Research Article
Print
Research Article
A new species of Kodormus Barber, with a redescription of the genus, taxonomic notes, and a key to the species of the genus (Hemiptera, Heteroptera, Reduviidae, Stenopodainae)
expand article infoHélcio R. Gil-Santana, Jean-Michel Bérenger§, Jader Oliveira|
‡ Laboratório de Diptera, Instituto Oswaldo Cruz, Rio de Janeiro, Brazil
§ Aix-Marseille Université, Marseille, France
| Universidade de São Paulo, Sao Paulo, Brazil
¶ Universidade Estadual Paulista “Julio de Mesquita Filho”, Araraquara, Brazil
Open Access

Abstract

Kodormus davidmartinsi sp. nov. is described. Taxonomic notes on the other species of Kodormus Barber, 1930, including the description of their male genitalia, are provided. The record of Kodormus bruneosus Barber, 1930 from Brazil and information about the female of the species are presented for the first time. A redescription of Kodormus and a key for its species are provided. Photographs of the holotypes of K. barberi (Costa Lima, 1941), K. bruneosus, and of a paratype of K. oscurus Maldonado & Bérenger, 1996 are presented.

Key words

Male genitalia, Neotropics, Nitornus, Rhyparoclopius, sexual dimorphism

Introduction

Approximately 114 genera belonging to the assassin bug subfamily Stenopodainae (Hemiptera: Heteroptera: Reduviidae) have been described, with the majority of them inhabiting the tropics (Gil-Santana and Oliveira 2016; Schuh and Weirauch 2020). Their diversity is greatest in Africa and America (Giacchi 1987), particularly in the Amazon basin of South America (Bérenger 2001). Currently twenty genera are recognized as valid in the New World, which are separated in the key presented by Gil-Santana and Oliveira (2016). The taxonomy, general morphology, and the scarcely available biological data for American Stenopodainae were reviewed by Giacchi (1987).

Many Stenopodainae appear to be closely associated with the soil, often being covered with soil or sand or various types of debris. Most species are known from light trap collect, with males being captured much more commonly than females, and little is known regarding their biology (Giacchi 1987; Bérenger 2001; Schuh and Weirauch 2020). In Kodormus Barber, 1930, only females of K. bruneosus were collected (Barber 1930; this work). Giacchi (1987) stated that the sexual dimorphism is quite developed among species of Stenopodainae, including several characteristics, such as females being larger and more robust than males, while the antennal vestiture of males is more developed and differentiated.

Kodormus currently includes three species: K. barberi (Costa Lima, 1941), K. bruneosus Barber, 1930, and K. oscurus Maldonado & Bérenger, 1996 (Maldonado 1990; Bérenger and Maldonado 1996). In the present paper, K. davidmartinsi sp. nov. from Brazil is described based on two male specimens. Additionally, morphological remarks of the female of a species of Kodormus (K. bruneosus) are given for the first time. Taxonomical notes on K. barberi, K. bruneosus, and K. oscurus, including the description of their male genitalia are provided. Kodormus bruneosus is recorded from Brazil for the first time. A redescription of Kodormus and a key for its species are also furnished.

Materials and methods

The male holotype of Kodormus bruneosus (Figs 51, 52) currently deposited in the National Museum of Natural History (NMNH), Smithsonian Institution, Washington, DC, USA, was directly examined and photographed by the third author. The holotype of Kodormus barberi (Figs 1, 2), deposited in the “Coleção Costa Lima” in the “Coleção Entomológica do Instituto Oswaldo Cruz” (CEIOC), Rio de Janeiro, Brazil; holotype and paratype (Figs 91104) of K. davidmartinsi sp. nov., and non-type specimens of K. barberi (Figs 3, 44–50) and K. bruneosus (Figs 5362, 68, 7385) were examined and their figures produced by the first author. Observations were made using a stereomicroscope (Zeiss Stemi) and a compound microscope (Leica CME). Measurements were made using a micrometer eyepiece. Photographs were taken using digital cameras (Nikon D5200 or D5600 with a Nikon Macro Lens 105). 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 h. The dissected structures were studied and photographed in glycerol and photographed using a digital camera (Sony DSC-W830). Drawings were made using a camera lucida. Images were edited using Adobe Photoshop CS6.

Photographs and scanning electron microscopy (SEM) images of a non-type specimen of K. barberi (Figs 543) and tibial pads of K. bruneosus (Figs 66, 67, 70) were obtained by the third author. The photographs were taken using a stereomicroscope (Leica 205A) with a digital camera. The SEM images were obtained by cleaning the specimen 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).

Non-type specimens of K. bruneosus and two male paratypes of K. oscurus were examined and imaged by the second author. Photographs of their habitus (Figs 86, 87, 105) were taken with a Canon 5D mark II with a Canon macro lens 100; the image comparing pedicels of a female and a male of K. bruneosus (Fig. 88) was taken with a Canon D40 with a Canon macrolens MP-E 65; photos were stacked using combineZ program. Photographs of the genital structure of a female of K. bruneosus (Figs 89, 90) and of a male of K. oscurus (Fig. 108) were taken with a Canon 5D mark II with a Laowa 25 mm f2.8 ultra macro lens, while the images of the phallus and paramere (Figs 106, 107, 109, 110) of the male of K. oscurus with Zeiss Axio Zoom V16 equipment. The SEM images of some structures of K. bruneosus (Figs 63–65, 69, 71, 72) were taken with TM 4000 Plus Hitachi tabletop microscope.

A photograph of a living specimen of K. barberi was taken by Dr. Ricardo Brugnera (Insetos do Brasil Project) (Fig. 4), which can be freely accessed with information about the data where it was found at https://www.inaturalist.org/observations/107772379.

The type specimens of Kodormus davidmartinsi sp. nov. will be deposited as follows: male holotype in the Collection of National Museum of the Federal University of Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ) and 1 male paratype in the “Coleção de Triatomíneos do Instituto Oswaldo Cruz”, Rio de Janeiro, Brazil (CTIOC) of the “Laboratório Nacional e Internacional de Referência em Taxonomia de Triatomíneos” (LNIRTT) at Oswaldo Cruz Institute, Rio de Janeiro, Brazil.

The general morphological terminology used mainly follows Giacchi (1987) and Schuh and Weirauch (2020). The [visible] segments of the labium are numbered as II to IV, given that the first segment is lost or fused to the head capsule in Reduviidae (Weirauch 2008; Schuh et al. 2009). The terminology applied to the male genital characteristics mainly follows Lent and Wygodzinsky (1979), Gil-Santana (2012), and Gil-Santana and Oliveira (2016).

Additional acronyms of the depository collections, not mentioned above, are the following:

J-MB Jean-Michel Bérenger private collection, France;

MNHN Museum national d’Histoire naturelle, Paris, France;

RBINS Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Belgium;

SEAG Société Entomologique Antilles-Guyane, Guyane.

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 ([]).

Taxonomic account

Kodormus Barber, 1930

Kodormus Barber, 1930: 151 [key], 213–214 [description]; Costa Lima 1940: 166, footnote [Kodormus considered identical to Otiodactylus Pinto, 1927]; Costa Lima 1941: 337–338 [Kodormus recognized as diverse from Otiodactylus but alleged as possessing the same characteristics of Ocrioessa Bergroth, 1918]; Costa Lima and Campos Seabra 1944: 507 [Kodormus recognized as distinct from Ocrioessa], 510 [key]; Costa Lima and Campos Seabra 1945: 159 [checklist]; Wygodzinsky 1949: 66 [catalog]; Putshkov and Putshkov 1985: 104 [catalog]; Giacchi 1985: 68–69 [redescription]; Maldonado 1990: 506 [catalog]; Wygodzinsky and Giacchi 1994: 7 [key], 8 [checklist]; Bérenger and Maldonado 1996: 37 [comparison with other genera]; Froeschner 1999: 227 [catalog]; Forero 2004: 166 [diagnosis], 167 [new record from Colombia], 191 [key]; Gil-Santana et al. 2015: 337 [citation], 341 [key]; Gil-Santana and Oliveira 2016: 501, 502 [citations], 505 [key].

