Polynoncus pilularius ( Germar, 1823 )

Costa-Silva, Vinícius, Strümpher, Werner P., Thyssen, Patricia J. & Vaz-de-Mello, Fernando Z., 2024, Taxonomic revision of the South American genus Polynoncus Burmeister, 1876 (Coleoptera: Scarabaeoidea: Trogidae), Journal of Natural History 58 (1 - 4), pp. 14-166 : 128-135

Polynoncus pilularius ( Germar, 1823)

( Figures 57–58 )

Trox pilularius Germar, 1823: 113 (original description); Dejean 1833: 148 (checklist – as ‘ pillularius ’); Curtis 1844: 445 (comments and records); Lacordaire 1856: 152 (checklist – as ‘ pillularius ’); Harold 1869: 1089 (checklist); Harold 1872: 139, 191 (redescription); Burmeister 1876: 262 (redescription); Preudhomme de Borre 1886: 75 (checklist); Bruch 1911: 193 (checklist); Luederwaldt 1911: 420 (list of necrophagous beetles from São Paulo); Arrow 1912: 59 (catalogue); Blackwelder 1944: 219 (checklist – as ‘ pilularia ’); Vaurie 1962: 127 (redescription)

Trox (Polynoncus) pilularius : Scholtz 1982: 16 (catalogue)

Polynoncus pilularius : Scholtz 1986a: 362 (systematics); Scholtz 1990: 1437 (redescription); Gómez 2008: 517 (key to Argentinean species); Zidek 2013: 15 (checklist); Zidek 2017: 106 (checklist); Smith 2017: 100 (lectotype designated); Costa-Silva and Diéguez 2020: 272 (records from Argentina)

Type specimen examined. LECTOTYPE. ( MNHN – Figure 57 ) First label [white, aged, handwritten]: ‘typ. / pilullarii’. Second label [green with black border, Ernst Germar’s handwriting]: “pilularius [underline] m. / 367* Buenos Ayres. Third label [white, aged, with black frame, typeset]: ‘Ex-Musӕo / E. Harold’. Fourth label [white, typeset]: ‘Museum Paris / ex Coll. / R. Oberthur’. Fifth label [red, typeset]: ‘LECTOTYPE’. Sixth label [red with black frame, Andrew Smith’s handwriting]: ‘ TROX / PILULARIUS / Germar 1824 / LECTOTYPE / A.B.T. SMITH [typeset]’ ( Figure 57c ). Type locality: ‘Buenos Ayres’ (Buenos Aires), Argentina.

Differential diagnosis. Polynoncus pilularius is most similar to P. brevicollis , and can easily be separated by having elytral tubercles coated, which is glabrous in P. brevicollis . Also, P. pilularius has well-developed tubercles near the lateral margin of elytra, while these tubercles are absent in P. brevicollis (or with just a few granules). Besides that, P. pilularius has the basal tubercles of the pronotum distinctly separate from the pronotal ridges, while they join the pronotal ridges in P. brevicollis . Polynoncus pilularius is also similar to P. gemmingeri , but the shape of scutellum is diagnostic to separate the two species. Polynoncus pilularius has a shield-shaped scutellum, which is elongated in P. gemmingeri . For details of description, see Vaurie (1962) and Scholtz (1990).

Geographic distribution. This species is widely distributed in the southern half of South America and occurs in Argentina, Bolivia, Brazil, Chile [?], Paraguay and Uruguay ( Figure 58 ).

Vaurie (1962) mentioned one specimen from ‘Chile’ without further information about the depository. We examined three specimens (1 BMHN and 2 MNHN) from the country without further data. One of these specimens, housed in MNHN,was collected by Charles Gaudichaud-Beaupré (1789–1854) during his third excursion to South America aboard the ship La Bonite between 1836–1837 ( Papavero 1971). According to Papavero (1971), the ship anchored in Valparaíso on 10 June 1836, where he spent several days making scientific observations/ collecting, before departing the country on 24 June.

However, it is important to point out that we noticed several mislabelled specimens collected by Gaudichaud-Beaupré housed in MNHN. Other than this isolated record for Chile, P. pilularius has never been reported or mentioned again from the country (eg Diéguez 2008).

Also, during the examination of specimens, we found one individual from Tefé city (Amazonas, Brazil) in the MZSP collection, collected by Ricardo von Diringshofen (or ‘Dirings’) in 1962. According to Ferreira et al. (2016), some of the specimens housed in Dirings’ collection were bought from Brazilian amateur collectors, and several mislabelled specimens were noticed in his collection ( Ferreira et al. 2016; Sousa and Seidel 2021). Also, according to our understanding of the geographical distribution of P. pilularius , we believe that this specimen housed in Dirings’ collection is another case of mislabelling of specimens. These records (from Chile and northern Brazil) should be treated as uncertain until additional data of this species from this locality or region arises.

