Tobochares sulcatus Short & García, 2007: 4; by original designation.

Small beetles, total body length 1.5–2.6 mm. Color yellowish brown, orange brown to dark brown. Body shape oval in dorsal view; moderately (Fig. 5B) to strongly convex in lateral view (e.g., Fig. 9E). Antennae with eight antennomeres (e.g., Kohlenberg and Short 2017: fig. 8). Maxillary palps curved inward, short (e.g., slightly shorter than the width of the head) and slender (Fig. 2A; e.g., T. goias sp. nov.), to very short (nearly half the width of the head) and stout (Fig. 2G; e.g., T. pemon sp. nov.). Elytra without sutural striae, but in T. akoerio sp. nov. and T. romanoae sp. nov. stria I more strongly impressed along posterior half of elytra (Fig. 8A and D, respectively); elytral punctures seemingly arranged in rows, in some species more evidently so (Fig. 8); in some species interserial punctures not longitudinally aligned (Figs 9, 10); serial punctures sometimes impressed into distinct grooves (e.g., Kohlenberg and Short 2017: fig. 2). Prosternum flat (e.g., Fig. 5C). Posterior elevation of mesoventrite either flat, bulging (Fig. 3E) or with a transverse or longitudinal ridge (Fig. 3D). Metaventrite densely pubescent, except for a median glabrous patch, which is either ovoid and wide (e.g., Kohlenberg and Short 2017: fig. 10) or longitudinal and narrow (Fig. 3D, E). Posterior femora mostly glabrous, with only few scattered setae, sometimes with hydrofuge pubescence along basal half of anterodorsal margin (Kohlenberg and Short 2017: fig. 22). Fifth abdominal ventrite apically evenly rounded, without stout setae (e.g., Kohlenberg and Short 2017: fig. 13). Aedeagus with basal piece usually very short (nearly one third of the length of parameres; Fig. 11).

Figure 2. 

Heads of Tobochares spp. A, B T. goias, black mark pointing to canthus emarginating the eye A dorsal view B anterolateral view C T. fusus D, E T. luteomargo, black mark pointing to straight anterior margin of the eye D dorsal view E anterolateral view F T. romanoae G T. pemon H T. communis I T. microps.

Figure 3. 

Characters of elytra and wings of Tobochares spp. A–E Scanning Electron Micrographs A–C elytra: white arrows point to serial punctures; black arrows point to interserial punctures A T. communis B T. atures C T. arawak D, E ventral view of mesoventrite: D T. communis: top white arrow points to posterior elevation of mesoventrite with low longitudinal elevation; bottom white arrow points to narrow longitudinal glabrous patch of the metaventrite E T. arawak: top white arrow points to posterior elevation of mesoventrite with weakly elevated bulge; bottom white arrow points to narrow longitudinal glabrous patch of the metaventrite. F–I detail of elytral punctation; black marks at top of each circle indicates serial punctures F T. pemon (all punctures relatively large; serial punctures longitudinally aligned; interserial punctures in irregular single row) G T. atures (all punctures longitudinally aligned; serial punctures larger than interserial punctures; interserial punctures more densely distributed) H T. kolokoe (serial punctures longitudinally aligned; interserial punctures forming one or two irregular rows) I T. canaima (serial punctures longitudinally aligned; interserial punctures forming two or three irregular rows) J, K hindwings J T. sipaliwini K T. microps.

This species group can be recognized by the oblique and posteriorly directed anterior margin of the eye in lateral view, which emarginates the eye in lateral view (Fig. 2A–C), the broad, ovoid to diamond-shaped glabrous patch on the metaventrite (Kohlenberg and Short 2017: fig. 10), the dense and uniform distribution of the hydrofuge pubescence on the abdominal ventrites (Kohlenberg and Short 2017: fig. 13A), and the moderate (in number and size) metatibial spines.

This species group includes the following species: Tobochares benettii sp. nov., T. canaliculatus Kohlenberg & Short, 2017, T. goias sp. nov., T. kasikasima Short, 2013, T. kusad Kohlenberg & Short, 2017, T. pallidus Kohlenberg & Short, 2017, T. sipaliwini Short & Kadosoe, 2011, T. striatus Short, 2013, and T. sulcatus Short & García, 2007; see Table 2.

Holotype (male): “Brazil: Amazonas: Rio Preto da Eva; -2.678466, -59.401714, 25 m; ca. 32 Km W of Rio Preto da Eva; seepage on sandstone with algae; 10.vi.2017; leg. Benetti and team; BR17-0610-01A” (INPA). Paratypes (8 exs.): Brazil: Amazonas: Same data as holotype (8, INPA, SEMC including DNA voucher SLE1264).

Tobochares benettii can be easily recognized from all other Tobochares species in the sulcatus species group by its elytral punctures seemingly uniformly distributed, not forming clear longitudinal rows, therefore completely lacking elytral striae (Fig. 4A). This character makes it similar to Tobochares goias, from which it can be distinguished by its uniform orange brown coloration along pronotum and elytra, with dark brown head (Fig. 4A–C), the posterior elevation of the mesoventrite forming a curved transverse ridge, which is medially prominent and acute and by characters of the aedeagus (Fig. 11A).

Figure 4. 

Habitus of Tobochares spp. in the sulcatus species group A–C T. benettii A dorsal view B lateral view C ventral view D–F T. goias D dorsal view E lateral view F ventral view. Scale bars: 1 mm.

Size and form : Body length 1.6–2.0 mm. Body elongate oval, strongly convex (Fig. 4A, B). Color and punctation. Head dark brown, pronotum and elytra uniformly orange brown; antennae, mouthparts and legs orange to yellowish brown, with paler tarsi (Fig. 4C). Ground punctation on head, pronotum and elytra moderately marked. Head: Eyes in dorsal view with anterior margin oblique, posteriorly directed, and outer margins continuous with outline of head (as in Fig. 2A); in lateral view, eyes emarginate to about half the length of eye (as in Fig. 2B). Maxillary palps slender, nearly as long as the width of the head, uniformly yellow in color (as in Fig. 2A). Thorax: Elytra with slightly defined rows of punctures, not forming grooves (Fig. 4A). Elevation of mesoventrite forming a somewhat transverse bulge (Fig. 4C). Metaventrite with distinct median, broad, ovoid glabrous area extending along posterior two thirds (Fig. 4C). Abdomen: Abdominal ventrites uniformly and very densely pubescent. Aedeagus (Fig. 11A) with basal piece 0.25 × the length of a paramere; widest point of parameres (near base) nearly 2/3 greatest width of median lobe (near base), with outer margins slightly sinuate, and rounded apex; median lobe roughly triangular, dorsally concave, apical region nearly half as wide as base, broadly rounded at apex; gonopore situated at apex of median lobe.

Named after Cesar J. Benetti, Brazilian specialist on aquatic beetles, in honor of his contributions to Neotropical beetle taxonomy and for all his assistance in the field.

Only known from the type locality in Amazonas State, Brazil, situated slightly north of the Amazon River (Fig. 13).

This only known series was collected on a vertical seepage on sedimentary rock (Fig. 14A, B).

Holotype (male): “Brazil: Goiás: Cristalina: -16.87004, -47.61716; 947 m; Cristalina Balneario Lajes; seepage on rock next to river; 4.iii.2018; Benetti and team; BR18-0304-02B.” (INPA). Paratypes (26 exs.): Brazil: Goiás: Same data as holotype (26, INPA, SEMC including DNA Voucher SLE1505).

Tobochares goias can be easily recognized from most species in the sulcatus species group by its elytral punctures seemingly uniformly distributed, not forming clear longitudinal rows, therefore completely lacking elytral striae (Fig. 4D). This character makes it similar to T. benettii, from which it can be distinguished by its dark brown coloration, with paler (yellowish to orange) lateral margins of pronotum and elytra (Fig. 4D, E), the posterior elevation of the mesoventrite forming a low and uniform curved transverse ridge (Fig. 4F), and by characters of the aedeagus (Fig. 11B).

