Helochares tectiformis Fernández, 1982 by original designation.

(Slightly modified from Girón and Short 2021). Body length 4.2–9.5 mm. Body shape oval in dorsal view (Figs 7, 14, 17, 24); slightly to moderately convex in lateral view, with dorsal outline nearly flat along anterior 1/2 of elytra, or somewhat evenly curved. Coloration usually uniformly dark brown (Figs 14, 17, 24), sometimes orange or pale brown (Fig. 7), often paler along margins (e.g., Fig. 14A–C); opalescent sheen frequent in larger and darker species; ground punctation from very shallow (e.g., Fig. 17D) to moderately marked (e.g., Fig. 14A). Shape of head trapezoid (Fig. 6D). Eyes relatively large, not emarginated anteriorly, usually projected from outline of head (Fig. 6D). Clypeus trapezoid, with anterior margin broadly and roundly emarginate (Fig. 6D); membranous preclypeal area sometimes visible. Labrum fully exposed. Mentum with lateral longitudinal crenulations, lateral oblique ridges, and transverse crenulations along antero-medial area. Antennae with nine antennomeres; cupule strongly asymmetric, with rounded outline; antennomere 9 slightly to 2× longer than antennomere 7. Maxillary palps slender, moderately long, 0.8 (N. pastinum sp. nov.) to 1.8× (N. yanomami sp. nov.) the width of head; inner margin of maxillary palpomere 2 weakly and evenly curved to nearly straight, outer margin evenly curved or curved along apical 1/2; maxillary palpomere 3 slightly longer than 4. Prosternum flat to broadly and weakly convex. Elytra without sutural striae, with ground punctures usually shallowly marked; usually at least one row of systematic punctures visible along midline of each elytron (e.g., Figs 7C, 14D, 24C, D), often with visible rows along lateral and posterior regions of elytra; serial punctures sometimes visible along posterior 1/2 of elytra. Posterior elevation of mesoventrite, usually simply bulging (Fig. 6B), sometimes bulge impressed posteriorly (Fig. 6A); bulge often extending anteriorly as low, shiny, and glabrous longitudinal ridge (Fig. 6A, B); anapleural sutures concave, separated at anterior margin by distance 0.6–0.9× the width of anterior margin of mesepisternum. Metaventrite with medial glabrous patch, sometimes very narrow and extending along entire length of metaventrite. Protibiae with spines of anterior row extremely reduced to tiny appressed denticles. Metafemora with tibial grooves well developed; hydrofuge pubescence covering basal 6/7 of anterior surface. Tarsomeres 1–4 with long, thick, and rather dense setae on ventral face, sometimes with only rows of short spines on metatarsomeres 2–4; metatarsomere 2 as long or slightly longer than 5 and as 3 and 4 combined. Fifth abdominal ventrite apically emarginate, with fringe of stout setae. Aedeagus divided (e.g., Figs 3–5); parameres separated from each other for most of their length; median lobe divided in dorsal and ventral plates; dorsal plate usually strongly sclerotized and elongated, often bifurcated, or otherwise shaped along apical region; ventral plate sometimes reduced, usually simple and of variable length; basal piece 0.3× or less than length of parameres, usually clearly noticeable; gonopore usually clearly visible.

Figure 3. 

Aedeagi of the Novochares abbreviatus species group along with schematic representations of the whole, the parameres (in blue), and the dorsal (in yellow) and ventral (in red) plates of the median lobe A N. abbreviatus B N. oculatus.

Figure 4. 

Aedeagi of the Novochares punctatostriatus, sallaei, and orchis species groups along with schematic representations of the whole, the parameres (in blue), and the dorsal (in yellow) and ventral (in red) plates of the median lobe A N. dentatus B N. orchis C N. sallaei.

Figure 5. 

Aedeagi of the Novochares tectiformis species group along with schematic representations of the whole, the parameres (in blue), and the dorsal (in yellow) and ventral (in red) plates of the median lobe A N. trifurcatus B N. tectiformis C N. mura.

Figure 6. 

Morphological features of Novochares spp. A, B mesoventrite C, D dorsal view of the head A N. tectiformis B N. cochlearis C Aulonochares ligulatus D N. pertusus.

Novochares includes medium sized, pale brown to nearly black species that are somewhat dorsoventrally compressed and highly polished (smooth, and often shiny) to the naked eye. Across the Americas the most similar genus is Aulonochares Girón & Short, 2019, from which it can be differentiated by the shape of the head [trapezoid in Novochares (posterior margin of clypeus nearly twice as wide as anterior margin; Fig. 6D), subquadrate in Aulonochares (posterior margin of clypeus only nearly 1.3× wider than anterior margin; Fig. 6C)], and the sculpture of the mentum (variously strigate in Novochares, punctate in Aulonochares). Some members of Helochares of the Americas may resemble Novochares in their external features, but the aedeagal form is completely different (tubular in Helochares, see fig. 7 in Short and Girón 2018; divided in Novochares, e.g., Figs 3–5).

Figure 7. 

Dorsal habitus of Novochares spp. A N. abbreviatus B N. pilatus C N. minor D N. tenedor.

Nearctic: U.S.A. (Florida). Neotropical: Argentina, Belize, Bolivia, Brazil (Amapá*, Amazonas, Espírito Santo, Goiás*, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará*, Paraíba, Pernambuco, Piauí, Rio de Janeiro, Rondônia*, Roraima*, São Paulo), Colombia, Costa Rica, Cuba, Dominican Republic*, Ecuador, French Guiana, Guatemala, Lesser Antilles (Grenada, Guadeloupe, St. Vincent), Mexico, Panama, Paraguay, Peru*, Puerto Rico, Suriname, Trinidad and Tobago*, Uruguay, Venezuela (new country or state records indicated with an asterisk).

The genus occupies a broad range of aquatic habitats, including both lentic and lotic situations. We are not aware of any seepage specialists. The most commonly collected species (e.g., those in the N. abbreviatus species group) are typically associated with open swamps and marshes where they may also be attracted to lights, sometimes in large numbers. Many species are also found in forested pools and swamps that contain abundant detritus. They may also be abundant in detrital pools in drying streambeds or along the margins of slow moving or quiet streams. It is also important to note that a number of Novochares species can co-occur in the same habitat at the same time.

The females of most if not all species in the genus carry their egg case around under their abdomen, a behavior also seen in other Helochares-group genera (such as the Neotropical Aulonochares, Radicitus, and Sindolus; Girón and Short 2021). Only the immature stages (eggs, larvae, pupae) of N. pallipes have been described to date (Fernández 1983).

Novochares abbreviatus species group

Species group diagnosis. Body length 5.5–8.1 mm. Coloration: Dorsal surfaces pale brown to yellowish brown (Fig. 7A, B), with paler (yellowish to orange) clypeus and margins of pronotum and elytra. Head: Maxillary palps 1.1–1.6× width of head, uniformly yellow or orange to brown in color (Fig. 7A, B). Thorax: Ground punctation on pronotum and elytra relatively dense and very shallowly impressed. Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum medially weakly and broadly convex. Posterior elevation of mesoventrite broadly and somewhat triangularly elevated, sometimes posteriorly transversely impressed, with low medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite shallow to moderately deep and broad, U-shaped. Aedeagus: (Figs 3, 8, 9) Overall shape sub-rectangular to sub-rhomboid, 2.5–3.0× longer than wide, joint basal margins of parameres truncate (e.g., Fig. 8M) to medially pointed (e.g., Fig. 8A, G); outer margin and apical region of each paramere variable, usually rounded at apex and laterally pointed; parameres longer than median lobe; parameres with apical region uniformly sclerotized and dorso-ventrally flattened; dorsal inner margin of each paramere usually medially broadly emarginate (see Fig. 5 in blue); dorsal plate of median lobe (in dorsal view) with basal apodemes usually reaching base of parameres; dorsal plate of median lobe usually with long and narrow neck (see Fig. 5 in yellow), variably expanded at apical region, usually with distal arms variable in shape and length; gonopore sitting proximal to base of median lobe; ventral plate of median lobe (in ventral view, see Fig. 5 in red) somewhat triangular, moderately sclerotized; dorsal surface of ventral plate of median lobe slightly concave (lateral regions curved dorsally); basal piece nearly (see Fig. 5 in green) 0.25–0.30× length of a paramere. In lateral view, aedeagus triangular, nearly straight at base, with ventral outline of parameres 3.0–3.6× longer than greatest width near base (e.g., Fig. 8C, F, I, L, O).

Figure 8. 

Aedeagi of the Novochares abbreviatus species group A–L N. abbreviatus A–C Nicaragua D–F Argentina G–I Venezuela J–L Guyana M–O N. latus P–R N. pilatus A, D, G, J, M, P dorsal view B, E, H, K, N, Q ventral view C, F, I, L, O, R lateral view.

Figure 9. 

Aedeagi and habitus of the Novochares abbreviatus species group A–E N. oculatus: A–C aedeagus, Mexico D, E paralectotype from Guatemala D aedeagus E habitus F–K miscellaneous forms of aedeagi in the abbreviatus species group F, G Argentina H, I Venezuela J, K Bolivia L, M, R N. pallipes N, O N. baca A, D, F, H, J, L, N dorsal view B, M ventral view C, G, I, K, O lateral view R oblique view.

Composition. This species group contains three previously described species: Novochares abbreviatus (Fabricius, 1801), N. oculatus (Sharp, 1882), and N. pallipes (Brullé, 1841) and three new species: N. baca sp. nov., N. latus sp. nov., and N. pilatus sp. nov.

Remarks. This is a widespread and certainly problematic species group. The external morphology of all the known species is annoyingly uniform, and the characteristics of the male genitalia, especially the apical region of the dorsal plate of the median lobe, when looking at long series of specimens from across the distributional ranges of N. abbreviatus and N. oculatus, exhibit gradual changes that blur the limits between both species. More thorough sampling and molecular data, perhaps in the context of international collaboration across the species group range, are needed to tackle the systematics and taxonomy of this species group.

