Research Article
Print
Research Article
A new genus and a new species of wasp-mimicking Harpactorini (Hemiptera, Heteroptera, Reduviidae, Harpactorinae), with an updated key to the Neotropical genera
expand article infoHélcio R. Gil-Santana, Jader Oliveira§|
‡ Laboratório de Diptera, Instituto Oswaldo Cruz, Rio de Janeiro, Brazil
§ Universidade Estadual Paulista “Julio de Mesquita Filho”, Araraquara, Brazil
| Universidade de São Paulo, São Paulo, Brazil
Open Access

Abstract

Quasigraptocleptes maracristinae gen. 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 Myocoris nigriceps Burmeister, 1835, Myocoris nugax Stål, 1872, Myocoris tipuliformis Burmeister, 1838 and Xystonyttus ichneumoneus (Fabricius, 1803) are presented. The intra-specific variability and sexual dimorphic characteristics among specimens of Q. maracristinae sp. nov. are recorded. General characteristics of Hiranetis Spinola, 1837, Graptocleptes Stål, 1866, Quasigraptocleptes gen. 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.

Keywords

Graptocleptes, Hiranetis, Parahiranetis, sexual dimorphism, wasp mimicry

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 (Gil-Santana et al. 2015). 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, 2017; Forero 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, 2022; Gil-Santana et al. 2013, 2015, 2017; Castro-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 and Parahiranetis 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 (Gil-Santana et al. 2015). 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 and Lozada 1992), and Parahiranetis Gil-Santana, 2015 (Gil-Santana 2015; Gil-Santana et al. 2017). However, in regard to Myocoris Burmeister, 1835 and Xystonyttus 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 and Z. 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 2016; Gil-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, 2022; Zhang et al. 2016; Forero and Mejía-Soto 2021), including in some wasp-mimicking 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, 2017; Gil-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 1975, 1986, 1987; Forero et al. 2008; Zhang et al. 2016; Gil-Santana et al. 2017; Castro-Huertas and Forero 2021). Among wasp-mimicking Neotropical Harpactorini, they have been described for Graptocleptes bicolor (Burmeister, 1838) (Gil-Santana et al. 2013), Hiranetis atra Stål, 1872 (Gil-Santana 2016), Parahiranetis salgadoi Gil-Santana, 2015 (Gil-Santana et al. 2017) and species of Acanthischium Amyot & Serville, 1843 (Castro-Huertas and Forero 2021). The genera to which the former three species belong, Graptocleptes Stål, 1866, Hiranetis and, Parahiranetis Gil-Santana, 2015, are considered to be closely related to each other (Stål 1872; Champion 1899; Gil-Santana 2015, 2016; Gil-Santana et al. 2017), allowing for important comparisons of several diagnostic traits of male genitalia.

Besides describing a new genus and new species, Quasigraptocleptes maracristinae gen. nov., sp. nov., photographs of the syntypes of Myocoris nigriceps Burmeister, 1835, Myocoris nugax Stål, 1872, Myocoris tipuliformis Burmeister, 1838 and Xystonyttus ichneumoneus (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.

Materials and methods

Syntypes of Myocoris nigriceps Burmeister, 1835 (catalog numbers MfN URI http://coll.mfn-berlin.de/u/915662 and MfN URI http://coll.mfn-berlin.de/u/bc0135) and Myocoris tipuliformis Burmeister, 1838 (catalog number MfN URI http://coll.mfn-berlin.de/u/f1f08c) deposited in the Hemimetabola Collection of the Museum für Naturkunde Berlin, Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany (MFNB), were directly examined by the first author (HRG-S), while their images (Figs 1–8, 13–16) were provided by Birgit Jaenicke and the copyright of these images is property of the MFNB. Images of a syntype [sex not determined] of Myocoris nugax Stål, 1872 (Figs 9–12), deposited in the Swedish Museum of Natural History, Stockholm, Sweden (NHRS) were provided by Gunvi Lindberg and are copyright (2022) of the NHRS. The photographs of the female syntypes of Zelus ichneumoneus Fabricius, 1803, deposited in the Natural History Museum of Denmark (ZMUC), Copenhagen, Denmark, were taken by Sree Gayathree Selvantharan (Figs 101–103, 106–108) and Anders Alexander Illum (Figs 104, 105), and provided by Lars Vilhelmsen.

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.

Photographs of paratypes of Quasigraptocleptes maracristinae sp. nov. (Figs 2024, 26, 34, 36, 38, 39, 42, 47–50, 5572, 8997) were taken by the second author (JO) using a stereoscope microscope (Leica 205A) with a digital camera. Scanning electron microscopy images (Figs 27, 28, 30–33, 35, 37, 40, 41, 4346, 5154, 73–76, 98–100) were also obtained by JO. Two males and a female paratypes of Quasigraptocleptes maracristinae sp. nov. were cleaned in an ultrasound machine. Subsequently, the samples were dehydrated in alcohol, dried in an incubator at 45 °C for 20 min, and fixed in small aluminum cylinders with transparent glaze. Sputtering metallization was then performed on the samples for 2 min at 10 mA in an Edwards sputter coater. After this process, the samples were studied and photographed using a high-resolution field emission gun scanning electron microscope (SEM; JEOL, JSM-6610LV), similarly as described by Rosa et al. (2010, 2014). All remaining figures were produced by the first author (HRG-S). The fixed adults, microscopic preparations, and genitalia were photographed using digital cameras (Nikon D5600 with a Nikon Macro Lens 105 mm and Sony DSC-W830). Drawings were made using a camera lucida. For clarity, the vestiture (setation) was omitted in the ink drawings of Figs 25, 29, 77. Images were edited using Adobe Photoshop CS6.

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., Figs 17–19). 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).