Dorsal surfaces of body dark brown with paler outer margins of pronotum and elytra (Fig. 4D, E); head slightly darker; mouthparts and antennae yellowish; legs yellowish to brown with paler tarsi (Fig. 4F). Ground punctation on head, pronotum and elytra moderately marked. Head: Eyes in dorsal view with anterior margin oblique, posteriorly directed, and outer margins continuous with outline of head; in lateral view, eyes emarginate to about half the length of eye (Fig. 2A, B). Maxillary palps slender, slightly shorter than width of head, uniformly yellowish in color (Fig. 2A). Thorax: Elytra with slightly defined rows of punctures, not forming grooves (Fig. 4D). Elevation of mesoventrite with a low transverse ridge (Fig. 4F). Metaventrite with distinct median, broad, ovoid glabrous area extending along posterior two thirds (Fig. 4F). Abdomen: Abdominal ventrites uniformly and very densely pubescent. Aedeagus (Fig. 11B) with basal piece 0.2 × the length of a paramere; widest point of parameres (near base) nearly as wide as basal width of median lobe, with outer margins very slightly sinuate, and rounded apex; median lobe fusiform, with widest point slightly apicad of midpoint, broadly rounded at apex; gonopore situated at apex of median lobe.

Named after the Brazilian state of Goiás, from which the species is known.

Only known from the type locality in the central Brazilian state of Goiás. This is the first and currently only species of Tobochares reported from south of the Amazon River (Fig. 12).

This species was collected on wet rock along the margins of the Ribeirão das Lejas. See Fig. 14C.

Brazil: Roraima: Caroebe Municipality, Reservoir by Usina de Jatapú, 0.872953°, -59.282170°, 185 m, 17.i.2018, large wall seep with algae, leg. Short, Benetti, and Santana, BR18-0117-01A (7, INPA, SEMC including DNA Voucher SLE1501).

Brazil: Roraima: Amajari Municipality, Serra do Tepequém, Igarape Preto Negro, Cachoeira Laje Preta, 3°36.381'N, 61°42.878'W, 618 m, 14.i.2018, leg. Short and Benetti, BR18-0114-04B (12, INPA, SEMC, including DNA voucher SLE1496); Caroebe Municipality, Reservoir by Usina de Jatapú, 0.872953, -59.282170, 185 m, 17.i.2018, large wall seep with algae, leg. Short, Benetti, and Santana, BR18-0117-01A (1, SEMC including DNA Voucher SLE1502).

Suriname: Sipaliwini: Sipaliwini Savanna Nature Reserve, 2°00.240'N, 55°58.259'W, 374 m, 4-Brothers Mountains, leg. Short and Baca, 30.iii.2017, seeps on granite SR17-0330-04A (1, SEMC including DNA Voucher SLE1244).

Members of this species group can be recognized by the straight anterior margin of the eye in lateral view (Fig. 2E), the longitudinally aligned and undifferentiated elytral punctures with about the same size and degree of impression (Fig. 5A), the short and narrow glabrous patch on the metaventrite (Fig. 5C), the scantiness of the hydrofuge pubescence on the abdominal ventrites (Fig. 5C), and the reduced (in number and size) metatibial spines.

Figure 5. 

Habitus of Tobochares spp. in the luteomargo and emarginatus species group A–C T. luteomargo A dorsal view B lateral view C ventral view. D–F T. fusus D dorsal view E lateral view F ventral view. Scale bars: 1 mm.

This species group currently contains only a single species, T. luteomargo sp. nov.

Holotype (male): “Venezuela: Bolivar State/ 7°41'23.6"N, 64°1'56.0"W, 134 m/ ca. 14 km E Rio Aro; 5.viii.2018/ leg. A. Short and M. García/ AS-08-073; rock outcropping” (MIZA). Paratypes (282 exs.): Venezuela: Bolívar: “6°35.671'N, 66°49.238'W; 80 m; Los Pijiguaos; morichal/ rock outcrop; 16.ix.2007; leg. Short, García, Joly; AS-07-015” (17, SEMC including DNA vouchers SLE1028); same, except “seeps and stream at night; 9.vii.2010; leg. Short and Téllez; VZ10-0709-03A” (1, SEMC); Same data as holotype (80, MALUZ, SEMC); “7°29'47.3"N, 65°51'44.8"W; 45 m; 2 Km E of Río Cuchivero; rock outcrop seeps; 6.viii.2008; leg. Short, Téllez, Arias; AS-08-075” (3, SEMC); “6°57.904'N, 66°36.392'W; 51 m; outcrop ca. 15 km NE of Los Pijiguaos; detritus flotation; 9.vii.2010; leg. Short and Téllez; VZ10-0709-01B” (2, SEMC including DNA voucher SLE1025); “7°27.501'N, 65°52.093'W; 45 m; Rock outcrop by Río Cuchivero; seeps; 10.vii.2010; leg. Short, Téllez, Arias; VZ10-0710-01A” (75, SEMC including DNA voucher SLE102); “7°37.443'N, 64°08.324'W; 90 m; Campamento Río Aro, by Río Aro; rock pools; 11.vii.2010; leg. Short, Téllez, Arias; VZ10-0711-01A” (2, SEMC); same, except “seep scrubbing; VZ10-0711-01B” (48, SEMC); same, except “seep/rock pools at night; VZ10-0711-01E” (44, SEMC including DNA voucher SLE451).

Tobochares luteomargo can be distinguished by the yellow band along the outer margins of the body (Fig. 5B).

Size and form : Body length 1.7–2.1 mm. Body elongate oval, moderately convex (Fig. 5B). Color and punctation: Dorsal surfaces of body dark brown, with clypeus (Fig. 2A), outer margins of pronotum, and outer margins of elytra yellowish brown (Fig. 5A, B); maxillary and labial palps yellow, remainder mouthparts orange; antennae yellowish brown; legs orange brown with paler (yellow) tarsi; ventral surfaces brown (Fig. 5C). Ground punctation on head, pronotum and elytra moderately marked (Figs 2A, 5A). Head: Eyes in dorsal view with anterior margin oblique and anteriorly directed (Fig. 2A, B), and outer margins slightly bulging from outline of head; in lateral view, eyes not anteriorly emarginate (Fig. 2B). Maxillary palps as long as 0.8 × width of head. Thorax: Elytra with all kinds of punctures similar in size and degree of impression, moderately aligned in rows, not forming grooves (Fig. 5A). Elevation of mesoventrite forming a very low transverse carina (Fig. 5C). Metaventrite with distinct median, longitudinal, glabrous area, 3 × longer than wide, extending along posterior half (Fig. 5C). Abdomen: Abdominal ventrites uniformly and very densely pubescent (Fig. 5C). Aedeagus (Fig. 11D). Basal piece 2 × longer than a paramere; greatest width of a paramere nearly 0.6 × greatest width of median lobe; outer margins of parameres slightly sinuate, inner margins strongly sinuate; apex of paramere sharply acute, pointing towards longitudinal midline of aedeagus; median lobe roughly sagittate, medially with narrow emargination extending along apical fourth; gonopore situated nearly at apical third of median lobe.

Named with the Greek words luteo, meaning yellow, and margo meaning margin, in reference to the striking yellow band surrounding the marginal areas of these beetles.

This species is known from several localities along the northwestern edge of the Guiana Shield in Bolívar State, Venezuela. See Fig. 13.

Tobochares luteomargo is found on rock seeps on granitic inselberg-like habitat. Some of these seeps may be very small, less than half a square meter in size. The largest series of this species were collected in seeps on which there was abundant apparent lichen growth, and specimens were often hiding under these growths (Fig. 15E).

The emarginatus species group can be recognized by the oblique and posteriorly directed anterior margin of the eye in lateral view, which emarginates the eye to about a quarter of its dorsal width (Fig. 2C; Kohlenberg and Short 2017: fig. 6C–F), the presence of a broad, ovoid to diamond-shaped glabrous patch on the metaventrite (Fig. 5F), the dense and uniform distribution of the hydrofuge pubescence on the abdominal ventrites (Fig. 5F), and the moderate (in number and size) metatibial spines.