(98 exs.). Bolivia: Santa Cruz: Ayacucho, 13–14.v.1969 leg. P. & P. Spangler (1, USNM); Santa Cruz, 11–12.v.1969, leg. P. & P. Spangler (1, USNM); 60 mi N. Santa Cruz, Saavedra Experiment Station, 3–5.i.1960, leg. Robert Cumming (1, USNM); Potrerillos del Guenda, Preserva Natura, 17°40'S, 63°27'W, 370 m, 17–22.x.2007, lights, leg. Cline & Wappes, BOL1Cline07 007 (1, SEMC); 3.7 km SSE Buena Vista, Hotel Flora y Fauna, 23–30.iv.2004, lights, leg. A.R. Cline (4, SEMC, TTU-Z), same data but 1–12.v.2004 (4, SEMC). Brazil: Bahia: 15 km E. Itabuna, 3.vii.1963, leg. P. & P. Spangler (2, USNM). Goiás: Divinópolis de Goiás, 2 km SE on GO-447, Roadside ponds, BR18-0222-02A (1, SEMC, DNA voucher, SLE2087). Pará: Murumuru, 1.5 km E, Muddy shallow marsh along road, BR18-0205-02A (1, SEMC, DNA voucher SLE 2083). Mato Grosso do Sul: Corumba (ca. 29 km SE) on BR-262, 25.vi.2018, leg. Hamada et al., drying marsh, BR18-0625-02A (1, SEMC); Miranda (ca. 9.5 km SW) on MS-339, Marsh area alongside stream, BR18-0626-03A (2, SEMC, including DNA voucher SLE2086). Rio Grande do Norte: Ceara-Mirim, 6–7.vii.1969, leg. P. & P. Spangler (1, USNM). Roraima: BR-401, ca. 6 km SW of Bonfim, 3°21.615'N, 59°53.361'W, 100 m, 12.i.2018, leg. Short, Benetti & Santana, large marsh with abundant vegetation, BR18-0112-02A (1, SEMC); BR-401, ca. 26 km NE of Boa Vista, 2°56.191'N, 60°28.017'W, 92 m, 12.i.2018, leg. Short, pooled up morichal, BR18-0112-06B (1, SEMC); Caroebe, Rio Jatapu, nr. Usina de Jatapu, 00°50.939'N, 59°18.262'W, 145 m, 17.i.2018, leg. A. Short, marginal pools of river, BR18-0117-02A (1, SEMC). São Paulo: Piracicaba, 12.xii.1965, leg. C.A. Triplehorn (1 male, USNM); same data but 6.x.1965 (1, USNM). Colombia: Amazonas: Leticia, 12–15.iii.1969, leg. P. & P. Spangler (1, USNM). Costa Rica: Guanacaste: nr. Carmona, laguna de crocodile, 34 m, 16.i.2003, leg. Short, Roughly, & Porras, HG light (3, SEMC); Rio Animas, River w/ volcanic rock bottom, small cascades, small isolated detrital backwaters, AS-04-040 (1, SEMC, DNA Voucher SLE1180). Puntarenas: Puntarenas, 22.vii.1965, leg. P.J. Spangler (1, USNM). Dominica: Cabrit Swamp, 10–13.v.1965, leg. D.R. Davis (1, USNM); Postsmouth, 19–21.x.1966 (1, USNM). Dominican Republic: La Toma N of San Cristobal, 9–10.vi.1969, leg. Flint & Gomez (1, USNM). Guadeloupe: Pointe-a-Pitre, 1936, Henri Stehle (1, USNM). Guyana: Good Hope (7 mi. NW, on road to Karasabai), 26.iv.1995, leg. Spangler & Perry (1, USNM); Region 9: Tributary of the Takatu River, NW of Kusad Mts., 2°50.563'N, 59°59.113'W, 109 m, 24.x.2013, leg. Short, Isaacs, & Salisbury, vegetated creek margins, GY13-1024-02B (1, CBDG); nr. Kusad Mts., 2°49.793'N, 59°48.361'W, 123 m, 25.x.2013, leg. Short, Isaacs, & Salisbury, large vegetated marsh, GY13-1025-01A (1, SEMC, DNA Voucher SLE1221); Pirara Ranch & River, 3°32.1'N, 59°40.5'W, 23–27.iv.1995, leg. O.S. Flint (1, USNM). Nicaragua: 13 mi N. Sn. Benito, 11.vii.1965, leg. P.J. Spangler (1, USNM); Rivas, Reserva Silvestri Domitila, 5–9.vi.2005, 400 ft., lights, W.D. Shepard (2, SEMC). Panama: Chiriqui Province, Las Lajas (8 km S.), 10 m, 5.vi.1983, leg. P.J. Spangler, roadside ditch (1, USNM). Panama Province: Pond at Panama Canal, Explosive Depot, 31.viii.2006, leg. W.D. Shepard & D. Post (3, SEMC). Paraguay: Central Department: San Bernardino, 10.iv.1980, leg. Spangler, Culzoni, & Wood (1, USNM); same locality but 22.vi.1969, leg. P. & P. Spangler; Aregua, 26–27.iv.1980, leg. P.J. Spangler (1, USNM). Peru: Huanuco: Tingo María, 19–24.iv.1969, leg. P. & P. Spangler (1, USNM); Loreto: SW Iquitos, Aguajal next to Iquitos-Nauta highway, Margins and saturated leaves of aguajal (palm swamp) PE20-0119-01A (1, SEMC, DNA voucher SLE2152). Puerto Rico: San Germán, 23.xii.1962, leg. P. & P Spangler (1, USNM); nr. La Cueva del Indio, 14.i.1963, leg. P.J. Spangler (1, USNM); Hwy 3, km 32.6 nr. Palmer, 10.i.1963 (1, USNM). Suriname: Commewinje: East-West Highway, ca. 6 km E. of Suriname River, 5°47.464’N, 55°06.730’W, leg. Short, SR13-0809-01A (3, NSCS, SEMC). Marowijne: East-West Highway, 15 Km E. Commewijne River, 4.iii.2012, leg. Short & Kadosoe, sandy/marshy roadside swale, SR12-0304-04 (1, SEMC, DNA voucher SLE1210). Saramacca: 1 km E. Sidiredjo, 5.iii.2012, leg. Short & Kadosoe, roadside swale, SR12-0305-02A (1, SEMC, DNA voucher 1215). Trinidad And Tobago: Trinidad, Debe, 17.vii.1969, leg. P. & P. Spangler (1, USNM). Venezuela: Anzoátegui: El Tigre, N of, river along highway, 9°5.808'N, 64°19.445'W, 236 m, 3.ii.2010, leg. García, shaded margins without vegetation, VZ10-0203-03A (1, SEMC); same data except leg. Short, vegetated backwater margins, VZ10-0203-03B (1, SEMC). Apure: Bruzual, edge of town, 8°2.534’N, 69°20.530’W, 83 m, 18.i.2009, leg. Short, Camacho, Miller, large marsh VZ09-0118-04X (1, SEMC). Barinas: Obispo, 25.ii.1969, leg. P. & P. Spangler (1, USNM); Libertad, E of, along side gravel road, 8°25.773’N, 69°35.202’W, 106 m, 19.i.2009, leg. Short, Camacho, Miller, forested canal, VZ09-0119-01X (2, SEMC); Ciudad Bolivia, approx. 13 km SE, large Hacienda, 8°19.394'N, 70°28.238'W, 173 m, 25.i.2012, leg. Short, Arias, & Gustafson, marsh, VZ12-0125-02A (1, SEMC, DNA voucher SLE1217). Bolívar: Gran Sabana, N. Santa Elena, Rio Guara at Rt. 10, 4°37.362’N, 61°5.679’W, 876 m, 17.vii.2010, leg. Short, Tellez, & Arias, marshy area, VZ10-0717-02A (2, SEMC). Cojedes: El Baul, 5 km S, 21.i.2012, leg. Short, Arias, & Gustafson, large marsh, VZ12-0121-03A (1, SEMC, DNA Voucher SLE1240). Delta Amacuro: Between Tucupita & Los Guires, 9°10.504’N, 61°54.610’W, 8 m, 3.ii.2010, leg. Short & García, marsh by road, VZ10-0203-01A (1, SEMC, DNA Voucher SLE1208); between Tucupita & Temblador, small pond along road, 8°46.439'N, 62°14.306'W, 19 m, 2.ii.2010, leg. Short, García, Joly, margins of vegetated pond, VZ10-0203-02A (1, SEMC). Guárico: San Fernando, 12.ii1969, leg. P.&P. Spangler (1, USNM). Monagas: S of Maturin, morichal at road crossing, 9°16.398'N, 62°56.246'W, 22 m, 2.ii.2010, leg. Short, García, & Joly, morichal margin, VZ10-0202-02A (2, SEMC). Sucre: El Pilar, approx. 5 km SE, 10°31.419'N, 63°7.070'W, 2 m, 29.i.2010, leg. Short & Garcia, marsh/swamp along road, VZ10-0129-04A (17, MIZA, SEMC, TTU-Z, including DNA voucher SLE1207). Trujillo: Sabana Grande, Rio Jirijara, 9°42.307'N, 70°32.570'W, 199 m, 29.i.2012, leg. Short, Arias, Gustafson, small muddy pool in river floodplain, VZ12-0129-02A (1, SEMC). Zulia: Puente del Zulia, lagoon on finca, 17.i.2012, leg. Short et al., large lagoon, V12-0127-01A (1, SEMC, DNA voucher SLE1237); Sabana de Machango, 10°2.581'N, 71°0.428'W, 35 m, 29.i.2012, leg. Short, Arias, & Gustafson, margin of artificial pond, VZ12-0129-03A (1, SEMC). Virgin Islands: St. Thomas, 20.i.1963, leg. P.J. Spangler (2, USNM).

The defining feature of this species is the shape of the apical region of the dorsal plate of the median lobe, which bears a small rounded “cup” at its apex with two or three small teeth along its distal margin (Fig. 8A–L). However, only specimens that very closely match the illustrated forms should be considered N. abbreviatus. Specimens that possess expanded apical “cups” or long distal arms are likely other species such as N. oculatus (Fig. 9A, B), N. baca (Fig. 9N), or other still undefined lineages.

Body length 5.5–7.0 mm. Coloration: Dorsal surfaces pale brown to yellowish brown, with paler (yellowish) clypeus and margins of pronotum and elytra (Fig. 7A). Head: Maxillary palps 1.2–1.6× width of head, uniformly orange to brown in color (Fig. 7A). Thorax: Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum medially weakly and broadly convex. Posterior elevation of mesoventrite broadly and somewhat triangularly elevated, sometimes posteriorly transversely impressed, with low medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite shallow to moderately deep and broad. Aedeagus: (Figs 3A, 8A–L) Outer margin and apical region of each paramere rounded at apex and laterally pointed to hook-shaped; dorsal plate of median lobe (in dorsal view) with long and narrow neck, ovally expanded at apical region, with two or three very short distal arms; ventral plate of median lobe triangular, strongly sclerotized.

Argentina, Bolivia, Brazil (Espírito Santo, Mato Grosso do Sul, Pará, Pernambuco, Piauí, Rio Grande do Norte, Roraima, São Paulo), Colombia, Costa Rica, Cuba, Dominica (new record), Dominican Republic (new record), French Guiana, Guadeloupe (new record), Guyana (new record), Nicaragua (new record), Panama, Paraguay, Peru (new record), Puerto Rico (new record), Suriname, St. Thomas (new record), Trinidad and Tobago (new record), Venezuela (Fig. 10A).

Figure 10. 

Distribution of Novochares abbreviatus species group A N. abbreviatus: examined specimens (red) and literature/unconfirmed records (blue) B N. oculatus: examined specimens (red) and literature/unconfirmed records (blue) C N. pallipes (red), N. latus (yellow), N. baca (blue), N. pilatus (green).

This species is a common element of the lentic water beetle fauna throughout much of the Neotropics. It is most frequently collected in open marshes, swamps, pond margins, or along the margins of larger rivers.

The body length measurements presented here correspond to confirmed males for the species. See extensive discussion and remarks under the species group for further history and information that relates to this name and taxon.

Among the thirteen specimens we sequenced from Costa Rica to southern Brazil, the maximum intraspecific pairwise divergence in COI was a relatively meager 3.7%, and along with relatively uniform morphology, this supports the conclusion that this species is extremely widespread throughout the Neotropical region as the literature suggests.

Holotype (male): “PERU: Madre de Dios: Tambopata/ -12.54034 S, -69.00074 W, 190m/ Kawsay Biological Station, 4.vi.2022/ Palm swamp; lots of detritus/ PE22-0604-01B, leg. Short et al.” (MHNSM). Paratypes (9 exs.): Brazil: Pará: ca. 25 km E of Alenquer, -1.96253, -54.50458, 44 m, 4.ii.2018, Short & Benetti, Palm swamp, lots of detritus, BR18-0204-01A (1, SEMC, DNA Voucher SLE1513); Vale do Paraíso, ca. 55 km N. Alenquer, -1.49292, -54.51566, 150 m, leg. Short & Benetti, seeps & pools by waterfall, BR18-0203-01D (1, INPA, DNA Voucher SLE1617); same data except pool with rocks and detritus on trail, BR18-0203-01E (1, SEMC, DNA Voucher SLE2081). Ecuador: Sucumbíos: Sacha Lodge, 0.5°S, 76.5°W, 270 m, 10–12.xi.1994, leg. Hibbs, ex. Malaise (1, SEMC). Peru: Madre de Dios: Same data as holotype (4, SEMC, MHNSM). Suriname: Sipaliwini: Sipaliwini Savannah Nature Reserve, North of Basecamp, 2°00.656'N, 55°59.070'W, 275 m, 1.iv.2017, leg. Short, grassy pools near river, SR17-0401-01A (1, NZCS).

The relatively long and straight outer margins of the parameres and the very long and narrow fork of the apex of the dorsal plate of the median lobe (Fig. 9N) help to distinguish this species from others in the species group.

Body length 6.0–6.2 mm. Coloration: Dorsal surfaces pale brown to yellowish brown, with paler (yellowish) clypeus and margins of pronotum and elytra. Head: Maxillary palps only slightly longer than width of head, uniformly yellow in color. Thorax: Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Posterior elevation of mesoventrite broadly and somewhat triangularly elevated, weakly posteriorly transversely impressed, with low medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite moderately deep and broad. Aedeagus: (Fig. 9N, O) Outer margin and apical region of each paramere rounded at apex and laterally pointed; dorsal plate of median lobe with long and narrow neck, ovally expanded at apical region, with two narrow, elongated, laminate (dorsoventrally oriented), distal arms, basally separated by width of arm; arms as long as base of fork; ventral plate of median lobe triangular, strongly sclerotized, apically broadly angulate, reaching mid-length of dorsal plate.

This species is named after Stephen Baca, long-time member of the Short Lab, who supported the lab and its members by providing assistance during fieldwork, lab work, and overall being a great friend, colleague, and mentor. The last name Baca, when read, sounds similar to the Spanish word vaca, cow in English; the distal region of the dorsal plate of the median lobe somewhat resembles the head of a cow with horns.

Known from a few but widely separated localities in Brazil (Pará), Ecuador, Peru, and Suriname (Fig. 10C).

This species has been collected in palm swamps, forested pools alongside rivers and waterfalls, and in the case of the Suriname locality, in a grassy pool alongside a river in an open savanna.

Holotype (male): “BRAZIL: Rondônia/ -8.92368, -62.12491; 82 m/ Tabajara (c. 7.5 km W) on RO-133/ 8.vii.2018. leg. Short; river/ w/sandy bottom and rocks/ BR18-0708-04A”, “DNA Voucher/ Extraction #/ SLE-2039” (INPA). Paratypes: (4 exs.): Brazil: Rondônia: same data as holotype (4, INPA, SEMC).

This taxon has one of the most distinctive aedeagal forms within the species group. The extremely broad, parallel sided parameres with a ‘birdhead’ form at the apex, paired with a uniquely short and deeply cleft apical region of the dorsal plate of the median lobe (Fig. 8M–O) make this species easy to recognize; we did not see any other variations that were similar to this one.

Body length 7.2–8.1 mm. Coloration: Dorsal surfaces pale brown, sometimes with weakly paler (yellowish) clypeus and margins of pronotum and elytra. Head: Maxillary palps nearly 1.3× width of head, uniformly yellow to orange in color. Thorax: Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum medially broadly convex. Posterior elevation of mesoventrite broadly and somewhat triangularly elevated, transversely weakly impressed posteriorly, with low medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite moderately deep and relatively narrow. Aedeagus: (Fig. 8M–O) Overall shape sub-rectangular; apical region of each paramere relatively broad, rounded at apex, with outer margin laterally pointed; dorsal plate of median lobe with long and narrow neck, bifurcated at apical region; each arm of the fork somewhat triangular, relatively broad, and short; ventral plate of median lobe triangular, strongly sclerotized, reaching to second 1/3 of dorsal plate.

Latus (L.) meaning broad, referring to the shape of each arm of the apical region of the dorsal plate of the median lobe.

Only known from the type locality in Brazil (Fig. 10C).

The single collection of this species was from the sandy margin of a forested creek with some detritus.

Paralectotype (male): “Helochares ocu-/latus D.S./ Paso Antonio. Guate/mala Champion [on card with specimen]”, “Sharp Coll./ 1905.-313”, “Paso Antonio,/ 400 ft./ Champion.”, “B.C.A. Col. I. 2./ Helochares/ oculatus,/ Sharp.”, “Brit.Mus./ USNM-1966/ EXCHANGE” (USNM; Fig. 9E).

