Belonuchus alternans ( Sharp, 1885 )

Márquez, Juan & Asiain, Julieta, 2022, Taxonomy of the Mexican species of Belonuchus Nordmann (Coleoptera: Staphylinidae), Zootaxa 5152 (1), pp. 1-129 : 20-21

Belonuchus alternans ( Sharp, 1885)

Figs. 2a, b , 14c , 15a , 18a , 21a , 26a

Philonthus alternans Sharp, 1885: 418 . Belonuchus alternans comb. nov. ex Philonthus ( Navarrete-Heredia et al. 2002) .

Total body length in males 10.27 mm (range 9.20–11.0 mm), in females 10.71 mm (range 9.30–11.4 mm). Black on head, antennae, mandibles, pronotum, scutellum and fourth (except its anterior border that is reddish) to sixth visible abdominal segments. Reddish on elytra, mesoventrite, metaventrite, legs (except procoxae that are black) and first three visible abdominal segments. Apical antennomere, maxillary and labial palpi, and prosternum brown; genital segment yellowish.

Head: transverse in both sexes; ratio length/width in males 0.715 (range 0.64–0.83), in females 0.825 (range 0.77–0.86). Dorsal surface with sulcate longitudinal midline visible from anterior margin of head to level of posterior margin of eyes; front with distinct and broad fovea between antennal insertions and a little posteriorly where it becomes narrower. Eyes slightly longer than 0.5 times the cephalic lateral length and slightly protruding. Antennomeres 2–3 almost with same length, 4–5 slightly elongate, 6 nearly as long as wide, 7–10 slightly transverse. Males with mandibles longer than head (ratio 1.3, range 1.00–1.48); females with mandibles slightly shorter than head (ratio 0.94, range 0.81–1.09); mandibles with two (basal and middle) clearly separated teeth; mandibular channel well developed, external margin well separated from internal margin; internal margin carinate until near the level of middle tooth, then extending higher as impressed line. Apical palpomere of maxillary and labial palpi nearly 1.5 times longer than preapical palpomere. Ventral surface with microsculpture less visible than on dorsal surface. Head wider than pronotum, both in males (ratio 1.21, range 1.14–1.28) and females (ratio 1.07, range 1.03–1.11).

Thorax: pronotum with five punctures in each dorsal row; slightly longer than wide, both in males (ratio 1.11, range 1.03–1.15) and females (ratio 1.16, range 1.03–1.22); almost as wide at anterior corners as at posterior corners in both sexes (ratio in males 1.05, range 1.04–1.10, in females 1.02, range 1.00–1.06). Scutellum with punctures denser than on elytra; elytra with sparse and fine punctures, with pale setae. Prosternum with an elevated area before anterior margin. Intercoxal process of mesoventrite with shape of acute shield; transverse discal ridge very widely U-shaped or V-shaped, not meeting margin of intercoxal process laterally. Males with profemora with external row of spines, starting near basal third and not reaching the apex, spines do not overlap with those of internal apex, which are among three to five of moderate size. Hind legs of males with trochanter modified like a hook, in large specimens slightly less than 1/2 the length of metafemur, in smaller specimens as long as 1/4 the length of metafemur and its apex is less convex; femur curved at level of its overlap with hook; tibia a little curved in apical third. Tarsomeres very slender and scarcely flattened dorsally.

Abdomen: first three visible tergites with posterior basal transverse carina, followed by a slightly visible slightly depressed area, with smooth microsculpture and some wide-superficial punctures; remaining area of tergites 1–3 and following tergites only with fine punctures. Visible sternites 1–2 with wide-superficial punctures denser than on tergites, covering more than anterior half, combined with fine punctures; sternites 3–4 with sparse wide-superficial punctures covering less than anterior half; sternite 5 with sparse wide-superficial and fine punctures; sternite 6 only with fine punctures. Male pregenital sternite evenly rounded at posterior margin ( Fig. 15a ). Male genital sternite very elongate (3.4 times longer than wide), more or less symmetrical, anterior portion occupies 42% and posterior portion occupies 48% of its length, apical emargination very deep ( Fig. 18a ). Male abdominal styli black, moderately widened at apex ( Fig. 14c ), although less wide and robust than in B. trochanterinus .