Taxonomy

Subfamily Harpactorinae

Tribe Harpactorini

Myocoris Burmeister, 1835

Myocoris Burmeister, 1835: 221 [key], 226 [description], 1838: 104 [diagnostic characteristics]; Stål 1859: 367 [key], 370 [diagnostic characteristics], 1866: 294 [key], 1872: 69 [key], 83 [catalog]; Walker 1873a: 49 [key], 63 [key], 1873b: 128 [catalog]; Lethierry and Severin 1896: 178 [catalog]; Wygodzinsky 1949: 42 [catalog]; Putshkov and Putshkov 1985: 52 [catalog]; Maldonado 1990: 236 [catalog]; Maldonado and Lozada 1992: 165 [key]; Forero 2011: 15 [checklist]; Gil-Santana 2015: 36 [key], 2016: 92 [citation]; Gil-Santana et al. 2017: 41 [citation]. Type species: Myocoris nigriceps Burmeister, 1835: 226, by subsequent designation, Wygodzinsky 1949: 42.

Cosmonyttus Stål 1866: 295 [key, in part, including M. nigriceps]; synonym proposed by Kirkaldy 1909: 388. Type species: Myocoris nigriceps Burmeister, 1835: 226, by monotypy.

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) described M. nugax and maintained only two other species included in Myocoris: M. nigriceps and M. tipuliformis. With exception of Walker (1873b), who considered 26 species as belonging to Myocoris, all other subsequent authors (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:

1 Humeral angles acutely spined (Figs 5, 13) 2
Humeral angles not acute, rounded (Fig. 9) M. nugax Stål, 1872
2 Only the basal portion of the first visible labial segment blackish (Fig. 3) M. nigriceps Burmeister, 1835
First visible labial segment entirely blackish (Fig. 14) M. tipuliformis Burmeister, 1838

Stål (1872) stated that M. nigriceps and M. tipuliformis were very related and similar species, while M. nugax although similar to M. tipuliformis, differed from the latter by the somewhat shorter neck, shorter postantennal spines, and humeral angle without acute prominence. Another difference in coloration between M. nigriceps and M. tipuliformis is in their legs. In M. tipuliformis the apical portion of the femora and basal portion of tibiae are blackish, while in M. nigriceps these portions are pale.

Myocoris nigriceps Burmeister, 1835

Figs 1–8

Myocoris nigriceps Burmeister, 1835: 226 [description], 1838: 105 [redescription]; Stål 1859: 370 [citation], 1872: 83 [catalog]; Walker 1873b: 128 [catalog]; Lethierry and Severin 1896: 178 [catalog]; Wygodzinsky 1949: 42 [catalog]; Maldonado 1990: 236 [catalog].

Type material examined

Myocoris nigriceps. Syntype [sex not determined; abdomen absent]: [printed label]: 2773 // [printed label]: Zool. Mus. / Berlin // [printed red label]: Typus // [handwritten label]: nigriceps / Br // [handwritten green label]: Bahia Sello. // [printed label]: [at left side]: QR CODE, [at right side]: MfN URI / http://coll.mfn- / berlin.de/u/ / 915662; Female syntype: [handwritten label]: 2773 // [printed label]: Zool. Mus. / Berlin // [printed red label]: Paratypus // [handwritten label]: Myocoris / nigriceps / Burm. / Paratypus // [handwritten green label]: Bahia / Sello // [printed label]: [at left side]: QR CODE, [at right side]: MfN URI / http://coll.mfn- / berlin.de/u/ / bc0135 (MFNB).

In the Hemimetabola Collection of the Museum für Naturkunde Berlin, Leibniz Institute for Evolution and Biodiversity Science, Berlin, Germany (MFNB) there are two type specimens of M. nigriceps. A specimen without abdomen, and sex not determined, labeled as “Typus”, catalog number MfN URI http://coll.mfn-berlin.de/u/915662 (Figs 1–4), and a female labeled as “paratypus”, catalog number MfN URI http://coll.mfn-berlin.de/u/bc0135 (Figs 5–8). Both of them are considered here as syntypes, following Art. 73.2 of the ICZN.

Figures 1–8. 

Myocoris nigriceps Burmeister, 1835, syntypes deposited in MFNB 1–4 sex not determined, catalog number MfN URI http://coll.mfn-berlin.de/u/915662 1 dorsal view 2 ventral view 3 lateral view 4 labels 5–8 female, catalog number MfN URI http://coll.mfn-berlin.de/u/bc0135 5 dorsal view 6 lateral view 7 head, anterior view 8 labels. The copyright of these images is property of the MFNB. Scale bars: 5.0 mm (1–3, 5, 6); 1.0 mm (7).

Myocoris nugax Stål, 1872

Figs 9–12

Myocoris nugax Stål, 1872: 83 [description]; Lethierry and Severin 1896: 178 [catalog]; Wygodzinsky 1949: 42 [catalog]; Maldonado 1990: 236 [catalog].

Xystonyttus nugax (Burm.); Haviland 1931: 150 [wrong combination and authorship; record from Guyana; comments on difficulties of determination: possible misidentification].

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).

Figures 9–12. 

Myocoris nugax Stål, 1872, syntype, sex not determined, deposited in NHRS, catalog number NHRS-GULI000000052, photographs provided by Gunvi Lindberg, 2022 Naturhistoriska riksmuseet. Made available by the Swedish Museum of Natural History under Creative Commons Attribution 4.0 International Public License, CC-BY 4.0, https://creativecommons.org/licenses/by/4.0/legalcode 9 dorsal view 10 ventral view 11 lateral view 12 labels.

Myocoris tipuliformis Burmeister, 1838

Figs 13–16

Myocoris tipuliformis Burmeister, 1838: 105 [description]; Stål 1859: 370 [citation], 1872: 83 [catalog]; Walker 1873b: 128 [catalog]; Lethierry and Severin 1896: 178 [catalog]; Wygodzinsky 1949: 42 [catalog]; Maldonado 1990: 236 [catalog].

Hiranetis tipuliformis; Stål 1860: 76 [catalog].