Figure 6. 

Habitus of Tobochares spp. in the communis species group A–C T. communis A dorsal view B lateral view C ventral view D–F T. microps D dorsal view E lateral view F ventral view. Scale bars: 1 mm.

This species group presently contains three species: T. canthus, T. emarginatus, and T. fusus sp. nov.

Figure 7. 

Habitus of Tobochares spp. in the communis species group A–C T. anthonyae A dorsal view B lateral view C ventral view D–F T. atures D dorsal view E lateral view F ventral view G–I T. pemon G dorsal view H lateral view I ventral view. Scale bars: 1 mm.

Holotype (male): “Brazil: Amapá: Oiapoque/ 3.85039, -51.81683; 17 m/ Oiapoque (ca. 1 km E); 18.vii.2018/leg. Short; Flotation of detritus/ex forest seep; BR18-0718-03C” (INPA). Paratypes (58 exs.): Brazil: Amapá: Same data as holotype (45, INPA, MNHN, SCC, SEMC including DNA voucher SLE1564); Oiapoque (4 km NE), 3.87234, -51.80315, 14 m, 18.vii.2018, leg. Short, root mats on rock at margin seep, BR18-0718-02A (10, INPA, SEMC); Oiapoque (ca. 5.5 km NE), Balneario, 18.vii.2018, leg. Short, margin of larger stream, BR18-0718-01C (1, SEMC). French Guiana: Savane Roche Virginie, near RN 2, 4.1883, -52.13982, 64 m, 10.iii.2020, leg. Short and Neff, rotting Clusia fruits, FG20-0310-01D (3, SEMC including DNA Voucher SLE2171).

Tobochares fusus is externally indistinguishable from T. emarginatus, given that both species share the same disposition and degree of impression of the elytral punctures, and a low transverse ridge on the posterior elevation of the metaventrite. However, they can be clearly differentiated by the general shape of the aedeagus, which is rather slender and nearly parallel sided in T. emarginatus (Kohlenberg and Short 2017: fig. 14I), whereas T. fusus has a fusiform aedeagus (see Fig. 11C).

Size and form : Body length 1.7–2.0 mm. Body elongate oval, moderately convex (Fig. 5D, E). Color and punctation: Dorsal surfaces of body dark brown, with slightly paler pronotum (Fig. 5D); antennae, mouthparts, and legs yellowish brown, with orange tibiae; ventral surfaces of meso-, metathorax and abdomen dark brown (Fig. 5F). Ground punctation on head, pronotum and elytra moderately marked. Head: Eyes in dorsal view with anterior margin oblique, posteriorly directed; canthus emarginating eye to about a quarter of its dorsal width in lateral view (Fig. 2C). Maxillary palps 0.7 × width of head. Thorax: Elytra with all kinds of punctures similar in size and degree of impression, seemingly evenly distributed, not forming grooves (Fig. 5D). Elevation of mesoventrite forming a low transverse ridge (Fig. 5F). Metaventrite with distinct median, broad, diamond-shaped glabrous area extending along posterior two thirds (Fig. 5F). Abdomen: Abdominal ventrites uniformly and densely pubescent (Fig. 5F). Aedeagus (Fig. 11C). Basal piece nearly 0.4 × length of a paramere; greatest width of a paramere nearly 0.7 × greatest width of median lobe; outer margins of parameres diverging along basal half, then broadly bending inward and tapering along apical fifth; apex of paramere oblique, pointing towards longitudinal midline of aedeagus; median lobe roughly triangular, apically rounded; gonopore situated at apex of median lobe.

Named with the Latin word fusus, meaning fusiform, in reference to the shape of the aedeagus of this species.

This species is known from two closely situated localities on either side of the Oiapoque River, the boundary between French Guiana and the Brazilian state of Amapá. See Fig. 13.

The series from Brazil were taken from a seepage habitat in a forested riparian corridor (Fig. 14F). A thin layer of saturated dead leaves was laying over granite, over which a thin film of water was seeping into an adjacent stream. The short series from French Guiana was collected from the rotting fruits of a Clusia on an otherwise dry forest floor and not adjacent to any aquatic habitat (Fig. 17C).

The Tobochares communis species group can be recognized by the straight anterior margin of the eye in lateral view (e.g., Fig. 2E), the usually longitudinally aligned elytral punctures, the short and narrow glabrous patch on the metaventrite (Fig. 3D, E), the high density of the hydrofuge pubescence on the abdominal ventrites, and the well-developed and numerous metatibial spines.

Tobochares akoerio sp. nov., T. arawak sp. nov., T. anthonyae sp. nov., T. atures sp. nov., T. canaima sp. nov., T. communis sp. nov., T. kappel sp. nov., T. kolokoe sp. nov., T. microps sp. nov., T. pemon sp. nov., and T. romanoae sp. nov.

Holotype (male): “Suriname: Sipaliwini District, 2.46554°N, 55.7700°; 800 m; Camp 2, Grensgebergte Rock; rock seepages; 12.iii.2012; leg. A. Short; SR12-0312-01A” (NZCS). Paratypes (5 exs.): Same data as holotype (5, SEMC).

Tobochares akoerio can be recognized by its strongly convex body in lateral view (Fig. 8E), accompanied by elytra with well-defined rows of serial punctures, moderately impressed, forming grooves along apical 3/4 of elytra (Fig. 8D, E); interserial punctures somewhat irregularly distributed (Fig. 8D, E). The general habitus of T. akoerio is similar to that of T. romanoae and T. canaima, especially by the uniformly dark coloration of the pronotum; T. akoerio can be distinguished from these two species by its strongly impressed striae, especially along the lateral regions of the elytra when compared to T. romanoae (compare Fig. 8D, E to 8A, B).

Figure 8. 

Habitus of Tobochares spp. in the communis species group A–C T. romanoae A dorsal view B lateral view C ventral view D–F T. akoerio D dorsal view E lateral view F ventral view. Scale bars: 1 mm.

Figure 9. 

Habitus of Tobochares spp. in the communis species group A–C T. kappel A dorsal view B lateral view C ventral view D–F T. kolokoe D dorsal view E lateral view F ventral view. Scale bars: 1 mm.

Size and form : Body length 2.0 mm. Body elongate oval, strongly convex (Fig. 8E). Color and punctation: Dorsal and ventral surfaces of body dark brown, with lateral margins of prothorax only very slightly paler (Fig. 8D, E); mouthparts and antennae yellow, with slightly darker antennal club and apical third of maxillary palpomere IV; legs orange with paler tarsi (Fig. 8F). Ground punctation on head, pronotum and elytra moderately marked (Fig. 8D). Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed), and outer margins slightly bulging from outline of head; in lateral view, eyes not emarginate. Thorax: Elytra with well-defined rows of serial punctures, moderately impressed, forming grooves along apical 3/4 of elytra (Fig. 8D); interserial punctures somewhat irregularly distributed (Fig. 8D). Metafemora mostly glabrous on anterior face, with narrow band of pubescence along basal third of dorsal margin (Fig. 8D). Elevation of mesoventrite forming a low transverse carina (Fig. 8D). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 8D). Abdomen: Abdominal ventrites uniformly and very densely pubescent. Aedeagus (Fig. 11K). Basal piece 0.4 × the length of a paramere; parameres nearly 1/3 as narrow as greatest width of median lobe, with outer margins widely and uniformly convex, and rounded apex; median lobe roughly triangular, rounded and slightly pinched at apex; gonopore situated nearly at midlength of median lobe.

Noun in apposition. Named after the Akoerio, an indigenous nomadic tribe, with only few people remaining in the South of Suriname.

The species is only known from an exposed rocky summit in the Grensgebergte Mountains along the border between Suriname and Brazil. See Fig. 13.

This species was collected on flowing seeps with moss and algae over granite. See Fig. 16F.

