Revision of the Plant Bug Genus Tytthus (Hemiptera, Heteroptera, Miridae, Phylinae)

Abstract The phyline plant bug genus Tytthus Fieber, previously containing 19 species, is revised. Isoproba Osborn and Drake, 1915, incorrectly placed in the subfamily Bryocorinae, tribe Dicyphini, is synonymized as a junior synonym of Tytthus Fieber, syn. n.; the only included species, Isoproba picea Osborn and Drake is transferred to Tytthus, comb. n., as the senior synonym of Tytthus hondurensis Carvalho, syn. n.; and Tytthus koreanus Josifov and Kerzhner, 1972 is synonymized with Tytthus chinensis (Stål 1860), syn. n.; and a lectotype for Tytthus parviceps is designated. The six new species Tytthus femoralis from Cuba, Ecuador, Guatemala, Jamaica, Mexico, and Peru,Tytthus fuscicornis from New Mexico (USA), Tytthus mexicanus from Mexico, Tytthus pallidus from Brazil and Panama, Tytthus uniformis from Arizona and New Mexico (USA), and Tytthus wheeleri from the eastern United States are described, bringing the total number of species for the genus to 24. A color adult habitus illustration of Tytthus wheeleri, color photographs for each species (except Tytthus juturnaiba Carvalho and Wallerstein), illustrations of male genitalia, scanning electron photomicrographs of selected structures of certain species, and an identification key are provided to facilitate species recognition. A phylogenetic analysis is offered to help infer relationships.

Prior to this study, 19 species of Tytthus were recognized worldwide. Of these, 16 occur in the New World, including two Holarctic and one circumtropical species (Afrotropical, Neotropical, and southern Oriental). Three additional species are known from eastern and southeastern Asia, Australia, and the Indo-Pacific. Tytthus was long confused with and considered a junior synonym of the remarkably similar-appearing orthotyline genus Cyrtorhinus Fieber, 1858 (Reuter 1875c), until Carvalho and Southwood (1955) showed that the genus belonged in the subfamily Phylinae based on pretarsal structures and male genitalia. Schuh (1974) included Tytthus in his new tribe Leucophoropterini, a group of predominately African, Australian, and South-Pacific taxa, based on the hairlike parempodia, the U-shaped endosoma, the relative small size of the male genitalia, the shape of the right paramere, and the simple female posterior wall. He also noted that Tytthus was not ant mimetic as are most other leucophoropterines. Kerzhner and Josifov (1999), however, conservatively placed Leucophoropterini in synonymy under Phylini after indicating that they were following Linnavuouri (1993), who thought it premature to split the tribe. Schuh (1995), however, maintains Leucophoropterini in the online version of his world mirid catalog (http://research.amnh.org/pbi/catalog/), until more information on relationships becomes available. More recently, Menard (2011) and Menard and Schuh (2011) gave convincing molecular and morphological evidence supporting the monophyly of Leucophoropterini, but also showed that Tytthus consistently grouped outside the tribe. As a consequence, I follow Menard (2011) and Menard and Schuh (2011) in transferring Tytthus from Leucophoropterini to nominate tribe Phylini.
In this paper, I revise the genus Tytthus; give a diagnosis and new host and distribution records for previously described species; synonymize the genus Isoproba Osborn and Drake as a junior synonym of Tytthus and transfer its only included species to Tytthus as the senior synonym of T. hondurensis Carvalho; and synonymize T. koreanus Josifov and Kerzhner under T. parviceps (Reuter). Six new New World species are described; 18 previously known species are redescribed; a revised identification key is provided to help distinguish the species of the genus; and a phylogenetic analysis is presented.

Methods
The male genital capsule was dissected and placed in room-temperature, 10% KOH solution for one to two days or until softened and cleared, after which it was rinsed in water and placed in a depression slide containing glycerol. The endosoma, right and left parameres, and phallotheca were dissected and pencil sketched using a Nikon E400 compound microscrope and drawing tube. Final illustrations were digitally rendered using Adobe Photoshop CS4.
Photomicrographs were taken using either an AMRAY 1810 or a Zeiss EVO/ MA15 scanning electron microscope. Specimens were glued to standard SEM stubs, sputter coated, and examined at 6-10 KV.