(8 exs.). Belize: Stann Creek, Sitte Point, Possum Point Biological Station, 24.iv.1987, leg. P.J. Spangler (1, USNM). Colombia: Magdalena: 8 km E Barranquilla, 19.iii.1969, leg. P. & P. Spangler (1, USNM). Mexico: Jalisco: Barra de Navidad, 23.iii.1971, leg. J.R. Zimmerman (1, USNM). Oaxaca: 31 km S Tuxtepec, Bethania, Ao. Chopan, 24.v.1981, blacklight, leg. P.J. Spangler (1, USNM); Sinaloa: Mazatlan, 17.vii.1963, leg. P.J. Spangler (1, USNM). Panama: Canal Zone, Barro Colorado Island, vi.1939, leg. J. Zetek (2, USNM), same locality but 29.v.1940, at light (1, USNM).

The precise morphological boundaries of this taxon are still uncertain. Its primary characteristic within the species group is its relatively large and oval distal cup of the dorsal plate of the median lobe, which bears two long arms (Fig. 9A, B). However, there is much variation and we recommend a relatively conservative approach in making confirmed species identifications at this time (see remarks for further discussion).

Body length 6.0–6.5 mm. Coloration: Dorsal surfaces pale brown to orange-brown, sometimes with weakly paler (orange) clypeus and margins of pronotum and elytra. Head: Maxillary palps nearly 1.3× width of head, uniformly yellow to orange in color. Thorax: Elytra without rows of serial punctures, faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures barely noticeable. Prosternum medially broadly convex. Posterior elevation of mesoventrite broadly elevated, with low medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite moderately deep and relatively narrow. Aedeagus: (Figs 3B, 9A–D) Outer margin and apical region of each paramere rounded at apex and laterally pointed to hook-shaped; dorsal plate of median lobe with long and narrow neck, obliquely explanate at apical region, somewhat diamond-shaped and laterally rounded, with two laminate and slender distal arms, narrowly separated at base, apically converging, and slightly dorsally pointing; arms nearly as long as base of fork; ventral plate of median lobe triangular, moderately sclerotized.

Belize (new record), Colombia (new record), Costa Rica, Guatemala, Mexico (confirmed record), Panama (Fig. 10B). All records from southern South America and the Caribbean need further review (Argentina, Brazil, Paraguay, and the Antilles [Grenada, St. Vincent]).

The identity of this species has been wrapped up in confusion with N. abbreviatus almost immediately after it was described (see above discussion of the abbreviatus group nomenclature). We examined the dissected male lectotype and a dissected male paralectotype male deposited in the USNM (Fig. 9D, E; it had been exchanged between USNM and the British Museum in the 1960s). We saw a number of other specimens that had an oculatus-like aedeagus but with a range of variation around the precise shape and size of the apical region of the dorsal plate of the median lobe, as well as the shape of the paramere apices (Fig. 9F, H). We did not have good sampling of these forms for molecular data, so were not able to use an integrated approach to explore this variation. Consequently, we have taken a very conservative approach and identified only male specimens which had genitalia that very closely matched the type material as N. oculatus. The remaining material will need to wait until more data, especially genetic data, is available. There are a number of published records of N. oculatus from Argentina, Paraguay, and southern Brazil (Fernández 1982a; Clarkson and Ferreira Jr 2014), and while it is very much possible that N. oculatus (like N. abbreviatus) is a very widespread Neotropical species, these reports should be treated as unconfirmed until molecular data from those localities is available.

(14 exs.). Argentina: Buenos Aires: Martinez, xii.1953 (2, CAS); San Isidro, xii.1955 (6, CAS, SEMC); Martinez, xii.1957 (2, USNM); Choya, “Sta. del Estero”, 10.iii.1962 (1, CAS). Entre Ríos: Río Paraná Ibicuy, Pto. Ibicuy, 10.xii.1979, leg. C.M. & S. Flint, Jr. (3, USNM).

The strongly curved parameres and the extremely large distinctive fork of the dorsal plate of the median lobe (Fig. 9L–R) serve to separate this species fairly easily from other members of the species group.

Body length 7.2–7.8 mm. Coloration: Dorsal surfaces pale brown to orange-brown, sometimes with weakly paler (orange) clypeus and margins of pronotum and elytra. Head: Maxillary palps nearly 1.3× width of head, uniformly yellow to orange in color. Thorax: Elytra without rows of serial punctures, faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures barely noticeable. Prosternum medially broadly convex. Posterior elevation of mesoventrite broadly elevated, with low medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite moderately deep and relatively narrow. Aedeagus: (Fig. 9L–R) Apical region of each paramere medially and dorsally curved; outer margin of apical region of paramere rounded, latero-ventrally weakly pointed; dorsal plate of median lobe with long and relatively broad neck, widened at apical region; neck with paired longitudinal, laminar elevations along second 1/2; distal arms laminate, dorso-ventrally oriented, 1/2 as long as neck of dorsal plate of median lobe; ventral plate of median lobe narrow and triangular, moderately sclerotized, reaching to beyond base of fork of dorsal plate of median lobe.

Argentina, Uruguay (Fig. 10C). Prior records from Brazil and Paraguay were erroneously reported and are removed from the known distribution of this species.

There has been some confusion about the distribution of this species in the literature. We have only been able to confirm the presence of this species in Argentina and Uruguay. Literature reports of this species occurring in Mato Grosso do Sul (Corumbá) and Paraguay (Río Alto Paraná) appear to be derived from the original description of Helochares (s. str.) parhedrus. This species was briefly synonymized with N. pallipes but later discovered to actually be a synonym of H. atratus (Fernández, 1982). However, because for a short time the distributional records of N. pallipes and H. parhedrus were fused, this muddied the literature. Fernández (1983) appears to have accidentally repeated the fused distribution which then carried into later papers. The record of this species from the state of Minas Gerais, Brazil derives from d’Orchymont (1926). However, later d’Orchymont (1939) noted that these records were erroneous and referred them to H. parhedrus, which later was synonymized with N. atratus. We are not aware of any verified Brazilian records of H. pallipes.

Holotype (male): “VENEZUELA: Bolivar State/ 6°35.617'N, 66°49.238'W, 80m/ Los Pijiguaos; outcrop/morichal/ 12.i.2009; leg. Miller & Short/ V09-0112-01C; detrital pools (MIZA). Paratypes: (68 exs.): Venezuela: Barinas: East of Santa Barbara, Rio Santa Barbara, 7°50.028'N, 71°11.188'W, 177 m, 25.i.2012, leg. Short, Arias, & Gustafson, big side pool of river, VZ12-0126-01B (1, SEMC, DNA Voucher SLE1241). Bolívar: same data as holotype (67, MIA, SEMC, TTU-Z, including DNA Voucher SLE1204).

This species is distinguished by a relatively extended and narrow expansion at the apex of the dorsal plate of the median lobe, which is also set with two modest arms (Fig. 8P, Q).

Body length 6.0–7.4 mm. Coloration: Dorsal surfaces pale brown to yellowish brown, with paler (yellowish) clypeus and margins of pronotum and elytra. Head: Maxillary palps 1.4–1.5× longer than width of head, uniformly yellow to orange in color. Thorax: Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum only weakly convex. Posterior elevation of mesoventrite broadly and somewhat triangularly elevated, posteriorly weakly transversely impressed, with low medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite moderately deep and broad. Aedeagus: (Fig. 8P–R) Outer margin and apical region of each paramere rounded at apex and laterally pointed; dorsal plate of median lobe with long and narrow neck, ovally expanded at apical region, with two narrow, elongated, laminate (dorsoventrally oriented), contiguous distal arms; arms as long as 1/2 length of base of fork; ventral plate of median lobe triangular, strongly sclerotized, apically pointed, reaching mid-length of dorsal plate.

Pilatus (L.) meaning large. In reference to the relatively large size of the aedeagus when compared with other species in the abbreviatus species group.

This species is found in several localities in central Venezuela (Fig. 10C).

The large main series of this species was collected in shallow detrital pools that were along the edge of a morichal that ran alongside a granite outcrop. Another specimen was collected in the side pools of a large river.

Novochares aperito species group

Species group diagnosis. Body length 4.9 mm. Coloration: Dorsal surfaces orange, with paler clypeus and margins of pronotum. Head: Maxillary palps slightly longer than width of head, uniformly orange in color. Thorax: Ground punctation on pronotum and elytra relatively dense and very shallowly impressed. Elytra without rows of serial punctures, each with very faint rows of weakly marked systematic punctures on lateral 1/2. Prosternum weakly medially convex. Posterior elevation of mesoventrite transversely elevated. Abdomen: Apical emargination of fifth ventrite relatively deep, U-shaped. Aedeagus: (Fig. 11A–C) Overall shape pear-like, joint basal margins of parameres truncate; apical region of each paramere somewhat hook-like; parameres longer than median lobe, with apex rounded; dorsal inner margin of each paramere sinuate; dorsal plate of median lobe (in dorsal view) with neck weakly defined, with arms weakly defined; gonopore placed near apex of dorsal plate of median lobe; ventral plate of median lobe (in ventral view) membranous; basal piece nearly 0.3× length of a paramere. In lateral view, aedeagus somewhat triangular, straight at base, with ventral outline of parameres nearly 3× longer than greatest width near base.

Figure 11. 

Aedeagi of Novochares spp. A–C N. aperito D–F N. orchis G–K N. minor G–J Peru K Venezuela A, D, G, K dorsal view B, E, H ventral view C, F, J oblique view I lateral view.

Composition. This group is composed of a single known species from Bolivia (known from a single male) with a very unusual and distinctive genitalia: N. aperito sp. nov.

Holotype (male): “BOLIVIA: Santa Cruz Dept./ Potrerillos del Guenda,/ Preserva Natural, 17°40'S, 63°27'W, 370m, 12–13-X-2007/ ex. BL/MV, A.R Cline & J.E./ Wappes BOL1Cline07 004.5” (SEMC).

See species group diagnosis.

Body length 4.9 mm. Coloration: Dorsal surfaces orange, with paler clypeus and margins of pronotum. Head: Maxillary palps slightly longer than width of head, uniformly orange in color. Thorax: Ground punctation on pronotum and elytra relatively dense and very shallowly impressed. Elytra without rows of serial punctures, each with very faint rows of weakly marked systematic punctures on lateral 1/2. Prosternum weakly medially convex. Posterior elevation of mesoventrite transversely elevated. Abdomen: Apical emargination of fifth ventrite relatively deep, U-shaped. Aedeagus: (Fig. 11A–C) 2.3× longer than wide, with outer lateral margins of parameres slightly convex along basal 1/2, then weakly sinuate up to apical region; apical region of each paramere rounded, with outer margin somewhat hook-like; at closest point, dorsal inner margins of parameres separated by distance 0.33× greatest width of a paramere; dorsal plate of median lobe with neck 0.7× as broad as base; arms of dorsal plate of median lobe not clearly indicated; apical region of dorsal plate of median lobe dorsally concave, apically pointed; ventral plate of median lobe membranous, neck of dorsal plate; basal piece 0.3× length of a paramere. In lateral view, aedeagus straight at base, with ventral outline of parameres 3× longer than greatest width near base; dorsal outline of aedeagus in lateral view slightly convex along basal 1/4, then oblique to apical region; ventral outline of aedeagus in lateral view straight.

Aperito (L.) meaning to open, referring to the distinctive shape of the apical region of the parameres which resemble a bottle opener.

Only known from the type locality in Bolivia (Fig. 13A).

Nothing is known about the habitat of this species as it was collected at lights.

Among the thousands of specimens studied here, we only found one specimen of this species, which might be an indicator of their rarity in nature, or a reflection of the lack of sampling in the lowlands of Bolivia. The specimen is in modest condition, with one maxillary palp and a few tarsi missing.

Novochares garfo species group

Species group diagnosis. Body length 4.9–6.5 mm. Coloration: Dorsal surfaces pale brown (orange to yellowish), with slightly paler margins of pronotum and elytra, sometimes also clypeus. Head: Maxillary palps slightly to 1.3× longer than width of head, uniformly orange or yellow in color. Thorax: Ground punctation on pronotum and elytra relatively dense and shallowly impressed. Elytra without rows of serial punctures, each with very faint rows (sometimes one dorsal and two or three lateral, more usually only lateral) of scarce and weakly marked systematic punctures. Prosternum medially broadly and weakly convex. Posterior elevation of mesoventrite broadly elevated, somewhat transverse, often with glabrous longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite relatively deep, U- or V-shaped. Aedeagus: (Fig. 12) Overall shape sub-rectangular or oval to pear-like, joint basal margins of parameres truncate to medially pointed or medially emarginate in dorsal view; outer margin and apical region of each paramere weakly to strongly pointed; parameres longer than median lobe, with apex rounded; parameres with apical region variable in degree of sclerotization; dorsal inner margin of each paramere straight to sinuate; dorsal plate of median lobe (in dorsal view) forming a narrow neck; notch between arms variable; arms usually parallel, variable in shape and length; gonopore placed at or near base of dorsal plate of median lobe; ventral plate of median lobe moderately sclerotized, triangular, slender, and apically rounded, not reaching base of fork of dorsal plate; basal piece nearly 0.3× length of a paramere. In lateral view, aedeagus somewhat triangular, straight to weakly oblique at base, with ventral outline of parameres 3–4× longer than greatest width.

Figure 12. 

Aedeagi of the Novochares garfo species group A–C N. bidens D–F N. furcatus G–J N. garfo K–M N. tenedor A, D, G, K dorsal view B, E, H, L ventral view C, I, M lateral view F, J oblique view.

Composition. The Novochares garfo species group is composed of four species: Novochares bidens sp. nov., N. furcatus sp. nov., N. garfo sp. nov., and N. tenedor sp. nov.