Type species

Kodormus bruneosus Barber, 1930: 214–216, by original designation.

Diagnosis

This genus can be separated from other genera of the New World by the following set of characters. Body somewhat elongated, ~ 2–3× as long as maximum width, slightly flattened dorsoventrally. Head large, anteocular portion longer than postocular; antennal scape shorter than anteocular portion; eyes prominent, shortly setose; labial segment II [first visible] shorter than the others combined; postocular region broad; ramose setigerous processes posterolaterally behind eyes; anterior lobe of pronotum with anterior angles prominent, anterior and lateral margins covered with a row of setigerous tubercules, and a pair of tubercles on its disc; pronotum wider across humeri than along midline; humeral angles protruding; prosternum behind coxae shorter than length of coxae; evaporatory area of metapleuron large, sooty black; fore femora strongly incrassate, robust, at least twice as thick as middle and hind femora; fore and hind tibiae curved, small tibial pads on apices of fore and middle tibia. Abdomen broad, with a more or less expanded connexival margins, which are denticulate and/or lobulated at posterolateral angles of segments II–VI; in male, posterior margin of the abdominal segment VII almost or completely covering the pygophore in dorsal view and with a slightly bilobate shape; in the (known) females, the genital area is visible from above and conical.

Redescription

Body somewhat elongated, ~ 2–3× as long as maximum width, slightly flattened dorsoventrally. General color pale to dark brownish with darkened and pale portions; a clear, generally whitish rounded or subrounded spot above the approximately mid-portion of the outer cell of the membrane of hemelytra. Integument dull, body and legs, except tarsi, generally covered with short, rounded tubercles, each with a short pale apical scale-like seta (setigerous tubercles), scale-like setae, and on some areas, simple setae too. Some glabrous areas, such as the interocular sulcus or forming lines on head, thoracic sterna and femora, subrounded to irregular areas on anterior lobe of pronotum, pleura and abdomen. The integument is generally rugous where there are setigerous tubercles and smooth in the glabrous portions. Simple erect or curved setae are present on labium, antennal segments II–IV, fore tibiae and tarsi. Head subcylindrical; a little longer than wide; shorter than pronotum; anteocular region ~ 2× longer than postocular region, the latter wider than the former. Mandibular plates (jugae sensu Barber 1930; Costa Lima 1941; tylus sensu Giacchi 1985) prominent, divergent, tapering. A small lateral protuberance on antenniferous with setigerous seta(e). Antenna inserted far from eye, somewhat anterior to middle point of anteocular portion, laterally; scape thickened at distal 2/3, somewhat curved at middle 1/3, shorter than anteocular region and covered with pale scale-like setae; pedicel, longer than other segments, > 2× longer and slenderer than the scape, straight at basal 1/2, somewhat curved at middle portion or distal 1/2 and slightly thickened apically; in male, with very numerous, pale to whitish, thin, long, erect to somewhat curved setae, forming a dense pubescence covering almost all the segment, except on anterodorsal surface, where these setae are scarcer and there are 2–4 irregular rows of sparse stout darkened and stiff long setae, in which one or two rows are composed by setae serrate at their distal portion, while the setae of the other rows are uniform; at distal portion of the pedicel the stout setae are less numerous and serrate setae are absent; apex covered by shorter curved pale setae. In the female, the pedicel has sparse scale-like setae and, at the distal portion, scattered straight or somewhat curved short pale setae. Flagellomeres much thinner, cylindrical, straight, subequal in length, each a little shorter than the scape, with few scattered long, erect, stouter setae and a pubescence of simple setae which is formed by thin, short to moderately longer setae on basiflagellomere and generally shorter and even thinner setae on distiflagellomere; the apex of the latter acutely pointed. Clypeus depressed, with a pair of more developed setigerous tubercles. Eyes globose, rounded in dorsal view; suboval in lateral view, extending somewhat on the lower surface of the head, with sparse scale-like setae among facets. Transverse sulcus not very deep, somewhat curved; more sinuous at lateral portions, reaching eyes at their inner posterior angle. Ocelli moderately large, prominently elevated, each ocellus separate from the other for a distance wider than the width of each of them. Labial segment II [first visible] thicker and shorter than the others combined, reaching approximately level of anterior portion of eye; its length subequal to that of the segment III; the latter thinner toward apex; segment IV slender, ~ 1/2 as long as segment III, tapering; its apex reaching stridulatory sulcus on approximately its middle 1/3. Postocular region of the head converging behind eyes to neck, rounded on dorsal view, with one or two conspicuous ramose setigerous processes posterolaterally at each side; above the latter, between eyes and posterior margin, a serial line of somewhat more developed setigerous tubercles. On ventral surface of head, 4–16 large conspicuous setigerous tubercles, generally grouped by transverse pairs, but sometimes, besides some pairs, an isolated tubercle is present at only one side. Two or more of these tubercles generally lie anteriorly to the eyes, and the more posterior pair lies between eyes, near their posterior margin. While in the most posterior pair, the tubercles are very close to each other or even contiguous; in the other pairs, they are clearly separated from each other. Thorax: pronotum wider than long, with anterolateral angles prominent; a pair of tubercles on disc of fore lobe; pronotum wider across humeri than along midline; humeral angles pointed or more prominent. Anterior collar with a variable number of somewhat more developed setigerous tubercles, which also form single rows on the lateral margins of fore lobe of pronotum and on the antero-lateral margin of propleura. Transverse furrow between fore and hind lobes of pronotum shallow, interrupted laterally by a pair of faint submedian ridges; the latter run on approximately the proximal 1/3 of the hind lobe. Fore lobe with a median very thin and somewhat deep midlongitudinal sulcus on the approximately distal 1/2 of fore lobe; sinuate linear ridges covered with setigerous tubercles, narrow and glabrous areas among them and between the most external ridges; lateral margin covered by a row of setigerous tubercles. Supracoxal lobes not prominent; anterior portion of fore supracoxal lobe with a group of conspicuous tubercles, similar to the ramose setigerous of the head. Scutellum subtriangular, longer than wide, with an erect apical tubercle; metascutum also with a short tubercle. Propleura moderately declivous, reaching ventral side laterally and posteriorly to fore coxae. Meso- and metapleura almost vertical; evaporatory area of metapleura large, sooty black. Anterior prosternal processes moderately elongated and curved downwards at apex. Prosternum behind coxae shorter than length of coxae. Stridulitrum long. Mesosternum flat; metasternum slightly prominent at median portion. Legs: fore coxae close, separated from each other by the prosternum, which surpasses fore coxae, by a short distance; middle coxae inserted somewhat less close to each other than the fore coxae; hind coxae inserted very distant from each other. Coxae with two or three ill-defined glabrous longitudinal lines; large setigerous tubercles on fore coxae, more numerous or only present anteriorly. Trochanters with glabrous areas; fore trochanters with two pairs of spiny tubercles on internal surface. Fore femora fusiform, strongly incrassate, at ≥ 2× thicker than middle and hind femora; ventrally with spiny, relatively small, rounded tubercles, including a basal group of 3–5 elements, a midline row with 5–8 elements and some others close to this row on anterior surface; at apex, a lateroventral pair of conspicuous setigerous processes. Fore and middle femora approximately as long as respective tibiae. Scale-like setae on femora and middle and hind tibiae very numerous and generally longer. Fore tibiae with smaller, less numerous or without tubercles and generally less setose than other tibiae. Tarsi with scattered scale-like setae dorsally and stout, straight or slightly curved setae, more numerous, sometimes forming tufts, ventrally. All femora with glabrous lines, which are larger and more evident on fore femora and straight, thinner, and less evident or partly interrupted on middle and hind femora. On fore femora, a ventral and two dorsal of these glabrous, somewhat shiny, lines are generally present. Mid and hind femora slender, straight, somewhat thickened subdistally, generally with some more developed subapical setigerous processes, ventrally. Fore and hind tibiae curved, middle tibiae slightly curved or sometimes straight; all of them compressed dorsoventrally, except at base, generally with a median shallow narrow longitudinal furrow on each lateral surface, except at base, with small tibial pads on fore and middle legs. Fore tarsi two- or three-segmented; middle and hind tarsi three-segmented. Ratio of tarsal segments (approximately): fore tarsi: 1:3 (when two-segmented) / 1:1.5:2.8 (when three-segmented); middle tarsi: 1:1.4–1.6:1.9–2.0; hind tarsi: 1:1.2–1.5:1.8–2.2. Hemelytra with discal cell closed, although the distal cross vein may be indistinct and the cell seems open; corium generally with sparse small scale-like setae, which are more numerous on lateral portion; membrane glabrous. Abdomen: suboval in shape, flattened; segments gradually widening to apex of segment V, then strongly shortening in the next two segments, towards apex; first tergite narrow, integument with shallow longitudinal ridges; tergites II–VI glabrous at median portion in variable extent; scars of dorsal abdominal gland openings on median anterior margins of tergites IV and V very small; connexival margins prominently denticulate and/or lobulated at posterolateral angles of segments II–VI; progressively larger from segment II to V, the latter, although variably in shape among the species, is always the largest, while that on segment VI has a dimension similar or slightly larger in comparison to the prominences on segments II and III. Sternite II (first visible) < 1/3 as long as the sternite III. Sternites II–VI with a median longitudinal narrow pronounced keel; spiracles on sternites II–VII elliptical, prominent, diagonally oriented in relation to the abdominal margin, approximately at medial point between the intersegmental furrows; their margins darkened, even in individuals in which the surrounding integument is pale. In male, posterior border of segment VII straight or curved at median portion, latero-distal margins curved or acute; eighth sternite slightly sinuous on median portion of posterior margin. In the (known) females, the genital area is visible from above and conical. Male genitalia: Genital capsule only visible in ventral view and when in situ, with parameres visible in posterior view; exposed portion of pygophore sub-rounded, covered with setigerous tubercles with scale-like setae; in dorsal view, between anterior and genital openings, a moderately narrow bridge; laterodorsal margin of pygophore, between the bridge and the insertion of parameres, with numerous variably long erect simple setae. Proctiger subsquared with several long setae on approximately its distal 1/3. Medial process of pygophore only visible via dorsal view, with adjacent sparse erect setae, directed upwards, situated just below the paramere apices, subtriangular, triangular or spiniform in anterior view; straight, elongated, thin, and with apex acute in lateral view. Paramere apices close in resting position; in ventral view only the posterior margins of their apices are visible. Parameres symmetrical, very curved in median portion, with a sclerotized moderately large subapical blunt prominence on internal surface; glabrous on approximately basal 1/3 and generally covered with scale-like setae on the exposed surface and scattered, straight, moderately short to longer, simple, erect, thin setae, which are somewhat more numerous around the subapical prominence. Phallus: articulatory apparatus short, with a short basal plate bridge and somewhat longer basal plate arms; pedicel longer than articulatory apparatus, slightly enlarged towards apex, with deep transverse ridges, curved in lateral view and subrectangular in dorsal and ventral views. Gonopore process slightly sclerotized, broad. Dorsal phallothecal sclerite subrectangular, moderately sclerotized. Struts as a pair of elongated arms, fused distally; subcylindrical in approximately basal 2/3 and somewhat enlarged towards apices, which are rounded. Endosoma formed only by its wall, which is smooth and very wrinkled, distal margin more coarsely rugous and sclerotized, shortly prolonged ventrally by an almost imperceptible fold which leans against the main portion of the endosoma.