Type material examined

Male syntype: [printed label]: 2774 // [printed label]: Zool. Mus. / Berlin // [printed red label]: Typus // [handwritten label]: tipulifor- / mis* ♂. // [handwritten green label]: Rio Bescke // [printed label]: [at left side]: QR CODE, [at right side]: MfN URI / http://coll.mfn- / berlin.de/u/ / f1f08c (MFNB).

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.

Figures 13–16. 

Myocoris tipuliformis Burmeister, 1838, male syntype deposited in MFNB, catalog number MfN URI http://coll.mfn-berlin.de/u/f1f08c 13 dorsal view 14 lateral view 15 head, anterior view 16 labels. The copyright of these images is property of the MFNB. Scale bars: 5.0 mm (13, 14); 1.0 mm (15).

Comments

The characteristics stated by Stål (1872) to separate the species of Myocoris were confirmed in their respective type specimens (Figs 1–3, 5–7, 9–11, 13–15). 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.

Quasigraptocleptes gen. nov.

Type species

Quasigraptocleptes maracristinae sp. nov., by present designation.

Diagnosis

Quasigraptocleptes gen. nov. can be separated from other genera of wasp-mimicking 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.

Quasigraptocleptes maracristinae sp. nov.

Figs 17–20, 21–24, 25–27, 28–33, 34–40, 41–44, 45–51, 52–54, 55–63, 64–67, 68–77, 78–81, 82–87, 88–94, 95–100

Type material

Brazil, Minas Gerais, Juiz de Fora Municipality, x. 1997, J. da Silva leg.: 1 male holotype (MNRJ), 2 males, 1 female paratypes (MNRJ), 6 male paratypes (CTJMSB); Paraná, Londrina Municipality, 25.ii.2004, Malaise trap, Rafael Barros leg.: 1 female paratype (CTJMSB).

Description

Male. Figs 1787. Measurements are given in Table 1.

Table 1.

Measurements (mm) of male type specimens (n = 9) of Q. maracristinae sp. nov.

Holotype Mean SD Maximum Minimum
Body length to tip of hemelytra 12.2 13.45 0.55 14.0 12.2
Body length to tip of abdomen 9.5 10.47 0.47 11.0 9.5
Head length (excluding neck) 1.3 1.42 0.19 1.7 1.0
Anteocular portion length1 0.32 0.43 0.08 0.5 0.32
Postocular portion length1 0.35 0.44 0.06 0.5 0.35
Head width across eyes 1.4 1.34 0.16 1.5 1.1
Interocular distance (synthlipsis) 0.8 0.8 0.08 0.9 0.6
Transverse width of eye 0.32 0.33 0.02 0.38 0.3
Length of eye 0.5 0.5 0.05 0.6 0.4
Ocellar tubercle width 0.9 0.82 0.09 0.9 0.6
Scape length 3.4 3.43 0.1 3.5 3.2
Pedicel length 1.0 0.97 0.04 1.0 0.9
Basiflagellomere length (n = 8)2 6.5 5.48 0.43 6.5 5.1
Basiflagell. max. width (n = 8)2 0.2 0.26 0.07 0.4 0.2
Distiflagellomere length (n = 3)2 1.5 0.00 1.5 1.5
Labial segment II length1 0.9 0.95 0.07 1.0 0.8
Labial segment III length1 0.8 0.82 0.08 0.9 0.7
Labial segment IV length1 0.2 0.21 0.03 0.3 0.2
Pronotum length (at midline) 2.0 2.27 0.11 2.4 2.0
Pronotum maximum width 1.9 2.37 0.19 2.5 1.9
Scutellum length 0.9 0.96 0.05 1.0 0.9
Fore femur length 3.2 3.39 0.16 3.7 3.2
Fore tibia length 3.5 3.6 0.15 4.0 3.5
Fore tarsus length (n = 7)2 0.4 0.4 0.06 0.5 0.3
Middle femur length 2.6 2.9 0.16 3.1 2.6
Middle tibia length 3.2 3.45 0.21 3.7 3.0
Middle tarsus length 0.5 0.52 0.03 0.6 0.5
Hind femur length 3.9 4.26 0.20 4.5 3.9
Hind tibia length 5.1 5.85 0.36 6.2 5.1
Hind tarsus length 0.5 0.57 0.05 0.65 0.5
Abdomen length3 5.3 5.58 0.35 6.5 5.3
Abdomen maximum width 1.2 1.52 0.13 1.7 1.2

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 (Figs 1724, 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 (Figs 1724). Scutellum with similar coloration of thorax, sometimes with the apex of the process, in variable extension paler or whitish (Figs 17–19, 47–50). 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 (Figs 17–19, 64–66). 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 6771). 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 (Figs 2024, 2628, 30–33). 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 38–40); distal (not thickened) portion of basiflagellomere and distiflagellomere covered with dense pubescence formed by short, thin, pale to whitish setae and with scattered short, darkened, stiff, semi-erect setae; the latter somewhat less numerous or not evident on distiflagellomere (Figs 38, 39, 41–43). Labium with scattered and somewhat curved, longer and thinner dark setae (Figs 28, 44). Eyes and ocelli glabrous (Figs 27, 28, 30). Neck almost completely or completely glabrous (Figs 2024, 2628, 30). Thorax. Prothorax covered with very numerous blackish thick setae on fore lobe of pronotum, anterior portions of propleura and hind lobe of pronotum; the latter with sparse long setae at dorsal portion or, almost glabrous, except on midline, where thinner, somewhat shorter and light yellowish to whitish setae form a faint midlongitudinal line on hind lobe; median portion of posterior margin of pronotum with some long thin darkened setae (Figs 2024, 45, 52). Scutellum with scattered thin dark or pale long setae more or less numerous; sometimes with midlongitudinal line of whitish setae on approximately its basal third, which may be a continuation of the line of whitish setae on hind lobe, or may be present only on the scutellum, while absent on pronotum (Figs 4652). The pale setae which form the midlongitudinal line on hind lobe of pronotum and/or scutellum are sometimes partially or completely covered by a small amount of white wax-like substance (e.g. Figs 21, 23, 47, 49). Posterior portion of propleura, mesopleura, metapleura and thoracic sterna with less numerous long darkened setae, which are shorter and thinner at center of mesosternum and metasternum (Figs 52, 55–59). In some specimens, there is a group of thin whitish setae basally covered with rounded, flocky patches of white wax-like substance, extending along midline of mesosternum and metasternum and sometimes extending to basal half of first visible sternite (Figs 55, 56, 58, 59). Dorsal sclerite below scutellum covered by numerous minute spiny setae (Figs 53, 54). Legs: coxae with numerous long thin setae on posterior and lateral surfaces, which are less numerous or absent on basal third; fore trochanters densely covered with pale long setae ventrally and with some scattered even longer thinner pale setae; middle and hind trochanters progressively less setose. All femora covered with scattered, long, straight, erect or semi-erect 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 6771). In some specimens, median portion of basal half of first visible sternite with whitish setae covered by white wax-like 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 2033). 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.

