Belonuchus rufipennis ( Fabricius, 1801 )

Márquez, Juan & Asiain, Julieta, 2022, Taxonomy of the Mexican species of Belonuchus Nordmann (Coleoptera: Staphylinidae), Zootaxa 5152 (1), pp. 1-129 : 59-62

Belonuchus rufipennis ( Fabricius, 1801)

Figs. 5d , 13d , 16j , 19i , 23f , 31b

Staphylinus rufipennis Fabricius, 1801: 596 . Staphylinus formosus Gravenhorst, 1806: 72 . Cafius apicalis Dejean, 1833: 63 . Belonuchus pallipes Melsheimer, 1844: 35 .

Total body length in males 8.82 mm (range 7.8–9.7 mm), in females 8.5 mm (range 6.7–9.5 mm). Black on head, pronotum, scutellum, posterior 2/3 or 1/2 of fifth visible abdominal segment to sixth visible segment. Reddish on mouth parts, first two and apical antennomeres, elytra, sterna, legs, first four visible abdominal segments and anterior third or half of fifth.

Head: transverse; ratio length/width in males 0.81 (range 0.75–0.87), in females 0.88 (range 0.85–0.92); convex in posterior half of dorsal surface and at posterior corners. Dorsal surface with sulcate longitudinal midline visible in anterior half of head; front moderately foveate at space between antennal insertions. Eyes large, more than 0.5 times the cephalic lateral length, slightly protruding laterally. Antennomeres 4–5 elongate, 6–10 transverse. Males with mandibles almost as long as head (ratio 1.06, range 0.91–1.18), females with mandibles shorter than head (ratio 0.80, range 0.73–0.85); each with two moderately separated teeth, basal tooth bigger than middle tooth; mandibular channel moderately developed, external margin weakly carinate and internal margin with carinate line only in basal third of its length, extending forward like impressed line beyond level of middle tooth. Apical palpomere of maxillary and labial palpi nearly 1.5 times longer than preapical palpomere. Head of males slightly wider than pronotum (ratio 1.14, range 1.11–1.20), in females head and pronotum almost equally wide (ratio 1.01, range 0.95–1.15).

Thorax: each dorsal row of pronotum with five punctures; pronotum slightly longer than wide (ratio in males 1.08, range 1.04–1.15, in females 1.14, range 1.08–1.23) and almost as wide at anterior corners as at posterior corners (ratio in males 1.09, range 1.07–1.12, in females 1.06, range 1.00–1.10). Scutellum with punctures denser than on elytra, where are somewhat sparse. Prosternum with anterior area slightly elevated transversally. Intercoxal process of mesoventrite shield-shaped; transverse discal ridge well developed, broadly V-shaped, not meeting margin of intercoxal process laterally. Profemur of males slightly widened toward apex, visible only when compared with that of females; external row of spines initiating after basal third and ending before apex, last three or four spines larger than previous spines and spines at internal side; internal row of spines initiating near half length of femur and reaching apex, small darker spines very numerous and close to each other. Metafemur of males with two rows of spines well developed at ventral surface, only near base of femur with some non-aligned spines, both rows can extend almost until apex or 2/3 of its length; spines as long and dark as spines of posterior tibiae ( Fig. 13d ). Tarsomeres dorsally flattened, although little evident because they are small.

Abdomen: first three visible tergites with posterior basal transverse carina well developed; adjacent area slightly depressed; anterior half of each tergite (including depression) with dense wide-superficial punctures that decrease in density toward posterior half, these punctures combined with moderately dense fine punctures; fourth tergite with similar punctures as previous tergites; tergites 5–6 only with fine punctures, although on visible segments 4–6 (particularly 5) with wide and dark setae that are sparse in other segments; visible sternites 1–4 with punctures as in tergites 1–3, but wide-superficial punctures denser. Male pregenital sternite moderately emarginate at posterior margin ( Fig. 16j ). Male genital sternite very elongate (3.19 times longer than wide), moderately symmetrical, anterior portion occupying 33% and posterior portion 67% of its length, apical emargination deep ( Fig. 19i ). Abdominal styli slender.