Key to the species of Kodormus

1 A single ramose setigerous process posterolaterally behind each eye (Figs 14, 94); fore tarsus with two segments only (Figs 32, 33, 95); connexivum of segment V lobulated at external margin (Figs 1, 3, 4, 39, 91–93, 96, 97) 2
A pair of ramose setigerous processes posterolaterally behind each eye; fore tarsus three-segmented (Fig. 68); connexivum of segment V acutely prominent at external margin (Figs 51, 5355, 73, 74, 86, 105) 3
2 Integument generally covered by more developed and larger tubercles, including dorsal surface of head and legs (Figs 5, 6, 11–14, 17, 26, 28, 29); processes on disc of fore lobe of pronotum, humeral angles and apex of scutellum long and conspicuous (Figs 17–25); latero-distal margins of abdominal segment VII acute (Figs 40, 41) ; medial process of pygophore spiniform in anterior view (Fig. 43) K. barberi (Costa Lima, 1941)
Integument with scattered small tubercles; processes on disc of fore lobe, humeral angles and apex of scutellum short (Fig. 92); latero-distal margins of abdominal segment VII rounded (Fig. 96); medial process of pygophore subtriangular in anterior view (Fig. 100) K. davidmartinsi sp. nov.
3 General coloration brownish; latero-distal angles of connexivum less prominent (Figs 51, 5355) K. bruneosus Barber, 1930
General coloration much darker than in preceding species; latero-distal angles of connexivum more prominent (Fig. 105) K. oscurus Maldonado & Bérenger, 1996

Kodormus barberi (Costa Lima, 1941)

Figs 1–4, 5–10, 11–16, 17–26, 27–33, 34–41, 42–50

Ocrioessa barberi Costa Lima, 1941: 339–341, figs 2, 5–6; Rodrigues et al. 2017: 188 [catalog of type specimens; present combination cited], fig. 75 [holotype, dorsal view], table 1 [citation; present combination cited].

Kodormus barberi; Costa Lima and Campos Seabra 1944: 507 [new combination]; Costa Lima and Campos Seabra 1945: 159 [checklist; new combination reinforced]; Wygodzinsky 1949: 66 [catalog]; Maldonado 1990: 506 [catalog]; Bérenger and Maldonado 1996: 35 [citation], figs 9, 37 [distinguishing features]; Gil-Santana and Alencar 2001: 173 [checklist; as a misidentification of K. davidmartinsi sp. nov.; see below]; Forero 2004: 166 [citation].

Notes

Costa Lima (1941) described Ocrioessa barberi based on a male holotype (Figs 1, 2) and a male paratype, both from southeast Brazil (States of Rio de Janeiro and São Paulo, respectively). It is noteworthy that Costa Lima (1941) argued that Kodormus bruneosus should belong to Ocrioessa, while O. barberi would be extremely close to this species. He additionally stated that O. barberi could not be subsumed to the other two species of Ocrioessa because both presented pads in fore tibiae (Barber 1930) while the latter were absent in fore tibiae of Kodormus, accordingly with its description (Barber 1930). Despite these statements, Costa Lima (1941) did not propose any formal synonym between Ocrioessa and Kodormus. Costa Lima and Campos Seabra (1944), however, concluded that Kodormus was really distinct from Ocrioessa, establishing the new combination, Kodormus barberi, which was reinforced by Costa Lima and Campos Seabra (1945).

Figures 1–4. 

Kodormus barberi (Costa Lima, 1941) 1, 2 male holotype deposited in CEIOC 1 dorsal view 2 labels 3, 4 non-type specimen, males, dorsal view 4 living specimen. Scale bars: 5.0 mm (1, 3).

Type material examined

Ocrioessa barberi Costa Lima, 1941. Brazil: Male holotype: [printed label] ANGRA – JUSSARAL / TRAV. [= Travassos] & ALMEIDA [leg.] / 8 [handwritten] 4 -[1]935 // [framed typewritten label] 4522 // [framed printed label] HOLOTYPO (CEIOC).

