Zelus Fabricius, 1803, p. 281, orig. descr.; Latreille, 1804, p. 260, list; Latreille, 1807, p. 129, list; Latreille, 1810, p. 433, type desig.; Lepeletier and Serville, 1825, p.815, list and descr.; Laporte, 1832, p. 9, type desig.; Burmeister, 1835, p. 225, descr.; Brulle, 1836, p. 316-317, descr.; Blanchard, 1840, p. 100, descr. and note; Blanchard, 1845, p. 433, 438, list and note; Herrich-Schaeffer, 1848, p. 88, descr. and note; Kolenati, 1857, p. 458-459, descr.; Stål, 1861, p. 148, descr.; Stål, 1862, p. 449-454, key and subgeneric descr. (with subgenus Zelus); Carpenter and Westwood, 1863, p. 565, note; Mayr, 1866, p. 138, list; Stål, 1866, p. 296, list; Stål, 1868, p. 107, restriction of definition; Stål, 1872, p. 69, 88, key and cat. (with subgenus Zelus); Walker, 1873, VII., p. 49, key, VIII., p. 131-136, cat.; Berg, 1879, p. 150, list (with subgenus Zelus); Uhler, 1886, p. 24, checklist; Provancher,1887, p. 179, note; Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 251, cat. and note; Kirkaldy, 1900a, p. 263, type verification; Kirkaldy, 1900b, p. 242, syn.; Kirkaldy, 1902, p. 149, note; Fracker, 1913, p. 223, 238-240, key and note (with subgenus Zelus); Van Duzee, 1916, p. 30, checklist (with subgenus Zelus); Van Duzee, 1917, p. 258-259, cat. (with subgenus Zelus); Blatchley, 1926, p. 567-568, key, descr. and note (with subgenus Zelus); Readio, 1927, p. 167, 168-169, key, descr. and note; Zimmerman, 1948, p. 137, note; Wygodzinsky, 1949a, p. 48, checklist; Fracker and Usinger, 1949, p. 277, key to nymphs; Alayo, 1967, p. 5, 35, list, key and note; Hart, 1986, key to North American species; Hart, 1987, key to Caribbean species; Maldonado, 1990, p. 325-332, cat.

Reduvius Fabricius, 1775 (type by subsequent desig., Cimex personatus Linnaeus, 1758), Lepeletier and Serville, 1825 (in part), p. 272, descr.; Perty, 1834 (in part), p. 173, list of species.

Arilus Hahn, 1831 (type by subsequent designation, Cimex carinatus Forster, 1771); Burmeister, 1835 (in part), p. 227-228, descr.; Herrich-Schaeffer, 1848 (in part), p. 33-35, descr.

Euagoras Burmeister, 1835 (type by subsequent designation, E. stollii Burmeister, 1835) (in part), p. 226, descr.; Amyot and Serville, 1843 (in part), p. 368, descr. (as Evagoras); Herrich-Schaeffer, 1848 (in part), p. 43-44, descr.; Stål, 1855 (in part), p. 189, list (as Eccagoras); Stål, 1861, p. 148, (in part) junior syn. of Zelus Fabr.; Mayr, 1866, p. 139, list; Walker, 1873, p. 49, 117, key and cat.; Provancher, 1887, p. 182, descr. (as Evagoras); Kirkaldy, 1900b, p. 242, junior syn. of Zelus Fabr.; Kirkaldy, 1903, p. 215-216, note.

Diplodus Amyot and Serville, 1843, p. 370, descr.; Burmeister, 1853, p. 91, list (included in Euagoras Burm.); Stål, 1860, p. 74, list; Stål, 1862, p. 450, descr. (as subgenus of Zelus); Stål, 1866, p. 296, key; Stål, 1872, p. 90, list (as subgenus of Zelus); Walker, 1873, VII., p. 49, VIII., p. 123, key and cat. (as Diploda); Berg, 1879, p. 151, list (as subgenus of Zelus); Uhler, 1886, p. 24, checklist; Provancher, 1887, p. 179, key and descr.; Kirkaldy, 1903, p. 232, note; Fracker, 1913, p. 239, 240, key and list (as subgenus of Zelus).

Darbanus Amyot and Serville, 1843 (type by monotypy, D. nigrolineatus); Provancher, 1872, p. 106, species descr.; Uhler, 1886, p. 24, checklist; Provancher, 1887, p. 179, 181, key and note; Van Duzee, 1912, p. 324; Fracker, 1913, p. 241, note.

Pindus Stål, 1862, p. 454, orig. descr. (as subgenus of Zelus); Stål, 1866, p. 296, key (as genus); Stål, 1872, p. 92, list and cat., as subgenus of Zelus); Walker, 1873, VII., p. 66, list and cat. (as genus); Berg, 1879, p. 150, list (as subgenus of Zelus) ; Thierry and Severin, 1896, p. 151, cat.; Fracker, 1913, p. 223, 240, key and list; Van Duzee, 1916, p. 30, checklist; Van Duzee, 1917, p. 261, cat.; Blatchley, 1926, p. 569, key.

Diplacodus Kirkaldy, 1900b, p. 242, new name for Diplodus A. and S. (preocc.).

Diplocodus Van Duzee, 1916, p. 30, checklist (new name for Diplacodus Kirkaldy, preocc.); Van Duzee, 1917, p. 260, cat.; Blatchley, 1926, p. 569, key.

Iquitozelus Bérenger, 2003, p. 23, orig. descr., syn. nov. (current study).

Male: Small to large, total length 8-25 mm (Suppl. material 2), with most of moderate sizes (11-18 mm); usually slender (length/width = 4.0-5.0), some species relatively robust (<3.5) to rather slender (>6.0). COLORATION: Colors and patterns of preserved specimens variably yellowish-brown, reddish-brown, orange-brown, and brownish-black to black, with most species uniformly colored. VESTITURE: Most species with moderately dense or dense, fine, short, recumbent and short, long, erect setae; some species with short, spine-like setae on head and pronotum; few species nearly glabrous. Setation on legs sparse in most species; profemur and tibia with dense sundew setae in some species. STRUCTURE: Head: Length much greater than width across eye. Postocular lobe usually longer than anteocular, tube-like posteriorly in most species. Ocellus raised, directed somewhat laterally. Eye variably sized, not protruding above or below dorsal or ventral surfaces of head, with one exception (Zelus grandoculus sp. n.). Antenna: Scape and basiflagellomere long and subequal in length, usually longer than head and pronotum combined; pedicel and distiflagellomere short and about 1/3 length of scape. Scape thickest; basiflagellomere usually thicker than pedicel, subequal in some species. Labium: Segment II longest, 1.3-2.2x length of segment I; segment III shortest, usually 0.5x length of segment I; variably curved between segments I and II. Thorax: Anterior pronotal lobe about 1/2 to 3/4 length of posterior lobe; anterolateral angles of pronotal collar rounded, with or without tuberculate protrusion; medial dorsal longitudinal sulcus usually shallow at collar, deepening through posterior 1/2; sometimes with subtuberculate elevation near posterior margin laterad to medial sulcus. Posterior pronotal lobe rugulose (not conspicuous in species with dense setation); slightly or greatly wider than anterior lobe; disc of most species elevated above humeral angle and posterior margin of lobe; humeral angle with tuberculate to long spinous lateral process, rounded and unarmed in small number of species. Scutellum in most species with angulate apex, slightly produced and projected upward in some species. Legs: long, slender in most species; femoral diameters generally subequal; pro- and metafemoral lengths subequal, greater than mesofemoral length. Hemelytron: Attaining or surpassing apex of abdomen, by large proportion in some species. Quadrate cell small to large; median vein conspicuous in some species and not visible in many. Cu and M of cubital cell subparallel in most species, converging in some. Abdomen: Lateral margins subparallel; ventral outline usually straight, in some species somewhat concave and abdomen appearing arched (see "Material and methods" for discussion of this character). Genitalia: Segment eight usually short, less than 1/2 length of pygophore; posterior margin generally slightly concave, straight in some species, never convex. Pygophore: ovoid to elongated; slightly to greatly expanded laterally close to base of paramere; dorsal bridge short to long. Medial process single, not bifurcating, of variable length and shape; triangular or cylindrical as most common configuration; apex blunt or with hooklike process. Paramere generally cylindrical, often swollen and bending apically, length variable. Phallus: Dorsal phallothecal sclerite generally semi-cylindrical, broad and shield-like in many species, elongated in some; dorsal surface lacking armature in most species, with projection, process or elevation in some species; lateral margins usually straight or convex, constricted or recurved in some species; apical part keeled in middle and/or curved dorsad; apex usually rounded or truncate, with or without medial emargination. Struts attached to dorsal phallothecal sclerite in majority of species; apical part recurved dorsad and often semi-circular; bridge connecting two sides in many species. Basal plate arm slender to heavy, separate or fused; basal plate bridge present, variable in width and degree of sclerotization; basal plate extension short, often extended onto basal plate arm.