Aedeagus: length 1.1 mm; apical half of elongate shape and basal half of oval shape; apex subacute; basal half evidently wider than apical half; internal sac visible ( Fig. 23f ).

Variability. In addition to the variation included in the description related to measurements and ratios, it was observed that in specimens of the central and northern parts of the country, antennomeres 1–2 and apical segment are almost as dark as remaining antennomeres, while in specimens from other states they are noticeably paler. Punctures in each dorsal row of prontoum vary from 5:6, 6:5 to 6:6. Scutellum of some specimens, presumably tenerals, is almost reddish or brown instead of black. The long and dark setae of visible abdominal segments 4–6 may be lacking to variable extent because they are often rubbed off. Posterior margin of male pregenital sternite varies in the depth of the emargination, from very weak to moderate.

Taxonomic comments. This species is similar to B. mixtecus and B. pollens , and their main differences have been indicated in the taxonomic comments of these species. The type material of this species was not studied, but we used the detailed taxonomic information given by Smetana (1995), so we had no doubts about the identification of this species.

Material examined (151 males, 136 females): Mexico: Chiapas: “ Cacaohatán, Unión Roja, La Boquilla, 550 m, NTP-80, 6-VIII-84, A. López col.” (19, IEXA). “carretera Tuxtla Gutiérrez- San Cristóbal de Las Casas , km 30.7, selva mediana perturbada, 961m , N16°45’, W92°57’, NTP-80 calamar, 20-XI a 3-XII-1999, J. Márquez col.” (3, MAAS). “La Victoria, ex carrion trap, 17-december-1982, M. A. Morón col.” (6, IEXA). “Tapachula, Ejido Raimundo Enríquez, 70 m, N14°51’52.18”, W92°19’56.66”. Toma 6/ Rata 2 (lluvia). 3-VII-2013, Y. Recinos y E. Chamé cols.” (1, CC-UAEH). Same data, except: “Toma 16/ Rata 1 (lluvias). 6-VII-2013 ” (1, CC-UAEH). Same data, except: “Toma 26/ Rata 2 (lluvias). 28-II-2013 ” (1, CC-UAEH). Same data, except: “Toma 9/ Rata 2 (lluvias). 4-VII-2013 ” (1, CC-UAEH) . Guerrero: “ Acahuizotla, Barranca Las Juntas , 900 m, trampa de intercepción de vuelo, 27 a 29-VIII-2003, Q. Santiago y L. Delgado cols.” (1, IEXA). Hidalgo: “ Atlapexco, rio Atlapexco , cultivos, 168 m , N21°00’2.6”, W98°20’52.2”, en tronco, 9-XII-2005, J. Márquez, F, Ramírez y J. Asiain cols.” (1, CC- UAEH). “ Chapulhuacán, Arroyo Blanco , salva alta, 239 m , N21°09.286’, W99°00.331’, NTP-80 #2, 30-III a 06-IV- 2009, J. Márquez, M. Rivero, M. Torres, M. Vargas y J. Sánchez cols.” (3, CC-UAEH). “ Chapulhuacán, Arroyo Blanco , salva alta, 276 m , N21°09.299’, W99° 0.317’, NTP-80 #1, 30-III a 06-IV-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 a 4- VI-2007, J. Asiain y J. Márquez cols.” (4, CC-UAEH). Same data, except: “NTP-80 (calamar), 14 a 17-VII-2007, J. Asiain, A.P. Martínez, I. A. Rodríguez y J. Márquez cols.” (1, CC-UAEH). “ Chapulhuacán, Puerto del Zopilote , bosque mesófilo de montaña, 1134 m, N21°09.9’ 7.9” , W98°55.8’ 04.0”, NTP-80 (calamar) #10, 28-VIII a 11-IX- 2011, J. Márquez col.” (1, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #3” (1, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #2” (1, CC-UAEH). “Huautla, Acatepec, selva baja, 315 m, N20°56’35.91”, W98.18° 16’15.85”, en mango, 9-VI-2006, J. Márquez y J. Asiain