﻿A new genus and a new species of wasp-mimicking Harpactorini (Hemiptera, Heteroptera, Reduviidae, Harpactorinae), with an updated key to the Neotropical genera

﻿Abstract Quasigraptocleptesmaracristinaegen. nov., sp. nov. (Hemiptera, Heteroptera, Reduviidae, Harpactorinae, Harpactorini) is described based on male and female specimens from Brazil. Photographs and comments about the syntypes of Myocorisnigriceps Burmeister, 1835, Myocorisnugax Stål, 1872, Myocoristipuliformis Burmeister, 1838 and Xystonyttusichneumoneus (Fabricius, 1803) are presented. The intra-specific variability and sexual dimorphic characteristics among specimens of Q.maracristinaesp. nov. are recorded. General characteristics of Hiranetis Spinola, 1837, Graptocleptes Stål, 1866, Quasigraptocleptesgen. nov. and Parahiranetis Gil-Santana, 2015, which seem to be closer genera, are compared, including those of the male genitalia of some species. A key to the species of Myocoris Burmeister, 1835, and an updated key to Neotropical wasp-mimicking Harpactorini genera are provided.


Introduction
The subfamily Harpactorinae has the largest number of genera and species of Reduviidae (Hemiptera, Heteroptera) and is represented by the tribes Apiomerini and Harpactorini in the Neotropical region . Harpactorini is the most diverse group within Reduviidae with about 52 genera in the Neotropics (Forero 2011;Gil-Santana 2015;Gil-Santana et al. 2015, 2017Forero and Mejía-Soto 2021). Several taxa of this tribe are recognized as being involved in mimicry systems with Hymenoptera, resembling bees or wasps in general body and wing coloration as well as characteristics of physical proportions (Champion 1899;Haviland 1931;Elkins 1969;Maldonado and Lozada 1992;Hogue 1993;Leathers and Sharkey 2003;Gil-Santana 2008, 2015, 2016, 2022Gil-Santana et al. 2013, 2015, 2017Castro-Huertas and Forero 2021). Maldonado and Lozada (1992) presented a key to Neotropical wasp-mimicking Harpactorinae genera, which in their view helps to quickly sort out specimens from unidentified material, although this is a somewhat artificial way of grouping genera. Gil-Santana (2015) has updated their key, including two additional genera to it, Coilopus Elkins, 1969 andParahiranetis Gil-Santana, 2015. Among the genera presented in this key, while Coilopus is regarded as mimicking species of Vespidae (Elkins 1969;Forero and Giraldo-Echeverry 2015), the remaining genera are considered as mimetic of Braconidae and/or Ichneumonidae and in need of a comprehensive revision in order to clarify their systematics . For most of the genera included in this group, more recent works have described or redescribed them or at least some of their species, providing figures and more comprehensive descriptions, allowing a better knowledge of their characteristics as follows: Acanthischium Amyot & Serville, 1843 (Castro-Huertas and Forero 2021), Coilopus (Elkins 1969;Gil-Santana and Forero 2009;Forero and Giraldo-Echeverry 2015), Graptocleptes bicolor (Burmeister, 1838) (Gil-Santana et al. 2013), Hiranetis Spinola, 1837(Gil-Santana 2016, Neotropiconyttus Kirkaldy, 1909 (Maldonado andLozada 1992), and Parahiranetis Gil-Santana, 2015(Gil-Santana 2015Gil-Santana et al. 2017). However, in regard to Myocoris Burmeister, 1835 andXystonyttus Kirkaldy, 1909, there are no published images of their species, and the information about their characteristics are restricted to short original descriptions or their mentioning in diagnosis or keys (e.g., Stål 1872; Maldonado and Lozada 1992). It is noteworthy that among other genera of Harpactorini, such as Zelus Fabricius, 1803, there are some species that are considered as being similar to braconids, e.g., Zelus vespiformis Hart, 1987 andZ. errans Fabricius, 1803, while most of the other numerous species of the same genus are not (Zhang et al. 2016). Therefore, only those genera mentioned before, in which all or most of the known included species have been recognized as wasp-mimetic (Maldonado and Lozada 1992;Gil-Santana 2015), will be considered as such for the purpose of our study.
