Neomacrocoris handlirschi Montandon
Creators: name: Sites, Robert W.
name: Mbogho, Aaron Y.
Publisher: name: Zenodo
Year: 2012
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Hemiptera
Naucoridae
Neomacrocoris
Neomacrocoris handlirschi

description: Neomacrocoris handlirschi (Montandon)
Figs. 10–14
Macrocoris handlirschi Montandon 1909: Bul. Soc. Rom. Stiinte, 18: 55.
Neomacrocoris handlirschi: Montandon 1913, Bul. Soc. Rom. Stiinte, 22: 332.
Neomacrocoris handlirschi var. armatus Poisson 1948: Rev. Zool. Bot. Afr. 41: 210-212. NEW SYNONYMY
Supplemental redescription. Based primarily on a macropterous male from Uganda determined by Montandon (see Additional material examined). Length 8.80, maximum width 5.84. General shape elongate oval, widest across embolia (Fig. 10); overall coloration dorsally light brown anteriorly with darker brown hemelytra and black scutellum; ventrally, lateral part of propleura yellow, legs and abdomen medium to light brown, meso- and metasterna dark brown.
Head. Length 1.58, maximum width 2.40. Yellowish brown with dark brown band at posterior margin, becoming brown coalescent punctures anteriorly, punctures continuing anteriorly as two convergent lines, additional brown punctures between midline and eye. Head projecting beyond eyes 9% of head length. Eyes dark brown; approximately twice as long as wide, length/width 1.02/0.56; inner margins slightly convergent, lateral margins convex; synthlipsis 1.12. Labrum yellowish brown, transverse, broadly rounded, width/length 0.70/0.36. Maxillary plate yellow, elongate, extending approximately to one-third length of labrum, bordering sides of rostral base. Labium with three visible segments darkening distally from yellowish-brown basal segment to dark orangebrown distal segment, short, extending 0.45 beyond labrum. Antenna short, yellow, hirsute, extending slightly beyond lateral margin of eye.
Thorax. Pronotum broad, highly convex, yellow; slight depression in middle of disc (not present in specimens from Tanzania); scattered dark brown punctation throughout, heavier and coalescent near middle; wide transverse band along posterior margin set off from rest of pronotum by series of nearly adjacent strong punctures, punctures giving perception of subtle furrow, devoid of brown punctation but with irregular row of large, dark brown to black, longitudinally elongate spots; maximum width 5.16, length at midline 2.36; posterior margin broadly, shallowly convex with subtle concavity at midline; lateral margins strongly convex, convergent anteriorly, distinctly explanate; posterolateral corners rounded; with scattered pale setae. Scutellum black, punctate, with scattered thin setae and shorter thick setae, width 3.32, length at midline 1.84, lateral margins slightly convex. Hemelytra medium brown mottled with darker brown maculation, mostly tuberculate, scattered thin setae and shorter thick setae, length 6.40 (chord measurement). Clavus distinct, yellow at commissure, with scattered thin setae and shorter thick setae, venation not evident. Embolium clearly delineated, tuberculate, lateral margin evenly convex throughout, mostly yellowish with darker infuscation distally and medially, maximum width 0.84, length 2.64. Membrane punctate. Hindwings well developed. Ventrally, prosternum with thin mid-ventral carina. Propleuron laterally with extensive yellow, glabrous area extending 2/3 distance to mesal margin, brown pruinose mesad of yellow glabrous area, mesally with elongate recumbent hairs on occlusal surface with coxa; propleura widely separated at midline. Mesosternum with medial carina with elongate golden-brown setae and series of 6-7 ventrally directed protuberances; meso- and metasterna mostly dark brown, pruinose.
Legs. All segments yellowish brown except brown protarsus, meso- and metacoxae; profemur with scattered brown spots in posterior 2/3, anterior margin with dense pad of pale elongate setae; protibia and tarsus with flattened inner surface, sparse row of pale spatulate setae; single tarsal segment; claw minute; middle and hind coxae covered with short, pale, recumbent setae; metaxyphus with pronounced transverse and longitudinal carinae, thus resembling head of Phillips screwdriver directed posteroventrad; meso- and metafemora with posteroventral row of dark, peg-like spines; spines of mesofemur becoming obsolete proximally and replaced by short, pale setae; spines of metafemur consistently developed; middle of posterior surface with row of short, pale setae; posterodorsal row of dark, peg-like spines; meso- and metatibiae with rows of stout reddish-brown spines, 2 transverse rows of long, stout spines at apex; meso- and metatibiae and tarsi with long, golden brown swimming hairs; claws slender, evenly curved, with basal tooth. Leg measurements as follows: foreleg, femur 2.32, tibia 1.60, tarsus 0.40; middle leg, femur 2.28, tibia 1.73, tarsomeres 1–3, 0.20, 0.32, 0.42; hind leg, femur 2.64, tibia 2.96, tarsomeres 1–3 0.22, 0.58, 0.62.
