Biodiversity Data Journal : Taxonomic paper
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Corresponding author: Guanyang Zhang (gyz151@gmail.com)
Academic editor: Pavel Stoev
Received: 17 Feb 2016 | Accepted: 16 Jun 2016 | Published: 08 Jul 2016
© 2016 Guanyang Zhang, Elwood R Hart, Christiane Weirauch.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Zhang G, Hart E, Weirauch C (2016) A taxonomic monograph of the assassin bug genus ZelusFabricius (Hemiptera: Reduviidae): 71 species based on 10,000 specimens. Biodiversity Data Journal 4: e8150. doi: 10.3897/BDJ.4.e8150
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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.
Harpactorinae, Heteroptera, natural enemy, Nearctic, Neotropical, new species, Reduviidae, synonymy, systematics, taxonomic revision, Zelus
Zelus Fabricius, 1803 is one of the largest reduviid genera (
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
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 (
A series of works by Stål greatly changed the generic limits of Zelus (
However, the subgeneric groups were raised to the generic rank by
Uhler listed a species of Diplodus from one of the U. S. Geological Survey expeditions (
The generic and subgeneric definitions of Stål were also used by
In his study of the Heteroptera of eastern North America,
In his Ph.D. dissertation
Recent taxonomic activities on Zelus spp. are scant.
During the course of this study, 10,626 specimens were examined and databased. Among those, 4,833 are males, 5,626 are females and the remainders are immatures or with sex undetermined (usually because of missing abdomen). Specimen loans were kindly provided by museums or collections listed in
List of museums/collections
Acronym | Museum/Collection | Manager/Curator |
AMNH | American Museum of Natural History, New York, USA | Ruth Salas/Randall T. Schuh |
BMNH | Natural History Museum, London, UK | Mick Webb |
BPBM | Bernice Pauahi Bishop Museum, Honolulu, USA | |
CAS | California Academy of Sciences, San Francisco, USA | Norman D. Penny |
CUIC | Cornell University Insect Collection, Ithaca, USA | Rick Hoebeke/James Liebherr |
FMNH | Field Museum of Natural History, Chicago, USA | James Boone |
FSCA | Florida State Collection of Arthropods, Florida Department of Agriculture, Gainesville, USA | Susan Halbert |
ICN | Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá, Colombia | Carlos Sarmiento |
IEXA | Instituto de Ecologia, Xalapa, México | Luis Cervantes |
INBIO | Instituto Nacional de Biodiversidad, San José, Costa Rica | James Lewis |
KU | Snow Entomological Museum, University of Kansas, Lawrence, USA | Zachary H. Falin |
LACM | Natural History Museum of Los Angeles County, California, USA | Weiping Xie/Brian Brown |
MEFLG | Museo Entomológico Francisco Luis Gallego, Medellín, Colombia | John Albeiro Quiroz |
NHMW | Natural History Museum of Vienna, Vienna, Austria | Herbert Zettel |
NHRS | Sweden Museum of Natural History, Stockholm, Sweden | Gunvi Lindberg |
RMNH | Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historie), Leiden, Netherlands | Yvonne van Nierop |
TAMU | Texas A&M University Insect Collection, College Station, USA | Edward G. Riley |
UCB | Essig Museum of Entomology, University of California, Berkeley, USA | Cheryl Barr |
UCD | Bohart Museum of Entomology, University of California, Davis, USA | Steve Heydon |
UCR | Entomology Research Museum, University of California, Riverside, USA | Douglas Yanega |
UMSP | University of Minnesota Insect Collection, St. Paul, USA | Philip J. Clausen |
UNAB | Museo Entomológico, Facultad de Agronomía, Universidad Nacional de Colombia, Bogotá, Colombia | Francisco Serna |
UNAM | Universidad Autonoma de México, Instituto de Biología, México | Harry Brailovsky |
USNM | United States National Museum of Natural History, Washington DC, USA | Michele Touchet/Thomas Henry |
ZMAN | Zoological Museum Amsterdam, Amsterdam, Netherlands | Willem Hogenes |
ZMUC | Copenhagen University Zoological Museum, Copenhagen, Denmark | Henrick Enghoff |
Information of type specimens of described, valid species, when available, were reported in the 'Materials' section of each species, as holotype, lectotype or neotype.
Distributions were based on specimen records captured in the current study. We have gathered the largest samples ever known of all species, which represent the best available knowledge of the distributions of the species of Zelus. Maps were created using the Simple Mapper tool through the PBI website (http://research.amnh.org/pbi/maps) based on the geo-referenced locality data. Because accuracy and error of geo-referencing are highly variable, distribution records shown on the maps are at best indicative. Besides, ambiguous localities and other localities provided only at the country or state level were not geo-referenced and thus not reflected on the distribution maps. It is, therefore, advisable to look up actual specimen data for locality information (
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
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
Length measurements
Width measurements
Description. Observations were recorded with the software DEscriptive Language for TAxonomy (DELTA) (
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
Male genitalic structure terms (Zelus errans Fabricius, 1803 is shown in the illustrations)
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.
This publication is registered in ZooBank. In accordance with the 2012 Amendment to the International Code of Zoological Nomenclature regarding electronically published works (
Zelus Fabricius, 1803, p. 281, orig. descr.; Latreille, 1804, p. 260, list; Latreille, 1807, p. 129, list; Latreille, 1810, p. 433, type desig.; Lepeletier and Serville, 1825, p.815, list and descr.; Laporte, 1832, p. 9, type desig.; Burmeister, 1835, p. 225, descr.; Brulle, 1836, p. 316-317, descr.; Blanchard, 1840, p. 100, descr. and note; Blanchard, 1845, p. 433, 438, list and note; Herrich-Schaeffer, 1848, p. 88, descr. and note; Kolenati, 1857, p. 458-459, descr.; Stål, 1861, p. 148, descr.; Stål, 1862, p. 449-454, key and subgeneric descr. (with subgenus Zelus); Carpenter and Westwood, 1863, p. 565, note; Mayr, 1866, p. 138, list; Stål, 1866, p. 296, list; Stål, 1868, p. 107, restriction of definition; Stål, 1872, p. 69, 88, key and cat. (with subgenus Zelus); Walker, 1873, VII., p. 49, key, VIII., p. 131-136, cat.; Berg, 1879, p. 150, list (with subgenus Zelus); Uhler, 1886, p. 24, checklist; Provancher,1887, p. 179, note; Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 251, cat. and note; Kirkaldy, 1900a, p. 263, type verification; Kirkaldy, 1900b, p. 242, syn.; Kirkaldy, 1902, p. 149, note; Fracker, 1913, p. 223, 238-240, key and note (with subgenus Zelus); Van Duzee, 1916, p. 30, checklist (with subgenus Zelus); Van Duzee, 1917, p. 258-259, cat. (with subgenus Zelus); Blatchley, 1926, p. 567-568, key, descr. and note (with subgenus Zelus); Readio, 1927, p. 167, 168-169, key, descr. and note; Zimmerman, 1948, p. 137, note; Wygodzinsky, 1949a, p. 48, checklist; Fracker and Usinger, 1949, p. 277, key to nymphs; Alayo, 1967, p. 5, 35, list, key and note; Hart, 1986, key to North American species; Hart, 1987, key to Caribbean species; Maldonado, 1990, p. 325-332, cat.
Reduvius Fabricius, 1775 (type by subsequent desig., Cimex personatus Linnaeus, 1758), Lepeletier and Serville, 1825 (in part), p. 272, descr.; Perty, 1834 (in part), p. 173, list of species.
Arilus Hahn, 1831 (type by subsequent designation, Cimex carinatus Forster, 1771); Burmeister, 1835 (in part), p. 227-228, descr.; Herrich-Schaeffer, 1848 (in part), p. 33-35, descr.
Euagoras Burmeister, 1835 (type by subsequent designation, E. stollii Burmeister, 1835) (in part), p. 226, descr.; Amyot and Serville, 1843 (in part), p. 368, descr. (as Evagoras); Herrich-Schaeffer, 1848 (in part), p. 43-44, descr.; Stål, 1855 (in part), p. 189, list (as Eccagoras); Stål, 1861, p. 148, (in part) junior syn. of Zelus Fabr.; Mayr, 1866, p. 139, list; Walker, 1873, p. 49, 117, key and cat.; Provancher, 1887, p. 182, descr. (as Evagoras); Kirkaldy, 1900b, p. 242, junior syn. of Zelus Fabr.; Kirkaldy, 1903, p. 215-216, note.
Diplodus Amyot and Serville, 1843, p. 370, descr.; Burmeister, 1853, p. 91, list (included in Euagoras Burm.); Stål, 1860, p. 74, list; Stål, 1862, p. 450, descr. (as subgenus of Zelus); Stål, 1866, p. 296, key; Stål, 1872, p. 90, list (as subgenus of Zelus); Walker, 1873, VII., p. 49, VIII., p. 123, key and cat. (as Diploda); Berg, 1879, p. 151, list (as subgenus of Zelus); Uhler, 1886, p. 24, checklist; Provancher, 1887, p. 179, key and descr.; Kirkaldy, 1903, p. 232, note; Fracker, 1913, p. 239, 240, key and list (as subgenus of Zelus).
Darbanus Amyot and Serville, 1843 (type by monotypy, D. nigrolineatus); Provancher, 1872, p. 106, species descr.; Uhler, 1886, p. 24, checklist; Provancher, 1887, p. 179, 181, key and note; Van Duzee, 1912, p. 324; Fracker, 1913, p. 241, note.
Pindus Stål, 1862, p. 454, orig. descr. (as subgenus of Zelus); Stål, 1866, p. 296, key (as genus); Stål, 1872, p. 92, list and cat., as subgenus of Zelus); Walker, 1873, VII., p. 66, list and cat. (as genus); Berg, 1879, p. 150, list (as subgenus of Zelus) ; Thierry and Severin, 1896, p. 151, cat.; Fracker, 1913, p. 223, 240, key and list; Van Duzee, 1916, p. 30, checklist; Van Duzee, 1917, p. 261, cat.; Blatchley, 1926, p. 569, key.
Diplacodus Kirkaldy, 1900b, p. 242, new name for Diplodus A. and S. (preocc.).
Diplocodus Van Duzee, 1916, p. 30, checklist (new name for Diplacodus Kirkaldy, preocc.); Van Duzee, 1917, p. 260, cat.; Blatchley, 1926, p. 569, key.
Iquitozelus Bérenger, 2003, p. 23, orig. descr., syn. nov. (current study).
Male: Small to large, total length 8-25 mm (
Female: Larger than male. Coloration usually similar to that of male and more variable in some species, but may differ between sexes dramatically in certain species. Eye and ocellus smaller than in male in some species. Basiflagellomere not swollen and about equal diameter as or smaller than pedicel. Lateral process on humeral angle, if present, usually more produced and longer than in male. Mesofemur slightly swollen in many species. Lateral margins of abdomen expanded in some species.
This genus is distinguished from other genera of the New World Harpactorini by the cylindrical head, the length of the head being at least 1.9X its width; the unarmed antenniferous tubercles; the second labial segment being at least 1.3x the length of the first segment; the long scape and basiflagellomere that are subequal in length and the short pedicel and distiflagellomere; the generally unarmed (i.e. no tubercles or spines) disc of the posterior pronotal lobe (except in Zelus tetracanthus Stål, 1862, Zelus lewisi sp. n. and Zelus minutus Hart, 1987); the humeral angle with or without process, and if present, usually not prominently projected; the legs with sundew setae and sticky glands (
Native to (except for Chile) and throughout the New World, including the Caribbean, with highest diversity in the Neotropics. One species (Z. renardii) has been introduced to Hawaii, the Polynesian islands, Jamaica, Philippines, Spain, Greece and Chile.
We provide a non-exhaustive account of the biology of various species of this genus. As with other harpactorines, species of Zelus generally do not show associations with or preferences of host plants, probably due to their generalist habits. However, two recent studies have found two species of Zelus that have both nymphs and adults occurring in the same plant species in relatively large quantities. In
Several species of Zelus are possibly mimics of various other insects. Zelus errans Fabricius, 1803, Zelus vespiformis Hart, 1987 and to some extent Zelus vagans Fabricius, 1803 and Zelus gracilipes sp. n. have wing and body color patterns similar to many braconid wasps, an intriguing form of mimicry seen also in a number of other Neotropical harpactorine genera. Zelus vagans shows areas of black and orange colors, however, the posterior pronotal lobe is medially dark and laterally orange. Zelus gracilipes also shows a uniformly orange posterior pronotal lobe, but the hemelytron is uniformly dark and lacks the banding pattern typical to a wasp mimic. Zelus nigromaculatus Champion, 1899 has an appearance similar to that of a typical vespid, the only species in this genus with that kind of color pattern. Zelus laticornis (Herrich-Schaeffer, 1853), Zelus grassans Stål, 1862 and Zelus ruficeps Stål, 1862 have red and dark markings on abdomens and orange or reddish dorsal surfaces, a pattern found in many species of pyrrhocorids (e.g., Dysdercus spp.) and coreids (e.g. Hypselonotus spp.). Interestingly, in Z. laticornis, it is only the females showing this coloration. Certain color forms of Z. longipes are possibly mimics of the milkweed bug, Oncopeltus fasciatus (Dallas).
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,
The genus that we are uncertain about its relationship with Zelus is Pronozelus Forero, erected by
Except for several pairs or complexes of closely related species, identification of males can be almost always unambiguously performed based on exposed genitalic structures such as paramere and medial process, further corroborated with phallic structures, external morphology and coloration. Identification of females of many species, where females appear to be as distinct as males, is straightforward based on coloration and external morphology. However, identification can be difficult for closely related species, where females are indistinguishable based on external morphology. In these cases, association of males and females and identification of females were primarily based on collecting event information. Sexual dimorphism presents another special challenge. While most species show limited sexual dimorphism that does not go beyond minor size and coloration differences, some species exhibit pronounced differences between the sexes (see Material and Methods for discussion of association of male and female specimens). Based on the observation that species in closely related genera do not exhibit strong sexual dimorphism, we here hypothesize that pronounced sexual dimorphism is a derived condition within Zelus.