Color images were captured using an EntoVision Imaging Suite that included a JVC KY-75 3CCD digital camera mounted to a Leica M16 zoom lens via a Leica zstep microscope stand. Multiple focal planes were merged using Cartograph 5.6.0 (Microvision Instruments, France) software. Plates of color habitus images (not to scale), SEM photomicrographs, and male genitalia were created using Adobe Photoshop CS4 and striped and numbered in Adobe Illustrator CS4.
Matrix code labels were attached to more than 1,000 specimens examined. These codes, referred to as unique specimen identifers (USIs), are a way to uniquely identify specimens and are stored in a database developed for the NSF Planetary Biodiversity Project awarded to R. T. Schuh (American Museum of Natural History, New York, NY) and G. Cassis (University of New South Wales, Sydney, Australia). The full code contains the prefix "AMNH_PBI, an eight-digit number, and the specimen depository, for example (AMNH_PBI 00162206) (USNM)." USI codes are included in the specimen data listed at the end of each species treatment. To save space, the "AMNH_PBI" prefix has been omitted. Data for several hundred additional specimens also were recorded, but matrix code labels were not available for these collections (e.g., BMNH, NMW) at the time the data were captured and, thus, were not entered into the database and therefore lack USI numbers.
Specimen measurements were taken as follows: Length (dorsal length from apex of clypeus to posterior margin of hemelytral membrane); length to base of cuneus (dorsal length from apex of clypeus to base of cuneus); width across hemelytra (widest dorsal width across hemelytra, usually just above each cuneus); head length (lateral length from posterior margin of pronotum to apex of clypeus); head width (dorsal width across eyes); interocular width (greatest dorsal width between eyes); labium (length from a base of labrum to apex of segment IV); antennal segment length (self explanatory); pronotum length (dorsal median length); pronotum basal width (dorsal width across posterior margin). I follow Cassis (2008) in using the term "endosoma" rather than "vesica" for the male intromittent organ. Other terminolgy follows conventional use in the contemporary literature, as defined in Nichols (1989).
The phylogenetic analysis was performed using Winclada (Nixon 1999(Nixon -2002, implementing the island hopping or the rachet function to run NONA (Goboloff 1999) and TNT (Goloboff et al. 2008) using the traditional and random addition sequence functions. All cladograms were generated using Winclada. 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.
Discussion. 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 (1923Knight ( , 1925Knight ( , 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, I. 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 (1952Carvalho ( , 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 I. 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 only in 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 T. hondurensis Carvalho (1984) is a junior synonym of T. 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 (T. alboornatus, T. wheeleri), two as brachypterous (T. montanus, T. piceus), and four (T. balli, T. fuscicornis, T. pubescens, and T. 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, T. mundulus, provides a good example of successful classical biological control (Hagen and Franz 1973, van den Bosch and Messenger 1973, Rosen 1985. 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 T. 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 T. mundulus and T. parviceps (Reuter) have been investigated for control of a tropiduchid, Numicia viridis Muir, on sugarcane (Carnegie and Harris 1969). Although T. mundulus was the better-known predator, T. parviceps was more easily reared and showed the greatest potential for controlling N. viridis. Jhansi et al. (2002) studied the biology and prey preferences of T. parviceps on planthoppers and leafhoppers on rice in India, including the brown planthopper, Nilaparvata lugens (Stål). The Holarctic species T. pubescens (Knight) and T. pygmaeus (Zetterstedt) are known to prey on leafhoppers and delphacids in England (Southwood and Leston 1959, Rothschild 1963. In coastal eastern North America, Döbel and Denno (1994) considered T. alboornatus (Knight) and T. 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. Diagnosis. 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 T. wheeleri, sp. n. In T. 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 T. 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.

Key to Species of Tytthus
Description. Macropterous male (n = 1, plus holotype in parentheses) (Fig. 2 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.
Distribution. Previously known only from Florida, New Jersey, New York, and Virginia Wheeler 1988, Hoffman 2000). New U. S. state records are Connecticut, Louisiana, South Carolina, and Texas. This distribution indicates that T. alboornatus should occur in all coastal states from at least New England to Texas.
Discussion. Tytthus alboornatus  is one of the smallest species in the genus, second in size only to the similar-appearing T. wheeleri . Like T. 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. Diagnosis. 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 T. 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 T. zwaluwenbergi is uniformly pale yellowish brown.
Description. Male (n = 5) ( 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.