Aedeagus: length 1.4 mm; elongate shape; apex slightly acute, somewhat rounded; in ventral view, basal half slightly wider than apical half; in lateral view the greater width of basal half is more noticeable; internal sac visible ( Fig. 21a ).

Variability. In addition to the variation in the measurements and ratios included in the description, only differences related to the sex and size of the specimens were observed.

Taxonomic comments. Belonuchus alternans is very similar to B. magnistylus , from which it is distinguished almost exclusively by the fact that the latter species has the male abdominal styli notably wider, pregenital sternite with the posterior margin emarginate and a different shape of the aedeagus. In B. alternans the abdominal styli of males are only moderately wider, the pregenital sternite is evenly rounded at posterior margin and the aedeagus is less wide at base. Additionally, B. alternans could be confused with B. trochanterinus and B. confusus for having a similar coloration pattern and size, as well as the wider male abdominal styli. It can be distinguished because B. alternans has the last three visible abdominal segments black (except the anterior border of the fourth visible segment that is reddish), male abdominal styli moderately wider and the genital sternite and the aedeagus of different shape in comparison with the other two species, which have only the last two visible abdominal segments black (except the anterior border of fifth visible segment that is reddish) and the male abdominal styli notably wider.

The syntypes of this species were not studied in person, but we analyzed some photos of these specimens ( Fig. 2a ; NHM, shared by M. Chani Posse) which, together with the original description ( Sharp, 1885), allowed us to identify this species.