Remarks. All four species in this group are relatively light brown/yellow in dorsal coloration, making them resemble the much more common members of the abbreviatus species group. Species in this group can be recognized by the relatively simple shape of the dorsal plate of the median lobe which resembles a bifid fork in all four species, as well as the shape of the parameres, which are typically straighter and with only a very small tooth near the apex (except N. tenedor, in which the apex is more curved and well developed).

Holotype (male): “BRAZIL, M.G./ Jacare, P.N. Zingu/ XI-1965, at lite/ M. Alvarenga” (USNM). Paratypes (2 exs.): Brazil: Mato Grosso: Tapirape Indian Village at confluence of R. Tapirape and R. Araguaia, 26–31.xii.1960, leg. B. Malkin, at light (1, FMNH). Mato Grosso: Same data as holotype (1, SEMC).

This species is easily distinguished from others in the species group by the extremely large and deeply cleft dorsal plate of the median lobe, and the unusually narrow and apically rounded parameres (Fig. 12A, B).

Body length 5.9–6.4 mm. Coloration: Dorsal surfaces pale (yellowish) brown, with slightly paler (orange) margins of pronotum and elytra. Head: Maxillary palps nearly 1.3× longer than width of head, uniformly orange in color. Thorax: Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum medially weakly convex. Posterior elevation of mesoventrite broadly elevated, posteriorly somewhat transverse, with low and glabrous longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite relatively deep, U-shaped. Aedeagus: (Fig. 12A–C) Overall shape sub-rectangular, 2.2× longer than wide, with outer lateral margins of parameres weakly sinuate; joint basal margins of parameres medially pointed in dorsal view; apical region of each paramere rounded, partly membranous, with outer margin weakly pointed; at closest point (near base of neck), dorsal inner margins of parameres separated by nearly 0.8× greatest width of a paramere; dorsal plate of median lobe with neck 0.28× as broad as base; arms of dorsal plate of median lobe dorsally concave, with inner margins nearly parallel, gradually narrowing towards apex, nearly 0.4× length of dorsal plate of median lobe; each arm acute at apex; notch between arms at base nearly as wide as base of an arm; basal piece nearly 0.3× length of a paramere. In lateral view, aedeagus straight at base, with ventral outline of parameres 4.2× longer than greatest width near base; dorsal outline of aedeagus in lateral view very weakly convex along basal 1/2, then concave along distal 1/2; ventral outline of aedeagus in lateral view sinuate.

Bidens (L.), meaning two-pronged fork, referring to the shape of the dorsal plate of the median lobe of this species.

Only known from the type locality in central Brazil (Fig. 13B).

Figure 13. 

Distribution of Novochares spp. A N. aperito (red), N. minor (yellow), N. orchis (blue) B N. bidens (red), N. furcatus (yellow), N. garfo (blue), N. tenedor (green).

Nothing is known about the habitat of this species.

Holotype (male): “BRAZIL: Mato Grosso do Sul/ -20.51369°, -55.42803°, 240 m/ Palmeiras (c. 7 km S) on MS-450/ 22.vi.2018; leg. Hamada & team/ Pond in field w/dense vegetation/ BR18-0622-01A” (INPA). Paratype (1 ex.): Brazil: Rondônia: Machadinho d’Oeste, Balneario São Jose, -9.44573, -61.98332, 103 m, 9.vii.2018, leg. Short, margins of various places along river, BR18-0709-01A (1, SEMC, DNA voucher SLE2097).

Among members of this species group, this species is most similar to N. garfo: both species share a relatively straight and parallel-sided dorsal plate of the median lobe with two relatively short arms at the apex (Fig. 12D, H). These arms are distinctly longer in N. furcatus (Fig. 12D), while they are barely noticeable in N. garfo (Fig. 12G). Additionally, the parameres are slightly narrower along the apical 1/3 in N. furcatus than in N. garfo.

Body length 5.6 mm. Coloration: Dorsal surfaces pale (yellowish) brown, with slightly paler (yellow) margins of pronotum and elytra. Head: Maxillary palps slightly longer than width of head, uniformly yellow in color. Thorax: Elytra without rows of serial punctures, each with very faint rows of scarce and weakly marked systematic punctures on lateral region. Prosternum broadly and weakly convex. Posterior elevation of mesoventrite somewhat transverse and broadly elevated. Abdomen: Apical emargination of fifth ventrite relatively deep, U-shaped. Aedeagus: (Fig. 12D–F) Overall shape pear-like, 2.2× longer than wide, with outer lateral margins of parameres evenly convex up to apical region; joint basal margins of parameres medially emarginate in dorsal view; apical region of each paramere rounded to truncate, partly membranous, with outer margin weakly pointed; at closest point (near base of neck), dorsal inner margins of parameres separated by distance slightly narrower than greatest width of a paramere; dorsal plate of median lobe with neck 0.3× as broad as base; base of arms of dorsal plate of median lobe dorsally concave; arms of dorsal plate nearly parallel, gradually narrowing towards apex, nearly 0.16× length of dorsal plate of median lobe; each arm acute and dorsally pointed at apex; notch between arms at base slightly narrower than base of an arm; basal piece nearly 0.32× length of a paramere. In lateral view, aedeagus weakly oblique at base, with ventral outline of parameres 4× longer than greatest width near base; dorsal outline of aedeagus in lateral view evenly convex along basal 2/3, then nearly straight along distal 1/3; ventral outline of aedeagus in lateral view nearly straight.

Furcatus (L.), meaning split in two, referring to the shape of the dorsal plate of the median lobe of this species.

Known from two localities in the Brazilian states of Mato Grosso do Sul and Rondônia (Fig. 13B).

One specimen was taken in the margins of an open pond, the other was taken along the margins of a river.

Holotype (male): “BRAZIL: Roraima: Caroebe/ 00°50.939'N, 59°18.262'W; 145m/ Rio Jatapu, nr. Usina de Jatapu;/ marginal pools of river; 17.i.2018/ leg. A. Short; BR18-0117-02A (INPA). Paratypes (11 exs.): Brazil: Amazonas: Manacapuru, -3.23037, -60.64269, 35 m, 9.vi.3017, leg. Benetti, margin of large marsh, BR17-0609-01A (1, SEMC, DNA voucher SLE1263); Presidente Figueiredo, (ca. 19 km E) on AM-240, Igarape Pantera, -2.04243, -59.84914, 17.i.2018, leg. Short, margin of small side stream, vegetation, and detritus, BR18-0617-01A (1, SEMC, DNA voucher SLE1931). Mato Grosso do Sul: Miranda (ca. 9.5 km SW) on MS-339, -20.32119, -56.42563, 131 m, 26.vi.2018, leg. Hamada & team, marshy area alongside stream, BR18-0626-03A (3, SEMC, including DNA voucher SLE2103). Pará: Rio Xingu Camp, ca. 60 km S. Altamira, Igarape Jabuti, 8–16.x.1986, leg. P. Spangler & O. Flint, malaise trap (2, USNM). Roraima: Same data as holotype (3, INPA, SEMC). Bolivia: Santa Cruz: 3.7 km SSE Buena Vista, Hotel Flora y Fauna, 1–12.v.2004, leg. A.R. Cline, MV+HG lights (1, SEMC).

See differential diagnosis of N. furcatus.

Body length 4.9–5.8 mm. Coloration: Dorsal surfaces pale (yellowish) brown, usually with slightly paler (yellow) clypeus and margins of pronotum and elytra. Head: Maxillary palps slightly longer than width of head, uniformly yellow in color. Thorax: Elytra without rows of serial punctures, each with very faint rows of scarce and weakly marked systematic punctures on lateral and posterior regions. Prosternum medially broadly and weakly convex. Posterior elevation of mesoventrite somewhat transverse and broadly elevated, with low and glabrous longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite relatively deep, U- or V-shaped. Aedeagus: (Fig. 12G–J) Overall shape pear-like, 2.1× longer than wide, with outer lateral margins of parameres evenly convex up to apical region; joint basal margins of parameres medially emarginate in dorsal view; apical region of each paramere rounded, partly membranous, with outer margin ventro-laterally pointed; at closest point (near base of neck), dorsal inner margins of parameres separated by distance 0.8× greatest width of a paramere; dorsal plate of median lobe with neck 0.38× as broad as base; base of arms of dorsal plate of median lobe dorsally concave; arms of dorsal plate parallel, parallel-sided along entire length, nearly 0.1× length of dorsal plate of median lobe; each arm acute and dorsally pointed at apex; notch between arms at base slightly narrower than base of an arm; basal piece nearly 0.31× length of a paramere. In lateral view, aedeagus weakly oblique at base, with ventral outline of parameres 3.3× longer than greatest width near base; dorsal outline of aedeagus in lateral view very weakly convex along basal 1/2, then concave along distal 1/2; ventral outline of aedeagus in lateral view nearly straight.

Garfo, meaning fork in Portuguese, in reference to the shape of the dorsal plate of the median lobe.

Brazil (Amazonas, Mato Grosso do Sul, Pará, Roraima), Bolivia (Fig. 13B).

This species has been collected in open marshes as well as along the margins of open rivers.

Holotype (male): “VENEZUELA: Apure State/ 7°37.289'N, 69°3.679'W, 83m/ side road ca. 10 km E. Mantecal/ leg. Short, García, & Camacho/ 18.i.2009; marshy area and pool by road; VZ09-0118-02X” (MIZA). Paratypes (59 exs.): Guyana: Region 6: Upper Berbice, Basecamp 1, 4°08.809'N, 58°14.232'W, 108 m, 22.ix.2014, leg. Short, Salisbury, La Cruz, margin of Berbice River, GY14-0922-02A (36, SEMC, CBDG, TTU-Z, including DNA voucher SLE1219). Region 9: Karanambu, 3°45.1'N, 59°18.6'W, Rupununi River, 2.iv.1994, leg. P.J. Spangler, colln #8 (1, USNM). Venezuela: Apure: Same data as holotype (1, SEMC, DNA voucher SLE1205). Barinas: SW of Batatuy, 8°10.259'N, 70°51.866'W, 275 m, 25.i.2012, leg. Short, Arias, & Gustafson, sandbar/gravel margin, VZ12-0125-03C (1, SEMC). Bolívar: Cuchivero, 30 km SE of Caicara, 4.viii.1987, leg. S. & J. Peck, Woodland, UV light, SBP87-108 (6, SEMC). Cojedes: Rio Tinaco, near, approx. 5 km NE of Tinaco, 9°44.160'N, 68°24.219'W, 170 m, 20.i.2012, leg. Short, Arias, & Gustafson, stream margins, VZ12-0120-02A (4, SEMC); El Pao, approx. 7.5 km, Rio Caiman Grande at San Brano, 9°39.246'N, 68°11.860'W, 137 m, 20.i.2012, leg. Short, Arias, & Gustafson, stream margins, VZ12-0120-03A (3, SEMC); Aparicion, at highway, lagoon/pond, 9°22.268'N, 69°23.062'W, 213 m, 22.i.2012, leg. Short, Arias, & Gustafson, pond, VZ12-0122-01A (2, SEMC). Portuguesa: Aparicion, Rio Are, 9°22.900'N, 69°23.153'W, 220 m, 22.i.2012, leg. Short & Arias, river margin, VZ12-0122-02A (5, SEMC). Zulia: Quebrada Riencito, 10.86041°N, 72.32210°W, 95 m, 30.xii.2008, leg. Short & García, along margin, VZ08-1230-01B (1, SEMC).

This taxon is unique among members of this species group in having the parameres strongly sinuate along the apical 1/3 (Fig. 12K, L). While the dorsal plate of the median lobe is slightly similar in form to N. bidens (Fig. 12A, B), the arms of the fork are less than 1/2 as long, and the plate itself is much shorter, sitting well below the apex of the parameres.

Body length 5.1–6.5 mm. Coloration: Dorsal surfaces orange to yellowish brown, usually with slightly paler (yellow) clypeus and margins of pronotum and elytra. Head: Maxillary palps slightly nearly 1.3× longer than width of head, uniformly orange in color. Thorax: Elytra without rows of serial punctures, each with very faint rows of scarce and weakly marked systematic punctures on lateral and posterior regions. Prosternum medially broadly and weakly convex. Posterior elevation of mesoventrite somewhat transverse and broadly elevated, with low and glabrous longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite relatively deep, U-shaped. Aedeagus: (Fig. 12K–M) Overall shape pear-like, 2.3× longer than wide, with outer lateral margins of parameres very weakly convex up to apical region; joint basal margins of parameres medially pointed in dorsal view; apical region of each paramere rounded, with outer margin laterally pointed; at closest point (along neck), dorsal inner margins of parameres separated by distance 0.54× greatest width of a paramere; dorsal plate of median lobe with neck 0.23× as broad as base; base of arms of dorsal plate of median lobe dorsally concave; arms of dorsal plate somewhat parallel, diverging at apex, nearly parallel-sided for most length, nearly 0.16× length of dorsal plate of median lobe; each arm acute and dorso-laterally pointed at apex; notch between arms at base nearly 1/2 as wide as base of an arm; basal piece nearly 0.3× length of a paramere. In lateral view, aedeagus straight at base, with ventral outline of parameres 3.9× longer than greatest width near base; dorsal outline of aedeagus in lateral view strongly convex at base, then nearly straight and oblique along basal 1/2, then sinuate along distal 1/2; ventral outline of aedeagus in lateral view nearly straight.

Tenedor, meaning fork in Spanish, in reference to the shape of the dorsal plate of the median lobe.

Guyana, Venezuela (Fig. 13B).

This species has been collected in a range of habitats, though most specimens seem to have been taken in riparian areas along the margins of streams and rivers. Other specimens have been collected in more typical marsh habitats.