Figures 17–20. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males, dorsal view 17 holotype 18–20 paratypes 20 head and pronotum. Scale bars: 5.0 mm (17–19); 0.5 mm (20).

Figures 25–27. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males, head, dorsal view. Scale bars: 0.5 mm (25–27).

Figures 21–24. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males, head and pronotum, dorsal view. Scale bars: 0.5 mm (21–24).

Male genitalia (Figs 6787). Pygophore darkened, blackish; paler or reddish at proximal portion, in paler specimens (Figs 6772); 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 (Figs 72–75, 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 following 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).

Figures 28–33. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males, head 28–30 posterolateral view 30 upper portion 31 anterior portion, including postantennal spines, dorsal view 32 postantennal spines, laterodorsal view 33 left postantennal spine, dorsal view. Scale bars: 0.5 mm (28–30); 0.1 mm (31–33).

Figures 34–40. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males, antenna, dorsal view 34, 35 scape 36, 37 apex of scape, pedicel and basal portion of basiflagellomere 38–40 arrows point to the portion of clear separation between the thickened and thinner portions of basiflagellomere 38 apex of pedicel and flagellomeres 39 apex of pedicel and basiflagellomere 40 apex of scape and approximately basal half of basiflagellomere. Scale bars: 0.5 mm (34–36, 38–40); 0.2 mm (37).

Figures 41–44. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., male 41 basiflagellomere, distal half, dorsal view 42 distal portion of basiflagellomere and distiflagellomere, lateral view 43 distiflagellomere, dorsal view 44 lower portion of head and labium, lateral view. Scale bars: 0.5 mm (42); 0.3 mm (41); 0.2 mm (44); 0.1 mm (43).

Female. Figs 88100. Measurements are given in Table 2. Similar to male in general. One specimen with a general dark coloration (Fig. 88) and other with reddish general coloration (Figs 8997); 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.

Table 2.

Measurements (mm) of female type specimens (n = 2) of Q. maracristinae sp. nov.

Mean SD Maximum Minimum
Body length to tip of hemelytra 15.3 0.28 15.5 15.1
Body length to tip of abdomen 12.1 0.14 12.2 12.0
Head length (excluding neck) 1.45 0.07 1.5 1.4
Anteocular portion length1 0.55 0.62 0.6 0.5
Postocular portion length1 0.55 0.62 0.6 0.5
Head width across eyes 1.45 0.07 1.5 1.4
Interocular distance (synthlipsis) 0.8 0.8 0.8
Transverse width of eye 0.34 0.01 0.35 0.33
Length of eye 0.5 0.5 0.5
Ocellar tubercle width 0.9 0.9 0.9
Scape length 4.25 0.07 4.3 4.2
Pedicel length 1.35 0.07 1.4 1.3
Basiflagellomere length 5.25 0.07 5.3 5.2
Basiflagell. max. width 0.15 0.07 0.2 0.1
Distiflagellomere length (n = 1)2 1.5 1.5
Labial segment II length1 1.2 1.2 1.2
Labial segment III length1 0.9 0.9 0.9
Labial segment IV length1 0.25 0.07 0.3 0.2
Pronotum length (at midline) 2.65 0.07 2.7 2.6
Pronotum maximum width 2.9 2.9 2.9
Scutellum length 1.2 1.2 1.2
Fore femur length 3.65 0.07 3.7 3.6
Fore tibia length 4.1 0.14 4.2 4.0
Fore tarsus length 0.5 - 0.5 0.5
Middle femur length 3.1 0.14 3.2 3.0
Middle tibia length 3.85 0.07 3.9 3.8
Middle tarsus length (n = 1)2 0.8 0.8
Hind femur length 4.8 0.14 4.9 4.7
Hind tibia length 5.25 0.07 5.3 5.2
Hind tarsus length 0.65 0.07 0.7 0.6
Abdomen length3 7.1 0.14 7.2 7.0
Abdomen maximum width 1.85 0.07 1.9 1.8
Figures 45–51. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males, thorax 45–50 dorsal view. 45 pronotum 46–51 scutellum 51 lateral view. Scale bars: 0.5 mm (45, 49); 0.25 mm (47, 48, 50); 0.1 mm (46, 51).

Figures 52–54. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., male, thorax, lateral view 52 legs and wings extracted 53, 54 sclerite below scutellum 54 under higher magnification. Scale bars: 0.3 mm (52); 0.1 mm (53, 54).

Figures 55–63. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males 55–59 thorax and first visible (s) sternite (s), ventral view 60–63 middle femur 60–62 lateral view 63 ventral view. Scale bars: 0.5 mm (55–59); 0.4 mm (60–63).

Comments

The genitalia of different males presenting a range of color variation (e.g. Figs 17–19) showed to have the same characteristics of structure (Figs 7287). 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.

Figures 64–67. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males 64–66 hind leg, lateral view. 67 abdomen, ventral view. Scale bars: 0.5 mm (64–67).