Holotype (male): “Guyana: Region VIII: 5°0.730'N, 59°38.965'W; 585 m; Upper Potaro Camp I; ca. 7 Km NW of Chenapau; top of falls on Potaro River; seeps with roots and algae; 12.iii.2014; leg. Short, Salisbury, La Cruz; GY14-0312-01B” (CBDG). Paratypes (127 exs.): Guyana: Region VIII: Same data as holotype (127, CBDG, SEMC).

Tobochares arawak can be recognized by its strongly convex body in lateral view (Fig. 10B), accompanied by elytral punctation uniform in size and degree of impression, with serial punctures seemingly aligned in rows, not impressed to form grooves (Fig. 10A); the interserial punctures are somewhat irregularly distributed in two or three rows (Fig. 10A). The general habitus and punctation of T. arawak are similar to those of T. canaima, T. kappel, and T. kolokoe. In T. kappel and T. kolokoe the interserial punctures form only one or two irregular rows (Fig. 3H). In T. canaima (Fig. 10D) the pronotal and elytral punctations are sharper than in T. arawak and the apodemes of the median lobe are one fourth the length of the median lobe in T. canaima (Fig. 11N), as opposed to half as long in T. arawak (Fig. 11M).

Figure 10. 

Habitus of Tobochares spp. in the communis species group A–C T. arawak A dorsal view B lateral view C ventral view D–F T. canaima D dorsal view E lateral view F ventral view. Scale bars: 1 mm.

Figure 11. 

Aedeagi of Tobochares spp. A T. benettii B T. goias C T. fusus D T. luteomargo E T. pemon F T. anthonyae G T. atures H T. communis I T. microps J T. romanoae K T. akoerio L T. kappel M T. arawak N T. canaima. Scale bars: 0.5 mm (A–D); 0.1 mm (E–N).

Size and form : Body length 1.6–1.8 mm. Body elongate oval, strongly convex (Fig. 10A, B). Color and punctation: Dorsal and ventral surfaces of body dark brown, with lateral margins of prothorax and elytra only slightly paler (Fig. 10A, B); mouthparts yellow, with slightly darker apical third of maxillary palpomere IV; antennae brown; legs reddish to dark brown with paler tarsi (Fig. 10C). Ground punctation on head, pronotum and elytra moderately marked (Fig. 10A, B). Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed), and outer margins slightly bulging from outline of head; in lateral view, eyes not emarginate (see Fig. 2E). Thorax: Elytra with slightly defined rows of shallow serial punctures, not forming grooves (Fig. 10A); interserial punctures somewhat irregularly distributed in two or three rows (Fig. 3I). Elevation of mesoventrite forming a very low transverse carina (Fig. 10C). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 10C). Abdomen: Abdominal ventrites uniformly and densely pubescent. Aedeagus (Fig. 11M). Basal piece 0.4 × the length of a paramere; parameres nearly 1/3 as narrow as greatest width of median lobe, with outer margins widely and uniformly convex, and rounded apex; median lobe roughly triangular, rounded and slightly pinched at apex; gonopore situated nearly at midlength of median lobe.

Noun in apposition. Named after the Arawak, an indigenous tribe of northern South America.

Tobochares arawak is only known from the Upper Potaro region in Guyana. See Fig. 13.

This species was collected in a wet seepage area along rocks at the margin of the Upper Potaro River. Specimens were collected by pulling back root mats and moss that were growing over the wet rock areas. See Fig. 15A, B.

Figure 12. 

Distribution of Tobochares spp., including all previous (red) and new (yellow) records.

Figure 13. 

Distribution of Tobochares spp.

Figure 14. 

Tobochares habitat in Brazil A, B type locality and habitat for T. benettii, seepage near Rio Preto de Eva (collecting event BR17-0610-01A) C type locality and habitat for T. goias, margin of Balneario Lejas (collecting event BR18-0304-02B) D habitat of T. kusad and T. sipaliwini, State of Roraima, near Usina de Jatapú reservoir (collecting Event BR18-0117-01A) E habitat and type locality of T. romanoae, and habitat of T. sipaliwini, State of Roraima, Serra do Tepequém, Igarape Preto Negro, Cachoeira Leje Preta (collecting event BR18-0114-04B) F type locality and habitat of T. fusus, State of Amapá, Calcoene (collecting event BR18-0721-02B).

Holotype (male): “Venezuela: Bolívar: 6°13'4.6"N, 67°14'26.4"W; 60 m; ca. 25 Km E of El Burro; rocky morichal; 12.i.2009; leg. Short et al.; VZ09-0113-01X” (MIZA). Paratypes (3 exs.): Venezuela: Bolívar: “6°13'4.6"N, 67°14'26.4"W; 60 m; ca. 25 Km E of El Burro; rocky morichal; 7.viii.2008; leg. Short, García, Joly; AS-08-077” (1, SEMC); same data as holotype (2, SEMC).

The general habitus and coloration of T. anthonyae is similar to that of several species in the communis group; nevertheless, the elytral punctation T. anthonyae is relatively distinct: all kinds of punctures are relatively large, similar in size and degree of impression, the serial punctures are aligned in rows and slightly impressed, forming shallow longitudinal grooves, and the interserial punctures are somewhat irregularly distributed in a single row (Fig. 7A). The relatively large punctures, similar in size and degree of impression may resemble those of T. communis, but in this species the serial punctures are not impressed to form grooves (Fig. 6A), as they are in T. anthonyae. In addition, the overall shape of the aedeagus, especially the shape of the median lobe of T. anthonyae is unique among members of the communis group: the median lobe gradually and slightly narrows towards a broadly rounded apex, and the gonopore is located near the apex of the median lobe (Fig. 11F).

Size and form : Body length 1.8–2.0 mm. Body elongate oval, moderately convex (Fig. 7B). Color and punctation: Dorsal and ventral surfaces of body, dark brown, with slightly paler margins of pronotum (Fig. 7A, B); mouthparts yellowish brown; antennae light brown; legs orange with yellow tarsi (Fig. 7C). Ground punctation on head, pronotum and elytra rather shallowly marked. Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed; e.g., Fig. 2D), and outer margins slightly bulging from outline of head; in lateral view, eyes not anteriorly emarginate (e.g., Fig. 2E). Thorax: Elytra with all kinds of punctures similar in size and degree of impression (Fig. 7A); serial punctures aligned in rows, slightly impressed, forming shallow longitudinal grooves; interserial punctures somewhat irregularly distributed in a single row (Fig. 7A). Metafemora mostly glabrous on anterior face, with hydrofuge pubescence along basal third of antero-dorsal margin (Fig. 7C). Elevation of mesoventrite forming a low longitudinal bulge (Fig. 7C). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 7C). Abdomen: Abdominal ventrites uniformly and very densely pubescent. Aedeagus (Fig. 11F). Basal piece 0.3 × the length of a paramere; greatest width of a paramere nearly 0.5 × greatest width of median lobe; outer margins of parameres nearly straight, only slightly curved inwards along apical region; apex of paramere rounded; median lobe roughly triangular, widely rounded at apex; gonopore situated at apical fourth of median lobe.

Named after Becky Anthony, program and meetings manager at the Entomological Society of America (ESA), in recognition of all her hard work in service to the society and the entomological community.

Only known from a single locality just south of the Orinoco River along the northwestern edge of the Guiana Shield. See Fig. 13.

The specimens were collected along a stream that was flowing over exposed granite.

