A taxonomic monograph of the assassin bug genus Zelus Fabricius (Hemiptera: Reduviidae): 71 species based on 10,000 specimens

Abstract The New World assassin bug genus Zelus Fabricius, 1803 (Insecta: Hemiptera: Heteroptera: Reduviidae: Harpactorinae: Harpactorini) is revised based on more than 10,000 specimens. Seventy-one species are recognized and twenty-four described as new: Zelus aithaleos sp. n., Zelus amblycephalus sp. n., Zelus antiguensis sp. n., Zelus auralanus sp. n., Zelus bahiaensis sp. n., Zelus banksi sp. n., Zelus casii sp. n., Zelus championi sp. n., Zelus cordazulus sp. n., Zelus fuliginatus sp. n., Zelus gilboventris sp. n., Zelus gracilipes sp. n., Zelus grandoculus sp. n., Zelus kartaboides sp. n., Zelus lewisi sp. n., Zelus panamensis sp. n., Zelus paracephalus sp. n., Zelus rosulentus sp. n., Zelus russulumus sp. n., Zelus spatulosus sp. n., Zelus truxali sp. n., Zelus umbraculoides sp. n., Zelus umbraculus sp. n., and Zelus xouthos sp. n. Five species, Zelus araneiformis Haviland, 1931, Zelus gradarius Bergroth, 1905, Zelus modestus (Stål, 1862), Zelus subfasciatus Stål, 1860 and Zelus vittaticeps Stål, 1866, are removed from Zelus and placed incertae sedis within Harpactorini. Nine new synonyms are recognized (senior synonym in parentheses): Zelus atripes Champion, 1898 syn. nov. (=Zelus conjungens [Stål, 1860]), Zelus dispar Fabricius, 1803 syn. nov. (=Zelus pedestris Fabricius, 1803), Zelus formosus Haviland, 1931 syn. nov. (=Zelus laticornis Herrich-Schaeffer, 1853), Zelus obscuridorsis (Stål, 1860) syn. nov. (=Zelus pedestris), Zelus pallidinervus Haviland, 1931 syn. nov. (=Zelus kartabensis Haviland, 1931), Zelus personatus Berg, 1879 syn. nov. (=Zelus versicolor Herrich-Schaeffer, 1848), Zelus trimaculatus Champion, 1898 syn. nov. (=Zelus means Fabricius, 1803), Zelus trimaculicollis (Stål, 1855) syn. nov. (=Zelus means), and Zelus tristis Haviland, 1931 syn. nov. (=Zelus laticornis). Zelus conjungens (Stål, 1860) stat. rev. Is resurrected from junior synonymy with zealous armillatus (Lepeletier & Seville, 1825). Zelus ambulans Stål, 1862 stat. rev. and Zelus cognatus (Costa, 1862) stat. rev. are resurrected from synonymy with Zelus exsanguis Stål, 1862. Iquitozelus Bérenger syn. nov. is synonymized with Zelus and its only species transferred to Zelus, hence resulting in a new combination, Zelus couturieri (Bérenger, 2003) comb. nov. Lectotypes, paralectotypes or neotypes are designated for a number of species. Habitus images, illustrations of male genitalia, distribution maps and measurements are provided for nearly all species. The three previously recognized subgenera of Zelus are found to be based upon superficial characters and these divisions do not reflect natural groupings. Using sets of characters, especially those of the male genitalia, eleven species groups are proposed. It is also hypothesized that Zelus is closely related to three other New World genera: Atopozelus Elkins, Ischnoclopius Stål and an undescribed genus "Hartzelus" [manuscript name]. Zelus is endemic to the New World, occurring naturally in the Caribbean and all but one of the continental countries, with introductions to Pacific islands, Europe and Chile.


Introduction
is one of the largest reduviid genera ) and the largest New World genus in the tribe Harpactorini (Reduviidae: Harpactorinae). Zelus is endemic to and widely distributed throughout the New World, ranging from southern Canada through central Argentina. One species, Zelus renardii Kolenati, 1856, has been introduced to Hawaii (Kirkaldy 1902, Zimmerman 1948, and was recently found in Chile (Curkovik et al. 2004, Elgueta andCarpintero 2004), Greece (Davranoglou 2011, van der Heyden 2015and Spain (Vivas 2012 (reviewed in Weirauch et al. 2012). Species of Zelus, among several other genera (e.g., Arilus Hahn, Sinea Amyot & Serville, and Montina Amyot & Serville), have been explored and studied as natural enemies in the Americas (Cohen andTang 1997, Cogni et al. 2002). Species of Zelus prey on a wide range of insects in cotton, corn, soybean, alfalfa crops and fruit trees in California and elsewhere (Ali and Watson 1978, McPherson et al. 1982, Cisneros and Rosenheim 1998, Virla et al. 2015, may reach population densities of up to 50,000 to 75,000/ha, and prevent outbreaks of lepidopteran larvae (Ables 1978). Hart (1972) conducted a taxonomic revision of Zelus with descriptions of twenty-five new species and twenty-six new synonyms, most of which remained unpublished (see  for treatments of twenty Canadian, US, northern Mexican and Caribbean species). The current state of taxonomy of Zelus remains unsatisfactory and impedes further research on the evolution and ecology of this group. Species identification is difficult in many instances, and misidentifications may arise. For example, Z. renardii was misidentified as Zelus cervicalis Stål, 1872 when it was reported as having been introduced to Chile ( Curkovik et al. 2004). This project was thus undertaken to provide a taxonomic monograph of the genus Zelus.
In the current study seventy-one species are treated and twenty-four described as new. Five species are removed from Zelus and placed incertae sedis within Harpactorini. Nine new synonyms are recognized. Three species are resurrected. Iquitozelus Bérenger is synonymized with Zelus. Habitus images, illustrations of male genitalia, distribution maps, and identification keys are provided. This work evaluates and maintains most of the manuscript new species names proposed in Hart (1972)'s systematic revision of Zelus. Four additional new species are discovered and described herein, some based on specimens more recently collected, but some arising from different views of species boundaries. The vast areas of the Amazon and many mountainous regions of Central America and South America remain poorly sampled and should represent an immediate frontier for new species discoveries in this genus.

Review of taxonomic history
The taxonomic history of Zelus is complex and the generic limit of Zelus has undergone constant fluctuations. The first species of Zelus, Z. longipes (Linnaeus), was described by Linnaeus in the 12th edition of Systema Naturae (Linnaeus 1767) under Cimex, a genus in which he also included various other Heteroptera that are now classified within a number of families. Fabricius (1775) transferred Z. longipes from Cimex to Reduvius, a genus that was established to accommodate most of the Reduviidae known at the time. It was again Fabricius who later in the first comprehensive treatment of Hemiptera (Fabricius 1803) erected the genus Zelus. In this work for each genus Fabricius selected one species for which he repeated the short generic description, expanded the species description and italicized the terms referring to the morphological structures described. Zelus longipes was treated by Fabricius this way for the genus Zelus. Therefore, many workers assumed that Fabricius had indicated Z. longipes as the type species of the genus (Kirkaldy 1900a).
Lepeletier and Serville (1825) expanded the limit of Reduvius to include nearly all then described Reduviidae and described many new species; several of them would now be considered members of Zelus. An erroneous designation of Zelus festinans as the type species of Zelus was made by Laporte (1832). He erected new genera and removed some species from Zelus, somewhat changing the generic limit of this genus. Perty (1834)  List of museums/collections look up actual specimen data for locality information (Suppl. material 1). Interactive specimen mapping is available using the Global Mapper module of "DiscoverLife" (http:// www.discoverlife.org/mp/20m?act=make_map).

Morphological methods -dissection, observation, imaging and measurement
Dissection. Male genitalia, including the eighth abdominal segment and pygophore with phallus enclosed, were removed, cleared in heated 10% potassium hydroxide (KOH) solution for 5-10 minutes, washed in distilled water, and stored in glycerol. To remove the genitalia, a specimen was softened by soaking the abdomen in water. This was achieved by stationing a pinned specimen on play clay with its abdomen pointing down and immersed in water, while the rest of the specimen was not submerged. This method avoided soaking the whole specimen or removing the entire abdomen. A 'genitalia hook' was made by melting the tip of a glass Pasteur pipette with a minuten insect pin inserted and fixed into it. The pygophore was carefully removed by holding the softened abdomen with forceps on one hand, and inserting the genitalia hook into the membranous connection between the seventh and eighth segments, breaking that membrane and pulling off the pygophore, a series of actions performed by the other hand.
Observation. Observations were made using a Nikon stereo dissecting microscope SMZ1500, illuminated by a Nikon NI-150 High Intensity Illuminator. Initial observations of morphological characters were made based on typically a small number of specimens (one to five) and intraspecific variations were subsequently examined based on a larger selection of geographical representation. Genitalia were observed in glycerol. Structures in this medium may look different from their dry state, especially for soft cuticles. For example, the apices of the parameres of Z. cervicalis, Zelus luridus Stål, 1862 and many other species usually appear shriveled in dry specimens, but are fully expanded in glycerol or 70% ethanol. The orientation of the dissected structures shown in illustrations does not necessarily reflect their natural condition.
Imaging and illustration. Dorsal and lateral habitus images of specimens were taken with a Microptics-USA system (now Visionary Digital, http://www.duninc.com/index.html) with a K2 lens and CF-2 or CF-4 objectives connected to a Canon EOS 1D digital SRL. Images were edited in Photoshop (CS3/CS4) to adjust levels and sharpness. Image background was removed and replaced using CorelDRAW or Photoshop. For most species illustrations of male genitalia were adapted from Hart (1972) except for several new species not included in that work.
Measurement. Measurements were made on a dissecting scope equipped with a two-axes movable stage (Mitutoyo Corp.), with the aid of two digital micrometers (Boeckeler®), which were connected to a Microcode II RS-232 digital readout (Boeckeler®). Most measurements were done in dorsal view, but various orientations were necessary for measuring appendages. Typically, five to ten specimens were measured for each species, but the number may be fewer for species without enough properly preserved specimens. All measurement values reported here are in mm, unless otherwise stated. In Suppl. material 2 measurement values are divided into length values and width values. Suppl. material 2 reports not only average values for each species but also values of all individual specimens. A total of twenty-nine measurements were captured as listed in the following.
Total length: length of body from clypeus to apex of hemelytron 2.
Head (length of head from clypeus to collar of anterior pronotal lobe) 4.
AntOc: Anteocular (length of anteocular region of head, from clypeus to anterior margins of eyes) 5.
PostOc: Postocular (length of postocular region of head, from posterior margins of eyes to collar of anterior pronotal lobe) 6.
Scut: Scutellum (only exposed part measured, from posterior margin of pronotum to apex of scutellum) 9.
Antn3: Antennal segment 3/Basiflagellomere (the basiflagellomere tends to be curled and in that case two or several consecutive measurements were taken and their sum was used) 12.
Lb1: 1st visible labial segment (this is actually homologous to the second labial segment in other heteropteran insects, and Lb2 and Lb3 are homologous to the third and fourth segments. See Weirauch 2008b) 20.
Lb3: 3rd visible labial segment Width measurements 1. Head (width from outer margin of one eye to that of the other) 2.
InterOcDi: Interocular distance (width from inner margin of one eye to that of the other) 3.
Abd: Abdomen (measured at the widest part of the abdomen) 6.

Descriptive taxonomy
Description. Observations were recorded with the software DEscriptive Language for TAxonomy (DELTA) (Dallwitz 1980, Dallwitz et al. 1999. Natural descriptive language was exported and edited. Observations and descriptions were done primarily based on male specimens, descriptions of females were restricted to non-genitalic characters different from males. Four major character systems were described: coloration, vestiture, structure (non-genitalic), and male genitalia. Description of abdominal vestiture was restricted to the ventral surface. Description of a body part usually started with the overall appearance of that body part such as length, width, and general shape. Structural components of that body part were then described in the order from anterior to posterior, medial to lateral, dorsal to ventral, and proximal to distal. Ratios were determined by comparing mean values across specimens measured. Ratio between the segments of labium or antenna was in reference to the respective first segment. Unless otherwise stated, all measurement values reported in the text are mean values. For closely related or very similar species, a full description was provided for one representative and only variable characters were described for other species. The singular form was usually used for paired structures except when referring to spatial relationships between these structures, e.g., "struts (of phallus) separate, sub-parallel", or when referring to the different pairs of legs at the same time, e.g., "femora and tibiae with alternating yellow and dark brown bands". Descriptions of coloration were mainly based on observations of dried specimens, but notes were made if images or observations of live specimens were available. We note that many dried specimens show browning or fading of colors. This was particularly evident in lightly or brightly colored species. We observed in a number of species that the ventral outline of abdomen is curved, a condition apparently resulting from the folding of segments three to six (sometimes two as well). It was not clear if this was a preservation artifact or a natural condition, but it has been consistently seen in several species. Thus, this character was described as observed.
Description of intraspecific variations. Intraspecific variations were described and indicated by terms or phrases as the following: sometimes, occasionally and in some specimens. When variations in coloration can be roughly delimited to several patterns, they were described and the frequency of the patterns sometimes mentioned as well. Intraspecific variations in male genitalic structures were usually not described or documented unless they are important for species diagnosis and identification, which is usually the case only for closely related species.
Association of males and females. For the majority of species, males and females show limited sexual dimorphism in size and coloration, and could be readily associated based on external morphology, corroborated by collecting data. However, sexual dimorphism is pronounced in a number of species. Males and females differ drastically in size, body configuration and coloration. Association of sexes for these species was based mainly on locality data and series of specimens of both sexes. Observations of mating reported in the literature were also consulted and used as corroborative evidence whenever available.
Terminology and abbreviation. External morphology and genitalic terminology followed Forero and Weirauch (2012), Davis (1966), Weirauch (2008a), Weirauch (2008b). The term 'posterior margin of foramen' was used to refer to the posterior margin of the foramen of the dorsal phallothecal sclerite. The foramen is located at the anterior part of the dorsal phallothecal sclerite and is an area that lacks sclerotization and surrounds the struts. Nomenclature (Annotated synonymic list). The synonymic list comprises abbreviated synonymies and included those names that previously appeared in the taxonomic literature or have affected the taxonomy of the species. Citation to ecological, agricultural or other non-taxonomic literature was presented when appropriate, but not meant to be exhaustive. Historical taxonomic publications were briefly annotated to indicate kind of taxonomic information or nomenclatural acts such as .orig descr., checklist, cat., note, fig. and key. When a species name is followed by the original author and year, there is no colon (:) separating the name and the author. A species name followed by a colon indicates that the author of the work is not the author of the name. Male genitalic structure terms (Zelus errans Fabricius, 1803 is shown in the illustrations)

Taxon treatments
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 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 andZelus 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.

Taxon discussion
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.  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.

Species groups
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.

Zelus tetracanthus species group.
Zelus minutus , 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 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.
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- Zelus tetracanthus species group, male gentitalic structures 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.

Zelus nugax species group.
Zelus grassans Stål, 1862, Zelus illotus Berg, 1879, Zelus impar Kuhlgatz, 1902, Zelus nugax Stål, 1862and 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  Zelus mimus species group, male genitalic structures 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.

Zelus puertoricensis species group.
Zelus bruneri De Zayas, 1960, Zelus puertoricensis Hart, 1987, Zelus subimpressus Stål, 1872and 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.
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 Zelus nugax species group, male genitalic structures Central America, but Z. cervicalis extends to northern South America. Comparative views of male genitalia are shown in Fig. 7 .

Zelus armillatus species group.
Zelus amblycephalus sp. n., Zelus annulosus (Stål, 1866)   Zelus renardii species group, male genitalic structures 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  This and the next species group (Zelus vagans species group) possess dense, spinelike 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 Zelus armillatus species group, male genitalic structures -pygophore 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. 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.

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 Zelus vagans species group, male genitalic structures 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. Zelus panamensis species group, male genitalic structures -phallus 11. Zelus erythrocephalus species group.
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. 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.

Diagnosis
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.

Distribution
South America (Fig. 18 Male: (Fig. 19a 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.

Diagnosis
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.

Distribution
Southern Mexico to northern South America and part of Brazil (Fig. 21  Zelus ambulans Stål, 1862, specimen record map 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. 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.

Diagnosis
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.

Taxon discussion
Champion (1898)  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.

Diagnosis
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.

Distribution
South America (Fig. 27) 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 reddishbrown, yellowish-brown ventrally. Dorsum of postocular lobe dark brown with yellowishbrown 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 yellowishbrown. Scutellum yellowish-brown to reddish-brown. Legs yellowish-brown to reddishbrown, 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.

Diagnosis
As with several members of the Zelus luridus species group, the coloration is greenishbrown, 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.

Etymology
Named after the type locality, Antigua, in Guatemala.

Zelus armillatus
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.