Tytthus balli
Tytthus balli is very similar to T. insperatus and T. uniformis based on the black head and antennal segments I and II, and the orange to brownish-orange pronotum and femora. From T. insperatus, it is distinguished by the uniformly browish-orange pronotum lacking a pale anterior margin and the infuscated apical half of each corium. From T. uniformis it is distinguished by the fuscous tibiae and infuscated hemelytra.
Description. Macropterous male (n = 5, plus holotype in parentheses) ( Fig. 7): Length to apex of hemelytron 2.58-2.73 mm (2.68  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. Hosts. 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].
Distribution. 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 Diagnosis. This species is very similar to T. 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 T. 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, T. entrerianus, T. femoralis, and T. mexicanus, also have fuscous "knees," a character that distinguishes them from all other species of the genus, except. T. chinensis and T. 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.
Description. Male (n = 15) ( 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. Hosts. Usinger (1939) reported T. 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.
Distribution. 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(Schuh , 1995Kerzhner and Josifov 1995;Cassis 1995). Based on specimens in the USNM, most or all records reported by Schuh (1984) from India and Sri Lanka should be applied to T. 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.
Discussion. 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 C. annulicornis Poppius (both in Deutsches Entomologisches Institut) and C. riveti Cheesman (BMNH) are junior synonyms. They characterized T. 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 T. riveti and agree that it follows the concept of T. chinensis outlined by Carvalho and Southwood (1955) and Carvalho (1956), including a uniformly dark pronotum.
In addition, Kerzhner and Josifov (1995) suggested that T. koreanus Josifov and Kerzhner (1972) could be a junior synonym of T. chinensis. Although I have not examined type material of T. 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 T. 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 T. chinensis.
Although most material of what I consider to be T. 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 T. 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 T. 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 T. 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 T. parviceps from Africa and the New World. As noted above, the endosoma of typical T. chinensis is C-shaped, whereas this very simple structure in African and New World specimens of T. parviceps is usually S-shaped. Until additional Tytthus columbiensis Carvalho http://species-id.net/wiki/Tytthus_columbiensis Fig. 11 Tytthus columbiensis Carvalho 1984: 202 (orig. descrip.) ;Schuh 1995: 248 (cat.) Diagnosis. 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 T. 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 T. 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.
Description. 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 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. Discussion. 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. Diagnosis. 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 T. femoralis based on the pale tibiae with fuscous knee spots and the apically fuscous hind femora. Tytthus entrerianus can be distinguished from T. femoralis by the dark brown or fuscous antennal segment II and having only apical third of the hind femur infuscated.

Tytthus entrerianus
Description. Male (n = 1, holotype in parentheses) (composite description based on Carvalho and Carpintero, 1986, and one USNM paratype) ( 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.
Female: None examined; described from 5 paratype ♀♀. Carvalho and Carpintero (1986)  Diagnosis. 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 T. entrerianus based on the pale tibiae with knee spots and the apically infuscated hind femora. It can be distinguished from T. entrerianus by the pale antennal segment II and the more extensively infuscated hind femur.
Description. Male (n =10, plus holotype in parentheses) ( 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 corium 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. 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.
Etymology. The specific epithet A "femoralis" denotes the dark brown to fuscous hind femur in contrast to the uniformly pale pro-and mesofemora.
Hosts. 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.
Distribution. So far recorded from Bolivia, Brazil, Colombia, Costa Rica, Cuba, Ecuador, Guatemala, Honduras, Jamaica, Mexico, Panama, and Peru. Diagnosis. 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 T. mexicanus and T. 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.
Description. 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. 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.
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. 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.
Distribution Diagnosis. 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 T. balli and T. 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.
Description. Holotype male (Fig. 17 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, recum-bent setae. Hemelytron: Macropterous, subparallel, cuneus and membrane fully developed, extending well beyond apex of abdomen; evenly set with short, recumbent setae.
Host. Unknown. The Maricopa specimen below from "on cotton" almost certainly represents an incidental or sitting record.
Distribution. Described from Tucson (Pima County), Arizona, and later reported from Buckeye (Maricopa County), Arizona, and Calexico (Imperial County), California (Carvalho and Southwood 1955  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 T. femoralis and T. 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. Description. Male (n = 4; plus holotype in parentheses) (Fig. 19): Length to apex of hemelytron 2.75-2.58 mm (2.55  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.