Material examined (448 males, 431 females): Mexico: Hidalgo: “Acaxochitlán, Tlazintla, bosque encinopino, 2149 m, N20°09’7.96”, W98°11’1.83”, NTP-80 (calamar), (without data) J. Islas col.” (8, CC-UAEH). “Chapulhuacán, El Infiernillo, selva mediana, 500-850 m, N21°7.54’, W98°58.001’, hojarasca cernida, 20-IV-2013, J. Márquez col.” (1, CC-UAEH). “Chapulhuacán, Tamaulipas, bosque mesófilo de montaña, 1133 m, N21°10.036’, W98°55.984’, NTP-80 (calamar) #3, 30-III a 06-IV-2005, J. Márquez, M. Rivero, M. Torres, M. Vargas y J. Sánchez cols.” (2, CC-UAEH). Same data, except: “ N21°09.9’ 7.9”, W98°55.8’ 04.0”, NTP-80 (calamar) #3, 12 a 26-III- 2011. J. Márquez col.” (2, CC-UAEH). Same data, except: “NTP-80 #7 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #6 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #4 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #1 (calamar)” (2, CC-UAEH). Same data, except: “NTP-80 #5 (calamar)” (2, CC-UAEH). Same data, except: “NTP-80 #8 (calamar)” (2, CC-UAEH). Same data, except: “NTP-80 #2 (calamar)” (1, CC- UAEH). Same data, except: “Coprotrampa” (4, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #3” (1, CC-UAEH). Same data, except: “NTP-80 #5 (calamar), 28-VIII a 11-IX-2011 ” (1, CC-UAEH). Same data, except: “NTP-80 #9 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #1 (calamar)” (1, CC-UAEH). “Huehuetla, Rio Blanco, bosque tropical perennifolio, 588 m, UTM (WG-384), 145 94809 E, 2257303 N, NTP-80 (calamar), 1 al 30-V-2010, C. Berriozábal col.” (9, CC-UAEH). “ Jacala de Ledezma, 1 km E de El Durazno, bosque pino-encino- Junniperus, 1694 m, N20°3’11.1”, W99°1’20”, NTP- 80 (calamar), 8 al 16-III-2008, J. Márquez y J. Asiain cols.” (4, CC-UAEH). “Jacala de Ledezma, 3 km antes de Las Moras, bosque mesófilo de montaña, 1678 m, trampa de intercepción de vuelo, 3-XII-2008, J. Márquez col. “ (17, CC-UAEH). “Juárez, bosque mesófilo perturbado, 1970 m, N20°47’9.6”, W98°49’5.2”, trampa de intercepción de vuelo #2, 7 al 14-III-2009, J. Márquez, M. Rivero, M. Torres, M. Vargas y J. Sánchez cols.” (3, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1” (1, CC-UAEH). Same data, except: “NTP-80 #3 (calamar)” (4, CC-UAEH). Same data, except: “NTP-80 #1 (calamar)” (1, CC-UAEH). “La Misión, Lomas del Pericón, bosque mesófilo de montaña, 1377 m, N21°60’46.0”, W99°6.0’ 15.6”, NTP-80 (calamar), 8 al 16-III-2008, J. Márquez y J. Asiain cols.” (38, CC-UAEH). Same data, except: “trampa de intercepción de vuelo” (4, CC-UAEH). Same data, except: “NTP-80 (calamar) #3, 26-VIII a 9- IX-2011, J. Márquez col.” (1, CC-UAEH). “La Misión, bosque mesófilo de montaña, 1337 m, N21°06’202”, W99°06’291”, NTP-80 (calamar) #1, 29-III al 05-IV-2009, J, Márquez, M. Rivera, M. Torres, M. Vargas y J. Sánchez cols.” (3, CC-UAEH). “La Misión, Los Naranjos, Palo Hueco, bosque mesófilo de montaña, 1276 m, N21°08’06”, W99°05’02”, NTP-80 (calamar) #6, 10 a 24-III-2011, J. Márquez col.” (2, CC-UAEH). “La Misión, Palo Hueco, 1.5 km al NE de Puerto de Naranjos, bosque mesófilo de montaña, 1197 m, N21°08’21”, W99° 05’4.6”, coprotrampa, 10 a 24-III-2011, J. Márquez col.” (3, CC-UAEH). “La Misión, camino al río Moctezuma, N21°8’30.6”, W99°5’49.1”, bosque tropical subcaducifolio, trampa de intercepción de vuelo, 23-VI a 5-VII-2017, J. Márquez y A. Lora cols.” (7, CC-UAEH). “Molango, Acuatitlán, bosque mesófilo de montaña, 1715 m, N20°45’38.4”, W88°42’50.7”, en tronco con hongos, 10-IV-2008, J. Márquez, J. Bueno