Remarks. During this study, we noticed several specimens from Bolivia being larger in size when compared with the individuals of P. pilularius from Argentina, Brazil, Paraguay and Uruguay. However, we conclude that the size of those specimens from Bolivia could be interpreted as population morphological variation and not necessarily a new species.

Examined non-type material (432 specimens). ARGENTINA: Buenos Aires, Balcarce , no date, J .M . Gallardo leg . (6 MACN); Bahía Blanca, no date or collector (1 MNHN); Buenos Aires , November 1944, no collector (1 SAMC); Buenos Aires, no date, M . Viana leg . (6 CVMD); Buenos Aires, Santa Catalina, no date or collector (1 MACN); Buenos Aires , Catamarca , date or collector (11 MACN); Carmen de Patagones , 7 January–1 September 1829, d’Orbygni leg . (2 MNHN); Felipe Solá , November 1961, A . Martinez leg . (5 MZSP); Gorchs , March 1917, G . Pellerano leg . (1 MACN); Gorchs , December 1918, G . Pellerano leg . (1 MACN); Hurlingham , March 1956, M . Viana leg . (2 CVMD); La Plata , February 1941, Mouros leg . (1 DZUP); La Plata, 1879, no collector (1 MNHN); La Plata , 9 September 1986, Sergio Roig leg . (1 IADIZA); La Plata , no date, A .R . Bezzi leg . (2 MLPA); La Plata, no date or collector (1 MNHN; 1 MLPA; 1 BMNH); La Plata , October 1948, Gaggero leg . (1 MLPA); Miramar, General Alvarado , 26 November 1946, Bachmann leg . (1 MACN); Miramar, General Alvarado , 16 November 1946, Bachmann leg . (1 MACN); Miramar, General Alvarado , 24 November 1946, Bachmann leg . (1 MACN); Miramar, General Alvarado , 25 November 1946, Bachmann leg . (1 MACN); Miramar, General Alvarado , 18 November 1946, Bachmann leg . (1 MACN); Olivos , December 1929, C . Bruch leg . [Bruch’s collection] (1 MACN); Pehuajó , 26 March 1938, Biraben-Scott leg . (1 MLPA); Rosas , ‘F .C. Sud’ [Ferrocarril al Sud – Train Station], no date, J .B . Daguerre leg . (3 MACN); San Miguel, Campo de Mayo , no date, Heritier leg . (1 SAMC); San Pedro , no date, A .G . Frers leg . (1 MACN); Buenos Aires, Sierra La Ventana , 27 September 1981, Sergio Roig leg . (2 IADIZA); Tolosa, no date or collector (2 MACN); Tornquist , Abra de la Ventana, 1 February 1947, Bachmann-Rossi leg . (3 MACN); Tornquist, Ernesto Tornquist Provincial Park , 31 January 1947, Bachmann-Rossi leg . (1 MACN); without specific locality, 15 January–15 March 1827, d’Orbygni leg . (1 MNHN); without specific locality or date, C . Berg leg . (2 MNHN); without specific locality, 1897[?], no collector (1 MNHN); without specific locality, 21 December 1908, J .B . Daguerre leg . (1 MACN); without specific locality, date or collector (13 MNHN; 2 BMNH; 2 MACN); without specific locality, date or collector [ Burmeister’s collection] (1 MACN); without specific locality, January 1977, J . Barriga leg . (1 CVMD); without specific locality, February 1961, A . Martinez leg . (1 MZSP); without specific locality, March 1930, no collector (1 MACN); without specific locality or date, A . Breyer leg . (2 MACN); without specific locality or date, A . Pellerano leg . (1 MACN); without specific locality or date, d’Orbigny leg . (1 MACN); without specific locality, June 1897, C . Bruch leg . [ Bruch’s collection] (1 MACN); without specific locality, October 1897, S . Vant leg . (1 MACN); without specific locality, October 1978, J . Barriga leg . (7 CVMD); Zelaya , 1936, J .A . Pereyra leg . (1 MACN); Zelaya , October 1978, J .E . Barriga leg . (1♀ and 1 unsexed CVMD) . Catamarca, Aconquija , El Sancho, 2500 m, 15 November 1915, P . Jorgensen and Joel Blamey leg . (2 OUMNH); Capilitas , 10 November 1946, R . Golbach leg . (1 IFML); El Arenal, 4 December 2003, collected actively, no collector (1 IFML); El Rodeo , 1296 