Most authors have only mentioned or given attention to the pattern of yellowish or straw-colored hemelytra with a median transverse black band, in relation to the alleged mimicry between Harpactorini and certain Ichneumonidae and Braconidae, as models (Champion 1899;Haviland 1931;Maldonado and Lozada 1992;Hogue 1993;Leathers and Sharkey 2003;Hespenheide 2010;Zhang et al. 2016). However, Gil-Santana (2015) and Gil-Santana et al. (2017) emphasized that other wasp-mimicking Harpactorini, like Parahiranetis salgadoi Gil-Santana, 2015, show a pattern of darkened to reddish general coloration with yellowish 'pterostigmata' on the hemelytra, which is similar to the similar coloration exhibited in the forewings by several other species of Ichneumonidae and Braconidae. This pattern was also observed for instance in Graptocleptes bicolor (Burmeister, 1838), G. haematogaster (Stål, 1860), and an undescribed species of Hiranetis Spinola, 1840(Gil-Santana 2016Gil-Santana et al. 2017). Yet, in some other species of wasp-mimicking Harpactorini, the hemelytra are almost or completely dark, such as in Hiranetis atra Stål, 1872(Gil-Santana 2016. Another common feature among all these species with a darkened general coloration on the hemelytra, with or without yellowish 'pterostigmata', is the presence of pale bands on the middle and hind femora (Gil-Santana 2015, 2016.
A variable degree of intraspecific variation in color, occasionally at extreme range, was documented in many harpactorines (e.g., Stål 1872; Champion 1899; Gil-Santana 2008, 2022Zhang et al. 2016;Forero and Mejía-Soto 2021), including in some waspmimicking Harpactorini (Champion 1899; Gil-Santana et al. 2013, 2017. However, at least in the species with the pattern of darkened or blackish hemelytra with yellowish pterostigmata, there is no variation in this pattern. The yellowish pterostigmata are always present (e.g., Gil-Santana et al. 2013, 2017. Numerous species of Harpactorini have shown sexual dimorphism. In several species belonging to Zelus Fabricius, 1803, for example, males and females differ drastically in size, body configuration, and coloration (Zhang et al. 2016). In Acanthischium Amyot & Serville, 1843 the pattern of coloration was found as being sexually dimorphic in most species of the genus (Castro-Huertas and Forero 2021). In addition to the bigger size and larger abdomen of females, males in several genera have larger eyes and/or a basally thickened antennal basiflagellomere. The latter has been considered to be among the diagnostic features at the generic level (Stål 1872;Champion 1899;Martin-Park et al. 2012;Gil-Santana et al. 2013, 2017Gil-Santana 2016). However, sexual dimorphism may also be limited to minor differences in coloration and size, as in many species of Zelus (Zhang et al. 2016).
The male genitalia has been found to provide useful diagnostic characteristics for distinguishing species within the genera of Harpactorini by diverse authors (e.g. Elkins 1954a, b;Hart 1975Hart , 1986Hart , 1987Forero et al. 2008;Zhang et al. 2016;Gil-Santana et al. 2017;Castro-Huertas and Forero 2021 (Fabricius, 1803) are provided to record the main characteristics of the genera to which they belong. An improved and updated version of the key to Neotropical wasp-mimicking Harpactorini genera, previously presented by Maldonado and Lozada (1992) and Gil-Santana (2015), is also provided. The original photographs were cropped, their lighting and contrast slightly adjusted but without modifying their characteristics, while the numbered scales of Figs 1-3, 5-7, 13-15, 101, 102, 104-107 were reduced or modified to similar simple scale bars in order to standardize them.
Observations were made using a stereoscope microscope (Zeiss Stemi) and a compound microscope (Leica CME). Measurements were made using a micrometer eyepiece. The total length of the head was measured excluding the neck, for better uniformity of this measurement. Dissections of the male genitalia were made by first removing the pygophore from the abdomen with a pair of forceps and then clearing it in 20% NaOH solution for 24 hours. The dissections were carried out on the genitalia of different males presenting the range of color variation recorded among them (e.g.,. The dissected structures were studied and photographed in glycerol. General morphological terminology mainly follows Schuh and Weirauch (2020). Currently, there is a lack of consensus about the terminology to be applied to female and male genitalia in Reduviidae (e.g., Rédei and Tsai 2011). Therefore, in order to maintain uniformity with previous works about species of Harpactorini, the terminology of the male and female genitalia structures generally follows Gil-Santana et al. (2013, 2017 and Gil-Santana (2016).
