypterous females (Fig. 37) lateral margins slightly more rounded than macropters, cuneus reduced to about as wide as long, membrane greatly reduced, extending only to the sixth or seventh abdominal tergite, exposing apex of abdomen.

Male genitalia: Left paramere (Fig. 167): Mitt-shaped; right arm long, wide, apically blunt; left arm shorter, apically pointed. Right paramere (Fig. 168): Elongate oval. Endosoma (Fig. 169): Strongly C-shaped, apically blunt. Phallotheca (Fig. 170): Relatively slender, apically acute.

Macropterous female (n = 10) (Fig. 36): Length to apex of hemelytron 2.60–2.85 mm, length to base of cuneus 1.85–2.05 mm, width across hemelytra 0.70–0.83 mm. Head: Length 0.35–0.40 mm, width across eyes 0.54–0.58 mm, interocular width 0.27–0.29 mm. Labium: Length 1.18–1.22 mm, extending to near base of ovipositor. Antenna: Segment I, length 0.26 mm; II, 0.85–0.93 mm; III, 0.51–0.53 mm; IV, 0.43–0.45 mm. Pronotum: Length 0.34–0.37 mm, basal width 0.69–0.70 mm.

Brachypterous female (n = 3) (Fig. 37): Length to apex of hemelytron 1.80–1.90 mm; length to base of cuneus 1.55–1.60 mm; length to apex of abdomen 2.20–2.40 mm; width across hemelytra 0.64–0.70 mm. Head: Length 0.35–0.40 mm, width across eyes 0.54–0.58 mm, interocular width 0.29 mm. Labium: Length 1.10–1.14 mm, extending to abdominal segment II. Antenna: Segment I, length 0.26–0.29 mm; II, 0.82–9.94 mm; III, 0.51–.56 mm; IV, 0.43–0.45 mm. Pronotum: Length 0.29–0.34 mm, basal width 0.51–0.59 mm.

Adults and nymphs have been taken in large numbers on switch grass, Panicum virgatum L. cv ‘Alamo’ [Poaceae], infested with Delphacidae (A. G. Wheeler, pers. comm.).

This species was described and known previously only from Guatemala (as Isoproba picea) and Honduras (as Tytthus hondurensis). Colombia, Costa Rica, Mexico, Panama, and the United States (Florida, Maryland, and South Carolina) represent new country records and considerable range extensions.