Tytthus montanus is similar to T. 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 T. wheeleri), pale yellow antennal segment I, redstreaked pro-and mesofemora, and dark brown metafemur (versus entirely pale yellow legs in T. wheeleri). It is also similar to T. 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).
Description. Macropterous male (n=1; holotype in parentheses) ( 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.
Macropterous female (n = 1) (Fig. 22 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. 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 (frac-ture still visible on most specimens) and a much abbreviated membrane lacking any trace of areoles (Fig. 23).
Host. 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.
Distribution. Described and previously known only from Montana. Arizona and Utah represent new state records.
Discussion. 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.  Diagnosis. 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 T. chinensis and T. parviceps; T. mundulus, however, lacks the fuscous knee spots found in these two species. All known specimens of this species are macropterous. Description. Male (n =5) (Fig. 24) 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 T. 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.
Female (n = 5) (Fig. 25 Diagnosis. 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 T. 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 T. juturnaiba. Description. 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. 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.
Hosts. No specific host known. Associated with grass savannas and marshes and pine-oak sand scrub.
Tytthus pallidus keys out with T. piceus based on the pale antennal segment I and the pale hemelytra with smoky-brown shading. It can be distinguished from T. 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.
Description. Holotype male (Fig. 28 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.

Diagnosis.
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 T. 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 T. 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 T. 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 genitalia: Not examined. See note below. Similar to male in overall appearance, differing primarily in the broader body form. Host. Unknown. Distribution. Described and known only from Panama. Discussion. 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.
Other specimens examined. 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). ( Ingham et al. 1995: 73 (note). Tytthus parviceps: Carvalho and Southwood 1955: 21 (descrip., n. comb Diagnosis. 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 T. chinensis and certain color forms of T. 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 T. pymaeus by the knee spots at the bases of the tibiae. From T. 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)  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.

Tytthus parviceps
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 T. 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.
Hosts. In the New World, T. 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 . 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).
Distribution. 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 Hilburn 1990, Wheeler and. 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 Henry 1992, Kerzhner andJosifov 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 T. chinensis.
Discussion. Tytthus parviceps, described from Egypt, is similar to T. 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 T. parviceps with greatly reduced yellow markings or an entirely dark pronotum are encountered, or when specimens of T. chinensis with more extensive yellow markings are found. Dissection of representative males of T. parviceps from different regions (list localities) shows that the endosoma is consistently S-shaped, whereas specimens of T. 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 C. pellicia from Cuba (labeled in Uhler's hand) in the USNM collection, and agree that Uhler's species is a junior synonym of T. parviceps as now defined.
Also, Josifov and Kerzhner (1972), in describing T. koreanus, initially indicated that it was similar to T. parviceps (Reuter) based on the extensive pale areas on the pronotum. However, they also added that the new species might be conspecific with T. 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 T. parviceps. Unfortunately, the type of T. 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 T. 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 T. parviceps. As a consequence, I am treating T. koreanus as a junior synonym of T. chinensis (which see) until additional studies can show otherwise.
Other Diagnosis. 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 T. 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.
Description. Male (n = 10; holotype in parentheses) ( 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.
Distribution. This species was described and known previously only from Guatemala (as I. picea) and Honduras (as T. hondurensis). Colombia, Costa Rica, Mexico, Panama, and the United States (Florida, Maryland, and South Carolina) represent new country records and considerable range extensions.

Tytthus pubescens (Knight) http://species-id.net/wiki/Tytthus_pubescens
Diagnosis. This species is distinguished by the fuscous to black head; brown to black pronotum, often entirely or with only the posterior angles or posterior half whitish to pale brown; the pale yellow to yellowish brown antennal segment I and black segments II-IV; the long, erect setae in both sexes on antennal segment II as long or longer than the diameter of the segment; the uniformly pale, translucent hemelytra; and pale brownish yellow legs. Males of this species are always macropterous (Fig. 38); both macropters ( Fig. 39) and brachypters (Fig. 40) are known for females, but brachypters are most common. This Holoarctic species is superficially similar to another Holarctic species, Tytthus pygmaeus, in general color and size. Tytthus pubescens is readily distinguished by the pale antennal segment I, the long erect and semierect setae on antennal segments I and II, and the often pale humeral pronotal angles.