Additional specimens

Kodormus barberi (Costa Lima, 1941). Brazil: Rio de Janeiro: Kodormus / barberi / (Costa Lima) [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // J. F. Zikán [printed vertically at left side] / Itatiaya [printed] 700 m [handwritten] / [printed] E. [State of] Rio [de Janeiro] – Brasil [Brazil] / [handwritten] 15.-X.-1935 Z. [?] // [framed printed label] CTIOC / N°. 855, 1 male; Kodormus / barberi / (Costa Lima) [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // J. F. Zikán [printed vertically at left side] / Itatiaya [printed] 700 m [handwritten] / [printed] E. [State of] Rio [de Janeiro] – Brasil [Brazil] / [handwritten] 13.-IX.-1941 Z. [?] // [printed label] Coleção [Collection] J. F. Zikan // [framed printed label] CTIOC / N°. 856, 1 male; São Paulo: [printed label] SALESÓPOLIS (BORACÉA) / S. PAULO – 24–IX–[1]946 / TRAVASSOS & VENTEL [leg.] // [framed printed label] CTIOC / N°. 851, 1 male; Kodormus / barberi / (Costa Lima) [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // [printed label] SALESÓPOLIS (BORACÉA) / S. PAULO – 24–9–[1]946 / TRAVASSOS &VANSOLINI [sic], [leg.] // [printed label] Instituto Osvaldo Cruz // [handwritten label] desenhado [drawn] // [framed printed label] CTIOC / N°. 852, 1 male; [printed label] SALESÓPOLIS (BORACÉA) / S. PAULO – 24–9–[1]946 / TRAVASSOS &VANSOLINI [sic], [leg.] // [framed printed label] CTIOC / N°. 853, 1 male; Kodormus / barberi / (Costa Lima) [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // [printed label] SALESÓPOLIS (BORACÉA) / S. PAULO – 24–9–[1]946 / TRAVASSOS &VANSOLINI [sic], [leg.] // [framed printed label] CTIOC / N°. 854, 1 male (CTIOC).

Diagnosis

Kodormus barberi can be separated from other species of the genus by the more developed and larger integumental setigerous tubercles, longer and more conspicuous processes on the disc of fore lobe of pronotum, humeral angles, scutellum and acute latero-distal margins of abdominal segment VII.

Description

Male. Figs 1, 350. Total length: 17.0–19.0 mm; maximum width of abdomen (between apices of connexival prominences of segment V): 5.5–6.8 mm. Coloration (Figs 1, 35, 11, 17, 20, 21, 24, 26, 34, 36, 41): generally brownish; scattered ill-defined and variable darkened and pale markings or portions along the body and legs; pedicel variably paler with apex darkened; apices of femora pale, more extensively on fore femora; apices of prominences of humeri, scutellum and metascutum paler; connexivum paler with prominences darkened; pale portions on fore femora and connexivum sometimes with a greenish to a yellowish tinge. Structure and vestiture: Dorsal surface of head with several large setigerous tubercles (Figs 5, 6, 11–14). Postocular region of the head with only one posterolateral ramose setigerous process at each side (Figs 5, 6, 9, 10, 14). Tubercles on disc of fore lobe, elevated, thick and spiniform (Fig. 11). Humeral angle with an elongated and thick process (Figs 17, 20–22). Process of scutellum moderately elongated (23–25). Coxae, femora and tibiae (except fore tibiae) generally covered by numerous large setigerous tubercles (Figs 2629, 34, 36). Middle tibiae slightly curved; straight in some individuals (Figs 1, 3, 34). Fore tarsi bi-segmented; the second segment ~ 3× as long as the first segment (Figs 32, 33). Hemelytra with distal cross vein variably distinct or not distinct; membrane of hemelytra varying from not reaching to slightly surpassing apex of abdomen (Figs 1, 3, 4). Connexival margins prominently lobulated at posterolateral angles of segments II–VI; short, but progressively larger from segment II to V, the latter, although variably in shape among the specimens, is always the largest, while that on segment VI has a dimension similar or slightly larger in comparison to the prominences on segments II–III (Figs 1, 3, 4, 39). Lateroapical margins of last abdominal segment prominent, acute or faintly curved (Figs 1, 3, 4, 40, 41). Male genitalia (Figs 4150): medial process of pygophore small, straight, spiniform in anterior view (Fig. 43).

Figures 5–10. 

Kodormus barberi, male, head 5–8 dorsal view 5 antennae, except right scape, excluded, the arrow indicates a posterolateral ramose setigerous process 6–8 antennae excluded 7, 8 anteocular portion 7 the arrow points to a mandibular plate 8 antenniferous 9, 10 posterolateral ramose setigerous process on postocular portion, pointed by an arrow 9 ventral view 10 dorsoposterior view. Scale bars: 2.0 mm (5); 0.5 mm (6, 9); 0.1 mm (7, 8, 10).

Figures 11–16. 

Kodormus barberi, male 11, 12 head and anterior portion of pronotum 11 laterodorsal view, the arrow indicates a tubercle of fore lobe of pronotum 12 lateral view, the arrow indicates an anterior prosternal process 13, 14 portions of head 13 anteocular portion, lateral view, the arrow indicates a mandibular plate 14 postocular portion, dorsolateral view, the arrow indicates a posterolateral ramose setigerous process 15, 16 antennal segments, lateral view 15 scape and basal portion of pedicel 16 scape and pedicel. Scale bars: 2.0 mm (11); 0.5 mm (12, 16); 0.2 mm (13–15).

Figures 17–26. 

Kodormus barberi, male 17–19 pronotum 17, 18 dorsal view 18 except humeral angles 19 central portion, lateral view, the arrow points to a tubercle of fore lobe 20–22 humeral prominences 20, 21 ventral view 22 dorsal view 23–25 scutellum and tubercle of metascutum 23 dorsal view 24, 25 lateral view 26 fore leg, lateral view. Abbreviations: s: apical tubercle of scutellum; m: tubercle of metascutum. Scale bars: 2.0 mm (17, 26); 1.0 mm (24); 0.5 mm (18–23, 25).

Figures 27–33. 

Kodormus barberi, male, fore leg portions, lateral view 27 coxa and basal portion of trochanter 28 trochanter and basal portion of femur 29 middle and distal portion of femur, arrows point to ventral spiny rounded tubercles 30 apex of tibia and basal portion of tarsus 31 apex of tibia, the arrows point to the apical pad 32, 33 tarsus. Scale bars: 0.5 mm (27–29); 0.2 mm (32); 0.1 mm (30, 31, 33).

Figures 34–41. 

Kodormus barberi, male 34–36 lateral view 34 middle leg 35 middle tarsus 36 hind leg 37 hind tarsus, lateroventral view 38–41 abdomen portions 38–40 dorsal view 38 tergite I, median portions of tergite II and basal half of tergite III 39 process of the connexival segment V and lateral portion of tergites V and VI 40, 41 apex of abdomen 41 ventral view. Scale bars: 5.0 mm (34, 36); 1.0 mm (41); 0.5 mm (37–40); 0.2 mm (35).

Figures 42–50. 

Kodormus barberi, male genitalia 42 paramere, inner view 43 medial process of pygophore, anterior view 44 phallus, lateral view 45, 46 dorsal view 45 articulatory apparatus 46, 47 pedicel 47 ventral view 48 dorsal phallothecal sclerite, struts and endosoma 49 struts 50 endosoma. Abbreviations: ba: basal plate arm; bb: basal plate bridge; dm: distal margin of endosoma; ds: dorsal phallothecal sclerite; ed: endosoma; pd: pedicel; st: struts. Scale bars: 0.3 mm (44); 0.2 mm (45–50); 0.1 mm (42); 0.03 mm (43).

Comments

In the original description of K. barberi, Costa Lima (1941) recorded the pads on fore tibiae as being absent. However, the examination of the holotype as well as additional non-type specimens allowed confirmation that a small pad is present on the fore and middle tibiae in this species (Fig. 31), while the fore tarsi are revealed to be bi-segmented (Figs 32, 33).

Distribution

Brazil (States of Rio Grande do Sul, Rio de Janeiro and São Paulo) (Costa Lima 1941; this work; Insetos do Brasil Project).

Kodormus bruneosus Barber, 1930

Figs 51–54, 55–62, 63–68, 69–74, 75–85, 86–90

Kodormus bruneosus Barber, 1930: 214–216 [description]; Costa Lima 1940: 166, footnote [Kodormus bruneosus considered as being possibly identical to Otiodactylus signatus Pinto, 1927]; Costa Lima 1941: 337–338 [K. bruneosus very different from Otiodactylus signatus; should be included in Ocrioessa]; Wygodzinsky 1949: 66 [catalog]; Villiers 1971: 684 [misspelled as “brunneosus”; recorded from French Guiana]; Giacchi 1985: 69 [redescription of the male]; Maldonado 1990: 506 [catalog]; Bérenger and Maldonado 1996: 35 [citation], fig. 8, 37 [distinguishing features]; Froeschner 1999: 227 [catalog]; Forero 2004: 166–167, fig. 5.108 [citation, new record from Colombia]; Forero 2006: 36, fig. 58 [new record from Colombia]; Gil-Santana and Husemann 2023: 407, fig. 26 [new records from Ecuador and Peru].