Female: Larger than male. Coloration usually similar to that of male and more variable in some species, but may differ between sexes dramatically in certain species. Eye and ocellus smaller than in male in some species. Basiflagellomere not swollen and about equal diameter as or smaller than pedicel. Lateral process on humeral angle, if present, usually more produced and longer than in male. Mesofemur slightly swollen in many species. Lateral margins of abdomen expanded in some species.

This genus is distinguished from other genera of the New World Harpactorini by the cylindrical head, the length of the head being at least 1.9X its width; the unarmed antenniferous tubercles; the second labial segment being at least 1.3x the length of the first segment; the long scape and basiflagellomere that are subequal in length and the short pedicel and distiflagellomere; the generally unarmed (i.e. no tubercles or spines) disc of the posterior pronotal lobe (except in Zelus tetracanthus Stål, 1862, Zelus lewisi sp. n. and Zelus minutus Hart, 1987); the humeral angle with or without process, and if present, usually not prominently projected; the legs with sundew setae and sticky glands (Zhang and Weirauch 2013); the profemur subequal in length and diameter to the metafemur; and the medial process of pygophore single, not bifurcating. Zelus is apparently closely related to three other genera, Atopozelus Elkins, Ischnoclopius Stål and "Hartzelus" [manuscript name], that share many of the aforementioned characters. It is separated from Atopozelus by the presence of paramere (lacking in Atopozelus). Ischnoclopius is distinguished from Zelus by its rather slender body form (length:width ratio greater than seven), the very long profemur, at least 0.6x of body length, and the very short paramere. An undescribed genus, "Hartzelus" (Gil-Santana and Berenger, pers. comm.), which would be in part based on species removed by us from Zelus, differs from Zelus in having a bifurcating medial process of the pygophore (single in Zelus) and generally more slender legs. No Old World species of Harpactorini are similar or appear to be closely related to Zelus. Confusion may potentially arise with members of genera that show a similar slender body form and slender legs (e.g., Euagoras and Vestula Stål), but these are distinguished from Zelus based on the characters listed above.

Native to (except for Chile) and throughout the New World, including the Caribbean, with highest diversity in the Neotropics. One species (Z. renardii) has been introduced to Hawaii, the Polynesian islands, Jamaica, Philippines, Spain, Greece and Chile.

We provide a non-exhaustive account of the biology of various species of this genus. As with other harpactorines, species of Zelus generally do not show associations with or preferences of host plants, probably due to their generalist habits. However, two recent studies have found two species of Zelus that have both nymphs and adults occurring in the same plant species in relatively large quantities. In Gil-Santana and Alves (2011), based on a multi-year study the authors observed forty-seven females, twenty-seven males and fifteen nymphs of Z. versicolor from Bidens rubifolia Kunth (Asterales, Asteraceae) in a single site in the city of Nova Friburgo, Brazil. Interestingly, they did not see individuals of the same species in other plants in the same site, which can be seen as evidence for host plant preference. In French Guiana Revel et al. (2010) counted as many as 405 individuals of Zelus annulosus (Stål, 1866) and its egg masses from several pubescent plant species, including (but not limited to) Hirtella physophora Mart. & Zucc. (Chrysobalanaceae), Cordia nodosa Lam. (Boraginaceae) and Tococa guianensis Aubl. (Melastomataceae); all three are myrmecophytes. They hypothesized an intriguing tri-party mutualistic relationship between the assassin bug, an ant (Allomerus decemarticulatus Mayr) and the plants.

Several species of Zelus are possibly mimics of various other insects. Zelus errans Fabricius, 1803, Zelus vespiformis Hart, 1987 and to some extent Zelus vagans Fabricius, 1803 and Zelus gracilipes sp. n. have wing and body color patterns similar to many braconid wasps, an intriguing form of mimicry seen also in a number of other Neotropical harpactorine genera. Zelus vagans shows areas of black and orange colors, however, the posterior pronotal lobe is medially dark and laterally orange. Zelus gracilipes also shows a uniformly orange posterior pronotal lobe, but the hemelytron is uniformly dark and lacks the banding pattern typical to a wasp mimic. Zelus nigromaculatus Champion, 1899 has an appearance similar to that of a typical vespid, the only species in this genus with that kind of color pattern. Zelus laticornis (Herrich-Schaeffer, 1853), Zelus grassans Stål, 1862 and Zelus ruficeps Stål, 1862 have red and dark markings on abdomens and orange or reddish dorsal surfaces, a pattern found in many species of pyrrhocorids (e.g., Dysdercus spp.) and coreids (e.g. Hypselonotus spp.). Interestingly, in Z. laticornis, it is only the females showing this coloration. Certain color forms of Z. longipes are possibly mimics of the milkweed bug, Oncopeltus fasciatus (Dallas).

Weirauch et al. (2012) studied predatory and mating behaviors of Z. renardii and Z. tetracanthus and discussed a possible link between biological attributes and invasion potential. Law and Sediqi (2010) experimentally demonstrated that sticky substance derived from egg mass coating improves predation success and substrate adhesion ability of Z. renardii first instar.

The generic limit of Zelus is now relatively well defined and the genus can be separated from all other but one genera of New World Harpactorini based on characters discussed in the diagnosis. Based on a molecular phylogeny, Zhang and Weirauch (2014) recovered the monophyly of Zelus, Atopozelus and "Hartzelus" (which includes Z. araneiformis, a species we remove from Zelus). In that analysis, Ischnoclopius was represented by a single species and placed as sister to Atopozelus. The genera Atopozelus, "Hartzelus", Ischnoclopius and Zelus together constitute a monophyletic group in the same study, and we here refer to this group the "Zelus clade". Without a cladistic analysis, questions remain if the characters used to diagnose Zelus are synapomorphies of that genus. It is almost certain that the unbifurcating medial process represents a symplesiomorphic state as that character can be seen in Atopozelus, Ischnoclopius and many other Neotropical harpactorines. The unarmed antenniferous tubercles are also plesiomorphic to Zelus, since all other genera of the Zelus clade exhibit that condition, but may be synapomorphic to the Zelus clade. We agree with Forero (2012) that Zelus is defined mainly by the absence of apomorphies seen in other genera. Future research should illuminate this issue by studying the distribution and the polarity of characters with a formal cladistic framework.

The genus that we are uncertain about its relationship with Zelus is Pronozelus Forero, erected by Forero (2012) to accommodate a new species, Pronozelus schuhi Forero, 2012. This species appears to possess all the characters diagnostic of Zelus, but also shows some peculiar characters. The principal characters separating Pronozelus from Zelus include the laterally expanded posterior pronotal lobe, the prominent, greatly expanded posterolateral rim of pygophore lateral to paramere socket, and the posterior pronotal lobe greater than 2.2x length of the anterior lobe. The conspicuous lateral expansion of the posterior pronotal lobe is not observed in any species of Zelus, but this character appears to be autapomorphic in the Zelus clade and does not support P. schuhi being phylogenetically separated from Zelus. We have not done an extensive survey of the condition of the posterolateral rim of the pygophore and cannot determine the distribution or polarity of the lateral prominence as exhibited in P. schuhi. In Zelus rosulentus sp. n., the posterolateral part of the pygophore also appears to be expanded, although not as prominent as that seen in P. schuhi. Finally, according to measurements done in this study, in Zelus spp. the posterior pronotal lobe frequently exceeds 2.2x length of the anterior lobe, thereby negating the use of that character as a basis for placing P. schuhi outside Zelus. Despite the foregoing discussion, we have opted to not transfer P. schuhi to Zelus or synonymize Pronozelus with Zelus. The polarity of the characters diagnostic to either genus has not been clearly defined. There remains a possibility, although we think a small one, that P. schuhi represents a lineage sister to Zelus.

Bérenger (2003)'s new species, Iquitozelus couturieri, exhibits all the characters diagnostic of Zelus, except for those of the male genitalia as the known specimens are all females. The main character that Bérenger used as the basis for erecting a new genus, i.e., the "foliaceous expansion of the VI connexivum segment", appears to be autapomorphic within the Zelus clade. Synonymy of Iquitozelus with Zelus is warranted and established here. We further postulate that I. couturieri is most closely related to Zelus amblycephalus sp. n., Zelus umbraculus sp. n. or Zelus umbraculoides sp. n. Further discussions regarding the status of Iquitozelus and the specific membership of Zelus couturieri syn. nov. (Bérenger, 2003) are presented in the treatment of that species.

Maldonado (1990) considered two unpublished, manuscript names invalid, and they are "Diplodus armiger" and "Diplodus melanophthalmus". They appeared in Dohrn (1860). We follow this treatment.