cols.” (1, CC-UAEH). “ Huautla , 2 km NW de Tohuaco Amatzintla , N21°07´23”, W98°16´51”, bosque tropical subcaducifolio, trampa cebada con champiñones, 17 a 28-IV-2015, R. Ramírez y J. Márquez cols.” (25, CC-UAEH). Same data, except: “trampa de intercepción de vuelo #3” (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.” (10, CC-UAEH). “ La Misión , Los Naranjos , Palo Hueco , BMM, 1276 m , N21°08’06”, W99°05’02”, NTP-80 (calamar) #8, 10 al 24-III- 2011, J. Márquez col.” (5, CC-UAEH). “ La Misión, Lomas del Pericón , bosque mesófilo de montaña, 1377 m , N21°6.0’ 46.0” W99°6.0’ 15.6”, NTP-80 (calamar) # 8, 26-VIII al 9-IX-2011, J. Márquez col.” (1, CC-UAEH). “La Misión, Lomas del Pericón, bosque mesófilo de montaña perturbado, 1377 m, N21°60’46.0”, W99°6.0’ 15.6”, NTP- 80 (calamar), 8 al 16-III-2008, J. Márquez y J. Asiain col.” (3, CC-UAEH). “La Misión, bosque mesófilo de montaña, 1334 m, N21°06’ 202”, W99°06’291”, NTP-80 (calamar) #2, 29-III al 05-IV-2009, J, Márquez, M. Rivera, M. Torres, M. Vargas y J. Sánchez cols.” (1, CC-UAEH). “ La Misión, Los Naranjos , Palo Hueco , bosque mesófilo de montaña, 1276 m , N21°08’06”, W99°05’02”, NTP-80 (calamar), 10 al 24-III-2011, J. Márquez col.” (19, 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.” (5, CC-UAEH). Same data except: “en NTP-80 (calamar)” (5, CC-UAEH). “ Pacula, camino Jacala-Pacula, 15.3 km hacia Pacula , selva baja caducifolia, 808 m , N21°02’20.3”, W99°16’32”, en trampa de intercepción de vuelo, 7 al 17-VI-2008, J. Márquez, I. Garrido y M. García cols.” (2, CC-UAEH). Same data, except: “NTP-80 (calamar), 7 a 17-VI-2008, J. Márquez col.” (12, CC- UAEH). “ Pacula, camino al Membrillar del Fraile, río Moctezuma , N99°23´28”, W 21°5´26.8”, 903 m, vegetación riparia, NTP-80 (calamar), 19-I a 25-II-2010, J. Islas col.” (7, 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 (calamar), 12-VIII-2010, C. Cornejo col.” (2, CC-UAEH). “ Tepehuacán de Guerrero, Chilijapa , bosque mesófilo de montaña, 1401 m, N21°01.2’ 5.1” , W98°51.8’ 2.1”, trampa de intercepción de vuelo #2, 11 al 25-III-2011, J. Márquez col.” (1, 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) #5, 16 a 30-IV-2011, J. Márquez col.” (2, CC-UAEH). Same data, except: “NTP-80 (calamar) #2” (1, CC-UAEH) . Jalisco: “ Casimiro Castillo, selva alta perenifolia, en excremento vacuno, 25-IX-1995.” (1, MAAS). “ La Huerta, carretera Barra de Navidad, Puerto Vallarta , Estación Biológica de Chamela , selva baja caducifolia, 77 m , N19°27’2.2”, W105°1’33”, en excremento, 10-XIII-2005, S. Bautista col.” (1, CC-UAEH). “ La Huerta, Estación Biológica de Chamela , selva baja caducifolia, 77 m , N19°27’2.2”, W105°1’33”, en excremento, 11 al 12-VII- 2006, S. Bautista y S. Pulido cols.” (8, CC-UAEH). “ Melaque, Coastecomate , selva baja caducifolia, en necrotrampa temporal (calamar), 23-IX-1995, J. L. Navarrete y J. Márquez cols.” (1, MAAS). Morelos: “ Ciudad Ayala, Unidad 10 de abril, 1360 m, trampa de intercepción de vuelo, 26 a 31-VII-2003, A. y L. Delgado cols.” (10, IEXA). “ Mexicapa, bosque de pino, NTP-80 calamar, zona 3, 27-XI-1993, A. Soria, C. Cabello y E. Córdoba cols.” (1, MAAS). “ Tlayacapan, San José de los Laureles, NTP-80, localidad 3, bosque mesófilo de montaña perturbado, V- 1996, J. Márquez col.” (2, MAAS). Same data, except: “localidad 