We find here that previous subgeneric groups are based on superficial resemblance and these are not adopted. Instead, we recognize eleven species groups in the current study, based primarily on characters of the male genitalia, but also on non-genitalic external morphology if those characters can be applied to both sexes. Several species for which only females are known are therefore not assigned to a species group. Although the groupings proposed here are not based on a cladistic analysis, they show a degree of congruence with the relationships recovered in the phylogenetic analysis based on molecular data in
1. Zelus tetracanthus species group.
Zelus minutus Hart, 1987, Zelus prolixus Stål, 1860, Zelus rosulentus sp. n. and Zelus tetracanthus Stål, 1862.
Members of this group have a rather broad, indistinct medial process, the base of which is nearly continuous with or inseparable from the ventral rim of the pygophore. We speculate that this character represents a plesiomorphic condition as it is seen in several other genera of the New World Harpactorini and thus the condition of the medial process does not necessarily support the monophyly of this group. Zelus tetracanthus and Z. minutus also both have tubercles on the disc of the posterior pronotal lobe, which are more pronounced in the former. Comparative views of male genitalia are shown in
2. Zelus luridus species group.
Zelus ambulans Stål, 1862, Zelus antiguensis sp. n., Zelus exsanguis Stål, 1862, Zelus grandoculus sp. n., Zelus luridus Stål, 1862 and Zelus spatulosus sp. n.
This is a group of species with primarily a North American distribution, with some species extending to northern Central America. The males show an apically expanded paramere and a triangular medial process that has a protrusion at the base but lacks any apical modifications. Notably, Z. spatulosus has a slender medial process, deviating greatly from the remainders of the group. It is placed in this group mainly because of the apically expanded paramere and the uniform coloration. Zelus ambulans and Z. exsanguis have the humeral angle elevated to about same level of and nearly continuous with the disc of the posterior pronotal lobe, a condition rarely seen in the genus. The coloration is quite homogenous among members of this genus, most of which have a uniform greenish (in live specimens) or dull brownish (in preserved specimens) habitus, with only Z. ambulans showing variable patterns or banding on the pronotum or legs. Comparative views of male genitalia are shown in
3. Zelus mimus species group
Zelus inconstans Champion, 1898 and Zelus mimus Stål 1862.
Members of this group, consisting of only two species, exhibit a highly unique paramere and a medial process of the pygophore. The paramere is slender and apically curved dorsad at an angle of nearly ninety degrees. The medial process, as is especially evident in Z. inconstans, possesses a simple posterior liplike fold at the apex; its lateral margins are subparallel and not broadened significantly at the base. Both have a semi-cylindrical dorsal phallothecal sclerite which is modified by a fold running obliquely toward the base from the middle of the lateral margins. Both species, being quite small, exhibit the usual reduction of setal tracts common to nearly all small species. Comparative views of male genitalia are shown in
4. Zelus nugax species group.
Zelus grassans Stål, 1862, Zelus illotus Berg, 1879, Zelus impar Kuhlgatz, 1902, Zelus nugax Stål, 1862 and Zelus pedestris Fabricius, 1803.
This is a group of smallish species with quite variable distributional ranges. The defining characters include a slender, laterally compressed medial process that is curved or recurved, and an acute apex of the dorsal phallothecal sclerite (except in Z. grassans). Zelus nugax has one of the widest distribution ranges in this genus, ranging from much of Mexico to northern South America. Zelus grassans is found primarily in Central America and the remaining two species mainly in northern South America. Comparative views of male genitalia are shown in
5. Zelus puertoricensis species group.
Zelus bruneri De Zayas, 1960, Zelus puertoricensis Hart, 1987, Zelus subimpressus Stål, 1872 and Zelus zayasi Bruner and Barber, 1937.
Members of this group are restricted to the Caribbean. They can be easily recognized by the rather slender body form. The posteriorly directed, robust medial process with a somewhat blunt apical protrusion is also distinctive of this group. The basal plate arms are widely separate and diverging and these features are rare in other species in the genus. They show resemblance to species of the Zelus renardii species group, especially to Z. cervicalis. Zelus bruneri was not physically examined, but the rather slender body form as seen in the original illustration places it within this group. Comparative views of male genitalia are shown in
6. Zelus renardii species group.
Zelus cervicalis Stål, 1872 and Zelus renardii Kolenati, 1856.
The two members of this group are very likely sister species since they share a number of unique characters: the apex of the medial process is greatly bent ventrad and hooklike, the lateral margin of the dorsal phallothecal sclerite is recurved dorsad and the basal part of the strut is absent. Both species are mainly distributed in North and Central America, but Z. cervicalis extends to northern South America. Comparative views of male genitalia are shown in
7. Zelus armillatus species group.
Zelus amblycephalus sp. n., Zelus annulosus (Stål, 1866), Zelus armillatus (Lepeletier & Serville, 1825), Zelus conjungens (Stål, 1860), Zelus janus Stål, 1862, Zelus leucogrammus (Perty, 1833), Zelus lewisi sp. n., Zelus litigiosus Stål, 1862, Zelus ruficeps Stål, 1862, Zelus sulcicollis Champion, 1899, Zelus umbraculoides sp. n. and Zelus umbraculus sp. n.
This is one of the two largest groups in the genus (the other being the Zelus panamensis species group). Species in this group are generally robust and large-sized (15-25 mm), and some are among the largest in the genus. The most distinctive character is that of the medial process, which has the apex slightly projected into two minute small lateral prongs or processes. This condition is different from that in several species groups listed below, where the apex of the medial process is hook-like and more strongly projected. The lateral spine of the humeral angle tends to be pronounced and somewhat broadened into a dentate effect. The pygophore is large, rounded, and somewhat shortened relative to the total length of the individual. The dorsal phallothecal sclerite having dorsolateral expansions or projections close to the basal arm is also unique to some species of this group. This condition, however, is not seen in Z. amblycephalus, Z. umbraculus, or Z. umbraculoides, which appear to be divergent from the remainders of the group, but the features of the medial process unambiguously place them in this species group. Comparative views of male genitalia are shown in
8. Zelus longipes species group.
Zelus bahiaensis sp. n., Zelus errans Fabricius, 1803, Zelus longipes (Linnaeus, 1767), 1803 and Zelus vespiformis Hart, 1987.
This and the next species group (Zelus vagans species group) possess dense, spine-like setae on the head and pronotum, and a rounded, unarmed humeral angle, both characters rather unique in Zelus and probably synapomorphies uniting the two groups. The former character is possibly homoplastic as it is also seen in two species in the Zelus armillatus species group. The medial process is slender and cylindrical and this condition is among the most extreme in the genus. It is semi-erect and posteriorly directed. The paramere exceeds the apex of the medial process. The dorsal phallothecal sclerite has subparallel margins and lacks obvious modifications or ornamentations (except for small lateral folds in Z. longipes). Some individuals of Z. errans and Z. vespiformis appear to be wasp mimics. Comparative views of male genitalia are shown in
9. Zelus vagans species group.
Zelus aithaleos, Zelus championi sp. n., Zelus fuliginatus sp. n., Zelus gracilipes sp. n. and Zelus vagans Fabricius, 1803.
Species of the Zelus vagans group share two characters also present in the preceding group (Zelus longipes species group): spinelike setae and rounded humeral angle. However, they differ in the structure of the male genitalia in significant ways. The medial process shapes like a somewhat laterally flattened cone. It is relatively broad at base, narrowing toward the apex, and is laterally compressed. The medial process is posteriorly directly, nearly horizontal. The paramere is removed from or barely reaching apex of the medial process. Furthermore, the phallus is elongated and slightly constricted toward the apex (not conspicuous in Z. gracilipes). Zelus vagans and Z. gracilipes also resemble wasps to some extent, but both not as perfectly as seen in Z. errans and Z. vespiformis. Comparative views of male genitalia are shown in
10. Zelus panamensis species group.
Zelus banksi sp. n., Zelus cordazulus sp. n., Zelus filicauda Bergroth, 1893, Zelus gilboventris sp. n., Zelus korystos Hart, 1986, Zelus nigromaculatus Champion, 1899, Zelus panamensis sp. n., Zelus truxali sp. n., Zelus varius (Herrich-Schaeffer, 1853) and Zelus xouthos sp. n.
This is another large group with ten species. Interestingly, most (seven) are new species. It is characterized by having an acute apical modification usually in the shape of a hook on the medial process and the conspicuous medial carination of the apical part of the dorsal phallothecal sclerite. The condition of the apical modification of the medial process differs from that in the Zelus armillatus species group in that it is much more prominent, usually acute and sometimes extending further ventrally. Rugulosity of the posterior pronotal lobe is highly pronounced relative to the other groups. Sexual dimorphism is pronounced in some species in this group (e.g., Z. gilboventris and Z. truxali). Most species in this group are concentrated in southern Central America and northern South America. Comparative views of male genitalia and habitus images are in
11. Zelus erythrocephalus species group.
Zelus auralanus sp. n., Zelus casii sp. n., Zelus chamaeleon Stål, 1872, Zelus erythrocephalus Fabricius, 1803, Zelus kartabenoides sp. n., Zelus kartabensis Haviland, 1931, Zelus laticornis (Herrich-Schaeffer, 1853), Zelus mattogrossensis Wygodzinsky, 1947, Zelus paracephalus sp. n., Zelus russulumus sp. n. and Zelus versicolor (Herrich-Schaeffer, 1848).
Two diagnostic characters identify members of this group. The medial process possesses a broad ridge-like projection or carina that initiates from the apex and extends ventrally or is removed from apex. The second feature is the apically oriented lateral sharp processes or projections on the dorsal phallothecal sclerite. These are not to be confused with the lateral expansion seen in the Zelus armillatus species group, where the direction of the expansion is laterad. In Z. auralanus and Z. versicolor, this process is short and somewhat dorsally directed, rather than apically directed. Three species, Z. kartabenoides, Z. kartabensis and Z. chamaeleon lack this structure. Their placement in this group is primarily based on the configuration of the medial process and the absence of characters of other groups. Also, the longitudinal ridge-like elevation or hook on the medial process is similar to the condition in another species, Z. laticornis, although the latter has a short modification. In this species group the parameres are usually somewhat bulbous and curved medially with moderate to long erect setae on the apical 1/2. The medial process is broadened at base, and usually anteroposteriorly compressed. Furthermore, the basal plate of the phallus is strongly curved in some members of this species group. Pronounced sexual dimorphism is seen in some species of this group. Notably, three species, Z. erythrocephalus, Z. paracephalus and Z. russulumus have purple, blue or greenish iridescence on the membrane of the hemelytron. Species of this group show a predominant southern South American distribution, with a few found only from the Amazons. Comparative views of male genitalia and habitus images are in
Because of the heavy emphasis on male genitalic characters for grouping species, four species described only from females are not placed in any of the species groups defined in the above. These are: Zelus fasciatus Champion, 1899, Zelus plagiatus (Signoret, 1852), Zelus sphegeus Fabricius, 1803 and Zelus means Fabricius, 1803. Zelus fasciatus is similar to the females of some of the species in the Zelus panamensis species group and also occurs in an overlapping geographical region (southern Central America). Zelus plagiatus and Z. sphegeus show resemblance to the females of Z. versicolor, which is in the Zelus erythrocephalus species group. Zelus means, by possessing a rounded humeral angle and spinelike setae, aligns most closely with the Zelus vagans species group and the Zelus longipes species group. A future cladistic analysis, including morphological and molecular data, is needed to test the monophyly of these species groups and may also have the potential to place these female-based species.
Five species are removed from Zelus: Z. araneiformis, Zelus gradarius Bergroth, 1905, Z. modestus (Stål, 1862), Zelus subfasciatus Stål, 1860 and Zelus vittaticeps Stål, 1866. These species represent an undescribed genus "Hartzelus" and will be treated in a separate study. They will be listed as Harpactorini incertae sedis until their generic placement is formally clarified.
Zelus aithaleos Zhang & Hart, sp. n., habitus
Zelus aithaleos Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Similar to male, except for the following. Larger than male, total length 13.87–17.61 mm (mean 16.27 mm,
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 (
A unicolourous near-black dorsum, including the head, the pronotum and the corium, separates Z. aithaleos from both sexes of Z. gracilipes, Z. vagans, and Z. means (known from females only), all of which have some orange, yellow or reddish colors. The dark dorsal profile is shared with Z. championi (only the male is known) and Z. fuliginatus. A longitudinal lateral patch of whitish recumbent setae on the postocular lobe serves to separate this species from Z. fuliginatus. It is distinguished by a dark abdomen from Z. championi, which has a brightly red abdomen.
From Greek aithales.
South America (
Zelus amblycephalus Zhang & Hart, sp. n., habitus
Zelus amblycephalus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
Can be recognized by the uniform pale coloration, the unpatterned legs (
Among species of the Zelus armillatus group (
Southern Mexico to northern South America and part of Brazil (
Zelus ambulans Stål, 1862, p. 451, orig. descr.; Stål, 1872, p. 91, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 259–260, Tab. XV. fig. 23, 23a, junior syn. of Z. exsanguis; Maldonado, 1990, p. 327. cat. and junior syn. of Z. exsanguis. stat. rev. (current study).
Diplodus ambulans: Uhler, 1886, p. 24, checklist; Walker, 1873, cat.
Zelus ambulans Stål, 1862, habitus
Zelus ambulans Stål, 1862, male genitalic structures
Male: (
Female: (
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 (
North and Central America (
Although this species shows very little morphological variations, color patterns within an area do vary considerably. The dark area at the posterior margin of the longitudinal medial sulcus of the anterior lobe, which serves to easily distinguish Z. ambulans from Z. exsanguis, is relatively constant. Other colors, specifically that of the posterior pronotal lobe and the femoral apices vary from quite light to very dark brown in any given locality. There is also an occasional specimen with somewhat darker hemelytron, but this does not show the wide range of variations of the aforementioned characters.
Most specimens examined have been collected from moderate to high altitudes.