m, January 1942, Dirings leg . (1 MZSP); El Taco , 1395 m, 28.707°S, 65.618°W, 4–7 February 2009, K . Sheldon, F .C. Ocampo leg . (1 IADIZA); La Merced , 20 February 1999, G . Arriagada leg . (2 CVMD). Chaco, Resistencia , December 1958, M . Viana leg . (1 CVMD); Resistencia , December 1958, M . Viana leg . (1♂ and 1 unsexed CVMD); Río Muerto , 26.125°S, 61.677°W, 31 October 2016, pitfall trap, C . Guerra-A leg. (1♀ CEMT) ; Roque Sáenz Peña , 1932, K .J . Hayward leg . (1 BMNH); without specific locality, 4 February 1986, no collector (2 SAMC); without specific locality, date or collector (1 MACN); without specific locality, December 1987, no collector (1 SAMC) . Córdoba, Alpa Corral, Eloisa , 1042 m, 32.632°S, 64.723°W, 5–7 February 2009, K . Sheldon, F .C. Ocampo leg . (1 IADIZA); Alta Gracia, December 1920, no collector [Bruch’s collection] (1 MACN); Bajo Grande , 5 November 1939, Maldonado leg . (1 MLPA); Calamuchita, Hotel El Sauce , December 1938, M . Viana leg . (4 DZUP); Calamuchita, El Sauce , January 1962, M . Viana leg . (1♂ and 2 unsexed CVMD); Cruz Alta , 1 February 1946, Duret leg . (1 CVMD); Fauna Nature Reserve Laguna La Felipa, 24 October 2002, horse carcass, no collector (1♂ IADIZA; 1 MNHN); Fauna Nature Reserve Laguna La Felipa , 26 December 2002, R . Gomez leg . (1 IFML); Fauna Nature Reserve Laguna La Felipa , 24 November 2002, sheep carcass, R . Gomez leg . (1 IFML); Laguna La Helvecia , 14 January 2003, R . Gomez leg . (1♂, 1♀ and 2 unsexed CVMD); Punilla, Copina , 1600 m, 14 April 1967, G . Barria leg . (1 CVMD); without specific locality, date or collector [Burmeister’s collection] leg. (1 MACN) ; without specific locality, date or collector (1 MLPA); without specific locality or date, A . Beyer leg . (1 MACN); without specific locality, 22 May 1924, J .B . Daguerre leg. (3 MACN). Corrientes, Santo Tomé , January 1923, G . Pellerano leg . (1 MACN); without specific locality or date, C . Bruch leg . [Bruch’s collection] (1 MACN) . Entre Rios, Colón , 16 February 1987, O . Mitre leg . (1 SAMC); Paraná, no date or collector (1 MACN); Paraná, no date or collector [Burmeister’s collection] (1 MACN); Pronunciamiento , 22 January 1987, O . Mitre leg . (4 SAMC); Pueblo Liebig , February 1996, Mateo Zelich leg . (1 CVMD); San Antonio de Padua de la Concordia , no date, H . Rodri . [sic!] leg. (1 MZSP); Villa Elisa , November 1973, L . Gontero leg . (1 MACN). Jujuy, Cerro Perales 6 February 1950, Willink y Monrós leg . (1 IFML). Misiones, Posadas , 10 November 1970, C . Porter leg . (1 SAMC); without specific locality, date or collector (2 MLPA); without specific locality or date, A . Breyer leg . (1 MACN). Rio Negro, village de Patagones [= Viedma ], 7 January–1 September 1829, A . d’Orbigny leg . (2 MNHN). Salta, Cachi , February 1983, M . Viana leg . (1♀ and 1 unsexed CVMD); Chicoana , November 1988, M . Viana leg . (1 CVMD); Corralito , October 1986, M . Viana leg . (1 CVMD); Corralito , October 1986, M .J . Viana leg . (1♀ CVMD); Cuesta del Obispo , November 1983, M . Viana leg . (1 CVMD); Guachipas , 1106 m, January 1951, A . Martinez leg . (1 MZSP); La Viña , December 1986, M . Viana leg . (1 CVMD); Pampa Grande, no date or collector (1 MACN); Piedra del Molino , 2640 m, January 1988, A . Martinez leg . (1♂ JZPC); Rosário de Lerma , January 1983, M .J . Viana leg . (1♂ CVMD); Ruiz de los Llanos [also known as ‘ El Tala’], February 1947, R . Golbach leg . (1 IFML); Sierra de Metán , 16–22 September 1977, Scrocchi leg . (1♂ IFML); Sierra de Metán , 2400 m, 13 October 1977, Dominguez leg . (2 IFML); without specific locality, date or collector (1 MACN). San Luis, Coronel Pringles , La Carolina, 26 September 1970, M . Viana leg . (1♂ CVMD); Villa