The genus PilobaliaBurmeister 1875 includes forty six species and four subspecies distributed in Andean environments of Peru, Bolivia, Chile and Argentina (Flores, 1997). This is the third most diverse genus of the Nycteliini tribe, a set of twelve genera and around 300 species endemic to the arid and semi-arid regions of South America (Flores, 1997; Flores, 1999; Flores & Vidal, 2000; Flores & Triplehorn, 2002).

The first species of the genus Pilobalia were described under various other genera by Perty (1830), Erichson (1834, 1847), Blanchard (1843), Waterhouse (1844) and Kirsch (1886). Subsequently, the genus Pilobalia was proposed by Burmeister (1875) providing a morphological diagnosis and listing the species known at that date. Fairmaire (1876; 1878; 1902; 1903) added five species and Gebien (1936) designated Pilobalia decorata (Erichson, 1834) as type species of the genus. In his revision of the Nycteliini tribe, Kulzer (1954) reviewed the genus, described twenty species, assigned subspecies rank to Pilobalia immaculata (Blanchard, 1843) and presented a key for all species known until then. Subsequent contributions of Kulzer (1955, 1958, 1959) added seven species and assigned subspecies rank to Pilobalia murinaKulzer, 1954. Later, Molinari (1968) and Peña (1973) described four and five species respectively, all of them from Argentina. More recently, Ferru & Pizarro-Araya (2007) offered the first Chilean records of Pilobalia torresiMolinari, 1968 and Pilobalia escobariPeña, 1973 based on specimens collected in the Altiplano of Tarapaca Region, including morphological redescriptions and environmental characterization of collection sites.

Within the tenebrionid fauna of Peru, the genus Pilobalia is currently represented by 25 species and one subspecies, with a high rate of endemism (about 96%) and with an imprecise number of undescribed species observed in collecting trips and entomological collections (Smith et al., 2015; Giraldo & Flores, 2016). Most of its species are distributed in Andean environments, including western Andean ranges (1000 m – 3800 m), inter-Andean valleys (1500m – 3300 m) and high plateaus or “Puna” (3800 m – 5000 m) (Giraldo & Flores, 2016) (Fig. 1a). Available published records account for its presence in particular habitats such as “lomas” vegetation in coastal desert (Aguilar, 1976), quinoa and potato crops (Escalante et al., 1981; Rasmussen et al., 2003) and high Andean Polylepis forests (Rossi et al., 2018).

The purpose of the present work is to update the knowledge of the genus Pilobalia, providing an updated checklist of Pilobalia species and new distribution records in Peruvian territory.

This study is based on direct and indirect examination of type and non-type specimens housed in the following collections: Natural History Museum Basel (NHMB, Christoph Germann), California Academy of Sciences, San Francisco, California, USA (CASC, Rachel Diaz-Bastin, Chris Grinter), Instituto Argentino de Investigaciones de Zonas Áridas, Mendoza, Argentina (IADIZA, Sergio Roig-Juñent, Gustavo Flores), Laboratorio de Entomología, Universidad Nacional Mayor de San Marcos, Lima, Perú (LEUSM, Eliana Quispitupac), Museo de Entomología Klaus Raven Büller, Universidad Nacional Agraria La Molina, Lima, Perú (MEKRB, Clorinda Vergara), Museo de La Plata, Universidad Nacional de La Plata, La Plata, Argentina (MLP, Nora Cabrera), Muséum National d'Histoire Naturelle, Paris, France (MNHN, Antoine Mantilleri, Christophe Rivier), Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Perú (MUSM, Mabel Alvarado), Laboratorio de Sanidad Vegetal, Servicio Nacional de Sanidad Agraria, Lima, Perú (SENASA, Graciano Tejada).