Description. Macropterous male (n = 10) (  : Fuscous to black, with a small, yellow, interocular spot near inner margin of each eye; eyes reddish brown. Labium: Segments I-III pale yellow; segment IV brown. Antenna: Segment I pale yellow; segment II-IV fuscous to black. Pronotum: Uniformly fuscous to black, some specimens brown at posterior angles, others pale brown on posterior half, sometimes with dark brown invading darker anterior half. Mesoscutum and Scutellum: Fuscous to black. Hemelytra: Pale translucent yellow to whitish. Ostiolar evaporative area (Fig. 84): Fuscous to black. Ventral surface: Thorax and abdomen uniformly fuscous to black. Legs: Coxae pale yellow, the bases brown; femora, tibiae, tarsi, and claws (Fig. 86) uniformly pale yellow.
Structure, texture, and vestiture: Head: Weakly shiny, impunctate; set with relatively long, erect and semierect setae on vertex and frons. Labium: Extending to apices of meso-or bases of metacoxae; segment I extending just past base of head to anterior margin of xyphyus before procoxae. Antenna: Segment I set with rather short, sparse, recumbent setae and two to four or more long, erect, subapical, bristlelike setae; segment II densely set with short, recumbent setae, intermixed with erect and semierect setae mostly subequal in length to diameter of segment. Pronotum: Nearly rectangular; anterior angles rounded; lateral margins straight, only slightly widening to posterior angles; posterior margin straight or only very slightly sinuate; set with relatively long, recumbent and semierect setae. Mesoscutum: Broadly exposed, even in brachypters; with a few scattered, semierect setae. Scutellum: weakly shiny, equilateral; set with relatively long, semierect setae. Hemelytron: Macropterous, subparallel; cuneus and membrane fully developed, extending beyond apex of abdomen; evenly set with recumbent setae.
Diagnosis. This highly variable species is distinguished by the black head, with relatively vague interocular spots; entirely pale (European material) to entirely fuscous to black (all North American material) pronotum, with intermediate color forms; very clear or translucent white hemelytra; the fuscous to black antennal segment I, with the apical one fourth pale yellow; antennal segments II-IV uniformly fuscous to black; and the uniformly pale yellow legs. All males and most females of this species are fully macropterous; only a few weakly brachypterous females, with the membrane extending to the apex of the abdomen, have been examined.
Tytthus pygmaeus is superfically similar to the Holarctic T. pubescens in overall size, coloration, and distribution. It can be distinguished from T. pubescens by the fuscous antennal segment I having only the apex pale, the short, recumbent setae on antennal segments I and II, and the variably fuscous to pale pronotum (but never fuscous with pale humeral angles as in T. pubescens).
Head: Length 0.29-0.32 mm, width across eyes 0.69-0.75 mm, interocular width 0.30-0.32 mm. Labium: Length 1.07-1.12 mm. Antenna: Segment I length  Coloration: Head: Shiny fuscous to black, with a relatively small, yellow, interocular spot near inner margin of each eye; eyes fuscous to dark reddish brown. Labium: Uniformly pale yellow, with apex of segment IV usually brown. Antenna: Segment I, fuscous to black, with only apical one fourth to one third pale yellow; segment II-IV black. Pronotum: Highly variable, ranging from uniformly pale yellow to entirely fuscous to black, with many intermediate forms, including all of disc dark and anterior half yellow and almost entirely dark with yellow across anterior margin and through middle of calli; some yellow specimens with only fuscous posterior angles. Mesoscutum and Scutellum: Uniformly fuscous to black. Hemelytron: Uniformly clear to very pale translucent white. Ostiolar evaporative area: Fuscous to black, even on palest specimens. Ventral surface: Thorax uniformly fuscous; abdomen fuscous along lateral margins and genital capsule, ventral area and sides pale yellow to whitish. Legs: Uniformly pale yellow.
Structure, texture, and vestiture: Head: Shiny, impunctate, much broader than long; set with numerous, long, erect and semierect setae on vertex and frons. Labium: Extending to apices of meso-or bases of metacoxae; segment I extending just beyond head to anterior margin of xyphus just before procoxae. Antenna: Segment I sparsely set with recumbent setae and two, long, erect, subapical, bristlelike setae; segment II, thickly set with short, recumbent and semierect setae. Pronotum: Trapeziform, anterior angles nearly weakly rounded; lateral margins straight, gradually widening to posterior angles; posterior margin nearly straight or only weakly sintuate. Mesoscutum: Broadly exposed. Scutellum: Equilateral; sparsely set with scattered relatively short, semierect setae. Hemelytra: Macropterous, cuneus and membrane fully developed, extending well beyond apex of abdomen; evenly set with recumbent setae.