1, bosque de pino-encino, IX-1995, J. Márquez col.” (3, MAAS). Same data, except: “ IV-1996, K. Villavicencio y J. Márquez cols.” (1, MAAS). “ Tlayacapan, San José de los Laureles , camino a Amatlán , bosque de pino-encino, 1874 m , N18°58.5’, W99°0.5’, NTP-80 (calamar), IX-1995, J. Márquez col.” (1, MAAS). “ Tlayacapan, en detritos de A. mexicana , 25-I-1999, J. Márquez col.” (1, MAAS). “ Tlayacapan, Santa Catarina , selva baja caducifolia y huertos, en hojarasca, 16-VIII-1998, J. Márquez y R. Toledo cols.” (1, MAAS). Puebla: “ Tlacuilotepec, cerca del río San Marcos , N20°19’22.5”, W98°02’35.4”, bosque tropical subcaducifolio, trampa con fruta, 13 a 25-X-2018, J. Márquez y J.D. Silva cols.” (1, CC-UAEH). Same data, except: “NTP-80, 26-VI a 17-VII-2018, J. D. Silva-Hurtado col.” (1, CC-UAEH). “ Xicotepec de Juárez, Hidroeléctrica Plata, selva mediana, 457 m, en plantas, 5-X-2002, en plátano podrido, J. Asiain y J. Márquez cols.” (1, MAAS). “ Xicotepec de Juárez, Hidroeléctrica Patla , bosque mesófilo de montaña, en necrotrampa temporal (calamar), 27-III-1998, O. Pérez col.” (1, MAAS). “km 4 carretera Tepexco - Izúcar de Matamoros , N18°39´22”, W98°39´53.2”, 1350 m, trampa de intercepción de vuelo # 1, 18-VII-2015, A. y L. Delgado cols.” (11, IEXA). Querétaro: “ Ayutla, Arroyo Seco, cerca de las cabañas Las Adjuntas , N21°24’11.1”, W99°34’38.9”, bosque tropical caducifolio, trampa de intercepción de vuelo, 15 a 17-VI-2016, J. Márquez col.” (1, CC-UAEH). Quintana Roo: “ A.N.P. Yum Balam, 2014, N21.36024°, W87.410656°, 169 mlnm, E. Chamé col. “ (1, CC-UAEH). “A.N.P. Yum Balam , sitio 1, necrotrampa, 04-09-2016 ” (1, CC-UAEH). Tamaulipas: “carretera antigua Ciudad Victoria, Huamave, camino a La Mina Virgen, selva baja caducifolia, 440 m, N23°42’9.4” W99°12’57.6”, en excremento vacuno, 2-XI- 2004, J. Márquez col.” (1, CC-UAEH). Veracruz: “ Catemaco, Pipiapan, Parque de la Flora y Fauna Silvestre Tropical, 600 m, necrotrampa, 28-X-1999, A. Díaz col.” (1, IEXA). “ Córdoba, San Rafael Calería, Ojo de Agua , N18°57’15”, W96°56’24”, selva mediana subcaducifolia, 1127 m, NTP-80 (calamar), 20-III a 17-IV-1999, Q. J. Santiago, J. Márquez y J. Asiain cols.” (1, MAAS). “ Huatusco, Coxcontla, bosque mesófilo de montaña perturbado, 1250 msnm, NTP-80 (calamar) #746, 3-VI-1995, M. A. Pensado y Q. Santiago cols.” (1, MAAS). Same data, except: “NTP-80 (calamar) #202, 22-X a 19-XI-1994 ” (1, MAAS). “ Ilamatlán , Xococapa , bosque tropical subperenifolio, 397 m , N20°48’ 24.0”, W98°20’24.6”, NTP-80 (calamar) # 4, 5-III a 1-IV-2006, F. Ramírez col.” (4, CC- UAEH). “ Totutla, Mata Oscura , 1 km S de Zacuapan , selva mediana subcaducifolia, 921 m , N19°12’23”, W96°50’32”, NTP-80 (calamar), 20-II al 19-III-1999, Q. Santiago y J. Márquez cols.” (6, MAAS). “ Totutla, El Encinal, bosque de encino perturbado, 750 msnm, NTP-80 (calamar) #683, 6-V al 3-VI-1995, M. A. Pensado y Q. Santiago cols.” (1, MAAS). “ Xalapa, San Antonio Paso del Toro , trampa de intercepción de vuelo #1, 4-IV-2015, L. Delgado col.” (8, IEXA). Same data, except: “ 2 a 30-V-2015, K. Cortes y Q. Santiago cols.” (15, IEXA). Yucatán: “ Conkal, NTP-80, 26-II a 26-III-2002, L. Peraza col.” (33, IEXA). “ Tzucacab, Tigre Grande , NTP-80, 20-VIII a 17-IX-2002, L. Peraza col. (5, IEXA). Same data, except: “ 14-I a 11-II-2002 ” (15, IEXA) .