Holotype (male): “PERU: Loreto: Maynas Province/ 3°50.723'S, 73°22.187'W, 113m/ ca. 10km SW Iquitos, nr. Facultad/ de Ciencias Biologicas UNAP/ leg. S. Baca, 18.i.2020/ seasonal pond; PE20-0118-03A” (MHNSM). Paratypes (41 exs.): Peru: Loreto: Same data as holotype (27, NZCS, SEMC, TTU-Z, including DNA voucher SLE2143); ca. 15 km SW Iquitos, on Iquitos-Nauta Highway, leg. S. Baca, 18.i.2020, flooded area with vegetation and detritus, PE20-0118-05A (10, SEMC); ca. 20 km SW Iquitos, on Iquitos-Nauta Hwy, 3°56.655'S, 73°23.853'W, 107 m, leg. S. Baca, 20.i.2020, shallow margins of lake, with vegetation/detritus; PE20-0120-01A (1, SEMC); ca. 60 km SW Iquitos, on Iquitos-Nauta Hwy, 4°16.279'S, 73°30.734'W, 95 m, leg. S. Baca, 20.i.2020, margin of small creek, inundated grass, PE20-0120-02A (1, SEMC). Suriname: Para: along Martin Luther King Hwy, blackwater marsh by road, 5.4204, -55.09876, SR12-0723-04A (1, SEMC, DNA voucher SLE535). Venezuela: Monagas State: S of Maturin, morichal at road crossing, 9°16.398'N, 62°56.246'W, 22 m, 2.ii.2010, leg. Short, García, & Joly, morichal margin, VZ10-0202-02A (1, MIZA).

See species group diagnosis.

Body length 4.2–5.4 mm. Coloration: Dorsal surfaces very dark brown, with paler (orange to yellow) margins of pronotum and elytra. Head: Maxillary palps slightly longer than width of head, orange to brown in color, usually paler (yellow) at ends of each palpomere (Fig. 7C). Thorax: Ground punctation on pronotum and elytra relatively dense and shallowly impressed. Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum only very weakly medially convex. Posterior elevation of mesoventrite broadly and somewhat transversely elevated; mesoventrite with medial longitudinal glabrous patch extending anteriorly. Abdomen: Apical emargination of fifth ventrite relatively deep, U-shaped. Aedeagus: (Fig. 11G–K) Overall shape pear-like, 2.1–2.3× longer than wide, with outer lateral margins of parameres convex up to apical region; apical region of each paramere rounded; at closest point (along neck), dorsal inner margins of parameres separated by distance slightly narrower than greatest width of a paramere; dorsal plate of median lobe with neck 0.5× as broad as base; base of arms of dorsal plate of median lobe dorsally concave; arms of dorsal plate somewhat parallel to slightly diverging, nearly parallel-sided for most length, nearly 0.23× length of dorsal plate of median lobe; each arm truncate at apex, sometimes obliquely so, inner margin extending beyond outer margin; notch between arms at base nearly 0.4× base of an arm; basal piece nearly 0.3× length of a paramere. In lateral view, ventral outline of parameres 3.5× longer than greatest width near base; dorsal outline of aedeagus in lateral view strongly convex at base, then very weakly convex along basal 1/2, then sinuate along distal 1/2; ventral outline of aedeagus in lateral view sinuate.

Minor, named after its small body size.

Peru, Suriname, Venezuela (Fig. 13A).

This species has been found primarily in open marshes and along the margins of vegetated creeks.

There are small differences in overall length/width of aedeagus and the shape of the apicolateral region of the arms of the dorsal plate of the median lobe between specimens from Peru and Venezuela. The small size and form of this species allow it to be easily confused with Sindolus Sharp, but that genus can easily be separated by the strongly raised longitudinal carina of the mesoventrite, which is absent in Novochares.

Holotype (male): “SURINAM/ Suriname Dist./ Krakka-Phedra Rd./ X-25-1962/ Borys Malkin”, “tiny pool in/forest, much/ fallen foliage” (USNM). Paratypes (57 exs.): Brazil: Amapá: Oiapoque (ca. 22 km S) on BR -156, 3.65822, -51.76958, leg. Short, forested detrital pools, BR18-0720-01B (1, SEMC, including DNA voucher SLE1851). Amazonas: Manaus Ducke Reserve, Igarape Barro Branco, -2.93079, -59.97514, 75 m, 6.vi.2018, leg. Short & team, stream margins, BR18-0606-02B (11, INPA, SEMC, TTU-Z); same data except forest pools/riparian area by stream, BR18-0606-02C (1, SEMC); same data except shallow pools, BR18-0606-02D (1, SEMC); same data except 9.vi.2018, muddy pools in swampy area by stream, BR18-0609-02B (1, SEMC). Rondônia: Tabajara (ca. 4.5 km W) on RO-133, -8.9217, -62.0978, 100 m, 8vii.2018, leg. Short, detrital pool/marsh by stream, BR18-0708-02B (2, SEMC). French Guiana: Piste de montagne de fer, 5.37641, -53.54782, 67 m, 3.iii.2020, leg. Short & Neff, large shallow detrital pool by road, FG20-0303-02A (4, SEMC); Forêt des Sables Blancs Park, 3.iii.2020, leg. Short & Neff, detrital puddle in forest, FG20-0303-03A (2, SEMC); Bagne des Annamites Park, Crique Anguille, 4.83287°N, -52.5145°W, 17 m, leg. Short & Neff, small sandy stream with detritus, FG20-0307-01B (5, SCC, SEMC). Suriname: Suriname: same data as holotype (25, SEMC, UNSM). Sipaliwini District: Camp 4 (low), Kasikasima; sandy/ creek, trail to Kasikasima, 2.97731°N, 55.38500°W, 200 m, 22.iii.2012, leg. A. Short, SR12-0322-02A (4, NSCS, SEMC, including SLE1214).

See species group diagnosis.

Body length 7.3–9.3 mm. Coloration: Dorsal surfaces dark brown and sheeny, with paler (brown or reddish brown) clypeus and margins of pronotum and elytra. Head: Maxillary palps nearly 1.3× longer than width of head, uniformly reddish brown in color (Fig. 24B). Thorax: Ground punctation on pronotum and elytra relatively dense and very shallowly impressed. Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum only very weakly medially convex. Posterior elevation of mesoventrite elevated as a triangular pyramid, with medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite relatively deep, V- or U-shaped. Aedeagus: (Figs 4B 11D, F) apical region of outer margin of each paramere pointed, dorsally directed; parameres longer than median lobe; at closest point, dorsal inner margins of parameres separated by distance 0.5× greatest width of a paramere; dorsal plate of median lobe with neck 0.5× as broad as base; arms of dorsal plate of median lobe abruptly broadening at mid-length, slightly converging at apex, with apex rounded; apical region of outer margin of each arm pointed; arms nearly 0.55× length of dorsal plate of median lobe; notch between arms at base nearly as broad as base of an arm; ventral plate of median lobe apically truncate, apex extending to apical third 1/4 of dorsal plate; basal piece 0.35× length of a paramere. In lateral view, aedeagus somewhat teardrop-shaped, with ventral outline of parameres 4× longer than greatest width near base; dorsal outline nearly evenly convex along entire length.

This species is named after the complex and intricate shape of the aedeagus, which we had informally named the “orchid one” during the course of this revision.

Known from Brazil (Amapá, Amazonas, Rondônia), French Guiana, and Suriname (Fig. 13A).

This species is most commonly found along detrital margins of streams and in forested pools associated with streams.

Novochares punctatostriatus species group

Species group description. Body length 4.7–8.8 mm. Coloration: Dorsal surfaces dark brown, usually with paler (orange to yellow) margins of head, pronotum, and elytra. Head: Maxillary palps slightly shorter to slightly longer than width of head, uniformly yellow to orange in color (Fig. 14). Thorax: Ground punctation on pronotum and elytra relatively dense and shallowly impressed. Elytra sometimes with defined rows of serial punctures, at least along posterior and lateral areas, not forming grooves. Prosternum flat to medially convex. Posterior elevation of mesoventrite transverse, usually blunt and low. Abdomen: Apical emargination of fifth ventrite small and shallow, slightly broader than deep. Aedeagus: (Figs 4A, 15) Overall shape sub-rectangular, approximately 2× as long as wide, joint basal margins of parameres medially emarginate and laterally rounded; outer margin of each paramere nearly straight or very weakly convex along basal 2/3, then concave and laterally pointed, then oblique to apex; parameres longer than median lobe; apex rounded; parameres with apical region cylindrical (hollow to apex) and lightly sclerotized; dorsal inner margin of each paramere sinuate; dorsal plate of median lobe (in dorsal view) with strongly sclerotized basal apodemes (nearly 0.25× length of plate); dorsal plate of median lobe nearly parallel-sided along basal 2/3, then bifurcating with parallel to distally converging arms; notch between arms variable; shape and orientation of arms variable; gonopore sitting proximal to base of fork of dorsal plate; ventral plate of median lobe (in ventral view) somewhat triangular, variable in length, shape of apex, and degree of sclerotization; dorsal surface of ventral plate of median lobe slightly concave (sides curved dorsally); basal piece nearly 0.25–0.40× length of a paramere, with distal margin straight. In lateral view, aedeagus triangular, strongly oblique at base, with ventral outline of parameres 3–5× longer than greatest width near base.

Figure 14. 

Dorsal habitus of Novochares spp. A N. dentatus B N. punctatostriatus C N. pertusus D N. triangularis.

Figure 15. 

Aedeagi of the Novochares punctatostriatus species group A–D N. dentatus E, F N. spangleri G, H N. geminus I N. triangularis J, K N. pertusus L–Q N. punctatostriatus A–C Venezuela D Ecuador L–N Suriname O Peru P, Q Brazil. A, E, G, I, J, L, O, P dorsal view H, M, Q ventral view C, F, N oblique view K lateral view.

Composition. The Novochares punctatostriatus species group is composed of six species: Novochares dentatus sp. nov., N. geminus sp. nov., N. pertusus sp. nov., N. punctatostriatus sp. nov., N. spangleri sp. nov., and N. triangularis sp. nov.

Remarks. This species group is united by several distinct morphological characters and was recovered as a monophyletic group by Short et al. (2021). However, our two-gene analysis did not recover the monophyly of this species group, instead recovering two distinct, sequentially diverging lineages (Fig. 1). These two subgroups can be morphologically differentiated: the lineage that contains N. dentatus, N. geminus, and N. spangleri have lateral teeth on the dorsal plate of the median lobe of the aedeagus combined with an extended ventral plate, while the other subgroup does not possess these features (Fig. 15).

Holotype (male): “VENEZUELA: Amazonas State/ 5°20.514'N, 67°45.315'W, 87m/ S. Communidad Porvenir/ 15.i.2009; leg. Miller & Short/ VZ09-0115-03B/ small streamlet (MIZA). Paratypes (57 exs.): Venezuela: Amazonas: 5 km N. Galipero, Pozo Azul, 25.i.1989, leg. Spangler, Faitoute, & Barr, “roots, stream edge” (1, USNM); same data as holotype (25, MIZA, SEMC, including DNA Voucher SLE1199); ca. 15 km S. Puerto Ayacucho, large rock outcrop, 14.ix.2007, leg. Short, pools at base of outcrop, AS-07-011a (2, SEMC); same locality but 8.viii.2008, leg. Short & García, pools at base of outcrop, AS-08-081a (4, SEMC); same locality but 14.i.2009, leg. Short, “rock pools et al”, VZ09-0114-03B (5, SEMC); Tobogan de la Selva, 5.i.2006, leg. Short, pools in rock w/sand, AS-06-011c (1, SEMC); nr. Iboruwa, 7.viii.2008, leg. Short, García, & Joly, AS-08-078 (6, SEMC); Puerto Ayacucho (39 km S.), Samariapo road, 15.xi.1987, leg. Spangler & Faitoute, “brook”, Collection #4 (1, USNM). Ecuador: Pastaza: AGIP platform Villano B, along transect 1 and 2, 24.v.2008, leg. Short, small forest stream, AS-08-008b (12, PUCE, SEMC, including DNA voucher SLE1188).

The distinctive rows of serial punctures on the lateral and posterior margins of the elytra serve to separate this species from all other Novochares except a few others in the punctatostriatus species group, particularly N. punctatostriatus which also occurs throughout much of the Amazonian region. However, as far as is known, the ranges of the two species do not overlap, with N. dentatus being more northern and western in distribution. The two species can be distinguished by the impression of the rows of serial punctures, which are more impressed and prominent in N. punctatostriatus, as well as the presence of small denticles on the aedeagus (Fig. 15A; denticles absent in N. punctatostriatus, e.g., Fig. 15L).

Size and form: Body length 4.7–6.5 mm. Coloration: Dorsal surfaces dark brown, with slightly paler margins of pronotum and elytra, occasionally paler clypeus; paler margin sometimes very wide. Head: Maxillary palps as long as to slightly longer than width of head, uniformly orange in color (Fig. 14A). Thorax: Each elytron with defined rows of serial punctures along posterior 1/3 and lateral 1/2. Prosternum medially weakly and broadly convex. Posterior elevation of mesoventrite transverse, low, and slightly curved (posteriorly concave). Aedeagus: (Figs 4A, 15A–D) lateral projection on apical region of outer margin of each paramere rounded to pointed; at closest point, dorsal inner margins of parameres separated by distance similar to greatest width of a paramere; dorsal plate of median lobe with small denticle on each lateral margin, proximal to base of fork; arms of dorsal plate of median lobe parallel to distally converging; each arm parallel sided, slightly wider on apical region, apically oblique with inner margin extending beyond outer margin; notch between arms at base nearly as broad as base of an arm; ventral plate of median lobe gradually more sclerotized distally, at widest point nearly as wide as maximum width of ventral face of a paramere, apically acuminate, apex extending beyond base of fork, not reaching apex of arms of dorsal plate, ventral surface of each side medially sharply elevated forming narrow medial slit; basal piece 0.3× length of a paramere.