The type specimens of Quasigraptocleptes maracristinae gen. nov., sp. nov. will be deposited as follows: male holotype, 2 male paratypes and 1 female paratype in the Entomological Collection of the "Museu Nacional da Universidade Federal do Rio de Janeiro", Rio de Janeiro, Brazil (MNRJ); 6 male paratypes and 1 female paratype in the Dr Jose Maria Soares Barata Triatominae Collection (CTJMSB) of the São Paulo State University Julio de Mesquita Filho, School of Pharmaceutical Sciences, Araraquara, São Paulo, Brazil.
When describing label data, a slash (/) separates the lines and a double slash (//) different labels, and comments or translations to English of the label data are provided in square brackets ([ ]). All measurements are in millimeters (mm). Morphological remarks. Head cylindrical, elongated, with sparse thin setae on ventral and anteocular portions; postantennal spines short, curved forward, apices acute or somewhat rounded. Legs: fore femora thickened, narrowing at apices and somewhat curved at basal half; fore tibia curved at apical third; middle and hind legs elongated, slender; hind femora curved approximately at basal half. Burmeister (1835) described Myocoris and two new species included in this genus: M. nigriceps and M. braconiformis. In his subsequent paper, "Some account of the Genus Myocoris…" (Burmeister 1838), he considered ten species as included in the genus, some of which were described as new in this occasion, such as M. tipuliformis. While several of these ten species were transferred to other genera or considered as synonyms of other species (Maldonado 1990), Stål (1872) (Lethierry and Severin 1896;Wygodzinsky 1949;Putshkov and Putshkov 1985;Maldonado 1990) followed Stål (1872). The three species currently included in Myocoris were described from Brazil and accordingly with Stål (1872) may be separated by the following key: Only the basal portion of the first visible labial segment blackish (Fig. 3)  Note. Stål (1872) described M. nugax based on an unspecified number of female specimens from Brazil and cited "Mus. Holm." (NHRS) as the depository of the type specimen(s). Currently, only one type specimen of M. nugax was found there and it is without its abdomen (G. Lindberg pers. comm.) (Figs 9-11). Consequently, it is not possible to be sure about its gender, but possibly it is a female taking into account that Stål (1872) mentioned only the female of the species in his description. It is considered a syntype accordingly to Art. 73.2 of the ICZN. Photographs of this specimen allow observing the shorter postantennal spine and the humeral angle without acute prominence, as described by Stål (1872) (Figs 9, 11).   The male type specimen deposited in the MFNB (Figs 13-15), labeled as "Typus" (Fig. 16), is herewith considered as a syntype, accordingly with Art. 73.2 of the ICZN.  (1872) to separate the species of Myocoris were confirmed in their respective type specimens (5)(6)(7)(9)(10)(11)(13)(14)(15). However, it is necessary to study more specimens, with a more profound taxonomical approach, to ascertain their value to this separation, and more importantly, to confirm the validity of these species, such as the case of M. nigriceps and M. tipuliformis, separated only by color differences, which may be subject of intraspecific variation, as commented above.
Diagnosis. Quasigraptocletes gen. nov. can be separated from other genera of waspmimicking Harpactorini by the combination of the characteristics presented in the key below, and specially by the postantennal spines, which are strongly curved backwards.
Description. Integument mostly shiny, smooth. Head gibbous, large, approximately as long as wide across eyes (neck excluded); with sparse long and short, straight or somewhat curved blackish setae; the latter much denser, forming pubescence of long blackish thick setae on postocular portion and gula. Clypeus straight in dorsal view, curved in lateral view. Antenna inserted at level of upper third of eye; scape and pedicel straight with shiny and smooth integument; flagellomeres with opaque integument; basiflagellomere variably curved; in males conspicuously thickened approximately in basal half; distiflagellomere thinner than the other segments and slightly curved. Postantennal spines strongly curved backwards and variably directed medially. Eyes globose, glabrous, projecting laterally, prominent in dorsal view, reaching closer to dorsal margin of head at interocular sulcus slightly behind its midportion; not reaching ventral margin of head, which is far from inferior margin of the eye. Interocular sulcus thin and shallow, curved laterally; just anterior to it, on midline, a small oval fossa, followed anteriorly by a very short thin shallow median sulcus, which sometimes is not evident. Ocelli and portion between them elevated, the former much closer to eyes than to each other. Labium stout, curved, reaching prosternum approximately at proximal part of its distal third; segment II (first apparent) thickest and longest, straight, surpassing level of posterior portion of eyes; segment IV shortest, triangular, tapering. Neck thin. Thorax. Anterior collar inconspicuous; anterolateral angles moderately prominent. Transverse sulcus not very deep, interrupted before middle by a pair of submedian shallow carina. Midlongitudinal sulcus on fore lobe of pronotum shallow or not evident at basal half, deeper at distal half, above transverse sulcus, almost or forming a narrow depression; disc of hind lobe smooth; lateral longitudinal sulci well marked at posterior half to posterior two-thirds of hind lobe of pronotum. Humeral angle moderately elevated, rounded at lateral margin. Scutellum with margins elevated, apex thin, acutely pointed or sometimes rounded at its tip. Mesosternum somewhat elevated laterally, with a median U-shaped carina posteriorly. Legs: coxae globose, slightly constricted apically; femora and tibiae slender, elongate and generally straight. Fore femur shorter than head and pronotum together, slightly thickened at basal portion and somewhat curved at midportion; middle and hind femora slightly dilated subapically and slightly narrower at the portion where submedian distal pale annuli are located; apices of all femora with a pair of lateral small tubercles. Fore and middle tibiae thickened apically, the former more than the latter; at apex of fore tibiae a dorsal spur and a mesal comb. Hemelytra long, surpassing abdomen by about half length of membrane. Abdomen: elongate; spiracles rounded.