Notes

Kodormus bruneosus was described based on three specimens: a male “Type” (Figs 51, 52), and as “Paratype”, a female and an additional male (Barber 1930). The use of the term “Paratype” in the singular must have been a typo. He probably meant to state both specimens as paratypes as he did in several other species described in the same paper. On the other hand, the male designated by him as the “Type” is regarded here as a holotype, following the Art. 73.1.1 of the International Code of Zoological Nomenclature (ICZN 1999), which defines that if an author states in the original publication that one specimen and only one is “the type” or uses some equivalent expression, that specimen is the holotype fixed by original designation.

Figures 51–54. 

Kodormus bruneosus Barber, 1930 51, 52 male holotype deposited in NMNH 51 dorsal view 52 labels and left hind leg glued to a card pinned with the specimen 53, 54 non-type specimens, males, dorsal view 53 specimen from Ecuador 54 specimen from Brazil. Scale bars: 5.0 mm (51, 53, 54).

Type material examined

Kodormus bruneosus Barber, 1930. Male holotype: Panama: [printed label] USNM_ENT, QR CODE / UCR_ENT 00007958 // [framed label] [handwritten] Kodormus / bruneosus / ♂ Barber / [printed] Det H G Barber // [almost completely printed label] CabimaPan [Cabima, Panama] / May 26 [handwritten].11 / AugustBusck // [red label] [almost completely printed label] TypeNo. / 43376 [handwritten] / U.S.N.M. (NMNH).

Additional specimens

Brazil: Maranhão: Balsas, 08°48'41"S, 46°21'49"W, x.1996, leg. M. Eklein, 1 male; Feira Nova do Maranhão, Retiro, 07°00'31"S, 46°26'41"W, 29–30.xi.1995, leg. M. Eklein, 1 male; Mato Grosso, Diamantino, Alto Rio Arinos, 14°25'S, 56°29'W, 30.iv.2002, E. Furtado, leg., 1 male; Pará: Kodormus / bruneosus / Barber [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // [printed label] Cachimbo E. [state of] Pará / Travassos-Oliveira / & Adão [leg.], 25/9-10-[1]956 // [framed printed label] CTIOC / N°. 847, 1 male; Kodormus / bruneosus / Barber [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // [printed label] Cachimbo E. [state of] Pará / Travassos-Oliveira / & Adão [leg.], 25/9-10-[1]956 // [framed printed label] CTIOC / N°. 848, 1 male; Kodormus / bruneosus / Baber [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // [printed label] Cachimbo E. [state of] Pará / Travassos-Oliveira / & Adão [leg.], 25/9-10-[1]956 // [printed label] Instituto Osvaldo Cruz // [handwritten label] desenhado [drawn] // [framed printed label] CTIOC / N°. 849, 1 male; Kodormus / bruneosus / Barber [handwritten] / Wygodzinsky det. [printed] ’64 [handwritten] // [handwritten label] Belém, Pará / M. Alvarenga / 1-1956 // [printed label] Instituto Osvaldo Cruz //[framed printed label] CTIOC / N°. 850, 1 male (CTIOC). Ecuador: Narupa, Napo Province, 1.200 m, 12.ii.1996, Juán Salvador leg., 1 male (MNRJ). French Guiana: Itoupé, DZ 570 m, 9.iii.2010, light trap, SEAG leg. 2 males and one female; N2, pk 79, 7.i.1996, PL, B. Hermier leg., 1 male; Degrad Corrèze, Route de régina, pk 62, 19.xii.1998, Kindl leg., 1 male; Degrad Kwata, iii.1995, PL, vesco JP leg., 1 male; Barrage petit Saut, 2.iii.1993, J–MB réc, 1 male; Grand Santi, PL, 29.iv.2000, P. Causse leg., 1 male; Laussat, PL, 11.ix.2010, light trap, SEAG leg., 1 female; D6, pk 37, 01.i.1998, light trap, B. Hermier leg., 1 female; Montagne des chevaux, PL, 22.xii.2008, light trap, SEAG leg., 1 female (J–MB). Peru: [red label with a smaller white label glued on it; both printed labels] Coll. R. I. Sc. N. B. [underlined by a black line] / Pérou [on the smaller white label] Peru 700 m / Chanchamayo / 20.X.1960 // [printed label] Kodormus / brunneus [sic] / Barber / JMaldonadoC.85 [1985] // Kodormus [printed] / bruneosus [handwritten] / Gil-Santana det. [printed] 19 [handwritten; 2019], 1 male (RBINS).

Diagnosis

Kodormus bruneosus may be separated from most of the other species of the genus by the denticulate latero-distal angles of connexival segments II–VI and from K. oscurus, which although has a somewhat similar connexival structure, presents more prominent connexival latero-distal angles, and by their general coloration, which is generally brownish in K. bruneosus and darker in K. oscurus.

Description

Male (Figs 5185). Total length 17.5–21.5 mm; maximum width of abdomen (between apices of connexival prominences of segment V): 5.5–8.8 mm. Coloration (Figs 51, 5355, 73, 74): generally brownish; in some individuals with some portions more darkened such as the fore lobe of pronotum, legs, prominences of connexivum, and ventral surface of abdomen. Antennal pedicel variably paler with apex darkened. Pale markings or portions variably scattered on head, apices of femora, basal portions of tibiae and sternites; the latter sometimes almost or completely paler. Structure and vestiture (Figs 51, 5374): Postocular region of the head with two ramose setigerous processes posterolaterally on each side, very close to each other, the most posterior one slightly above of the other. Setigerous tubercles on serial line of postocular region of head, anterior collar and single rows on the lateral margins of fore lobe of pronotum variable in size and coloration among individuals, larger and pale to whitish or smaller and darker. Tubercles on disc of fore lobe flat, rounded. Humeral angle short, spiniform (Figs 51, 5355). Process of scutellum short. Membrane of hemelytra varying from not reaching to slightly surpassing apex of abdomen (Figs 51, 5355). Fore tarsus three-segmented (Fig. 68). Lateroapical margins of connexivum more or less prominent among individuals; that on segment V is sometimes apically curved downward (Figs 51, 5355, 73, 74). Male genitalia (Figs 7485): medial process of pygophore enlarged; triangular in anterior view (Fig. 77).

Figures 55–62. 

Kodormus bruneosus, male 55 specimen from Brazil, dorsal view 56–62 antennal segments or portions, lateral view 56 scape 57–60 pedicel 57 basal portion 58, 59 middle portion 59 somewhat distally 60 apical portion 61 basiflagellomere 62 distiflagellomere Scale bars: 5.0 mm (55); 0.2 mm (56–62).

Figures 63–68. 

Kodormus bruneosus, male 63–65 fore femur, lateral view 63, 64 anterior surface 63 median portion, arrows point to ventral spiny rounded tubercles 64 portion of the integument 65 a spiny ventral rounded tubercle 66, 67 fore tibia, ventral view 66 apical pad 67 tenent hairs of a portion of the pad 68 apex of fore tibia and tarsus, lateral view, tibial pad pointed by arrows. Scale bars: 0.5 mm (63); 0.2 mm (68); 0.1 mm (64–66); 0.01 mm (67).

Figures 69–74. 

Kodormus bruneosus, male 69, 70 apex of middle tibia 69 lateral view, the arrows point to the distal pad 70 tibial pad, ventral view 71, 72 lateral view 71 middle tarsus 72 hind tarsus 73, 74 abdomen 73 dorsal view 74 ventral view. Scale bars: 2.0 mm (73, 74); 0.5 mm (71, 72); 0.1 mm (69, 70).

Figures 75–85. 