Holotype (male): Venezuela, “T.F. Amazonas/ Puerto Ayacucho (40km S)/ El Tobogán, Caño Coromoto/ 26 Jan 1989, stream edge/ at upper shelter”, “collected by/ PJSpangler/RAFaitoute & CBBarr” (MIZA). Paratypes (357 exs.): Venezuela: Amazonas: “40 Km S of Puerto Ayacucho, at Tobogán; upper seep; 18.i.1989; leg. Spangler, Faitoute, Barr” (34, USNM); same, except “; colln #1; collected by pouring water over stream bank and washing riparian insects into seine; 19.vi.1989; leg. Spangler and Faitoute” (3, USNM); same, except “sandy margins; colln. #10; 23.ii.1986; leg. P. Spangler” (38, USNM); same, except “colln. #14; 25.ii.1986” (16, USNM); “40 Km S of Puerto Ayacucho, El Tobogán, Caño Coromoto; seep, at upper shelter; 26.i.1989; leg. Spangler, Faitoute, Barr” (55, USNM); same data as holotype (174, SEMC, MIZA, USNM); “5°62'N [Sic!], 66°23'W; 1250 m; Cerro Guanay; Exp. Terramar; 5–12.ii.1995; leg. J. Clavijo” (1, MIZA); Tobogán de la Selva; old “Tobogancito” on seepage area with detritus; 8.viii.2008; leg. Short, García, Joly; AS-08-080b (16, SEMC including DNA voucher SLE1032); same, except “Tobogán de la Selva; wet rock covered with detritus; upstream slide; 14.i.2009; leg. Short, García, Miller, Joly; VZ09-0114-01F” (16, SEMC); same, except “partly shaded wet rock with algae; leg. Short and Miller; VZ09-0114-01G” (2, SEMC). Bolívar: “6°13'4.6"N, 67°14'26.4"W; 60 m; ca. 25 Km E of El Burro; rocky morichal; 12.i.2009; leg. Short and Téllez; VZ09-0113-01X” (1, SEMC).

Figure 15. 

Tobochares habitat in Guyana and Venezuela A, B habitat for T. arawak, Guyana, upper Potaro River (collecting event GY14-0312-01B) C, D habitat for T. luteomargo, Venezuela, rock outcrop by Río Cuchivero, (collecting event VZ10-0710-01A) E habitat for T. luteomargo, Venezuela Campamento Río Aro (collecting event VZ10-0711-01B) F type locality and habitat for T. atures, Venezuela, Tobogan de la Selva (collecting event VZ09-0114-01F).

The general habitus and coloration of T. atures is similar to that of several species in the communis group, nevertheless, the elytral punctation in T. atures is relatively distinct: all the elytral punctures are shallowly impressed, longitudinally aligned, and have two different sizes: the serial punctures, which are slightly impressed, are larger, whereas the interserial punctures are smaller and denser (Fig. 3B, G). In addition, the overall shape of the aedeagus, especially the shape of the median lobe of T. atures is unique among members of the communis group: the median lobe gradually and slightly narrows towards a rounded apex, and the gonopore is located at the apical third of the median lobe (Fig. 11G).

Size and form : Body length 2.0–2.2 mm. Body elongate oval, moderately convex (Fig. 7D). Color and punctation: Dorsal and ventral surfaces of body dark brown, with prothorax and lateral margins of elytra slightly paler (Fig. 7D, E); mouthparts and antennae yellow, with slightly darker apical third of maxillary palpomere IV; legs orange to reddish brown with paler tarsi (Fig. 7E). Ground punctation on head, pronotum and elytra rather shallowly marked. Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed), and outer margins slightly bulging from outline of head (e.g., Fig. 2D); in lateral view, eyes not emarginate (Fig. 2E). Thorax: Elytra with longitudinal rows of shallow punctures, not forming grooves; punctures in two different sizes: serial punctures larger, interserial punctures smaller and denser (Fig. 3B, G). Metafemora mostly glabrous on anterior face (Fig. 7F). Elevation of mesoventrite forming a low transverse carina (Fig. 7F). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 7F). Abdomen: Abdominal ventrites uniformly and very densely pubescent (Fig. 7F). Aedeagus (Fig. 11G). Basal piece nearly 0.5 × the length of a paramere; parameres nearly 0.4 × greatest width of median lobe, with outer margins weakly and uniformly convex, and rounded apex; median lobe roughly triangular, widely rounded at apex; gonopore situated nearly at apical third of median lobe.

Noun in apposition. Named after Atures, the municipality where the type locality is situated.

This species is known from several localities along the northwestern edge of the Guiana Shield in Venezuela. See Fig. 13.

Most specimens were collected on granite seepages that were adjacent to permanent streams. See Fig. 15F.

Holotype (male): “Venezuela: Bolívar: 5°51'N, 62°33'W; 1700 m; Auyan-tepui; Intercept trap; 7–14.ii.1994; leg. J.L. García, A. Chacón” (MIZA). Paratypes (7 exs.): Venezuela: Bolívar: Same data as holotype (6, MIZA, SEMC); “5°46'50"N, 62°31'36"W; 2170 m; Auyan-tepui; yellow trap; 20.iv.1994; leg. L. Mesner, J.L. García” (1, MIZA).

Tobochares canaima can be recognized by its strongly convex body in lateral view, accompanied by elytral punctation uniform in size and degree of impression, with serial punctures seemingly aligned in rows, not impressed to form grooves; the interserial punctures are somewhat irregularly distributed in two or three rows (Fig. 10D, E). The general habitus and punctation of T. canaima are similar to those of T. arawak, T. kappel, and T. kolokoe. In T. kappel and T. kolokoe the interserial punctures form only one or two irregular rows (e.g., Fig. 3H). In T. arawak the pronotal and elytral punctation is shallower than in T. canaima (compare Fig. 10A vs. Fig. 10B) and the apodemes of the median lobe are half the length of the median lobe in T. arawak (Fig. 11M), as opposed to one fourth of the length in T. canaima (Fig. 11N).

Size and form : Body length 1.9–2.1 mm. Body elongate oval, moderately convex (Fig. 10E). Color and punctation: Dorsal and ventral surfaces of body dark brown, with anterolateral margins of prothorax slightly paler (Fig. 10D, E); mouthparts yellow to orange; antennae light brown; legs reddish to dark brown, distally paler (orange), with paler (yellow) tarsi (Fig. 10F). Ground punctation on head, pronotum and elytra sharply marked; pronotal punctation dense (Fig. 10D). Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed; e.g., Fig. 2D), and outer margins slightly bulging from outline of head; in lateral view, eyes not emarginate (e.g., Fig. 2E). Thorax: Elytra with serial punctures similar in size and degree of impression to interserial punctures, and only seemingly aligned longitudinally, not forming grooves; interserial punctures irregularly distributed in two or three rows (Fig. 10D). Metafemora mostly glabrous on anterior face (Fig. 10F). Elevation of mesoventrite forming a very low transverse carina (Fig. 10F). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 10F). Abdomen. Abdominal ventrites uniformly and very densely pubescent (Fig. 10F). Aedeagus (Fig. 11N). Basal piece nearly 0.45 × the length of a paramere; greatest width of a paramere nearly 0.6 × greatest width of median lobe; outer margins of parameres straight to widely and uniformly convex; apex of paramere obliquely rounded; median lobe roughly sagittate, rounded at apex; gonopore situated nearly at midlength of median lobe.

Noun in apposition. Named after the Canaima National Park in Venezuela, where the type locality is situated.

This species is known from the famous Auyan-tepui, which is also home to Angel Falls, the highest waterfall in the world. Collected at elevations of 1700–2170 m, this species is one of the relatively few water beetle taxa known from the “Pantepui Province”, which comprises areas of the Guiana Shield which are greater than 1500 m in elevation. See Fig. 13.

The only known series was collected in a flight intercept trap and a yellow pan trap. Nothing further is known about the habitat or biology of this species.