The primary source of distribution records were about 580 specimens examined in Peruvian entomological collections, namely LEUSM, MEKRB, MUSM and SENASA. Direct examination of type specimens of eight Argentinean and two Peruvian species was carried out in IADIZA, through temporary loans from MLP (Cabrera et al., 2010). Indirect examination of type specimens housed at CASC, MNHN and NHMB was carried out through images sent by their respective curators, including two Peruvian and two Bolivian (CASC), four Peruvian (MNHN) and four Peruvian species (NHMB). Also specimens from Argentina and Bolivia were examined for comparative purposes at IADIZA. When type specimens were not accessible, dichotomic key, original descriptions and image plates presented by Kulzer (1954, 1955, 1958, 1959) were useful for species determination.

The checklist includes distribution status (endemic or new record), synonymy, type specimens examined (directly or indirectly), type localities (incorrectly spelled in quotation marks, doubtful with a question mark) and material examined, as available for each species. The distribution records of specimens that were not assigned to described species due to their poor condition, doubtful resemblance to known species, or preliminary assignment to undescribed species were summarized in a table indicating regions and provinces according to Peruvian territorial jurisdictions, number of specimens found for each of them, as well as geographic coordinates and altitude data. Overall, type and non-type localities were employed to plot distribution maps for the genus in Peru. The frequency of distribution records according to its latitude and altitude were plotted with histograms and their patterns were interpreted taking into account habitat selection of genus and collecting effort performed in Peruvian territory.

Distribution maps were elaborated using SimpleMappr (Shorthouse, 2010). Histogram plots for latitude and altitude data were elaborated with PAST software (Hammer et al., 2001).

Checklist of Pilobalia species recorded in Peru

Pilobalia alboscripta Fairmaire, 1903 ENDEMIC (Fig. 1b)

Type specimen examined: Syntype (MNHN).

Type locality: La Libertad (Otuzco, Chuquizongo “Choquisongo” 2100 m).

Material examined: La Libertad, Otuzco, Sinsicap, 07°42'47.59"S 78°44'24.81"W, 2380 m, 12-III-2022, E. Gamboa, 1 (MUSM).

Pilobalia baeriFairmaire, 1902 ENDEMIC

Type specimen examined: Syntype (MNHN).

Type locality: La Libertad (Santiago de Chuco 3000 m).

Pilobalia beckiKulzer, 1954 ENDEMIC (Fig. 2a)

Type specimens examined: Holotype (CASC), paratype (MLP).

Type locality: Junin (Junin Lake).

Pilobalia blancasiKulzer, 1958 ENDEMIC (Figs. 1b, 2b)

Type specimen examined: Holotype (NHMB).

Type localities: Lima (Atocongo, Quilmaná “Quilmara”), San Martin (Rioja).

Material examined: Lima, Cañete, Quilmaná, 300 m, 11-XI-1951, Jordan, 1 (MUSM); Lima, 15-IV-1958, F. Blancas, 2 (MUSM), lomas de Atocongo, 04-IX-1948, F. Blancas, 2 (MUSM), 30-X-1958, P. Aguilar, 2 (MEKRB), 12-X-1969, R. García, 1 (MUSM). San Martín, Rioja, 4 (MUSM). PERU (no data), 2 (MUSM).

Pilobalia crassicornis Kulzer, 1954 ENDEMIC (Fig. 1b)

Type localities: Junin (Tarma, Acobamba 2900 m and Carpapata “Carpapaca” 2300 m). Material examined: Junín, Chupaca, Comunidad Ñahuinpuquio, 3200m., 07-XII-2003, Rivera & Montgomery, 1 (MEKRB); Huancayo, Pucará, 3310 m., 30-IX-2005, S. Mayta, 1 (MEKRB); Jauja, 23-II-1998, J. Paucar, 1 (MEKRB).

Pilobalia crassicostaKulzer, 1954 ENDEMIC (Fig. 1c)

Type localities: Arequipa, Junin (Huacapistana “Huacapistrana”).

Material examined: Ayacucho, 08-IV-2004, A. Martínez, 1 (MEKRB). Junín, 20-XII-1960, Vargas, 1 (SENASA).

Pilobalia decorata decorata (Erichson, 1834) (Fig. 1b)

= Pilobalia nebulosa (Waterhouse, 1844)

Type localities: Southern Peru, Tacora and Titicaca plateaus.