Kodormus bruneosus, male genitalia 75 genital capsule, dorsal view 76 left paramere, inner view 77 medial process of pygophore, anterior view 78–80 phallus 78, 79 lateral view 80 ventral view 81–85 dorsal view 81 articulatory apparatus and basal portion of pedicel 82 basal portion of basal plate arms, pedicel and basal portion of phallothecal sclerite and struts 83 dorsal phallothecal sclerite, struts and endosoma 84 dorsal phallothecal sclerite and struts (endosoma extracted) 85 endosoma. Abbreviations: ba: basal plate arm; bb: basal plate bridge; br: bridge; dm: distal margin of endosoma; ds: dorsal phallothecal sclerite; ed: endosoma; pa: paramere; pd: pedicel; pt: proctiger; py: pygophore; st: struts; vf: ventral fold of endosoma Scale bars: 0.5 mm (75); 0.3 mm (76, 78, 79); 0.2 mm (77); 0.1 mm (80–85).

Female (Figs 86–90): Total length: 22–23 mm; maximum width of abdomen between apices of connexival prominences of segment V: 9–11 mm. Similar to male in general (Figs 86, 87). Antennal pedicel with scattered very short and sparse scale-like setae, and a few thin, long setae distally (Fig. 88, A). Abdomen very wide, with a maximum width on segment V (Figs 86, 87). Membrane of hemelytra not reaching apex of abdomen (Fig. 86); genital area visible from above, cone shaped and acute (Fig. 86). Female genitalia: external genitalia as in Figs 89, 90.

Figures 86–90. 

Kodormus bruneosus 86, 87 female 86 dorsal view 87 ventral view 88 antennal pedicels of a female (A) and of a male (B), lateral views 89, 90 female genitalia, external view 89 posterior view 90 ventral view. Scale bars: 5.0 mm (86, 87); 1.0 mm (88); 0.5 mm (89, 90).

Comments

Barber (1930) recorded the tibial pad as absent at the apex of fore tibia in K. bruneosus. Giacchi (1985), when redescribed the male of this species, did not mention the presence or absence of pads on the tibiae. However, we have recorded the presence of tibial pads at apices of fore and middle tibiae in all specimens of K. bruneosus studied here (Figs 6670). Our observation is in accordance with Weirauch (2007) who also recorded tibial pads (as fossula spongiosa) as present both in fore and middle tibiae of K. bruneosus.

The description of K. bruneosus by Barber (1930) seems to have been based only on the male type (s), because no detail was given concerning the female cited as “Paratype”. There was no mention about differences between sexes and neither about the genital portions. While Barber (1930) recorded the pedicel as densely setose, as seen in males (Figs 57–60), he did not mention that, accordingly with the females examined here, it is remarkably less setose in the females (Fig. 88, A). Additionally, the females were generally larger, with wider abdomens (Figs 86, 87).

When recording K. bruneosus from Colombia, Forero (2006) listed Brazil as a country of occurrence of the species too. However, this supposed record was based on Wygodzinsky and Giacchi (1994), who actually recorded only Kodormus from Brazil, not specifying any species of the genus. Their record may possibly have been based on K. barberi, the only species recorded from Brazil so far. This assertion was confirmed to the first author (HRG-S) by D. Forero (pers. inform.). Therefore, the first proven record of K. bruneosus from this country is provided here.

Distribution

Panama, Guyana (Barber 1930), French Guiana (Villiers 1971), Trinidad and Tobago, Venezuela (Giacchi 1985), Bolivia (Maldonado 1990), Colombia (Forero 2004, 2006), Ecuador and Peru (Gil-Santana and Husemann 2023).

New record

Brazil (States of Maranhão, Mato Grosso and Pará).

Kodormus davidmartinsi sp. nov.

Figs 91–95, 96–104

Kodormus barberi; Gil-Santana and Alencar 2001: 173 [checklist; misidentification].

Notes

Gil-Santana and Alencar (2001) based on a male specimen from a Natural Reserve in Linhares, Espírito Santo State, Brazil, included Kodormus barberi in a checklist of Reduviidae of this locality. However, a re-examination of the specimen from Linhares made it clear that it belongs to the new species, K. davidmartinsi sp. nov., with the designation of this specimen as the holotype. An additional specimen from the same locality was included as a paratype.

Type material examined

Brazil, Espírito Santo: Linhares, Reserva Natural Vale, 19°09'S, 40°04'W, José Simplício dos Santos leg., male holotype, xi.1990 (MNRJ); same locality and collector, 1 male paratype, 11.xii.1987, CTIOC n° 13832 (CTIOC).

Diagnosis

Kodormus davidmartinsi sp. nov. and K. barberi may be separated from other species of the genus by the presence of connexival margins of segments III–V lobulated. These species may be separated from each other by the larger lobulated portion of connexival segment V in K. davidmartinsi sp. nov. Additionally, K. davidmartinsi sp. nov. has smaller integumental setigerous spiniferous processes, shorter processes of disc of fore lobe of pronotum, humeral angles, scutellum and rounded latero-distal margins of abdominal segment VII. In male genitalia, the medial process of pygophore in anterior view, is subtriangular in K. davidmartinsi sp. nov. and spiniform in K. barberi.

Description

Male. Figs 91104. Measurements (mm) (holotype / paratype): Total length: 20.5 / 21.5; head: total length (excluding neck, lateral view): 3.2 / 3.1; maximum width across eyes: 2.4 / 2.7; length of anteocular portion: 1.6 / 1.6; length of postocular portion: 0.8 / 0.7; interocular space (synthlipsis): 1.0 / 1.1; transverse width of right eye: 0.7 / 0.8; length of right eye: 0.8 / 0.9; lengths of antennal segments: scape: 1.3 / 1.4; pedicel: 3.0 / 3.0; basiflagellomere 0.8 / 0.6; distiflagellomere: 0.8 [approx.; very curved] / absent; lengths of labial segments: II [first visible]: 1.6 / 1.7; III: 1.4 / 1.6; IV: 0.9 / 0.9. Thorax: pronotum: length of fore lobe (at midline): 1.9 / 1.9; length of hind lobe (at midline): 1.8 / 1.8; width at posterior margin: 4.7 / 5.0. Fore legs: length of femur: 5.4 / 5.6; maximum width of femur at mid portion: 1.3 / 1.4; length of tibia: 5.2 / 5.4; length of pad: 0.1 / 0.2; length of tarsus: 0.7 / 0.8; middle legs: length of femur: 6.5 / 7.2; maximum width of femur at mid portion: 0.45 / 0.45; length of tibia: 6.4 / 7.5; length of pad: 0.2 / 0.2; length of tarsus: 1.5 / 1.5; hind legs: length of femur: 10.0 / 11.5; maximum width of femur at mid portion: 0.5 / 0.5; length of tibia: 12.8 / 13.8; length of tarsus: 1.8 / 1.9. Abdomen: length: 11.2 / 11.7; maximum width (measured between outer margins of anterior portion of sternite V): 6.1 / 6.5; maximum distance between outer margins of lobulated prominence of connexival segment V: 9.2 / 8.8. Coloration (Figs 91–93, 96, 97): generally dark brownish with ill-defined scattered pale portions or markings on head, femora and sternites; hemelytra, except basal portion, paler, more in the paratype. Antenna pale brownish; scape with irregular scattered dark small markings; apex of pedicel dark. The following portions variably paler: glabrous areas of head and fore femora, more intensively in the holotype; inferior margin of fore supracoxal lobe; upper portions of lateral surfaces and apices of femora; a pair of linear markings at basal portions of tibiae; median portion of tergites I–V (Fig. 96), progressively in less extent from the former to the latter; margins of sternites between segments II–VI; on the latter, at each lateral side, a pair of rounded (paratype) or irregular (holotype) small markings on basal margin of these segments and another similar markings between basal and distal margins (Fig. 97). Structure and vestiture (Figs 9197): Postocular region of the head with only one ramose setigerous process posterolaterally at each side (Fig. 94). Processes of scutellum and humeri short. Fore trochanters with two pairs of spiny tubercles on internal surface. Fore femora with a basal group from two to four spiny, relatively small, rounded tubercles, a midline row with eight spiny rounded tubercles and two (holotype) and four (paratype) others close to this row on anterior surface. Fore tarsi bi-segmented; the second segment ~ 3× as long as the first segment (Fig. 95). Connexival margins of segments III–VI lobulated; those on segments III–V have the external margin sinuated at median portion and a short spiny prominence at latero-distal angle in holotype, while in paratype the external margin is faintly curved without spiny prominences (Figs 91–93, 96, 97). Membrane of hemelytra not reaching apex of abdomen (Figs 91–93). Lateroapical margins of abdomen curved, slightly prominent (Fig. 96). Male terminalia (Figs 98–104): medial process of pygophore subtriangular in anterior view (Fig. 100).