Diagnosis
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.

Taxon discussion
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.

Diagnosis
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.

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

Diagnosis
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.

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

Distribution
South America (Fig. 41). Known from the type locality in Brazil. 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; Zelus banksi Zhang & Hart, sp. n., specimen record map 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.

Diagnosis
Recognized by the following combination of characters: the posterior pronotal lobe usually orangish-brown; the rather long paramere, apex obliquely truncate; and the medial process nearly straight, curvature small. Among the males of the Zelus panamensis species group (Fig. 12), Z. banksi has the longest paramere, which is longer than the medial process.

Etymology
Named after N. Banks, the collector of the type specimen.

Distribution
Southern Central America and northern South America (Fig. 44). Countries with specimen records: Colombia, Costa Rica and Panama

Distribution
Known only from Cuba.

Taxon discussion
Two male specimens are known from Cuba, which were not physically examined, but the original description and illustration provide a strong basis for placing this species in the Zelus puertoricensis species group. This is confirmed by the narrow, elongated body form; the flat and rectangular pronotum; the general genitalic shape indicated in the figure. The much smaller size and flat postocular lobe negates it being a male of Z. subimpressus. It is more likely to be the male of Z. zayasi. posterior lobe with short, erect, spine-like setae, some apically curved; pleura with short to moderately long, recumbent and semi-erect setae, some covered with white waxy exudation; scutellum with recumbent setae. Legs with sparse setation. Corium and clavus with short, recumbent setae. Abdomen with sparse, short, recumbent setae, intermixed with few longer setae. Dorsal, outer surface of enlarged part of paramere with dense, long, erect setae. STRUCTURE: Head: Elongated. Postocular lobe very long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye prominent; lateral margin much wider than postocular lobe; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head. Thorax: Anterolateral angle with inconspicuous subtuberculate 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 projection. Scutellum moderately long; apex angulate, slightly projected upward. Legs: Slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell moderately sized; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 46) Pygophore: Ovoid; expanded laterally near base of paramere in dorsal view. Medial process expanded laterally; rather broad; moderately long; anteroposteriorly compressed; erect; curved at middle; apex emarginate, with pair of subapical, lateral, hooklike processes; lateral elevations running from below base of medial process through middle. Paramere: Short, not reaching apex of medial process; base slightly constricted; strongly curved ventrad. Phallus: Dorsal phallothecal sclerite somewhat squarish; lateral longitudinal blade-like heavy sclerotization pressed against phallothecal sclerite; apical portion of phallothecal sclerite gradually tapering, slightly convex, laterally angulate, angulation ending anteriorly in sharp, dorsad projection; apex rounded, medially emarginate; posterior margin of foramen concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally separate. Basal plate arm moderately robust; basally fused; in lateral view strongly curved at midpoint; bridge extremely short; extension of basal plate expanded onto arm.

Diagnosis
Recognized by the uniform dark brown coloration; the extremely long postocular lobe; and the rather broad medial process, apex emarginate in the middle and bearing a pair of ventrally directed projections.

Etymology
Named after Casi.

Distribution
South America (Fig. 47). A single specimen is known from the State of Amazonas, Brazil.

Figs 48, 49, 50
Male: (Fig. 48a, b, c) Medium-sized, total length 9.81-13.08 mm (mean 11.78 mm, Suppl. material 2), very slender, body length/width=6.2. COLORATION: Yellowishbrown to dark brown, some specimens with dark spots or bands on legs. Anteocular lobe yellowish-brown to reddish-brown, dark brown between eye and antennal insertion, some specimens with dark brown mid-dorsal areas. Dorsum of postocular lobe dark brown, variably shaped medial longitudinal line and area between ocelli and eye yellowish-brown, ventral surface yellowish-brown. Labial segments I & II yellowishbrown; segment III reddish to dark brown. Antennal segments brown, sometimes scape darker on dorsal surface or pedicel darker apically. Anterior pronotal lobe yellowishbrown to brown, collar and setal tracts darker, some specimens with dark brown spot on pro-episternum. Posterior pronotal lobe yellowish-brown to brown. Pleura yellowishbrown. Sternites yellowish-brown; meso-sternum with dark brown area anterior to meso-coxa. Scutellum yellowish-brown to brown, apex lighter. Legs yellowish-brown, many specimens with dark brown raised spots or bands on femora and tibiae (see "Taxon Discussion" below). Corium and clavus reddish-brown, veins yellowish-brown; membrane yellowish-brown. Dorsum of abdomen yellowish, reddish, or dark brown; connexival margins and ventral surface yellowish-brown. Pygophore yellowish-brown; some specimens with medial process apically reddish-brown or brown. VESTITURE: Moderately setose. Pubescence of short recumbent and short to long erect setae. Anteocular lobe with short recumbent and erect setae over entire surface, more dense dorsally; postocular lobe with short to moderate recumbent and moderate to long erect setae, erect setae more dense posteriorly. With short to moderate recumbent setae over entire surface, confined to setal tracts on dorsum of anterior pronotal lobe, longer erect setae on lateral surface; scutellum with short recumbent and short to moderate semi-erect and erect setae over surface. Legs with short to long semi-erect to erect setae. Corium and clavus with short, recumbent setae. Abdomen with short recumbent and some short to moderate erect setae over ventral and lateral surfaces. Exposed surface of pygophore with short recumbent and short to long erect setae; short to moderately stiff erect setae on apical half of parameres. STRUCTURE: Head: Cylindrical, L/W = 2.83. Postocular lobe moderately long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye moderately sized; lateral margin only slightly wider than postocular lobe; dorsal and ventral margins removed from surfaces of head. Labium: I: II: III = 1.0: 2.0: 0.5. Basiflagellomere diameter larger than that of pedicel. Thorax: Anterolateral angle bearing small projection; medial longitudinal sulcus evident only on posterior 1/2, deepening anterior to transverse sulcus of pronotum. Posterior pronotal lobe with finely rugulose surface; disc slightly elevated above humeral angle; humeral angle armed, with dentate projection. Scutellum long; apex angulate, not projected. Legs: Slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell small, elongate; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 49) Pygophore: Ovoid. Medial process cylindrical; slender; long; laterally somewhat compressed; erect; nearly straight; basally without protrusion; apex in posterior view modified, hooklike. Paramere: Cylindrical; moderately long, achieving apex of medial process; directed toward medial process; basally narrower; curved dorsad; apical part enlarged. Phallus: Dorsal phallothecal sclerite shield-shaped; lateral margin recurved dorsad; apical portion of phallothecal sclerite gradually tapering, flat, lateral margin recurved; apex rounded, medially emarginate; posterior margin of foramen broadly concave. Struts attached to dorsal phallothecal sclerite; apically missing. Basal plate arm moderately robust; basally fused; in lateral view basally strongly curved; bridge short; extension of basal plate small, marginally expanded onto arm.

Diagnosis
The rather slender body form makes this species easy to separate from other species that occur in the same geographic region. Males can also be recognized by the paramere apically greatly enlarged; the medial process apically curved ventrad, hooklike; the lateral margin of the dorsal phallothecal sclerite recurved. Zelus cervicalis is most similar to Z. renardii and the two share a number putatively synapomorphic characters of structures of male genitalia. The more slender body separates both sexes of Z. cervicalis from Z. renardii. Males of Z. cervicalis also have the apex of medial process not bent as strongly as that in Z. renardii.  Taxon discussion  stated that, based on male genitalic characters and pilosity, Z. cervicalis and Z. renardii are closely related species, and we agree with that view. We also corroborate, using a larger specimen sample, the western and eastern parapatric a b c d Figure 49.   delimited two populations of Z. cervicalis, i.e., a South Atlantic and Gulf Coast population and a Mexico-Central America population, the latter also extending to southeastern Arizona and northern Colombia. Most individuals of the South Atlantic and Gulf Coast population have unicolorous legs, or, at most, only a few brownish to reddish spots. Specimens of the Mexico-Central America population have heavily spotted or banded legs. This pattern is also recovered in the current study. However, contrary to Hart's claim that "occasional specimens from either population may occur that do not conform to the normal pattern for that population", we found that all specimens of the Mexico-Central America population have spotted or banded legs.

Zelus cervicalis
This condition also appears in a small number of specimens in other populations (e.g., UCR_ENT 00016129, UCR_ENT 00039079, UCR_ENT 00042740, UCR_ENT 00042741, UCR_ENT 39522, UCR_ENT 00039519, UCR_ENT 00039531, UCR_ENT 00039525, UCR_ENT 00039561, UCR_ENT 00039560, UCR_ENT 00039559, UCR_ENT 00039557, and more specimens from Texas). We also observed that compared to populations in other US states, specimens from southern Texas tend to have spotted legs, but the density of spots is lower than that in the Mexico-Central America population. By examining previously unstudied Mexican specimens from southern Sonora and northern Sinaloa, we also support Hart's second theory that the Arizona specimens are in continuity with the remainder of the population. The male genitalia are also variable in a number of respects between the two populations ( Fig.  49), mainly in the shape of the paramere, the elevation of the lateral margins of the dorsal phallothecal sclerite near the base, and the relative massiveness of the basal plate arms. Hart remarked that the Mexico-Central American specimens show more similarities to the Gulf Coast specimens as one proceeds southward through Central America.
The images of the lectotype of Z. cervicalis are available on the 'Types of Heteroptera' website of the Swedish Museum of Natural History.   Lethierry and Severin, 1896, p. 151, cat.;Champion, 1898, p. 260, note;Wygodzinsky, 1949a, p. 48-49, checklist;Maldonado, 1990, p. 326 Male: (Fig. 51a, b) Medium-sized, total length 12.50-14.06 mm (mean 13.45 mm, Suppl. material 2); robust. COLORATION: Usually entire surface of body black, some specimen with white posterior margin of posterior pronotal lobe, some with posterior pronotal lobe uniformly orangish-brown. Scape with or without band. Profemur with single pale brown band, protibia without band; meso-and metafemora with two or three bands, tibiae with single band. VESTITURE: Sparsely setose. Dorsum of head with dense, short, stout, recumbent, black setae and sparse, long, fine, erect setae, as long as width of eye in dorsal view; ventral surface with moderately dense, short, recumbent, Zelus chamaeleon Stål, 1872, specimen record map fine setae and spare, moderately long, erect setae. Moderately dense, short, stout, recumbent setae on pronotum; anterior lobe also with sparse, long, erect setae; scutelum with dense, short to long, semi-erect setae; pleura with short to long, semierect or erect setae; sternites with dense, long, erect setae. Corium and clavus with short, recumbent setae. Abdomen with moderately dense, short, recumbent to subadpressed setae, intermixed with long, erect setae. With dense, short, adpressed setae, mixed with sparse, very long, moderately stout, semi-erect setae; apical half with dense, moderately long, stout, semi-erect setae. STRUCTURE: Head: Somewhat globular, L/W = 1.93. Postocular lobe short; in dorsal view narrowing till abrupt posterior constriction, very short behind constriction. Eye moderately sized; lateral margin much wider than postocular lobe; dorsal and ventral margins removed from surfaces of head. Labium: I: II: III = 1: 1.6: 0.4. Basiflagellomere diameter slightly larger than that of pedicel. Thorax: Anterolateral angle with inconspicuous subtuberculate projection; medial longitudinal sulcus evident only on posterior 1/2, deepening anterior to transverse sulcus of pronotum. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with spinous processes. Scutellum moderately long; apex angulate, slightly projected upward in some specimens. Legs: Slender. Hemelytron: Surpassing apex of abdomen by about twice length of abdominal segment seven; quadrate cell moderately large; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 52) Pygophore: Ovoid; mid-lateral fold adjacent to paramere insertion; expanded laterally near base of paramere in dorsal view. Medial process triangular; short, shorter than paramere; erect; straight; apex in posterior view acute, with hooklike projection extending towards downward, ending as transverse bridge. Paramere: Cylindrical; short, not reaching apex of medial process; directed posteriad, slightly curved towards medial process; basally narrower; slightly curved ventrad; apical part enlarged. Phallus: Dorsal phallothecal sclerite shieldshaped; lateral ridge-like dorsad expansion continuous with basal arm; apical portion of phallothecal sclerite gradually tapering, convex, medially keeled; apex truncate; posterior margin of foramen deeply concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally separate. Basal plate arm robust; separate; converging; in lateral view slightly curved; bridge short; extension of basal plate expanded onto arm.
Female: (Fig. 51c, d) Similar to male, except for the following. Larger than male, total length 13.45-15.02 mm (mean 14.39 mm, Suppl. material 2). More variable than in male. Dorsum of postocular lobe always black; pale, slender, medial longitudinal stripe; anteocular lobe usually black, if with part of surface red, clypeus always black. Dorsum of anterior pronotal lobe almost always black, collar sometimes red, lateral surface black or mixed with red. Dorsal surface of posterior pronotal lobe of four major color patterns: entirely black, entirely red or orange, anterior portion red and posteriorly black, mostly black with medial red circular patch; last pattern most common (eight out of fourteen); lateral surface always red or orange (when dorsal surface is orange). Variable amounts of black and red on pleura. Corium and clavus brownish-black or yellowish-brown; membrane always dark brown. Abdomen segment usually red, anterior black stripe; entirely black in some specimens. Hemelytron slightly surpassing apex of abdomen.

Diagnosis
Can be recognized by the following combination of characters: the long, erect, fine setae on head, anterior pronotal lobe, pleura and sternites; the stout and short head and the nearly hemispherical postocular lobe; the short paramere, not exceeding medial process; the medial process short and triangular, apex as hooklike process, extending ventrally as transverse ridge; and the apical surface of dorsal phallothecal sclerite medially with keel-like elevation.

Distribution
South America (Fig. 53). Known only from Colombia.   Zelus championi, Zhang & Hart, sp. n., specimen record map sclerite; apically separate, connected by bridge. Basal plate arm slender; separate; somewhat converging; in lateral view very slightly curved; bridge moderately long; extension of basal plate small, laterally not greatly expanded onto arm.

Diagnosis
The strongly contrasting black dorsal surface and red abdomen is distinctive of this species. The features of the genitalia are rather similar to those of other species in the Zelus vagans species group (Fig. 11), but the apex of the medial process is more strongly bent ventrad. Other diagnostic characters shared with members of the Zelus vagans species group and the Zelus longipes species group include the unarmed rounded humeral angle and the spine-like setae on pronotum.

Etymology
This species epithet is a patronym, in honor of entomologist George C. Champion , who authored several volumes on Rhyncophora (Heteroptera) in the Biologia Centrali Americana series.

Distribution
Central and South America (Fig. 56). Countries with records: Costa Rica, Ecuador and Panama

Ecology
No natural history or ecological knowledge is known, but we hypothesize that the strikingly contrasting black and red coloration is at the same time cryptic and aposematic, and may also be mimetic. Based on observations of other species, we know that low vegetation is a common habitat of members of this genus. In a dense forest, predators from above may confuse the black dorsum of Z. championi with dark forest background, while the strong contrast formed by black and red colors is highly visible to predators (e.g., lizards) at the same level or approaching from below. Like many assassin bugs, species of Zelus may inflict a painful bite when attacked. Besides, harpactorines, including Zelus spp., emit a foul odor when handled. We do not know if vertebrate predators are deterred by this odor, but it is strong enough to be detected by a human even meters away. Hence Z. championi may be well defended against predators and the contrasting coloration serves as a signal for unpalatability. Of course, many other species of Zelus are dull colored, but expected to have the same kind of physical or chemical defense. There may be other ecological factors that determine the coloration of Z. championi. We suspect that mimicry is one. Many other unpalatable insects show similar contrasting bright red and black color patterns. Zelus championi may participate in Müllerian mimicry with those species.

Taxon discussion
The type specimen of this species was originally described as the male of Z. inconstans, a species very similar in general form to Z. championi. On the basis of pubescence, pronotal armature and whitish exudation, Champion himself questioned the conspecificity of this male with the three females of the original type series. As more material was available for the present work, his doubts have been substantiated, the male of Z. inconstans identified and this particular specimen found to be a male of a new species. The two species belong to different species groups, as verified by pubescence and genitalic characters.

Description
The following is a translation of the original description: "Closely related to the preceding species [Z. ambulans]; differing in that the spines of the humeral angle of the pronotum are conspicuously directed obliquely upward; dorsal surface of the head black, longitudinal line and transverse sulcus yellowish; first and second antennal segments testaceous, apex black. --length 15 mm."