Material examined: Arequipa, 10-X-1996, O. Marín, 2 (MEKRB), 06-VIII-1999, J. Huanca, 1 (MEKRB), Castilla, Orcopampa, 15°17'29.90"S 72°19'17"W, 3855 m, 10-II-2018, P. Sanchez, 1 (MUSM). Cusco, Canas, Langui, 14°27'32.39"S 71°17'34.8"W, 4296 m, 28-II-2022, A. Giraldo, 3 (MEKRB); Canchis, Sicuani, 11-II-2002, I. Pardo, 1 (MEKRB); Chumbivilcas, Chilloroya, pajonal, 4020 m, 14°30'09.63"S 71°46'52.51"W, VI-2012, J. Jara, 10 (MEKRB), Livitaca, 14°29'15.46"S 72°05'0.07"W, 3841 m, 21-III-2021, E. Quispe, 3 (MUSM), 14°32'46.90"S 72°04'05.87"W, 3841 m, 21-III-2021, E. Quispe, 1 (MUSM), Urasana, 4029 m., V-2012, J. Jara, 1 (MEKRB); Cusco, 08-X-1961, Molledo, 2 (SENASA), 06-I-1962, Molledo, 5 (SENASA), 05-VI-1974, C. Martinez, 1 (SENASA), 13-III-1971, P. Aguilar, 2 (MEKRB), 20-IV-1971, M. Buitrón, 2 (MEKRB), 10-VIII-1995, C. Marhuay, 1 (MEKRB), 04-IV-1998, N. Elguera, 2 (MEKRB), 03-VI-2001, M. Cárdenas, 1 (MEKRB), Sacsayhuamán, 3585 m, 27-IV-1977, R. García, 1 (MUSM), V-1979, N. Castro, 1 (MUSM); Paucartambo, Huancarani, 3500 m, 03-III-1975, G. Lamas, 1 (MUSM); Quispicanchi, Huaro, 13°45'33.3"S 71°39'59.6"W, 4246 m, 12-I-2021, M. Gill, 4 (MUSM); Urubamba, Chincheros, 08-VII-1996, W. Catalán, 2 (MEKRB). Puno, Azángaro, Arapa, II-1948, F. Blancas, 8 (MUSM), Azángaro, II-1948, F. Blancas, 1 (MUSM), San Antón, 14°34'31.40"S 70°19'01.27"W, 4062 m, 20-II-2022, J. Aragon, 12 (MUSM); Chucuito, 06-II-2013, J. Huanca, 11 (MEKRB), Huacullani, Comunidad Cóndor de Aconcahua, 16°39'41.72"S 69°21'52.03"W, 4183 m, 01-III-2010, E. Rázuri, 1 (MUSM), Comunidad de Cotoco, 16°41'23.20"S 69°18'18.69"W, 4070m., 05-III-2010, E. Rázuri, 1 (MUSM), Comunidad de Ingenio, 16°39'41.72"S 69°19'01.41"W, 4226m., 02-III-2010, E. Rázuri, 1 (MUSM), Comunidad de Yorohoco, 16°34'05.64"S 69°19'34.19"W, 3856 m, 06-III-2010, E. Rázuri, 1 (MUSM), Juli, 16°16'08.39"S 69°30'54"W, 4060 m, 23-II-2022, A. Giraldo, 1 (MEKRB); El Collao, Ilave, 3900 m, II-1948, Weyrauch, 11 (MUSM); Huancane, Miajachi, 28-II-1948, F. Blancas, 6 (MUSM); Lampa, 0.5 Km NE of Paratia, 15°27'07"S 70°35'38"W, II-2020, G. Sarabia, 3 (MEKRB); Puno, 1 (no data) (MUSM), 1940, Soukup, 1 (SENASA), 3900 m, II-1948, F. Blancas, 5 (MUSM), 15-IV-1948, J.E. Wille, 2 (SENASA), 12-II-1952, F. Blancas, 1 (MUSM), 23-II-1968, M. Delgado, 4 (SENASA), 3850 m, 27-IV-1974, R. García, 6 (MUSM), 31-VIII-1981, R.Ortiz, 3 (MEKRB), 26-31-XII-1981, G. Mamani, 3 (MEKRB), 13-I-1982, F. Wong, 4 (MEKRB), 20-I-1983, E. Deza, 4 (MEKRB), II-2020, H. Coaquira, 3 (MEKRB), Hacienda Umayo, III-1948, F. Blancas, 3 (MUSM), Lago Titicaca, 27-XI-1952, F. Blancas, 1 (MUSM), Pichacani, 1.3 km northeast to Ayuncora, 16°04'40.79"S 70°00' 39.59"W, 3978 m, 15-I-2022, A. Giraldo, 2 (MEKRB), Pichacani, 5.8 km northeast to Puente Bello, 16°18'17.99"S 70°13'01.19"W, 4459 m, 24-II-2022, A. Giraldo, 2 (MEKRB), Toramipata, 3825m., 15°48'08"S 69°59'36"W, II-2015, J. Ugarte, 3 (MEKRB); San Román, Juliaca, III-1956, 1 (MUSM); Yunguyo, Cuturapi, 16°16'11.99"S 69°12' 25.19"W, 3869 m, 02-III-2022, A. Giraldo, 1 (MEKRB).