Except for several pairs or complexes of closely related species, identification of males can be almost always unambiguously performed based on exposed genitalic structures such as paramere and medial process, further corroborated with phallic structures, external morphology and coloration. Identification of females of many species, where females appear to be as distinct as males, is straightforward based on coloration and external morphology. However, identification can be difficult for closely related species, where females are indistinguishable based on external morphology. In these cases, association of males and females and identification of females were primarily based on collecting event information. Sexual dimorphism presents another special challenge. While most species show limited sexual dimorphism that does not go beyond minor size and coloration differences, some species exhibit pronounced differences between the sexes (see Material and Methods for discussion of association of male and female specimens). Based on the observation that species in closely related genera do not exhibit strong sexual dimorphism, we here hypothesize that pronounced sexual dimorphism is a derived condition within Zelus.

We find here that previous subgeneric groups are based on superficial resemblance and these are not adopted. Instead, we recognize eleven species groups in the current study, based primarily on characters of the male genitalia, but also on non-genitalic external morphology if those characters can be applied to both sexes. Several species for which only females are known are therefore not assigned to a species group. Although the groupings proposed here are not based on a cladistic analysis, they show a degree of congruence with the relationships recovered in the phylogenetic analysis based on molecular data in Zhang and Weirauch (2014) and many of the characters are putative synapomorphies of the groups. A brief discussion of the species groups is presented below.

1. Zelus tetracanthus species group.

Zelus minutus Hart, 1987, Zelus prolixus Stål, 1860, Zelus rosulentus sp. n. and Zelus tetracanthus Stål, 1862.

Members of this group have a rather broad, indistinct medial process, the base of which is nearly continuous with or inseparable from the ventral rim of the pygophore. We speculate that this character represents a plesiomorphic condition as it is seen in several other genera of the New World Harpactorini and thus the condition of the medial process does not necessarily support the monophyly of this group. Zelus tetracanthus and Z. minutus also both have tubercles on the disc of the posterior pronotal lobe, which are more pronounced in the former. Comparative views of male genitalia are shown in Fig. 2.

2. Zelus luridus species group.

Zelus ambulans Stål, 1862, Zelus antiguensis sp. n., Zelus exsanguis Stål, 1862, Zelus grandoculus sp. n., Zelus luridus Stål, 1862 and Zelus spatulosus sp. n.

This is a group of species with primarily a North American distribution, with some species extending to northern Central America. The males show an apically expanded paramere and a triangular medial process that has a protrusion at the base but lacks any apical modifications. Notably, Z. spatulosus has a slender medial process, deviating greatly from the remainders of the group. It is placed in this group mainly because of the apically expanded paramere and the uniform coloration. Zelus ambulans and Z. exsanguis have the humeral angle elevated to about same level of and nearly continuous with the disc of the posterior pronotal lobe, a condition rarely seen in the genus. The coloration is quite homogenous among members of this genus, most of which have a uniform greenish (in live specimens) or dull brownish (in preserved specimens) habitus, with only Z. ambulans showing variable patterns or banding on the pronotum or legs. Comparative views of male genitalia are shown in Fig. 3.

3. Zelus mimus species group

Zelus inconstans Champion, 1898 and Zelus mimus Stål 1862.

Members of this group, consisting of only two species, exhibit a highly unique paramere and a medial process of the pygophore. The paramere is slender and apically curved dorsad at an angle of nearly ninety degrees. The medial process, as is especially evident in Z. inconstans, possesses a simple posterior liplike fold at the apex; its lateral margins are subparallel and not broadened significantly at the base. Both have a semi-cylindrical dorsal phallothecal sclerite which is modified by a fold running obliquely toward the base from the middle of the lateral margins. Both species, being quite small, exhibit the usual reduction of setal tracts common to nearly all small species. Comparative views of male genitalia are shown in Fig. 4.

4. Zelus nugax species group.

Zelus grassans Stål, 1862, Zelus illotus Berg, 1879, Zelus impar Kuhlgatz, 1902, Zelus nugax Stål, 1862 and Zelus pedestris Fabricius, 1803.

This is a group of smallish species with quite variable distributional ranges. The defining characters include a slender, laterally compressed medial process that is curved or recurved, and an acute apex of the dorsal phallothecal sclerite (except in Z. grassans). Zelus nugax has one of the widest distribution ranges in this genus, ranging from much of Mexico to northern South America. Zelus grassans is found primarily in Central America and the remaining two species mainly in northern South America. Comparative views of male genitalia are shown in Fig. 5.

5. Zelus puertoricensis species group.

Zelus bruneri De Zayas, 1960, Zelus puertoricensis Hart, 1987, Zelus subimpressus Stål, 1872 and Zelus zayasi Bruner and Barber, 1937.

Members of this group are restricted to the Caribbean. They can be easily recognized by the rather slender body form. The posteriorly directed, robust medial process with a somewhat blunt apical protrusion is also distinctive of this group. The basal plate arms are widely separate and diverging and these features are rare in other species in the genus. They show resemblance to species of the Zelus renardii species group, especially to Z. cervicalis. Zelus bruneri was not physically examined, but the rather slender body form as seen in the original illustration places it within this group. Comparative views of male genitalia are shown in Fig. 6.

6. Zelus renardii species group.

Zelus cervicalis Stål, 1872 and Zelus renardii Kolenati, 1856.

The two members of this group are very likely sister species since they share a number of unique characters: the apex of the medial process is greatly bent ventrad and hooklike, the lateral margin of the dorsal phallothecal sclerite is recurved dorsad and the basal part of the strut is absent. Both species are mainly distributed in North and Central America, but Z. cervicalis extends to northern South America. Comparative views of male genitalia are shown in Fig. 7 .

7. Zelus armillatus species group.

Zelus amblycephalus sp. n., Zelus annulosus (Stål, 1866), Zelus armillatus (Lepeletier & Serville, 1825), Zelus conjungens (Stål, 1860), Zelus janus Stål, 1862, Zelus leucogrammus (Perty, 1833), Zelus lewisi sp. n., Zelus litigiosus Stål, 1862, Zelus ruficeps Stål, 1862, Zelus sulcicollis Champion, 1899, Zelus umbraculoides sp. n. and Zelus umbraculus sp. n.

This is one of the two largest groups in the genus (the other being the Zelus panamensis species group). Species in this group are generally robust and large-sized (15-25 mm), and some are among the largest in the genus. The most distinctive character is that of the medial process, which has the apex slightly projected into two minute small lateral prongs or processes. This condition is different from that in several species groups listed below, where the apex of the medial process is hook-like and more strongly projected. The lateral spine of the humeral angle tends to be pronounced and somewhat broadened into a dentate effect. The pygophore is large, rounded, and somewhat shortened relative to the total length of the individual. The dorsal phallothecal sclerite having dorsolateral expansions or projections close to the basal arm is also unique to some species of this group. This condition, however, is not seen in Z. amblycephalus, Z. umbraculus, or Z. umbraculoides, which appear to be divergent from the remainders of the group, but the features of the medial process unambiguously place them in this species group. Comparative views of male genitalia are shown in Figs 8, 9.

8. Zelus longipes species group.

Zelus bahiaensis sp. n., Zelus errans Fabricius, 1803, Zelus longipes (Linnaeus, 1767), 1803 and Zelus vespiformis Hart, 1987.

This and the next species group (Zelus vagans species group) possess dense, spine-like setae on the head and pronotum, and a rounded, unarmed humeral angle, both characters rather unique in Zelus and probably synapomorphies uniting the two groups. The former character is possibly homoplastic as it is also seen in two species in the Zelus armillatus species group. The medial process is slender and cylindrical and this condition is among the most extreme in the genus. It is semi-erect and posteriorly directed. The paramere exceeds the apex of the medial process. The dorsal phallothecal sclerite has subparallel margins and lacks obvious modifications or ornamentations (except for small lateral folds in Z. longipes). Some individuals of Z. errans and Z. vespiformis appear to be wasp mimics. Comparative views of male genitalia are shown in Fig. 10.

9. Zelus vagans species group.

Zelus aithaleos, Zelus championi sp. n., Zelus fuliginatus sp. n., Zelus gracilipes sp. n. and Zelus vagans Fabricius, 1803.

Species of the Zelus vagans group share two characters also present in the preceding group (Zelus longipes species group): spinelike setae and rounded humeral angle. However, they differ in the structure of the male genitalia in significant ways. The medial process shapes like a somewhat laterally flattened cone. It is relatively broad at base, narrowing toward the apex, and is laterally compressed. The medial process is posteriorly directly, nearly horizontal. The paramere is removed from or barely reaching apex of the medial process. Furthermore, the phallus is elongated and slightly constricted toward the apex (not conspicuous in Z. gracilipes). Zelus vagans and Z. gracilipes also resemble wasps to some extent, but both not as perfectly as seen in Z. errans and Z. vespiformis. Comparative views of male genitalia are shown in Fig. 11.