y M. García cols.” (2, CC-UAEH). “Molango, Acuatitlán, bosque mesófilo de montaña, 1715 m, N20°45’38.4”, W88°42’50.7”, NTP-80 (calamar) #2, 15 a 29-IV-2011, J. Márquez col.” (27, CC-UAEH). Same data, except: “NTP-80 #1 (calamar)” (25, CC-UAEH). Same data, except: “NTP-80 #4 (calamar)” (22, CC-UAEH). Same data, except: “NTP-80 #7 (calamar)” (19, CC-UAEH). Same data, except: “NTP-80 #3 (calamar)” (6, CC-UAEH). Same data, except: “NTP-80 #6 (calamar)” (10, CC- UAEH). Same data, except: “NTP-80 #5 (calamar)” (13, CC-UAEH). Same data, except: “NTP-80 #8 (calamar)” (4, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1” (2, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #3” (3, CC-UAEH). Same data, except: “NTP-80 #3 (calamar), 30-IX a 21-X- 2011 ” (1, CC-UAEH). Same data, except: “hojarasca cernida, 12-IV-2011 ” (1, CC-UAEH). “Nicolás Flores, cerca de Villahermosa, bosque de encino, 2221 m, N20°43’49.7”, W99°07’35.2”, NTP-80 #3 (calamar), 21 a 28-II-2009, J. Márquez, M. Rivero, M. Torres y J. Sánchez cols.” (1, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #2” (4, CC-UAEH). “Pacula, camino al Membrillar del Fraile, río Moctezuma, N99°23´28”, W21°5´26.8”, 903 m, vegetación riparia, NTP-80 (calamar), 19-I a 25-II-2010, J. Islas col. “ (7, CC-UAEH). “Pacula, Puerto Grande, bosque de encino, 2109 m, N20°53.850’, W99°19.562’, NTP- 80 #3 (calamar), 28-III al 04-IV-2009, J. Márquez, M. Rivero, M. Torres, M. Vargas y J. Sánchez cols.” (2, CC-UAEH). Same data, except: “NTP-80 #1 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #2 (calamar)” (1, CC-UAEH). “Tenango de Doria, El Potrero, bosque mesófilo de montaña, 2185 m, N20°18’49.5”, W98°13’43.3”, NTP-80 #1 (calamar), 18 a 25-IV- 2009, J. Márquez, M. Rivero, M. Torres, M. Vargas y J. Sánchez cols.” (1, CC-UAEH). “Tenango de Doria, camino a La Viejita, bosque mesófilo de montaña, 1647 m, N20°19.0’ 37.9”, W98°14’ 39.7”, NTP-80 #2 (calamar), 29-III a 12-IV-2011, J. Márquez col.” (1, CC-UAEH). Same data, except: “NTP-80 #1 (calamar)” (2, CC-UAEH). Same data, except: “NTP-80 #7 (calamar)” (3, CC-UAEH). Same data, except: “NTP-80 #5 (calamar)” (4, CC-UAEH). Same data, except: “NTP-80 #3 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #3 (calamar), 1 a 15-IX- 2011.” (1, CC-UAEH). Same data, except: “NTP-80 #4 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #1 (calamar)” (3, CC-UAEH). Same data, except: “NTP-80 #6 (calamar)” (3, CC-UAEH). Same data, except: “NTP- 80 #5 (calamar)” (1, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #2, 5 a 12-IV-2013, N. Gutiérrez col. “ (2, CC-UAEH). Same data, except: “trampa de intercepción de vuelo # 1” (3, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1, 11 a 16-VI-2013 ” (1, CC-UAEH). Same data, except: “trampa de intercepción de vuelo # 3” (1, CC-UAEH). Same data, except: “NTP-80 (calamar), 12-VIII-2010, C. Cornejo col.” (9, CC-UAEH). “Tepehuacán de Guerrero, Chilijapa, bosque mesófilo de montaña perturbado, 1323 m, N21°1.91’, W98°51.812’, NTP-80 (calamar), 22 a 24-IV-2004, J. Asiain, J. Islas, J. Márquez cols.” (1, CC-UAEH). “Tepehuacán de Guerrero, Chilijapa, bosque mesófilo de montaña, 1401 m, N21°1.2’ 5.1”, W98°51.8’ 2.1”, NTP-80 (calamar) #2, 11 a 25-III-2011, J. Márquez col.” (13, CC-UAEH). Same data, except: “NTP-80 #3 (calamar)” (7, CC- UAEH). Same data, except: “NTP-80 #1 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #9 (cal.)” (3, CC-UAEH). Same data, except: “NTP-80 #6 (calamar)” (6, CC-UAEH). Same data, except: “NTP-80 #8 (calamar)” (11, CC-UAEH). Same data, except: “NTP-80 #7 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #5 (calamar)” (7, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #2.” (1, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1.” (5, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1, 27-VIII a 10-IX-2011 ” (2, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #2.” (4, CC- UAEH). Same data, except: “trampa de intercepción de vuelo #3.” (1, CC-UAEH). Same data, except: “NTP-80 (calamar) #1” (1, CC-UAEH). Same data, except: “NTP-80 #2 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #10 (calamar)” (4, CC-UAEH). Same data, except: “coprotrampa, 12 a 26-III-2011.” (2, CC-UAEH). Same data, except: “Coprotrampa” (12, CC-UAEH). Same data, except: “NTP-80 (calamar)” (4, CC-UAEH). Same data, except: “NTP-80 (calamar) #4” (22, CC-UAEH). Same data, except: “en hojarasca cernida, 19-IV-2013 ” (1, CC-UAEH). “Tepehuacán de Guerrero, Otongo, 650 m, 2 marzo 1981, ex carrion trap, M. A. Morón” (1, IEXA). Same data, except: “ 1120 m, may-june, 1981” (1, IEXA). “Tianguistengo, camino a Santa Mónica, bosque mesófilo de montaña 1868 m, N20°43’5.4”, W98°39’40.3”, trampa de intercepción de vuelo, 7 a 13-VI-2009, J. Sánchez, M. Rivero, M. Vargas y J. Márquez cols.” (3, CC-UAEH). “Tepehuacán de Guerrero, San Juan Ahuehueco, selva mediana, 1016 m, N21°01.766’, W98°54.285’, NTP-80 (calamar) #1, 29-III al 05-IV-2009, J. Márquez, M. Rivero, M. Torres, M. Vargas y J. Sánchez cols.” (1, CC-UAEH). “Tlahuiltepa, 500 m al sureste de Tlaxcantitla, N20°51’41.03”, W98°53’17.03”, NTP-80 #1, 17 al 14-III-2009, J. Márquez, M. Rivero, M. Torres, M. Vargas y M. Sánchez cols.” (6, CC-UAEH). “Tlanchinol, bosque mesófilo de montaña, en necrotrampa temporal, 15 al 19-V- 1996, J. Márquez col.” (6, MAAS). Same data, except: “NTP- 80 (cal.), 29-IV a 29-V de 1995, A. Soria col.” (2, MAAS). “ 3 km N de Tlanchinol, bosque mesófilo de montaña, 1450 m, N21°1.328’, W98°38.77’, en hojarasca, 1- V-2003, J. Asiain y J. Márquez cols.” (2, CC-UAEH). Same data, except: “NTP- 80 (calamar) 16-IV a 1-V-2003, J. Asiain y J. Márquez cols.” (6, CC-UAEH). “Tlanchinol camino a Apantlazol, El Pozo, bosque mesófilo de montaña, 1391 m, N20°59’6.73”, W98°37’39.86”, NTP-80 (calamar) #6, 1 a 22-X-2011, J. Márquez col.” (1, CC-UAEH). Same data, except: “NTP-80 #4 (calamar)” (1, CC-UAEH). Same data, except: “NTP-80 #1(calamar)” (1, CC- UAEH). Same data, except: “trampa de intercepción de vuelo #2” (2, CC-UAEH). Same data, except: “NTP-80 #2 (calamar), 16 a 30-IV-2011.” (8, CC-UAEH). Same data, except: “NTP-80 #4 (calamar)” (7, CC-UAEH). Same data, except: “NTP-80 #6 (calamar)” (13, CC-UAEH). Same data, except: “NTP-80 (calamar) #5” (8, CC-UAEH). Same data, excep: “NTP-80 (calamar) #1” (10, CC-UAEH). Same data, except: “NTP-80 (calamar) #2” (8, CC- UAEH). Same data, except: “trampa de intercepción de vuelo #3” (27, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1” (9, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #2” (6, CC-UAEH). “Tlanchinol, La Cabaña, bosque mesófilo de montaña, 1462 m, N21°01.333’, W98°38.650’, trampa de intercepción de vuelo #4, 15 a 22-IV-2006, C. Ortiz y M. C. Pedraza cols.” (1, CC-UAEH). Same data, except: “ 1478 m, N21°0.343’, W98°38.600’, trampa de intercepción de vuelo # 5, 16 a 23-VII-2005, M.C. Pedraza, E. Pedraza y C. Ortiz cols.” (1, CC-UAEH). Same data, except: “ 1485 