Pilobalia decorata immaculata (Blanchard, 1843) NEW COUNTRY RECORD (Fig. 1c)

= Pilobalia opaca (Kirsch, 1886)

= Pilobalia alticola (Kirsch, 1886)

Type localities: Bolivia (mountain range near to Potosí; Sajama 4500 m).

Material examined: Arequipa, Caylloma, Caylloma, 15°11'38.15"S 71°52'17.56"W, 4900 m, 15-VIII-2021, A. Aliaga, 2 (MUSM), 15-II-2022, E. Medina, 2 (MUSM), Pampa de Toccra, 15°47'31.19"S 71°22'40.8"W, 4325 m, 18-I-2022, A. Giraldo, 2 (MEKRB), San Antonio de Chuca, 15°49'22.8"S 71°13'44.39"W, 4358 m 27-II-2022, A. Giraldo, 1 (MEKRB). Cusco, Chumbivilcas, Chilloroya Alto, Mina Constancia, 14°31’34’’S 71°46’52’’W, 4139 m, 11-VI-2013, L. Huerto, 1 (MUSM), 14°35’21’’S 71°44’13’’W, 4380 m, 11-VI-2013, L. Huerto, 1 (MUSM), Pumaccocha, 4029 m., V-2012, J. Jara, 7 (MEKRB). Puno, Puno, 1944, 2 (MUSM), 3800 m, II-1948, F. Blancas, 2 (MUSM), 3900 m, II-1948, Weyrauch, 1 (MUSM), 23-II-1968, M. Delgado, 11 (SENASA), Isla Amantani, 3900 m., III-1948, F. Blancas, 10 (MUSM); Yunguyo, Cuturapi, 16°16'11.99"S 69°12'25.19"W, 3869 m, 23-II-2022, A. Giraldo, 8 (MEKRB).

Pilobalia decorata titschackiKulzer, 1954 ENDEMIC (Fig. 1d)

Type localities: Apurimac (37 km South of Andahuaylas), Ayacucho (Huacaña 3700 m “Huacana”, Querobamba 3500 m, Tayapampa 4000 m).