Figures 91–95. 

Kodormus davidmartinsi sp. nov., male 91–94 dorsal view 91, 92 holotype 93 paratype 94 postocular portion of the head, the arrow points to a posterolateral ramose setigerous process 95 fore leg, apex of tibia and tarsus, lateral view. Scale bars: 5.0 mm (91–93); 0.5 mm (94); 0.2 mm (95).

Figures 96–104. 

Kodormus davidmartinsi sp. nov., male 96, 97 abdomen of the paratype 96 dorsal view 97, 98 ventral view 98 eighth sternite 99–104 male genitalia 99 pygophore and parameres (proctiger and phallus extracted), dorsal view 100 medial process of pygophore, anterior view 101 phallus, lateral view 102–104 dorsal view 102 pedicel and basal portion of phallothecal sclerite and struts 103 dorsal phallothecal sclerite and struts (endosoma extracted) 104 endosoma. Abbreviations: ba: basal plate arm; br: bridge; dm: distal margin of endosoma; ds: dorsal phallothecal sclerite; ed: endosoma; pa: paramere; pd: pedicel; mp: medial process of pygophore; py: pygophore; st: struts. Scale bars: 2.0 mm (96, 97); 0.5 mm (98, 99); 0.2 mm (101–104); 0.1 mm (100).

Etymology

The new species is named in honor of Dr. David dos Santos Martins, researcher of the Instituto Capixaba de Pesquisa, Assistência Técnica e Extensão Rural (INCAPER), Vitória, Espírito Santo, Brazil, for his great contribution to the knowledge of the entomofauna of the State of Espírito Santo where the new species was found.

Distribution

Brazil (State of Espírito Santo).

Kodormus oscurus Maldonado & Bérenger, 1996

Figs 105–110

Kodormus oscurus Maldonado & Berénger, 1996 in Bérenger and Maldonado 1996: 35–37, figs 1–7 [description]; Forero 2004: 166 [citation].

Notes

Kodormus oscurus was described based on 10 males collected in Bolivia (Bérenger and Maldonado 1996).

Type material examined

Bolivia: Ixiamas, Beni, oct. 1993, 270 m, Bleuzen leg., 1 male paratype; Pucara, Beni, oct. 1993, 750 m, Bleuzen leg., 1 male paratype (J-MB).

Diagnosis

Kodormus oscurus is separated by its general coloration, which is generally dark brown, while in the other species of Kodormus, it is generally brownish. Kodormus oscurus seems closer to K. bruneosus, based on the denticulate latero-distal angles of connexival segments II–VI, but in the latter species these angles are less prominent.

Description

Male. Figs 105–110. Total length: 19.5–25 mm. Coloration (Fig. 105): Generally dark brownish; hemelytra and sternites reddish brown, slightly paler than dark brownish portions. Structure and vestiture (Fig. 105): Postocular region of the head with two ramose setigerous processes posterolaterally at each side, very close to each other, the most posterior one slightly above of the other. Setigerous tubercles on serial line of postocular region of head very numerous, pale and conspicuous; those on the lateral margins of fore lobe of pronotum forming an irregular row, most of them pale. Tubercles on disc of fore lobe flat, rounded. Humeral angle short, spiniform. Process of scutellum short. Membrane of hemelytra varying from not reaching to reaching apex of abdomen. Fore tarsus three-segmented. Lateroapical margins of connexivum conspicuously prominent; those on segments IV and V are apically curved downward in some individuals. Male genitalia (Figs 106–110): medial process of pygophore enlarged; triangular in anterior view (Fig. 108).

Figures 105–110. 

Kodormus oscurus Maldonado & Bérenger, 1996, male paratype 105–107 dorsal view 106–110 male genitalia 106, 107 parameres 106 right 107 left 108 medial process of pygophore, anterodorsal view 109, 110 dorsal view 109 phallus 110 apex of dorsal phallothecal sclerite, struts and endosoma, arrows point to the distal margin of endosoma. Abbreviations: dm: distal margin of endosoma; ds: dorsal phallothecal sclerite; pd: pedicel; st: struts. Scale bars: 5.0 mm (105); 0.5 mm (108, 109); 0.2 mm (106, 107); 0.1 mm (110).

Distribution

Bolivia (Bérenger and Maldonado 1996).

Discussion

Because most species of Stenopodainae are known from light trap collecting, with males being captured much more commonly than females (Giacchi 1987; Bérenger 2001; Schuh and Weirauch 2020), the female of many species remains unknown. In Kodormus, only females of K. bruneosus were collected. Barber (1930) included a female “Paratype” in his description of K. bruneosus, but did not provide any specific comments about it, such as differences in size or in the antennal vestiture, as it commonly occurs among reduviids (Giacchi 1987). In this study, females of K. bruneosus from French Guiana were examined by the second author and shown to be generally larger, with wider abdomens. Additionally, the antennal vestiture of the pedicel was quite diverse, much more differentiated in the males, as described above. Nevertheless, there is a need to obtain females of the other species to a better knowledge of range of variation or sexual dimorphism between males and females of species of Kodormus.

Barber (1930) in his description of Kodormus recorded the pad as absent at the apex of fore tibia. Although the presence or absence of a pad on the fore and/or middle tibiae is an important feature for separating the genera of Stenopodainae, there is no further mention about the presence or absence of the pad on tibiae of Kodormus in the redescription of this genus by Giacchi (1985), and in diagnosis and/or keys in which it was included (Costa Lima and Campos Seabra 1944; Wygodzinsky and Giacchi 1994; Forero 2004; Gil-Santana et al. 2015; Gil-Santana and Oliveira 2016). Only in the morphological comparative study of Weirauch (2007: 159) it was recorded on fore and middle tibiae of K. bruneosus. However, we confirmed the presence of the pad on fore and middle tibiae in all species of Kodormus studied here (e.g., Figs 31, 66, 6870). Therefore, this feature was included in accordance with our observations in the description of the genus presented above. On the other hand, the fore tarsi are bi-segmented in K. barberi and K. davidmartinsi sp. nov. (Figs 32, 33, 95) while in K. bruneosus (Fig. 68) and K. oscurus it is tri-segmented, as usual in reduviids. Among New World Stenopodainae, the bi-segmented fore tarsi are included among the diagnostic characteristics of Rhyparoclopius Stål, 1868 (Forero 2004; Gil-Santana 2012; Gil-Santana et al. 2015; Gil-Santana and Oliveira 2016). Interestingly, when describing Kodormus, Barber (1930) stated that this genus had no close affinity to any other genus of Stenopodainae, but seemed close to Rhyparoclopius in the broader character of the body, while Gil-Santana and Oliveira (2016) highlighted that these two genera were among those which have the fore femora moderately to strongly incrassate (at least twice as thick as the mid and hind femora), and together with Otiodactylus Pinto, 1927, all three have the antennal scape shorter than the length of the anteocular portion of the head. Bérenger and Maldonado (1996) considered that among Neotropical genera with enlargement of the connexival margins, such as Rhyparoclopius and Otiodactylus, the latter seemed to be the closest to Kodormus. Nevertheless, the significance of these similiarities needs a more thorough evaluation, including cladistic studies in order to clarify the systematic relationship among the genera of Stenopodainae.