Distribution. Brazil, States of Minas Gerais and Paraná. Etymology. The name of the new genus was composed by the Latin word quasi, meaning almost, nearly, like, and Graptocleptes, in reference to its apparent proximity to the latter genus. The gender is neutral.  Table 1. Coloration: general coloration black, brownish or reddish (Figs 17-19). Head black or reddish; antennal segments black, dark brownish or reddish; labium completely dark or with distal half of second visible segment and last segment pale or entirely reddish; neck black, mostly or completely dark yellow or reddish 26,34,36,38,39,42). Thorax completely blackish, dark brown, brownish or reddish, sometimes blackish on fore lobe and brownish on hind lobe; sometimes humeral angles and/or posterior margin of hind lobe of pronotum slightly paler . Scutellum with similar coloration of thorax, sometimes with the apex of the process, in variable extension paler or whitish (47)(48)(49)(50). Hemelytra generally black with veins concolorous or paler, brownish to dark gray with the veins of the corium darker; a yellowish spot on external and mid-distal portions of corium reaching adjacent part of membrane, especially in basal portion of distal cell of membrane and just posterior to it; in paler specimens, the portion medial to the yellowish spot slightly paler (Figs 17-19). Legs. All coxae blackish (Fig. 55) or with fore coxae pale on posterior surface (Fig. 56), or completely pale (Figs 57, 58), or all coxae almost or completely pale or reddish (Fig. 59). All trochanters blackish or dark brownish (Figs 55, 56), or fore or the latter and middle trochanters partially or completely pale (Figs 57,58) or all of them almost or completely pale or reddish (Fig. 59). Femora generally blackish, brownish or reddish. Fore femora frequently with dorsal surface paler, with dark yellowish tinge or generally reddish or dark yellowish in variable extent along the segment (Figs 17-19). Middle and hind femora with one pale or yellowish submedian distal annulus or also with an additional pale or yellowish subbasal or basal annulus (Figs 17,60,64,65); sometimes the subbasal annulus is fainter, darker and/or incomplete, and as such, only evident on hind leg (Fig. 17); in specimens with reddish general coloration, the portion proximal to the submedian distal annuli is sometimes variably darker (Figs 19, 63, 66); additionally, in some specimens with a pair of annuli, the apex of these femora are also paler (Figs 18,61,62,65). Tibiae completely dark or variably pale brownish or reddish on its basal or distal portion, or almost completely or completely pale brownish to pale reddish (64)(65)(66). Tarsi in general with a similar coloration to the apex of the respective tibia. Abdomen. Sternites generally pale reddish, reddish or dark reddish with some or most segments partially or entirely darkened to blackish (Figs 67-71). Vestiture. Head covered by long and short, straight or somewhat curved blackish setae, which are denser, forming pubescence of long blackish thick setae on postocular portion, even more numerous on gula, and sparser or absent in the area anterior to transverse sulcus (26)(27)(28)(30)(31)(32)(33). Antenna: scape with sparse short, stiff, slightly curved, dark setae, which become more numerous on mesal surface, approximately in distal two-thirds (Figs 34, 35) and a few longer blackish thin setae scattered along the segment; pedicel, except at glabrous base, covered with numerous short, stiff, obliquely semi-erect dark setae (Figs 36, 37), and a few (about eight to ten) very much thinner isolated elements (interpreted as trichobothria), which are present laterally on basal two-thirds of external surface and dorsally on distal third; thickened portion of basiflagellomere, except at extreme base (which is glabrous), completely covered with short, stiff, dark, adpressed setae, and with scattered stiff, darkened, semi-erect setae and a pubescence formed by longer, very thin, pale setae, which are almost imperceptible in this portion (Figs  darkened setae and dense, erect, mostly pale, brush-like setae ventrally, which are even more numerous on basal portion and absent on hind femur. All tibiae with scattered long thick blackish setae and covered with shorter dark setae on ventral surface, which become progressively more numerous towards apex, where they also cover lateral and dorsal surfaces. Tarsi covered with shorter dark setae. Hemelytron: clavus and corium generally covered by numerous curved, short, very thin, pale setae, which become much less numerous, adpressed and even shorter on distal half of corium; membrane glabrous. Abdomen: number of setae on sternites varying among individuals, generally with scattered long thin setae, which are light on reddish portions and dark on the blackish segments, and