Material examined: Apurímac, Cotabambas, 14°07'37.65"S 72°15'34.45"W, 3798m., 22-24-I-2017, J. Suárez, 1 (MUSM), 14°03'43.37"S 72°18'36.89"W, 4326m., 29-30-I-2017, J. Suárez, 1 (MUSM), Chalhuahuacho, C. C. Ccahuapirhua, 14°07'17.53"S 72°21'55.38"W, 4384 m, 23-III-2021, R. Coronel, 1 (MUSM), 14°08'05.93"S 72°20'27.92"W, 4030 m, 23-III-2021, J. Aragon, 1 (MUSM), 14°08'13.02"S 72°19'43.95"W, 3876 m, 24-III-2021, J. Aragon, 1 (MUSM), Chalcobamba, Qda. Itaña, 14°04'08.02"S 72°22'51.19"W, 4419 m, 03-III-2021, J. Aragon, 1 (MUSM), Ferrobamba, 14°05'18.54"S 72°20'51.01"W, 4376 m, J. Aragon, 1 (MUSM), 14°06'26.73"S 72°15'08.01"W, 3994 m, 18-III-2021, R. Coronel, 1 (MUSM), Coyllurqui, Huancuire, 14°01'05.37"S 72°20'36.04"W, 4367 m, 09-IV-2021, R. Coronel, 1 (MUSM), 14°01'24.64"S 72°20'48.85"W, 4442 m, 09-IV-2021, J. Aragon & A. Elme, 1 (MUSM), 14°02'21.48"S 72°23'06.43"W, 4351 m, 12-IV-2021, R. Coronel, 1 (MUSM), Pumamarca (Sallahue), 14°01'46.67"S 72°19'36.28"W, 4349 m, 24-II-01-III-2020, N. Zenteno & A. Silva, 1 (MUSM), Mina Las Bambas, sector Sagrapeña, 14°04'37.24"S 72°18'33"W, 4265 m, 01-III-2020, L. Perez, 2 (MUSM), Tambobamba, C. C. Antuyo, 14°04'32.38"S 72°16'03.88"W, 4244 m, 29-II-01-III-2020, N. Zenteno & A. Silva, 1 (MUSM); Grau, Progreso, Palcca, 14°01'52.88"S 72°26'28.02"W, 4148 m, 12-III-2021, R. Coronel, 2 (MUSM). Cusco, 1962, Molleda, 1 (MEKRB); Chumbivilcas, Urasana, 4029 m, V-2012, J. Jara, 21 (MEKRB), 5 (IADIZA).

Pilobalia dorsoplicata Fairmaire, 1876 ENDEMIC

Type locality: Peru (without further details).

Note: After examining the only type specimen housed at British Museum, Kulzer (1954) pointed out that an accurate determination of the species is unreliable, since the elytral pattern is completely lost and the "Peru" locality is too imprecise.

Pilobalia freyiKulzer, 1954 ENDEMIC

Type specimen examined: Holotype (NHMB).

Type locality: Lima? (Pativilca River “Rio Pativilia” 3300-3600 m Huaraz to Huacho route).

Pilobalia gracilipesKulzer, 1954 ENDEMIC

Type locality: Junin (Tarma River, Palca 2800 m).

Pilobalia haagiFairmaire, 1878 ENDEMIC

Type locality: Peru (without further details).

Pilobalia lineataKulzer, 1958 ENDEMIC

Type locality: Peru (without further details).

Pilobalia löffleriKulzer, 1955 ENDEMIC

Type localities: Ancash (Cohuptal? 4600 m in Huaraz zone, dry slope west to Huaraz 4200 m, highland between Quilcayhuanca “Quilquaihuanca” and Shallapa “Schallaptal”).

Pilobalia michelbacheri Kulzer, 1954 ENDEMIC (Figs. 1c, 2c)

Type specimen examined: Holotype (CASC).

Type localities: Lima (Matucana, San Miguel de Viso “Viso, Rio Rimai” 2700 m).

Material examined: Lima, Huarochirí, 28-V-1974, P. Hocking, 1 (MUSM), carretera central km 70, 26-IV-1945, 2 (MUSM), Huanza, 11°37'20.54"S 76°25'48.54"W, 3910 m, 15-VII-2021, M. Rodriguez, 1 (MUSM), 11°37'51.34"S 76°26'53.01"W, 3900 m, 15-III-2021, M. Rodriguez, 2 (MUSM), Matucana, 23-VI-2001, S. Díaz, 1 (MEKRB), San Bartolomé, 04-VI-2003, E. Arango, 1 (MEKRB), San Jerónimo de Surco, V-2014, A. Solano, 2 (MEKRB); Lima, Chosica, 13-IV-2003, W. Soto, 1 (MEKRB).