Dentatus (L.), meaning toothed, in reference to the small, lateral, tooth-like projections on the dorsal plate of the median lobe.

Venezuela, Ecuador (Fig. 16B).

Figure 16. 

Distribution of Novochares punctatostriatus species group A N. punctatostriatus (red), N. pertusus (yellow) B N. dentatus (red), N. triangularis (yellow), N. spangleri (blue), N. geminus (green).

In Venezuela, this species was typically found along the margins of streams that were flowing on or near granite outcrops. The series from Ecuador was taken from a small forested stream with lots of detritus, though it did not appear to be associated with any rocky substrate.

Specimens from Ecuador tend to be smaller and more yellowish than specimens from Venezuela (specimens from the “Tobogancito” locality are similar in size to specimens from Ecuador). The eyes of specimens from Venezuela are relatively larger and more prominent than those of specimens from Ecuador.

Holotype (male): “BRAZIL: Mato Grosso do Sul/ -20.72281, -55.69127; 225 m/ Aquidauana (c. 27 m S) on/ MS-174; leg. Hamada & team;/27.vi.2018; seepage & debris nr. stream margin; BR18-0627-01E” (INPA). Paratypes (2 exs.): Brazil: Mato Grosso do Sul: Same data as holotype (2, SEMC, including DNA Voucher SLE2092).

Among species of the punctatostriatus species group, N. geminus is one of three species that lack distinct rows of elytral serial punctures, the others being N. spangleri and N. triangularis. Those two species can be separated by the shape of the forked projections of the median lobe, which are broader and swollen at the apex (Fig. 15 E, F, I), while they are slender and not expanded apically in N. geminus (Fig. 15G, H).

Size and form: Body length 5.9–6.0 mm. Coloration: Dorsal surfaces dark brown, with slightly paler margins of pronotum and elytra. Head: Maxillary palps as long as to slightly longer than width of head, uniformly orange in color. Thorax: Elytra without defined rows of serial punctures, each with one dorsal and a few lateral sparse rows of systematic punctures. Prosternum medially weakly and broadly convex. Posterior elevation of mesoventrite transverse, low, and blunt. Aedeagus: (Fig. 15G, H) lateral projection on apical region of outer margin of each paramere pointed; at closest point, dorsal inner margins of parameres separated by distance nearly 1/2 greatest width of a paramere; dorsal plate of median lobe with small denticle on each lateral margin, proximal to base of fork; arms of dorsal plate of median lobe parallel, diverging at apex; each arm parallel sided, with inner margins sinuate, apex rounded and pointing outwards; notch between arms at base nearly as broad as base of an arm; ventral plate of median lobe lightly sclerotized, at widest point nearly as wide as dorsal plate of median lobe, apically acuminate, apex nearly reaching apex of arms of dorsal plate, ventral surface of each side medially sharply elevated forming narrow medial slit; basal piece 0.35× length of a paramere.

Geminus (L.), meaning twin, in reference to the split apex of the dorsal plate of the median lobe.

Brazil (Mato Grosso do Sul) (Fig. 16B).

The only known series was taken from the margin of a rocky stream, where a seep was flowing over large rocks.

Holotype (male): “BRAZIL: Goiás/ Chapada dos Veadeiros/ 18–24km N. of Alto Paraíso/ 1400–1500m, 25.x.1985 leg. S.E. Miller” (USNM). Paratypes (14 exs.): Brazil: Goiás: Same data as holotype (14, USNM, SEMC).

The distinctive rows of serial punctures on the lateral and posterior margins of the elytra serve to separate this species from all other Novochares except a few others in the punctatostriatus species group. It is most similar to N. punctatostriatus, to which it is probably most closely related, by the form of the aedeagus which lacks small lateral teeth on the dorsal plate of the median lobe (Fig. 15J–Q). The comparatively long arms of the fork of the dorsal plate of the median lobe, and the more basal positioning of the ventral plate (Fig. 15J, K) serve to distinguish this species from N. punctatostriatus (Fig. 15L–Q).

Size and form: Body length 6.0–7.3 mm. Coloration: Dorsal surface of head very dark brown to nearly black; pronotum and elytra dark brown, with broad paler margins. Head: Maxillary palps as long as to slightly longer than width of head, uniformly orange in color (Fig. 14C). Thorax: Elytra with defined rows of serial punctures, except on central elytral disc. Prosternum medially broadly and very weakly convex. Posterior elevation of mesoventrite transverse, blunt, and low. Aedeagus: (Fig. 15J, K) lateral projection on apical region of outer margin of each paramere pointed; at closest point, dorsal inner margins of parameres separated by distance slightly narrower than greatest width of a paramere; lateral margins of dorsal plate of median lobe smooth, lacking denticles; arms of dorsal plate of median lobe nearly parallel, somewhat converging at apex; each arm with outer and inner margins slightly sinuate, apical region broader than base of arm, apex roundly truncate and oblique, with inner margin extending beyond outer margin; notch between arms at base narrower than base of an arm, slightly wider at basal 1/3, narrowing towards apex; ventral plate of median lobe strongly sclerotized, at widest point nearly as wide as dorsal plate of median lobe, with apex narrowly rounded, not reaching base of fork of dorsal plate; basal piece 0.28× length of a paramere.

Pertusus (L.), meaning perforated, in reference to the rows of fine serial punctures on the elytra.

Only known from the type locality in Brazil (Goiás) (Fig. 16A).

Nothing is known about the habitat of this species.

Holotype (male): “PERU: Madre de Dios/ Rio Tambopata Res.; 290m/ 30 air km SW of Puerto Maldonado/ 16–20.xi.1979; subtropical moist forest/ leg. J.B. Heppner” (USNM). Paratypes (262 exs.): Brazil: Amapá: Oiapoque (ca. 22 km S) on BR-156, leg. Short, forested detrital pools, BR18-0720-01B (1, SEMC, including DNA Voucher SLE2094). Amazonas: Tapauá, Humaita (ca. 240 km N) on BR-319, -5.50298, -62.12392, 54 m, 11.vii.2018, leg. Short, margin of stream, BR18-0712-01B (2, INPA, SEMC, including DNA Voucher SLE1969). Pará: Altamira (ca. 60 km S), Rio Xingu Camp, 52°40'W, 3°50'S, 12.x.1986, leg. P. Spangler & O. Flint (10, USNM). Rondônia: Machadinho d’Oeste, Tabajara (ca. 7.5 km W) on RO-133, -8.92368, -62.12491, 82 m, 8.vii.2018, leg. Short, river with sandy bottom and rocks, BR18-0708-04A (1, SEMC, including DNA Voucher SLE2037); Novo Uniao, Vale do Cachoeiras, -10.91764, -62.377, 359 m, 10.vii.2018, leg. Short, small sandy-bottom stream margin, BR18-0710-02A (1, SEMC, including DNA Voucher SLE2090). São Paulo: Piracicaba, dates between 10.ii.1965 and 2.xii.1965, blacklight, C.A. Triplehorn (68, USNM). French Guiana: Route de Petit Saut, Crique Maman Lézard, 5.06701°N, -52.99783°W, 39 m, 1.iii.2020, leg. Short & Neff, margin of creek with detritus, FG20-0301-01A (1, SEMC); Same data but detrital pools in drying creakbed, FG20-0301-01B (1, SEMC); Carbet ONF Montagne de fer, Piste de montagne de fer (formerly road Degrad Florian), Crique Petit Laussat, 5.40697°N, -53.55468°W, 10 m, leg. Short, detrital pools, FG20-0302-01C (3, SEMC); Piste de montagne de fer (formerly Degrad Florian road), tributary of Crique Florian, 5.29688°N, -53.52458°W, 25 m, leg. Short & Neff, small pools in stream channel with sand and detritus, FG20-0303-01A (3, SEMC); same data but leg. Short, margin of clearwater creek, FG20-0303-01B (2, SEMC); St. Laurent du Maroni, Sentier des Malgaches, 5.48627°N, -54.00238°W, 14 m, leg. Short & Neff, pond margins in secondary forest, FG20-0304-01A (1, SEMC); Piste de montagne de fer (formerly road Degrad Florian), 5.40697°N, -53.55468°W, 10 m, leg. Short & Neff, forested detrital pools, FG20-0305-01A (7, SCC, SEMC); Carbet communal St-Elie, Route de Saint-Elie, tributary of Crique Toussaint, 5.29653°N, -53.05205°W, 42 m, leg. Short & Neff, margins of clearwater stream, FG20-0305-03B (2, SEMC); Paracou, Station de recherche CIRAD, Crique Verlot, 5.27966°N -52.92846°W, 8 m, leg. Short & Neff, forested detrital pools, FG20-0306-01A (1, SEMC); Bagne des Annamites Park, Crique Anguille, 4.83287°N, -52.5145°W, 17 m, leg. Short & Neff, small sandy stream with detritus, FG20-0307-01B (2, SEMC). Guyana: Region 6: Upper Berbice circa 1 km west of Basecamp 1, 4°09.143'N, 58°11.207'W, 105 m, 22.iv.2014, leg, Short, Salisbury and La Cruz, margins of creek, GY14-0921-03H (1, SEMC). Region 8: Konawaruk River, Basecamp 2 (NARIL camp), 14.ix.2014, leg. Salisbury & La Cruz, small puddle along road, GY14-0914-03 (1, SEMC); Konawaruk River, Basecamp 2 (NARIL basecamp), 5°07.539'N, 59°06.732'W, 80 m, 15.ix.2014, leg. Salisbury and La Cruz, unnamed clear water creek, slow flowing and shallow, GY14-0915-02 (1, SEMC). Region 9: along road to Parabara, 2°09.557'N, 59°17.569'W, 268 m, 1.xi.2013, leg. Short, Isaacs and Salisbury, forest pools near Mushai Wao, GY13-1101-02A, (2, SEMC); Parabara, trail to mines, 2°05.095'N, 59°14.174'W, 250 m, 2.xi.2013, leg. Short, Isaacs and Salisbury, detrital pools in forest, GY13-1102-01A, (1, SEMC); Parabara north side of river, 2°06.492'N, 59°13.653'W, 274 m, 3.xi.2013, detritus margins and leaf packs, GY13-1103-02A (1, SEMC); Karaawaimin Taawa, Basecamp and surroundings, 2.42284N, 59.06157W, 11-13.iii.2022, leg. Short & Edward, detrital pools near camp, GY22-0311-01A (1, SEMC); pooled up sandy creek GY22-0311-01D (2, SEMC); Karaawaimin Taawa, Trail from Camp 1 to Camp 2, 14.iii.2022, leg. Short & Edward, palm swamp, GY22-0314-04A (4, SEMC); Karaawaimin Taawa, Camp 2 and surroundings, 15.iii.2022, leg. Short & Edward, stream with palm detritus, rocks, and sand, GY22-0315-01A (1, SEMC); Karaawaimin Taawa, Camp 3 and surroundings, 16-18.iii.2022, leg. Short & Edward, small pool in streambed, GY22-0316-01D (1, SEMC); same data but forest pools, GY22-0316-01C (3, SEMC); Karaawaimin Taawa, Camp 4 and surroundings, 18-21.iii.2022, leg. Short & Edward, small stream, GY22-0318-01C (8, SEMC); same data but pools in creekbed, GY22-0318-01D (8, SEMC); same data but second small stream, GY22-0318-01E (18, CBDG, SEMC); Karaawaimin Taawa, Trail between Camps 3 and 4, 21.iii.2022, leg. Short & Edward, small pool in forest, GY22-0321-01B (3, SEMC). Peru: Cuzco: Pilcopata, 600 m, 8-10.xii.1979 premonate moist forest, leg. J.B. Heppner (3, USNM); Pilcopata (ca. 3 km NE), on nearby mountain road, 30.v.2022, leg. Short et al., pools and roadside ditches, PE22-0530-01B (1, SEMC); same data but small stream and adjacent grassy pool, PE22-0530-01C (1, SEMC). Madre de Dios: same data as holotype (22, USNM, SEMC); Manu Pakitza, 12°7'S, 10°58'W, 250 m, 18.viii.1988, UV light, leg. O. Flight & N. Adams (1, USNM); same locality but 14–23.ix.1988, malaise traps, “trail 2, 1^st stream” (1, USNM); Amazonas Lodge, N Atalaya, 12°52.2'S, 71°22.6'W, 480 m, 10-13.xi.2007, leg. D. Brzoska, flight intercept trap, PER1B07 002 (2, SEMC); Villa Carmen Biological Station (ca. 2 km N of Pilcopata), South of Rio Piñipiñi, 26.v.2022, leg. Short et al., small streams in bamboo thicket, PE22-0526-01A (1, SEMC); same data but large marshy pool along trail with abundant detritus, PE22-0526-01E (1, SEMC); Villa Carmen Biological Station (ca. 2 km N of Pilcopata), North of Rio Piñipiñi, 28.v.2022, leg. Short et al., small detrital pools, PE22-0526-02F (3, MHNSM, SEMC). Suriname: Saramacca: Coesewijne Savanna, 6.iii.2012, leg. Short, forested pool in muddy road, SR12-0306-03B (9, SEMC). Sipaliwini: Camp 1 on Kutari River, 2°10.521'N, 56°47.244'W, 228 m, 20.viii.2010, leg. Short and Kadosoe, forest stream, CI-RAP Survey, forested swamp, SR10-0819-01A (7, SEMC); Iwaana Saamu, forest swamp, 26.viii.2010, leg. Short, SR10-0826-01A (1, SEMC); Camp 3, Werehpai, 2°21.776'N, 56°41.861'W, 237 m, 3-7.ix.2010, leg. Short and Kadosoe, pooled up detrital creek, SR10-0903-01A (3, SEMC, including DNA Voucher SLE452); same data except detrital forest pools, SR10-0903-02A (1,SEMC); same data except sandy forest creek, SR10-0904-01A (3, SEMC); Upper Palumeu, Camp 1, 2.47700°N, 55.62941°W, 275 m. leg. A. Short, 10–16.iii.2012, Flight Intercept Trap, SR12-0310-TN1 (3 SEMC); Raleighfallen Nature Reserve, trail to Raleighfallen, 04°42.480'N, 56°13.159'W, 24 m, 27.vii.2012, SR12-0727-03A (1, SEMC); same data but leg. C. McIntosh, detrital pools near creek in forest, SR12-0727-03D (2, SEMC); Raleighvallen Nature Reserve Voltzberg trail, 04°40.910'N, 56°11.138'W, 78 m, 30.vii.2012, SR12-0730-01A (1, SEMC); same data but detrital pools along stream, SR12-0730-01B (2, SEMC); Raleighfallen Nature Reserve, Fungu island, 04°43.459'N, 56°12.658'W, 30 m, 1.viii.2012, SR12-0801-01D (2, SEMC); Raleighvallen Nature Reserve, base of Voltzberg, 4°40.432’N, 56°11.079’W, 86 m, 16.iii.2016, leg. Short et al., pooled up stream, SR16-0316-01B (2, SEMC); Raleighvallen Nature Reserve, Lolopaise area, 4°42.48'N, 56°13.15908'W, 24 m, intermittent stream pools, 19.iii.2016, leg. Toussaint et al., SR16-0319-02C (1, SEMC); Raleighvallen Nature Reserve, Coppename River, Voltzberg trail, 17.iii.2016, leg. A. Short, detrital pools in stream bed, SR16-0319-01A (10, NZCS, SEMC); Kabalebo Nature Resort, Moi Moi Creek, leg. Short, detrital pool, SR19-0310-01G (3, SEMC, including DNA Voucher SLE1802); same data except leg. Short and class, SR19-0310-01M (2, SEMC). Suriname: Krakka-Phedra Road, 25.x.1962, leg. B. Malkin (12, USNM).