Tytthus uniformis is most similar in overall appearance to T. balli and T. insperatus. It can be distinguished from T. balli by uniformly brownish-orange pronotum and hemelytra; T. balli is usually infuscated on the posterior half of the pronotum, and the inner half of the clavus and apical third of the corium are dark brown. It is distinguished from T. insperatus by the uniformly pale orange pronotum and legs; T. insperatus has a dark brown pronotum, each femur has a narrow dorsal and lateral red stripe, and the hind tibiae are fuscous. Coloration: Head (Figs 89-91): Semishiny fuscous to black, pale spot on either side of vertex obsolete or indistinct. Labium: Antenna: Uniformly black, segment I very narrowly pale at apex on some specimens. Pronotum: Uniformly pale brownish orange, posterior angles sometimes slightly infuscated; collar pale or whitish; mesoscutum and scutellum uniformly pale brownish orange. Hemelytron: Uniformly pale, translucent, brownish orange; membrane translucent smoky brown. Ostiolar evaporative area (Fig. 92): Brownish orange to orange. Ventral surface: Thoracic segments orange to reddish orange; abdomen in males reddish orange dorsally, paler orange ventrally, with genital capsule becoming fuscous or black; abdomen in females infuscated dorsally, gradually fading to pale orange ventrally. Legs: Uniformly pale brownish orange; claw (Fig. 94).
Structure, texture, and vestiture: Head: Wider than long, interocular width subequal to length, impunctate, semishiny; set with a few, scattered, erect, nearly bristlelike dark setae. Labium: Extending beyond metacoxae to base of abbdomen. Pronotum: About 2.5 times as wide at base as long, lateral margins weakly concave, posterior margins moderately flared; set with short, semierect and recumbent setae. Mesoscutum: Broadly exposed. Scutellum: Slightly wider across base than on sides; set with short, semierect and recumbent setae. Hemelytron: Entire in males, cuneus nearly three times as long as wide at base, membrane fully developed, extending well beyond apex of abdomen; set with short, semierect and recumbent setae.
Etymology. Named for the uniformly pale brownish-orange pronotum, hemelytra, and legs.
Host. Big sacaton, Sporobolus wrightii Munro ex Scribn. [Poaceae].  Maw et al. 2000: 121 (list). Diagnosis. This species is distinguished by the black head, pronotum, and scutellum; black antennae; dark translucent smoky brown hemelytra; and pale yellowish brown legs, with the hind femora infuscated on the distal third to one half. All known specimens are macropterous.
Tytthus vagus is similar to T. femoralis in having dark antennae, pale infuscated hemelytra, and apically infuscated hind femora. It is readily distinguished from T. femoralis in lacking distinct fuscous knee spots on the tibiae. It keys out with T. panamensis and T. juturnaiba, but is separated by the combination of dark antennae and infuscated hind femora.
It is similar to T. montanus in overall brown coloration with the basal area of the corium and clavus pale, but is distinguished from that species by the much smaller size, dark antennal segment I, and pale tibiae. It is also similar to T. alboornatus in overall coloration, but differs in lacking a narrow pale area across the apex of the corium and the dark antennal segment I and pale segment II. All specimens of T. wheeleri at hand, except one macropterous female, are strongly brachypterous and lack a cuneus and membrane on each hemelytron. Macropterous male: Unknown. Coloration: Overall coloration dark brown. Head: Dark shiny brown, becoming paler yellowish brown ventrally, yellow spot on inner margin of each eye absent; eyes fuscous, often with a reddish tinge. Antenna: Segment I dark brown; II, pale yellowish brown, sometimes weakly red tinged at apex; III and IV pale yellowish brown. Pronotum: Dark shiny brown; scutellum dark shiny brown with apical half to one third pale yellowish brown. Hemelytron: Dark brown pale translucent yellowish brown to whitish on basal one third to half of corium. Ventral surface: Thorax dark brown, usually red tinged; ostiolar auricle brown, often red tinged; abdomen dark brown laterally, becoming paler Discussion. All specimens of this new species are brachypterous, except for one macropterous female collected in Highlands Co., Florida (March). That a macropterous female was discovered indicates that macropterous males eventually should be found. Given the small size and considering how difficult it is to collect this species from the crowns of certain bunch grasses, it is not surprising that few undetermined specimens of this species were found in collections.