Pilobalia oberthüriKulzer, 1954 ENDEMIC (Fig. 2d)

Type specimens examined: Holotype, allotype, two paratypes (MNHN).

Type locality: Cajamarca (Cajabamba 4000 m).

Pilobalia oblonga (Blanchard, 1843) NEW COUNTRY RECORD (Fig. 1c)

Type locality: Puna highlands of mountain range (no mention of any country).

Material examined: Moquegua, Mariscal Nieto, Carumas, 16°38'52"S 70°20'48"W, III-2015, M. Deza, 5 (MEKRB), 16°39'41.20"S 70°21'53.43"W, 4371 m, 22-23-II-2021, L. Ramirez, 1 (MUSM), 16°54'16.54"S 70°30'26.00"W, 4659 m, 22-23-II-2021, L. Ramirez, 3 (MUSM). Puno, El Collao, Capaso, Challapalca, 17°13'15.6"S 69°46' 37.2"W, 4305 m, 23-II-2022, A. Giraldo, 3 (MEKRB). Tacna, Candarave, Laguna Suches, 16°56'57.69"S 70°24'09.87"W, 4462 m, pajonal, VIII-2015, Y. Callohuari, 1 (MEKRB); Tarata, near to Ticaco, 17°25'44.39"S 69°57'46.79"W, 4247 m, 23-II-2022, A. Giraldo, 1 (MEKRB).

Pilobalia ornataKulzer, 1958 ENDEMIC

Type locality: Ancash (Laguna Conococha “Cenococha” 4150 m).

Pilobalia philippi Kulzer, 1958 ENDEMIC

Type locality: La Libertad (Otuzco, Chuquizongo “Choquisongo” 2100 m).

Pilobalia ruficollisKulzer, 1954 ENDEMIC (Fig. 1d)

Type locality: La Libertad (Huamachuco).

Material examined: Cajamarca, Cajamarca, 1969, J. Alata, 3 (SENASA), Cumbemayo, 3300 m., 30-VI-2002, M. Diéguez, 6 (MUSM); Hualgayoc, El Tambillo, 1877, A. Raimondi, 3 (MUSM).

Pilobalia soror Kulzer, 1958 ENDEMIC (Fig. 3a)

Type specimen examined: Holotype (NHMB).

Type locality: Peru (without further details).

Pilobalia subnudaKulzer, 1954 ENDEMIC (Fig. 3b)

Type specimen examined: Paratype (MLP)

Type locality: Lima? (Pativilca River “Pativilia” 3300-3600 m Huaraz to Huacho route).

Pilobalia tenella Kulzer, 1954 ENDEMIC (Fig. 3c)

Type specimen examined: Paratype (MNHN).

Type locality: La Libertad (Otuzco, Chuquizongo “Choquisongo” 2100 m).

Pilobalia tristisKulzer, 1954 ENDEMIC

Type localities: Apurimac (Mamara), Cusco, Madre de Dios.

Pilobalia unicolorKulzer, 1954 ENDEMIC (Figs. 1d, 3d)

Type specimen examined: Holotype (NHMB).

Type locality: Cusco (Cajea? 3900 m).

Material examined: Cusco, Paucartambo, 18-IV-1993, L. González, 1 (MEKRB).

Pilobalia voogdiKulzer, 1954 ENDEMIC

Type locality: Huánuco 1900 m.

Pilobalia weyrauchiKulzer, 1954 ENDEMIC (Fig. 1d)

Type locality: Junin (Tarma 3300-3800 m).

Material examined: Junín, Huancayo, 21-IV-1978, J. Alcázar, 9 (MEKRB), 24-XI-2012, C. Huaripata, 1 (MEKRB), Pucará, 30-IX-2005, S. Mayta, 2 (MEKRB); Jauja, 7 Km NE Jauja, 3390 m., 22-XI-2004, N. Lara, 1 (MEKRB).