m, N21°1.357’, W98°38.556’, trampa de intercepción de vuelo # 3, 17 a 24- IX-2005 ” (1, CC-UAEH). Same data, except: “ 1477 m, N21°01’15.94”, W98°38’40.14”, en hojarasca cernida, 8- VI-2007, J. Márquez y J. Asiain cols.” (1, CC-UAEH). Same data, except:“ 1462 m, N21°1.338’, W98°650’, trampa de intercepción de vuelo, 13 al 20- V-2006, C. Ortiz y M.C. Pedraza cols.” (1, CC-UAEH). Same data, except: “ 1478 m, N21°01.343’, W98°650’” (1, CC-UAEH). “Tlanchinol, cerca de la Cabaña, bosque mesófilo de montaña, 1450 m, N21°01.328’, W98°38.770’, en hongos de repisa de troncos, 16-IV-2003, J. Márquez y J. Asiain cols.” (2, CC-UAEH). “Zacualtipán, camino a Sto. Domingo, bosque mesófilo de montaña, 1830 m, N20°37.749’, W98°34.851’, NTP-80 (calamar), 30-VII al 30-X-2003, J. Márquez y J. Asiain cols.” (2, CC-UAEH). “Zacualtipán camino a Tizapán, bosque mesófilo de montaña, 1790 m, N 20°38’44.5”, W98°36’7.2”, NTP-80 (calamar) #3, 14 a 28-IV-2011, J. Márquez col.” (8, CC-UAEH). Same data, except: “NTP-80 #9 (calamar)” (9, CC- UAEH). Same data, except: “NTP-80 (calamar) #10” (5, CC-UAEH). Same data, except: “NTP-80 (calamar) #6” (4, CC-UAEH). Same data, except: “NTP-80 (calamar) #7” (1, CC-UAEH). Same data, except: “NTP-80 (calamar) #8” (11, CC-UAEH). Same data, except: “NTP-80 (calamar) #1” (4, CC-UAEH). Same data, except: “NTP-80 (calamar) #5” (9, CC-UAEH). Same data, except: “NTP-80 (calamar) #2” (1, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1” (3, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #3” (6, CC- UAEH). Same data, except: “trampa de intercepción de vuelo #3, 20-IX a 4-X-2011 ” (1, CC-UAEH). Same data, except: “NTP-80 (calamar) #10, 20-IX a 4-X-2011 ” (2, CC-UAEH). Same data, except: “NTP-80 (calamar) #2 “ (1, CC-UAEH). “Zimapán, P.N. Los Mármoles, Minas Viejas, bosque de encino, 1892 m, N20°55’5.2”, W99°12’41.1”, NTP-80 (calamar), 11-X al 17-XI-2007, J. Márquez, J. Asiain y S. Sierra cols.” (4, CC-UAEH). Same data, except: “coprotrampa, 18-VIII al 20-IX-2007.” (1, CC-UAEH). Same data, except: “necrotrampa.” (1, CC-UAEH). Same data, except: “NTP-80 (calamar), 13-VII al 18-VIII-2007.” (5, CC-UAEH). “P. N. Los Mármoles, Zimapán, La Encarnación, bosque de encino, 2412 m, N20°52’1”, W99°12’50.3”, Coprotrampa, 30-V al 23-IV-2007, J. Márquez, J. Asiain y S. Sierra cols.” (5, CC-UAEH). Same data, except: “NTP-80 (calamar), 23-VI al 13-VII-2007.” (1, CC-UAEH).”Zimapán, El Salto a 2 km al E de Durango, bosque de encino, 2099 m, N20°54.023 ”, W99°12.933’, NTP-80 #1 (calamar), 28-III a 4-IV-2009, J. Márquez, M. Rivero, M. Torres, M. Vargas y J. Sánchez cols.” (1, CC- UAEH). Same data, except: “14Q0477632, UTM 2311197, 2155 m, NTP-80 #3 (calamar)” (1, CC-UAEH). “Zimapán, Minas Viejas, bosque de encino, 1897 m, N20°55’23”, W99°12’28”, en excremento vacuno, 20-X-2006, J. Asiain y J. Márquez cols.” (1, CC-UAEH). “Zimapán, P. N. Los Mármoles, bosque de pino-encino, 2444 m, N20°48’12.22”, W99°14’41”, coprotrampa, del 22-VI al 12-VII-2007, J. Márquez, J. Asiain y S. Sierra cols.” (2, CC-UAEH). “Zacualtipán, La Mojonera, N20º37’59”, W98º35’48”, 1886 m, bosque mesófilo de montaña perturbado, NTP-80 (calamar), 28-II a 29-IV-2004, J. Asiain y J. Márquez cols.” (1, CC-UAEH). Same data, except: “mayo-junio, 2004” (1, CC-UAEH). “Zacualtipán, camino a Santo Domingo, N20º37’44.9”, W98º34’51.06”, 1830 m, bosque mesófilo de montaña, NTP-80 (calamar), 28-II a 29-IV-2004, J. Márquez y J. Asiain cols.” (11, CC- UAEH). Oaxaca: “Candelaria