Figures 68–77. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., males 68–71 abdomen and apical portion of membrane of hemelytra, ventral view 72–76 ventral view 72, 73 apex of abdomen with pygophore in situ, black arrows point to distal margin of segment VIII 74 medial process of pygophore 75, 76 lateral portion of pygophore and left paramere 77 pygophore without parameres, schematic outline, dorsal view. Abbreviations: db: dorsal bridge; mp: medial process of pygophore; pa: paramere. Scale bars: 0.5 mm (68–71, 77); 0.2 mm (72); 0.1 mm (73–76).

Figures 78–81. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., male genitalia 78–80 phallus 78 dorsal view 79 lateral view 80 ventral view 81 dorsal phallothecal plate and struts, dorsal view. Abbreviations: ba: basal plate arm; bb: basal plate bridge; dp: dorsal phallothecal plate; fp: flat process of endosoma; m: median process of endosoma; pd: pedicel; sl: sclerotized subtriangular lobe of endosoma wall; sn: not sclerotized lobe; sp: small sclerotized portion; st: strut; u: U-shaped process of endosoma. Scale bars: 0.2 mm (78–80); 0.1 mm (81).

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, 2022; Zhang 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 wax-like material have been registered in some Harpactorini species, such as Cosmoclopius curacavensis Cobben & Wygodzinsky, 1975 (Cobben and Wygodzinsky 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, 2017; Gil-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 inter-individual variation.

In the male genitalia, while the variation in color of the pygophore (Figs 6772) is compatible with the general intra-specific variability in coloration, the uniformity of the other characteristics (Figs 7287) 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.

Figures 82–87. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., male genitalia 82–86 dorsal view 82 articulatory apparatus 83 phallus, middle portion 84 flat processes of endosoma 85 phallus, apical portion 86 apical subtriangular lobes of endosoma wall; 87 U-shaped and median processes of endosoma, ventral view. Abbreviations: ba: basal plate arm; bb: basal plate bridge; fp: flat process of endosoma; m: median process of endosoma; pd: pedicel; sl: sclerotized subtriangular lobe of endosoma wall; sn: not sclerotized lobe; sp: small sclerotized portion; u: U-shaped process of endosoma. Scale bars: 0.2 mm (82, 87); 0.1 mm (83–86).

Figures 88–94. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., females 88–89 dorsal view. 89 head and anterolateral angles of pronotum 90 head, anterolateral view 91 left postantennal spine (pointed by an arrow), inner surface, lateral view 92–94 antennal segments, dorsal view 92 scape and pedicel 93 apex of scape, pedicel and basal portion of basiflagellomere 94 basiflagellomere. Scale bars: 5.0 mm (88); 0.5 mm (89, 92–94); 0.2 mm (90); 0.1 mm (91).

Figures 95–100. 

Quasigraptocleptes maracristinae gen. nov., sp. nov., female 95 postocular portion of head, pronotum and basal portion of scutellum, dorsal view 96–99 ventral view 96 thorax and first visible sternites 97 abdomen 98 patches of minute, short, adpressed, thin, whitish setae, covered with a variable amount of white wax on left side of sternites VI–VII 99 main portion of the patch of left side of sternite VI in higher magnification 100 female genitalia, posterior view. Scale bars: 0.5 mm (95–97); 0.1 mm (98–100).

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 1975, 1986, 1987; Forero 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.

Xystonyttus Kirkaldy, 1909

Cosmonyttus Stål, 1868: 103; Stål 1872: 83 (not Stål 1866: 295); Lethierry and Severin 1896: 178 [catalog; including erroneously Stål 1866: 295]. Type species: Zelus ichneumoneusFabricius 1803: 286, by monotypy.

Xystonyttus Kirkaldy, 1909: 388 [as a new name for “Cosmonyttus, Stål, 1872 (not 1866)”]; Wygodzinsky 1949: 48 [catalog]; Putshkov and Putshkov 1985: 66 [catalog]; Maldonado 1990: 324 [catalog]; Maldonado and Lozada 1992: 165 [key]; Forero 2011: 16 [checklist]; Gil-Santana 2015: 37 [key], 2016: 92 [citations]; Gil-Santana et al. 2017: 41 [citation]. Type species: Zelus ichneumoneusFabricius 1803: 286, by original designation.

Morphological remarks

Head gibbous, large, as long as wide across eyes, densely covered with long setae on ventral and postocular portions; postantennal spines elongated, curved forward, apices acute. Legs: fore and hind femora curved at median portion; fore femora thickened, narrowing at apices; fore tibia curved at apical third; middle and hind legs elongated, slender. Hemelytra long, surpassing the abdomen by somewhat more than half of the length of the membrane.

Xystonyttus ichneumoneus (Fabricius, 1803)

Figs 101–108

Zelus ichneumoneus Fabricius, 1803: 286 [description; “Habitat in America meridionali”].

Cosmonyttus ichneumoneus Stål, 1868: 103–104 [redescription; color varieties “a”, “b” and “c”; record from Suriname]; Stål 1872: 83 [catalog]; Lethierry and Severin 1896: 178 [catalog; cited as being from Guyana].

Myocoris ichneumoneus Walker, 1873b: 129 [catalog; record from Brazil].

Xystonyttus ichneumoneus Kirkaldy, 1909: 388 [type species to Xystonyttus, new name to Cosmonyttus Stål, 1872]; Haviland 1931: 150 [record from Guyana]; Wygodzinsky 1949: 48 [catalog; cited as being from [British] Guyana]; Maldonado 1990: 324 [catalog; cited as being originally described from [British] Guyana]; Gil-Santana et al. 2003: 12 [comment on its etymology].