Diagnosis. This species is readily distinguished by the uniformly yellow to testaceous coloration, except for the dark brown eyes and a vague brown area at the middle of the head. Coloration: Uniformly yellow to testaceous, except for the dark brown eyes and a vague brown area on the middle of the head.
Structure, texture, and vestiture (after Usinger 1944): Head half again as broad as long, 11.5; 8, smooth, shining, and strongly convex above. Eyes slightly less than half as wide as interocular space, 2.75: 6. First antennal segment shorter than interocular space, 5: 6, second segment three times as long as first, third and fourth segments broken off. Rostrum nearly reaching apices of middle coxae. Pronotum somewhat duller than head, clothed with short, sparse, decumbent hairs; broader across humeri than width of head, 15: 11.5, and less than half as long as broad, 6.5: 15; front margin shallowly concave, lateral margins feebly sinuate, and hind margin slightly concave. Scutellum longer than pronotum at middle, 7: 6, subdepressed, the disk very sparsely clothed with appressed hairs. Hemelytron simple, distinctly but sparsely clothed with appressed hairs; costal margin slightly, evenly arcuate. Legs slender, clothed with short, inconspicuous, pale hairs. Claws with simple hairlike setae rather than arolia.
Hosts. Recorded from Boerhavia sp. (Nyctaginaceae) by Usinger (1944) and Schuh (1984). Usinger (1944) noted that the various species of the genus prey on delphacid eggs, but since "delphacids have not been reported on Canton Island and since both Cyrtorhinus and the cicadellid Nesaloha cantonis Oman were collected on Boerhaavia, it is possible that this new mirid is a predator on Nesaloha." Distribution. This species has been reported from Baker Island, Howland Island, and the Phoenix Islands (Canton Island) in the central Pacific (Usinger 1944, Schuh 1984. Discussion. Tytthus zwaluwenburgi was described from only three specimens (holotype, paratype, and one teneral specimen). Schuh (1984)

Phylogeny
Menard (2011) and Menard and Schuh (2011) have shown that Tytthus does not belong in Leucophoropterini based on both molecular sequence data and morphology and, thus, they transferred Tytthus to nominate tribe Phylini where it shows a relationship with several New World genera, including Criocoris Fieber, Semium Reuter, and Spanagonicus Berg. These finding are supported by Kelton's (1959) work that showed similarities in the male genitalia among these genera. The relationship of Tytthus to New World genera, the restriction to the Western Hemisphere of 17 of its 24 species, and the Holarctic distribution of two other species suggest a New World origin. Only the far-eastern T. chinensis, the Indo-Pacific T. mundulus, and T. zwaluwenbergi found on a few central Pacific islands are restricted to the Old World.
The morphology-based phylogeny of Tytthus presented here should be considered tentative because of the limited number of informative characters available to infer relationships. Although most species possess distinctions that allow relatively easy separation, characters, such as antennal, hemelytral, and leg coloration, are very homoplastic and did not offer much resolution. The male genitalia are relatively simple structures (e.g., Figs 109-116) and lack an apparent secondary gonopore, which is shared with several outgroup taxa, including species of the genera Criocoris Fieber and Semium Reuter (Kelton 1955, Menard 2011. As a consequence, the matrix has fewer characters than taxa. For this analysis, the matrix (Table 1) contained 27 species-group taxa and 23 characters, 17 of which were multistate. Characters (Table 2) were first processed using WinClada (Nixon 1999(Nixon -2002 to run NONA (Goboloff 1999), using the default settings for the ratchet function. Three taxa (T. chinensis, T. parviceps, and T. pygmaeus) were scored polymorphic for pronotal coloration. Four characters (6, antennal segment II setae; 10, pronotal color; 13, hemelytral color; 19, knee spots) were scored additive because they reflected clear relationships among certain taxa, whereas homologies for all remaining characters were uncertain and, thus, scored as nonadditive. The genera Criocoris, Plagiognathus Fieber, and Semium were used as outgroups. The analysis with all characters activated resulted in nine most parsimonius cladograms, with a length of 117 steps, a ci of 47, and an ri of 60. Generation of a strict consensus (Fig. 198) tree collapsed 10 nodes, reflecting the considerable homoplasy in the data. Manipulation of certain characters with very low consistencies, led to the conclusion that character 17 (hind femur color) was very uniformative, which significantly skewed the results. By deactivating, character 17, only four trees  were