Diplodus annulosus Stål, 1866, p. 299, orig. descr.; Walker, 1873, p. 126, cat.
Zelus annulosus: Stål, 1872, p. 92, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Fracker and Bruner, 1924, p. 170, note; Wygodzinsky, 1949a, p. 48, checklist; Maldonado, 1990, p. 326, cat.
Zelus annulosus (Stål, 1866), habitus
Zelus annulosus (Stål, 1866), male genitalic structures
Male: (
Female: Similar to male, except for the following. Larger than male, total length 21.19–22.72 mm (mean 21.91 mm,
Recognized by the following combination of characters: the posterior pronotal and corium dark green; the legs with four to five alternative yellow and black bands; the head, pronotum, scutellum and corium with moderately dense, black, erect, spine-like setae; the rather long and slender legs, the profemur 1/2 of body length; the rather long postocular lobe, enlarged at posterior 3/4; and the quadrate cell on corium rather slender, length more than 2x width.
Males can also be recognized by the long paramere, reaching apex of medial process; the apex of paramere recurved; the medial process apically with two lateral sharp projections; the membranous sclerite between paramere and medial process, not distinctly protruding posteriorly; and the dorsal phallothecal sclerite with lateral expansion close to basal arm, sharp, dorsad.
South America (
Zelus antiguensis Zhang & Hart, sp. n., habitus
Zelus antiguensis Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
As with several members of the Zelus luridus species group, the coloration is greenish-brown, rather uniform. The medial process is triangular, its base distinct from the rest of pygophore ventral rim and apex without modification. Can be distinguished from males of other species of the Zelus luridus species group (
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 (
Named after the type locality, Antigua, in Guatemala.
Southern Mexico and Guatemala (
Reduvius armillatus Lepeletier and Serville, 1825, p. 278, orig. descr.
Diplodus armillatus: Amyot and Serville, 1843, p. 370, descr.; Stål, 1860, p. 75, list; Walker, 1873, p. 123, cat.
Euagoras armillatus: Herrich-Schaeffer, 1853, p. 91, list.
Zelus armillatus: Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Wygodzinsky, 1949a, p. 49, checklist; Mayr, 1866, p. 138-139, senior syn. of Z. brasiliensis, Z. aurantiacus, Z. guttifer and Z. conjungens; Berg, 1879, p. 151-152, list and nymphs (subgenus Diplodus); Costa Lima, 1940, 218, list (subgenus Diplocodus); Wygodzinsky, 1957, p. 268, note; Wygodzinsky, 1960; p. 307, list; Maldonado, 1990, p. 326, cat.; Van der Heyden, p. 85-90, new record (misidentification, should be Zelus atripes).
Reduvius brasiliensis Lepeletier and Serville, 1825, p. 278, orig. descr.
Diplodus brasiliensis: Amyot and Serville, 1843, p. 370, descr.; Stål, 1860, p. 75, note; Mayr, 1866, p. 138- 139, junior syn. of Z. armillatus; Walker, 1873, p. 123, cat.
Euagoras brasiliensis: Herrich-Schaeffer, 1853, p. 91, list.
Zelus brasiliensis: Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, junior syn. of Z. armillatus.
Arilus aurantiacus Herrich-Schaeffer, 1848, p. 35-36 Tab. CCLXI. fig. 809, orig. descr. and fig.; Mayr, 1866, p. 138-139, junior syn. of Z. armillatus; Stål, 1872, p. 90, junior syn. of Z. armillatus.
Euagoras aurantiacus: Herrich-Schaeffer, 1853, p. 91, list (aurantius (sic)).
Ploeogaster aurantiacus: Herrich-Schaeffer, 1853, p. 168, list.
Arilus guttifer Herrich-Schaeffer, 1848, p. 36, Tab. CCLXI, fig. 810, orig. descr. and fig.; Mayr, 1866, p. 138-139, junior syn. of Z. armillatus; Stål, 1872, p. 90, junior syn. of Z. brasiliensis.
Euagoras guttifer: Herrich-Schaeffer, 1853, p. 92, list.
Ploeogaster guttifer: Herrich-Schaeffer, 1853, p. 168, list.
Diplodus guttifer: Stål , 1860, p. 74, descr.; Walker, 1873, p. 126, cat.
Zelus guttifer: Stål 1862, p. 453, note.
Arilus guttifer , Mayr, 1866, p. 138-139, junior syn. of Z. armillatus .
Zelus armillatus (Lepeletier & Serville, 1825), habitus, males
Zelus armillatus (Lepeletier & Serville, 1825), habitus, females
Zelus armillatus (Lepeletier & Serville, 1825), habitus, females
Zelus armillatus (Lepeletier & Serville, 1825), male genitalic structures
Male: (
Female: (
The large and robust body, the dorsal coloration usually bright, yellow or red with black, the medial process short and relatively slender are characteristic to Z. armillatus. Male genitalic structures of Z. armillatus and Z. janus are nearly identical, but these two species do not overlap in range and are sufficiently different in non-genitalic morphological characters, which allow them to be easily separated.
The only species with which Z. armillatus is sympatric which may cause some identification problems is Z. conjungens. It may be distinguished from that species by the characters discussed under Z. conjungens.
South America (
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.,
Zelus auralanus Zhang & Hart, sp. n., habitus
Zelus auralanus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
Can be readily recognized by the uniformly brown dorsal coloration; the darkened tibial apex; the humeral angle elevated to level of disc; the dorsal setae on head and pronotum appearing somewhat golden, shining when viewed under magnification. Males can also be recognized by the gradually enlarged paramere; the triangular medial process, curved slightly posteriad in the middle, apex with a hooklike projection; and the dorsal phallothecal sclerite with short, dorsad projections sub-laterally.
The species epithet indicates the somewhat reddish tone of the coloration.
South America (
Zelus bahiaensis Zhang & Hart, sp. n., habitus
Zelus bahiaensis Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Unknown.
Recognized by the following combination of characters: the anterior pronotal lobe dark brown and the posterior pronotal lobe orange; the 1A an Pcu not intersecting, short crossvein between them; the long and slender, cylindrical medial process; the medial process apically folded posteriad; and the rather long paramere.
Named after the Brazilian state Bahia, where the holotype was collected.
South America (
Zelus banksi Zhang & Hart, sp. n., habitus
Zelus banksi Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Unknown.
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 (
Named after N. Banks, the collector of the type specimen.
Southern Central America and northern South America (
Zelus bruneri De Zayas, 1960, p. 125–127, orig. descr. and fig; Alayo, 1967, p. 5, 36, 37, list, key and note; Hart, 1987, p. 296-297, note and key; Maldonado, 1990, p. 326, cat.
Known only from Cuba.
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.
Zelus casii Zhang & Hart, sp. n., habitus
Zelus casii Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Unknown.
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.
Named after Casi.
South America (
Several characters of Z. casii are highly unique among all species of Zelus. It has an extraordinarily long postocular lobe. The medial process is very broad, has lateral ridge-like elevations, and the apex is emarginate.
Zelus cervicalis Stål, 1872, p. 90, orig. descr. (subgenus Zelus); Uhler, 1876, p. 61, list (reprint); Uhler, 1886, p. 24, checklist; Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 255, cat.; Banks, 1910, p. 16, cat.; Torre-Bueno and Engelhardt, 1910, p. 150, note; Van Duzee, 1912, p. 324, senior syn. of Z. marginata (Provancher); Fracker, 1913, p. 239, 240, key and list (subgenus Zelus); Torre-Bueno, 1913, p. 60, list; Barber, 1914, p. 506, list; Van Duzee, 1916, p. 30, checklist (subgenus Zelus) ; Van Duzee, 1917, p. 260, cat. (subgenus Zelus); Dozier, 192,0 p. 357, list; Blatchley, 1926, p. 569, key and note (subgenus Zelus); Readio, 1927, p. 169, 170, key and descr.; Wygodzinsky, 1949a, p. 48, checklist; Elkins, 1951, p. 410, list; Sibley, 1951, p. 92, list; Kelton, 1968, p. 1071, note; Snow, 1906, p. 180, list; Van Duzee, 1909, p. 177, list; Osborne and Drake, 1915, p. 531, note; Brimley, 1938, p. 73, list; Elliott, 1938, p. 39, list; Tenhet and Howe, 1939, p. 24, note; Drew and Schaeffer, 1962, p. 106, list; Oliver, 1964, p. 316, note; Whitcomb and Bell , 1964, p. 22, List and note; Hart, 1986, p. 542-543, lectotype desig., redescription, note, fig. and key; Maldonado, 1990, p. 326, cat.
Evagoras marginata Provancher, 1887, p. 182–183, orig. descr.; Van Duzee, 1912, p. 324, junior syn. of Z. cervicalis; Kelton, 1968, p. 1071, note.
Zelus marginatus: Lethierry and Severin, 1896, p. 152, cat.; Banks, 1910, p. 16, cat.
Zelus pictipes Champion, 1898, p. 255, Tab. XV, fig. 14, orig. descr, and fig.; Fracker, 1913, p. 239, 240, key and list; Van Duzee, 1916, p. 30, checklist (subgenus Zelus) ; Van Duzee, 1917, p. 259, cat. (subgenus Zelus); Readio, 1927, p. 169, 170, key and descr.; Wygodzinsky, 1949a, p. 50, checklist; Snow, 1906, p. 180, list; Elkins, 1951, p. 410, list; Sibley, 1951, p. 92, list; Drew and Schaeffer, 1962, p. 106, list; Hart, 1986, p. 542, lectotype desig. and junior syn. of Z. cervicalis.
Zelus cervicalis Stål, 1872, habitus
Zelus cervicalis Stål, 1872, male genitalic structures
Male: (
Female: (
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.
South Atlantic and Gulf Coast states of the United States, southeastern Arizona, most of Mexico, Central America and Northern Colombia (
Hart (1986) 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 distribution pattern for Z. renardii and Z. cervicalis found by Hart. Based mainly on the coloration of the legs, Hart (1986) 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. 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 (
The images of the lectotype of Z. cervicalis are available on the 'Types of Heteroptera' website of the Swedish Museum of Natural History.
Zelus chamaeleon Stål, 1872, p. 90–91, orig. descr. and cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 260, note; Wygodzinsky, 1949a, p. 48–49, checklist; Maldonado, 1990, p. 326, cat.
Zelus chamaeleon Stål, 1872, habitus
Zelus chamaeleon Stål, 1872, male genitalic structures
Male: (
Female: (
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.
South America (
Zelus championi, Zhang & Hart, sp. n., habitus
Zelus championi, Zhang & Hart, sp. n., pygophore, lateral and posterior views
Male: (
Female: Unknown.
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 (
This species epithet is a patronym, in honor of entomologist George C. Champion (1851-1927), who authored several volumes on Rhyncophora (Heteroptera) in the Biologia Centrali Americana series.
Central and South America (
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.
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.
Diplodus cognatus Costa, 1864, p. 81, orig. descr.; Uhler, 1886, p. 24, checklist; Walker, 1873, p. 125, cat.
Zelus cognatus: Stål, 1872, p. 91, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 259–260, junior syn. of Z. exsanguis Stål. stat. rev. (current study).
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."
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.
Diplodus conjungens Stål, 1860, p. 75, orig. descr.; Mayr, 1866, p. 138–139, junior syn. of Z. armillatus; Walker, 1873, p. 125, cat.
Zelus conjungens: Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, junior syn. of Z. armillatus; Wygodzinsky, 1949a, p. 49, checklist. stat. rev. (current study).
Zelus atripes Champion, 1898, p. 259, Tab. XV. fig. 22, orig. descr. and fig.; Wygodzinsky, 1949a, p. 48, checklist; Maldonado, 1990, p. 326, cat. syn. nov. (current study).
Zelus conjungens (Stål, 1860), habitus
Zelus conjungens (Stål, 1860), male genitalic structures
Male: (
Female: (
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.
Southern Central America, northern South America and Southern Brazil (
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.
Zelus cordazulus Zhang & Hart, sp. n., habitus
Zelus cordazulus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
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.
Named after the type locality "Cordillera" in Peru.
South America (
Iquitozelus couturieri Bérenger, 2003, p. 23, orig. descr. and fig.
Zelus couturieri (Bérenger, 2003), comb. nov. (current study).
The connexivum segment VI with foliaceous expansion is unique among all females of Zelus spp.
South America (
Zelus errans Fabricius, 1803, p. 282, orig. descr.; Stål, 1868, p.108, descr., note and senior syn. of Z. cursitans; Stål, 1872, p.88, cat.; Walker, 1873, p. 135, cat.; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 326, cat.
Zelus cursitans Fabricius, 1803, p. 284, orig. descr.; Blanchard, 1840, p. 101, descr.; Stål, 1868,p. 108, junior syn. of Z. errans.
Zelus errans Fabricius, 1803, habitus
Zelus errans Fabricius 1803, male genitalic structures
Male: (
Female: (
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 (
South America (
Zelus errans is quite closely related to Z. vespiformis, and they are possibly sister species. These two species appear to be allopatric, with their boundaries roughly lying across central Colombia and southern Venezuela.
Zelus erythrocephalus Fabricius, 1803, p. 283, orig. descr.; Blanchard, 1840, p. 101, cat. (erytrocephalus sic.); Stål, 1872, p. 92, cat. (subgenus Diplodus); Bergroth, 1893, p. 63, note; Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 257, note; Brindley, 1931, p. 137, 151, list and note; Wygodzinsky, 1949a, p. 49, checklist; Zimsen, 1964, p. 338, list; Maldonado, 1990, p. 327, cat.
Euagoras erythrocephalus: Burmeister, 1835, p. 227, list.
Diplodus erythrocephalus: Stål, 1868, p. 283, descr.; Walker, 1873, p. 125, cat.; Van Duzee, 1901, p. 351, note.
Zelus erythrocephalus Fabricius, 1803, habitus
Zelus erythrocephalus Fabricius, 1803, male genitalic structures
Male: (
Female: (
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.
South America (
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.