thicker, longer, and also more numerous on parts adjacent to genitalia and on the latter (Figs 67-71). In some specimens, median portion of basal half of first visible sternite with whitish setae covered by white waxlike substance (Figs 56, 59); in one individual, a pair of lateral narrow stripes of sparse whitish setae on distal half of sternite VII, extending to basal exposed portion of pygophore (Fig. 69). Structure. Antennal basiflagellomere variably slightly curved, from 1.6 to 1.8 times longer than scape, conspicuously thickened approximately in basal half, which is clearly separated in relation to the distal thinner portion (Figs 38-40). Postantennal spines variable in length and thickness among individuals; slightly or strongly directed medially towards their apices, which are blunt to subacute (Figs 20-33). Abdominal segment VIII with only its distal margin of ventral surface visible externally (Figs 72, 73); sclerotized only on ventral portion, which is subrectangular in shape and has both basal and distal margins curved, the latter more than the former, and more prominent laterally. Male genitalia (Figs 67-87). Pygophore darkened, blackish; paler or reddish at proximal portion, in paler specimens (Figs 67-72); suboval in ventral view, somewhat enlarged laterally just below the insertions of the parameres (Fig. 73); with an enlarged, somewhat arrow-shaped apex (medial process, mp), in which lateral margins are acutely pointed and the median portion is rounded 77); between anterior and posterior genital openings, a relatively narrow dorsal (transverse) bridge (db) (Fig. 77); ventrolateral margins of exposed portion of pygophore with numerous, long, erect setae (Figs 72,73,75,76). Parameres (pa) symmetrical, rod-like in shape; apices rounded, paler at basal third, becoming darker to blackish in apical half; glabrous in basal two-thirds and with long, stout, dark setae in apical third (Figs 72,73,75,76). Phallus (Figs 78-80): articulatory apparatus with basal plate arms (ba) and basal plate bridge (bb) narrow and forming a subsquared set, except in apical portion, where the arms are curved (Fig. 82); pedicel (pd) (= basal plate extension) short (Figs 78, 80, 82). Dorsal phallothecal plate (dp) weakly sclerotized (Figs 78, 81); subrectangular in dorsal view, somewhat expanded laterally at basal portion and with small acute spines on lateral margins (Figs 78, 79, 81); medially to the latter, a pair of somewhat depressed subrectangular areas on the disc; struts (st) with curved lateral arms, which are thicker basally, and subparallel median arms slightly converging towards apices (Fig. 81). Endosoma wall smooth on basal half, becoming progressively more densely, minutely, spiny towards apex; at distal third: a pair of small more sclerotized lateral portions (sp); an apical pair of prominent sclerotized subtriangular lobes (sl), between which, ventrally, a shallower not sclerotized lobe (sn) (Figs 78-80, 85, 86). The fol-lowing endosomal processes were observed: 1 -a pair of elongate, parallel, flat, medial and weakly sclerotized processes (fp), wrapped in a smooth portion of endosoma wall, dorsally (Figs 78, 79, 83-85); 2 -a larger U-shaped basal process (u) formed by diffuse thickening (Figs 80, 87); 3 -a median subspherical process (m), situated between the lateral arms of the basal process and formed by a dense grouping of small thickenings (Figs 80, 83, 87).     Table 2. Similar to male in general. One specimen with a general dark coloration (Fig. 88) and other with reddish general coloration (Figs 89-97); both with only the submedian distal yellowish annuli on middle and hind femora evident. Structure. Head: basiflagellomere slightly thicker in basal portion (Fig. 94), but much thinner as a whole than that in males, and becoming progressively thinner toward apex, without a clear separation between more or less thickened portions (Fig. 94), uniformly covered with pubescence formed by thin, pale setae (blackish, stiff, adpressed, short setae that completely cover thicker portion in male are absent); approximately 1.2 times longer than scape. Sternites IV-VII with fusiform or elongated patches of minute, short, adpressed, thin, whitish setae, covered with a variable amount of white wax, present on midlateral portions of basal half (sternites IV, V) or, although with more numerous setae on basal portions, extending along the segment on sternites VI and VII; in one specimen, also sparsely on basal portion of the genital tergite 9 (Figs 97-99). External genitalia in posterior view (Fig. 100): tergite 9 with very long, sparse, strong blackish setae at median and lateral portions and numerous shorter, thinner setae at distal margin; tergite 10 with sparse short setae; gonocoxite 8 and gonapophysis 9 with numerous short to somewhat longer setae.