10. Zelus panamensis species group.

Zelus banksi sp. n., Zelus cordazulus sp. n., Zelus filicauda Bergroth, 1893, Zelus gilboventris sp. n., Zelus korystos Hart, 1986, Zelus nigromaculatus Champion, 1899, Zelus panamensis sp. n., Zelus truxali sp. n., Zelus varius (Herrich-Schaeffer, 1853) and Zelus xouthos sp. n.

This is another large group with ten species. Interestingly, most (seven) are new species. It is characterized by having an acute apical modification usually in the shape of a hook on the medial process and the conspicuous medial carination of the apical part of the dorsal phallothecal sclerite. The condition of the apical modification of the medial process differs from that in the Zelus armillatus species group in that it is much more prominent, usually acute and sometimes extending further ventrally. Rugulosity of the posterior pronotal lobe is highly pronounced relative to the other groups. Sexual dimorphism is pronounced in some species in this group (e.g., Z. gilboventris and Z. truxali). Most species in this group are concentrated in southern Central America and northern South America. Comparative views of male genitalia and habitus images are in Figs 12, 13.

11. Zelus erythrocephalus species group.

Zelus auralanus sp. n., Zelus casii sp. n., Zelus chamaeleon Stål, 1872, ​Zelus erythrocephalus Fabricius, 1803, Zelus kartabenoides sp. n., Zelus kartabensis Haviland, 1931, Zelus laticornis (Herrich-Schaeffer, 1853), Zelus mattogrossensis Wygodzinsky, 1947, Zelus paracephalus sp. n., Zelus russulumus sp. n. and Zelus versicolor (Herrich-Schaeffer, 1848).

Two diagnostic characters identify members of this group. The medial process possesses a broad ridge-like projection or carina that initiates from the apex and extends ventrally or is removed from apex. The second feature is the apically oriented lateral sharp processes or projections on the dorsal phallothecal sclerite. These are not to be confused with the lateral expansion seen in the Zelus armillatus species group, where the direction of the expansion is laterad. In Z. auralanus and Z. versicolor, this process is short and somewhat dorsally directed, rather than apically directed. Three species, Z. kartabenoides, Z. kartabensis and Z. chamaeleon lack this structure. Their placement in this group is primarily based on the configuration of the medial process and the absence of characters of other groups. Also, the longitudinal ridge-like elevation or hook on the medial process is similar to the condition in another species, Z. laticornis, although the latter has a short modification. In this species group the parameres are usually somewhat bulbous and curved medially with moderate to long erect setae on the apical 1/2. The medial process is broadened at base, and usually anteroposteriorly compressed. Furthermore, the basal plate of the phallus is strongly curved in some members of this species group. Pronounced sexual dimorphism is seen in some species of this group. Notably, three species, Z. erythrocephalus, Z. paracephalus and Z. russulumus have purple, blue or greenish iridescence on the membrane of the hemelytron. Species of this group show a predominant southern South American distribution, with a few found only from the Amazons. Comparative views of male genitalia and habitus images are in Figs 14, 15.

Because of the heavy emphasis on male genitalic characters for grouping species, four species described only from females are not placed in any of the species groups defined in the above. These are: Zelus fasciatus Champion, 1899, Zelus plagiatus (Signoret, 1852), Zelus sphegeus Fabricius, 1803 and Zelus means Fabricius, 1803. Zelus fasciatus is similar to the females of some of the species in the Zelus panamensis species group and also occurs in an overlapping geographical region (southern Central America). Zelus plagiatus and Z. sphegeus show resemblance to the females of Z. versicolor, which is in the Zelus erythrocephalus species group. Zelus means, by possessing a rounded humeral angle and spinelike setae, aligns most closely with the Zelus vagans species group and the Zelus longipes species group. A future cladistic analysis, including morphological and molecular data, is needed to test the monophyly of these species groups and may also have the potential to place these female-based species.

Five species are removed from Zelus: Z. araneiformis, Zelus gradarius Bergroth, 1905, Z. modestus (Stål, 1862), Zelus subfasciatus Stål, 1860 and Zelus vittaticeps Stål, 1866. These species represent an undescribed genus "Hartzelus" and will be treated in a separate study. They will be listed as Harpactorini incertae sedis until their generic placement is formally clarified.

Figs 16, 17, 18

Male: (Fig. 16) Medium-sized, total length 13.47 mm (n=1, Suppl. material 2); slender. COLORATION: Entirely dark, nearly black; inconspicuous, light-colored, thin, medial longitudinal stripe on postocular lobe. Membrane of hemelytron semi-translucent. VESTITURE: Densely setose. Dorsum of anteocular lobe with moderately dense, short, recumbent and sparse, short , erect, spine-like setae. Dorsum of postocular lobe nearly glabrous; spine-like setae anteriorly between eyes; stripe of longitudinal whitish recumbent setae laterally. Ventral surface of head with moderately dense, recumbent setae, intermixed with erect setae. Scape nearly glabrous. Pronotum with dense, short, erect, stout, spine-like setae, also on lateral surfaces and pleura; scutellum with dense, apically curved, stout setae. Legs with sparse setation. Sundew setae on profemur sparse. Abdomen with moderately dense, short, semi-erect, fine setae. Ventral surface of pypophore with sparse, long, erect setae; posteroventral rim with long, erect setae; Paramere apically with dense, short to long, erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.30. Postocular lobe short; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye moderately sized; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head. Labium: I: II: III = 1: 1.7: 0.5 . Basiflagellomere diameter slightly larger than that of pedicel. Thorax: Anterolateral angle rounded, without projection; medial longitudinal sulcus evident throughout, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle rounded, without projection. Scutellum long; apex angulate, slightly projected upward. Legs: Very slender. Hemelytron: Greatly surpassing apex of abdomen by about 3x length of abdominal segment seven; quadrate cell large and broad; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 17) Pygophore: Elongate ovoid; mid-lateral fold adjacent to paramere insertion; slightly expanded laterally near base of paramere in dorsal view. Medial process somewhat laterally compressed, cone-shaped; anterior and posterior surfaces angulate medially; long, nearly as long as paramere; posteriorly directed, nearly horizontal; basally slightly curved; apex in posterior view blunt, without modification. Paramere: Cylindrical; moderately long, not reaching apex of medial process; directed posteriad, slightly curved towards medial process; nearly straight; apical part slightly enlarged, depression along inner side. Phallus: Dorsal phallothecal sclerite elongated; slightly constricted near middle; apical 1/3 of phallothecal sclerite tapering to apex, dorsal surface strongly convex; apex medially notched; posterior margin of foramen deeply concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally separate. Basal plate arm extremely slender; separate; subparallel; bridge short; extension of basal plate well developed, only slightly expanded laterally.

Female: Similar to male, except for the following. Larger than male, total length 13.87–17.61 mm (mean 16.27 mm, Suppl. material 2). Abdomen expanded beyond margins of wings. Metafemoral diameter smallest, mesofemoral diameter significantly larger than that of profemur. Occasional specimens with orange posterior pronotal lobe and mesopleuron. Setae on some specimens golden.

The nearly colorless cells of the membrane of the hemelytron contrast markedly with the dark veins, making Z. aithaleos an easily recognizable species in this genus. Also recognized by the following combination of characters: the postocular lobe short, 1.7x of the length of anteocular lobe in males and 1.2x in females; the anterior pronotal lobe short, abbreviated; the pronotum strongly convex; the humeral angle of pronotum rounded, unarmed; the cranium, the pronotum, the pleura and the scutellum with spinelike, short, stout setae (the last two characters also seen in the Zelus longipes species group and the Zelus vagans species group).

Males can also be recognized by the medial process laterally compressed, posteriorly directly and almost horizontal (also seen in the Zelus vagans species group). Within the Zelus vagans species group (Fig. 11), the medial process of Z. aithaleos is comparatively long, exceeding 1/2 length of the main body of the pygophore, whereas all other species in this group have the medial process less than 1/2 length of the pygophore. The basal plate arm is remarkably more slender than those in the same species group.

A unicolourous near-black dorsum, including the head, the pronotum and the corium, separates Z. aithaleos from both sexes of Z. gracilipes, Z. vagans, and Z. means (known from females only), all of which have some orange, yellow or reddish colors. The dark dorsal profile is shared with Z. championi (only the male is known) and Z. fuliginatus. A longitudinal lateral patch of whitish recumbent setae on the postocular lobe serves to separate this species from Z. fuliginatus. It is distinguished by a dark abdomen from Z. championi, which has a brightly red abdomen.

From Greek aithales.

South America (Fig. 18). Countries with specimen records: Bolivia, Brazil, Paraguay and Peru.