Zelus exsanguis Stål, 1862, p. 452, orig. descr.; Stål, 1872, p. 91, cat. (subgenus Diplodus); Walker, 1873, p. 124, cat.; Uhler, 1894, p. 283, list; Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 259–260, senior syn. of Z. luridus Stål, Z. ambulans Stål and Z. cognatus Costa; Banks, 1910, p. 16, cat.; Fracker, 1913, p- 239, 240, 241, key, list and note (subgenus Diplodus); Van Duzee, 1916, p, 30, checklist (subgenus Diplodus); Van Duzee, 1917, p. 260, cat. (subgenus Diplodus); Wygodzinsky, 1949a, p. 49, checklist; Hart, 1986, p. 539, redescription, lectotype desig., note, fig. and key; Maldonado, 1990, p. 327, cat.
Diplodus exsanguis: Uhler, 1886, p. 24, checklist.
Zelus exsanguis Stål, 1862, habitus
Zelus exsanguis Stål, 1862, male genitalic structures
Male: (
Female: (
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 (
Mexico to Panama (
Zelus fasciatus Champion, 1899, p. 257, Tab. XV. fig. 18, orig. descr. and fig.; Kuhlgatz, 1902, p. 266, note; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 327, cat.
Zelus fasciatus Champion, 1899, habitus
Male: unknown.
Female: (
The rather unique dorsal color pattern easily distinguishes this species from all other species in the genus.
Southern Central America and northern South America (
Zelus filicauda Bergroth, 1893, p. 63, orig. descr.; Lethierry and Severin, 1896, p. 152, cat.; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 327, cat.
Zelus filicauda Bergroth, 1893, habitus
Zelus filicauda Bergroth, 1893, male genitalic structures
Male: (
Female: Unknown.
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 (
South America (
Zelus fuliginatus Zhang & Hart, sp. n., habitus
Male: (
Female: (
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 (
The species epithet means 'soot' or painted black, referring to the black dorsal coloration of this species.
Northern South America (
Zelus gilboventris Zhang & Hart, sp. n., habitus
Zelus gilboventris Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
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.
South America (
Zelus gracilipes Zhang & Hart, sp. n., habitus
Male: (
Female: (
Recognized by the posterior pronotal lobe orangish-brown; the legs uniformly blackish-brown, 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 (
From Latin gracilis, meaning slender.
South America (
Zelus grandoculus Zhang & Hart, sp. n., habitus
Zelus grandoculus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Unknown.
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 (
The species epithet combines grandis, meaning large, with oculus, meaning eye, to indicate the prominently large-sized compound eye.
Known only from type locality in Guatemala (
Zelus grassans Stål, 1862, p, 450, orig. descr.; Stål, 1872, p. 91, cat.; Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 256–257, Tab. XV, fig. 16, 17, note and fig.; Kuhlgatz, 1902, p. 266, note; Fracker, 1913, p. 239, 240, key and list (subgenus Diplodus); Wygodzinsky, 1949a, p. 49, checklist; Hart, 1986, p. 546-547, redescription, note, fig. and key; Maldonado, 1990, p. 327, cat.
Diplodus grassans: Walker, 1872, p. 124, cat.; Uhler, 1886, p. 24, checklist.
Zelus grassans Stål, 1862, male genitalic structures
Male: (
Female: (
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.
From Mexico to Panama (
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.
Zelus illotus Berg, 1879, p. 153–154, orig. descr. (subgenus Diplodus); 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. and fig.; Zikan and Wygodzinsky, 1948, p. 17, list; Wygodzinsky, 1949a, p. 48, checklist; Wygodzinsky, 1949b, p. 336, note; wygodzinsky, 1957, p. 264, 268, list and junior syn. of Z. obscuridorsis; Hart, 1987, p. 297, junior syn. of Z. illotus Berg.
Zelus illotus Berg, 1879, habitus
Zelus illotus Berg, 1879, male genitalic structures
Male: (
Female: (
Among species of the Zelus nugax species group (
South America and adjacent islands of the Caribbean (
Zelus impar Kuhlgatz, 1902, p. 264–266, Tab. IV, fig. 6, 6a, 6b, orig. descr. and fig.; Wygodzinsky, 1949a, p. 49, checklist; Hart, 1987, p. 297, redescription, note, fig., key and neotype desig.; Maldonado, 1990, p. 327, cat.
Zelus impar Kuhlgatz & Melichar, 1902, habitus
Zelus impar Kuhlgatz & Melichar, 1902, male genitalic structures
Male: (
Female: Unknown.
Recognized by the slender, curved, laterally compressed, and apically tapered medial process (shared with members of the Zelus nugax species group;
Panama and Northern South America and adjacent islands of the Caribbean (
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.
Zelus inconstans Champion, 1898, p. 254–255, Tab. XV., fig. 13, orig. descr. and fig.; Wygodzinsky, 1947, p. 43, note; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 328, cat.
Zelus inconstans Champion, 1899, habitus
Zelus inconstans Champion, 1899, male genitalic structures
Male: (
Female: (
The humeral angle rounded and the pronotum lacking conspicuous spine-like setae are diagnostic of this species. Most other species with unarmed humeral angle also have 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 (
Southern Central America and northern South America (
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.
Zelus janus Stål, 1862, p. 452, orig. descr.; Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, (in part), p. 257–258, Tab. XV. fig. 19, note, fig. and senior syn. of Z. litigiosus; Fracker, 1913, p. 239, 240, key and list (subgenus Diplodus); Fracker and Bruner, 1924, p. 170–171, note; Wygodzinsky, 1949a, p. 49, checklist; Hart, 1986, p. 543, redescription, lectotype desig., note, key and fig.; Maldonado, 1990, p. 328, cat.
Diplodus janus: Uhler, 1886, p. 24, checklist; Walker, 1873, p. 124, cat.
Zelus janus Stål, 1862, habitus
Male: (
Female: (
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.
Southern Texas to Central America (
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.
Zelus kartabensis Haviland, 1931: 137, 148, 152 (checklist, fig. and orig. descr. [subgenus Diplodus]); Wygodzinsky, 1949a: 49 (checklist); Maldonado, 1990, p. 328, cat.
Zelus pallidinervus Haviland, 1931: 137, 148, 153 (checklist, fig and orig. descr. [subgenus Diplodus]); Wygodzinsky, 1949a: 50 (checklist); Maldonado, 1990, p. 330, cat. syn. nov. (current study).
Zelus kartabensis Haviland, 1931, habitus
Zelus kartabensis Haviland, 1931, male genitalic structures
Male: (
Female: (
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.
South America (
Zelus kartaboides Zhang & Hart, sp. n., habitus
Zelus kartaboides Zhang & Hart, sp. n., male, genitalic structures
Male: (
Female: Unknown.
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.
The specific epithet indicates that this species is rather similar to Z. kartabensis.
South America (
Zelus korystos Hart, 1986, p. 303-304, figs. 25–27, orig. descri., note, fig. and key; Maldonado, 1990, p. 328, cat.
Zelus korystos Hart, 1986, habitus
Zelus korystos Hart, 1986, male genitalic structures
Male: (
Female: Unknown.
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.
South America and adjacent islands of the Caribbean (
Euagoras laticornis Herrich-Schaeffer, 1853, p. 123, Tab. CCCIX. Fig. C, orig. descr. and fig.
Zelus laticornis: Stål, 1872, p. 92, cat.; Lethierry and Severin, 1896, p. 152, cat.; Wygodzinsky, l949a, p. 49, checklist; Maldonado, 1990, p. 328, cat.
Darbanus laticornis: Walker, 1873, p. 127, cat.
Zelus formosus Haviland, 1931, p. 137, 151–152, list and orig. descr. (subgenus Diplodus); Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 327, cat. syn. nov. (current study).
Zelus tristis Haviland, 1931, p. 137, 154, list and orig. descr. (subgenus Diplodus); Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 331, cat. syn. nov. (current study).
Zelus laticornis (Herrich-Schaeffer, 1853), habitus
Zelus laticornis (Herrich-Schaeffer, 1853), male genitalic structures
Male: (
Female: (
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.
Southern Central America (Panama) and South America (
The type material of Z. laticornis (under the name Euagoras laticornis) was destroyed during World War II. The female holotype of Zelus formosus Haviland, 1931 is deposited in the Natural History Museum, London.
Reduvius leucogrammus Perty, 1834, p. 174, pl. 34, fig. 14, orig. descr. and fig.
Zelus leucogrammus: Stål, 1872, p. 90, cat. (subgenus Diplodus); Berg, 1879, p. 152–153, cat., descr. and nymph (subgenus Diplodus); Lethierry and Severin, 1896, p. 152, cat.; Costa Lima, 1940, p. 7, 218, 224 illus., biol. notes (subgenus Diplocodus); Wygodzinsky, 1949a, p. 49, checklist; Wygodzinsky, 1957, p. 264, list (Z. leucogrammus (sic.)); Wygodzinsky, 1960, p. 307, locality; Maldonado, 1990, p. 328, cat.
Zelus leucogrammus (Perty, 1833), habitus
Zelus leucogrammus (Perty, 1833), male genitalic structures
Male: (
Female: (
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.
South America (
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.
Zelus lewisi Zhang & Hart, sp. n., habitus
Male: (
Female: (
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 (
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.
Central America (
Zelus litigiosus Stål, 1862, p. 453, orig. descr.; Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 257, Tab. XV. fig. 20, 20a, fig. and junior syn. of Z. janus; Maldonado, 1990, p. 328, cat.
Diplodus litigiosus: Walker, 1873, p. 124, cat; Uhler, 1886, p. 24, checklist.
Zelus litigiosus Stål, 1872, habitus
Zelus litigiosus Stål, 1872, male genitalic structures
Male: (
Female: (
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.
Southwestern Mexico (
Cimex longipes Linnaeus, 1767, p. 724, orig. descr.; Gmelin, 1788, p. 2197, list (Reduvius); Turton, 1806, p. 690, descr.
Reduvius longipes: Fabricius, 1775, p. 730, descr.; Fabricius, 1781, p. 378, descr.; Fabricius, 1787, p. 309, list; Fabricius, 1794, p. 196, descr.
Zelus longipes: Fabricius, 1803, p. 283, descr.; Stål, 1872, p. 88-89, cat. (subgenus Zelus); Blanchard, 1840, p. 101, descr.; Stål, 1862, p. 449-450, descr.; Uhler, 1878, p. 427, list; Uhler, 1886, p, 24, checklist; Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 253, note; Kirkaldy, 1900a, p. 263, note; Fracker, 1913, p. 239, 240, key and list (subgenus Zelus); Cotton, 1917, p. 170-173, note; Barber, 1923, p. 27-28, note and syn.; Wygodzinsky, 1949a, p. 49, checklist; Wolcott, 1950 (1948), p. 212, list and note; Elkins, 1951, p. 410, list; Guagliumi, 1953, p. 16, note; Barber, 1954, p. 13-14, list; Elkins, 1954, p. 44, 45, note and fig.; Simmonds, 1956, p. 232, note; Alayo, 1967, p. 5, 36-37, list and note; Hart, 1986, p. 543-546, redescription, note, fig. and key; Hart, 1987, p. 304, note and key; Maldonado, 1990, p. 328, cat.
Euagoras longipes: Walker, 1873, p. 117-118, cat.
Reduvius rubidus Lepeletier and Serville, 1825, p. 278, orig. descr,; Guerin-Meneville, 1857, p. 411-412, descr. and list (subgenus Evagoras).
Evagoras rubidus: Amyot and Serville, 1843, p. 368-369, descr. and senior syn. of Evagoras speciosus Burmeister. Stål, 1862, p. 449, junior syn. (in part) of Z. longipes. Walker, 1873, p. 117, junior syn. of Euagoras longipes.
Euagoras rubidus: Walker, 1873, p. 118, cat.
Zelus rubidus: Stål, 1872, p. 89, cat. and descr. (subgenus Zelus); Uhler, 1886, p. 24, checklist; Lethierry and Severin, 1896, p. 153, cat.; Champion, 1898, p. 252-253, cat. and note; Fracker, 1913, p. 238, 240, key and list (subgenus Zelus); Ballou, 1913, p. 65, note; Jones, 1914, p. 462, note; Osborne and Drake, 1915, p. 531, list; Cotton, 1917, p. 173, note; Ritchie, 1917, p. 94, note; Gibson, 1919, p. 276, list; Dash, 1920, p. 31, note; Barber, 1923, p. 27, junior syn. of Z. longipes; Bruner, 1926, p. 78, descr.; Gowdey, 1927, p. 16-17, note; Martorell, 1939, p. 189, list; Wygodzinsky, 1949a, p. 49, checklist and junior syn. of Z. longipes; Alayo, 1967, p. 36-37, note.
Reduvius phalangium Fabricius, 1794, p. 1966, orig. descr.; Zirnsen, 1964, p. 338, list; Hart, 1986, p. 543, junior syn. of Z. longipes.
Zelus phalangium: Fabricius, 1803, p. 283, descr.; Stål, 1872, p. 92, cat.; Lethierry and Severin, 1896, p. 153, cat.; Fracker, 1913, p. 240, descr. and list; Wygodzinsky, 1949a, p. 50, checklist.
Diplodus phalangium: Uhler, 1886, p. 24, checklist.
Zelus bilobus Say, 1832, p. 12, orig. descr.; LeConte, 1859, p. 306, descr.; Stål, 1862, p. 449, list (as variety of Z. longipes); Stål, 1872, p. 88, cat. (subgenus Zelus); Uhler,1876, p. 61, list; Uhler, 1886, p. 24, checklist; Lethierry and Severin, 1896, p. 151, cat.; Champion, 1898, p. 253, note; Van Duzee, 1909, p. 176, list; Torre-Bueno and Engelhardt, 1910, p. 150, list; Fracker, 1913, p. 239, 240, key and list (subgenus Zelus); Barber, 1914, p. 505, list; Van Duzee, 1916, p. 30, checklist (s .g. Zelus); Dozier, 1917, p. 542, note; Van Duzee, 1917, p. 259, cat. (subgenus Zelus); Dozier, 1920, p. 357, note; Blatchley, 1926, p. 568, 569, key and descr. (subgenus Zelus); Readio, 1927, p. 169-170, key, descr. and note; Miller, 1929, p. 462, note; Creighton 1936a p. 94, note; Creighton, 1936b, p. 382, note; Elliott , 1938, p. 39, key and list; Wygodzinsky, 1949a, p. 48, checklist; Elkins, 1951, p. 410, list; Sibley, 1951., p. 92, list ; Oliver, 1964, p. 316, note; Whitcomb and Bell , 1964, p. 22, list; Davis, 1969, p. 81, fig. and note (sic. Zellus bilobatus); Hart, 1986, p. 543, junior syn. of Z. longipes.