Holotype (male): “Suriname: Sipaliwini District: 4°40.432'N, 56°11.079'W; 86 m; Raleighvallen Nature Reserve, base of Voltzberg; flotation of roots and debris from seepage; 17.iii.2016; leg. Short and Girón; SR16-0317-01C” (NZCS). Paratypes (187 exs.): Brazil: Amapá: 2.60342, -51.33237; 73 m, Calcoene (ca. 42 km NW) on BR-156, Large rock outcrop; 21.vii.2018, leg. Short; Floating rootmats/dirt from seepage, BR18-0721-02B (1, SEMC including DNA voucher SLE1566); 3.87281, -51.78939; 10 m, Oiaqpoque (ca. 5.5 km NE), balneario, 18.vii.2018; leg. Short; seepage area, BR18-0718-01B (1, SEMC, DNA voucher SLE1568). Roraima: Amajari Municipality, Serra do Tepequém, Igarape Preto Negro, Cachoeira Leje Preta, 3 36.381'N, 61 42.878'W, 618 m, 14.i.2018, leg. Short and Benetti, BR18-0114-04B (1, SEMC, DNA voucher SLE1494); Caroebe Municipality, Reservoir by Usina de Jatapú, 0.872953, -59.282170, 185 m, 17.i.2018, large wall seep with algae, leg. Short, Benetti, and Santana, BR18-0117-01A (1, INPA, DNA voucher SLE1503). Guyana: Region VIII: “5°10.514'N, 59°28.970'W; 440 m; Kaieteur National Park; rock savanna, large seepage area; 16.iii.2014; leg. A. Short; GY14-0316-01C” (51, CBDG, SEMC); same, except “flotation of wet leaves and roots; GY14-0316-01B” (SEMC, 6); same, except “seepage on granite with some vegetation; 21.iii.2014; leg. Short, Salisbury, La Cruz; GY14-0321-02A” (9, SEMC including DNA voucher SLE1022). Suriname: Sipaliwini District: “2.46554°N, 55.7700°W; 800 m; Camp 2, Grensgebergte Rock; rock seepages; 12.iii.2012; leg. A. Short; SR12-0312-01A” (23, NZCS, SEMC including DNA Voucher SLE481); “4°40.966'N, 56°11.262'W; 96 m; Raleighvallen Nature Reserve, plateau below Voltzberg; seepage; 28.vii.2012; leg. Short, Maier, McIntosh; SR12-0728-01B” (12, SEMC); same, except “SR12-0728-01C” (6, SEMC); same, except “small seep at margin of plateau; 15.iii.2016; leg. A. Short; SR16-0315-03B” (2, SEMC); “3°47.479'N, 56°8.968'W; 320 m; CSNR: near Kappel airstrip; temporary rivulets in tire tracks; 13.viii.2013; leg. Short, Bloom, Kadosoe; SR13-0813-02A” (1, SEMC); same, except “pond in forest on trail to Tafelberg; 13.viii.2013; leg. Short and Bloom; SR13-0813-05A+B” (1, SEMC); CSNR: Tafelberg Summit, nr. Caiman Creek Camp, leg. Short & Bloom, 19.viii.2013, large seepage area, SR13-0819-01A (1, SEMC, DNA voucher 1047); “3°47.479'N, 56°8.968'W; 320 m; CSNR: near Kappel airstrip; seepage flowing into canal/ ditch on S side of airstrip; 24.viii.2013; leg. Short and Bloom; SR13-0824-02A” (1, SEMC); same, except “SR13-0824-02B” (19, SEMC); “4°40.432'N, 56°11.079'W; 86 m; Raleighvallen Nature Reserve, base of Voltzberg; seepage spot on side of rock; 16.iii.2016; leg. A. Short; SR16-0316-01A” (1, SEMC); same data as holotype (15, SEMC); “2°00.240'N, 55°58.259'W; 374 m; Sipaliwini Savanna Nature Reserve; 4-Brothers mountains; seeps on granite; with algae; 30.iii.2017; leg. Short and Baca; SR17-0330-04A” (6, SEMC); same, except “seeps on granite; by flotation; SR17-0330-04B” (11, SEMC). Venezuela: Bolívar: “6°04'54.7"N, 61°23'52.7"W; 509 m; along La Escalera, Highway 10; rock seepages; 14.vii.2010; leg. Short, Téllez, Arias; VZ10-0714-01B” (3, SEMC including DNA Voucher SLE1026); “6°35.617'N, 66°49.238'W; 80 m; Los Pijiguaos; morichal/rock outcrop; 16.ix.2007; leg. A. Short, M. García, L. Joly; AS-07-015” (7, MIZA, SEMC); same, except “seeps and stream at night; 9.vii.2010; leg. Short and Téllez; VZ10-0709-03A” (2, SEMC including DNA voucher SLE1036); “5°40'24.8"N, 61°24'11.3"W; 1330 m; unnamed river; small side stream; 2.viii.2008; leg. Short, García, Joly; AS-08-066” (3, SEMC); “6°57.904'N, 66°36.392'W; 51 m; outcrop ca. 15 Km NE of Los Pijiguaos; detritus flotation; 9.vii.2010; leg. Short and Téllez; VZ10-0709-01B” (4, SEMC); 7°29'47.3"N, 65°51'44.8"W; 45 m; 2 km E of Río Cuchivero; rock outcrop seeps; 6.viii.2008; leg. Short, Téllez, Arias; AS-08-075 (1, SEMC, DNA voucher SLE1031).

By its elytral punctation with all punctures about the same size and degree of impression, T. communis is similar to T. anthonyae and T. microps. From T. microps, T. communis can be easily distinguished by the relative size of their eyes: in T. communis the eyes are separated by a distance 4.5 × larger than the largest diameter of the eye in dorsal view (Fig. 2H), whereas in T. microps the eyes are separated by a distance 6 × larger than the largest diameter of the eye in dorsal view (Fig. 2I). In T. communis the dorsal punctation is sharper, with all punctures seemingly longitudinally aligned, not forming grooves (Fig. 6A), whereas in T. anthonyae the dorsal punctation is relatively shallow, with serial punctures aligned in rows and slightly impressed, forming shallow longitudinal grooves (Fig. 7A).

Size and form : Body length 1.8–2.0 mm. Body elongate oval, somewhat parallel sided, moderately convex (Fig. 6A, B). Color and punctation: Dorsal and ventral surfaces of body dark brown, with anterior and lateral margins of prothorax, and outer margins of elytra slightly paler (Fig. 6A); mouthparts yellow (especially maxillary palps) to orange; antennae yellowish brown; legs orange brown with paler (yellow) tarsi (Fig. 6C). Ground punctation on head, pronotum and elytra sharply marked. Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed), and outer margins slightly bulging from outline of head (Fig. 2H); in lateral view, eyes not emarginate (e.g., Fig. 2E). Thorax: Elytra with all kinds of punctures similar in size and degree of impression, all seemingly longitudinally aligned and uniformly distributed, not forming grooves (Fig. 6A). Metafemora mostly glabrous on anterior face (Fig. 6C). Elevation of mesoventrite forming a longitudinal bulge (Fig. 3D). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 3D). Abdomen: Abdominal ventrites uniformly and very densely pubescent (Fig. 6C). Aedeagus (Fig. 11H). Basal piece nearly 0.4 × the length of a paramere; greatest width of a paramere nearly 0.6 × greatest width of median lobe; outer margins of parameres nearly straight along basal 2/3, then curved inwards; apex of paramere rounded; median lobe roughly sagittate, narrow along apical third, rounded at apex; gonopore situated nearly at midlength of median lobe.

Named with the Latin word communis meaning common, highlighting the abundance and wide distribution of the species, which is the most commonly found.

This species is the most widely distributed of all known Tobochares, occurring from the northwest margin of the Guiana Shield in Venezuela all the way to its eastern edge in the state of Amapá, Brazil (Fig. 12).

This species is strongly associated with seepage habitats on exposed granite. Many of the seepages on which this species has been collected are seasonal, although others are adjacent to permanent streams. See Fig. 16D, F.

Figure 16. 

Habitat of Tobochares in Suriname A, B habitat for T. kappel, Petrominas Falls (collecting event SR13-0813-04C) C habitat for T. microps, Tafelberg Tepui Summit (collecting event SR13-0820-01B) D habitat for T. kappel and T. communis, seepage along Kappel Airstrip (collecting event SR13-0824-02B) E habitat for T. microps, Tafelberg Tepui Summit (collecting event SR13-0815-01A) F habitat for T. akoerio and T. communis Grensgebergte Mountains (collecting event SR12-0312-01A).

Holotype (male): “Suriname: Sipaliwini District: 3°47.479'N, 56°8.968'W; 320 m; CSNR: near Kappel airstrip, wet rocks on sides of waterfall; 13.viii.2013; leg. Short and Bloom; SR13-0813-04B” (NZCS). Paratypes (53 exs.): Suriname: Sipaliwini District: same data as holotype (49, NZCS, SEMC); same, except “Petromina Falls; leg. Short, Bloom, Kadosoe; SR13-0813-04C” (4, SEMC).