Taxon discussion
This species was originally described from a single specimen from Mexico. The original description did not indicate its sex. Champion's synonymy was apparently based on the description and not upon examination of the specimen. Attempts to locate the holotype were unsuccessful. From the above original description it is impossible to determine whether this species may be synonymous with Z. exsanguis or is a separate species. It is reasonably certain, however, that it belongs to the Zelus luridus species group, as the comparison with Z. ambulans precludes any similarity to other reduviid genera or even groups in this genus within Mexico. Male: (Fig. 57a, b, c) Large, total length 16.64-19.80 mm (mean=18.34 mm, Suppl. material 2); robust. COLORATION: Variably yellowish-brown to brownish-black. Dorsal surface of head brown, mixed brownish-black; ventral surface brown. Anterior pronotal lobe variably brown to brownish-black, nearly entirely brownish-black in some specimens, never entirely yellowish-brown, with at least brownish-black spots. Posterior pronotal lobe usually brownish-black in center, margins yellowish-brown, entirely yellowish-brown in occasional specimens. Corium and clavus with proximal portion brownish-black, distal portion yellowish-brown, entire surface yellowish-brown in some specimens. Legs with or without bands. Lateral and ventral surfaces varying from most Zelus conjungens (Stål, 1860), specimen record map yellowish-brown with brownish-black spots to nearly entirely brownish-black. VESTITURE: Moderately setose. Very similar to that in Z. armillatus; adpressed setae more sparse. STRUCTURE: Head: Cylindrical, L/W = 2.29. Postocular lobe long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye smallish; lateral margin only slightly wider than postocular lobe; dorsal and ventral margins removed from surfaces of head. Labium: I: II: III = 1: 1.4: 0.4. Basiflagellomere diameter smaller than that of pedicel. Thorax: Anterolateral angle bearing small projection; medial longitudinal sulcus shallow near collar, deepening posteriorly. Posterior pronotal lobe with finely rugulose surface; disc slightly elevated above humeral angle; humeral angle armed, with short tuberculate processes. Scutellum moderately long; apex blunt, very slightly projected upward. Legs: Robust. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell large and broad; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 58) Pygophore: Ovoid; slightly expanded laterally near base of paramere in dorsal view; posteriorly expanded sac-like sclerite between paramere and medial process. Medial process robust; broad; short; posteriorly directed; basally slightly protruding; apex in posterior view truncate, with very inconspicuous lateral prongs. Paramere: Cylindrical; moderately long, not reaching apex of medial process; directed posteriad; slightly curved dorsad; apical part not enlarged. Phallus: Dorsal phallothecal sclerite shield-shaped; lateral expansion arising close to base; apical portion of phallothecal sclerite not distinctly tapered, flat, laterally angulate; apex rounded; 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 slightly curved; bridge moderately long; extension of basal plate expanded onto arm.

Diagnosis
Among species of the Zelus armillatus species group, the medial process in Z. conjungens is the broadest, being more than 2x the diameter of paramere and more than 1.5x the diameter of ocellus. Other characters helpful for identification may include the lateral processes on apex of medial process minute, inconspicuous. Most similar to Z. armillatus, but can be separated by characters aforementioned, and also the lateral expansion on dorsal phallothecal sclerite close to basal arm not as sharp process. In females the mesofemur is swollen nearly throughout, much thicker than profemur, which will serve as a basis for separation from the females of Z. armillatus.

Distribution
Southern Central America, northern South America and Southern Brazil (Fig. 59). Countries with specimen records: Costa Rica, Colombia, Ecuador, Panama, Venezuela and Brazil.

Taxon discussion
Mayr (1866) synonymized Z. conjungens with Z. armillatus, the former as the junior synonym. We here resurrect this species on the basis of the characters described in the diagnosis. Two disjunct populations are recognized for this species, one in Southern Brazil and another in Central America and Northern South America. The latter represents a species formerly described by Champion, Z. atripes, which is here considered conspecific with Z. conjungens, as in both species the females show a swollen femur and the medial processes of males are broad. However, male genitalia of the two populations are somewhat different and may warrant a further closer examination. In any area in which the species occurs there is apparently a great range of color and pattern variations, but there does seem to be a general trend toward the lighter colorations in Colombia, Costa Rica and Panama.
Female: (Fig. 60c, d) Similar to male, except for the following. Larger than male, total length 15.56-18.20 mm (mean 16.82 mm, Suppl. material 2). Dorsum nearly uniformly brown, lateral and ventral surfaces and legs yellowish; single dark spot on each abdominal segment.

Diagnosis
The nearly uniform brown dorsal coloration; the dorsum with short, erect, somewhat spine-like setae; and the posterior margin of the pronotum smoothly convex. The paramere short, broad; the apical part of medial process laterally compressed and ridged on the anterior surface; the posterolateral rim of pygophore with lightly sclerotized expansion below paramere; and the basal plate arm separate, not fused. In females the dorsum is nearly uniformly brown, the lateral and ventral surfaces yellowish, the legs apically reddish-brown, not conspicuously banded and the abdominal segment with single dark spot.

Etymology
Named after the type locality "Cordillera" in Peru.

Distribution
South America (Fig. 62). Known only from Peru.

Diagnosis
The connexivum segment VI with foliaceous expansion is unique among all females of Zelus spp.

Distribution
South America (Fig. 64). Known only from Peru.

Taxon discussion
Bérenger (  differences. The posterior pronotal lobe disc in Z. couturieri appears to be only slightly above and nearly continuous with humeral angle. In the other three species, the discs are clearly above humeral angle. The posterior margin of the posterior pronotal lobe in Z. couturieri is not developed, whereas it is well defined in other species. Furthermore, the lateral process of the humeral angle is conspicuously darkened, also unique to Z. couturieri. Based the foregoing observations, it is likely that Z. couturieri represents a distinct a species, and not the female of Z. umbraculoides or Z. umbraculus. The specimens were not physically examined as Bérenger (2003) has provided a detailed description of this species. Male: (Fig. 65a, b) Medium-sized, total length 13.62-17.91 mm (mean 14.10 mm, Suppl. material 2); slender. COLORATION: Many specimens with wasp-like habitus, with alternating black and yellow areas; anterior pronotal lobe usually dark brown, posterior lobe and proximal portion of corium yellowish; some specimens with nearly entire dorsal surface dark. Legs vary from nearly uniformly yellow or blackish-brown to yellow-brown banded. VESTITURE: Densely setose. Head with dark, erect, spine-like setae dorsally and light, recumbent setae ventrally. Anterior pronotal lobe with short, dark, spine-like setae, confined to setal tracts; posterior pronotal lobe with short, spine- Medial process cylindrical; extremely slender; moderately long; semi-erect; apex in posterior view blunt, slightly folded posteriad. Paramere: Cylindrical; long, surpassing medial process; directed posteriad, slightly curved towards medial process; slightly curved ventrad; apical part very slightly enlarged. Phallus: Dorsal phallothecal sclerite elongated; apical portion of phallothecal sclerite not distinctly tapered, convex, laterally indistinctly angulate; apex truncate, medially emarginate; posterior margin of foramen deeply concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally almost completely fused. Basal plate arm slender; separate; converging; in lateral view very slightly curved; bridge moderately long; extension of basal plate small, marginally expanded onto arm.

Zelus errans
Female: (Fig. 65c, d, e) Similar to male, except for the following. Larger than male, total length 18.30-20.18 mm (mean 18.97 mm, Suppl. material 2). Coloration pattern more variable than in male.

Diagnosis
May be confused with Z. vespiformis and Z. gracilipes, species that have similar appearances. Distinguished from Z. vespiformis by the more elongated body; the longer medial process (Fig. 10), the Cu and Pcu of quadrate cell subparallel, and the Cu-Pcu2 (posterior cross vein) less than 1/2x length of Cu. Males of Z. errans can be separate from Z. gracilipes by features of the genitalia, the latter belonging to a different species group. Females typically have the anterior membrane of the hemelytron semitranslucent, whereas the entire membrane is colored or opaque in Z. gracilipes.

Diagnosis
Recognized by the following combination of characters: the dorsal coloration nearly uniformly dark brown, the head reddish-brown, and the membrane with indistinct iridescence. Most similar to Z. paracephalus and Z. russulumus; can be distinguished from both by the rather slender medial process. Females of Z. erythrocephalus, Z. paracephalus and Z. russulumus are difficult to separate.

Taxon discussion
Zelus erythrocephalus and two other species in the same species group, Z. paracephalus and Z. russulumus superficially resemble Z. panamensis, a species in a different group. All have a orange, reddish head and a uniformly dark dorsum. These can be separated from Z. panamensis based on male genitalia and iridescence on membrane. Male: (Fig. 71a, b) Medium-sized, total length 13.79-16.41 mm (mean 14.98 mm, Suppl. material 2); slender. COLORATION: Anteocular lobe reddish-brown, most specimens with central dorsal area darker reddish-brown and with lighter mid-dorsal line, dark areas between compound eyes and antennal insertions. Postocular lobe yellowish-brown to reddish-brown, dorsal surface dark brown to brownish-black with mid-dorsal and circumocellar areas yellowish-brown to reddish-brown. Rostrum yellowish-brown to reddish-brown. Antennae reddish-brown, flagellomeres darker, base and apex of scape and pedicel dark brown. Anterior pronotal lobe yellowish-brown to reddish-brown, some specimens with dark brown areas on longitudinal medial sulcus and anterolateral margins. Posterior lobe reddish-brown dorsally with yellowish-brown lateral and posterior margins, lateral surfaces yellowish-brown, dark brown areas at anterior or dorsolateral margins and humeral angle. Scutellum yellowish-brown to reddish-brown. Legs yellowish-brown, most specimens with wide reddish-brown to brownish-black band at apices of femora and small dark area at apices of tibiae. Hernelytron brown with costal margin and veins of corium yellowish-brown. Abdomen venter reddish-brown. VESTITURE: Moderately setose. Anteocular lobe with short recumbent setae dorsally and laterally, short to moderately long erect setae on ventral surface. Postocular lobe with short, recumbent setae on dorsal and lateral surfaces, moderately erect setae scattered over surface, long, fine setae laterally. Anterior pronotal lobe with short, recumbent setae confined to setal tracts, long silky erect setae laterally. Posterior pronotal lobe with recumbent and sparse, erect setae. Meso-and metapleural surfaces with long silky erect setae. Scutellum with silky, erect setae. Clavus and corium of hemelytron with recumbent setae. Dorsum of abdomen with short erect setae, remainder of surface with short recumbent and short to moderately long erect setae, margin of connexivum fringed with erect setae. Exposed surface of pygophore with short to long erect setae. Apex of paramere with long erect setae. Female: (Fig. 71c, d) Similar to male, except for the following. Larger than male, total length 15.12-19.36 mm (mean 17.00 mm, Suppl. material 2). Coloration rather similar to that in male; more uniform on head. Lateral process on humeral angle spinous, usually longer. Pro-and mesofemoral diameters larger than that of metafemur.

Diagnosis
As with some species of the Zelus luridus species group, Z. exsanguis has a rather uniform greenish-brown coloration. Can be distinguished from most other species of the same species group by the humeral angle elevated to same level of and nearly continuous with disc. This is also seen in Z. ambulans, but the two species can be easily separated on the basis of coloration (Figs 22,71). Males can be distinguished from species of the Zelus luridus species group by the greatly enlarged apical part of the paramere (Fig. 72a, b), the medial process moderately broad, and the apex of the medial process somewhat narrowed.

Distribution
Mexico to Panama (Fig. 73) Taxon discussion  discussed the confusion over the use of the name Z. exsanguis. As this species appears highly similar to several other species, incorrect identification is common in museum specimens. Almost without exceptions specimens from the US identified as Z. exsanguis are actually Z. luridus. Champion (1898)

Diagnosis
The rather unique dorsal color pattern easily distinguishes this species from all other species in the genus.

Distribution
Southern Central America and northern South America (Fig. 75). Countries with specimen records: Colombia, Costa Rica and Panama. Male: (Fig. 76) Medium-sized, total length 12.71 mm (n=1, Suppl. material 2); slender. COLORATION: Nearly uniformly dark brown. Head dark brown; yellowish patch between eye and ocellus; inconspicuous, slender, medial yellow area on postocular lobe; ventral surface very slightly lighter than dorsal. Anterior pronotal lobe dark brown; posterior lobe slightly lighter than anterior lobe, somewhat reddish-brown; remainder of body surface and legs dark reddish-brown. VESTITURE: Sparsely setose. Head with moderately dense, short, recumbent on entire surface; dorsum also with short, spinelike setae, denser on anteocular lobe; ventral surface also with sparse, long, erect setae. Pronotum dorsal and lateral surfaces, pleura and sternites with short, recumbent setae and short to long, erect setae; recumbent setae dense on lateral surface of pronotum and pleuron; scutellum with sparse setation. Legs with sparse setation on femora and moderately dense setation on tibiae. Corium and clavus with sparse, short, a b

Diagnosis
Recognized by the entire body dark brown, the posterior pronotal lobe slightly lighter and somewhat reddish, the legs without bands; the humeral angle projected into spinous process. Distinguished among species of the Zelus panamensis species group by the curved medial process (Fig. 12).

Diagnosis
Recognized by the strongly contrasting black dorsum and yellow abdomen, the rather short postocular lobe, and the Sc not reaching apex of cubital cell. Other diagnostic characters shared with members of the Zelus vagans species group and the Zelus longipes species group include the unarmed rounded humeral angle and the spine-like setae on pronotum. Males can also be separated from other species of the Zelus vagans species group by the medial process apically tapered, somewhat pointed ( Fig.  11).

Etymology
The species epithet means 'soot' or painted black, referring to the black dorsal coloration of this species.

Distribution
Northern South America (Fig. 81). Countries with records: Colombia and Ecuador.  apically not evident or missing; basally separate throughout. Basal plate arm robust; basally fused; in lateral view slightly curved; bridge extremely short; extension of basal plate well expanded laterally onto arm, covering more than 1/2 of arm, curved.
Female: (Fig. 82c, d) Different from male as outlined below. Larger than male, total length 14.52-17.41 mm (mean 16.23 mm, Suppl. material 2). Dorsal surface, including hemelytron, lighter colored, nearly uniformly pale brown; quadrate cell and proximal margin of postcubital cell yellowish; lateral and ventral surfaces yellowish; legs without or with inconspicuous bands. Basiflagellomere subequal in diameter to pedicel. Process on humeral angle spinous, long.

Diagnosis
The posterior pronotal lobe usually orangish-brown; the medial process rather long, much longer than paramere; and the anterior side of medial process keeled medially at apex. In females the dorsal surface is nearly uniformly brown, the lateral and ventral surfaces yellowish, and the quadrate cell and proximal margin of postcubital cell yellowish.

Distribution
South America (Fig. 84  Male: (Fig. 85a, b) Large, total length 15.43-16.74 mm (mean 16.09 mm, Suppl. material 2); slender. COLORATION: Hed, anterior pronotal lobe, hemelytron, and legs dark brown to brownish-black; very inconspicuous, light-colored, rather thin, medial longitudinal stripe on postocular lobe. Ventral surface of head yellowish in some specimens. Ventral surface of abdomen in some specimens reddish-brown. Posterior pronotal lobe and mesopleuron orange or reddish-brown. Setae on corium golden. Abdominal segments 2-5 reddish-brown in some specimens. Variations minimal between specimens. VESTITURE: Densely setose. Dorsum of anteocular and anterior part of postocular with moderately dense, short, erect, spine-like setae; rest with sparse, short, erect or recumbent setae; posterior part of postocular nearly glabrous. Pronotum with dense, short, erect, spine-like setae on dorsal and lateral surfaces. Pleura with mixed spine-like and fine setae; scutellum with dense, short to long, semierect to recument setae. Legs with sparse setae; sundew setae on profemur sparse and randomly arranged. Corium and clavus with dense, recumbent, stout setae. Abdomen with moderately dense, short, semi-erect, fine setae. Apically with moderately long, erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.28. Postocular lobe in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye prominent; 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: 1.5: 0.4. Basiflagellomere diameter slightly larger than that of pedicel. Thorax: Anterolateral angle rounded, without projection; medial longitudinal sulcus evident throughout, deepening posteriorly. Disc distinctly elevated above humeral angle; humeral angle rounded, without projection. Scutellum moderately long; apex angulate, 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. 86) Pygophore: Ovoid; mid-lateral fold adjacent to paramere insertion; slightly expanded laterally near base of paramere in dorsal view. Medial process somewhat cone-shaped; moderately long; posteriorly directed, in less than forty-five degree with body axis; nearly straight; apex in posterior view blunt. Paramere: Cylindrical; long, nearly reaching apex of medial process; directed toward medial process; nearly straight; apex oblique; apical part not enlarged. Phallus: Dorsal phallothecal sclerite elongated; apical 1/3 of phallothecal sclerite tapering to apex, strongly convex, laterally rounded, not forming angle; apex with small medial emargination; basal part expanded laterally; posterior margin of foramen broadly concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally fused in part. Basal plate arm slender; separate; subparallel; in lateral view very slightly curved; bridge moderately long; extension of basal plate small and confined to apex of basal plate arm.
Female: (Fig. 85c, d) Similar to male, except for the following. Larger than male, total length 19.62-20.74 mm (mean 20.06 mm, Suppl. material 2). Dorsal coloration variable, ranging from nearly entirely orange to predominantly dark brown.