Euagoras speciosus Burmeister, 1835, p. 227, orig. descr.; Herrich-Schaeffer, 1848, p. 45, Tab. CCLXIV. fig. 817, descr. and fig; Hart, 1986, p. 543, junior syn. of Z. longipes.
Evagoras speciosus: Amyot and Serville, 1843, p. 368, junior syn. of Evagoras rubidus Le P. and Serv.
Zelus speciosus: Stål, 1862, p. 449, syn. (as variety of Z. longipes) Stål, 1872, p. 89, cat. (subgenus Zelus); Berg, 1879, p. 151, note; Uhler, 1886, p. 24, checklist; Lethierry and Severin, 1896, p. 153, cat.;, Kirkaldy, 1909, p. 32, list and syn.; Wygodzinsky, 1949a, p. 50, checklist.
Zelus speciosus var. stolli Lethierry and Severin, 1896, p. 152, nomen nudum; Champion, 1898, p. 253, syn. (=Zelus rubidus). Euagoras tricolor Herrich-Schaeffer, 1848, p. 45-46, Tab. CCLXIV, fig. 818, orig. descr. and fig.; Stål, 1862, p. 450, syn. (as variety of Zelus longipes); Stål, 1872, p. 89, syn. (as variety of Zelus speciosus); Champion, 1898, p. 253, junior syn. of Zelus rubidus; Fracker and Bruner, 1924, p. 170, list; Bruner, 1926, p. 79, descr; Wygodzinsky, 1949a, p. 50, junior syn. of Zelus speciosus.
Zelus mactans Stål, 1861, p. 148, orig. descr.; Stål, 1872, p. 88, cat. (subgenus Zelus); Uhler, 1886, p. 24, checklist; Lethierry and Severin, 1896, p. 152, cat.; Fracker, 1913, p. 239, 240, key and list (subgenus Zelus); Barber, 1923, p. 28, note; Wygodzinsky, 1949a, p. 49, checklist; Alayo, 1967, p. 36, key and note; Hart, 1986, p. 544, lectotype desig. and junior syn. of Z. longipes.
Diplodus mactans: Walker, 1873, p. 125, cat.
Velia agavis Blasguez, 1870, p. 289, 290, fig. 14, orig. descr.; Champion, 1898, p. 253, junior syn. of Z. rubidus; Kirkaldy, 1909, p. 32, junior syn. (as variety of Z. speciosus); Fracker, 1913, p. 240, junior syn. of Z. rubidus.
Zelus longipes (L., 1767), habitus
Zelus longipes (L., 1767), male genitalic structures
Male: (
Female: (
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 (
Southern parts of US, Mexico, Central America, the Caribbean, Northern South America, Paraguay and Southern Brazil (
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 luridus Stål, 1862, p. 452, orig. descr.; Stål, 1872, p. 91, cat. (subgenus Diplodus) Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 259–260, junior syn. of Z. exsanguis; Uhler, 1904, p. 364, list; Wirtner, 1904, p. 206, list; Torre-Bueno and Brimley, 1907, p. 437, list; Torre-Bueno, 1910, p. 32, note; Torre-Bueno and Engelhardt, 1910, p. 150, note; Torre-Bueno, 1913, p. 60, note (subgenus Diplodus); Barger, 1914, p. 506, list; Van Duzee, 1916, p. 30, junior syn. of Z. exsanguis; Hart, 1986, p. 537, redescription, lectotype desig, note, key, fig. and stat. rev.; Maldonado, 1990, p. 329, cat.
Diplodus luridus: Uhler, 1872a, p. 471, checklist; Uhler, 187213, p. 420, note; Walker, 1873, p. 124, cat.; Uhler, 1876, p. 61, note; Uhler, 1877, p. 429, note; Uhler, 1878, p. 427, note; Uhler, 1886, p. 24, checklist; Provancher, 1887, p. 181, note; Van Duzee, 1894, p. 183, list; Gillette and Baker, 1895, p. 60, list.
Diplocodus luridus: Van Duzee, 1912, p. 324, note.
Darbanus georgiae Provancher, 1872, p. 106, orig. descr.; Uhler, 1886, p. 24, checklist; Provancher, 1887, p. 181, note; Kelton, 1968, p. 1070, note; Hart, 1986, p. 53, junior syn. of Z. luridus.
Zelus georgiae: Lethierry and Severin, 1896, p. 152, cat.; Banks, 1910, p. 16, cat.; Van Duzee, 1916, p. 30, junior syn. of Z. exsanguis.
Zelus acanthogonius Say & Uhler, 1878, p. 427, manuscript name.
Evagoras viridis Uhler, 1878, p. 427, manuscript name
Darbanus palliatus Provancher, 1887, p. 182, orig. descr.; Kelton, 1968, 1070, note; Hart, 1986, p. 53, junior syn. of Z. luridus.
Zelus palliatus: Lethierry and Severin, 1896, p. 153, cat.; Banks, 1910, p. 16, cat.; Van Duzee, 1916, p. 30, junior syn. of Z. exsanguis.
Reduvius sp. Emmons, 1854, p. 168, P1. 7, fig. 3, note and fig.
Zelus exsanguis [misapplication of name due to Champion's synonymy of Z. luridus under Z. exsanguis]: Wirtner, 1904, p. 206, list; Barber, 1906, p. 285, list; Snow, 1907, p. 159, list; Parshley, 1914, p. 144, list; Hussey, 1922, p. 24, list; Britton, 1923, p. 687, list; Blatchley, 1926, p. 570–571, descr. and note (Diplodus); Readio, 1926, 167, P1. X, fig. 5, 6 and 7, note and fig.; Readio, 1927, p. 169, 171–177, P1. XIII, descr., notes and fig.; Leonard, 1928, p. 105, list; Brimley, 1938, p. 73, list; Harris, 1943, p. 151, list; Procter, 1946, p. 319, list; Elkins, 1951, p. 410, list; Sibley, 1951, p. 92, list; Hall, Downe, MacLellan and West, 1953; p. 199–204, note; Dome and West, 1954, p. 181–184, behavior; West and De Long, 1955, p. 97–101, biology; Davis, 1961, p. 351, note; Drew and Schaeffer, 1962, p. 106, list; Whitcomb and Bell, 1964, p. 22, list; Kelton, 1968, p. 1070– 1071, note; Yonke and Medler, 1970, p. 441–443, biology.
Zelus luridus Stål, 1862, habitus
Zelus luridus Stål, 1862, male genitalic structures
Male: (
Female: (
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 (
North America (
This is one of the most commonly collected species in this genus.
Zelus mattogrossensis Wygodzinsky, 1947, p. 431–434, orig. descr. and fig.; Wygodzinsky, 1948, p. 17, cat.; Wygodzinsky, 1949a, p. 49, checklist; Wygodzinsky, 1960, p. 308, note; Maldonado, 1990, p. 329, cat.
Zelus mattogrossensis Wygodzinsky, 1947, male genitalic structures
Male: (
Female: (
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 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.
South America (
Zelus means Fabricius, 1803, p. 282. orig. descr.; Stål, 1868, p. 107, descr.; Stål, 1872, p. 89, cat.; Walker, 1873, p. 134, cat; Lethierry and Severin, 1896, p.l62., cat.; Champion, 1898, p. 254, note; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 329, cat.
Eccagoras trimaculicollis Stål 1855, p. 189, orig. descr.
Zelus trimaculicollis Stål, 1866, p. 298, descr.; Stål, 1872, p. 89, cat.; Lethierry and Severin, 1896, p. 153, cat.; Champion, 1898, p. 254, note; Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 331, cat. syn. nov. (current study).
Zelus trimaculatus Champion, 1898, p. 254, orig. descr.; Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 331, cat. syn. nov. (current study).
Zelus means Fabricius, 1803, habitus
Male: Unknown.
Female: (
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.
Southern Central America and South America (
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.
Zelus mimus Stål, 1862, p. 451, orig. descr. (subgenus Diplodus); Stål, 1872, p. 91, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 257, 261, note, list and senior syn. of Z. umbratilis; Kuhlgatz, 1902, p. 266, note; Fracker, 1913, p. 239, 240, list (subgenus Diplodus); Williams, 1918, p. 163–173; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 329, cat.
Diplodus mimus: Walker, 1873, p. 124, cat.; Uhler, 1886, p. 24, checklist.
Zelus umbratilis Stål, 1862, p. 451, orig. descr. (subgenus Diplodus); Stål, 1872, p. 91, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 153, cat.; Champion, 1898, p. 261, junior syn. of Z. mimus.
Diplodus umbratilis: Walker, 1873, p. 124, cat.; Uhler, 1886, p. 24, checklist.
Zelus mimus Stål, 1862, habitus
Zelus mimus Stål, 1862, male genitalic structures
Male: (
Female: (
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 (
Southern Mexico and Central America (
Zelus minutus Hart, 1987, p. 299-301, figs. 16-21, orig. descr., note, fig. and key; Maldonado, 1990, p. 330, cat.
Zelus minutus Hart, 1987, habitus
Male: (
Female: Similar to male, except for the following. Larger than male, total length 10.70 mm (
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.
Southern Central America, South America and adjacent islands of the Caribbean (
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.
Zelus nigromaculatus Champion, 1898, p. 261–262, Tab. XV, Fig. 26, orig. descr. and fig.; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 330, cat.
Zelus nigromaculatus Champion, 1899, habitus
Zelus nigromaculatus Champion, 1899, male, genitalic structures
Male: (
Female: (
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 (
Central America (
Zelus nugax Stål, 1862, p. 450–451, orig. descr. (subgenus Diplodus); Stål, 1872, p. 91, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 152, cat.; Champion, 1898, p. 257, 2, note and list; Kuhlgatz, 1902,, p. 266, note; Fracker, 1913, p. 239, 240, key and list (subgenus Diplodus); Fracker and Bruner, 1924, p. 171, list; Haviland, 1931, p. 137, 152, list and note; Leonard and Mills, 1931, p. 309–323, note; Wygodzinsky, 1947, p. 431, note; Wygodzinsky, 1949a, p. 49, checklist; Guagliumi, 1953, p. 16, note; Wygodzinsky, 1957, p. 268, junior syn. of Zelus obscuridorsis; Sucre, et. al., 1966, p. 31, note; Hart, 1986, p. 540, redescription, note, fig., key and lectotype desig.; Maldonado, 1990, p. 330, cat.
Diplodus nugax: Walker, 1873, p. 124, cat; Uhler, 1886, p. 24, checklist; Hart, 1986, p. 540, lectotype desig.
Zelus rufigeniculatus Haviland, 1931, p. 137, 148, 153–154, list, fig. and orig. descr. (subgenus Diplodus); Wygodzinsky, 1949a, p. 50, checklist; Hart, 1986, p. 540, junior syn. of Z. nugax.
Zelus nugax Stål, 1862, habitus
Zelus nugax Champion, 1899, male genitalic structures
Male: (
Female: (
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).
Mexico, Central America and South America (
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 (
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.
Zelus panamensis Zhang & Hart, sp. n., habitus
Zelus panamensis Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
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.
Named after the country Panama, where the holotype was collected.
Southern Central America and northern South America (
Zelus paracephalus Zhang & Hart, sp. n., habitus
Zelus paracephalus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
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 (
South America (
Zelus pedestris Fabricius, 1803, p. 288, orig. descr.; Stål, 1872, p. 91, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 330, cat.
Diplodus pedestris: Stål, 1868, p. 109, descr. and note; Walker, 1873, p. 125, cat.
Zelus dispar Fabricius, 1803, p. 291, orig. descr.; Stål, 1872, p. 92, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 151, cat.; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 326, cat. syn. nov. (current study).
Diplodus dispar: Stål, 1868, p. 109–110, descr. and note; Walker, 1873, p. 125, cat.
Diplodus obscuridorsis Stål, 1860, p. 75, orig. descr.; Stål, 1868, p. 109, note; Walker, 1873, p. 125, cat.
Zelus obscuridorsis: Stål, 1872, p. 91, cat. and descr. (subgenus Diplodus); kthierry and Severin, 1896, p. 152, cat.; Wygodzinsky, 1949a, p. 50, checklist; Wygodzinsky, 1957, p. 26, note and senior syn. of Z. nugax, Z. illotus and Z. carvalhoi; Wygodzinsky, 1960, p. 307–308, list. syn. nov. (current study).
Zelus pedestris Fabricius, 1803, habitus
Zelus pedestris Fabricius, 1803, male genitalic structures
Male: (
Female: (
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.
South America and adjacent islands of the Caribbean (
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.
Diplodus plagiatus Signoret, 1862, p. 584–585, orig. descr.; Walker, 1873, p. 126, cat.
Zelus plagiatus: Stål, 1872, p. 92, cat.; Lethierry and Severin, 1896, p. 153, cat.; Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 330, cat.
Zelus plagiatus (Signoret, 1852), habitus
Male: unknown.
Female: (
The dorsum predominantly yellow with the posterior pronotal lobe partly black and two black spots on hemelytra is distinctive of this species.
South America (
Euagoras prolixus Stål, 1860, p. 74, orig. descr.; Walker, 1873, p. 118, cat.
Zelus prolixus: Stål, 1872, p. 89, cat. (subgenus Zelus); Lethierry and Severin, 1896, p. 153, cat.; Champion, 1898, p. 255, note; Wygodzinsky, 1949a; p. 50, checklist; Elkins, 1954, p. 39, 40, note; Hart, 1987, p. 297-299, redescription, note, fig. and key; Maldonado, 1990, p. 330, cat.