Abdomen. Connexiva III–VI exposed laterally beyond hemelytra, each yellow anteriorly, brown posteriorly, margins smooth, with dense fringe of pale setae; posterolateral angles of II–III square, IV–VI slightly acuminate. (Terminal segments missing from Uganda specimen; VIII and aedeagus slide-mounted). Tergum VII of Tanzanian specimens (L-1174) posterior margin with left medial lobe broadly right-angled, with 0–5 denticles beneath the left margin; right medial lobe (pseudostrigil) with strong dorsal reflexion, large, transverse, with dense mat of denticles directed caudad on elliptical caudal surface narrowing toward midline, group of larger denticles on ventral surface of anterolateral corner (Fig. 12). Tergum VIII with left medial lobe broadly rounded (Uganda) to slightly digitate (Tanzania); right lobe triangular, broadly spatulate, with posterolateral area swollen and with long setae. Ventrally medium brown except narrow marginal glabrous yellow band; densely covered with short, pale, recumbent setae. Sternum V with posterior margin concave and mediosternite VI displaced asymmetrically to left. Genital operculum evenly rounded.
Genitalia. Pygophore brown, elongate setae generally scattered and with a dense brush posteriorly. Parameres lacking. Phallosoma with basal oblique striations hidden behind pygophore; abrupt dextral gibbosity; shaft beyond gibbosity straight and angled to left; apex with left corner rounded (Fig. 11); sclerotized vesica distad of phallosoma.
FIGURES 10–13. Neomacrocoris handlirschi (10) habitus of Montandon determined male from Uganda, (11) aedeagus of specimen in Fig. 10, (12) medial lobes of 7th abdominal tergum of male, (13) terminal abdominal sterna of female including subgenital plate. Sizes are not proportionate among figures. Fig. 11 photographed from Poisson slide at USNM; Figs. 12, 13 of specimens from L-1174, Tanzania.
Macropterous female. LECTOTYPE Ƥ. Length 9.08, maximum width 5.68. Similar to male in general structure and coloration with following exceptions: Abdominal mediosternites with quadrilateral glabrous patches widening posteriorly on V–VI, elongate glabrous patch covering middle 1/2 of subgenital plate (VII); long pale setae especially dense near margin of glabrous areas. Mediosternite VI with posterolateral corners slightly expanded and elevated (East African populations). Subgenital plate broad basally, gradually narrowing to elongate, laterally cupped, tongue-like lobe in distal 1/3; lateral margins sinuate (Fig. 13); lateral fringe of elongate golden brown setae at base of elongate lobe; length at midline 1.70, width at base 1.80.
Discussion. This species originally was described as Macrocoris handlirschi from specimens from South Africa (Montandon 1909). Four years later, this and two other species of Macrocoris were transferred to the new genus Neomacrocoris by Montandon (1913). Poisson (1948) reported that the species has a wide size range of 8.5–10 mm length and 5.2–5.3 mm width, which was slightly greater than the size range given in the original description. N. handlirschi was distinguished from N. angusticeps by Montandon (1914) by its smaller size, the pronotum with more distinct black punctation and posterolateral corners more blunt and narrowly rounded, and a superficial transverse furrow of the pronotal disc. Poisson (1948) also noted a slight indication of a posterior transverse furrow of the pronotal disc.
Neomacrocoris handlirschi var. armatus was described by Poisson (1948) from seven localities in the Belgian Congo (Luebo, Matadi, Boma, Luluabourg, Mayidi, Haut Uelé, and Katanga) and was distinguished from the nominate form by the condition of the 7th abdominal tergum. Because this variety was described prior to 1961, it is recognized as a valid subspecies by the Code of Zoological Nomenclature (ICZN 1999). Specifically, the left lobe has 1–8 denticles in N. handlirschi var. armatus, whereas it is without denticles in the nominate subspecies. In a population that we sampled from a small stream in Gombe National Park in northwestern Tanzania are males with from zero to five denticles on the left medial lobe of the 7th abdominal tergum, thereby presenting characteristics of both subspecies. Thus, because both character states are present in the same population, the subspecific designation based solely on presence or absence of denticles is not valid and we hereby synonymize N. handlirschi var. armatus Poisson with the nominate subspecies.