Tobochares kappel can be recognized by its strongly convex body in lateral view, accompanied by elytral punctation uniform in size and degree of impression, not impressed to form elytral striae; the serial punctures are seemingly aligned in rows and the interserial punctures are somewhat irregularly distributed and moderately dense (in one or two rows, Fig. 9A, B). The general habitus and punctation of T. kappel are similar to those of T. arawak, T. canaima, and T. kolokoe. In T. arawak and T. canaima the interserial punctures are highly dense (forming two or three irregular rows, Fig. 3I). In T. kappel the coloration of pronotum and elytra gradually becomes paler (orange) towards the outer margins (Fig. 9A, B), and the legs are orange in color (Fig. 9C), whereas in T. kolokoe only the anterolateral margins of the pronotum are slightly paler and the legs are reddish brown in coloration (Fig. 9D–F). In addition, the serial punctures are equally impressed along the entire surface of the elytra in T. kappel (Fig. 9B) whereas in T. kolokoe the serial punctures become more impressed along the postero-lateral areas of the elytra (Fig. 9E).

Size and form : Body length 1.6–1.9 mm. Body elongate oval, strongly convex (Fig. 9A, B). Color and punctation: Dorsal and ventral surfaces of body dark brown, prothorax (especially its lateral areas) and outer margins of elytra slightly paler (Fig. 9A, B); mouthparts yellow (especially maxillary palps); antennae yellowish brown; legs orange brown with paler (yellow) tarsi (Fig. 9C). Ground punctation on head, pronotum and elytra moderately marked. Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed; e.g., Fig. 2D), and outer margins slightly bulging from outline of head; in lateral view, eyes not emarginate (e.g., Fig. 2E). Thorax: Elytral punctation uniform in size and degree of impression, not impressed to form elytral striae; serial punctures seemingly aligned in rows; interserial punctures somewhat irregularly distributed in one or two rows (Fig. 9A). Metafemora mostly glabrous on anterior face (Fig. 9C). Elevation of mesoventrite forming a very low transverse carina (Fig. 9C). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 9C). Abdomen: Abdominal ventrites uniformly and very densely pubescent (Fig. 9C). Aedeagus (Fig. 11L). Basal piece nearly 0.5 × the length of a paramere; greatest width of a paramere nearly 0.5 × greatest width of median lobe; outer margins of parameres weakly convex; apex of paramere narrowly rounded; median lobe roughly sagittate, pinched and rounded at apex; gonopore situated nearly at midlength of median lobe.

Noun in apposition. Named after the Kappel airstrip, the locality where the species has been collected.

Known from two very closely situated localities adjacent to Kappel Airstrip, at the foot to Tafelberg Tepui in Suriname. See Fig. 13.

Series were collected from two seepage habitats: one vertical seepage on sandstone adjacent to a large waterfall, and the second from a small mostly horizontal seepage on sandstone seeping into an adjacent stream. See Fig. 16A, B, D.

Holotype (female): “Suriname: Sipaliwini District: CSNR: Tafelberg Summit, Arrowhead Basin; flotation of wet moss on rocks; 20.viii.2013; leg. Short and Bloom; SR13-0820-04A” (NZCS).

Tobochares kolokoe can be recognized by its strongly convex body in lateral view (Fig. 9E), accompanied by elytral punctation uniform in size and degree of impression, not impressed to form elytral striae; the serial punctures are seemingly aligned in rows and the interserial punctures are somewhat irregularly distributed and moderately dense (in one or two rows, e.g., Fig. 3H). The general habitus and punctation of T. kolokoe are similar to those of T. arawak, T. canaima, and T. kappel. In T. arawak and T. canaima the interserial punctures are highly dense (forming two or three irregular rows, Fig. 3I). In T. kolokoe only the anterolateral margins of the pronotum are slightly paler than the general coloration of the pronotum and the legs are reddish brown in coloration (Fig. 9E, F), whereas in T. kappel the coloration of pronotum and elytra gradually becomes paler (orange) towards the outer margins (Fig. 9B), and the legs are orange in color (Fig. 9C). In addition, the serial punctures become more impressed along the postero-lateral areas of the elytra in T. kolokoe (Fig. 9E), whereas in T. kappel the serial punctures are equally impressed along the entire surface of the elytra (Fig. 9B).

Size and form : Body length 1.9 mm. Body elongate oval, strongly convex (Fig. 9E). Color and punctation: Dorsal and ventral surfaces of body dark brown, with prothorax (especially its anterolateral margins) slightly paler (Fig. 9D, E); mouthparts yellow (especially maxillary palps) to orange brown; antennae yellowish brown; legs orange brown with paler (yellow) tarsi (Fig. 9F). Ground punctation on head, pronotum and elytra moderately marked. Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed; e.g., Fig. 2D), and outer margins slightly bulging from outline of head; in lateral view, eyes not emarginate (e.g., Fig. 2E). Thorax: Elytral punctation uniform in size and degree of impression, not impressed to form elytral striae (Fig. 9D); serial punctures seemingly aligned in rows; interserial punctures somewhat irregularly distributed in one or two rows (e.g., Fig. 3H). Metafemora mostly glabrous on anterior face (Fig. 9F). Elevation of mesoventrite forming a very low transverse carina (Fig. 9F). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 9F). Abdomen: Abdominal ventrites uniformly and very densely pubescent (Fig. 9F).

Noun in apposition. Named with the Surinamese word kolokoe meaning lucky, as this species is known from a single female specimen.

This species is only known from the summit of Tafelberg Tepui, a low-elevation sandstone massif in the center of Suriname (Fig. 13).

The lone specimen of this species was collected by floating saturated moss that was growing on rocks by a waterfall. The males of this species remain unknown.

Holotype (male): “Suriname: Sipaliwini District/ N3 53.359' W56 10.052', 879m/ CSNR: Tafelberg Summit, near/South Rim, small seepage area/leg. Short and Bloom; 20.viii.2013/SR13-0820-01B” (NZCS). Paratypes (43 exs.): Suriname: Sipaliwini District: Same data as holotype (38, SEMC including DNA voucher SLE1041); “3°55.600'N, 56°11.300'W; 600 m; CSNR: Tafelberg Summit, near Augustus Creek Camp; muddy stream pools; 15.viii.2013; leg. Short and Bloom; SR13-0815-01A” (1, SEMC DNA voucher SLE1051); same, except “pool in rock; SR13-0820-01C” (2, SEMC including DNA voucher SLE 1043); CSNR: Tafelberg Summit, Arrowhead Basin; flotation of wet moss on rocks; 20.viii.2013; leg. Short and Bloom; SR13-0820-04A (2, SEMC including DNA vouchers SLE1038 and SLE1040).

Tobochares microps is unique among Tobochares species by the reduced size of the eyes, which are separated by a distance 6 × larger than the largest diameter of the eye in dorsal view (Fig. 2I; the ventral face of the eye is only slightly wider than the antennal club, Fig. 6F), whereas in all other species in the genus the eyes are separated by a distance approximately 4.5 × larger than the largest diameter of the eye in dorsal view (e.g., Fig. 2H; the ventral face of the eye is nearly twice as wide as antennal club; e.g., Fig. 6C). Tobochares microps is polymorphic for hindwings, with individuals exhibiting either full size or brachypterous wing forms (e.g., Fig. 3K), a condition so far unique in Tobochares. By its elytral punctation with all punctures about the same size and degree of impression (Fig. 6D), T. microps is similar to T. communis and T. anthonyae. Besides the shape of the eyes, T. microps, can be distinguished from T. communis by its relatively shallower punctation and smaller size (compare Fig. 6D to Fig. 6A). From T. anthonyae, in which the dorsal punctation is also relatively shallow (Fig. 7A), T. microps can be recognized by the serial punctures seemingly longitudinally aligned and uniformly distributed, not impressed to form grooves (serial punctures clearly aligned in rows and slightly impressed, forming shallow longitudinal grooves in T. anthonyae; Fig. 7A).