The distinctive rows of serial punctures on the lateral and posterior regions of the elytra serve to separate this species from all other Novochares except a few others in the punctatostriatus species group. This species is the most commonly encountered and widespread of the six known species in the group, occurring from the eastern Guiana Shield (Guyana, Suriname), west to the foothills of the Peruvian Andes, and south São Paulo, Brazil. It is also one of the largest species in the group. The lack of lateral denticles on the dorsal plate of the median lobe separates this from other punctatostriatus species group taxa except for N. pertusus sp. nov. (see diagnosis of that species).

Size and form: Body length 5.2–8.8 mm. Coloration: Dorsal surface of head dark brown, sometimes with gradually paler clypeus and labrum; pronotum and elytra dark brown, with broad pale margins. Head: Maxillary palps as long as to slightly longer than width of head, uniformly orange in color (Fig. 14B). Thorax: Ground punctation on pronotum and elytra relatively dense and shallowly impressed. Elytra with defined rows of serial punctures, except on central elytral disc. Prosternum flat to medially weakly and broadly convex. Posterior elevation of mesoventrite transverse, blunt, and low. Aedeagus: (Fig. 15L–Q) lateral projection on apical region of outer margin of each paramere pointed and strongly protruded; at closest point, dorsal inner margins of parameres separated by distance slightly narrower than greatest width of a paramere; lateral margins of dorsal plate of median lobe smooth, lacking denticles; arms of dorsal plate of median lobe slightly converging at apex; each arm with outer and inner margins straight at base, curved at apex (outer margin concave, inner margin convex); apical region of arm broader than base of arm, apex broadly rounded and oblique, with inner margin extending beyond outer margin; notch between arms very narrow, much narrower than base of an arm, narrowing even more towards apex; ventral plate of median lobe moderately sclerotized, at widest point seemingly wider than dorsal plate of median lobe, with apex broadly to narrowly rounded, reaching to slightly surpassing base of fork of dorsal plate; basal piece 0.35× length of a paramere.

Puctatostriatus (L.) in reference to the distinct rows of elytral serial punctures.

Brazil (Amapá, Amazonas, Rondônia, São Paulo), French Guiana, Guyana, Peru, and Suriname (Fig. 16A).

This species is most commonly found in the detrital margins of densely forested lowland streams. Some collections have also been made in forested detrital pools, especially those that are riparian in origin. It has also been collected at lights and in malaise traps.

This relatively distinct and widespread taxon is very likely a complex of very closely related species. The pairwise genetic divergence in COI is nearly 9% among the ten individuals we sequenced from Peru to Suriname and French Guiana. This is the largest observed intraspecific divergence among any Novochares examined here, though not unprecedented in Acidocerinae: both Helochares maculicollis Mulsant, 1844 and H. normatus (LeConte, 1861) showed intraspecific variation of greater than 9% (Short and Girón 2018) but these too also likely represent species complexes. We examined the aedeagus from a variety of populations, and indeed there are subtle variations. The apex of the ventral plate of the median lobe varies in length and shape. The width of the notch between the arms of the dorsal plate of the median lobe can be very narrow and be essentially not visible to be as broad as 1/4 the width of an arm at base. However, we were not able to correlate this variation with any structure in the molecular tree. We have taken a conservative approach, and favor treating all these populations as one widespread, somewhat variable species.

Holotype (male): “PERU: Cusco: Paucartambo/ -12.91411S, -71.37492W, 585m/ c. 3km NE of Pilcopata; 30.v.2022/ grassy pools/ditch along road/ PE22-0530-01C; leg. Short et al.” (MHNSM). Paratypes (6 exs.): Peru: Cusco: same data as holotype (1, SEMC); same data except mountain side pools and ditches, PE22-0530-01B (2, SEMC). Madre de Dios: Villa Carmen Biological Station (ca. 2 km N of Pilcopata), North of Rio Piñipiñi, 26.v.2022, leg. Short et al., small muddy pools in landslide, PE22-0526-02B (1, SEMC); same data except 28.v.2022, detrital pools formed by seeps, PE22-0526-02F (2, MHNSM, SEMC).

Among members of the punctatostriatus species group, N. spangleri is one of three species that lack distinct rows of elytral serial punctures, the others being N. pertusus and N. triangularis. It is most similar to N. triangularis, to which the aedeagal form is very similar, though it can be distinguished by the apices of the forked projection of the dorsal plate of the median lobe being slightly more swollen, with the outer margins of the parameres weakly curved (apices of the forked projection of the dorsal plate of the median lobe being not as swollen and the outer margins of the parameres strongly curved in N. triangularis; compare Fig. 15E vs. Fig. 15I). The elytral ground punctation is extremely fine and almost not noticeable in N. spangleri, while it is distinctly coarser in N. triangularis. Additionally, N. triangularis has only very narrow paler margins of the pronotum, these margins are much more broadly pale in N. spangleri. See diagnosis of N. pertusus for separation from that species.

Body length 5.6–6.0 mm. Coloration: Dorsal surface of head, labrum, pronotum, and elytra dark brown, gradually paler towards margins. Head: Maxillary palps slightly shorter than width of head, uniformly yellow in color. Thorax: Ground punctation on pronotum and elytra relatively dense and very shallowly impressed. Elytra without rows of serial punctures, except for rows of very weak serial punctures along lateral regions of each. Prosternum medially convex. Posterior elevation of mesoventrite transverse, blunt, and low. Aedeagus: (Fig. 15E, F) lateral projection on apical region of outer margin of each paramere rounded to weakly pointed; at closest point, dorsal inner margins of parameres separated by distance slightly greater than greatest width of a paramere; dorsal plate of median lobe with small denticle on each lateral margin, proximal to base of fork; arms of dorsal plate of median lobe distally weakly converging; each arm slightly wider at base and widened on apical region, apically oblique with inner margin extending beyond outer margin; notch between arms at base 1/2 as wide as base of an arm; ventral plate of median lobe moderately sclerotized, at widest point as wide as dorsal plate of median lobe, apically acuminate, apex extending beyond base of fork, not reaching apex of arms of dorsal plate; basal piece 0.4× length of a paramere.

Named after Paul J. Spangler, longtime curator at the US National Museum of Natural History, Smithsonian Institution, and specialist on aquatic beetles, who collected and sorted a lot of the specimens included in this contribution.

Peru (Fig. 16B).

This species was collected primarily in forested riparian habitats.

Holotype (male): “BOLIVIA: Santa Cruz Dept./Potrerillos del Guenda,/Preserva Natural, 17°40'S, 63°27'W, 370m, 17-22-X-2007/ ex BL/MV, J. & F. Romero/ BOL1Cline07 007.5” (SEMC). Paratypes (25 exs.): Brazil: Goiás: Chapada dos Veadeiros, 18–24 km. N. of Alto Paraíso, 1400–1500 m, 25.x.1985, leg. S.E. Miller (5, USNM). Minas Gerais: Pedra Azul, 800 m, xi.1972, leg. M. Alvarenga (2, CMNH). São Paulo: Piracicaba, 12.xii.1965, leg. C.A. Triplehorn (10, USNM). Bolivia: Santa Cruz: Same data as holotype except leg. A.R. Cline & J. Wappes, “BOL1Cline07 007” (1, SEMC). Paraguay: Cordillera: Compania Naranjo, 266 m, leg. Brzoska; 2.xii.2012 (9, SEMC).

See differential diagnoses of N. geminus and N. spangleri.

Body length 5.4–6.7 mm. Coloration: Dorsal surfaces very dark brown, with paler margins of pronotum and elytra. Head: Maxillary palps as long as to slightly longer than width of head, uniformly orange in color (Fig. 14D). Thorax: Ground punctation on pronotum and elytra relatively dense and shallowly impressed. Elytra without defined rows of serial punctures, each with one dorsal and a few lateral sparse rows of systematic punctures; rows of systematic punctures more evident along posterior region. Prosternum sometimes medially convex. Posterior elevation of mesoventrite transverse, low, and slightly curved (posteriorly concave). Aedeagus: (Fig. 15I) lateral projection on apical region of outer margin of each paramere rounded to weakly pointed; at closest point, dorsal inner margins of parameres separated by distance similar to greatest width of a paramere; dorsal plate of median lobe with small denticle on each lateral margin, proximal to base of fork; arms of dorsal plate of median lobe distally converging; each arm nearly parallel sided, slightly wider on apical region, apically oblique with inner margin extending beyond outer margin; notch between arms at base slightly narrower than base of an arm; ventral plate of median lobe weakly sclerotized, at widest point nearly as wide as dorsal plate of median lobe, apically acuminate, apex extending beyond base of fork, not reaching apex of arms of dorsal plate; basal piece 0.4× length of a paramere.

Figure 17. 

Dorsal habitus of Novochares spp. A N. guadelupensis B N. pichilingue C N. cochlearis D N. quadrispinus.

Triangularis (L.), in reference to the triangular shape of the ventral plate of the median lobe.

Bolivia, Brazil (Goiás, Minas Gerais, São Paulo), Paraguay (Fig. 16B).

Little is known about the habitat of this species, there is no ecological information on the labels. Some specimens were collected at lights.

Novochares sallaei species group

Species group diagnosis. Body length 4.7–8.0 mm. Coloration: Dorsal surfaces brown to dark brown or reddish brown. Aedeagus: (Figs 18–21) Overall shape and relative length variable, joint basal margins of parameres broadly rounded, truncate or medially pointed; outer margin and apical region of each paramere variable; parameres usually longer than median lobe, with apex rounded or truncate; parameres with apical region variable in degree of sclerotization; dorsal inner margin of each paramere highly variable; dorsal plate of median lobe (in dorsal view) highly variable, sometimes forming a narrow neck; notch between arms variable; shape and orientation of arms variable; position of gonopore variable; ventral plate of median lobe (in ventral view) somewhat triangular, variable in length, shape of apex, and degree of sclerotization, sometimes absent; basal piece 0.20–0.55× length of a paramere, rarely strongly reduced (see cochlearis). In lateral view, aedeagus somewhat triangular, straight to strongly oblique at base, with ventral outline of parameres 1.5–4.2× longer than greatest width.

Figure 18. 

Aedeagi of the Novochares sallaei species group A, B N. bisinuatus C–E N. yanomami F–I N. fernandezae J, K N. unguis L N. clavieri M–P N. cochlearis A, C, F, J, L, M, O, P dorsal view B, H, N lateral view D, G ventral view E, I, K oblique view M, N Brazil O Venezuela P Suriname.

Figure 19. 

Aedeagi of the Novochares sallaei species group A–H N. guadelupensis I–L N. kawsay A–C Guadeloupe D–G Suriname H Guyana I–K Ecuador L Peru A, D, E, H, I, L dorsal view B, F, J ventral view C, G, K lateral view.

Figure 20. 

Aedeagi of the Novochares sallaei species group A–E N. chaquensis F N. garciai G–I N. dicranospathus J–L N. atratus M–R N. sallaei A–C Argentina D Costa Rica E Trinidad M–P Guatemala Q, R Mexico A, D–G, J, M, Q dorsal view B, H, K, N, R ventral view C, I, L, O lateral view P oblique view.

Figure 21. 

Aedeagi of the Novochares sallaei species group A–D N. pastinum E, F N. quadrispinus G–J N. tridentis K, L N. pichilingue A, E, G, K dorsal view B ventral view C, F lateral view D, I (dorsal) J (ventral) L oblique view.

Composition. The Novochares sallaei species group is composed of 17 species: N. atratus (Bruch, 1915), N. bisinuatus sp. nov., N. chaquensis (Fernández, 1982) [= N. carmona (Short, 2005) syn. nov.], N. clavieri sp. nov., N. cochlearis (Fernández, 1982), N. dicranospathus sp. nov., N. fernandezae sp. nov., N. garciai sp. nov., N. guadelupensis (d’Orchymont, 1926), N. kawsay sp. nov., N. pastinum sp. nov., N. pichilingue (Fernández, 1989), N. quadrispinus sp. nov., N. sallaei (Sharp, 1882), N. tridentis sp. nov., N. unguis sp. nov., and N. yanomami sp. nov.