Loxicha, Portillo del Rayo, VI- 1990, 950 m, bosque tropical y cafetal, C. Deloya López col.” (2, IEXA). Puebla: “Honey, Chila de Juárez, río San Marcos, N20.277778°, W98.213611°, bosque mesófilo de montaña, 1500 m, NTP-80 (calamar), 8 a 15-VIII-2019, J. D. Silva, R. Ramírez y J. Márquez cols.” (m1, CC-UAEH). Same data except: “en carpotrampa” (1, CC-UAEH). “Zacapoaxtla, Ex-hacienda Apulco, cascada La Gloria, N19°54´2”, W97°37´00”, 1,508 m, dentro de cañada bosque mesófilo de montaña rodeado por pinos, trampa de intercepción de vuelo, 18 a 28-V-2021, familia Márquez-Asiain cols.” (14, MAAS). Same data except: “NTP-80 (calamar)” (2, MAAS). Querétaro: “Pinal de Amoles, cascada del Chuveje, 1350 m, micotrampa, 6 a 10- VIII-2002, Q. Santiago y L. Delgado cols.” (1, IEXA). Veracruz: “Banderilla, La Martinica, bosque mesófilo de montaña, 1550 m, en hongo Armillariella polymyces # 42, 19-VI-2001, L. Delgado col.” (1, IEXA). “Córdoba, San Rafael, Calería, Ojo de Agua, selva media subcaducifolia, 1127 m, en lima podrida, 24-I-1999, Q. Santiago, J. Asiain y J. Márquez cols.” (5, MAAS). Same data, except: “Q. Santiago y J. Márquez cols.” (1, MAAS). “Coscomatepec, Tecoac, bosque mesófilo de montaña, 1750 m, NTP-80 (calamar), 6-V a 3-VI-1995, M. A. Pensado y Q. J. Santiago cols.” (1, QJSJ). “Huatusco, Coxcontla, bosque mesófilo de montaña perturbado, 1250 m, NTP-80 (calamar) #200, 22-X a 19-XI- 1994, M. A. Pensado y Q. Santiago cols.” (3, QJSJ; 1, MAAS). Same data, except: “NTP-80 (calamar) #419, 17-XII-1994 a 14-I-1995 ” (1, MAAS; 1, QJSJ). Same data, except: “NTP-80 (calamar) #853, 29-VII a 26-VIII-1995 ” (1, MAAS). Same data, except: “ 6-V a 3-VI-1995 ” (1, QJSJ). Same data, except: “ 3- VI a 1-VII-1995 ” (1, QJSJ). Same data, except: “ 23-IX a 21-X-1995 ” (2, QJSJ). “Huayacocotla, San Antonio, bosque mesófilo de montaña, 1435 m, N20°40’44.6”, W98°23’06.3”, NTP-80 (calamar) #2, 01-V a 28-V-2006, F. Ramírez col.” (1, CC-UAEH). Same data, except: “NTP-80 (calamar) #1” (1, CC-UAEH). “Huayacocotla, Helechales, bosque mesófilo de montaña, 1936 m, N20°37.5’ 25.0”, W98°27.6’ 49”, NTP-80 (calamar) #3, 19-III a 2-IV-2011, J. Márquez col.” (17, CC-UAEH). Same data, except: “NTP-80 (calamar) #10” (29, CC-UAEH). Same data, except: “NTP-80 (calamar) #4” (2, CC-UAEH). Same data, except: “NTP-80 (calamar) #6” (11, CC-UAEH). Same data, except: “NTP-80 (calamar) #7” (24, CC-UAEH). Same data, except: “NTP-80 (calamar) #3” (8, CC- UAEH). Same data, except: “NTP-80 (calamar) #1” (68, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #1” (3, CC-UAEH). “Xalapa, La Herradura, cafetal y bosque tropical, 1292 m, N19°31’07”, W96°53’45”, NTP-80 (calamar), 26-XII-1998 a 26-I-1999, Q. Santiago col.” (4, MAAS). “Xalapa, Macuiltepetl, Arca Verde, N19°32´17.72”, W96°55´14.52”, 1535 m, pitfall, 2-VII-2013, V. Cruz, F. Sandoval y E. Montes de Oca cols.” (1, QJSJ). Same data, except: “ N19°32´49.2”, W96°55´12.12”, 1538 m, NTP-80” (1, QJSJ). Same data, except: “ N19°33´7.98”, W96°56´22.62”, NTP-80” (1, QJSJ). Same data, except: “ N19°33´7.86”, W96°56´23.64”, 1428 m, coprotrampa” (2, QJSJ). Same data, except: “ N19°32´47.46”, W96°55´12.72”, 1532 m, NTP-80” (1, QJSJ). Same data, except: “ N19°32´54.12”, W96°55´10.8”, 1546 m, NTP-80” (2, QJSJ). Guatemala: Unknown state: “ Guatemala / M. Cameron. Bequest. B. M. 1955-147” (m 3, NHM). “ Guatemala, L. Cowcald/ M. Cameron. Bequest. B. M. 1955-147/ Hesperus . Philonthus trochanterinus Shp. ” (m 1, NHM).