Size and form : Body length 1.6–1.7 mm. Body elongate oval, moderately convex (Fig. 6E). Color and punctation: Dorsal and ventral surfaces of body dark brown, with anterior and lateral margins of prothorax slightly paler (Fig. 6D, E); mouthparts yellow (especially maxillary palps) to orange; antennae yellowish brown; legs orange brown with paler (yellow) tarsi (Fig. 6F). Ground punctation on head, pronotum and elytra moderately marked. Head: Eyes in dorsal view with anterior margin slightly oblique (anteriorly directed), and outer margins nearly half the greatest length of eye, slightly bulging from outline of head (Fig. 2I); in lateral view, eyes not anteriorly emarginate (e.g., Fig. 2E). Thorax: Elytra with all punctures about the same size and degree of impression, all seemingly aligned in rows and uniformly distributed, not forming grooves (Fig. 6D). Metafemora mostly glabrous on anterior face (Fig. 6F). Elevation of mesoventrite forming a broad bulge (Fig. 6F). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 6F). Abdomen: Abdominal ventrites uniformly and very densely pubescent. Aedeagus (Fig. 11I). Basal piece nearly 0.4 × the length of a paramere; greatest width of a paramere nearly 0.5 × greatest width of median lobe; outer margins of parameres uniformly weakly convex; apex of paramere rounded; median lobe roughly sagittate, narrow along apical third, rounded at apex; gonopore situated basad of midlength of median lobe.

Named with the combination of the Latin words micro, meaning small, and ops, meaning eyes, highlighting the small eyes of the members of the species.

This species is only known from the summit of Tafelberg Tepui, a low-elevation sandstone massif in the center of Suriname (Fig. 13).

Most specimens of this species were collected in seepage habitats by directly floating them out of saturated moss that was on the rock. Several specimens were collected in shallow pools on rock that were adjacent to seepages or streams. See Fig. 16C.

Holotype (male): “Venezuela: Bolívar: 5°51'N, 62°33'W; 1700 m; Auyan-tepui; Intercept trap; 7–14.ii.1994; leg. J.L. García, A. Chacón” (MIZA). Paratypes (6 exs.): Same data as holotype (6, MIZA, SEMC).

The general orange coloration with dark head of T. pemon is quite distinct among Tobochares, and particularly among members of the communis group. In addition, its elytral punctation is relatively unique, including all kinds of punctures being similar in size and degree of impression, with serial punctures aligned in rows, but not forming grooves, and with interserial punctures somewhat irregularly distributed in a single row (Figs 3F, 7G, H). In addition, the median lobe of the aedeagus of T. pemon is unique, as it is uniformly broad throughout and apically broadly emarginate (Fig. 11E); the median lobe in other species typically narrows towards the apex and is usually rounded, except for T. luteomargo, which has an emarginated median lobe, but in this case the emargination is deep and very narrow (Fig. 11D).

Size and form : Body length 1.7–1.8 mm. Body elongate oval, moderately convex (Fig. 7H). Color and punctation: Dorsal surfaces of body orange brown, with lateral margins of prothorax slightly paler (Fig. 7G, H); ventral surfaces of body (except prosternum) dark brown; mouthparts orange brown; antennae light brown; legs, including tarsi orange brown (Fig. 7I). Ground punctation on head, pronotum and elytra rather shallowly marked. Head: Eyes in dorsal view with anterior margin slightly oblique (anteriorly directed; Fig. 2D); in lateral view, eyes not anteriorly emarginate (e.g., Fig. 2E). Thorax: Elytra with all kinds of punctures similar in size and degree of impression; serial punctures aligned in rows, not forming grooves; interserial punctures somewhat irregularly distributed in a single row (Fig. 3F). Metafemora mostly glabrous on anterior face (Fig. 7I). Elevation of mesoventrite forming a low transverse carina (Fig. 7I). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 7I). Abdomen: Abdominal ventrites uniformly and very densely pubescent (Fig. 7I). Aedeagus (Fig. 11E) with basal piece nearly 0.6 × the length of a paramere; greatest width of a paramere nearly 0.7 × greatest width of median lobe; outer margins of parameres straight and slightly converging along basal 2/5, then uniformly and widely convex; apex of paramere rounded; median lobe roughly rectangular, with wide and short medial emargination at apex; gonopore situated at apical fourth of median lobe.

Noun in apposition. Named after the Pemon, an indigenous tribe that inhabits La Gran Sabana region in Venezuela, where Auyan Tepui is located.

This species is known from the famous Auyan-tepui, which is also home to Angel Falls, the highest waterfall in the world. Collected at an elevation of 1700 m, this species is one of the relatively few water beetle taxa known from the “Pantepui Province”, which comprises areas of the Guiana Shield which are greater than 1500 m in elevation (Fig. 13).

The only known series was collected in a flight intercept trap. Nothing further is known about the habitat or biology of this species.

Holotype (male): “Brazil: Roraima: Amajari; 3°36.381'N, 61°42.878'W; 618 m; Serra do Tepequém, Igarape Preto Negro, Cachoeira Leje Preta; at edge of seepage, root mats and moss; 14.i.2018; leg. Short and Benetti; BR18-0114-04B”; DNA voucher SLE1493 (INPA).

Tobochares romanoae can be recognized by its elytra with well-defined and moderately impressed rows of serial punctures; the impressed stria I is more strongly impressed along the posterior half of the elytra, resembling a well-developed sutural stria (Fig. 8A); interserial punctures somewhat irregularly distributed (Fig. 8A, B). The general habitus of T. romanoae is similar to that of T. akoerio and T. canaima, especially by the uniformly dark coloration of the pronotum; T. romanoae (Fig. 8A) can be distinguished from T. canaima by the impressed striae in T. romanoae (compare with Fig. 10D); from T. akoerio the moderately convex body T. romanoae allows its recognition (compare Fig. 8B to 8E). The only known specimen was extracted for DNA, so the dark brown coloration might not reflect the true coloration of the species.

Size and form : Body length 2.6 mm. Body elongate oval, moderately convex. Color and punctation: Dorsal and ventral surfaces of body, antennae and legs (except yellowish tarsi) dark brown; mouthparts light brown (Fig. 8A–C). Ground punctation on head, pronotum and elytra moderately marked. Head: Eyes in dorsal view with anterior margin oblique (anteriorly directed), and outer margins bulging from outline of head; in lateral view, eyes not anteriorly emarginate. Thorax: Elytra with all kinds of punctures similar in size and degree of impression; serial punctures aligned in rows, impressed as to form longitudinal grooves (Fig. 8A, B); elytral stria I more strongly impressed along posterior half of elytra, resembling well-developed sutural stria (Fig. 8A); interserial punctures somewhat irregularly distributed (Fig. 8A, B). Metafemora mostly glabrous on anterior face, with hydrofuge pubescence along basal half of antero-dorsal margin (Fig. 8C). Elevation of mesoventrite forming a very low transverse carina (Fig. 8C). Metaventrite with distinct median, longitudinal, narrow glabrous area extending along posterior half (Fig. 8C). Abdomen: Abdominal ventrites uniformly and very densely pubescent (Fig. 8C). Aedeagus (Fig. 11J). Basal piece nearly 0.5 × the length of a paramere; greatest width of a paramere nearly 0.4 × greatest width of median lobe; outer margins of parameres very weakly convex; apex of paramere rounded; median lobe roughly triangular, rounded at apex; gonopore situated at midlength of median lobe.

Named after Rosina Romano, Director of Meetings and Membership at the Entomological Society of America (ESA), in recognition of all her hard work and dedication to the society and entomological community.

This species is only known from the summit of Serra do Tepequem, a low-elevation sandstone massif in northern Brazil, near the border with Venezuela (Fig. 13).

The only known specimen was collected by floating rootlets and moss that were present on a rocky seepage formed along the margin of the Igarape (stream) Preto Negro (Fig. 14E).