Diagnosis
Recognized by the posterior pronotal lobe orangish-brown; the legs uniformly blackishbrown, without bands; the humeral angle rounded; and the head and pronotum with spine-like setae (last two characters shared with species of the Zelus vagans species group and the Zelus longipes species group). Males can also be recognized by the medial process posteriorly directed and apical portion laterally compressed (diagnostic of Zelus vagans species group). The medial process is comparatively shorter than those in other species of the Zelus vagans species group (Fig. 11). Zelus gracilipes may be confused with Z. vespiformis and Z. errans, which also tend to show a combination of black and orange colors. The males of Z. gracilipes can be separated on the basis of the male genitalia. The females of Z. gracilipes have the entire membrane colored or opaque, whereas the known females of Z. errans have the anterior 1/2 clear or semi-translucent. Some females of Z. gracilipes exhibit entirely orange-brown hemelytron, a character also found in some specimens of Z. vespiformis.
As the ranges the these two do not appear to overlap, locality data may help in most cases separate the species. Additionally, Z. gracilipes is generally more slender than Z. vespiformis.

Etymology
From Latin gracilis, meaning slender.

Distribution
South America (Fig. 87 Rostrum yellowish-brown. Scape and pedicel yellowish-brown with small dark brown areas at base and apex. Flagellomeres dark reddish-brown. Anterior pronotal lobe reddish-brown dorsally; lateral surface yellowish-brown. Posterior lobe reddish-brown dorsally with margins and lateral surface yellowish-brown, humeral angle dark brown, lateral surfaces yellowish-brown. Scutellum yellowish-brown. Legs yellowish-brown with dark reddish-brown areas near apices of femora and tibiae. Hemelytron yellowishbrown to reddish-brown, costal margin of discal cell and adjacent area of corium dark brown. Abdominal dorsal surface yellowish-brown to brown, remainder of surface yellowish-brown. VESTITURE: Moderately setose. Head with recumbent setae on entire surface, more dense dorsally, long erect setae ventrally and ventrolaterally. Anterior pronotal lobe with scattered patches of recumbent and semi-erect setae dorsally, semi-erect and erect setae longer and more dense on lateral surface. Posterior lobe with recumbent setae over entire surface, some erect setae lateroventrally. Scutellum with erect and semi-erect setae. Corium and clavus with recumbent setae. Abdomen with short erect setae over entire surface, longer erect setae on margins of connexivum and ventrally on terminal segments. Exposed area of pygophore with erect setae. Apical half of paramere with short erect setae. STRUCTURE: Head: Cylindrical, L/W = 1.75. Postocular lobe moderately long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye unusually large; lateral margins much wider than postocular lobe; margins beyond dorsal and ventral outlines of head in lateral view. Ocellus elevated, large, diameter over 1.3x ocular-ocellar distance. Labium: I: II: III = 1: 1.9: 0.6. Basiflagellomere diameter larger than that of pedicel. Thorax: Anterolateral angle with inconspicuous subtuberculate projection; medial longitudinal sulcus appearing after anterior third. Posterior pronotal lobe with finely rugulose surface; disc slightly elevated above

Diagnosis
This is the only species in the genus with the margins of the eye exceeding outlines of the head both dorsally and ventrally. Compared to other species of the Zelus luridus species group (Fig. 3), the paramere is short, more slender and its apex very slightly expanded.

Etymology
The species epithet combines grandis, meaning large, with oculus, meaning eye, to indicate the prominently large-sized compound eye.

Distribution
Known only from type locality in Guatemala (Fig. 90).

Diagnosis
Can be distinguished by the following combination of characters: Humeral angle without or with minute processes; head and legs predominantly reddish; abdominal segments usually banded; pronotum with erect, nearly spine-like setae. Males can also be recognized by the paramere greatly curved at middle and distinctly tapered apically; the medial process curved and directed posteriad; and the dorsal phallothecal sclerite constricted and the apex truncate, without emargination. The only species within the range of Z. grassans with which the females may be confused is Z. ruficeps. The pronotal armature readily separate them, that of Z. ruficeps consisting of broad dentate lateral processes while those of Z. grassans are as given above.

Distribution
From Mexico to Panama (Fig. 93). Countries with specimen records: Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua and Panama.

Taxon discussion
There is a great amount of size and color variations in Z. grassans. The dorsal coloration varies from almost entirely yellowish with only a narrow transverse dark band near the posterior margins of the pronotal lobes, to almost entirely dark brown. The legs show a similar range of coloration, from almost entirely light to entirely dark. The contrasting black and orange or red colors and the banded abdomen in many specimens of Z. grassans suggest that they may be mimics of Dysdercus, whose members have similarly strongly contrasting red and black colors. They have been observed to co-occur on the same plant (Zhang, unpublished), indicating that this may be a case of aggressive mimicry.  Lethierry and Severin, 1896, p. 152, cat.;Wygodzinsky, 1949a, p. 49, checklist;Wygodzinsky, 1949b, p. 336, note;Wygodzinsky, 1957, p. 264, 268, list and junior syn. of Z. obscuridorsis; Hart, 1987, p. 297, redescription, note, fig, key, lectotype desig. and stat. rev.; Maldonado, 1990, p. 330, cat. Zelus carvalhoi Wygodzinsky, 1947, p. 428-431, orig. descr. andfig.;Zikan and Wygodzinsky, 1948, p. 17, list;Wygodzinsky, 1949a, p. 48, checklist;Wygodzinsky, 1949b, p. 336, note;wygodzinsky, 1957, p. 264, 268, list  Male: (Fig. 94a, b) Small, total length 9.96-11.91 mm (mean 11.03 mm, Suppl. material 2); slender. COLORATION: Dorsal 1/2 brown to dark brown, ventral surface yellowishbrown. Rostrum yellowish-brown. Antennae reddish-brown to dark brown, bases and apices darker than shaft on scape and pedicel. Anterior pronotal lobe light to dark brown, yellowish-brown lateroventrally. Posterior lobe light to medium reddish-brown; humeral angle usually darkened, brownish-black; lateral surface lighter ventrally. Legs yellowish-brown, femora with reddish-brown to brownish-black bands at apices and basad to apical swelling, tibiae with at least two such dark bands, tibiae darkened toward apex. Hemelytron brown to dark brown. Dorsum of abdomen reddish-brown to dark brown, connexival margins and remainder of surface yellowish-brown. VESTITURE: Entire surface of head with short recumbent setae, short to moderate semi-erect and erect setae on lateroventral and ventral surfaces. Anterior pronotal lobe entire surface with short recumbent setae, confined to setal tracts dorsally, some longer erect setae laterally. Posterior lobe entire surface with short recumbent setae, some erect setae laterally. Recumbent setae over clavus and corium. Abdomen with short erect setae dorsally, lateral and ventral surfaces with short recumbent and scattered erect setae. Exposed surface of pygophore with short recumbent and short to long erect setae. STRUCTURE: Head: Elongated, L/W = 2.79. Postocular lobe long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye smallish; lateral margin only slightly wider than postocular lobe; dorsal and ventral margins removed from surfaces of head. Labium: I: II: III = 1: 2.0: 0.4. Basiflagellomere diameter larger than that of pedicel. Thorax: Anterolateral angle subtuberculate to tuberculate; medial longitudinal sulcus shallow near collar, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with dentate or short spinous projection. Scutellum short; apex angulate. Legs: Very slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell small, elongate; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 95) Pygophore: Ovoid; not expanded laterally in dorsal view. Medial process slender; long, slightly shorter than paramere; laterally somewhat compressed; semi-erect; apically recurved; apex in posterior view acute, without modification. Paramere: Cylindrical; long, achieving apex of medial process; directed posteriad, basal half sharply curved towards medial process; apically slightly recurved; apical part not enlarged. Phallus: Dorsal phallothecal sclerite shield-shaped; small indentation of lateral margin at about midpoint; apical portion of phallothecal sclerite distinctly tapered, dorsal surface folded in middle, laterally rounded, not forming angle; posterior margin of foramen inversely Vshaped. Struts apical portion missing or not evident; basally separate. Basal plate arm moderately robust; basally fused; in lateral view very slightly curved; bridge extremely short; extension of basal plate heavily sclerotized, laterally expanded onto arm.

Zelus illotus
Female: (Fig. 94c) Similar to male, except for the following. Larger than male, total length 12.35-14.28 mm (mean 13.68 mm, Suppl. material 2). Coloration lighter than in male; pronotal coloration not as variable, usually concolorous yellowish-brown to dark brown.

Diagnosis
Among species of the Zelus nugax species group (Fig. 5), males of Z. illotus can be recognized by the paramere slender and long, curved in middle and recurved apically and the medial process also strongly recurved. The males of Z. pedestris and Z. nugax have straight blade-like medial processes. The females of this species cannot be consistently separated based on any character or combination of characters yet discovered from females of Z. pedestris and Z. nugax. Most females of Z. impar have almost no erect setae on the dorsal surface of the posterior pronotal lobe while most females of the other two species have readily noticeable erect setae on this area.

Diagnosis
Recognized by the slender, curved, laterally compressed, and apically tapered medial process (shared with members of the Zelus nugax species group; Fig. 5). Distinguished from Z. nugax and Z. pedestris by the recurved medial process. Similar to Z. illotus in having a recurved medial process, but is differentiated by the straight paramere.

Distribution
Panama and Northern South America and adjacent islands of the Caribbean (Fig. 99).
Countries with records: Colombia, Panama (record not mapped), Trinidad and Tobago and Venezuela.

Taxon discussion
Hart (1987) designated a neotype for Z. impar because the original type material of that species was destroyed during World War II. This neotype specimen was at that time deposited in the private collection of J. C. Elkins, Houston,Texas. This specimen was eventually transferred to and deposited at TAMU, instead of AMNH as the author had indicated.

Diagnosis
The Zelus inconstans Champion, 1899, specimen record map spine-like setae on pronotum and are in the Zelus longipes species group. Males can also be recognized by the paramere apically greatly projected dorsad (Fig. 101a), nearly 90 degree; and the apex of the medial process folded posteriorly and ventrally. Very similar to Z. mimus, but the medial process is much shorter and broader.

Distribution
Southern Central America and northern South America (Fig. 102). Countries with records: Colombia, Panama and Peru.

Taxon discussion
Coloration variations are mainly seen on the pronotum. The two males from Panama show similar coloration, the central 1/3 of the pronotum being lighter than the margins. The Colombian male, however, has a nearly unicolorous pronotum. The females from Panama have pronotal coloration ranging from reddish-brown to brown, the lighter color apparently being more common. The single female from Colombia has the lighter pronotum while that of the Peruvian specimen is dark brown. The lectotype and two paralectotypes exhibit the lighter coloration. The banding patterns of the legs appear to be somewhat variable in Panama, the only area from which several specimens are available for comparison. Male: (Fig. 103a,  Female: (Fig. 103c, d) Similar to male, except for the following. Larger than male, total length 19.01-21.96 mm (mean 20.53 mm, Suppl. material 2). Whitish area on distal part of corium smaller, almost non-existent in some specimens; posterior pronotal lobe, corium and clavus entirely yellowish-brown in some specimens.

Diagnosis
Among closely related species in the Zelus armillatus species group with overlapping distribution ranges, Z. janus is the only species with the humeral angle elevated to about same level as the disc of the posterior pronotal lobe. This species is much larger than and the coloration different from two other species sharing this feature, Z. exsanguis and Z. ambulans, both from a different species group. Other characters useful for diagnosis include the dorsal surface usually mostly brown and the lateral and ventral surfaces yellowish-brown; the abdominal segment each with single dark spot anteriorly; and in males the medial process narrower, relatively short.

Distribution
Southern Texas to Central America (Fig. 105). Countries with records: USA (Texas), Belize, Guatemala, Honduras, Mexico and Nicaragua.

Taxon discussion
It is rather difficult to distinguish Z. janus from Z. armillatus on the basis of the male genitalia alone, the only two species in the genus where such distinction cannot be made. As the humeral angle of the posterior pronotal lobe are raised to the level of and are nearly continuous with the disc, however, it is quite easy to separate specimens of these species. There is further divergence in coloration and pattern between the two which may be seen in the descriptions of these species. A sympatric and closely related species, Z. litigiosus, also has the disc elevated above the humeral angle and is easily distinguished. Zelus janus has a somewhat more uniform brown dorsal coloration, whereas the color pattern in Z. litigiosus is more variable.  Male: (Fig. 106a,  Zelus kartabensis Haviland, 1931, specimen record map yellowish. Legs brown; meso or metafemora sub-basally, medially, or sub-apically with yellowish band(s); meso and metatibiae occasionally with inconspicuous medial yellow band; fore leg never banded. VESTITURE: Sparsely setose. Dorsum primarily consisting of moderately dense, short, erect or recumbent setae; short spine-like setae also on dorsum of head, more concentrated on anteocular lobe, and on pronotum. Sundew setae on profemur sparsely and randomly distributed. Microtrichia throughout posterior margin of membrane of hemelytron. Abdominal venter with short, recumbent setae, intermixed with sparse, moderately long, erect setae. Setae on pypophore short to long, recumbent to erect; paramere apical 1/2 with dense, very long, almost as long as paramere, erect, apically curved setae, directed mediad. Female: (Fig. 106c, d) Different from male as outlined below. Larger than male, total length 15.49-17.45 mm (mean 16.53 mm, Suppl. material 2). Mostly yellowish to greenish. No dark dorsal markings of anteocular lobe; remainder of dark cranial markings less pronounced than in male. Posterior margin of posterior pronotal lobe and corium dark. Femora unbanded. Spine-like setae more conspicuous than in male.
Basiflagellomere not swollen basally. Humeral angle nearly elevated to level of pronotal disc; lateral processes spinous.

Diagnosis
The dorsal surface of posterior pronotal lobe uniformly dark brown. The paramere gradually enlarged, somewhat club-shaped; the medial process with ridge-like medial elevation through apical 1/2.
Zelus kartabensis is most similar to Z. kartaboides, and the differences between the two species are presented in the diagnosis of the latter species.

Distribution
South America (Fig. 108). Countries with records: Brazil, Guyana and Suriname. Male: (Fig. 109) Medium-sized, total length 11.22-11.61 mm (mean 11.6 mm, Suppl. material 2); Coloration, vestiture and structure including genitalia very similar to Z. kartabensis except for the following. COLORATION: Lateral surface and posterior margin of posterior pronotal lobe, and scutellum yellowish, lighter than remainder of body surface. GENITALIA: (Fig. 110) Pygophore: Posterolateral rim of pygophore in smaller angle with body long-axis in lateral view, nearly horizontal; lateral protrusion on posterodorsal rim of pygophore more pronounced, proximal side of arch extending down as process. Medial process short. Paramere: Paramere more strongly curved, banana-like; diameter uniform throughout, apex not enlarged, base slightly constricted. Phallus: Struts apically diverging, V-shaped.

Diagnosis
The posterior margin of the pronotum and the scutellum yellowish, strongly contrasting to the remaining dark brown dorsal surface, makes this species easily recognizable among all species of Zelus, including the very similarly looking Z. kartabensis. It can also be recognized by the medial process with ridge-like medial elevation through apical 1/2 (also in Z. kartabensis and Z. chamaeleon). It is separated from Z. kartabensis by the paramere uniquely shaped like a banana and its diameter uniform throughout. Some specimens of Z. armillatus, Z. conjungens, Z. longipes and Z. ruficeps also exhibit a predominantly dark brown pronotum with posterior and/or dorsolateral margins yellow or orange, but they are much more larger and robust than Z. kartaboides and the male genitalic structures are very different.

Etymology
The specific epithet indicates that this species is rather similar to Z. kartabensis.