Notes

The general similarity of X. ichneumoneus (Figs 101, 102, 104, 105) with ichneumonids certainly led Fabricius (1803) to name it as such (Gil-Santana et al. 2003). 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) cited “Dom. Smidt. Mus. Dom. de Sehestedt” when stating about the specimens of Zelus ichneumoneus examined by him. In fact, “most of the numerous South American species that Fabricius described and for which a certain Smidt is cited as a collector are found only in the Copenhagen Museum, in Tönder Lund’s and Sehestedt’s famous collections (…). The only information (…) on Smidt is that he visited besides several West Indian islands, certain places on the South American mainland, such as Essequibo and Demerara in the present British Guiana; therefore all of the South American species cited as having been collected by Smidt can with certainty be considered as coming from the vicinity of the named localities, and this is just what one who is familiar with American Hemiptera and their distribution immediately perceives” (Stål 1868: 3, translation by G. C. Steyskal in Papavero 1971: 21). Stål (1868: 104), however, when recording the distribution of X. ichneumoneus maintained the original statement of Fabricius (1803) (“America meridionalis. Dom Smidt. (Mus. Sehestedt)”), adding: “Surinam. (Mus. Holm.)”. The latter refers to the specimen(s) examined and considered as belonging to X. ichneumoneus by himself in Stockholm Museum (currently, Swedish Museum of Natural History, Stockholm, Sweden), while Stål (1872) restricted the occurrence of the species to Suriname only. In any case, Haviland (1931) recorded X. ichneumoneus from Kartabo, Bartica District, Guyana, confirming the presence of the species in this country.

Figures 101–108. 

Zelus ichneumoneus Fabricius, 1803, female syntypes deposited in ZMUC 101–103 syntype catalog number ZMUC 00 103079 101 dorsal view 102 ventral view 103 labels 104–108 syntype catalog number ZMUC 00 103080 104 dorsal view 105 ventral view 106 head, pronotum, fore legs, lateral view 107 head and most part of pronotum, dorsal view 108 labels. Scale bars: 5.0 mm (104, 105); 2.0 mm (101, 102, 106, 107).

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.

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.

Discussion

Among the Neotropical Harpactorini, Quasigraptocleptes gen. nov. seems closer to Graptocleptes and Hiranetis, while the latter have been considered allied genera (Stål 1872; Champion 1899) and also close to Parahiranetis (Gil-Santana 2015; Gil-Santana et al. 2017).

The main diagnostic characteristics that separate Hiranetis, Graptocleptes, Parahiranetis and Quasigraptocleptes gen. nov. according to Spinola (1840a, b), Stål (1866, 1872), Maldonado and Lozada (1992), Gil-Santana et al. (2013, 2017), Gil-Santana (2015, 2016) and this work, are the following:

  1. Graptocleptes : head elongate, approximately 1.1 to 1.3 times as long as width across eyes; legs thicker; fore femur shorter than head and pronotum together, somewhat thicker basally or with uniform thickness.
  2. Hiranetis : head gibbous, swollen ventrally, approximately as long as width across eyes; legs elongated, slender; fore femur subequally longer than head and pronotum together, thicker basally.
  3. Parahiranetis : head elongate, approximately 1.3–1.7 times as long as width across eyes; legs elongated, slender; fore femur subequally longer than head and pronotum together, thicker basally.
  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 (Figs 2033), 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, 2017; Gil-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

Based on Stål 1866, 1872; Elkins 1969; Maldonado and Lozada 1992; Gil-Santana 2015; Castro-Huertas and Forero 2021.

1 Pronotum greatly inflated and covering scutellum posteriorly Coilopus Elkins, 1969
Pronotum not inflated; scutellum not covered by the posterior portion of pronotum and visible from above 2
2 Hind lobe of pronotum with a pair of elevated submedial longitudinal carinae and spines on its posterior margin; fore trochanter with a ventral spine in most species Acanthischium Amyot & Serville, 1843
Hind lobe of pronotum different; fore trochanter without spines 3
3 Postantennal spines curved and directed forward 4
Postantennal spines absent, as tubercles, straight directed vertically, semivertically or strongly curved backwards 6
4 Head generally sparsely setose (Figs 3, 11, 14) Myocoris Burmeister, 1835
Head quite setose to very densely setose, especially on ventral and post-ocular portions 5
5 Fore femora almost entirely thickened, somewhat narrowing at apex only; fore tibiae curved at apical third (Fig. 106) Xystonyttus Kirkaldy, 1909
Fore femora thicker only basally; fore tibiae straight Neotropiconyttus Kirkaldy, 1909
6 Postantennal spines strongly curved backwards and completely or partially directed medially (Figs 2033) Quasigraptocleptes gen. nov.
Postantennal spines absent, or straight directed vertically, semivertically, or as tubercles 7
7 Fore femora slender, elongated; clearly thicker basally 8
Fore femora stouter; not or only slightly thicker basally Graptocleptes Stål, 1866
8 Head gibbous, swollen ventrally, subequally as long as wide across eyes; in dorsal view, postocular portion clearly separated from a distinct neck Hiranetis Spinola, 1837
Head elongate, not swollen ventrally, approximately 1.3 to 1.7 times as long as wide across eyes; in dorsal view, postocular portion narrowing gradually to form neck Parahiranetis Gil-Santana, 2015

Acknowledgements

We thank the “Laboratório de Microscopia Avançada LMA, Instituto de Química, Unesp”, Araraquara, São Paulo, Brazil” for allowing the use of the scanning electron microscope facility. The first author (HRG-S) thanks Jürgen Deckert, former curator of MFNB, for allowing him to examine specimens of this institution. The second author (JO) thanks FAPESP (“Fundação de Amparo à Pesquisa do Estado de São Paulo”) and Dr João Aristeu da Rosa for support and for providing the facilities of the Parasitology Laboratory, and is very grateful to the Brazilian entomologist Rafael Barros for donating a female paratype of Q. maracristinae sp. nov. We also especially thank the Natural History Museum of Denmark, Anders Alexander Illum, Sree Gayathree Selvantharan, and Lars Vilhelmsen (ZMUC), Gunvi Lindberg (NHRS), as well as Birgit Jaenicke (MFNB), for the photographs that they provided. We are also grateful to an anonymous reviewer, Manuel Baena, and Nikolay Simov for their valuable comments and suggestions.