Figs 19, 20, 21

Male: (Fig. 19a, b) Large, total length 15.0–17.5 mm (mean 15.7 mm, Suppl. material 2); slender, body length/width = 4.5. COLORATION: Yellowish or greenish-brown. Head yellowish-brown to brown. Antennae and femoral apices reddish. Anterior pronotal lobe uniformly yellowish-brown or brown. Posterior pronotal lobe yellowish-brown or brown with lateral processes and surrounding area darker, brown to dark brown. Clavus and corium brown, veins yellowish-brown, membrane brown. Abdomen pale brown. Pygophore yellowish-brown. VESTITURE: Sparsely setose; sparse, short, erect setae over most of integument. Short spinelike setae on dorsal surface of head, with some short recumbent setae dorsally on posterior lobe, sparse short recument and erect setae on lateral and ventral surfaces. Anterior pronotal lobe with short, spine-like setae dorsally and short, erect and recumbent setae laterally; posterior lobe with short, recumbent and erect setae. Scutellum with short erect and recumbent setae. Corium and clavus with short, recumbent setae. Microtrichia throughout posterior margin of membrane of hemelytron. Abdominal venter with short, erect setae, interspersed with long setae. Sparse, moderately long setae on apical 1/2 of paramere. STRUCTURE: Head: Cylindrical. In dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower; dorsal outline in lateral view gradually sloping. Eye prominent; dorsal and ventral margins removed from outlines of head. Labium: I: II: III = 1: 1.4: 0.3. Segment I surpassing anterior margin of eye. Antenna: Basiflagellomere diameter larger than that of pedicel. Thorax: Anterior pronotal lobe with indistinct collar, anterolateral angle rounded, without projection; medial longitudinal sulcus evident throughout, deepening posteriorly; slightly raised inconspicuous protuberance laterad to longitudinal medial sulcus anterior to transverse pronotal sulcus, more apparent in lateral view. Posterior pronotal lobe with smooth surface; disc distinctly elevated above humeral angle, slightly convex; humeral angle slightly expanded and wider than abdomen, armed with spinous process; margin indistinct, convex. Scutellar apex angulate, not projected. Legs: Moderately robust. Hemelytron: Surpassing apex of abdomen by about length of abdominal segment seven; costal margin somewhat concave; Sc surpassing level of apex of cubital cell; quadrate cell small and slender; 1A and Pcu intersecting; Cu and M of cubital cell subparallel. Abdomen: Segments of sub-equal sizes; segment seven much shorter than preceding segments, posterior margin in lateral view slightly concave. GENITALIA: (Fig. 20) Segment eight with nearly straight posterior margin, slightly concave in middle. Pygophore: Ovoid; posterolateral rim in lateral view straight above paramere, concave below paramere. Slender. Medial process cylindrical, slender; long; posteriorly directed, in less than forty-five degree with body axis; nearly straight; apex rounded. Paramere: Cylindrical; long, surpassing medial process; directed posteriad; not distinctly curved; apical part very slightly enlarged. Phallus: Surface flat; laterally indistinctly angulate; apex truncate; posterior margin of foramen broadly concave; basal arm short. Struts attached to dorsal phallothecal sclerite; basally separate. Basal plate arm robust; separate; basally converging; in lateral view nearly straight, very slightly curved; bridge moderately long, slender; extension expanded and extended onto arm.

Female: (Fig. 19c, d) Similar to male, except for the following. Pleura and abdominal segments with patches of whitish exudation. Basiflagellomere diameter smaller or subequal to that of pedicel. Hemelytron barely surpassing apex of abdomen.

Can be recognized by the uniform pale coloration, the unpatterned legs (Fig. 19), and the relatively large size (>15mm, Suppl. material 2). Males can also be recognized by the apex of medial process with two minute prongs; the long, somewhat recurved paramere, which, viewed laterally, is at least 1.5x length of the medial process; and the dorsal phallothecal sclerite without lateral expansion close to the basal arm.

Among species of the Zelus armillatus group (Fig. 8), only Z. annulosus also possesses a paramere much longer than the medial process, but the two can be easily separated by the general aspects of coloration. Both in general appearance and in the appearance of certain characters of the male genitalia, this species appears to be most closely related to Z. umbraculus and Z. umbraculoides, which have short parameres and are known only from male specimens.

Southern Mexico to northern South America and part of Brazil (Fig. 21). Countries with records: Brazil, Colombia, Costa Rica, Ecuador, Mexico, Panama and Suriname.

Zelus ambulans Stål, 1862, p. 451, orig. descr.; Stål, 1872, p. 91, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 259–260, Tab. XV. fig. 23, 23a, junior syn. of Z. exsanguis; Maldonado, 1990, p. 327. cat. and junior syn. of Z. exsanguis. stat. rev. (current study).

Diplodus ambulans: Uhler, 1886, p. 24, checklist; Walker, 1873, cat.

Figs 22, 23, 24

Male: (Fig. 22a, b) Medium-sized, total length 12.89–15.19 mm (mean 14.27 mm, Suppl. material 2); slender. COLORATION: Dorsal surface generally brown. Anteocular lobe yellowish-brown to light reddish with darker brown areas on lateral surfaces between compound eyes and antennal insertions, some specimens with dark brown areas on posterodorsal surface. Dorsal surface of postocular lobe dark brown with wide yellowish-brown mid-dorsal and circumocellar areas, remainder of surface yellowish-brown. Rostrum yellowish-brown to reddish-brown, some specimens with segment I and apex of segment II darker reddish-brown. Antennal segments I and II with varying dark brown areas at base and apex, remainder of I and II yellowish-brown to dark reddish-brown, III and IV dark reddish-brown. Anterior lobe yellowish-brown with varying dark brown areas on dorsolateral margins, anterolateral angles of collar, medial sulcus, and small patches at posterodorsal margin. Posterior lobe yellowish-brown with posterior 1/2, except for posterior margin, darkening brown in some specimens. Scutellum yellowish-brown to dark brown. Legs yellowish-brown, apical 1/5 of femora with brown to brownish-black band and apex of tibiae darkening to dark reddish-brown. Hemelytron brown with yellowish-brown costal margins and veins of clavus and corium yellowish-brown except for veins bounding discal cells. Abdominal venter yellowish-brown. VESTITURE: Moderately setose. Short recumbent setae predominating dorsally; short to moderate erect setae over entire body. Recumbent and erect setae over entire sure of head, recumbent setae predominating dorsally. Postocular lobe with recumbent and scattered erect setae over entire surface, recumbent setae more dense dorsally, erect setae more dense lateroventrally. Anterior pronotal lobe with erect and recumbent setae confined to setal tracts dorsally, erect setae laterally. Posterior lobe vestiture consisting of recumbent and scattered erect setae over entire surface. Scutellum with moderate to long, silky setae. Clavus and corium with short recumbent setae, longer near base of clavus. Abdomen with short, stiff, erect setae dorsally, remainder of surface with recumbent and scattered erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.30. Postocular lobe moderately long, posterior 1/2 width constant, lateral margins subparallel. Eey moderately prominent; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head. Ocellus only slightly elevated. Labium: I: II: III = 1: 1.8: 0.4. Thorax: Anterolateral angle with inconspicuous subtuberculate projection; medial longitudinal sulcus evident only on posterior 1/2, deepening to transverse sulcus of pronotum. Posterior pronotal lobe with finely rugulose surface; disc about same level of, and continuous with, humeral angle; humeral angle armed, with spinous process. Scutellum short; apex blunt to subtuberculate. Legs: Slender. Femoral diameters subequal. Hemelytron: Surpassing apex of abdomen by about length of abdominal segment seven; quadrate cell small; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 23) Pygophore: Elongate ovoid; not expanded laterally in dorsal view. Medial process triangular, relatively broad, moderately long, semi-erect, nearly straight, curved slightly posteriad apically; apex in posterior view blunt, without modification. Paramere: Cylindrical; moderately long, nearly reaching apex of medial process; slightly curved ventrad; apical part enlarged. Phallus: Dorsal phallothecal sclerite elongated, somewhat flattened; medially slightly constricted; apical portion of phallothecal sclerite not distinctly tapered, slightly convex; apex truncate, medially emarginate; posterior margin of foramen broadly concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally separate. Basal plate arm moderately robust, separate, converging, in lateral view nearly straight, very slightly curved; bridge short; extension of basal plate small and confined to apex of basal plate arm.

Female: (Fig. 22c) Similar to male, except for the following. Larger than male, total length 15.43–18.59 mm (mean 16.55 mm, Suppl. material 2). Generall coloration slightly lighter; legs more or less uniformly colored, apices somewhat reddish, without dark bands.