Distribution
South America (Fig. 111). Countries with records: Brazil, Colombia, Ecuador and Peru. Male: (Fig. 112) Medium-sized, total length 10.42-11.55 mm (mean 10.98 mm, Suppl. material 2); slender. COLORATION: Head dark brown; inconspicuous yellowish patch between eye and ocellus; medial, yellow stripe on postocular lobe; ventral surface yellowish-brown, lighter than dorsum. Pronotum and scutellum dark brown. Abdomen yellowish-brown. VESTITURE: Sparsely setose. Dorsum of head with moderately dense, short, recumbent setae and sparse, short, erect, somewhat spine-like setae; ventral surface with sparse, short, recumbent setae and few moderately long, erect, fine setae. Pronotum with very sparse, short, erect setae over dorsal surface, some setae curved apically, appearing recumbent; moderately dense, short to moderately long, recumbent setae on lateral surface and pleura, intermixed with semi-erect 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, erect setae, intermixed with sparse, long, erect setae. Apical half of dorsal surface with moderately dense, medium-length, semi-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 prominent; lateral margin 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 larger than that of pedicel. Thorax: Anterolateral angle bearing small, somewhat acute projection; medial longitudinal sulcus evident throughout, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with spinous processes. Scutellum moderately long; apex angulate, very slightly projected upward. Legs: Slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell small, elongate; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 113) Pygophore: Elongate ovoid; lightly sclerotized expansion below paramere; not expanded laterally in dorsal view. Medial process cylindrical; slender; moderately long; laterally compressed towards apex; anterior surface towards apex ridged; minute spicules on posterior surface; posteriorly directed; curved at middle; apex in posterior view acute, with small hooklike projection. Paramere: Cylindrical; moderately long, not reaching medial process; directed posteriad; basally slightly narrower; slightly curved ventrad; apical part not enlarged. Phallus: Dorsal phallothecal sclerite somewhat ovoid; sclerotization reduced (yet not absent) on dorsal surface close to posterior margin of foramen; expansion of lateral margin at about mid-portion small; apical portion of phallothecal sclerite gradually tapering, distinctly keeled medially; apex acute; posterior margin of foramen concave. Struts attached to dorsal phallothecal sclerite; apically separate, connected by bridge; basally separate. Basal plate arm robust; basally fused; in lateral view nearly straight, very slightly curved; bridge extremely short; extension of basal plate expanded laterally onto arm, covering more than 1/2 of arm, curved.

Diagnosis
Recognized by the nearly uniformly dark brown dorsum; the abdomen light-colored, pale yellowish-brown; the posterolateral rim with lightly sclerotized expansion between paramere and medial process; the medial process curved at middle; the anterior surface of the medial process carinate; the apex of the medial process hooklike, the curvature of paramere small; the dorsal phallothecal sclerite with strong carination at apical part, the lateral expansion close to basal arm. Most similar to Z. filicauda, but the medial process is shorter and not as strongly curved and the paramere curvature is weaker.

Distribution
South America and adjacent islands of the Caribbean (Fig. 114). Countries with records: Ecuador, Guyana and Trinidad and Tobago.  Male: (Fig. 115a, b, c, d) Small, total length 9.94-11.44 mm (mean 10.82 mm, Suppl. material 2); slender. COLORATION: Head mostly yellowish; some specimens with submedial stripes on anteocular lobe; variable brown areas on dorsal surface of postocular lobe, anteriorly broad, narrowing and fusing posteriorly, medially separated by yellowish stripe. Anterior pronotal lobe dark brown; posterior lobe variable, dark brown, orange, or medially and laterally yellowish-brown; pleura brown, mixed with yellow parts. Proportion of dark brown and yellow on posterior pronotal lobe variable, some specimens entirely dark and some entirely yellowish or orange. Scutellum broadly medially yellowish, lateral parts dark brown, some specimens nearly entirely dark or yellowish. Hemelytron uniformly dark brown, corium in some specimens yellowish. Profemur and protibia dark brown, sometimes with single inconspicuous yellowish band; meso and metafemora dark brown, with two or three yellow bands, sometimes basal band rather broad; meso-and meta-tibiae usually dark brown with single yellow band. VESTITURE: Moderately setose. Body surface with mostly short, recumbent setae, erect setae sparse. STRUCTURE: Head: Cylindrical, L/W = 2.24. Postocular lobe long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye moderately sized; lateral margin much wider than postocular lobe; dorsal and ventral margins removed from surfaces of head. Labium: I: II: III = 1: 1.8: 0.5. Basiflagellomere diameter slightly larger than that of pedicel. Thorax: Anterolateral angle bearing small projection; medial longitudinal sulcus evident only on posterior 1/2, deepening anterior to transverse sulcus of pronotum. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with dentate projection. Scutellum moderately long; apex angulate, slightly projected upward in some specimens. Legs: Moderately robust. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell large and broad; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 116 Female: (Fig. 115e, f) Different from male as outlined below. Larger than male, total length 12.38-14.14 mm (mean 13.67 mm, Suppl. material 2). Head, dorsum of pronotum and corium reddish orange, entirety or portion of posterior pronotal lobe dark brown in some specimens; membrane dark brown; lateral surface of pronotum, pleura and abdomen yellowish, with dark stripes; legs reddish, with dark bands. Hemelytron attaining apex of abdomen.

Diagnosis
The strongly convex pronotum distinguishes this species from most other species of the genus. The males can be distinguished by the relatively small size (mean 10.82 mm); the dorsum of the posterior pronotal lobe usually with lighter colored, pale brown, with medial stripe; the broad, pentagonal, apically angulate medial process; the short, blade-like process on dorsal phallothecal sclerite; and the ridge mesad to the blade-like process. In females the head, pronotum and corium are usually orangish-brown to reddish.

Diagnosis
The black dorsal and red ventral coloration is distinctive of this species. Other diagnostic characters include the legs uniformly black and the posterior pronotal lobe with medial depression.

Taxon discussion
Zelus leucogrammus is one of the most distinctive species among Zelus. It can be easily recognized the red and black coloration, the medial depression on the posterior pronotal lobe. Variations in coloration are minimal and is usually seen in the size the dark area on the posterior pronotal lobe.
According to Dr. Heinz Wundt at ZSM (pers. comm.), the type material for this species was destroyed during World War II. The original description lists this species from the Amazon River. As this is such a distinctive species, it is not felt that neotype is needed. Male: (Fig. 121a, b Female: (Fig. 121c, d, e, f) Larger than male, total length 22.52-24.06 mm (mean 23.29 mm, Suppl. material 2). Coloration variable; yellowish or reddish with dark spots or markings.

Diagnosis
Recognized by the large and slender body and the posterior pronotal lobe bearing a pair of tubercles. Males can be easily recognized by the black coloration with white markings on scutellum and abdomen and females yellowish or reddish with black spots and markings. Among males of the Zelus armillatus group (Fig. 8), the paramere of Z. annulosus is more than 2x longer than the medial process. Zelus amblycephalus and Z. annulosus also have long parameres, but these are apically curved, whereas it is straight in Z. annulosus.

Etymology
The specific epithet is a patronym, named after Dr. James Lewis, in honor of his contribution to the curation of Heteroptera of Costa Rica at INBio. Without his and his fellow scientists' work the discovery of this species would not have been possible.

Distribution
Central America (Fig. 123). Countries with records: Costa Rica and Panama.  Male: (Fig. 124a, b, c, d) Large, total length 17.10-18.80 mm (mean 18.15 mm, Suppl. material 2); robust. COLORATION: Brown, brownish-black, sometimes with orange or red. Dorsal and lateral surfaces of head usually brownish-black, ventral surface yellowish-brown; variable amount of yellowish-brown on anteocular lobe; yellowishbrown patch usually between eye and ocellus and medially on postocular lobe. Scape and pedicel with yellow and black bands. Areas of setal tracts on anterior pronotal lobe lighter than glabrous surface, difference subtle in some specimens. Posterior lobe usually uniformly brown, orange or red; lateral surfaces lighter in some specimens. Scutellum dark brown. Corium and clavus usually uniform, orange, brown or dark brown; some specimens with distal part lighter; membrane dark brown. Legs usually yellowish-brown with black bands, usually one on tibiae and two or three on femora; completely black in some dark specimens. VESTITURE: Densely setose. Head with both recumbent and erect setae dorsally, and predominantly short, recumbent setae ventrally. Anterior pronotal lobe with long erect setae, mainly occupying setal tracts; posterior pronotal lobe with fine, erect setae. Abdomen with short, recumbent setae, interspersed with long, erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.25. Postocular lobe long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye smallish; lateral margin only slightly wider than postocular lobe; dorsal margin removed from postocular transverse groove, ventral margin attaining ventral surface of head. Labium: I: II: III = 1: 1.4: 0.4. Basiflagellomere diameter subequal to that of pedicel. Thorax: Anterolateral angle bearing small protuberance; medial longitudinal sulcus evident throughout, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with short tuberculate process. Scutellum moderately long; apex angulate. Legs: Slender. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell large and broad; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 125) Pygophore: Rounded; slightly expanded laterally near base of paramere in dorsal view; posteriorly expanded sac-like sclerite between paramere and medial process. Medial process cylindrical; slender; moderately long; posteriorly directed; straight; apex in posterior view rounded, with very inconspicuous lateral prongs. Paramere: Cylindrical; moderately long, not reaching apex of medial process; directed posteriad; nearly straight; apical part not enlarged. Phallus: Dorsal phallothecal sclerite somewhat squarish; sharp laterally oriented process close to posterior margin of foramen and basal arms; apical portion of phallothecal sclerite not distinctly tapered, flat, lateral margin narrowly angulate; apex rounded; posterior margin of foramen broadly concave, medially deeper. Struts attached to dorsal phallothecal sclerite; apically fused; basally mostly separate, moderately fused. Basal plate arm moderately robust; separate; diverging; in lateral view very slightly curved; bridge short; extension of basal plate small and confined to apex of basal plate arm.

Diagnosis
Among species in the Zelus armillatus species group occurring in overlapping geographical regions, Zelus litigiosus can be easily distinguished from Z. janus by the elevated disc of the posterior pronotal lobe. It can be separated from Z. sulcicollis by the flat or slightly convex disc of the posterior pronotal lobe, and that being depressed in Z. sulcicollis.

Diagnosis
Although highly variable, the black and red coloration is distinctive of Z. longipes. The combination of size, coloration, rounded humeral angle, and raised anterior pronotal lobe serves to separate this species from any others that may cause confusions. Males can also be recognized by the long and slender medial process, the apex slightly folded posteriad, and the long paramere, exceeding apex of medial process. Among the Zelus longipes species group (Fig. 10), Z. bahiaensis also has a long paramere clear exceedingly medial process, but the two species can be readily separated on the basis of coloration.

Distribution
Southern parts of US, Mexico, Central America, the Caribbean, Northern South America, Paraguay and Southern Brazil (Fig. 129)

Taxon discussion
Zelus longipes is a highly variable species. In any given area there is a wide range of color and color pattern variations. The dorsal coloration can vary from nearly entirely orange brown, through various patterns of orange-brown and brownish-black, to almost completely black. The habitus images provided here only represent a subset of the range of variations. The most typical form is one with alternating orange and black areas on the dorsum, abdominal venter orange or reddish-brown, each segment black anteriorly, and legs black with two yellow rings medially.    Zelus longipes (L., 1767), specimen record map

Figs 130, 131, 132
Male: (Fig. 130a, b) Medium-sized, total length 13.21-15.27 mm (mean 13.89 mm, Suppl. material 2), slender. COLORATION: Yellowish-brown through reddish-brown to brownish-black, colors more vibrant in live individuals. Anteocular lobe uniformly yellowish-brown to reddish-brown or with dorsum variably reddish-brown to brownishblack with lighter lateral and ventral surfaces. Postocular lobe reddish-brown to brownish-black dorsally and laterally with mid-dorsal and occasionally circumocellar areas yellowish-brown, ventral surface yellowish-brown. Rostrum uniformly yellowishbrown to light reddish-brown. Antennae light reddish-brown, base and apex of scape and pedicel sometimes slightly darker than shaft. Anterior pronotal lobe yellowish-brown to brownish-black, light specimens usually with darker coloration on anterior portion of medial longitudinal sulcus and/or with setal tracts darker, many darker specimens with lateral margins and/or collar yellowish-brown, remainder of surface yellowish-brown. Dorsum of posterior lobe reddish-brown to brownish-black, lateral margins and usually posterior margins yellowish-brown, lateral processes of lighter specimens usually brownish-black, remainder of surface yellowish-brown. Scutellum reddish-brown to brownish-black with apex and sometimes posterior mid-dorsal surface yellowish-brown. Femora yellowish-brown to light reddish-brown, usually with dark bands or dorsal markings near apices, tibiae yellowish-brown to dark reddish-brown, usually slightly darker than femora. Hemelytron yellowish-brown to reddish-brown, some darker specimens with lighter veins in clavus and corium and/or lighter area along costal margin. Dorsum of abdomen reddish-brown, lateral and ventral surfaces yellowish-brown to light reddish-brown. VESTITURE: Moderately setose. Anteocular lobe with short, recumbent and erect setae dorsally, erect setae predominating on vertex, sparse short erect and semi-erect setae on ventral half. Postocular lobe with mostly recumbent, some erect setae dorsally; lateroventral and ventral surfaces with moderate to long erect and scattered recumbent setae. Anterior pronotal lobe with short erect and recumbent setae over surface, confined to setal tracts dorsally, long erect setae laterally. Dorsum of posterior lobe with short recumbent setae, some short recumbent setae laterally, long erect setae lateroventrally. Lateral surface of scutellum with moderate to long erect setae, elevated dorsal surface nearly bare. Clavus and corium with inconspicuous short erect and recumbent setae. Abdomen dorsally with short, sparse, erect setae, remainder of surface with short erect and recumbent setae and some scattered longer erect setae. Exposed area of pygophore with short recument and short to long erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.44. Postocular lobe moderately long; in dorsal view anteriorly gradually narrowing, posterior portion somewhat constricted. Eye moderately sized; lateral margin only slightly wider than postocular lobe; in lateral view removed from both dorsal and ventral surfaces of head. Labium: I: II: III = 1: 1.8: 0.4. Thorax: Anterolateral angle bearing small protuberance; medial longitudinal sulcus shallow near collar, deepening anterior to transverse sulcus of pronotum. Posterior pronotal lobe with rugulose surface; disc elevated above humeral angle; humeral angle armed, with tuberculate to long spinous lateral processes. Scutellum moderately long; apex angulate, slightly projected upward. Legs: Slender. Profemoral diameter slightly larger than mesofemoral diameter, metafemoral diameter slightly less than that of mesofemur. Hemelytron: Surpassing apex of abdomen by about length of abdominal segment seven; quadrate cell small, elongate; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 131 Female: (Fig. 130c,

Diagnosis
Recognized by the following combination of characters: Yellow-green to green-black; apices of femora with reddish or brown bands; disc elevated above humeral angle. As with other members of the Zelus luridus species group, the medial process is triangular, its base distinct from rest of the ventral rim of pygophore and apex without modification (Fig. 131a, b). Zelus luridus has the pronotal disc noticeably elevated above and not continuous with the humeral angle, thus distinguishing it from both Z. exsanguis and Z. ambulans. Males of the two differ in significant ways in paramere and medial process a b c d Figure 131.  (Fig. 3). Among males of the Zelus luridus species group, Z. luridus is similar to Z. antiguensis, and Z. grandoculus in having the paramere apex not greatly expanded. The medial process is narrower than that in Z. antiguensis. The eyes are not as prominent as those of Z. grandoculus. Among species with overlapping distributions, Z. luridus may be confused with Z. renardii. However, Z. renardii usually has the corium reddish, whereas it is greenish or dark brown in Z. luridus.

Distribution
North America (Fig. 132). Countries with records: Canada, Mexico and USA.

Taxon discussion
This is one of the most commonly collected species in this genus.  discussed intraspecific variations and species identity confusions, which we briefly summarize here. Individuals in the western population (i.e., Southeastern Arizona) are larger than those from the eastern population. Eastern males have parameres somewhat more enlarged apically (Fig. 131a), and the apical half of the dorsal phallothecal sclerite is less expanded laterally (Fig. 131c). Since Champion (1898) synonymized Z. luridus and Z. ambulans under Z. exsanguis, almost all specimens collected from the US were labeled as Z. exsanguis. Zelus luridus has the disc clearly elevated above, and not continuous with, the humeral angle which distinguishes it from Z. exsanguis.

Diagnosis
Distinguished by the small size; the robust form, the humeral angle rounded, without projection; the profemur much longer than the metafemur (1.20x); the profemoral length being less than 20.0x the profemoral width (16.94x). The paramere base not distinctly a b Figure 134. Zelus mattogrossensis Wygodzinsky, 1947, specimen record map constricted; the medial process slender, apex angulate and bearing subapical medial protrusion; the presence of blade-like process on dorsal phallotheca and the process not extending beyond mid-point.

Distribution
South America (Fig. 135). Countries with records: Bolivia, Brazil and Paraguay.

Diagnosis
The humeral angle rounded; the pronotum with spine-like setae; and the colors usually consisting of yellow, orange, red and black. The anterior pronotal lobe is rather small, margins not laterally expanded, nearly continuous with lateral margins of posterior lobe, dorsally nearly flat, not bulging. The small anterior lobe together with the regularly sized posterior lobe gives the pronotum a triangular appearance. Most similar to Z. fuliginatus, but the postocular lobe is shorter.