Poisson (1949, 1955, 1959) discussed the comparative morphology of the species and provided illustrations of the male 7th and 8th terga (1948, 1949) and aedeagus (1949). It is these papers that have contributed most to the currently accepted species concept of N. handlirschi, principally because they provided the only illustrations of the diagnostic male structures. Montandon (1909) described the species from a series collected in southern South Africa but did not designate a type specimen. Generally, males are required for identification of species of Neomacrocoris because diagnostic attributes usually are associated with the aedeagus and pseudostrigil. Montandon (1909) indicated that specimens of the type series would be deposited in Vienna and his personal collection. The following museums including Vienna were queried for syntypes of N. handlirschi: AMNH, BMNH, CMNH, DMSA, FMNH, HNHM, MCSN, MNB, MNHN, MNIN, NHMW, NMWC, RMCA, RMNH, SDEI, SEMC, SMNH, UMC, USNM. The only syntype is housed in NHMW (Vienna) and is a female, which we here designate as a lectotype. Although a male syntype was not available upon which to anchor the species, a male specimen of N. handlirschi from Uganda that was determined by Montandon in 1913 is housed in the USNM and with which we have many conspecific specimens from our recent naucorid survey in Tanzania. This specimen also is in agreement with Poisson’s concept of the species. It is based on this male specimen that we accept the species concept of Poisson and provide further diagnostic information here.
Attributes previously not reported that we have found to characterize the species are pronotum flattening and short, thick setae on the scutellum, embolium, clavus, corium, and membrane. Female specimens from our collections from Tanzania and others from Zambia, Zimbabwe, and western and northeastern Democratic Republic of the Congo, have the posterolateral corners of mediosternite VI slightly expanded and deflexed. Although the female lectotype’s pronotum is slightly flattened and several thick setae are at least on the left clavus, the posterolateral corners of mediosternite VI are not expanded. We provisionally consider the inconsistency in expression of mediosternite VI of females to be a regional adaptation and not necessarily an attribute representative of this widely distributed species. More specimens from throughout the range are needed to resolve this inconsistency.
This species is nearly indistinguishable from several species of similar size and shape without examination of male genitalia and pseudostrigil, although consistency of embolar curvature, pronotum flattening, and presence of thick setae on the hemelytra and scutellum can be used. N. handlirschi has a pronounced gibbosity on the right side of the phallosoma (Fig. 11) as does N. bondelaufa (Fig. 2); however, the top of the apex appears flattened from a dorsal view, whereas in N. bondelaufa, the flattened appearance is on the left side of the apex. Another similar and common species with which it co-occurs is N. parviceps, in which the right side of the phallosoma has an elongate flange rather than a gibbosity, and the abdominal sterna are reddish orange rather than brown as in N. handlirschi. Poisson (1949) provided an illustration of the aedeagus of N. handlirschi, although he reversed the orientation of his illustration such that the gibbosity is on the left, probably because his slide-mounted specimen (which we examined) was upside-down. He also gave attributes of male terga 7 and 8 to distinguish N. handlirschi from N. parviceps, including more denticles of the left lobe and the shape of the right corner of tergum 7, and longer and sparser hair of the right medial lobe of tergum 8. Among other diagnostic features of N. handlirschi (which it shares with N. ndugai) is that the pronotum is flatter in profile than that of N. bondelaufa, N. parviceps, and N. schaeferi (Figs. 14, 15). In addition to elongate thin hairs on the hemelytra, shorter, thicker hairs are sparsely but generally distributed, including on the embolium, corium, clavus, membrane, and scutellum (Fig. 10), whereas the thicker hairs are restricted to the embolium and adjacent areas of the corium in N. bondelaufa, N. parviceps, N. reavelli, and N. schaeferi.
Diagnosis. This species may be distinguished from all congeners except N. ndugai by the slight flattening of the pronotum and by the presence of short, thick setae sparsely distributed over the surface of the scutellum and hemelytra. In fresh specimens, the thick setae can be found sparsely over the entire dorsal surface of the hemelytra, including the clavus and membrane; however, the hairs apparently are fragile and can be less evident on older or mishandled specimens. The condition of the apex of the phallosoma distinguishes N. handlirschi from N. ndugai. The female subgenital plate is gently sinuate to the distal spatulate process (Fig. 13), which differs from only N. hungerfordi, N. transvaalensis, and N. vuga among the known females.