Quasigraptocleptes maracristinae
Comments. The genitalia of different males presenting a range of color variation  showed to have the same characteristics of structure (Figs 72-87). The females were slightly larger than the males (Tables 1, 2).  The minimum body length in females (to tip of hemelytra/tip of abdomen: 15.1/12.0) is greater than the maximum body length in males (14.0/11.0). Many of the other measurements are proportionally greater in females, in accordance with their bigger size (Tables 1, 2), including the antennal scape and pedicel, slightly longer than in males. One apparent exception is the basiflagellomere, which was longer in most males (5.1 to 6.5 mm in length; n = 8) than in females (5.2 to 5.3 mm in length; n = 2) and showed to be generally thicker approximately in basal half in males (maximum width: 0.2-0.4 mm) (Figs 17-19, 38-40), but thinner in females (maximum width: 0.1-0.2 mm) (Figs 88, 94). This thickened region in males is completely covered by blackish, stiff, adpressed, and short setae (Figs 38-40), which are absent in females    ( Fig. 94). While the males presented a wide range of variation in coloration and markings, the two females examined presented different patterns of coloration, but similar to some of the males. Also, the patches of minute, short, adpressed, thin, whitish setae, covered with a variable amount of white wax, present on sternites IV-VII and even on the basal portion of the genitalia (Fig. 97), were absent in the males, with the exception of the record of a narrow stripe of whitish setae on distal half of sternite VII, extending to the basal portion of the exposed portion of pygophore in a single male (Fig. 69). However, because the relatively low number of specimens examined, especially of females (only two), it is not possible to be sure in what extent most of these differences are intraspecific or sexually dimorphic characteristics. Similarly, in relation to coloration, only the examination of more specimens of both sexes will allow ascertaining the range of variation and if there is any sexual dimorphism. Distribution. Brazil, in states of Minas Gerais and Paraná. Etymology. The new species is named in honor of Dr. Mara Cristina Pinto (Faculty of Pharmaceutical Sciences, UNESP, Araraquara, São Paulo, Brazil), a friend and eternal mentor of the second author (JO), as a tribute and recognition for her contributions to the studies of Medical Entomology, especially those on sandflies and also for all her meritorious performance as a teacher and knowledge as a transforming agent. The taxon's homage is a way of rewarding all her remarkable contributions to Brazilian entomology which she has been studying for more than 33 years.