Distribution
Southern Central America and South America (Fig. 137). Countries with records: Bolivia, Brazil, Costa Rica, Colombia, Ecuador and Peru.

Taxon discussion
On the basis of the spine-like setae on the head and pronotum and the rounded humeral angle, Zelus means would be placed in either the Zelus longipes species group or the Zelus vagans species group. Due to its close resemblance to Z. fuliginatus, this species is most likely part of the Zelus vagans species group.
This species appears to have a highly variable color pattern in all areas of its distribution. The clavus and corium range from entirely brownish-black to almost entirely light yellowish-brown. The dorsum of the pronotal lobe may be entirely brownish-black, but is usually variably patterned brownish-black and reddish-brown.

Diagnosis
Dark brown coloration predominating dorsally in most specimens, posterior pronotal lobe laterally yellowish. Among species that have overlapping distributions (Southern Mexico and Central America), the coloration of Z. mimus is unique. Males can also be recognized by the paramere apically greatly projected dorsad (Fig. 139a); and the apex of the medial process folded posteriorly. Similar to Z. inconstans, but the medial process is much longer and more slender.

Distribution
Southern Mexico and Central America (Fig. 140) Male: (Fig. 141) Small, total length 7.80-10.13 mm (mean 9.00 mm, Suppl. material 2); slender. COLORATION: Yellowish-brown to brownish-black, shining. Antennae yellowish-brown to brown, apex of scape and pedicel and base of pedicel darker than remainder of segments. Anterior pronotal lobe yellowish-brown to brownish-black, shining. Posterior pronotal lobe yellowish-brown to brownish-black, pattern variable, medial and lateral areas lightest in dark specimens. Scutellum yellowish-brown, apex lighter than surrounding area. Legs yellowish-brown with two to four brown rings on apical 1/2 of femora, variable brown areas on tibiae. Clavus and corium yellowishbrown to brown, darkest along costal margin, membrane nearly clear, veins dark brown. Dorsum of abdomen yellowish-brown to dark brown, pattern variable, lateral surfaces yellowish-brown. Pygophore yellowish-brown to brown, parameres dark brown to brownish-black. VESTITURE: Moderately setose. Scattered erect and recumbent setae over entire surface of head, longer ventrally. Anterior pronotal lobe with very sparse a b c d Figure 142.
Female: Similar to male, except for the following. Larger than male, total length 10.70 mm (Suppl. material 2).

Diagnosis
This species can be readily recognized by the small size (<10.2 mm) and the disc of the posterior lobe with spinous tubercles. The medial process is broadly triangular, apex without modification (shared with the Zelus tetracanthus species group). Can be recognized among the Zelus tetracanthus species group by the relatively long paramere, exceeding the medial process.

Distribution
Southern Central America, South America and adjacent islands of the Caribbean (Fig.  143). Countries with records: Brazil, Colombia, Ecuador, Panama, Suriname, Trinidad & Tobago and Venezuela.

Taxon discussion
Zelus minutus shows some variations through its range. The Panamanian specimens have noticeably shorter paramere and the apex of the dorsal phallothecal sclerite is somewhat less emarginate.

Diagnosis
Recognized by the conspicuous black and yellow color pattern, resembling vespid wasp; and the meso-and metafemora with at least three dark bands three yellow bands. Among males of the Zelus panamensis group (Fig. 12) Zelus nigromaculatus Champion, 1899, specimen record map the shortest paramere and the longest medial process and the medial process is the most erect in this species group.

Diagnosis
The slender, cylindrical paramere and the laterally compressed medial process can separate males of this species from most other species of the genus. Difficult to distinguish from Z. pedestris, but the paramere apex is generally more truncate, the dorsal phallothecal sclerite usually without lateral indentation, and the basal plate arms are separate in most specimens (see taxon discussion).

Taxon discussion
This is by far the most widespread species of the Zelus nugax species group, and one of the most widespread members of the genus. In Ecuador, Mexico and Central America, Z. nugax is apparently among the most common species of reduviids, occurring in second growth foliage and tall grasses throughout its range. Other than a variation in color from yellowish-brown to dark brown in any given area and a more noticeably produced scutellar apex in South America, there are little readily apparent external variations in the species. In the internal male genitalia, however, we find some noticeable geographic variations. In South America the basal plate arms are apparently always separate, while there seems to be tendency toward fusion of these arms as one progresses northward to Mexico.
The following discussion on some of the diagnostic characters may be useful for separating species when confusions arise, but as we have not clearly defined the boundaries between Z. nugax and Z. pedestris, reliable identification may not be achieved at all times. We will discuss this in the next paragraph. The straight bladelike medial process of Z. nugax distinguishes it from Z. impar and Z. illotus, both of which have similar appearances to Z. nugax, but both have recurved medial processes. The less rounded pygophore and more blunted paramere separate this species externally from most male specimens of Z. pedestris. Although it is difficult to separate the females of Z. nugax from Z. illotus and Z. pedestris, there appears to be some differences among these species. The normal lack of erect setae on the dorsal surface of the posterior pronotal lobe of Z. illotus and lower posterior margin of the anterior pronotal lobe of Z. pedestris usually serve as diagnostic characters for someone with large series and a familiarization with all three species.
It remains unresolved if Z. nugax and Z. pedestris are distinct species. They are currently delimited based on several characters of the male genitalia, which, however, do not appear to be fixed. With regards to four characters (paramere apex shape, fusion of basal plate arms, lateral indentation on dorsal phallothecal sclerite, and basal plate arm extension) used in delimitation, the following combinations have been observed: (1) paramere apex acute, basal plate arms fused, phallothecal indentation present and basal plate arm extension present and expanded laterally (observations based on specimens from Santa Catarina, Brazil); (2) paramere apex truncate, slightly enlarged, somewhat diamond-shaped, basal plate arms fused, indentation absent and extension present, laterally expanded (La Molina, Peru); and (3) paramere apex truncate, slightly enlarged, basal plate arms separate, phallothecal indentation absent and extension absent or inconspicuous (El Valle, Panama). Following the phylogenetic species concept sensu Wheeler and Platnick (Wheeler and Platnick 2000), we would call each population a different species. We restrained from doing this as we have not thoroughly examined the ranges of variations.
Both the types of Z. nugax and Z. pedestris are females, further complicating the application of the names as we have not been able to distinguish females of the two species. The type of Z. nugax is from Mexico and that of Z. pedestris from South America (country not recorded). Zelus nugax and Z. pedestris overlap broadly in northern South America, but Z. nugax appears to be absent South of Peru and in most of Brazil and Z. pedestris not recorded from North and Central Americas. This geographic pattern currently serves as an additional means to apply the names. Male: (Fig. 150a, b) Medium-sized, total length 11.25-12.90 mm (mean 11.96 mm, Suppl. material 2); slender. COLORATION: Dark brown. Head orangish or reddish; remainder of body surface nearly uniformly dark brown; legs not banded or with inconspicuous bands. VESTITURE: Sparsely setose. Entire surface of head with short, recumbent setae; sparse, short, erect, spine-like setae on dorsal surface, denser on anteocular lobe; few long, erect, fine setae on ventral surface. Pronotum with sparse, short, erect, spine-like setae on dorsum, very sparse on anterior lobe, setal tracts indistinct; lateral surface of pronotum and pleura with moderately long, semi-erect or Zelus panamensis Zhang & Hart, sp. n., specimen record map recumbent setae; scutellum with moderately long, erect, spine-like setae. Legs with very sparse setation; sundew setae on profemur very sparse. Corium and clavus with mix of short, recumbent or erect setae and long, erect, fine setae. Abdomen with moderately dense, short recumbent setae, intermixed with sparse, long, erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.35. 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 in lateral view. Labium: I: II: III = 1: 1.9: 0.4. Basiflagellomere diameter larger than that of pedicel. Thorax: Anterolateral angle bearing small, somewhat acute 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 projection. Scutellum moderately long; apex angulate, slightly projected upward. 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. 151) Pygophore: Elongate ovoid; lightly sclerotized expansion below paramere; not expanded laterally in dorsal view. Medial process cylindrical; slender; moderately long, about as long as paramere; laterally slightly compressed towards apex; minute spicules on posterior surface; semi-erect; very slightly curved at middle; apex in posterior view acute, with small hooklike projection. Paramere: Cylindrical; moderately long, not reaching medial process; directed posteriad; basally slightly narrower; slightly curved ventrad; apical part very slightly enlarged. Phallus: Dorsal phallothecal sclerite shield-shaped; sclerotization reduced (yet not absent) on dorsal surface close to posterior margin of foramen; expansion of lateral margin at about mid-portion pronounced, covering ventral surface of endosoma; 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 moderately robust; basally fused; in lateral view slightly curved; bridge extremely short; extension of basal plate expanded laterally onto arm, covering more than 1/2 of arm.

Zelus panamensis
Female: (Fig. 150c, d) Similar to male, except for the following. Larger than male, total length 13.40-15.28 mm (mean 14.43 mm, Suppl. material 2). Dorsal coloration similar to that in male; lateral surface of pronotum, pleura in some specimens dark brown; abdomen always orangish or reddish. Basiflagellomere subequal in diameter to pedicel. Process on humeral angle spinous, long.

Diagnosis
Recognized by the orangish or reddish head and the dark brown remainder of the body.
The short, nearly straight medial process and the short paramere separate males of this species from all other species of the same species group. The yellowish or reddish ventral surface and the usually yellowish or reddish (blackish-brown in some specimens) lateral surface distinguishes females of this species from others in the same group.

Etymology
Named after the country Panama, where the holotype was collected.

Diagnosis
The dorsal coloration nearly uniformly dark brown, the head reddish-brown, the membrane with indistinct iridescence are characteristic of this species. Most similar to Z. erythrocephalus and Z. russulumus; males can be distinguished from both by the rather wide medial process and the uniquely shaped paramere (Fig. 14). Females of Z. erythrocephalus, Z. paracephalus and Z. russulumus are difficult to separate.

Distribution
South America (Fig. 155). Countries with specimen records: Brazil, Colombia, Ecuador and Peru.

Diagnosis
The slender, cylindrical paramere and the laterally compressed, blade-like medial process can separate this species from most other species of the genus. Different from Z. illotus and Z. impar by the straight medial process, contrasting with the recurved medial processes of the other two species. Difficult to distinguish from Z. nugax, but the paramere apex is generally rounded, the dorsal phallothecal sclerite usually with lateral indentation, the basal plate arms are fused and the basal plate arm extension is present and laterally expanded.

Distribution
South America and adjacent islands of the Caribbean (Fig. 158). Countries with records: Argentina, Bolivia, Brazil, Colombia, Ecuador, Guyana, Paraguay, Peru, Suriname and Trinidad and Tobago.

Taxon discussion
Some helpful, although not consistently reliable, characters used to distinguish females of Z. pedestris from those of Z. nugax and Z. illotus are given in the taxon discussion section of Z. nugax. Uncertainties in species limits between Z. pedestris and Z. nugax are also discussed in that species. The application of the name Z. pedestris is restricted to specimens from South America.
The coloration of the posterior pronotal lobe is the most obvious variable character in the males. This varies from a very light to a medium reddish-brown in any given geographic area. This pronotal variation is absent in the females, the dorsal surface varying from a relatively uniform light to medium brown in any geographic area.   Signoret, 1862, p. 584-585, orig. descr.;Walker, 1873, p. 126, cat. Zelus plagiatus: Stål, 1872Lethierry and Severin, 1896, p. 153, cat.;Wygodzinsky, 1949a, p. 50, checklist;Maldonado, 1990, p. 330

Diagnosis
The dorsum predominantly yellow with the posterior pronotal lobe partly black and two black spots on hemelytra is distinctive of this species. Zelus plagiatus (Signoret, 1852), specimen record map

Diagnosis
Distinguished by the greenish coloration; the veins of membrane darker than the cells; the smallish size; the rather slender body and very delicate legs; the head somewhat a b Figure 162.

Distribution
South America and adjacent islands of the Caribbean (Fig. 163). Argentina, Brazil, Ecuador, Guyana, Paraguay, Suriname, Trinidad and Tobago and Venezuela.

Diagnosis
The rather slender body form of Z. puertoricensis is characteristic of the Zelus puertoricensis species group (total length/width more than 8x). Both sexes of Z. puertoricensis have the dorsal and ventral surfaces of the postocular lobe nearly parallel through the anterior 2/3 of the lobe. This contrasts with the sloping configuration of the dorsal surface in Z. subimpressus.
Males can be recognized by the robust, posteriorly directed medial process, apex bent and the short, cylindrical paramere. This is smaller in Z. puertoricensis than in Z. subimpressus (Fig. 6). Additionally, the medial process of Z. puertoricensis appears to be longer and more delicate than that of Z. subimpressus. The dorsal phallothecal sclerite is much narrower than in Z. subimpressus.

Materials
Holotype: Male: (Fig. 167a, b) Medium-sized, total length 10.57-12.98 mm (mean 12.01 mm, Suppl. material 2); robust. COLORATION: Anteocular lobe yellowish-brown, some specimens with darker areas on either side of mid-dorsal line. Postocular lobe yellowish-brown, usually with variable brownish-black areas dorsally but always with mid-dorsal area and area anterior to ocellus yellowish-brown. Labium yellowish-brown, some specimens with brown labrum. Antennae yellowish-brown to light reddish-brown. Anterior pronotal lobe uniformly yellowish-brown or yellowish-brown with reddish-brown to dark brown setal tracts. Posterior pronotal lobe yellowish-brown to dark brown, margins light yellowish-brown, lateral surfaces yellowish-brown. Scutellum yellowishbrown to brown, apex lighter in color. Legs yellowish-brown, some specimens with a b Figure 168. apices of tibiae reddish-brown. Hemelytron yellowish-brown to dark brown, veins of clavus and corium usually lighter in color than surrounding area. Dorsum of abdomen reddish-brown to brown, connexival margins yellowish-brown, lateral and ventral surfaces usually yellowish-brown, some specimens with reddish-brown areas laterally. Pygophore yellowish-brown. VESTITURE: Moderately setose. Short recumbent and variable erect setae over surface. Anteocular lobe with short recumbent setae dorsally and laterally, short erect setae on tylus and ventral surface; postocular lobe with recumbent setae dorsally, longer erect setae on lateral surface and on dorsal and ventral surfaces of posterior half. Dorsal surface of anterior pronotal lobe with short recumbent setae confined to setal tracts, remainder of surface with longer recumbent and erect setae; posterior lobe with short recumbent and erect setae and some longer erect setae lateroventrally; scutellum with semi-erect setae over surface. Abdominal dorsal setae very short, erect, lateral and ventral surfaces with short recumbent and some short to moderately long erect setae. Exposed surface of pygophore with short recumbent and some short to moderate erect setae; erect setae over apical 3/5 of parameres. STRUCTURE: Head: Cylindrical, L/W = 2.61. Postocular lobe moderately long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye smallish; lateral margin only slightly wider than postocular lobe; dorsal and ventral margins removed from surfaces of head. Labium: I: II: III = 1: 1.6: 0.4. Basiflagellomere diameter larger than that of pedicel. Thorax: Medial longitudinal sulcus shallow near collar, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc slightly elevated above humeral angle; humeral angle armed, with dentate projection. Scutellum long; apex angulate, not projected. Legs: Robust. Hemelytron: Attaining apex of abdomen; quadrate cell small; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 168) Pygophore: Ovoid. Medial process cylindrical; slender; long; laterally somewhat compressed; erect; nearly straight; basally without protrusion; apex in posterior view modified, hooklike. Paramere: Cylindrical; moderately long, not reaching apex of medial process; directed toward medial process; basally narrower; curved dorsad; apical part enlarged. Phallus: Dorsal phallothecal sclerite shieldshaped; lateral margin recurved dorsad; apical portion of phallothecal sclerite gradually tapering, flat, lateral margin recurved; apex medially notched; posterior margin of foramen broadly concave. Struts not attached to base of dorsal phallothecal sclerite; apically missing. Basal plate arm robust; separate; converging; in lateral view basally strongly curved; bridge moderately long; extension of basal plate small, marginally expanded onto arm, lateral margins recurved.

Diagnosis
Can be recognized by the reddish corium; the remainder of the body surface greenish; the humeral angle with small subtuberculate projection. More robust than a very similar species, Z. cervicalis. Males can be recognized by the paramere apically greatly enlarged; the medial process apically curved ventrad, somewhat hooklike, more strongly than in Z. cervicalis, the only species that may be confused with; and the lateral margin of the dorsal phallothecal sclerite recurved.

Distribution
Western and Southwestern US, most of Mexico and Central America (Fig. 169).
Countries The current study revealed specimen records of Z. renardii from French Polynesia, representing the first report of this species from that region (Fig. 169, Suppl. material 1).