Among the species of Zelus luridus group, Z. ambulans has the humeral angle elevated to level of, and continuous with, disc of the posterior pronotal lobe, a condition that is also present in Z. exsanguis, but it can be separated from that species by the yellowish veins on corium, contrasting to the brown corium, whereas the entire corium is more or less uniformly colored in Z. exsanguis.

Among species of the Zelus luridus species group (Fig. 3) males of Z. ambulans can be recognized by the relatively slender medial process (Fig. 23a) and the paramere barely reaching the medial process. The apical enlargement of the paramere is smaller than that in Z. spatulosus and Z. exsanguis, but larger than that in Z. grandoculus, Z. luridus and Z. antiguensis.

North and Central America (Fig. 24). Countries with records: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panama.

Champion (1898) synonymized Z. ambulans, Z. luridus and Z. cognatus under Z. exsanguis. Hart (1986) recognized all three as valid species, but did not formally reinstate Z. ambulans, probably because it is outside the geographic focus of that particular study. Zelus ambulans remained a synonym of Z. exsanguis in Maldonado (1990)'s catalogue or Reduviidae. We here resurrect Z. ambulans from synonymy. Champion (1898)'s figures of Z. exsanguis actually depict Z. ambulans.

Although this species shows very little morphological variations, color patterns within an area do vary considerably. The dark area at the posterior margin of the longitudinal medial sulcus of the anterior lobe, which serves to easily distinguish Z. ambulans from Z. exsanguis, is relatively constant. Other colors, specifically that of the posterior pronotal lobe and the femoral apices vary from quite light to very dark brown in any given locality. There is also an occasional specimen with somewhat darker hemelytron, but this does not show the wide range of variations of the aforementioned characters.

Most specimens examined have been collected from moderate to high altitudes.

Diplodus annulosus Stål, 1866, p. 299, orig. descr.; Walker, 1873, p. 126, cat.

Zelus annulosus: Stål, 1872, p. 92, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Fracker and Bruner, 1924, p. 170, note; Wygodzinsky, 1949a, p. 48, checklist; Maldonado, 1990, p. 326, cat.

Figs 25, 26, 27

Male: (Fig. 25) Large, total length 14.57 mm (n=1, Suppl. material 2); very slender. COLORATION: Yellowish with dark brown patches; green on posterior pronotal lobe and corium. Most surface of head yellowish, dark stripe between eye and antennal insertion, on postocular lobe behind ocellus, and on lateral surface. Scape dark brown with three yellowish bands. Labium yellowish, dark band on first and second segments. Anterior pronotal lobe yellowish, anterior medial brown patch, anterolateral angle dark brown, connected to dark brown patch on lateral surface. Posterior pronotal lobe, anterior part of corium green; rest of hemelytron brown to dark brown. Pleura yellowish with dark brown patch. Femora and tibiae with alternating yellow and dark brown bands, six of each on femora, four of each on tibiae, yellow band smaller, more so on tibiae. VESTITURE: Moderately setose. Entire dorsal surface, including corium and clavus, with dark, dense, short to moderately long, erect, spine-like setae. Ventral surface of head, pleura with short, semi-erect to recumbent setae. Abdomen with moderately dense, short, semi-erect to recumbent setae, intermixed with long, erect setae. Sundew setae on profemur sparse. STRUCTURE: Head: Elongated, L/W = 2.13. Postocular lobe very long; in dorsal view distinctly narrowing through anterior 1/2, posterior 1/2 constant, tube-like. Eye prominent; lateral margin much wider than postocular lobe; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head. Labium: I: II: III = 1: 1.5: 0.4. Basiflagellomere diameter subequal to that of pedicel. Thorax: Anterolateral angle bearing small protuberance; medial longitudinal sulcus shallow near collar, deepening posteriorly. Posterior pronotal lobe with finely rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with short tuberculate processes. Scutellum moderately long; apex angulate. Legs: Slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell small and slender; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 26) Pygophore: Ovoid; not expanded laterally in dorsal view; broad lightly sclerotized expansion between paramere and medial process. Medial process expanded laterally; short; semi-erect; basally slightly protruding; apex in posterior view truncate, with small sharp lateral projections. Paramere: Cylindrical; long, nearly reaching apex of medial process; directed toward medial process; apically recurved. Phallus: Dorsal phallothecal sclerite shield-shaped; sharp, dorsad projection arising close to base; apical portion of phallothecal sclerite not distinctly tapered, flat, laterally distinctly angulate, ridge-like; apex truncate, not emarginate; posterior margin of foramen broadly inversely v-shaped. Struts attached to dorsal phallothecal sclerite; apically separate, not connected by bridge; basally fused. Basal plate arm moderately robust; separate; converging; in lateral view slightly curved; bridge moderately long; extension of basal plate expanded onto arm.

Female: Similar to male, except for the following. Larger than male, total length 21.19–22.72 mm (mean 21.91 mm, Suppl. material 2). Some dry-preserved specimens have posterior pronotal lobe and corium not green but brown, probably a result of preservation artifact.

Recognized by the following combination of characters: the posterior pronotal and corium dark green; the legs with four to five alternative yellow and black bands; the head, pronotum, scutellum and corium with moderately dense, black, erect, spine-like setae; the rather long and slender legs, the profemur 1/2 of body length; the rather long postocular lobe, enlarged at posterior 3/4; and the quadrate cell on corium rather slender, length more than 2x width.

Males can also be recognized by the long paramere, reaching apex of medial process; the apex of paramere recurved; the medial process apically with two lateral sharp projections; the membranous sclerite between paramere and medial process, not distinctly protruding posteriorly; and the dorsal phallothecal sclerite with lateral expansion close to basal arm, sharp, dorsad.

South America (Fig. 27). The Colombian and Brazilian Amazonia and Frech Guiana. Countries with records: Brazil, Colombia, French Guiana.

Figs 28, 29, 30

Male: (Fig. 28a, b) Medium-sized, total length 13.69–16.28 mm (mean 14.98 mm, Suppl. material 1); slender. COLORATION: Dorsal surface of anteocular lobe reddish-brown, yellowish-brown ventrally. Dorsum of postocular lobe dark brown with yellowish-brown mid-dorsal line and circumocellar areas, ventral surface yellowish-brown. Rostrum light reddish-brown. Scape and pedicel light reddish-brown with dark brown areas near base and apex. Anterior pronotal lobe reddish-brown with yellowish-brown anteroventral area. Dorsal surface of posterior lobe reddish-brown with yellowish-brown lateral and posterior margins, humeral angle dark brown and lateral surfaces yellowish-brown. Scutellum yellowish-brown to reddish-brown. Legs yellowish-brown to reddish-brown, femoral and tibial apices darker reddish-brown. Hemelytron brown, veins of clavus and corium slightly lighter in color than surrounding areas. Abdomen with dorsal surface reddish-brown, ventral surface yellowish-brown. VESTITURE: Moderately setose. Anteocular lobe with recumbent setae over dorsal surface, some erect setae ventrally. Postocular lobe with recumbent setae predominating dorsally, long silky erect setae posterodorsally and over lateral surface. Anterior pronotal lobe with recumbent and short erect setae on faint setal tracts dorsally and scattered on lateral surfaces, long silky erect setae on anterior margins. Posterior pronotal lobe with short recumbent and erect setae over entire surface, some long erect setae mid-dorsally on anterior half. Scutellum with moderate to long setae. Corium and clavus of hemelytron with recumbent setae. Abdomen with sparse, short, erect setae over entire surface, some short recumbent and longer erect setae on lateral and ventral surfaces, moderate to long erect setae posteroventrally on segment seven. Pygophore with short to moderate setae over exposed surface. STRUCTURE: Head: Cylindrical, L/W = 2.29. Postocular lobe relatively short; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye moderately sized; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head. Ocellus greatly elevated. Labium: I: II: III = 1: 1.8: 0.6. Basiflagellomere diameter larger than that of pedicel. Thorax: Anterolateral angle rounded, without projection; medial longitudinal sulcus shallow near collar, deepening slightly in posterior half. Posterior pronotal lobe with finely rugulose surface; disc slightly elevated above humeral angle; humeral angle armed, with short tuberculate processes. Scutellum moderately long; lateral depressions deep; apex slightly produced. Legs: Slender. Pro- and mesofemoral diameters subequal, metafemoral diameter slightly smaller. Hemelytron: Surpassing apex of abdomen by about length of abdominal segment seven; quadrate cell small, elongate; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 29) Pygophore: Elongate ovoid; lateral margin above paramere insertion slightly expanded laterally in dorsal view. Medial process triangular, broad, short, erect; nearly straight; apex in posterior view blunt, without modification. Paramere: Cylindrical over basal half, slightly compressed and enlarged dorsoventrally over apical half; moderately long, nearly reaching medial process; not distinctly curved; apical part very slightly enlarged. Phallus: Dorsal phallothecal sclerite somewhat squarish; apex blunt, medially very slightly emarginate, not distinctly tapered; surface flat; posterior margin of foramen broadly concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally mostly separate, moderately fused. Basal plate arm slender; separate; converging; in lateral view nearly straight, very slightly curved; bridge moderately long; extension of basal plate small and confined to apex of basal plate arm.