Comments. The variation in color and number and extension of pale markings recorded among the specimens of Q. maracristinae sp. nov. studied here are considered as intra-specific variability. It is in accordance with the intraspecific variation in color, occasionally at extreme range, previously documented in many harpactorines (e.g., Stål 1872; Champion 1899; Gil-Santana 2008, 2022Zhang et al. 2016), including in some wasp-mimicking Harpactorini (Champion 1899; Gil-Santana et al. 2013, 2017. The wax-like substance was sometimes absent from portions where it was observed on other specimens. It may be lost during the manipulation of the individuals, which may also include loss of the thin fragile setae associated with it (HRG-S pers. obs.; Gil-Santana et al. 2017). Body parts covered with patches of setae with whitish waxlike material have been registered in some Harpactorini species, such as Cosmoclopius curacavensis Cobben &Wygodzinsky, 1975 (Cobben andWygodzinsky 1975), Harpactor angulosus (Lepeletier & Serville, 1825) (Pikart et al. 2014), various species of Heza Amyot & Serville, 1843 (Maldonado 1976), Sphedanolestes zhengi Zhao, Ren, Wang & Cai, 2015(Zhao et al. 2015, and Parahiranetis salgadoi (Gil-Santana et al. 2017). It is noteworthy that the wax-like substance may be absent when specimens are examined and described, and thus the extent of their existence may remain unknown (Gil-Santana et al. 2017). Similarly, records of the presence or absence of a wax-like substance may be an additional feature of systematic or taxonomic importance, in the same way as suggested for the "extensive sericeous areas on the abdominal sterna" of Heza ventralis Stål, 1872 (Maldonado 1976). Therefore, as stressed by Gil-Santana et al. (2017), future studies on Harpactorini should include careful handling of the specimens after collection, to avoid unintentional removal of these substances from their bodies. It is also recommended that this information should be included in the records and/or descriptions whenever present. Differences in the structure and vestiture of the basiflagellomere were clear-cut enough to be considered sexually dimorphic in Q. maracristinae sp. nov. Despite the small number of females, adults can be sexed readily with the naked eye, by observing the basiflagellomeres of their antennae. The females examined were larger than males in many of the morphological characteristics measured, what can be confirmed by studying more specimens in the future. In any case, the two sexual differences pointed out in Q. maracristinae sp. nov. (i.e., females larger than males and the latter with basiflagellomere thickened) are concordant with several observations in the literature (Champion 1899;Martin-Park et al. 2012;Gil-Santana et al. 2013, 2017Gil-Santana 2016). Additionally, the thickened portion of the basiflagellomere in males was completely covered by short, stiff, adpressed, blackish setae, which were absent in females. Although fewer females were examined, their coloration showed similar patterns of variation of some of the males, therefore only with the examination of more specimens will be possible to ascertain possible sexual variation in coloration patterns. Yet, in the females, patches of minute, short, adpressed, thin, whitish setae, covered with a variable amount of white wax, were present on sternites IV-VII and even on the basal portion of the genitalia, while in the males they were absent (with the exception of a single male in which only a narrow stripe of whitish setae was present on distal half of sternite VII, extending to the basal portion of the exposed portion of pygophore). In other wasp-mimicking harpactorines, such as Parahiranetis salgadoi, similar patches of setae covered with white wax on sternites were observed in both sexes (Gil-Santana et al. 2017). Therefore, it is necessary to examine more specimens in order to ascertain if the absence/presence of these patches on sternites in males and females of Q. maracristinae sp. nov., respectively, expresses another sexual dimorphism or if it is merely an interindividual variation.
In the male genitalia, while the variation in color of the pygophore (Figs 67-72) is compatible with the general intra-specific variability in coloration, the uniformity of the other characteristics  is in accordance with the assumption that all specimens belong to the same species.
Yet, the male genitalia of Q. maracristinae sp. nov. showed similarities to those of G. bicolor (Gil-Santana et al. 2013), H. atra (Gil-Santana 2016, and P. salgadoi (Gil-Santana et al. 2017), such as: -parameres similar in shape and somewhat similar in vestiture; -pygophore with a somewhat large medial process that is medially rounded at the apex, but in G. bicolor and H. atra it is subtriangular in shape, while in P. salgadoi and Q. maracristinae sp. nov. it is somewhat arrow-shaped, with the lateral margins acutely pointed (Figs 72-74); -pedicel (pd) (= basal plate extension) short; -struts with subparallel median arms and curved basal lateral arms, although with different shapes in each species; -a pair of elongate, parallel, flat, weakly sclerotized endosomal processes, although with different locations and shapes in each of these species.
The presence of a somewhat laterally expanded basal portion with small acute spines on lateral margins of the dorsal phallothecal plate was recorded in P. salgadoi and Q. maracristinae sp. nov. U-shaped and median subspherical endosomal processes very similar to those of Q. maracristinae sp. nov. (Figs 80, 87) were recorded in H. atra and P. salgadoi. Yet, variable, different, or not well evident spiny lobes or portions of endosoma wall were recorded in each of these species, making their comparison difficult.
On the other hand, the general shape and peculiarities of the dorsal phallothecal plate were different in all species (Gil-Santana et al. 2013;Gil-Santana 2016;Gil-Santana et al. 2017;this study).