Group 3: xanthomelas

Included species: B. basiventris ( Sharp, 1885) , B. connexus ( Say, 1830) , B. longitarsus sp. nov., B. rufiventris ( Sharp, 1887) , and B. xanthomelas Solsky, 1868 .

Diagnosis of the xanthomelas group: head without temporal carina; males with mandibles expanded at base ( Fig. 10c ), mandibles as long as head or slightly longer ( Fig. 10c ); females with mandibles shorter than head ( Fig. 6c ); pronotum with five or less punctures in each dorsal row; elytra reddish; trochanters and femora of posterior legs without modifications in both sexes; tarsomeres somewhat flattened dorsally; abdomen reddish with black or completely black; aedeagus with reduced paramere ( Figs. 23g –i ).

Key to Mexican species of Belonuchus of the xanthomelas group

1 Elytra and, usually, legs reddish, remaining body black ( Figs. 5g, h ); aedeagus 1.1 mm long, cone shaped, apex very acute, basal half moderately wider than apical half ( Fig. 23h )................................................... B. connexus

- Elytra, legs and entire abdomen or some part of abdomen reddish, remaining body black; aedeagus not exactly as above... 2

2(1) Abdomen with some segments reddish and others black ( Figs. 5f , 6c, d, e )........................................ 3

- Abdomen entirely reddish or reddish-brown ( Figs. 6a, b )...................................................... 4

3(2) First two visible abdominal segments irregularly black, with minor portions reddish ( Fig. 5f ); male pregenital sternite with moderately deep, broadly U- or V-shaped emargination at posterior margin ( Fig. 16k ); aedeagus as in Fig. 23g. B . basiventris

- First two visible abdominal segments as reddish as following two ( Figs. 6c, d, e ); male pregenital sternite with very broad and shallow emargination at posterior margin ( Fig. 16m ); aedeagus as in Fig. 24b........................... B . xanthomelas

4(2) Pronotum with two punctures in each dorsal row ( Fig. 6b ); eyes slightly longer than 0.5 times the cephalic lateral length; coxae black or dark-brown; third visible abdominal tergite with posterior basal transverse carina well developed; male pregenital sternite deeply emarginate at posterior margin (Fig. 17d); aedeagus as in Fig. 24a......................... B . rufiventris