Female: (Fig. 28c, d) Similar to male, except for the following. Larger than male, total length 15.77–18.15 mm (mean 16.91 mm, Suppl. material 2). Coloration very similar to that in male, slightly lighter overall. Anteocular lobe varying from reddish-brown to brown dorsally, legs of some specimens unicolorous. Ocellar elevation not pronounced; middle of mesothoracic femora slightly swollen.

As with several members of the Zelus luridus species group, the coloration is greenish-brown, rather uniform. The medial process is triangular, its base distinct from the rest of pygophore ventral rim and apex without modification. Can be distinguished from males of other species of the Zelus luridus species group (Fig. 3) by the base of the medial process extended, the apex of the paramere not greatly enlarged, and the phallothecal sclerite rather short.

Zelus antiguensis is similar in appearance to Z. luridus and might easily be confused with that species. The comparatively broader medial process and posterior protrusion of the base of the medial process are readily evident in Z. antiguensis (Fig. 3). This species also shows an internal folding of the dorsolateral apical areas of the dorsal phallothecal sclerite, such folding being absent in Z. luridus. Generally, in both sexes the head is more pubescent and the pronotum more flattened dorsally than is normally found in Z. luridus. These two species do not overlap in distribution.

Named after the type locality, Antigua, in Guatemala.

Southern Mexico and Guatemala (Fig. 30).

Reduvius armillatus Lepeletier and Serville, 1825, p. 278, orig. descr.

Diplodus armillatus: Amyot and Serville, 1843, p. 370, descr.; Stål, 1860, p. 75, list; Walker, 1873, p. 123, cat.

Euagoras armillatus: Herrich-Schaeffer, 1853, p. 91, list.

Zelus armillatus: Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Wygodzinsky, 1949a, p. 49, checklist; Mayr, 1866, p. 138-139, senior syn. of Z. brasiliensis, Z. aurantiacus, Z. guttifer and Z. conjungens; Berg, 1879, p. 151-152, list and nymphs (subgenus Diplodus); Costa Lima, 1940, 218, list (subgenus Diplocodus); Wygodzinsky, 1957, p. 268, note; Wygodzinsky, 1960; p. 307, list; Maldonado, 1990, p. 326, cat.; Van der Heyden, p. 85-90, new record (misidentification, should be Zelus atripes).

Reduvius brasiliensis Lepeletier and Serville, 1825, p. 278, orig. descr.

Diplodus brasiliensis: Amyot and Serville, 1843, p. 370, descr.; Stål, 1860, p. 75, note; Mayr, 1866, p. 138- 139, junior syn. of Z. armillatus; Walker, 1873, p. 123, cat.

Euagoras brasiliensis: Herrich-Schaeffer, 1853, p. 91, list.

Zelus brasiliensis: Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, junior syn. of Z. armillatus.

Arilus aurantiacus Herrich-Schaeffer, 1848, p. 35-36 Tab. CCLXI. fig. 809, orig. descr. and fig.; Mayr, 1866, p. 138-139, junior syn. of Z. armillatus; Stål, 1872, p. 90, junior syn. of Z. armillatus.

Euagoras aurantiacus: Herrich-Schaeffer, 1853, p. 91, list (aurantius (sic)).

Ploeogaster aurantiacus: Herrich-Schaeffer, 1853, p. 168, list.

Arilus guttifer Herrich-Schaeffer, 1848, p. 36, Tab. CCLXI, fig. 810, orig. descr. and fig.; Mayr, 1866, p. 138-139, junior syn. of Z. armillatus; Stål, 1872, p. 90, junior syn. of Z. brasiliensis.

Euagoras guttifer: Herrich-Schaeffer, 1853, p. 92, list.

Ploeogaster guttifer: Herrich-Schaeffer, 1853, p. 168, list.

Diplodus guttifer: Stål , 1860, p. 74, descr.; Walker, 1873, p. 126, cat.

Zelus guttifer: Stål 1862, p. 453, note.

Arilus guttifer , Mayr, 1866, p. 138-139, junior syn. of Z. armillatus .

Figs 31, 32, 33, 34, 35

Male: (Fig. 31) Large, total length 17.15–19.02 mm (mean 17.87 mm, Suppl. material 2); robust. COLORATION: Highly variable, with varying combinations and amounts of yellow, yellowish-brown and brownish-black; margins of posterior pronotal lobe and corium yellowish, rest brownish-black as most common pattern, amount of black varies, sometimes almost entirely black; legs uniformly black or apically reddish, yellow-black banded in some specimens. Abdominal dorsal surface dark brown, segments with yellowish-brown posterior and lateral margins; lateral and ventral surfaces dark brown to brownish-black with lighter mid-ventral line or yellowish-brown with variable darker areas. Pygophore yellowish-brown to brownish-black, pattern variable. VESTITURE: Densely setose. Short recumbent and short to long erect setae over entire surface of head. Anterior pronotal lobe with short recumbent and short to long erect setae on lateral surface, short to long erect setae confined to tracts dorsally; posterior lobe with recumbent to erect setae on lateral surface, erect setae on dorsal surface; scutellum with dense moderate to long semi-erect to erect setae, denser on apex. Corium and clavus with short, recumbent or erect setae. Scattered short erect setae on abdominal dorsum, lateral and ventral surfaces with short recumbent setae interspersed with erect setae of varying lengths. Recumbent and erect setae on exposed surface of pygophore; apical 1/3 of parameres with erect setae on dorsal surface. STRUCTURE: Head: Cylindrical, L/W = 2.40. Postocular lobe relatively short; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye moderately sized; dorsal and ventral margins removed from outlines of head. Labium: I: II: III = 1: 1.5: 0.4. Basiflagellomere diameter subequal to that of pedicel. Thorax: Anterolateral angle rounded, without projection; medial longitudinal sulcus evident throughout, deepening posteriorly. Posterior pronotal lobe with finely rugulose surface; disc slightly elevated above humeral angle; humeral angle armed, with spinous processes. Scutellum short; apex rounded, not projected. Legs: Robust. Hemelytron: Surpassing apex of abdomen by about length of abdominal segment seven; quadrate cell large and broad; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 34) Pygophore: Ovoid; posteriorly expanded sac-like sclerite between parameres and medial process. Medial process cylindrical; slender; moderately long, almost as long as exposed part of parameres; posteriorly directed, in less than forty-five degree with body axis; nearly straight; basally without protrusion; apex in posterior view rounded, with minute projection. Paramere: Cylindrical; long, not reaching apex of medial process; directed posteriad; not distinctly curved; apical part not enlarged to very slightly enlarged. Phallus: Sharp laterally oriented process close to posterior margin of foramen and basal arms; apical portion of phallothecal sclerite not distinctly tapered, flat, laterally angulate; apex truncate; posterior margin of foramen broadly concave. Struts attached to dorsal phallothecal sclerite; apically separate, not connected by bridge; basally mostly separate, moderately fused. Basal plate arm robust; separate; converging; in lateral view nearly straight, very slightly curved; bridge short; extension of basal plate expanded onto arm.

Female: (Figs 32, 33) Similar to male, except for the following. Larger than male, total length 19.0–24.7 mm (mean 21.94 mm, Suppl. material 2). Coloration variations more extensive than in male.

The large and robust body, the dorsal coloration usually bright, yellow or red with black, the medial process short and relatively slender are characteristic to Z. armillatus. Male genitalic structures of Z. armillatus and Z. janus are nearly identical, but these two species do not overlap in range and are sufficiently different in non-genitalic morphological characters, which allow them to be easily separated.

The only species with which Z. armillatus is sympatric which may cause some identification problems is Z. conjungens. It may be distinguished from that species by the characters discussed under Z. conjungens.

South America (Fig. 35). Countries with specimen records: Argentina, Bolivia, Brazil, Colombia, Ecuador, Guyana, and Paraguay.

Zelus armillatus is a very common, widespread, variable species in South America. It is known to occur in nearly all areas of the continent from central Argentina and northward, at altitudes from sea level to several thousand feet, and dry temperate to moist tropical areas. The coloration and markings of Z. armillatus are highly variable throughout the range and appear to be as variable in any given area (e.g., Fig. 33d) as they are between areas. This fact is responsible for the several descriptions based upon color forms of this species. The drawings of Herrich-Schaeffer (1848) illustrate two of the common variations encountered, although the dorsal coloration, as well as that of the legs, may vary from almost entirely yellowish-brown through various combinations of that color and brownish-black to almost entirely brownish-black.