According to the updated checklist presented here, the genus Pilobalia in Peru is now represented by 26 species and two subspecies. It has been possible to examine type specimens of eleven species, which represents a 39% of the recorded species and subspecies in Peru. In the Peruvian collections, specimens corresponding to 10 species and two subspecies were found, namely P. alboscripta, P. blancasi, P. crassicornis, P. crassicosta, P. decorata decorata, P. decorata immaculata, P. decorata titschacki, P. michelbacheri, P. oblonga, P. ruficollis, P. unicolor and P. weyrauchi. Otherwise, sixteen species are currently known only from their type specimens and are not properly represented in Peruvian collections. Among these species with poorly known distribution, P. baeri, P. becki, P. freyi, P. gracilipes, P. löffleri, P. oberthüri, P. ornata, P. philippi, P. subnuda, P. tenella, P. tristis and P. voogdi have more or less well-defined type localities, so their distributions must be restricted and hitherto overlooked by collecting surveys in Peru. For P. haagi, P. lineata and P. soror, the label data of type specimens only indicates “Peru” without further details, so its precise distribution will only be defined with more collecting effort. The status of P. dorsoplicata is even more uncertain following notes by Kulzer (1954), with a type specimen extensively damaged and bearing label data "Peru", so this species should be declared nomen dubium according to International Code of Zoological Nomenclature.

Distribution records of undetermined specimens of genus Pilobalia are presented in Table I. These records include 275 specimens from 15 regions and 41 provinces of the Peruvian territory. A high number of specimens not reliably assigned to described species is not surprising for a genus of wingless darkling beetles prone to speciation by vicariant orographic barriers and this biological trait partly explains why its systematics studies are still in progress.

Taking into account all the available distribution records, distribution maps were elaborated, including 240 localities (Fig. 1). These records place the northern limit of the Nycteliini tribe to around 6°S, some 7° further north than on the map presented by Flores & Triplehorn (2002). As expected according to previous works (Peña 1973, Giraldo & Flores 2016), records are mostly located along the different altitude levels of the Andes mountain range, with very few to the west (coastal desert) and east (tropical forests) of the Andes. The records in the coastal desert are located between 12–13°S, these correspond specifically to Pilobalia blancasi collected in patches of fog-dependent vegetation called "lomas", as documented in early works by Aguilar (1976) and by the specimens found in the collections. The label data of some specimens suggest that they were collected in tropical areas between 5–13°S, from 150 to 2200 meters of altitude. At first glance, tropical areas do not have suitable habitats for the genus, however the presence of semi-arid patches or dry micro-habitats cannot be ruled out, which should be verified with a greater collection effort in these places.

The histogram plot of latitude data shows higher frequencies (0.57) for southern latitudes from 12°30'0" S (Fig. 4a). This pattern could be explained by the fact that the Southern Peruvian Andes have a wider area, mostly covered with highland grasslands, which are also easier places for collecting, using a combination of pitfall trapping and direct searching under rocks. The histogram plot of altitude data shows higher frequencies (0.76) for altitudes above 3000 m (Fig. 4b). This means that the genus Pilobalia would have largest populations and species number inhabiting highland plains and interandean valleys, while ravines covered with cacti and shrubs at lower altitudes would be secondary habitats. Also, as stated above, collecting is easier in larger areas and more difficult in narrow places, where populations could also be smaller.

Acknowledgments

The author wishes to thank Gustavo Flores (IADIZA), Eliana Quispitupac (LEUSM), Clorinda Vergara (MEKRB), Mabel Alvarado (MUSM), and Graciano Tejada (SENASA) for the collaboration provided during the visit to their respective entomological collections. Also, thanks to Nora Cabrera (MLP) for the loan of type specimens to IADIZA, which were examined during two author's visits, the first during the Fourth International Tenebrionoidea Symposium (November 25-26th, 2015) and the second for the internship titled “Implementación de investigaciones sobre Sistemática de tenebriónidos sudamericanos (Coleoptera, Tenebrionidae)” (FONDECYT–Peru, September 3–16, 2017). Photos of type specimens were kindly provided by Rachel Diaz-Bastin and Chris Grinter (CASC), Christophe Rivier and Antoine Mantilleri (MNHN), and Christoph Germann (NHMB).

REFERENCES

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Notas de autor

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