Holotype (male): We examined images of the holotype, including the dissected aedeagus. The specimen is from Buenos Aires Province, Argentina and deposited in MACN.

(9 exs.): Argentina: Entre Ríos: Río Paraná Ibicuy, Pto. Ibicuy, 10.xii.1979, leg. C.M. & O.S. Flint, Jr. (2, USNM). Brazil: Bahia: 5 km W. Ilheus, 4.vii.1969, leg. P. & P. Spangler (3, USNM). Espírito Santo: Muniz Freire, 19.vi.1908 (1, CMNH). Rio de Janeiro: Araruama, xi.1981, leg. Moacir Alvarenga (3, USNM).

The dorsal plate of the median lobe of the aedeagus is very unusual and distinct in that each side of the fork is bilobed and projected dorsally (Fig. 20J, L). The aedeagus is generally most similar to N. chaquensis, which may also be very dark brown to black.

Body length 6.0–7.2 mm. Coloration: Dorsal surfaces brown to dark brown, usually with slightly paler (orange) clypeus, margins of head, pronotum, and elytra. Head: Maxillary palps nearly 1.2× width of head, uniformly orange in color. Thorax: Ground punctation on pronotum and elytra relatively dense and very shallowly impressed. Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum only very weakly medially convex. Posterior elevation of mesoventrite weakly, broadly, and somewhat triangularly elevated. Abdomen: Apical emargination of fifth ventrite shallow to deep, U-shaped. Aedeagus: (Fig. 20J–L) Overall shape pear-like, 4.5× longer than wide, with outer lateral margins of parameres weakly and evenly convex; apical region of each paramere rounded, with outer margin very weakly pointed; at closest point, dorsal inner margins of parameres separated by distance 0.6× greatest width of a paramere; dorsal plate of median lobe with neck 0.65× as broad as base; lateral margins of dorsal plate of median lobe strongly laterally projected at base of fork; arms of dorsal plate of median lobe diverging, dorsally concave, nearly 0.2× length of dorsal plate of median lobe; each arm uniformly wide along basal 1/2, then narrowing to acute apex; notch between arms at base nearly as wide as base of an arm; gonopore placed at base of dorsal plate of median lobe; ventral plate of median lobe weakly sclerotized, triangular, acute at apex, apex extending to base of neck of dorsal plate; basal piece 0.3× length of a paramere. In lateral view, aedeagus oblique at base, with ventral outline of parameres 3.4× longer than greatest width near base; dorsal outline of aedeagus in lateral view nearly straight along second 1/3.

Argentina, Brazil (Bahia, Espírito Santo, Minas Gerais, Rio de Janeiro) (Fig. 22B). Colombia, Ecuador, Paraguay, and the Brazilian state of Mato Grosso do Sul are removed from the distribution, see remarks.

Figure 22. 

Distribution of Novochares sallaei species group A N. cochlearis (red: circles, examined specimens; squares, literature/unconfirmed records), N. dicranospathus (yellow), N. pastinum (blue) B N. atratus (red: circles, examined specimens; squares, literature/unconfirmed records), N. clavieri (yellow), N. bisinuatus (blue), N. garciai (green) C N. chaquensis (red: circles, examined specimens; squares, literature/unconfirmed records), N. pichilingue (yellow) D N. fernandezae (red), N. tridentis (yellow), N. quadrispinus (blue), N. unguis (green).

Little is known about the habitat of this species.

The Colombian specimens identified by Gonzalez-Rodriguez et al. (2017) as N. atratus are in fact N. chaquensis, as can be determined by their figure 24 of the aedeagus, which matches N. chaquensis perfectly. Colombia is thereby removed from the range of this species. Additionally, there is a historical record from Ecuador that was published by Knisch (1925). This Ecuador record was based on two specimens from “Gualaquiza” from “Dr. Festa”. No sex is indicated, though in records of some other species in the same paper, the sex is sometimes indicated, suggesting that the specimens mentioned here may have been unsexed. Because this record is likely a misidentification and so far outside the known range of this species, which is otherwise not known from northern South America or anywhere in the Andean region, we remove Ecuador from the reported range of this species.

The records from Paraguay (Rio Alto Parana) and Mato Grosso do Sul (Corumba) in Brazil derive from specimens cited by d’Orchymont (1939) under H. parhedrus, which is now a synonym of N. atratus. However, all the specimens cited from these localities were females, so there is no way to confirm they are in fact this species and so we remove them from the currently known distribution.

Holotype (male): “BRASIL: Goiás, Sta./ Isabel, R. Araguaia,/ Isla do Bananal/ I,8–11,1961./ B. Malkin leg.” (FMNH). Paratype (1 ex.): Brazil: Goiás: Same data as holotype (1, SEMC).

The aedeagal form of this species is most similar to N. yanomami, but that species has a deeply forked dorsal plate of the median lobe that extends to the apex of the parameres (Fig. 18C); it is not forked and it is much shorter in N. bisinuatus (Fig. 18A).

Body length 5.5–6.3 mm. Coloration: Dorsal surfaces brown to dark brown, with slightly to strongly paler (orange) margins of clypeus, head, pronotum, and elytra. Head: Maxillary palps nearly 1.2× width of head, uniformly orange in color. Thorax: Ground punctation on pronotum and elytra relatively dense and moderately impressed. Each elytron with eight or nine well-defined rows of serial punctures on dorsal surface, with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures; elytral ground punctation moderately impressed. Prosternum flat. Posterior elevation of mesoventrite transverse, curved, and posteriorly concave. Abdomen: Apical emargination of fifth ventrite relatively broad and deep, U-shaped. Aedeagus: (Fig. 18A, B) Overall shape pear-like, nearly 2× longer than wide; lateral projection on apical region of outer margin of each paramere pointed; at closest point, dorsal inner margins of parameres separated by distance 0.2× greatest width of a paramere; dorsal plate of median lobe with neck 0.2× as broad as base; apex of dorsal plate of median lobe medially emarginate, not forming distinct arms; gonopore placed at base of dorsal plate of median lobe; ventral plate of median lobe moderately sclerotized, triangular, apically acute, apex nearly extending to apex of dorsal plate; basal piece 0.37× length of a paramere. In lateral view, aedeagus nearly straight at base, somewhat teardrop-shaped, with ventral outline of parameres 3× longer than greatest width near basal 1/3; dorsal outline nearly straight along basal 1/2.

Referring to the bisinuate apex of the dorsal plate of the median lobe.

Known only from the type locality in central Brazil (Fig. 22B).

Nothing is known about the habitat of this species.

Helochares chaquensis Fernández, 1982: Holotype male from Argentina (Chaco: San Bernardo) deposited in MLP (not seen).

Helochares carmona Short, 2005: Paratypes (2 exs.): Costa Rica: Guanacaste Province: nr. Carmona, laguna de Cocodrilo, 16.i.2003, leg. Short, Roughley, & Porras, HG-vapor light (2, SEMC).

(34 exs.). Argentina: Formosa Province: P.N. Rio Pilcomayo, 50 km NW Clorinda, 19.xii.1990, FM#90-293, marsh edge, S & J Peck, UV light (3, FMNH). Bolivia: Santa Cruz: 60 mi. N. Santa Cruz, Saavedre Exp. Sta., 3–5.i.1960, leg. R. Cumming (4, USNM), same data but 27–20.xii.1959 (1, USNM). Brazil: Mato Grosso: Jacare, Xingu National Park, xi.1965, leg. M. Alvarenga, at light (3, UNSM). Mato Grosso do Sul: Corumba, Paraguay River, -18.95184, -57.66642, 101 m, 25.vi.2018, leg. Hamada & team, macrophytes along river margins and in floating island mats, BR18-0625-01A (4, INPA, SEMC). São Paulo: Piracicaba, 12.xii.1965, leg. C.A. Triplehorn (2, USNM). Ecuador: Napo: Limoncocha, 15.vi.1977, leg. P.J. Spangler & D.R. Givens #129 (1, USNM). Guyana: Mazaruni-Potaro District, Kartabo Point, 1.i.1983, leg. W.E. Steiner (1, USNM). Panama: Darién: Cana ANCON station, 500 m, 7°45.323'N, 77°41.069'W, 3–9.vi.1996, leg. S. Lingafelter, blacklight (1, SEMC). Peru: Madre de Dios: Parque Manu, Pakitza, Cocha Salvador, 12°07'S, 70°59'W, 250 m, 21.ix.1989, leg. R.A. Faitoute, colln. #50 (2, USNM). Trinidad And Tobago: Trinidad, Piarco, 15–16.vii.1969, leg. P. & P. Spangler (4, USNM). Venezuela: Apure: 5 km N. San Juan de Payara, 350’, 25.vii.1988, leg. C. & L. O’Brien & G. Wibmer (2, CAS). Aragua: El Limon, 450 m, 25.v.1977, leg. J. Clavijo, at light (3, MIZA). Guárico: Corozo Pando (8 km. N.), 17–18.vi.1984, leg. F.W. Eiland & V. Linares, blacklight (2, USNM). Zulia: 9°51.833'N, 72°43.285'W, 96 m, btw Machiques & Tukuko, 29.i.2009, leg. Short, García, & Camacho, roadside marsh, VZ09-0129-03Z (1, SEMC).

This widespread species has several less common aedeagal features, particularly the straight, untoothed apices of the parameres (Fig. 20A). The shallow, generally smooth fork at the apex of the dorsal plate of the median lobe is somewhat similar to N. garciai, but that species has strongly sinuate outer margins of the parameres, as well as the dorsal plate being constricted medially (Fig. 20F).

Body length 5.2–7.0 mm. Coloration: Dorsal surfaces brown to dark brown, usually with slightly paler (brown to orange) margins of pronotum, and elytra. Head: Maxillary palps 1.2–1.3× width of head, orange to brown in color, with apex of each palpomere paler. Thorax: Ground punctation on pronotum and elytra relatively dense and shallowly impressed. Elytra without rows of serial punctures, each with very faint rows (one dorsal and two or three lateral) of scarce and weakly marked systematic punctures. Prosternum medially broadly convex. Posterior elevation of mesoventrite broadly and roundly elevated, with medial longitudinal ridge extending anteriorly. Abdomen: Apical emargination of fifth ventrite relatively deep, U- to V-shaped. Aedeagus: (Fig. 20A–E) Overall shape pear-like, 4.5× longer than wide, with outer lateral margins of parameres straight to weakly and evenly convex along basal 3/4; apical region of each paramere rounded, with outer margin smooth, not pointed; at closest point, dorsal inner margins of parameres separated by distance 0.5–1.0× greatest width of a paramere; dorsal plate of median lobe with neck 0.35–0.50× as broad as base; arms of dorsal plate of median lobe diverging, dorsally concave, nearly 0.1× length of dorsal plate of median lobe; each arm uniformly wide along basal 2/3, then narrowing to rounded to acute apex; notch between arms at base slightly wider than base of an arm; gonopore placed at base of dorsal plate of median lobe; ventral plate of median lobe weakly sclerotized, nearly membranous, triangular, irregular at apex, apex extending to second 1/3 of neck of dorsal plate; basal piece 0.3× length of a paramere. In lateral view, aedeagus oblique at base, with ventral outline of parameres 4.2× longer than greatest width near base; dorsal outline of aedeagus in lateral view straight nearly along basal 2/3.

Argentina, Bolivia (new record), Brazil (Mato Grosso, Mato Grosso do Sul, São Paulo), Colombia (new record), Costa Rica (new record), Ecuador (new record), Guyana (new record), Panama (new record), Peru (new record), Trinidad and Tobago (new record), Venezuela (new record) (Fig. 22C). The only previously published locality that we have not examined specimens or images is the type locality in Argentina.

Little is known about this species: most specimens were taken at light traps. One series of specimens was collected in floating macrophytes on the Paraguay River, others were collected in marshes.

This species occurs from Costa Rica south to Argentina. Despite this vast range, it is rather uncommonly collected, with just a smattering of records known to us. Specimens from Colombia identified by Gonzalez-Rodriguez et al. (2017) as N. atratus are in fact this species.

We examined the aedeagus of a paratype of the hitherto Costa Rican endemic N. carmona, which we found to be an exact match to N. chaquensis leading us to synonymize the two species.

Holotype (male): Brazil: Pará: Alenquer/ -1.96253, -54.50458; 44m/ ca. 25 km E of Alenquer;/ Palm swamp, lots of detritus/ 4.ii.2018; Short & Benetti;/ BR18-0204-01A (INPA). Paratypes (11 exs.): Brazil: Amapá: Tartarugalzinho (22 km S) on BR-156, 1.30747, -50.93803, 41 m, 23.vii.2018, leg. Short, marsh/pond, BR18-0723-02A (1, SEMC, DNA voucher SLE2085). Pará: Same data as holotype (8, INPA, SCC, SEMC, including DNA voucher SLE1514). French Guiana: St. Laurent du Maroni, Sentier des Malgaches, 5.48627, -54.00238, 14 m, 4.iii.2020, leg. Short & Neff, pond in secondary forest, FG20-0304-01A (1, SEMC, DNA voucher SLE2420). Peru: Madre de Dios: Tambopata, Kawsay Biological Station, -12.54034 S, -69.00074W, 190 m, 4.vi.2022, leg. Short et al., palm swamp with lots of detritus (1, MHNSM).

This species, with its moderately sinuate outer paramere margins and Y-shaped dorsal plate of the median lobe (Fig. 18L), is at first glance most similar to N. garciai (Fig. 20F), and also to N. fernandezae (Fig. 18F) and N. unguis (Fig. 18J). However, these species all differ in the specific shapes of the median lobe and parameres.