References

  • Burmeister H (1835) Handbuch der Entomologie. Zweiter Band. Erste Abtheilung. Schnabelkerfe. Rhynchota. Theodor Christian Friedrich Enslin, Berlin, 400 pp.
  • Castro-Huertas V, Forero D (2021) Revision and phylogenetic analysis of the genus Acanthischium Amyot & Serville, 1843 (Hemiptera: Reduviidae: Harpactorinae). Insect Systematics & Evolution 52(5): 524–574. https://doi.org/10.1163/1876312X-bja10018
  • Champion GC (1899) Insecta Rhynchota. Hemiptera-Heteroptera, Vol II. In: Godman FD, Salvin O (Eds) Biologia Centrali Americana. Taylor and Francis, London, 193–304.
  • Cobben RH, Wygodzinsky P (1975) The Heteroptera of the Netherlands Antilles – IX. Reduviidae (Assassin Bugs). Studies on the Fauna of Curaçao and other Caribbean Islands 48: 1–62.
  • Elkins JC (1954a) A new American Harpactorine genus (Hemiptera, Reduviidae). Texas Reports on Biology and Medicine 12: 39–48.
  • Elkins JC (1954b) A synopsis of Atrachelus (Hemiptera, Reduviidae). Proceedings of the Entomological Society of Washington 56: 97–120.
  • Elkins JC (1969) A new genus of hemipteran wasp mimics (Reduviidae, Harpactorinae). Journal of the Kansas Entomological Society 42: 456–461.
  • Fabricius JC (1803) Systema Rhyngotorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Carolum Reichard, Brunsvigae, 314 pp. https://doi.org/10.5962/bhl.title.11644
  • Forero D (2011) Classification of Harpactorinae, assassin bugs Hemiptera: Heteroptera: Reduviidae. Boletín del Museo Entomológico Francisco Luís Gallego 1: 9–24.
  • Forero D, Giraldo-Echeverry N (2015) First record of the assassin bug genus Coilopus Elkins, 1969 (Hemiptera: Heteroptera: Reduviidae) from Colombia. Check List 11(3): 1634. https://doi.org/10.15560/11.3.1634
  • Forero D, Mejía-Soto A (2021) A striking sexually dimorphic new species of Castolus (Hemiptera: Heteroptera: Reduviidae) from Colombia, with new records from Neotropical countries and taxonomic notes on the genus. Zootaxa 5048(4): 538–560. https://doi.org/10.11646/zootaxa.5048.4.4
  • Forero D, Gil-Santana HR, van Doesburg PH (2008) Redescription of the Neotropical genus Aristathlus (Heteroptera, Reduviidae, Harpactorinae). In: Grozeva S, Simov N (Eds) Advances in Heteroptera research: festschrift in honor of 80th anniversary of Michail Josifov. Pensoft, Sofia-Moscow, 85–103.
  • Gil-Santana HR (2008) New records, and nomenclatural and biological notes on Reduviidae (Hemiptera: Heteroptera) from Bolivia and Brazil. Zootaxa 1785(1): 43–53. https://doi.org/10.11646/zootaxa.1785.1.2
  • Gil-Santana HR (2015) Parahiranetis salgadoi, a new genus and species of Harpactorini (Hemiptera: Heteroptera: Reduviidae), with a key to Neotropical wasp-mimicking harpactorine genera. Acta Entomologica Musei Nationalis Pragae 55: 29–38.
  • Gil-Santana HR (2016) First description of the male of Hiranetis atra Stål and new country records, with taxonomic notes on other species of Hiranetis Spinola (Hemiptera, Heteroptera, Reduviidae, Harpactorinae). ZooKeys 605: 91–111. https://doi.org/10.3897/zookeys.605.8797
  • Gil-Santana HR (2022) New records, taxonomic notes, and the description of a new species of Harpactorinae (Hemiptera: Heteroptera: Reduviidae) from French Guiana. Zootaxa 5105(3): 381–400. https://doi.org/10.11646/zootaxa.5105.3.3
  • Gil-Santana HR, Forero D (2009) A new species of Notocyrtus, a new synonym of Coilopus, and new records and notes on other Harpactorini (Hemiptera: Heteroptera: Reduviidae: Harpactorinae) from South America. Zootaxa 2148(1): 55–67. https://doi.org/10.11646/zootaxa.2148.1.5
  • Gil-Santana HR, Costa LAA, Forero D, Zeraik SO (2003) Sinopse dos Apiomerini, com chave ilustrada para os gêneros (Hemiptera-Heteroptera, Reduviidae, Harpactorinae). Publicações Avulsas do Museu Nacional 97: 1–24.
  • Gil-Santana HR, Davranoglou L-R, Neves JA (2013) A new synonym of Graptocleptes bicolor (Burmeister), with taxonomical notes (Hemiptera: Heteroptera: Reduviidae: Harpactorini). Zootaxa 3700: 348–360. https://doi.org/10.11646/zootaxa.3700.3.2
  • Gil-Santana HR, Forero D, Weirauch C (2015) Assassin bugs (Reduviidae excluding Triatominae). In: Panizzi AR, Grazia J (Eds) True bugs (Heteroptera) of the Neotropics, Entomology in Focus 2. Springer Science+Business Media, Dordrecht, 307–351. https://doi.org/10.1007/978-94-017-9861-7_12
  • Gil-Santana HR, Salomão AT, Oliveira J (2017) First description of the male and redescription of the female of Parahiranetis salgadoi Gil-Santana (Hemiptera, Reduviidae, Harpactorinae). ZooKeys 671: 19–48. https://doi.org/10.3897/zookeys.671.11985
  • Hart ER (1975) A new species of Ischnoclopius Stål, with notes on the systematic position of the genus (Hemiptera: Reduviidae). Proceedings of the Entomological Society of Washington 94: 162–165.
  • Hart ER (1986) Genus Zelus Fabricius in the United States, Canada, and Northern Mexico (Hemiptera: Reduviidae). Annals of the Entomological Society of America 77(3): 419–425. https://doi.org/10.1093/aesa/79.3.535