Taxon discussion
Zelus renardii is almost certainly sister species of Z. cervicalis. The two share two unique characters: the lateral margins of the dorsal phallothecal sclerite recurved and the medial process apically strongly hooked.

Diagnosis
The uniquely reddish coloration of the entire body makes this species easy to recognize. The medial process is highly reduced and rather indistinct, separating Z. rosulentus from other members of the Zelus tetracanthus species group (Fig. 2).

Etymology
The specific epithet indicates the reddish-pink coloration of this species.

Distribution
South America (Fig. 172). Only known from Ecuador.

Materials
Lectotype: Male: (Fig. 173a, b) Medium-sized, total length 13.24-16.83 mm (mean 14.93 mm, Suppl. material 2); robust. COLORATION: Orangish to reddish, with black areas. Head orangish or reddish-brown; dark areas on postocular lobe in some specimens. Anterior pronotal lobe orangish or reddish-brown; posterior margin dark, sometimes occupying more than 1/2 of surface. Posterior pronotal lobe, corium and clavus orangish-brown; sometimes with dark areas on each. Scutellum dark brown; margins yellowish-brown. Membrane dark brown. Legs not distinctly banded; femoral apical portion usually reddish, occasionally dark, single small black band subapically, sometimes very faint black marking medially. Lateral and ventral surfaces orangish to reddish-brown; variable black areas on pleura and abdomen; dark stripe along posterior margin of each segment, width variable; pygophore usually reddish. VESTITURE: Densely setose. Dorsal surface of head with fine to stiff erect setae and some recumbent setae, lateral and ventral surfaces with erect and recumbent setae. Anterior pronotal lobe with recumbent and erect setae, confined to setal tracts, erect setae predominant; posterior pronotal lobe with erect setae and some recumbent setae. Abdomen with short, recumbent setae, interspersed with erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.21. Postocular lobe relatively short; in dorsal view distinctly narrowing through anterior 1/2, posterior 1/2 constant, tube-like. Eye smallish; lateral margin only slightly wider than postocular lobe; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head in lateral view. Labium: I: II: III = 1: 1.4: 0.4. Basiflagellomere diameter subequal to that of pedicel. Thorax: Anterolateral angle bearing small protuberance; medial longitudinal sulcus distinct throughout. Posterior pronotal lobe with rugulose surface; disc distinctly elevated above humeral angle; humeral angle armed, with short tuberculate processes. Scutellum moderately long; apex angulate. Legs: Moderately robust. Hemelytron: Slightly surpassing apex of abdomen, not more than length of abdominal segment seven; quadrate cell large and broad; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 174) Pygophore: Rounded; slightly expanded laterally near base of paramere in dorsal view; posteriorly expanded sac-like sclerite between parameres and medial process. Medial process cylindrical; slender; moderately long; posteriorly directed; nearly straight; apex in posterior view rounded, with small sharp lateral projections. Paramere: Cylindrical; moderately long, not reaching apex of medial process; directed posteriad; nearly straight; apical part not enlarged. Phallus: Dorsal phallothecal sclerite somewhat squarish; lateral expansion arising close to base; apical portion of phallothecal sclerite not distinctly tapered, flat, lateral margin narrowly angulate; apex rounded; posterior margin of foramen broadly inversely V-shaped. Struts attached to dorsal phallothecal sclerite; apically fused; basally separate throughout. Basal plate arm moderately robust; basally fused; in lateral view very slightly curved; bridge extremely short; extension of basal plate small, marginally expanded onto arm.
Female: (Fig. 173c, d) Similar to male, except for the following. Larger than male, total length 16.46-20.01 mm (mean 17.84 mm, Suppl. material 2). Usually lighter than male and larger areas of red and yellow.

Diagnosis
The combination of relatively large size, stout body, the reddish head and parts of body can separate both sexes of this species from other species of Zelus. Among males of the Zelus armillatus species group, Z. ruficeps has some of the most delicate medial process. Females may be confused with Z. grassans, but are much larger and the humeral angle is equipped with dentate process, contrasting to the nearly rounded humeral angle of Z. grassans.

Distribution
Southern Mexico to Northern South America (Fig. 175) Female: (Fig. 176c, d) Similar to male, except for the following. Larger than male, total length 17.52-19.39 mm (mean 18.23 mm, Suppl. material 2). Spinous process on humeral angle long.

Diagnosis
The dorsal coloration nearly uniformly dark brown, the head reddish-brown, and the membrane with blue or green iridescence. Most similar to Z. erythrocephalus and Z. paracephalus; can be distinguished by the medial process wider than that in Z. erythrocephalus and narrower than Z. paracephalus. Females of Z. erythrocephalus, Z. paracephalus and Z. russulumus are difficult to separate.

Etymology
The species epithet refers to the reddish-brown area on the membrane.

Distribution
South America (Fig. 178)  Zelus spatulosus Zhang & Hart, sp. n., specimen record map dark reddish-brown, remainder variably banded, light to medium reddish-brown, basal area, apex and two bands on meso and metafemora dark reddish-brown, remainder light reddish-brown, tibiae medium to dark reddish-brown with light reddish-brown band about half distance from base. Abdomen reddish-brown, connexival margins lighter, anterior margins of segments three to seven dark reddish-brown. VESTITURE: Moderately setose. Anteocular lobe with short, recumbent to erect setae. Postocular lobe with short, recumbent setae, some long, erect setae on lateral and ventral surfaces. Anterior pronotal lobe with moderately long, recumbent to semi-erect setae, confined to setal tracts dorsally, long erect setae laterally. Posterior pronotal lobe with short, recumbent setae, long erect setae laterally. Scutellum with recumbent to semierect setae. Dense recumbent to semi-erect setae over clavus and corium. Abdomen with short, erect setae on dorsum, dense, short, recumbent and moderate to long erect setae over lateral and ventral surface, with exception of bare dark areas. Exposed surface of pygophore with short semi-erect and long erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.12. Postocular lobe moderately long; in dorsal view anteriorly gradually narrowing, posterior portion constant, slightly narrower. Eye moderately sized; lateral margin only slightly wider than postocular lobe; dorsal margin attaining postocular transverse groove, ventral margin removed from ventral surface of head in lateral view. Labium: I: II: III = 1: 1.7 : 0.6. Thorax: Anterolateral angle of collar rounded, collar not well defined medially; medial longitudinal sulcus evident only on posterior 1/2, deepening anterior to transverse sulcus of pronotum. Posterior pronotal lobe with finely rugulose surface; disc slightly elevated above humeral angle; humeral angle armed, with dentate process. Scutellum moderately long; apex angulate, not projected. Legs: Rather slender. Metafemur slightly more slender than pro-and mesofemora. Hemelytron: Surpassing apex of abdomen by more than length of abdominal segment seven; quadrate cell small; Cu and M of cubital cell converging towards R. GENITALIA: (Fig. 180) Pygophore: Elongate ovoid; not expanded laterally in dorsal view. Medial process laterally compressed; slender; long; semi-erect; straight; apex in posterior view blunt, without modification. Paramere: moderately long, reaching medial process; apical part greatly expanded. Phallus: Dorsal phallothecal sclerite elongated; apical portion tapered, slightly convex, lateral margins recurved upward; apex with rounded protuberances, posterior margin of foramen inversely V-shaped.
Struts not readily evident. Basal plate arm moderately robust; basally fused; in lateral view nearly straight, very slightly curved; bridge short; extension of basal plate reduced.

Diagnosis
Zelus spatulosus has a rather distinctly enlarged apical part of the paramere (Fig.  180a), the extent not compared by any other species of Zelus. Also, the slender form of the medial process is distinctive among the Zelus luridus species group.

Etymology
The specific epithet is from spatula, referring to the rather broad apical part of the paramere.

Distribution
Central America (Fig. 181). Known only from the type locality in Belize.

Taxon discussion
The primary basis for placing Z. spatulosus in the Zelus luridus group is the expanded paramere (Fig. 3). However, this species shows several characters which depart significantly from the remaining species as described in the following. The medial process is slender and laterally compressed, the struts are not evidently visible, the dorsal phallothecal sclerite is distinctly shaped, with a narrow basal portion and expanded apical part, and the basal plate arms are fused. These characters are so distinct that they are possibly either autapomorphic or homoplasious. As no characters are found to unite Z. spatulosus with species of other species group, its placement in the Zelus luridus species group appears to be the best decision to take.

Diagnosis
This species can be recognized by the pronotum bicolorous, anterior lobe yellowish and posterior lobe dark brown; the humeral angle with long spinous process. Similar to females of Z. truxali, but the legs are not banded and the anterior pronotal lobe appears to be more humped than that in Z. truxali.

Distribution
Central America and South America (Fig. 183). Only know from two countries: Guyana and Panama.

Diagnosis
The rather slender body form of Z. subimpressus is characteristic of the Zelus puertoricensis species group (total length/width more than 8x). Both sexes are readily distinguished from Z. puertoricensis by having a sloping dorsal surface on the postocular lobe. Males can be recognized by the robust, posteriorly directed medial process, apex bent and the short, cylindrical paramere. This is larger in Z. subimpressus than in Z. puertoricensis (Fig. 6).

Distribution
The Caribbean, the islands of Cuba and Hispaniola (Fig. 186). Countries with records: Cuba, Dominican Republic and Haiti. Male: (Fig. 187a, b Female: (Fig. 187c, d) Similar to male, except for the following. Larger than male, total length 20.59-21.91 mm (mean 21.41 mm, Suppl. material 2).

Diagnosis
Recognized by the dorsal coloration nearly uniformly brown, somewhat reddish and the posterior pronotal lobe medially depressed. Among males of the Zelus armillatus species group occurring in the same geographic range, the medial process and paramere of Z. sulcicollis are longer than that in Z. litigiosus, Z. ruficeps and Z. janus.

Distribution
Southern Mexico and Northern Central America (Fig. 189) fig. 27, note and fig.; Fracker, 1913, p. 240, key and list (subgenus  Male: (Fig. 190a) Medium-sized, total length 11.34-13.73 mm (mean 12.36 mm, Suppl. material 2); slender. COLORATION: Greyish-brown to black. Anteocular lobe dark reddish-brown to brownish-black, variable yellowish-brown areas usually present dorsolaterally anterior to compound eyes, dorsally and medially at antennal bases, and on lateral and ventral surfaces. Postocular lobe dark reddish-brown with lighter areas between ocelli and compound eyes, along mid-dorsal line, longitudinally on lateral and ventral surface. Labium reddish-brown to brownish-black with anterior surface of segment I lighter in color, yellowish-brown to reddish-brown. Antennal segments I and II reddish-brown, outer dorsolateral surface of I darker, especially near base, base and apex of II darker, segments III and IV yellowish-brown to reddish-brown. Anterior pronotal lobe reddish-brown to brownish-black, occasionally with variable lighter areas, especially near margins, lateral surface reddish-brown to brownish-black, usually lighter ventrally. Posterior pronotal lobe reddish-brown to brownish-black with dark yellowishbrown posterior margin and/or darker reddish-brown to brownish-black apices on lateral and dorsal processes. Scutellum variably yellowish-brown to brownish-black, usually with lighter apex. Legs yellowish-brown to reddish-brown, usually with variable darker reddish-brown spots or bands, especially near dorsal surfaces of femoral apices. Hemelytron brown to dark brown, veins in area anterior to basal and discal cells lighter. Abdomen variable, dark yellowish-brown to reddish-brown, darker areas usually at posterior of segments of connexivum and mid-laterally at anterior of segments 3-7.
Abdomen yellowish-brown with variable reddish-brown areas on lateral and posterior margins and posterior surface. VESTITURE: Moderately setose. Short recumbent and short to long erect setae, many specimens covered with white waxlike exudation. Anteocular lobe with short recumbent setae on entire surface, longer erect setae on tylus and ventral surface; postocular lobe with short recumbent setae over entire surface, moderate to long erect setae on posterodorsal and ventral surfaces. Entire surface of anterior pronotal lobe with short recumbent and erect setae, confined to setal  Female: (Fig. 190b, c) Similar to male, except for the following. Larger than male, total length 13.22-15.63 mm (mean 14.43 mm, Suppl. material 2). Often darker than male. Tuberculate processes of posterior pronotal lobe usually more pronounced, apex often produced. Mid femur slightly swollen on central 1/4, pro-and mesofemoral diameters subequal, about 1.4-1.5x diameter of hind femur. Hemelytron attaining apex of abdomen.

Diagnosis
Recognized by the disc of the posterior pronotal lobe with large conspicuous tubercles and the greyish-black coloration. Males can also be recognized by the medial process broadly triangular; the paramere not exceeding medial process; and the dorsal phallothecal sclerite apically with deep emargination.

Distribution
North America, Central America and parts of South America (Fig. 192). Countries with records: Brazil, Costa Rica, Curaçao, Honduras, Mexico, Panama, Paraguay, USA and Venezuela.

Taxon discussion
Besides Z. tetracanthus, two other species, Z. minutus and Z. lewisi also exhibit tuberculate processes on posterior pronotal lobe, but these species can be easily separated on the basis of coloration and body shape. Zelus tetracanthus is a rather widely distributed species, found nearly throughout North America, ranging from southern Canada to parts of Central America. A few records are from Brazil and Paraguay. As the collecting events of these specimens are unrelated, it is highly unlikely these specimens are mislabelled (unless they were each independently mislabelled, which is not likely Male: (Fig. 193a, b) Medium-sized, total length 11.59-12.25 mm (mean 11.91 mm, Suppl. material 2); slender. COLORATION: Two major patterns recognized, one of predominantly brown to orangish-brown, the other of dark brown to reddish-brown. In former, posterior lobe lighter colored than anterior lobe, orangish-brown; meso-and meta-femora with two minor bands. In latter, posterior pronotal lobe same color as anterior lobe, or only slightly lighter, somewhat reddish; legs not banded. In both patterns, medial longitudinal lighter colored stripe on postocular lobe. VESTITURE: Sparsely setose. Short, recumbent setae on entire surface of head; 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, recumbent setae on lateral surface and pleura, intermixed with short, 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. STRUCTURE: Head: Cylindrical, L/W = 2.30. 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 attaining postocular transverse groove, ventral margin removed from ventral surface of head in lateral view. Labium: I: II: III = 1: 1.9: 0.4. Basiflagellomere diameter larger than that of pedicel. Thorax: Anterolateral angle bearing small, somewhat acute 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 moderately long; 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. 194) Pygophore: Elongate ovoid; lightly sclerotized expansion below paramere; not expanded laterally in dorsal view. Medial process cylindrical; slender; moderately long, slightly longer than paramere; laterally slightly compressed towards apex; semi-erect; basally slightly curved, apex recurved; apex in posterior view acute, with small hooklike projection. Paramere: Cylindrical; moderately long, not reaching medial process; directed posteriad; slightly curved ventrad; apical part not enlarged. Phallus: Dorsal phallothecal sclerite shield-shaped; 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 truncate; posterior margin of foramen broadly concave. Struts apical portion missing; basally separate. Basal plate arm slender; separate; in lateral view slightly curved; bridge short; extension of basal plate expanded laterally onto arm, covering more than 1/2 of arm, curved.

Diagnosis
The uniquely slender, recurved medial process can distinguish this species among the members of the Zelus panamensis species group (Fig. 12). In females the anterior pronotal lobe yellowish and posterior lobe brown; the lateral and ventral surface of the body yellowish is unique among females of all species. Females are very similar to Z. sphegeus, but are separate from that species by the banded legs and the anterior pronotal lobe nearly flat, not distinctly elevated.

Etymology
Named after F. S. Truxal, the collector of the type specimen.

Diagnosis
Recognized by the body surface greenish-brown; the area of hemelytron adjacent to the quadrate cell dark brown, inversely U-shaped in appearance; the head short and stout (L/W=<2.1) ; the ocellus situated on conspicuous elevation (the same set of characters are also present in Z. umbraculus). Characters separating Z. umbraculoides and Z. umbraculus are discussed in the diagnosis of the latter.

Etymology
The specific epithet indicates its close resemblance to Z. umbraculus, another new species described in the current study.

Taxon discussion
Hart (1972) did not discover this species and included some of the specimens under Z. umbraculus. Upon a close examination of these and several new specimens collected after 1972, we found several major differences that can reliably separate Z. umbraculoides and Z. umbraculus (see diagnosis of the latter). Although specimen records are still sparse, the two species do not overlap in distribution. Zelus umbraculoides is known from southern Peru and south of Peru in Bolivia, Paraguay and southern Brazil, while Z. umbraculus from Ecuador and northern Peru.