Figs 36, 37, 38

Male: (Fig. 36a, b, c, d) Medium-sized, total length 12.64–14.37 mm (mean 13.61 mm, Suppl. material 2); slender. COLORATION: Nearly entire surface medium dark brown; apices of femora dark colored; abdomen light brown to yellowish-brown. VESTITURE: Densely setose. Body surface covered with short, adpressed to recumbent setae, dorsal setae on head and pronotum golden, shining; ventral surface of head and abdomen with longer, erect setae; sometimes setae covered with white waxy exudation. Dense, long, erect setae on paramere; bush of short, erect, spine-like setae on posterior surface of medial process. Corium and clavus with short, recumbent setae. STRUCTURE: Head: Cylindrical, L/W = 2.63. Postocular lobe long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye prominent; much wider than postocular lobe; dorsal and ventral margins removed from surfaces of head. Labium: I: II: III = 1: 2.0: 0.4. Basiflagellomere diameter slightly larger than that of pedicel. Thorax: Anterolateral angle bearing small projection; medial longitudinal sulcus shallow near collar, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc about same level as humeral angle; humeral angle armed, with dentate or spinous process. Scutellum long; apex blunt, not projected. Legs: Slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell small, relatively broad; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 37) Pygophore: Ovoid; slightly expanded laterally near base of paramere in dorsal view. Medial process triangular; slender; long; anteroposteriorly compressed; semi-erect; curved at middle; apex in posterior view acute, with small hooklike projection. Paramere: Cylindrical; moderately long, slightly exceeding medial process; directed posteriad, slightly curved towards medial process; basally narrower; curved ventrad; apical part very slightly enlarged. Phallus: Dorsal phallothecal sclerite somewhat pandurate, medially strongly constricted; laterally with dorsally directed small sharp projection at mid-portion; apical portion of phallothecal sclerite gradually tapering, lateral margin narrowly angulate, angulation ending anteriorly in sharp, dorsad projection; apex rounded, medially emarginate; posterior margin of foramen inversely V-shaped. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally almost completely fused. Basal plate arm moderately robust; separate; converging; in lateral view severely curved, nearly semi-circular; bridge short; extension of basal plate expanded onto arm.

Female: (Fig. 36e, f) Similar to male, except for the following. Larger than male, total length 17.21–18.32 mm (mean 17.85 mm, Suppl. material 2). Eye moderate size, smaller than in male.

Can be readily recognized by the uniformly brown dorsal coloration; the darkened tibial apex; the humeral angle elevated to level of disc; the dorsal setae on head and pronotum appearing somewhat golden, shining when viewed under magnification. Males can also be recognized by the gradually enlarged paramere; the triangular medial process, curved slightly posteriad in the middle, apex with a hooklike projection; and the dorsal phallothecal sclerite with short, dorsad projections sub-laterally.

The species epithet indicates the somewhat reddish tone of the coloration.

South America (Fig. 38). Countries with specimen records: Bolivia, Brazil, Ecuador and Peru.

Figs 39, 40, 41

Male: (Fig. 39) Medium-sized, total length 12.35 mm (n=1); slender. COLORATION: Much of body surface including head, anterior pronotal lobe, membrane, legs dark brown; very slender lighter colored medial longitudinal stripe on postocular lobe. Posterior pronotal lobe and corium orange. Pleura, abdomen reddish-brown. VESTITURE: Moderately setose. Dorsum of anteocular and anterior part of postocular with moderately dense, short, erect, spine-like setae, posterior part of postocular nearly glabrous; ventral surface of head with sparse, short, erect or recumbent setae. Pronotum with dense, short, erect, spine-like setae on dorsum and lateral surfaces, anterior lobe also intermixed with sparse, long, fine setae. Pleura with spine-like setae, sparse on metapleuron; intermixed with short to long, erect, fine and short, recumbent setae; scutellum with spine-like setae and erect, fine setae. Legs with very sparse setation. Corium and clavus with short, recumbent setae. Abdomen with moderately dense, short, semi-erect, fine setae, intermixed with sparse, long setae. Bush of moderately long, erect setae flanking medial process on posteroventral rim of pygophore; paramere apically with sparse, short, erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.27. Postocular lobe long; in dorsal view distinctly narrowing through anterior 2/3, posterior 1/3 constant, tube-like. Eye moderately sized; lateral margin much wider than postocular lobe; dorsal margin removed from postocular transverse groove, ventral margin attaining ventral surface of head. Labium: I: II: III = 1: 1.8: 0.6. Basiflagellomere diameter slightly larger than that of pedicel. Thorax: Anterolateral angle rounded, without projection; medial longitudinal sulcus evident throughout, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle rounded, without projection. Scutellum moderately long; apex slightly pointed, not projected. Legs: Very slender. Hemelytron: Greatly surpassing apex of abdomen by about 3x length of abdominal segment seven; quadrate cell large and broad; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 40) Pygophore: Ovoid; mid-lateral fold adjacent to paramere insertion; slightly expanded laterally near base of paramere in dorsal view. Medial process cylindrical; very slender; moderately long, nearly half length of paramere; posteriorly directed; basal 2/3 straight, apically curved; apex in posterior view blunt, folded posteriad, marginally narrower. Paramere: Cylindrical; long, surpassing medial process; directed posteriad; slightly curved ventrad; apical part very slightly enlarged. Phallus: Dorsal phallothecal sclerite elongated; apical portion of phallothecal sclerite not distinctly tapered, convex, laterally angulate; apex truncate, medially emarginate; posterior margin of foramen deeply concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally mostly separate, moderately fused. Basal plate arm slender; separate; converging; in lateral view apically curved; bridge short; extension of basal plate expanded onto arm.

Female: Unknown.

Recognized by the following combination of characters: the anterior pronotal lobe dark brown and the posterior pronotal lobe orange; the 1A an Pcu not intersecting, short crossvein between them; the long and slender, cylindrical medial process; the medial process apically folded posteriad; and the rather long paramere.

Named after the Brazilian state Bahia, where the holotype was collected.

South America (Fig. 41). Known from the type locality in Brazil.

Figs 42, 43, 44

Male: (Fig. 42) Medium-sized, total length 10.81–12.62 mm (mean 12.00 mm, Suppl. material 2); slender. COLORATION: Head uniformly brown; postocular lobe with very faint longitudinal medial stripe. Anterior pronotal lobe and hemelytron brown; posterior lobe yellowish-brown. Remainder of body surface mostly yellowish-brown, parts of pleura darker. Femora with two or three yellowish bands; tibiae with single band. VESTITURE: Sparsely setose. Short, recumbent setae on entire surface; very short, erect, spine-like setae on dorsum, denser on anterior lobe; few moderately long, erect, fine setae on ventral surface. Pronotum with sparse, recumbent setae and short, erect setae over dorsal surface; denser, long recumbent setae on lateral surface and pleura, intermixed with semierect or erect setae; scutellum with sparse, semi-erect and recumbent setae. Legs with sparse setation on femora and moderately dense setation on tibiae. Corium and clavus with mix of sparse, short, recumbent and erect setae. Abdomen with moderately dense, short recumbent setae, intermixed with sparse, short to long, erect setae. Apical half of dorsal surface of paramere with moderately dense, medium-length, semi-erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.25. Postocular lobe long; in dorsal view distinctly narrowing through anterior 2/3, posterior 1/3 constant, tube-like. Eye prominent; lateral margin much wider than postocular lobe; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head. Labium: I: II: III = 1: 2.1: 0.5. Basiflagellomere diameter larger than that of pedicel. Thorax: Anterolateral angle bearing small projection; medial longitudinal sulcus evident throughout, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with dentate or spinous process. Scutellum short; apex angulate, slightly projected upward in some specimens. Legs: Slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell small; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 43) Pygophore: Elongate ovoid; not expanded laterally in dorsal view. Medial process cylindrical; slender; moderately long, nearly as long as paramere; laterally compressed towards apex; anterior surface towards apex ridged; minute spicules on posterior surface; semi-erect; very slightly curved at middle; apex in posterior view acute, with small hooklike projection. Paramere: Cylindrical; long, achieving apex of medial process; directed posteriad, slightly curved towards medial process; basally slightly narrower; nearly straight; apical part slightly enlarged, obliquely truncate. Phallus: Dorsal phallothecal sclerite somewhat ovoid; sclerotization reduced (yet not absent) on dorsal surface close to posterior margin of foramen; apical portion of phallothecal sclerite gradually tapering, distinctly keeled medially, laterally indistinctly angulate; apex acute; posterior margin of foramen broadly concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally separate throughout. Basal plate arm robust; basally fused; in lateral view mid-portion curved; bridge extremely short; extension of basal plate expanded laterally onto arm, covering more than 1/2 of arm, curved.

Female: Unknown.