Zelus prolixus (Stål, 1860), habitus
Zelus prolixus (Stål, 1860), male genitalic structures
Male: (
Female: (
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 dorsoventrally flattened; and the eye somewhat elongated. Males can be separated from most species of Zelus by the broadly triangular medial process. The short paramere not exceeding the medial process separates Z. prolixus from Z. minutus.
South America and adjacent islands of the Caribbean (
Zelus puertoricensis Hart, 1987, p. 294, orig. descr., key and fig.; Maldonado, 1990, p. 330, cat.
Zelus puertoricensis Hart, 1987, habitus
Zelus puertoricensis Hart, 1987, male genitalic structures
Male: (
Female: (
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 (
The Caribbean, islands of Puerto Rico and Hispaniola (
Zelus renardii Kolenati, 1857, p. 460, Tab. III. fig. 2, orig. descr. and fig.; Stål, 1872, p. 91, cat.; Kirkaldy, 1908, p. 195, list and senior syn. of Z. laevicollis Champion and Z. perigrinus Kirkaldy; Banks, 1910b, p. 16, cat.; Fracker, 1913, p. 239, 240, key and checklist; Van Duzee, 1914, p. 13, list; Van Duzee, 1916, p. 30, checklist (subgenus Diplocodus); Van Duzee, 1917, p. 261, cat. (subgenus Diplocodus); Muir, 1920, p. 285–286, note; Muir, 1921, p. 119, note; Horton, 1922, p. 385, note; Hawaiian Sugar Planter's Association, 1924, p, 29, note; Readio, 1927, p, 169, 178–179, key, descr. and note; Williams, 1931, p. 101, note; Haldaway and Look, 1942, p. 257–258, note; Ewing and Ivy, 1943, p, 604–606, note; Clancy, 1946, p. 326, note; Van Zwaluwenburg, 1946, p. 15, note; Zimmerman, 1948, p. 137–138 , note and fig.; Wygodzinsky, 1949a, p. 50, checklist; Elkins, 1951, p. 410, list; Sibley, 1951, p. 92, list; Nishida, 1955, p. 172, note; Atkins, et. al., 1957, p. 258, note; Nielsen and Henderson, 1959, p. 159, note; Wene and Sheets, 1962, p. 397, note; Butler, 1966, p. 1306–1307, note; Wygodzinsky, 1966, p. 66, note; Lingren, Ridgway and Jones, 1968, p. 615, note; Parencia, 1968, p. 276, note; Nutting and Spangler, 1969, p. 765, note; Hart, 1986, p. 540-542, redescription, note, fig. and key; Maldonado, 1990, p. 331, cat.
Diplodus renardii: Uhler, 1894, list.
Zelus laevicollis Champion, 1898, p. 260–261, Tab. XV. fig. 24, orig. descr. and fig.; Banks, 1910, p. 16, cat.; Fracker, 1913, p. 239, 240, key and list; Osborne and Drake, 1915, p. 531, note; Van Duzee, 1916, p. 30, checklist (subgenus Diplocodus); Van Duzee, 1917, p. 261, cat. (subgenus Diplocodus); Readio, 1927, p. 177–178, descr.; Wygodzinsky, 1949a, p. 49, checklist; Elkins, 1951, p. 410, list; Sibley, 1951, p. 92, list; Young and Sifuentes, 1960, p. 1109–1111, biology; Hart, 1986, p. 540, junior syn. of Z. renardii.
Zelus peregrinus Kirkaldy, 1902, p. 149–150, orig. descr.; Swezey, 1905, p. 232–234, biology; Kirkaldy, 1907a, p. 247, note; Kirkaldy, 1907b, p. 156–518, biology; Kirkaldy, 1908, p. 195, junior syn. of Z. renardii; Severin, et. al., 1914, p. 197, note; Fullaway, 1918, p. 12, note; Clausen, 1940, p. 589–590, note (sic. Zellus).
Diplocodus exsanguis: Van Duzee, 1914, p. 13, list (probable misidentification).
Zelus renardii Kolenati, 1857, habitus
Zelus renardii Kolenati, 1857, male genitalic structures
Zelus renardii Kolenati, 1857, specimen record map. Pacific islands (left) and American continent are not to the same scale.
Male: (
Female: (
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.
Western and Southwestern US, most of Mexico and Central America (
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.
Zelus rosulentus Zhang & Hart, sp. n., habitus
Zelus rosulentus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: unknown.
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 (
The specific epithet indicates the reddish-pink coloration of this species.
South America (
Zelus ruficeps Stål, 1862, p. 453–454, orig. descr.; Stål, 1872, p. 90, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 153, cat.; Champion, 1898, p. 256, Tab. 15. fig. 15, note and fig.; Kuhlgatz, 1902, p. 266, note; Fracker, 1913, p. 239, 240, list (subgenus Diplodus); Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 331, cat.
Diplodus ruficeps: Walker, 1873, p. 124, cat.; Uhler, 1886, p. 24, checklist.
Zelus ruficeps Stål, 1862, habitus
Zelus ruficeps Stål, 1862, male genitalic structures
Male: (
Female: (
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.
Southern Mexico to Northern South America (
Zelus russulumus Zhang & Hart, sp. n., habitus
Zelus russulumus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
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.
The species epithet refers to the reddish-brown area on the membrane.
South America (
Zelus spatulosus Zhang & Hart, sp. n., habitus
Zelus spatulosus Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: unknown.
Zelus spatulosus has a rather distinctly enlarged apical part of the paramere (
The specific epithet is from spatula, referring to the rather broad apical part of the paramere.
Central America (
The primary basis for placing Z. spatulosus in the Zelus luridus group is the expanded paramere (
Zelus sphegeus Fabricius, 1803, p. 287, orig. descr.; Stål, 1872, p. 91, cat.; Lethierry and Severin, 1896, p. 153, cat.; Haviland, 1931, p. 137, 153, list and note; Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 331, cat.
Diplodus sphegeus, Stål, 1868, p. 109, descr.; Walker, 1873, p. 125, cat.
Zelus sphegeus Fabricius, 1803, habitus
Male: unknown.
Female: (
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.
Central America and South America (
Zelus subimpressus Stål, 1872, p. 91, orig. descr. (subgenus Diplodus); Lethierry and Severin, 1896, p. 153, cat.; Fracker, 1913, p. 239, 240, key and list (subgenus Diplodus); Fracker and Bruner, 1924, p. 170, note; Bruner 1926, p. 78, 79, key and note; Leonard, 1933, p. 319, note; Wolcott, 1950, p. 212, note; Wygodzinsky, 1949a, p. 50, checklist; Alayo, 1967, p. 37, note; Hart, 1987, p. 294, redescription, note, fig. and key; Maldonado, 1990, p. 331, cat.
Diplodus subimpressus: Uhler, 1886, p. 24, checklist; Gundlach, 1894, p. 598, checklist.
Zelus subimpressus Stål, 1872, habitus
Zelus subimpressus Stål, 1872, male genitalic structures
Male: (
Female: (
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 (
The Caribbean, the islands of Cuba and Hispaniola (
Zelus sulcicollis Champion, 1898, p. 258–259, Tab. XV. fig. 21, orig. descr. and fig.; Fracker, 1913, p. 239, 240, key and list (subgenus Diplodus); Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 331, cat.
Zelus sulcicollis Champion, 1899, habitus
Zelus sulcicollis Champion, 1899, male genitalic structures
Male: (
Female: (
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.
Southern Mexico and Northern Central America (
Zelus tetracanthus Stål , 1862, p. 454, orig. descr. (subgenus Pindus) ; Stål , 1872, p. 92, cat. (subgenus Pindus); Lethierry and Severin, 1896, p. 153, cat.; Champion, 1898, p. 362, Tab. XV, fig. 27, note and fig.; Fracker, 1913, p. 240, key and list (subgenus Pindus); Wygodzinsky, 1949a, p. 50, checklist; Hart, 1986, p. 536-537, redescription, note, fig. and key; Maldonado, 1990, p. 331, cat.
Diplodus tetracanthus: Walker, 1873, p. 124, cat.; Uhler, 1886, p. 24, checklist.
Pindus socius Uhler, 1872b, p. 420, orig. descr.; Uhler, 186, p. 62, list (reprint); Uhler, 1877, p. 1329, list; Uhler, 1894, p. 284, list.; Hart, 1986, p. 536, junior syn. of Z. tetracanthus.
Diplodus socius: Uhler, 1886, p. 24, checklist; Gillette and Baker, 1895, p. 60, list.
Zelus socius: Lethierry and Severin, 1896, p. 153 cat.; Barber, 1906, p. 28,6, list; Hart, 1907, p. 237, list; Torre-Bueno, 1908, p. 235, list (subgenus Pindus); Van Duzee, 1909, p. 177, list; Banks, 1910b, p. 16, cat.; Fracker, 1913, p. 240, key and list (subgenus Pindus); Torre-Bueno, 1913, p. 60, list (subgenus Pindus); Barber, 1914, p. 506, list; Parshley, 1914, p. 144, list; Van duzee, 1916, p. 30, checklist (subgenus Pindus); Van Duzee, 1917, p. 261, cat. (subgenus Pindus); Parshley, 1921, p. 5, list; Anonymous, 1923, p. 120-135, note; Blatchley, 1926, p. 569, 571, key and descr. (subgenus Pindus); Readio, 1926, p. 168, 177, note and fig.; Readio, 1927, p. 169, 179-181, P1. XIV. fig. 1,2,5, descr., notes and fig.; Leonard, 1928, p. 105, list; Essig, 1929, p. 357, note; Knowlton, 1932, p. 12, note; Harris, 1937, p. 174, list; Brimley, 1938, p. 73, list; Procter, 1946, p. 319, list; Wygodzinsky, 1949a, p. 50, checklist; Elkins, 1951, p. 410, list; Sibley, 1951, p. 92, list; Elkins, 1954, p. 47, note; Atkins, et. al., 1957, p. 251-259, note; Drew and Schaeffer, 1962, p. 106, list; Wene and Sheets, 1962, p. 395-398, note; Whitcomb and Bell, 1964, p. 22, list; Butler, 1966, p. 1306-1307, note; Nutting and Spangler, 1969, p. 763-769, note and photo.
Diplocodus socius: Van Duzee, 1914, p. 13, list and note.
Zelus audax Banks, 1910, p. 325, orig. descr.; Fracker, 1913, p. 240, note; Van Duzee, 1916, p. 30 checklist (subgenus Pindus); Van Duzee, 1917, p. 261, cat. (subgenus Pindus); Britton, 1923, p. 687, list (subgenus Pindus); Blatchley, 1926, p. 569, 571-572, key and descr. (subgenus Pindus); Readio, 1927, p. 169, 181, descr.; Leonard, 1928, p. 105, list; Wygodzinsky, 1949a, p. 48, checklist; Hart, 1986, p. 536, junior syn. of Z. tetracanthus and lectotype desig.
Zelus occiduus Torre-Bueno, 1913a, p. 22, orig. descr. (subgenus Pindus); Van Duzee, 1916, p. 30, checklist; Van Duzee, 1917, p. 261, cat. (subgenus Pindus) ; Readio, 1927, p. 169, 181-182, key and descr.; Wygodzinsky, 1949a, p. 50, checklist; Elkins, 1951, p. 410, list; Hart, 1986, p. 536, junior syn. of Z. tetracanthus and lectotype desig.
Zelus angustatus Hussey, 1925, p. 66-67, orig. descr.; Blatchley, 1926, p. 569, 572, descr. and note (subgenus Pindus); Readio, 1927, p. 169, 182., key and descr.; Blatchley, 1928, p. 6, note (subgenus Pindus); Wygodzinsky, 1949a, p. 48, checklist; Elkins, 1951, p. 410, list; Hussey, 1953, p. 9-11, note; Hart, 1986, p. 536, junior syn. of Z. tetracanthus.
Zelus tetracanthus Stål, 1862, habitus
Zelus tetracanthus Stål, 1862, male genitalic structures
Male: (
Female: (
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.
North America, Central America and parts of South America (
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). Further investigations should examine if the South Amerian populations are native or introduced.
Zelus truxali Zhang & Hart, sp. n., habitus
Zelus truxali Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: (
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.
Named after F. S. Truxal, the collector of the type specimen.
South America (
Zelus umbraculoides Zhang & Hart, sp. n., habitus
Zelus umbraculoides Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Unknown.
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.
The specific epithet indicates its close resemblance to Z. umbraculus, another new species described in the current study.
South America (
Zelus umbraculus Zhang & Hart, sp. n., habitus
Zelus umbraculus, Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Unknown.
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 (
The specific epithet is from Latin "umbra", referring to the shadow in the anterior part of the membrane.
South America (
Zelus vagans Fabricius, 1803, p. 284, orig. descr.; Stål, 1868, p. 108, descr. and note; Stål, 1872, p. 88, cat.; Walker, 1873, p. 134, cat.; Lethierry and Severin, 1896, p. 153, cat.; Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 332, cat.
Zelus vagans Fabricius, 1803, habitus
Zelus vagans Fabricius, 1803, male genitalic structures
Male: (
Female: Unknown.
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.
South America (
Euagoras varius Herrich-Schaeffer, 1853, p. 122, orig. descr.
Zelus varius: Stål , 1872, p. 92, cat . (subgenus Diplodus); Lethierry and Severin, 1896, p. 153, cat.; Wygodzinsky 1949a, p. 50, checklist; Maldonado, 1990, p. 332, cat.
Diplodus varius: Walker, 1873, p. 126, cat.
Zelus varius (Herrich-Schaeffer, 1853), habitus
Zelus varius (Herrich-Schaeffer, 1853), male genitalic structures
Male: (
Female: Similar to male, except for the following. Larger than male, total length 13.71–14.03 mm (mean 13.87 mm,
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.
South America (
Euagoras versicolor Herrich-Schaeffer, 1848, p. 46-47, Tab. CCLXIV, fig. 820, orig. descr. and fig.; Herrich-Schaeffer, 1853, p. 92, cat.; Walker, 1873, p. 118, note.