Thus, in agreement with previous studies (Elkins 1954a, b;Hart 1975Hart , 1986Hart , 1987Forero et al. 2008;Zhang et al. 2016), the features of the male genitalia of Q. maracristinae sp. nov. that should especially be taken into consideration for comparative purposes are the shape of the medial process of the pygophore and the features of the dorsal phallothecal plate. Fabricius (1803) recorded the species from South America ("America meridionali") without specifying a country or location. The citations of Guyana ("British Guiana") as the country from which the species was described, apparently were first stated by Lethierry and Severin (1896). Possibly, it was assumed because Fabricius (1803)  In the Natural History Museum of Denmark (ZMUC), Copenhagen, Denmark, there are two type specimens of X. ichneumoneus, both of them females, and considered here as syntypes, following Art. 73.2 of ICZN. In the syntype catalogued as ZMUC 00 103079, the head, fore legs, and a portion of the prothorax are missing (Figs 101, 102), while the other syntype, numbered as ZMUC 00 103080, is quite well preserved, with only the antennae and most tarsi missing (Figs 104-107). In the original label attached to the latter syntype (Fig. 108) the word "ichneumoneus" is clearly legible. Both specimens seem to belong to the same species judging by the remaining portions of the syntype ZMUC 00 103079 (Figs 101, 102), while the characteristics observed in the syntype ZMUC 00 103080 (Figs 104-107) agree very well with the original description (Fabricius 1803: 286). The photographs presented here are helpful to ascertain not only the characteristics of the species but also those regarded as establishing limits between the recognized genera of Neotropical wasp-mimicking Harpactorini.

Xystonyttus
The concise description of the coloration of the specimens of X. ichneumoneus, as recorded by Haviland (1931), without mention of any variation, was quite similar not only with the original description of the species (Fabricius 1803) but also with that observed in the syntypes (Figs 101, 102, 104-107). The color variations attributed to X. ichneumoneus by Stål (1866), however, are in need to be reviewed by examining series of specimens with more comprehensive approaches, since they may be variations of this single species or may represent two or more different species.
4. Quasigraptocleptes gen. nov.: head gibbous, swollen ventrally, approximately as long as width across eyes; legs thicker; fore femur shorter than head and pronotum together, somewhat thicker basally.
Additionally, while Quasigraptocleptes gen. nov. has conspicuous postantennal spines strongly curved backwards and completely or partially directed medially , in most species of the other three genera mentioned above, these spines are absent or present as small postantennal tubercles or in some species of Graptocleptes as straight vertical or semivertical spines (Stål 1872; Maldonado and Lozada 1992;Gil-Santana et al. 2013, 2017Gil-Santana 2015, 2016, 2022. Historically, only the pattern of yellowish or straw-colored hemelytra with a median transverse black band has received attention in regard to the supposed mimicry between Harpactorini and species of hymenopteran Ichneumonidae and Braconidae (Champion 1899;Haviland 1931;Maldonado and Lozada 1992;Hogue 1993;Leathers and Sharkey 2003;Hespenheide 2010). However, some species that have similar patterns of blackish wings with yellowish 'pterostigmata' and pale bands on the middle and hind femora arise as possible candidates for mimetic complexes, including Q. maracristinae sp. nov. As suggested by Gil-Santana et al. (2017), these would be the wasp-mimicking harpactorines Graptocleptes bicolor, G. haematogaster (Stål, 1860), an undescribed species of Hiranetis and Parahiranetis salgadoi as well as ichneumonoid wasps and a species of the cerambycid beetle, all of them recorded from southeastern Brazil.
Although there are records of color variation in some wasp-mimicking Harpactorini, at least in the species with the pattern of darkened or blackish hemelytra with yellowish pterostigmata, there is no variation in this pattern. The yellowish pterostigmata are always present (e.g., Gil-Santana et al. 2013, 2017. The specimens of Q. maracristinae sp. nov. studied here were concordant with this assumption, taking into account that, in spite of a considerable range of variation in color, the mentioned pattern of the hemelytra was present in all of them. Additionally, as emphasized by Gil-Santana (2015) and Gil-Santana et al. (2017), it is necessary to elucidate which species or groups of insects share the same color pattern as in Q. maracristinae sp. nov. (i.e., blackish to reddish coloration with yellowish 'pterostigmata' on wings and/or yellowish markings on legs) are involved in possible mimicry complexes.
Key to the Neotropical genera of wasp-mimicking Harpactorini Fore femora almost entirely thickened, somewhat narrowing at apex only; fore tibiae curved at apical third ( Fig. 106)