generated, with a length of 110, a ci of 46, and and ri of 59 The resulting consensus tree (Fig. 199) collapsed 9 nodes and resulted in a topology not too dissimilar to the previous analysis. The genus Tytthus is defined by a combination of the yellow spots on interocular area, bell-shaped or campanulate pronotum, simple S-or C-shaped endosoma, and host specificity (eggs of Delphacidae). In all trees , three monophyletic species groups were hypothesized: the balli group (T. balli, T. insperatus, and T. table 2. Characters and character states used in analysis of the genus Tytthus. The four left columns represent 1 character number 2 number of steps 3 consistency index, and 4 retention index. Character statistics are from one of nine trees (Fig. 194) generated by WinClada to run NONA using the ratchet function. uniformis), the chinensis group (T. pymaeus, T. mundulus, T. pubescens, T. parviceps, T. chinensis, and T. mexicanus), and the alboornatus group (alboornatus, columbiensis, montanus, and wheeleri). The balli group was defined by one synapomorphy, the pale orange pronotum (character 9, state 3). These species are also share a broad black head, a strongly campanulate pronotum, and orange or largely orange scutellum, hemelytra, and legs. The chinensis group was defined by one synapomorphy, the brushlike setae on antennal segment II (character 5, state 1), and the broad swordlike left arm of the left paramere (character 21, state 3). Within this group, T. chinensis, T. mexicanus, and T. parviceps shared the fuscous knee spots (character 18, state 1), found elsewhere only in T. entrerianus and T. femoralis. The alboornatus group was defined by one synapomorphy, the largely dark brown hemelytra with pale areas at the base and apex of the corium (character 12, state 3). Also, within this group, T. alboornatus and T. wheeleri possess the most extreme hemelytral brachyptery in the genus; brachyptery is found to a lesser extent in two other members of the group, T. montanus, and T. piceus, the latter of which falls outside the group. Tytthus entrerianus and T. neotropicalis always came out as sister species, based on two synapomorphies, the pale fore and middle femora (characters 14, state 2, ♀ character 15, state 2), as did T. pallidus and T. piceus, based on the color of the hemelytra (character 12, state 1) and the pale antennal segment I (character 3, state 3). Tytthus zwaluwenburgi is the most problematic species of the genus. Although it came out as the sister species to T. amazonicus, this relationship, supported only by color characters (character 3, state 4 ♀ character 9, state 2), is un- likely to be correct. Its pale coloration and extreme isolation on islands in the central Pacific preclude any meaningful hypothesis about its relationship to other species. The characters for most remaining taxa exhibited considerably more homoplasy and, thus, their hypothesized position among the species was less stable.
The same character matrix (Table 1) as above, with 27 species-group taxa and 23 characters (with the same four characters scored as additive and character 17 deactivated), also was run using TNT (Goloboff et al. 2010). A first pass using the traditional analysis setting resulted in 29 trees with a length of 116, a ci of 46, and an ri of 60. An additional run using the random addition sequence function resulted in only one most parsimonius tree (Fig. 200), with a length of 109, a ci of 45, and an ri of 59. This tree was far more resolved than either of the above strict consensus trees (Fig. 198,199) generated by Nona and most closely resembled tree number one (Fig. 194) of the four generated by Nona with character 17 deactivated. Tytthus uniformis, T. balli, and T. insperatus were hypothesized as a monophyletic group, as were T. mundulus, T. pubescens, T. parviceps, T. chinensis, and T. mexicanus. In sequence, T. panamaensis was hypothesized as sister to the remainder of the taxa, followed by the holoarctic T. pygmaeus. Tytthus juturnaiba and T. vagus formed a sister pair, as did T. pallidus and T. piceus and T. entrerianus and T. neotropicalis. Also, as in all of the previous analyses, T. columbiensis, T. montanus, T. alboornatus, and T. wheeleri formed a monophyletic group. The problematic T. zwaluwenburgi still grouped with T. amazonicus, a doubtful relationship.
As noted in my discussion of T. chinensis and T. parviceps, molecular data likely will help resolve some of the unclear relationships among the species of this intriguing genus of egg predators. If T. juturnaiba proves to be a junior synonym of T. neotropicalis, the removal of that species and its conflicting character information should help reduce the level of homoplasy in certain characters and likely will yield a better hypothesis of relationships.