Diplodus versicolor , Stål, 1860, p. 75, cat.
Zelus versicolor: Stål, 1862, p. 451, note; Stål, 1872, p. 92, cat. (subgenus Diplodus); Lethierry and Severin, 1896, p. 153, cat.; Wygodzinsky, 1949a, p. 50, checklist; Maldonado, 1990, p. 332, cat.
Euagoras nigrispinus Herrich-Schaeffer, 1848, p. 47-48, Tab. CCLXII, fig. 816, orig. descr. and fig.; Herrich-Schaeffer, 1853, p. 92, cat. Gil-Santana, 2008, p. 48, Figs 15-21, junior syn. of Zelus versicolor (Herrich-Schaeffer, 1848).
Diplodus nigrispinus: Stål, 1860, p. 75, cat.; Stål, 1868, p. 109, note; Walker, 1873, p. 126, cat.
Zelus nigrispinus: Mayr, 1866, p. 139, cat.; Stål, 1872, p. 91, cat. (subgenus Diplodus); Berg, 1879, p. 154; Lethierry and Severin, 1896, p. 152, cat.; Wygodzinsky, 1949a, p. 49, checklist; Maldonado, 1990, p. 330, cat; Gil-Santana, 2008, p. 48, junior syn. of Z. versicolor.
Zelus personatus Berg, 1879, p. 150-151, orig. descr. (subgenus Zelus); Lethierry and Severin, 1896, p. 153, cat.; Wygodzinsky, 1949a, p. 50, checklist; Wygodzinsky, 1957, p. 264, 268-269, list and note; Maldonado, 1990, p. 330, cat. syn. nov. (current study).
Zelus versicolor (Herrich-Schaeffer, 1848), habitus
Zelus versicolor (Herrich-Schaeffer, 1848), male genitalic structures
Male: (
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.
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.
South America (
The type specimens of Z. versicolor and Z. nigrispinus were destroyed during World War II.
Zelus vespiformis Hart, 1987, p. 301, figs. 22-27, orig. descr., note, fig. and key; Maldonado, 1990, p. 332, cat.
Zelus vespiformis Hart, 1987, habitus
Zelus vespiformis Hart, 1987, male genitalic structures
Male: (
Female: Similar to male, except for the following. Larger than male, total length 16.48–18.29 mm (mean 17.47 mm,
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.
Central America and Northern South America (
Zelus xouthos Zhang & Hart, sp. n., habitus
Zelus xouthos Zhang & Hart, sp. n., male genitalic structures
Male: (
Female: Similar to male, except for the following. Larger than male, total length 14.69 mm (n=1,
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.
From Greek xutho, meaning yellowish-brown, referring to the yellowish coloration.
Central America (
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.
Zelus zayasi Bruner and Barber, 1937, p. 186–188, orig. descr. and fig. (subgenus Zelus); Wygodzinsky, 1949a, p. 50, checklist; Zayas, 1960, p. 125, 126, note; Alayo, 1967, p. 36, 37, key and list.; Hart, 1987, p. 296, key and note; Maldonado, 1990, p. 332, cat.
Zelus zayasi Bruner and Barber, 1937, habitus
Male: unknown.
Female: (
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.
The Caribbean. Known only from Cuba (
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.
Key to species groups of Zelus (males only) |
||
1 | Apex of medial process without ventrally or posteriorly directed hook, projection or fold ( |
2 |
– | Apex of medial process with ventrally or posteriorly directed hook, projection or fold ( |
5 |
2 | Medial process triangular, broad at base, if slender, paramere apically enlarged; anteroposteriorly compressed ( |
3 |
– | Medial process with lateral margins subparallel or narrowly triangular; base not conspicuously broadened; laterally compressed or blade-like throughout or in apical 1/2 ( |
4 |
3 | Medial process basally nearly continuous with rest of pygophore, base not protruding posteriorly. Paramere apically not enlarged or slightly enlarged ( |
Zelus tetracanthus species group |
– | Medial process basally readily separable from rest of pygophore; base prominent and protruding posteriorly. Paramere expanded apically in some species ( |
Zelus luridus species group |
4 | Medial process slender, laterally compressed, dorsally oriented, at about forty-five degree angle to body axis (except in Z. grassans) ( |
Zelus nugax species group |
– | Medial process stout, posteriorly oriented, nearly parallel to body axis ( |
Zelus vagans species group |
5 | Apex of medial process strongly curved ventrally to form hook-like process or lip-like fold ( |
6 |
– | Apex of medial process not curved or weakly curved ventrally ( |
7 |
6 | Medial process laterally compressed; apex strongly curved ventrally, clearly hook-like. Paramere apically expanded, slightly curved dorsally ( |
Zelus renardii species group |
– | Medial process cylindrical or broad at base; not laterally compressed; apex curved, lip-like rather than hook-like ( |
Zelus mimus species group |
7 | Apex of medial process with distinct, acute, hook-like process or ridge-like elevation ( |
8 |
– | Apex of medial process with indistinct process or weak curvature, not forming sharp projection ( |
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 ( |
Zelus erythrocephalus species group |
– | Medial process long, slender, stalk-like; somewhat laterally compressed; lateral margins subparallel, base not broadened or slightly broadened; gradually curving and directed posteriorly in apical 1/3 or 1/2; apex with short, acute, hook-like process or sharp fold ( |
Zelus panamensis species group |
9 | Medial process extremely slender, usually long ( |
Zelus longipes species group |
– | Medial process stout, relatively short ( |
10 |
10 | Large (>13 mm), robust. Apex of medial process with pair of minute projections ( |
Zelus armillatus species group |
– | Medium-sized (<12 mm), elongate. Apex of medial process with inconspicuous folding ( |
Zelus puertoricensis species group |
Key to males of the Zelus tetracanthus species group |
||
1 | Posterior pronotal lobe disc bears pair of tubercles. | 2 |
– | Posterior pronotal lobe disc without tubercles. | 3 |
2 | Smallish, 7.8-10.1 mm. Legs delicate. Head and parts of pronotum orangish-brown. Paramere clearly exceeds medial process ( |
Zelus minutus ( |
– | Length 11.3-13.7 mm. Greyish-brown to brownish-black. Paramere removed from or reaching medial process, never exceeding ( |
Zelus tetracanthus ( |
3 | Entire surface reddish-brown ( |
Zelus rosulentus |
– | Pale brown ( |
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. | 2 |
– | Humeral angle clearly below level of, and not continuous with, disc. | 3 |
2 | Medial longitudinal sulcus of anterior pronotal lobe with dark brown area near posterior margin. Femora usually with dark apical bands. Parameres in fresh or relaxed specimens not achieving apex of medial process ( |
Zelus ambulans ( |
– | 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 ( |
Zelus exsanguis ( |
3 | Paramere spatulate apically, compressed; apical portion about 2x width of cylindrical basal portion ( |
Zelus spatulosus ( |
– | Paramere slightly expanded apically, apical portion less than 2x width of base. Medial process triangular, base broad and protruding posteriorly. | 4 |
4 | Eyes prominent, extending below ventral surface of head. |
Zelus grandoculus ( |
– | Eyes moderate, not extending below ventral surface of head. | 5 |
5 | Anterior pronotal lobe usually 1/2 or greater of length of posterior pronotal lobe; if slightly less than 1/2, profemoral length less than 20x profemoral diameter. Protrusion at base of medial process moderate ( |
Zelus luridus ( |
– | Anterior pronotal lobe less than 1/2 length of posterior pronotal lobe; profemoral length 20x or more profemoral diameter (C.A.). Protrusion at base of medial process more prominent ( |
Zelus antiguensis ( |
Key to males of the Zelus nugax species group |
||
1 | Paramere curved strongly medially and recurved apically ( |
Zelus grassans ( |
– | Paramere straight or only slightly curved. Medial process dorsally directed. Brown, abdomen without banding. | 2 |
2 | Medial process exhibiting pronounced curvature posteriorly about 1/3 of distance from base, then recurved toward dorsum about 3/4 distance from base. | 3 |
– | Medial process blade-like, straight; any curvature toward posterior weak. | 4 |
3 | Labial segment I and coxae medium brown to brownish-black. Paramere nearly straight, any curvature weak ( |
Zelus impar ( |
– | Labium and coxae light yellowish-brown. Paramere medially curved ventrally ( |
Zelus illotus ( |
4 | Basal plate arms fused. Pygophore as in |
Zelus pedestris ( |
– | Basal plate arms not fused, separate in specimens from S.A., tendency to fuse in C.A. and Mexico. Pygophore as in |
Zelus nugax ( |
Key to males of the Zelus vagans species group |
||
1 | Dorsal surface entirely black. Abdomen brown, yellow or red. | 2 |
– | Dorsal surface with orange and dark brown areas on pronotum or hemelytron. Abdomen typically orange or reddish with terminal segments dark brown; variations exist. | 4 |
2 | Abdomen brown, not brightly yellow or red. Cells of membrane conspicuously less pigmented than veins. Postocular lobe with longitudinal lateral patch of whitish recumbent setae. |
Zelus aithaleos ( |
– | Abdomen brightly yellow or red. Cells of membrane same color as veins. Postocular lobe without lateral whitish setae. | 3 |
3 | Abdomen red. Paramere medially curved ventrad ( |
Zelus championi ( |
– | Abdomen yellow. Paramere slightly bent near base, straight in remaining part ( |
Zelus fuliginatus ( |
4 | Dorsal surface of pronotum medially dark brown, laterally orange-brown. Paramere removed from apex of medial process, medially curved ventrad ( |
Zelus vagans ( |
– | Coloration of pronotum not as described above; anterior lobe dark brown and posterior lobe orange-brown in some specimens. Paramere straight, apex oblique ( |
Zelus gracilipes ( |
Key to males of the Zelus renardii species group |
||
1 | Relatively robust. Humeral angle of pronotum widened; body length 5.5x or less of width through humeral angles. Apical hook of paramere more prominent ( |
Zelus renardii ( |
– | Very slender. Humeral angle not conspicuously widened; body length greater than 5.5x width through humeral angles. Apical hook of paramere less prominent ( |
Zelus cervicalis ( |
Key to males of the Zelus mimus species group |
||
1 | Medial process long and slender ( |
Zelus mimus ( |
– | Medial process short and broad ( |
Zelus inconstans ( |
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. | 2 |
– | 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. | 4 |
2 | Medial process very slender, diameter less than that of paramere ( |
Zelus erythrocephalus ( |
– | Medial process broad, diameter greater than that of paramere. | 3 |
3 | Medial process twice as broad as paramere ( |
Zelus paracephalus ( |
– | Medial process only slightly broader than paramere. Paramere bent ventrally near base, not recurved ( |
Zelus russulumus ( |
4 | Medial process very broad; lateral margins parallel or converging basally. | 5 |
– | Medial process triangular; lateral margins converging apically, base broader than apex. | 6 |
5 | Length less than 11.4 mm. Medial process apex broader than base, apex acute, medially not notched ( |
Zelus laticornis ( |
– | Length more than 12 mm. Medial process lateral margins parallel, apex notched in middle ( |
Zelus casii ( |
6 | Posterior surface of medial process with ridge-like elevation, extending ventrally, not across width of medial process ( |
7 |
– | Posterior surface of medial process with hook-like process, across width of medial process; if extending ventrally, as pair of dentate processes. | 8 |
7 | Posterior margin of posterior pronotal lobe yellowish, much lighter than remaining surface of the lobe. Paramere strongly curved downward, width more or less uniform, abruptly constricted at base ( |
Zelus kartaboides ( |
– | Posterior pronotal lobe uniformly colored, dark brown. Paramere with basal 1/2 nearly straight, only curved in apical 1/2, gradually expanded toward apex ( |
Zelus kartabensis ( |
8 | Humeral angle raised to level of, and nearly continuous with, disc. Short, light, shining recumbent setae dorsally on head and pronotum. Medial process long, longer than parameres; base very broad ( |
Zelus auralanus ( |
– | Pronotal disc clearly elevated above humeral angle. Medial process narrowly triangular, shorter or subequal in length to paramere. Paramere constricted at base and somewhat bent ventrad. | 9 |
9 | Humeral angle rounded. Medial process long, upright ( |
Zelus mattogrossensis ( |
– | Humeral angle with spinous or dentate process. Medial process relatively short, semi-erect. | 10 |
10 | Medial process posterior surface with pair of processes near apex ( |
Zelus versicolor ( |
– | Medial process apex hook-like, entire, not discontinuous as pair of projections ( |
Zelus chamaeleon ( |
Key to males of the Zelus panamensis species group |
||
1 | Paramere achieving or surpassing apex of medial process. | 2 |
– | Paramere not achieving apex of medial process. | 3 |
2 | Reddish-brown. Femora without banding. Paramere curved ventrally, apex recurved slightly dorsally ( |
Zelus xouthos ( |
– | Anterior pronotal lobe dark brown, posterior lobe yellowish-brown. Femora dark brown, with two yellowish bands. Paramere straight ( |
Zelus banksi ( |
3 | Medial process shorter than or at most subequal in length to paramere. | 4 |
– | Medial process at least 1.1x length of paramere. | 6 |
4 | Head orangish or reddish. |
Zelus panamensis ( |
– | Head brown or dark brown. | 5 |
5 | Postocular lobe with longitudinal yellowish stripe. Medial process about as long as paramere ( |
Zelus truxali ( |
– | Postocular lobe without stripe. Medial process less than 0.8x length of paramere ( |
Zelus korystos ( |
6 | Paramere diameter constant through apical 3/4 or only slightly expanding. | 7 |
– | Paramere expanded apically, enlarged portion in lateral view much greater than diameter of medial process. | 9 |
7 | Medial process bent in middle ( |
Zelus filicauda ( |
– | Medial process nearly straight, curving gradually. Paramere diameter constant or weakly expanding. | 8 |
8 | Medial process posteriorly directed, at less than forty-five degree angle to horizontal axis ( |
Zelus gilboventris ( |
– | Medial process erect, at larger than forty-five degree angle to horizontal axis ( |
Zelus nigromaculatus ( |
9 | Medial process narrowed apically in lateral view ( |
Zelus cordazulus ( |
– | Medial process slightly expanded subapically in lateral view ( |
Zelus varius ( |
Key to males of the Zelus armillatus species group |