Several works, mostly authored by J.C. Giacchi, summarized by Gil-Santana et al. (2015), have described the male genitalia of many species of American Stenopodainae. Differences between or among species in the same genus, involving mostly the parameres, the shape of the dorsal phallothecal plate, struts and the processes or sclerotizations of the endosoma, have been recorded (e.g., Giacchi 1969; Gil-Santana 2012). However, in the two species of Nitornus Stål, 1859, which are quite diverse in several morphological features, the male genitalia of both was very similar (Gil-Santana 2016). Among the species of Kodormus examined here, the structure of the male genitalia, including the pygophore, parameres and phallus, was also very similar. The medial process of pygophore was the only structure which was recorded as being slightly different in shape (in its anterior view), more prominently in K. barberi. Thus, it seems that the male genitalia of Kodormus has limited taxonomic utility.

Acknowledgements

The authors are grateful to Talitta Guimarães Simões (NMNH) for the support and providing the equipment to photograph the holotype of K. bruneosus and to Thomas Henry (NMNH) for granting access to this specimen; to Ricardo Brugnera (Insetos do Brasil Project) for allowing us to reproduce his iNaturalist photograph of a live specimen of K. barberi. HRG-S is grateful to Jane Costa, Márcio E. Felix, and Cláudia L. Rodrigues (CEIOC); M. Dolsan de Almeida and Solange Ribeiro Peixoto (CTIOC); and Jérôme Constant and Wouter Dekoninck (RBINS) for providing access to the specimens of their institutions. For donating the respective specimens, HRG-S is grateful to Renato de Jesus (former manager of the Reserva Natural Vale; holotype of K. davidmartinsi sp. nov.), David dos Santos Martins (INCAPER; paratype of K. davidmartinsi sp. nov.), and the late entomologist Eurides Furtado (a non-type specimen of K. bruneosus from Mato Grosso state). The second author thanks Hitachi High-Tech Corporation, from which he received partial support for his work and he is also grateful to SEAG for donating the specimens of K. bruneosus collected by them. The third author thanks the Laboratório de Microscopia Avançada LMA, Instituto de Química, Unesp”, Araraquara, São Paulo, Brazil for allowing the use of the scanning electron microscope facility to obtain the SEM images made by him.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Funding

The third author (JO) extends appreciation to the funding agency São Paulo Research Foundation (FAPESP) under the process number 19/02145-2.

Author contributions

Conceptualization: HRGS. Data curation: JMB, HRGS. Formal analysis: JO, HRGS. Investigation: JMB, HRGS. Methodology: HRGS, JO, JMB. Project administration: HRGS. Supervision: JO. Writing – original draft: HRGS. Writing – review and editing: JO, JMB, HRGS.

Author ORCIDs

Hélcio R. Gil-Santana https://orcid.org/0000-0002-0544-348X

Jean-Michel Bérenger https://orcid.org/0000-0002-8497-2792

Jader Oliveira https://orcid.org/0000-0002-2588-1911

Data availability

All of the data that support the findings of this study are available in the main text.

References

  • Barber HG (1930) Essay on the subfamily Stenopodinae of the New World. Entomologica Americana n.s. 10: 149–238.
  • Bérenger J-M (2001) Hétéroptères Reduviidae nouveaux ou peu connus de Guyane Française. I, Stenopodainae. Bulletin de la Société Entomologique de France 106(1): 5–18. https://doi.org/10.3406/bsef.2001.16722
  • Bérenger J-M, Maldonado CJ (1996) Une nouvelle espèce du genre Néotropical Kodormus Barber [Heteroptera, Reduviidae, Stenopodainae]. Revue Française d’Entomologie (n.s. ) 18: 35–37.
  • Costa Lima AM (1940) Insetos do Brasil. V.2. Hemípteros. Escola Nacional de Agronomia, Rio de Janeiro, Brazil, 351 pp.
  • Costa Lima A (1941) Considerações sobre alguns reduviideos da subfamília “Stenopodinae”. Revista Brasileira de Biologia 1: 337–342.
  • Forero D (2004) Capítulo 5. Diagnosis de los gêneros neotropicales de la família Reduviidae (Hemiptera: Heteroptera), y su distribución em Colombia (excepto Harpactorinae). In: Fernández F, Andrade G, Amat G (Eds) Insectos de Colombia (Vol. 3). Universidade Nacional de Colombia, Bogotá, 128–275.
  • Froeschner RC (1999) True bugs (Heteroptera) of Panama: A synoptic catalog as a contribution to the study of Panamanian biodiversity. Memoirs of the American Entomological Institute 61: 1–393.
  • Giacchi JC (1969) Revision del genero Stenopoda Laporte, 1833 (Hemiptera, Reduviidae, Stenopodainae). Physis 29: 1–26.
  • Giacchi JC (1985) Revision de los stenopodainos americanos. VII. Redescripcion de los generos Pnirontis Stal, 1859; Pnohirmus Stal, 1859; Ctenotrachelus Stal, 1868; Ocrioessa Bergroth, 1918 y Kodormus Barber, 1930 (Heteroptera, Reduviidae). Physis. Secc. C 43: 61–70.
  • Giacchi JC (1987) Aportes a la morfologia y taxonomia de los stenopodainos americanos. (Heteroptera, Reduviidae). II. Consideraciones acerca de la subfamilia. Physis. Secc. C 45: 19–30.
  • Gil-Santana HR (2012) Rhyparoclopius aokiae sp. nov., a remarkable Stenopodainae (Hemiptera: Heteroptera: Reduviidae) from Mato Grosso do Sul, Brazil, with taxonomical notes on other species of Rhyparoclopius Stål. Zootaxa 3478(1): 93–104. https://doi.org/10.11646/zootaxa.3478.1.11
  • Gil-Santana HR, Alencar J (2001) Reduviidae da Reserva Florestal de Linhares, Espírito Santo, Brasil (Hemiptera-Heteroptera). Revista Brasileira de Zoociências 3: 171–176.
  • Gil-Santana HR, Husemann M (2023) New records of Reduviidae (Hemiptera: Heteroptera) from Costa Rica, Ecuador, French Guiana, Paraguay and Peru. Revista Chilena de Entomologia 49(2): 395–412. https://doi.org/10.35249/rche.49.2.23.23
  • Gil-Santana HR, Oliveira J (2016) Pratigi aristeui, a new Neotropical genus and species of Stenopodainae (Hemiptera: Heteroptera: Reduviidae). Acta Entomologica Musei Nationalis Pragae 56: 491–506.
  • 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
  • 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/
  • Lent H, Wygodzinsky P (1979) Revision of the Triatominae (Hemiptera: Reduviidae) and their significance as vectors of Chagas’ disease. Bulletin of the American Museum of Natural History 163: 123–520.
  • Maldonado CJ (1990) Systematic catalogue of the Reduviidae of the World. Caribbean Journal of Science, Special publication No. 1, University of Puerto Rico, Mayagüez, Puerto Rico, 694 pp.
  • Putshkov VG, Putshkov PV (1985) A Catalogue of the Assassin-Bugs Genera of the World (Heteroptera, Reduviidae). Published by the authors, Kiev, 137 pp.
  • Rodrigues JMS, Cordeiro IRS, Moreira FFF (2017) Additions and corrections to the type list of Heteroptera (Insecta: Hemiptera) of the Costa Lima Collection, Oswaldo Cruz Foundation. Zootaxa 4311(2): 183–210. https://doi.org/10.11646/zootaxa.4311.2.2
  • 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): 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.]
  • Villiers A (1971) Reduviidae [Heteroptera] récoltés en Guyane Française par la mission du Muséum national d’Histoire naturelle. Annales de la Société entomologique de France (n.s. ) 7: 679–684. https://doi.org/10.1080/21686351.1971.12277966
  • Weirauch C (2007) Hairy attachment structures in Reduviidae (Cimicomorpha, Heteroptera), with observations on the fossula spongiosa in some other Cimicomorpha. Zoologischer Anzeiger 246(3): 155–175. https://doi.org/10.1016/j.jcz.2007.03.003
  • Wygodzinsky P (1949) Elenco sistematico de los reduviiformes americanos. Instituto de Medicina Regional de la Universidad Nacional de Tucumán. Monografia 1: 1–102.
  • Wygodzinsky P, Giacchi JC (1994) Key to the genera of Stenopodainae of the New World (Insecta, Heteroptera, Reduviidae). Physis Secc. C 49: 5–9.
login to comment