Figures

FIGURE 2. Dorsal view of males of: a) Belonuchus alternans (syntype, NHM©), b) B. alternans, c) B. apiciventris, d) B. cariniventris (lectotype), e) B. colon, f) B. confusus (holotype), g) B. julietitae (paratype), h) B. linuzensis (paratype). Scale bars = 2.5 mm.

FIGURE 14. Dorsal (left) and ventral (right) view of the first four visible abdominal segments of males of: a) Belonuchus singularis, b) B. tepoztecus. Abdominal styli of male of: c) B. alternans, d) B. confusus, e) B. singularis, f) B. magnistylus (right: detail of left stylus in ventral view), g) B. trochanterinus (right: detail of left stylus in ventral view).

FIGURE 15. Male pregenital sternite of: a) Belonuchus alternans, b) B. oxyporinus, c) B. tepoztecus, d) B. jalappensis, e) B. iridescens, f) B. platypterus, g) B. cyanipennis, h) B. apiciventris, i) B. linuzensis, j) B. metafemoralis, k) B. rufisternus, l) B. tenuistylus, m) B. bidens, n) B. pollens, o) B. longitarsus.

FIGURE 18. Male genital sternite of: a) Belonuchus altenans, b) B. apiciventris, c) B. cariniventris, d) B. colon, e) B. confusus, f) B. julietitae, g) B. magnistylus, h) B. linuzensis, i) B. metafemoralis, j) B. oxyporimimus, k) B. oxyporinus, l) B. pulcher, m) B. rufisternus, n) B. singularis, o) B. tenuistylus. Scale bars = 0.5 mm.

FIGURE 21. Ventral (left) and lateral (right) view of the aedeagus of: a) Belonuchus alternans, b) B. apiciventris, c) B. cariniventris, d) B. colon, e) B. confusus, f) B. julietitae, g) B. linuzensis, h) B. magnistylus, i) B. metafemoralis. Scale bar = 0.5 mm (all figures).

FIGURE 26. Geographical distribution in Mexico and Guatemala of: a) Belonuchus alternans, B. apiciventris, B. magnistylus; b) B. cariniventris, B. confusus, B. julietitae, B. rufisternus, B. singularis and B. zunilensis.