||
1 | General coloration greenish or yellowish pale brown, rather uniform. Legs without banding. | 2 |
– | Not as described above, consisting of typically two or more different colors; if uniformly colored, then dark brown or blackish-brown. Legs typically banded; if unbanded, then uniformly blackish-brown. | 4 |
2 | Paramere long, achieving or surpassing apex of medial process ( |
Zelus amblycephalus ( |
– | Paramere removed from apex of medial process. | 3 |
3 | Disc of posterior pronotal lobe strongly bulging. Paramere slender; diameter of basal 1/2 less than that of medial process in lateral view ( |
Zelus umbraculoides ( |
– | Disc slightly bulging, nearly flat. Diameter of paramere greater than that of medial process in lateral view ( |
Zelus umbraculus ( |
4 | Disc of posterior pronotal lobe depressed in middle. | 5 |
– | Disc bulging or flat. | 6 |
5 | Dorsal coloration predominantly brownish-black, with various reddish areas; venter reddish; white waxy exudation usually conspicuous dorsally and laterally (Southern S.A.). |
Zelus leucogrammus ( |
– | Entire surface testaceous-brown (Mexico and C.A.). |
Zelus sulcicollis ( |
6 | Paramere long, more than 2x length of median process. | 7 |
– | Paramere short, less than 2x length of median process. | 8 |
7 | Dorsal coloration brownish-black; femora without or with few indistinct bands. Disc of posterior pronotal lobe bears pair of tubercles (C.A.). |
Zelus lewisi ( |
– | Dorsal coloration yellow-green; legs conspicuously annulated. Disc without tubercles (Northern S.A.). |
Zelus annulosus ( |
8 | Humeral angle raised nearly to level of, and almost continuous with, disc. |
Zelus janus ( |
– | Humeral angle clearly below level of disc. | 9 |
9 | Medial process relatively robust. | 10 |
– | Medial process delicate. | 11 |
10 | As viewed posteriorly, diameter of medial process less than 1.5x ocellar diameter, slightly larger than that of paramere ( |
Zelus armillatus ( |
– | Medial process broad, diameter near middle at least 1.5x ocellar diameter ( |
Zelus conjungens ( |
11 | Head reddish-brown. Paramere very delicate ( |
Zelus ruficeps ( |
– | Head dark brown, sometimes with stripes. Apical portion of paramere obviously thicker than medial process ( |
Zelus litigiosus ( |
Key to males of the Zelus longipes species group |
||
1 | Paramere more than 2x length of medial process in lateral view, surpassing medial process by a moderately large margin. | 2 |
– | Paramere less than 2x length of medial process, achieving or slightly surpassing medial process. | 3 |
2 | Coloration consisting of reddish-brown and brownish-black, locations and relative amounts highly variable. Paramere diameter more or less constant ( |
Zelus longipes ( |
– | Coloration mainly of yellow and dark brown, posterior pronotal lobe orangish-brown. Paramere medially slightly constricted ( |
Zelus bahiaensis ( |
3 | Body elongated. Medial process long ( |
Zelus errans ( |
– | Body not as elongated as that of Zelus errans. Medial process short ( |
Zelus vespiformis ( |
Key to males of the Zelus puertoricensis species group Zelus bruneri and Zelus zayasi are not included in the key. No physical specimens were examined for the former and male is not known for the latter. |
||
1 | Postocular lobe with dorsal and ventral surfaces nearly parallel through anterior 2/3; height at middle of lobe and through ocelli subequal. Medial process short ( |
Zelus puertoricensis ( |
– | Dorsal surface sloping downward from ocelli, height at middle of lobe less than 0.9x that through ocelli. Medial process moderately long ( |
Zelus subimpressus ( |
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. |
||
1 | Anterior pronotal lobe dorsally with short, dense, erect, spine-like setae on well-defined setal tracts. Humeral angle usually rounded, some armed. | 2 |
– | 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. | 4 |
– | Dorsal surface with reddish-brown areas on pronotum and/or scutellum, often with yellowish-brown to reddish-brown areas on clavus and corium and/or basal 1/2 of membrane. | 5 |
4 | Veins and cells of membrane same color; cranial setae dark (northern S. A.). |
Zelus fuliginatus ( |
– | Cells of membrane conspicuously less pigmented than veins; postocular lobe with longitudinal lateral patch of whitish recumbent setae (Bolivia, Peru, Paraguay, southern Brazil). | 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 ( |
– | Profemoral length at least 20x profemoral diameter. Quadrate cell elongate, if broad, then Pcu more than 1/2 length of Cu. | 6 |
6 | Body elongated. Cu and Pcu of quadrate cell subparallel, Cu-Pcu2 (posterior cross vein) less than 1/2x length of Cu. | 7 |
– | Body not as elongated as that of Zelus errans or Z. gracilipes. Cu and Pcu of quadrate cell somewhat converging anteriorly, Cu-Pcu2 (posterior cross vein) more than 1/2x length of Cu. |
Zelus vespiformis ( |
7 | Entire membrane colored or opaque. |
Zelus gracilipes ( |
– | Anterior 1/2 of membrane clear or semi-translucent. |
Zelus errans ( |
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 minutus |
11 | Large, 22-25 mm. Head, pronotum and abdomen yellow or reddish with dark spots or irregularly shaped patches (C.A.). |
Zelus lewisi ( |
– | Medium-sized, 13-16 mm. Pronotum and abdomen more or less unicolorous, greyish (N.A., C.A., southern S.A.). |
Zelus tetracanthus ( |
12 | Posterior pronotal lobe, as viewed from behind, with posterior margin sloping sharply downward on either side of scutellum (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 |
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. | 14 |
– | Coloration other than above. Length variable. | 17 |
14 | Membrane and clavus with bluish iridescence. |
Zelus erythrocephalus ( |
– | Membrane shining, but not showing bluish iridescence. | 15 |
15 | Veins of corium anterior to membrane reddish-brown. Postocular lobe with dark semierect to erect setae dorsally. Scutellum bearing light recumbent setae dorsally. |
Zelus russulumus ( |
– | At least one of the characters not as described above. | 16 |
16 | Postocular lobe with light-colored, inconspicuous setae dorsally (northern S.A.). |
Zelus paracephalus ( |
– | Postocular lobe with conspicuous dark semi-erect to erect setae (southern C.A. and northern S.A.). |
Zelus panamensis ( |
17 | Length of postocular lobe less than 0.77x width of head through compound eyes. Postocular lobe with long erect setae over surface. Metafemoral length less than 20x metafemoral width (northern S.A.). |
Zelus chamaeleon ( |
– | Length of postocular lobe at least 0.80x width of head through compound eyes; if long setae present on postocular lobe, then not over entire surface. | 18 |
18 | Anterior portion of posterior pronotal lobe broadly sulcate on dorsal surface; at least 18.5 mm in length. | 19 |
– | Posterior pronotal lobe not conspicuously depressed medially; if slight depression present, not larger than 17.0 mm in length | 20 |
19 | General coloration brown; female profemur approximately 0.95x or more diameter of mesofemur (Mexico and C.A.). |
Zelus sulcicollis ( |
– | Dorsal coloration reddish-brown and brownish-black; white waxy exudation usually conspicuous dorsally; profemur approximately 0.90x or less diameter of mesofemur (southern and central S.A.). |
Zelus leucogrammus ( |
20 | Humeral angle of pronotum rounded; dorsal surface, except dorsum of abdomen, reddish-brown and brownish-black, pattern variable; erect setae predominating on dorsum of pronotum; posterior portion of anterior pronotal lobe conspicuously raised above level of anterior margin of posterior lobe (N.A. C.A., S.A. and Caribbean). |
Zelus longipes ( |
– | 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). |
Zelus puertoricensis ( |
– | Dorsal surface sloping downward from ocelli; height at middle of lobe less than 0.9x that through ocelli (Greater Antilles). |
Zelus subimpressus ( |
23 | Length at least 5.5x width. | 24 |
– | Length less than 5.5x width. | 26 |
24 | Length 12.5 mm or greater (Southern U.S., Mexico, C.A. and northwestern S.A.) |
Zelus cervicalis ( |
– | Length less than 12.5 mm (9.11-12.00 mm). | 25 |
25 | Anteocular lobe at least 1.25x length of postocular lobe. 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 ( |
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. | 28 |
27 | Profemoral length at least 20x profemoral width (S.A. and southern C.A.). |
Zelus inconstans ( |
– | Profemoral length less than 20x profemoral width (Brazil, Bolivia and Paraguay). |
Zelus mattogrossensis ( |
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. |
Zelus conjungens ( |
– | Mesofemoral diameter less than that of profemur, if mesofemoral diameter greatest, then never exceeding 1.1x that of profemoral diameter. | 30 |
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 ( |
– | 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 |
31 | Humeral angle raised to level of, and nearly continuous with, disc (Mexico and C.A.). |
Zelus janus ( |
– | 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 litigiosus ( |
– | Setal tracts and dorsum of posterior pronotal lobe with few, if any, recumbent setae; tracts not usually of contrasting color. Length:width ratio usually less than 3.5:1. | 33 |
33 | Pubescence distributed evenly over lateral surface of abdominal segments (S.A.). |
Zelus armillatus ( |
– | Pubescence, except for some scattered setae, restricted to anterior 1/2 of lateral surface of abdominal segments (Mexico, C.A., and northern S.A.). |
Zelus ruficeps ( |
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 ( |
– | Dorsal surface usually not dark brown, but if so, then setae not as above. If humeral angle raised, coloration not as dark above. | 35 |
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 xouthos |
– | Profemoral length less than 22x profemoral width. | 37 |
37 | Medial sulcus of anterior pronotal lobe with dark brown areas near posterior margin (Mexico and C.A.). |
Zelus ambulans ( |
– | Medial sulcus of anterior pronotal lobe same color as surrounding area of lobe near posterior margin (Mexico and C.A.). |
Zelus exsanguis ( |
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 ( |
– | 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 ( |
– | At least one of the characters not as described above. | 40 |
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 renardii ( |
– | Humeral angle with short to moderate, conical, spinous lateral processes, usually darker than surrounding area. Posterior pronotal lobe noticeably rugulose. | 42 |
42 | Dorsum of postocular lobe with long erect setae on posterior 1/2, some longer than ocular-ocellar distance (C.A.). |
Zelus antiguensis ( |
– | Any erect setae on dorsum of postocular lobe shorter than ocular-ocellar distance (Canada, U.S. and Mexico). |
Zelus luridus ( |
43 | Length greater than 4.5x width. Humeral angle with very short inconspicuous, spinous lateral processes. Dorsum predominately dark brown to brownish-black except for light to dark reddish-brown to brown posterior pronotal lobe, but lighter than anterior lobe (Mexico and C.A.). |
Zelus mimus ( |
– | 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. | 45 |
– | Coloration of pronotum not as described above, brown or stramineous coloration predominating dorsally, anterior pronotal lobe same color as or lighter than posterior lobe. | 49 |
45 | Anterior pronotal lobe and anterior part of posterior lobe with dark bands; wasp-like appearance. |
Zelus nigromaculatus ( |
– | Anterior pronotal lobe and anterior portion of posterior pronotal lobe of same yellowish color. | 46 |
46 | Transverse dark band on pronotum does not cover posterior margin of posterior pronotal lobe, which is instead yellowish. | 47 |
– | Transverse dark band on pronotum covers posterior margin of posterior pronotal lobe. | 48 |
47 | Femoral yellowish-brown with only small darker markings near apices (central and northern S.A.) |
Zelus kartabensis ( |
– | Femora yellowish-brown on basal 1/3 to 1/2; remainder dark brown to brownish-black, usually with one or two narrow yellowish bands (S.A.). |
Zelus plagiatus ( |
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 ( |
– | 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.). |
Zelus fasciatus ( |
49 | Length >14 mm. Anterior pronotal lobe lighter than or same color as posterior lobe. | 50 |
– | Length <13 mm. Anterior and posterior pronotal lobes same color. |
Zelus illotus ( |
50 | Anterior pronotal lobe yellowish, lighter than posterior lobe. | 51 |
– | Anterior pronotal lobe brown, same color as posterior lobe. | 52 |
51 | Femora unicolorous or with indistinct single bands. Anterior pronotal lobe elevated. |
Zelus sphegeus ( |
– | Meso- and Metafemora with two or three dark bands. Anterior pronotal lobe nearly flat, not elevated. |
Zelus truxali ( |
52 | Meso- and metafemora apical half more or less continuously reddish-brown, not broken into bands. Abdominal sternites each bearing a small black spot. |
Zelus cordazulus ( |
– | Meso- and metafemora apical half with three indistinct but visible reddish-brown bands; abdominal sternites yellow, without black spots. |
Zelus gilboventris ( |
Financial support is provided primarily by the NSF grant “Partnership in Enhancing Expertise in Taxonomy” (PEET) #0933853 awarded to C. Weirauch. Additional financial support comes from the Robert van den Bosch Scholarship (Center for Biological Control, University of California, Berkeley) and the Dissertation Year Fellowship (University of California, Riverside) awarded to G. Zhang. The following individuals assisted with imaging, databasing or measurements: Tracy Castaneda, Nick Duncan, Isaac Esquivel, Rochelle Hoey-Chamberlain, Erika Olson, Elizabeth Romero, Kaleigh Russell, Laura Soto, Amanda Villareal and Haris Yu. Jean-Michel Bérenger kindly provided the habitus images of Z. couturieri. The editorial and technical staff (Lyubomir Penev and Teodor Georgiev) of Pensoft provided critical technical support.
This file contains specimen records of species of Zelus, including type and non-type materials examined by Zhang et al. (Biodiversity Data Journal) or recorded from literature.
Measurements of species of Zelus Fabricius, 1803.