  • Hart ER (1987) The genus Zelus Fabricius in the West Indies (Hemiptera: Reduviidae). Annals of the Entomological Society of America 80(2): 293–305. https://doi.org/10.1093/aesa/80.2.293
  • Hogue CL (1993) Latin American Insects and Entomology. University of California Press, Los Angeles, USA, 536 pp.
  • ICZN [International Comission on Zoological Nomenclature] (1999) International code of zoological nomenclature. 4th edn. The International Trust for Zoological Nomenclature, London. https://www.iczn.org/the-code/
  • Leathers JW, Sharkey MJ (2003) Taxonomy and life history of Costa Rican Alabagrus (Hymenoptera: Braconidae), with a key to World species. Contributions in Science 497: 1–78. https://doi.org/10.5962/p.214390
  • Lethierry L, Severin G (1896) Catalogue général des Hémiptères. Tome III. Hétéroptères. R. Friedländer & Fils, Libraires-Éditeurs, Berlin, 275 pp.
  • Maldonado CJ (1990) Systematic catalogue of the Reduviidae of the World. Caribbean Journal of Science, Special publication No. 1, University of Puerto Rico, Mayagüez, 694 pp.
  • Maldonado CJ, Lozada RPW (1992) Key to the group of neotropical wasp-mimetic harpactorine genera and the description of a new species (Hemiptera: Reduviidae). Proceedings of the Entomological Society of Washington 94: 162–165.
  • Martin-Park A, Delfín-González H, Coscarón MC (2012) Revision of genus Repipta Stål, 1859 (Hemiptera: Heteroptera: Reduviidae: Harpactorinae) with new species and distribution data. Zootaxa 3501(1): 1–54. https://doi.org/10.11646/zootaxa.3501.1.1
  • Papavero N (1971) Essays on the history of Neotropical Dipterology, with special reference to collectors (1750–1905). Vol. 1. Universidade de São Paulo, São Paulo, 216 pp. https://doi.org/10.5962/bhl.title.101715
  • Pikart TG, Souza GK, Ribeiro RC, Zanuncio JC, Serrão JE (2014) Epidermis associated with wax secretion in the Harpactor angulosus (Hemiptera: Reduviidae). Annals of the Entomological Society of America 107(1): 227–233. https://doi.org/10.1603/AN13003
  • Putshkov VG, Putshkov PV (1985) A catalogue of the Assassin-bugs genera of the World (Heteroptera, Reduviidae). Published by the authors, Kiev, 137 pp.
  • Rédei D, Tsai J-F (2011) The assassin bug subfamilies Centrocnemidinae and Holoptilinae in Taiwan (Hemiptera: Heteroptera: Reduviidae). Acta Entomologica Musei Nationalis Pragae 51: 411–442.
  • Rosa JA, Mendonça VJ, Rocha CS, Gardim S, Cilense M (2010) Characterization of the external female genitalia of six species of Triatominae (Hemiptera, Reduviidade) by scanning electron microscopy. Memorias do Instituto Oswaldo Cruz 105(3): 286–292. https://doi.org/10.1590/S0074-02762010000300007
  • Rosa JA, Mendonça VJ, Gardim S, Carvalho DB, Oliveira J, Nascimento JD, Pinotti H, Pinto MC, Cilense M, Galvão C, Barata JMS (2014) Study of the external female genitalia of 14 Rhodnius species (Hemiptera, Reduviidae, Triatominae) using scanning electron microscopy. Parasites & Vectors 7(1): 17. https://doi.org/10.1186/1756-3305-7-17
  • Schuh RT, Weirauch C (2020) True Bugs of the World (Hemiptera: Heteroptera). Classification and natural history, 2nd edn. Siri Scientific Press, Manchester, 767 pp. [32 pls]
  • Spinola M (1840a) [1837] Essai sur les genres d’insectes appartenants à l’ordre des Hémiptères, Lin. ou Rhyngotes, Fab. et à la section des Hétéroptères, Dufour. Yves Gravier, Gènes, 383 pp. https://doi.org/10.5962/bhl.title.65481
  • Stål C (1859) Till kännedomen om Reduvini. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 16(8): 363–386.
  • Stål C (1860) Bidrag till Rio de Janeiro-Traktens Hemipter-Fauna. Kongliga Svenska Vetenskaps-Akademiens Handlingar 2(7): 1–84.
  • Stål C (1866) Bidrag till Reduviidernas kännedom. Öfversigt af Kongliga Vetenskaps-Akademiens Förhandlingar 23(9): 235–302.
  • Stål C (1868) Hemiptera Fabriciana. Kongliga Svenska Vetenskaps-Akademiens Handlingar 7: 93–131.
  • Stål C (1872) Enumeratio Hemipterorum. Bidrag till en företeckning öfver alla hittils kända Hemiptera, jemte systematiska meddelanden. 2. Kongliga Svenska Vetenskaps-Akademiens Handlingar 10(4): 1–159.
  • Walker F (1873a) Catalogue of the specimens of Hemiptera Heteroptera in the Collection of the British Museum (Part VII). Printed for the Trustees of the British Museum, London, 213 pp.
  • Walker F (1873b) Catalogue of the specimens of Hemiptera Heteroptera in the Collection of the British Museum (Part VIII). Printed for the Trustees of the British Museum, London, 220 pp.
  • Wygodzinsky P (1949) Elenco sistematico de los reduviiformes americanos. Instituto de Medicina Regional de la Universidad Nacional de Tucumán. Monografia 1: 1–102.
  • Zhang G, Hart ER, Weirauch C (2016) A taxonomic monograph of the assassin bug genus Zelus Fabricius (Hemiptera: Reduviidae): 71 species based on 10,000 specimens. https://doi.org/10.3897/BDJ.4.e8150
  • Zhao P, Ren S, Wang B, Cai W (2015) A new species of the genus Sphedanolestes Stål, 1866 (Hemiptera: Reduviidae: Harpactorinae) from China, with a key to Chinese species. Zootaxa 3985(4): 591–599. https://doi.org/10.11646/zootaxa.3985.4.8
login to comment