- Pronotum with five punctures in each dorsal row ( Fig. 6a ); eyes small, 0.38 times the cephalic lateral length; coxae reddish; third visible abdominal tergite with posterior basal transverse carina incomplete, only clearly visible at sides; male pregenital sternite moderately emarginate at posterior margin ( Fig. 15–o ); aedeagus as in Fig. 23i.................... B . longitarsus

Figures

FIGURE 5. Dorsal view of males (except figure g) of: a) B. mixtecus (holotype), b) B. pollens (syntype, NHM©), c) B. pollens, d) B. rufipennis, e) B. basiventris (syntype, NHM©), f) B. basiventris, g) B. connexus (female holotype of B. xanthopus, synonym of B. connexus, shared by A. Kovalev©), h) B. connexus. Scale bars = 2.5 mm.

FIGURE 6. Dorsal view of males (except figure c) of: a) Belonuchus longitarsus (holotype), b) B. rufiventris, c) B. iteratus (male lectotype and female paralectotype, NHM©), d) B. xanthomelas (NHM©, Biologia Centrali-Americana), e) B. xanthomelas, f) B. ephippiatus, g) B. moroni (paratype). Scale bars = 2.5 mm.

FIGURE 10. Dorsal view of the head of males of: a) Belonuchus julietitae, b) B. singularis, c) B. connexus, d) B. platypterus, e) B. bidens, f) B. nigerrimus, g) B. optatus, h) B. cyanipennis, i) B. godmani. Dorsal view of the head and pronotum of the holotype female of: j) B. optatus.

FIGURE 13. Ventral view of the left posterior leg of males of: a) B. trochanterinus, b) B. tenuistylus, c) B. zunilensis, d) B. rufipennis, e) B. mixtecus, f) B. pollens, g) B. ephippiatus, h) B. aenigmaticus, i) B. iridescens, j) B. viridipennis. Tarsomeres of posterior leg of males of: k) B. colon, l) B. ephippiatus. Yellow ovals: median longitudinal groove.

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 16. Male pregenital sternite of: a) Belonuchus moroni, b) B. dichrous, c) B. flavipennis, d) B. aenigmaticus, e) B. nigerrimus, f) B. juanmorronei, g) B. godmani, h) B. magnistylus, i) B. trochanterinus, j) B. rufipennis, k) B. basiventris, l) B. connexus, m) B. xanthomelas, n) B. ephippiatus, o) B. optatus.

FIGURE 19. Male genital sternite of: a) Belonuchus tepoztecus, b) B. trochanterinus, c) B. zunilensis, d) B. bidens, e) B. erichsoni, f) B. jalappensis, g) B. mixtecus, h) B. pollens, i) B. rufipennis, j) B. basiventris, k) B. connexus, l) B. longitarsus, m) B. rufiventris, n) B. xanthomelas, o) B. ephippiatus, p) B. moroni. Scale bars = 0.5 mm.

FIGURE 23. Ventral (left) and lateral (right) view of the aedeagus of: a) Belonuchus bidens, b) B. erichsoni, c) B. jalappensis, d) B. mixtecus, e) B. pollens, f) B. rufipennis, g) B. basiventris, h) B. connexus (at center, detail of the apex of the median lobe), i) B. longitarsus (at center, detail of the apex of the median lobe). Scale bar = 0.5 mm (all figures).

FIGURE 24. Ventral (left) and lateral (right) view (except e) of the aedeagus of: a) Belonuchus rufiventris, b) B. xanthomelas (at center, detail of the apex of the median lobe), c) B. ephippiatus, d) B. moroni, e) B. cifuentesi (left: detail of paramere, right: ventral view; from Rodríguez and Navarrete-Heredia, 2016), f) B. dichrous, g) B. flavipennis, h) B. aenigmaticus, i) B. iridescens, j) B. nigerrimus, k) B. optatus, l) B. platypterus. Scale bar = 0.5 mm (all figures, except e).

FIGURE 31. Geographical distribution in Mexico and Guatemala of: a) Belonuchus pollens; b) B. rufipennis.