Research Article |
Corresponding author: Hélcio R. Gil-Santana ( helciogil@uol.com.br ) Academic editor: Nikolay Simov
© 2023 Hélcio R. Gil-Santana, Jean-Michel Bérenger, Jader Oliveira.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Gil-Santana HR, Bérenger J-M, Oliveira J (2023) 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). ZooKeys 1181: 265-298. https://doi.org/10.3897/zookeys.1181.108463
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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.
Male genitalia, Neotropics, Nitornus, Rhyparoclopius, sexual dimorphism
Approximately 114 genera belonging to the assassin bug subfamily Stenopodainae (Hemiptera: Heteroptera: Reduviidae) have been described, with the majority of them inhabiting the tropics (
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 (
Kodormus currently includes three species: K. barberi (Costa Lima, 1941), K. bruneosus Barber, 1930, and K. oscurus Maldonado & Bérenger, 1996 (
The male holotype of Kodormus bruneosus (Figs
Photographs and scanning electron microscopy (SEM) images of a non-type specimen of K. barberi (Figs
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
A photograph of a living specimen of K. barberi was taken by Dr. Ricardo Brugnera (Insetos do Brasil Project) (Fig.
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 (
The general morphological terminology used mainly follows
Additional acronyms of the depository collections, not mentioned above, are the following:
J-MB Jean-Michel Bérenger private collection, France;
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 ([]).
Kodormus
Barber, 1930: 151 [key], 213–214 [description];
Kodormus bruneosus Barber, 1930: 214–216, by original designation.
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.
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
1 | A single ramose setigerous process posterolaterally behind each eye (Figs |
2 |
– | A pair of ramose setigerous processes posterolaterally behind each eye; fore tarsus three-segmented (Fig. |
3 |
2 | Integument generally covered by more developed and larger tubercles, including dorsal surface of head and legs (Figs |
K. barberi (Costa Lima, 1941) |
– | Integument with scattered small tubercles; processes on disc of fore lobe, humeral angles and apex of scutellum short (Fig. |
K. davidmartinsi sp. nov. |
3 | General coloration brownish; latero-distal angles of connexivum less prominent (Figs |
K. bruneosus Barber, 1930 |
– | General coloration much darker than in preceding species; latero-distal angles of connexivum more prominent (Fig. |
K. oscurus Maldonado & Bérenger, 1996 |
Ocrioessa barberi
Costa Lima, 1941: 339–341, figs 2, 5–6;
Kodormus barberi; Costa Lima and Campos Seabra 1944: 507 [new combination];
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).
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).
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.
Male. Figs
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).
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).
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).
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).
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).
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).
In the original description of K. barberi,
Brazil (States of Rio Grande do Sul, Rio de Janeiro and São Paulo) (
Kodormus bruneosus
Barber, 1930: 214–216 [description];
Kodormus bruneosus was described based on three specimens: a male “Type” (Figs
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).
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 (
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.
Male (Figs
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).
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).
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).
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
The description of K. bruneosus by
When recording K. bruneosus from Colombia,
Panama, Guyana (
Brazil (States of Maranhão, Mato Grosso and Pará).
Kodormus barberi;
Brazil, Espírito Santo: Linhares, Reserva Natural Vale, 19°09'S, 40°04'W, José Simplício dos Santos leg., male holotype, xi.1990 (
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.
Male. Figs
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).
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).
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.
Brazil (State of Espírito Santo).
Kodormus oscurus
Maldonado & Berénger, 1996 in
Kodormus oscurus was described based on 10 males collected in Bolivia (
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).
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.
Male. Figs
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).
Bolivia (
Because most species of Stenopodainae are known from light trap collecting, with males being captured much more commonly than females (
Several works, mostly authored by J.C. Giacchi, summarized by
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 (
The authors have declared that no competing interests exist.
No ethical statement was reported.
The third author (JO) extends appreciation to the funding agency São Paulo Research Foundation (FAPESP) under the process number 19/02145-2.
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.
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
All of the data that support the findings of this study are available in the main text.