Distribution. The species occurs from eastern Sudan (Linnavuori 1981) and the Democratic Republic of Congo and Uganda south to South Africa. As such, N. handlirschi and N. parviceps might have the widest distribution of any species of Neomacrocoris. In Tanzania, N. handlirschi was reported from the Usambara Mountains in the northeastern part of the country (Poisson 1963). We collected N. handlirschi syntopically with N. bondelaufa at L-1202 and with N. parviceps at L-1174 and L-1202. All three species were collected together in the Msanzi River in Rukwa Region (L-1202). Specimens were collected in densely vegetated stream margins with muddy or gravelly substrate.
Published records. BELGIAN CONGO as N. handlirschi armatus (Poisson 1948), CENTRAL AFRICAN REPUBLIC (Linnavuori 1981), CONGO EX-BELGE, TANGANYIKA (Poisson 1963), [DEMOCRATIC REPUBLIC OF CONGO / UGANDA] Lac Albert (Poisson 1949), RUANDA (Poisson 1955), SOUTH AFRICA (Montandon 1909), SUDAN (Linnavuori 1981), UGANDA (Montandon 1914).
Type material examined. [SOUTH AFRICA]: LECTOTYPE by present designation: Algoa Bay, Capland, Dr. Brauns, 25 5 98 / Macrocoris handlirschi Montandon type 1909 (NHMW, Ƥ).
Additional material examined. BURUNDI: Coll. Mus. Tervuren, Churongwe, 25-II-1979, J. Decelle (RMCA, 13). CENTRAL AFRICAN REPUBLIC: Béboura-Bossanggoa, 1 June 1973, Linnavuori (NMWC, 13). CONGO: Poisson to Drake Coll 1979 (USNM, 1 3). CONGO BELGE: P. N. U., [= Parc National de l’Upemba], Gorges de la Pelenge, (1.150m.), 18-20-VI-1947, Mis. G. F. De Witte.526a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13); P. N. U., Kamitungulu (1700m), 21-1-1948, Mis. G. F. De Witte: 1259a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13); P. N U, Kanonga (695m), 13-27-IX-1947, Mis. G. F. De Witte: 785a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA, 1Ƥ); P. N. U., Kaziba (1.140m.), 7-12- II-1948, Mis. G. F. De Witte.1258a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi (Mont.) (RMCA 13); P. N. U., Lusinga (1760 m), 20-III-1947, Mis. G. F. De Witte: 79a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13); P. N. U., Lusinga (riv. Kagomwe), 8-vi-1945, G. F. De Witte: 81 / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13); same data, 88 (RMCA 23, 1Ƥ); P. N. U., Lusinga (Kamalongiru), 22-vi-1945, G. F. De Witte: 154 / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13, 2Ƥ); P. N. U., Lusinga (1760 m), 8-iv-1947, Mis. G. F. De Witte: 203a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13); P. N. U., Lusinga (galerie), 7-20-vi-1945, G. F. De Witte: 191 / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 1Ƥ); P. N. U., Masombwe-riv Kanakakazi (1129m), 4.16. X.1948, Mis. G. F. De Witte: 1911a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13); ex P. N. U., Masombwe- Kipepe affl., Tumbwe affl. Grde. Kafwe (1.120m.), 4-16-X-1948, Mis. G. F. De Witte.1918a / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA, 1Ƥ); P. N. U., Riv. Kagoma (affl. Lusinga), 12-vii-1945, G. F. De Witte: 212 / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA 13); P. N. U., Riv. Kipangaribwe (affl. Lusinga), 2-4-vii-1945, G. F. De Witte: 181 / Coll. Mus. Tervuren / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (RMCA); Poisson to Drake Coll 1979 (USNM, 23); P.N.U., Lusinga (Kamalongiru), 22 June 1945, G. F. de Witte:154 / R.
Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (NMWC, 13); P.N.U., Lusinga (1760 m), 20 March 1947, Mis. G. F. de Witte: 79a / R. Poisson det., 1951, Neomacrocoris handlirschi handlirschi Mont. (NMWC, 1Ƥ); P.N.A., 22-III-1954, P. Vanschuytbroeck & H. Synave, 8149-50 / Sector Tshiaberimu Kirungu (lieudit), 2.720 m (RMCA, 23, 1Ƥ); P.N.A., 29-VIII-3-IX-1952, P. Vanschuytbroeck & J. Kekenbosch, 919 / Massif Ruwenzori Kolonge, 2.210 m (RMCA, 2Ƥ); P.N.A., 18-VI-1957, P. Vanschuytbroeck, VS-53 / Massif Ruwenzori Kolonge, 2.150 m, riv. Nyamwamba affl. dr. Butahu (RMCA, 1Ƥ); P.N.A., 20-VII-1954, P. Vanschuytbroeck & H. Synave, 9847 / Massif Ruwenzori, Litongo (lieu-dit), 1.575 m, ex P.N.A. (RMCA, 13); Coll. Mus. Congo, Kaniama, -1931, R. Massart / Macrocoris flavicollis Sign., Det R. Poisson 1958 (RMCA, 13). [DEMOCRATIC REPUBLIC OF CONGO]: Coll. Mus. Congo, Bambesa, 9-V-1938, P. Henrard / R. Poisson det., 1954, Neomacrocoris handlirschi Mont. (RMCA, 13); Musée du Congo, Boma, -1937, Dr Dartevelle / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 2Ƥ); Musée du Congo, Dilolo, -VIII-IX- 1931, G. F. De Witte / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 13, 1Ƥ); Musée du Congo, Haut Uelé: Moto, 1920, L. Burgeon / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det. 5306, M. (RMCA, 13); Musée du Congo, Kasai, L. Achten / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 13); Musée du Congo, Katanga: Kansenla, 16-VII-1931, G. F. De Witte / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 1Ƥ); Musée du Congo, Katanga: Kazenza, 19 August 1931, G. F. de Witte, R. Det. 7126 B (RMCA, 2Ƥ) Musée du Congo, Katanga: Kanzenze, 20-viii-1931, G. F. De Witte / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det. 5306, M (RMCA, 23, 1Ƥ); Musée du Congo, Katanga 19/20, Kanzenze VIII-1931, G. F. De Witte / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det. 5306, M (RMCA, 1Ƥ); Musée du Congo, Katanga; Kapiri, VI-1925, G. F. De Witte / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det. 5306, M. (RMCA, 13, 1Ƥ); Musée du Congo, Tanganika: Moero, Kiambi-VI-VII-1930, Dr P. Gérard / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 13); Coll. Mus. Congo, Kivu: Mont Kahuzi, Km. 82, IX-51 / II-1952, H. Bomans / Neomacrocoris handlirschi Mont., R. Poisson det., 1960 (RMCA, 1Ƥ); Musée du Congo, Luebo, 16-VIII-1921, Dr H. Schouteden / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 1Ƥ); Musée du Congo, Luluabourg, 14-IV-1923, P. Callewaert / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 13, 1Ƥ); Coll. Mus. Congo, Mayidi, 1942, Rév P Van Eyen / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 1Ƥ); same data, 1945, (RMCA, 23, 1Ƥ); Coll. Mus. Congo, Matadi, -1937, Dr. Dartevelle / R. Poisson det., 1945, Neomacrocoris handlirschi Mont. / R. Det., 5306, M. (RMCA, 13); Matadi, 1937, Dr. Dartevelle / Poisson to Drake Coll 1979 (USNM, 13). TANZANIA: Kigoma Region, Gombe National Park, Mitumba stream, S 04º 38.405', E 029º 37.823', 776 m, 29 July 2010, colls R. W. Sites & A. Mbogho, heavily vegetated narrow forest stream, L-1174. Rukwa Region, Msanzi River at Msanzi, S 08º 11.411', E 031º 31.295', 1778 m, 2 August 2010, colls R. W. Sites & A. Mbogho, rocks, mud, gravel, sand, marginal grasses, L-1202. RUANDA: Kagogo, 1900 m., f. Ruhengeri, 29- I-1953, P. Basilewsky, Coll. Mus. Congo / R. Poisson det., 1954, Neomacrocoris handlirschi Mont. Ƥ (RMCA, 1Ƥ). S. RHODESIA: Inyangombi Falls Inyanga, 15 ii 1948, Mr. & Mrs. Omer-Cooper, B.M. 1969-138 (BMNH, 13, 3Ƥ; UMC, 1Ƥ); Inyamaziwa, Inyanga, TMSA (DMSA, 23, 4Ƥ; UMC, 1Ƥ). [UGANDA]: Albert Nyanza N.E. Riv. Waki, Alluoud 1909 / Neomacrocoris handlirschi Montand det. Montandon 1913 / Poisson to Drake Coll 1979 (USNM, 13). [ZAMBIA]: Kaombe, 11062010 #1 (PRC, 1Ƥ). [ZIMBABWE]: Salisbury, Mashonald. 1898 (BMNH, 13).
description: Published as part of Sites, Robert W. & Mbogho, Aaron Y., 2012, Revision of the African genus Neomacrocoris (Hemiptera: Heteroptera: Nepomorpha: Naucoridae), pp. 1-39 in Zootaxa 3555 on pages 10-15, DOI: 10.5281/zenodo.211898