Diagnosis
Recognized by the body surface greenish-brown; the area of hemelytron adjacent to the quadrate cell dark brown, inversely V-shaped in appearance; the head short and stout (L/W=<2.1) ; the ocellus situated on conspicuous elevation (the same set of characters are also present in Z. umbraculus). Zelus umbraculus can be separated from Z. umbraculoides by several characters listed below (Figs 8,9). Zelus umbraculus: (1) Disc of the posterior pronotal lobe slightly bulging, nearly flat; (2) paramere moderately robust, apical 1/2 greater; (3) apex of the dorsal phallothecal sclerite rounded; and (4) basal plate arms apically separate. Zelus umbraculoides: (1) Disc of the posterior pronotal lobe clearly bulging; (2) paramere relatively slender; (3) apex of the dorsal phallothecal sclerite truncate, and (4) basal plate arms apically touching. The disc of the pronotal lobe is best observed from a posterior angle with the head facing down.

Etymology
The specific epithet is from Latin "umbra", referring to the shadow in the anterior part of the membrane.

Distribution
South America (Fig. 201). Countries with records: Ecuador and Peru.

Diagnosis
Can be easily identified by the unique coloration pattern, the posterior pronotal lobe medially black and laterally orange. Distinguished among members of the Zelus vagans species group by the smaller size; the postocular lobe covered with recumbent setae. The paramere is similar to that in Z. championi in showing ventrally directed curvature, but is shorter than in Z. championi and reaching to only about mid-point of medial process.
Female: Similar to male, except for the following. Larger than male, total length 13.71-14.03 mm (mean 13.87 mm, Suppl. material 2). Entire body nearly uniformly pale brown or dark brown.

Diagnosis
Recognized by the following combination of characters: The posterior margin of posterior pronotal lobe with expansion laterad to scutellum; the rather long, spinous process on humeral angle; the smallish body size; the paramere short; the apical part of medial process compressed laterally, anterior side ridge-like; and the apex of medial process re-expanded, not acute.
Eye prominent; lateral margin 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 rounded, without 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 projection. Scutellum moderately long; 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. 209) Pygophore: Ovoid; expanded laterally near base of paramere in dorsal view. Medial process triangular; long; anteroposteriorly compressed; erect; basally without protrusion; apex in posterior view acute; posterior surface with pair of sharp ventrally directed processes. Paramere: Bulbous, moderately long, slightly exceeding medial process; curved toward medial process; basally constricted; slightly curved ventrad; apical part enlarged. Phallus: Dorsal phallothecal sclerite somewhat squarish; laterally with small blade-like heavily sclerotized process; apical portion of phallothecal sclerite not distinctly tapered, laterally indistinctly angulate, angulation anteriorly connected with membranous, dorsad expansion; apex rounded, medially emarginate; 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 moderately robust; separate; subparallel; in lateral view severely curved, nearly semi-circular; bridge long; extension of basal plate expanded onto arm.
Female: Similar to male, except for the following. Larger than male, total length 14.43-15.84 mm (mean 15.01 mm, Table 4.2). Dorsum dark brown, a large number of specimens with anterior 1/2 of posterior pronotal lobe yellow and some also with entire anterior pronotal lobe yellow; lateral surface and abdomen yellowish. Hemelytron slightly surpassing apex of abdomen.

Diagnosis
Recognized by the following combination of characters: the posterior pronotal lobe usually lighter than the anterior pronotal lobe, orangish or reddish-brown; the paramere bulbous, basally constricted, curved towards medial process; the medial process triangular, apex with pair of processes; the dorsal phallothecal sclerite with submedial ridge-like dorsad projection continuous from basal arm; and the basal plate arms subparallel and strongly curved. In females the posterior pronotal lobe is often bicolorous, anterior portion yellowish and posterior portion brown.
Female: Similar to male, except for the following. Larger than male, total length 16.48-18.29 mm (mean 17.47 mm, Suppl. material 2). Some specimens almost entirely dark brown, only with medial part of or entire posterior pronotal lobe, mesopleuron and mesosternum brownish orange or orange.

Diagnosis
The generally wasp-like coloration pattern can separate this species from most other species of the Zelus, but not from a few species that may also display a wasp-like appearance. Males can be recognized by the rather slender and short medial process, it being much shorter than that in Zelus errans, the only species that may cause confusion.  Male: (Fig. 214) Medium-sized, total length 12.59 mm (n=1, Suppl. material 2); slender. COLORATION: Entire surface brown, dorsum of head, posterior pronotal lobe, corium, clavus, apices of femora, parts of tibiae somewhat reddish. VESTITURE: Sparsely setose. Short, recumbent setae on dorsal surface of head, long recumbent setae on ventral surface; 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, moderately long recumbent setae on lateral surface and pleura, intermixed with semi-erect 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 short, recumbent setae. Abdomen with moderately dense, short recumbent setae, intermixed with sparse, short to long, erect setae. STRUCTURE: Head: Cylindrical, L/W = 2.43. 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 in lateral view. Labium: I: II: III = 1: 2.1: 0.4. Basiflagellomere diameter slightly larger than that of pedicel. Thorax: Anterolateral angle with inconspicuous subtuberculate projection; medial longitudinal sulcus evident throughout, deepening posteriorly. Posterior pronotal lobe with rugulose surface; disc about same level as humeral angle; humeral angle armed, with spinous processes. Scutellum moderately long; apex angulate. Legs: Very slender, femoral diameters subequal. Hemelytron: Surpassing apex of abdomen by about length of abdominal segment seven; quadrate cell small and slender; Cu and M of cubital cell subparallel. GENITALIA: (Fig. 215) Pygophore: Ovoid; not expanded laterally in dorsal view. Medial process cylindrical; slender; moderately long, much longer than paramere; minute spicules on posterior surface; semi-erect; basally slightly curved; apex in posterior view acute, with small hooklike projection. Paramere: Cylindrical; moderately long, slightly exceeding medial Zelus xouthos Zhang & Hart, sp. n., specimen record map process; directed posteriad; apical part appearing somewhat truncate dorsally. Phallus: Dorsal phallothecal sclerite elongated; medial, longitudinal sulcus on dorsal surface; apical portion of phallothecal sclerite gradually tapering, distinctly keeled medially, laterally flat, not forming angle; apex acute; posterior margin of foramen broadly concave, accuminate in middle. Basal plate arm slender; separate, briefly touching; converging; in lateral view slightly curved; bridge short; extension of basal plate expanded laterally onto arm, covering more than 1/2 of arm.

Description
Female: Similar to male, except for the following. Larger than male, total length 14.69 mm (n=1, Suppl. material 2).

Diagnosis
Recognized by the following combination of characters: the slender body and delicate legs, the dorsal coloration somewhat reddish; the humeral angle elevated nearly to the level of and nearly continuous with pronotal disc; the paramere uniquely shaped, long, exceeding medial process, apex somewhat obliquely truncate; and the medial process apically not compressed.

Etymology
From Greek xutho, meaning yellowish-brown, referring to the yellowish coloration.

Distribution
Central America (Fig. 216). Known only from Guatemala.

Taxon discussion
This species appears to be the most divergent of the Zelus panamensis species group. Although three other species within the same geographical range of this species, Z. janus, Z. exsanguis, Z. ambulans, also have an elevated humeral angle, their much greater sizes and features of the male genitalia should eliminate any confusion in recognizing Z. xouthos.

Diagnosis
The extremely slender body form separates this species from most other species of Zelus. Can be distinguished from Z. puertoricensis by the lack of conspicuous lateral process on the humeral angle and from Z. subimpressus by the parallel dorsal and ventral surfaces on the anterior 2/3 of the postocular lobe. The dorsal surface in Z. subimpressus is sloping.

Distribution
The Caribbean. Known only from Cuba (Fig. 218).

Taxon discussion
Based on the description and figure of Z. bruneri, it is possible that Z. bruneri is the male of Z. zayasi. Because the specimens of Z. bruneri were not physically examined, we restrain from making this association and a formal synonymy between these two species. Zelus zayasi Bruner and Barber, 1937, specimen record map -Medial process cylindrical or broad at base; not laterally compressed; apex curved, lip-like rather than hook-like (Fig. 4). Paramere apically not expanded, strongly curved dorsally.

Zelus mimus species group 7
Apex of medial process with distinct, acute, hook-like process or ridge-like elevation (Figs 12, 14). 8 -Apex of medial process with indistinct process or weak curvature, not forming sharp projection (Figs 6, 8, 10). 9 8 Medial process dorsally directed and apical 1/3 not curving posteriorly; usually anteroposteriorly compressed; base broad in some species; ridge-like elevation on posterior surface, usually extending ventrally and elevated, removed from apex and appearing as pair of projections in some species (Fig. 14). Paramere usually bulbous in apical 1/2 or 2/3, curved medially.
Zelus longipes species group -Medial process stout, relatively short (Figs 6, 8). Paramere removed from or reaching apex of medial process. Humeral angle armed with projection.

Zelus prolixus
Key to males of the Zelus luridus species group 1 Humeral angle raised to level of, and usually continuous with, disc of posterior pronotal lobe. Zelus ambulans (Fig. 22a, b) -Medial longitudinal sulcus of anterior pronotal lobe same color as surrounding area at posterior margin, any dark areas present very small. Femora apically reddish-brown, not forming distinct bands. Paramere in fresh or relaxed specimens achieving or surpassing apex of medial process (Fig. 72a, b) (Mexico and C.A.).

Key to males of the Zelus erythrocephalus species group
1 Head reddish-brown. Dorsal surface of pronotum and corium brownish-black. Membrane of hemelytron pale brown or blue or green iridescent. Legs brownish-black, without banding or with inconspicuous bands.

-
Head brown, yellow, or black, sometimes with stripes. Dorsal surface uniformly brown or with light-colored areas, mainly on pronotum or corium. Membrane brown, not iridescent. Legs with or without banding.
7 -Posterior surface of medial process with hook-like process, across width of medial process; if extending ventrally, as pair of dentate processes.

8
-Dorsal surface sloping downward from ocelli, height at middle of lobe less than 0.9x that through ocelli. Medial process moderately long (Fig. 185a).

Key to females of Zelus
This key is not organized by species group, as females do not readily display characters placing them to species groups. It is especially challenging or sometimes impossible to distinguish females of species in several species groups. Large series of co-occurring males and females will be helpful in these cases. Coloration is heavily used to key out females, and readers are reminded that some species exhibit a wide range of color variations and the exemplar habitus images provided in this work are not inclusive of all variants.

-
Anterior pronotal lobe dorsally with fine setae or nearly bare, any spine-like setae sparse; setal tracts not necessarily well-defined. Humeral angle usually armed, some rounded. 8 2 Humeral angle armed with spinous process. Body elongate.

Zelus annulosus
-Humeral angle rounded, without lateral process. 3 3 Dorsal surface of head, pronotum, scutellum and hemelytron nearly unicolorous, dark brown to black. Cells of membrane occasionally lighter in color than veins.

-
Dorsal surface with reddish-brown areas on pronotum and/or scutellum, often with yellowish-brown to reddishbrown areas on clavus and corium and/or basal 1/2 of membrane.

Zelus aithaleos
5 Profemoral length less than 20x profemoral diameter. Quadrate cell short and broad; Pcu of quadrate cell less than 1/2 length of Cu.
Zelus means (Fig. 136) 8 Compound eyes extending below ventral surface of head. Zelus grandoculus -Compound eyes not extending below ventral surface of head. 9 9 Posterior margin of pronotal disc with two tubercles. Humeral angle with spinous process. 10 -Posterior margin of pronotal disc unarmed. Humeral angle armed or unarmed. 12 10 Tubercles of pronotal disc pronounced. More than 13 mm in length. 11 -Tubercles very small, minute in some specimens. Thirteen mm or less in length. (Southern C.A. and Northern S.A.)

Zelus varius
-Posterior pronotal lobe, with posterior margin straight or sloping gradually (not appreciably more than forty-five degrees) downward on either side of scutellum.

13
Head surface reddish-brown, any darker cranial markings with indistinct outlines; remainder of dorsal surface of body primarily dark brown to brownish-black. 13.0-18.5 mm in length.
Zelus longipes (Fig. 127e, f) -At least one of the characters not as described above. 21

21
Length at least 5x width. Profemoral diameter equal to or greater than that of mesofemur. Mesofemoral diameter less than 1.2x that of metafemur. Tuberculate to small spinous lateral processes on humeral angle. Interocular distance less than 1.15x interocellar distance.

22
-At least one of the characters not as described above. 23

22
Postocular lobe with dorsal and ventral surfaces nearly parallel through anterior 2/3; height at middle of lobe and through ocelli subequal (Greater Antilles).
Body length at least 7.2x width (Cuba). Zelus bruneri -Anteocular lobe less than 1.25x length of postocular lobe. Body length less than 7.2x width (S.A.) Zelus prolixus (Fig. 161c, d) 26 Humeral angle rounded. Less than 14 mm in length. 27 -Humeral angle with at least small tubercles or spines; if tubercles and spines not readily evident, at least 14 mm in length.
Zelus mattogrossensis (Fig.  133c, d) 28 Length usually less than 4.0x width; if length greater than 4.0x and less than 4.5x width, profemoral diameter greater than that of mesofemur.

29
-Length usually at least 4.0x width; if length less than 4.0x width, mesofemoral diameter greatest. 34 29 Length usually at least 4.0x width; if length less than 4.0x width, mesofemur enlarged, diameter greatest.

30
Length of anteocular lobe at least 1.1x that of postocular lobe. Rostral segment II less than 1.35x length of segment I. Dorsal coloration uniform, pale brown, somewhat greenish. Legs not banded (Mexico, C.A. and

S.A.).
Zelus amblycephalus (Fig.  19c, d) -Anteocular lobe less than 1.1x length of postocular lobe. Rostral segment II at least 1.35x length of segment I. Dorsal surface is either variously patterned, with contrasting pale and dark regions or uniformly colored, dark brown to black. Legs usually with bands or annulations, or uniformly dark brown to black.

31
Humeral angle raised to level of, and nearly continuous with, disc (Mexico and C.A.). Zelus janus (Fig. 103c, d) -Humeral angle below level of disc; disc clearly elevated. 32 32 Setal tracts of anterior pronotal lobe and dorsum of posterior lobe with conspicuous light-colored shining recumbent setae, especially dorsolateral area of lobe; some erect setae also present; setal tracts usually of contrasting color. Length:width ratio approximately 4:1 (Mexico and C.A.).
Zelus ruficeps (Fig. 173c, d) 34 Dorsal surface dark brown to brownish-black. Golden, shining, short recumbent setae dorsally, especially on head and pronotum. Humeral angle raised to level of, and nearly continuous with, disc (central and southern S.A.).
Zelus auralanus (Fig. 36e, f) -Dorsal surface usually not dark brown, but if so, then setae not as above. If humeral angle raised, coloration not as dark above.

35
Humeral angle raised to level of, and nearly continuous with, disc. General dorsal coloration yellowish-brown. 36 -Humeral angle below level of disc. If dorsal coloration yellowish-brown as above, then disc clearly differentiated. 38 36 Profemoral length at least 22x profemoral width. Zelus exsanguis (Fig. 71c, d) 38 Length of rostral segment II less than 1.8x that of segment I. Profemoral length less than 22x profemoral width. Pronotum covered with erect setae dorsally, some setae subequal in length to diameter of shaft of antennal segment I. Humeral angle with very small tuberculate or subtuberculate lateral process (Mexico, C.A., and northern S.A.).
Zelus grassans (Fig. 91d) -At least one of the characters not as described above. 39

39
Length of rostral segment II less than 2x that of segment I. Profemoral and mesofemoral lengths less than 20x and 11x that of respective widths. Length of posterior pronotal lobe greater 2.4x that of anterior lobe.
Zelus laticornis (Fig. 115e, f) -At least one of the characters not as described above. 40 Profemoral length less than 17x that of profemoral width.
Length of rostral segment II less than 2.1x that of segment I.

41
-Profemoral length at least 17x of width. Rostral segment II usually greater than 2.1x that of segment I. 43

41
Humeral angle with short, inconspicuous subtuberculate to nearly dentate lateral processes, usually same color as surrounding area. Posterior pronotal lobe nearly smooth (Guatemala, Mexico and western and southwestern U.S.).
Zelus mimus (Fig. 138c, d, e, f) -At least one of the characters not as described above. 44

44
Posterior pronotal lobe with single broad dark brown transverse band posteriorly or pair of bands anteriorly and posteriorly, remaining dorsal surface of pronotum yellowish, with or without band.
Zelus plagiatus (Fig. 159) 48 Pubescence of lateral surface of posterior pronotal lobe consisting almost entirely of erect setae, many longer than diameter of shaft of antennal segment II (S.A.).
Zelus versicolor (Fig. 208c, d) -Majority of setae on lateral surface of posterior pronotal lobe semi-erect to recumbent, no erect setae as long as diameter of shaft of antennal segment II (southern C.A. and northern S.A.).