Elena , 23 October 1971, M . Viana leg . (2 CVMD); Villa Elena , November 1972, M . Viana leg . (1♀ CVMD); Villa Elena , November 1976, M . Viana leg . (2♂ and 6 unsexed CVMD); without specific locality, date or collector (9 MACN) . Santa Cruz, Las Horquetas , 19 November 1939, Biraben-Bezzi leg . (2 MLPA). Santa Fé, Colonia Florencia, Rio Tapenagá , 1903, E .R . Wagner leg . (1 MNHN); Golondrina , August 1924, K .J . Hayward leg . (1♂ BMNH); Rio las Garzas, 1918, no collector (4 BMNH); Rio las Garzas , Ocampo 25 km, 1903, E .R . Wagner leg . (1 BMNH); Rosário, no date or collector (1 MACN); Santa Fé, Rosário , no date, A . Stevenin leg . (1 MACN); without specific locality or date, C. Bruch [Bruch’s collection] leg. (1 MACN) ; without specific locality or date, Weiser leg . [Bruch’s collection] (3 MACN) . Santiago del Estero, Icaño, Edges of the Rio Salado , 1909, E .R . Wagner leg . (2 MNHN); Icaño, Guarda Escolta , November–December 1909, E .R . Wagner leg . (1 MNHN); Rio Salado, no date or collector (1 SAMC); Río Salado , no date, Wagner leg . (6 IFML); without specific locality or date, Wagner leg. (1 MLPA). Tucumán, 5 km E El Infiernillo , 7 December 2003, under horse carcass, R . Gomez leg . (1 CVMD; 1 IFML); 5 km E El Infiernillo, 7 December 2003, no collector (1 CVMD); Burruyacú, Villa Padre Monti , 21 January–7 February 1948, R . Golbach leg . (1 IFML); Choromoro, Alto de las Salinas , 25 February 1900, no collector (1 IFML); Colalao , January 1954, M . Viana leg . (1♂ CVMD); Experimental Reserve Horco Molle , May 1977, Carpintero leg . (1 CVMD); Lules , December 1957, A . Martinez leg . (3 MZSP); San Javier , April 1955, Duret leg . (2 CVMD); San Javier Ligustral , 26.798°S, 65.333°W, 11 December 2012, B . Maldonado leg . (1 IADIZA); San Javier 26.771°S, 65.326°W, 12 December 2012, B . Maldonado leg . (3 IADIZA); San Javier 26.783°S, 65.329°W, 10 December 2012, B . Maldonado leg . (1 IADIZA); San Javier Ligustral , 26.798°S, 65.333°W, 8 December 2012, B . Maldonado leg . (1 IADIZA); Tacanas , January 1949, M . Arnau leg . (17 IFML); Tacanas , 24 August 1973, L . Stange leg . (1♂ IFML); Taficillo December 1929, no collector (1 IFML); without specific locality, December 1946, R . Golbach leg . (4 IFML); without specific locality, November–December 1944, R . Golbach leg . (1 IFML); without specific locality, date or collector (1 MACN); without specific locality, 1940, no collector (1 MACN); without specific locality, February 1902, no collector (2 MACN) . Without specific locality, no date or collector (1 CVMD) . BOLIVIA: Chiquisaca, Sucre , January–February 1833, d’Orbygni leg . (3 MNHN). Cochabamba, Chapare , 400 m, September 1947, no collector [Justus’ collection] (2 DZUP); without specific locality, 17 September–12 October 1899, P . Germain leg . (4 MNHN). La Paz, Nor Yungas, Caranavi , 1500 m, October 1990, P . Arnaud leg . (2♀ CEMT) . Potosí, Betanzos , 3345 m, 1 February 2008, Marcos Ferru leg . (7♂, 3♀ and 8 unsexed CVMD); Betanzos , 3345 m, 16 February 2008, under Vulpes sp . carcass, Marcos Ferru leg . (3♂ and 3♀ CVMD) . Without specific locality, date or collector (2 BMNH); without specific locality, 1933, d’Orbygni leg . (1 MNHN); without specific locality, date or collector (8 MNHN). BRAZIL: Amazônas [dubious locality], Tefé , June 1962, Dirings leg . (1 MZSP). Paraná, Castro , 1907, E . Garbe leg . (3 MZSP); Curitiba , 900 m, 14 March 1968, Depto . Zoo . Exp. (2 DZUP) ; Ponta Grossa, Vila Velha Park , 7 October 2007, P . Grossi leg . (1♀ CEMT); without specific locality, date or collector (1♂ BMNH) . Rio Grande do Sul, Bagé, Embrapa Pecuária , January 2006, pitfall trap, L .D . Audino leg . (1 CEMT); Bagé, Embrapa Pecuária , January 2006, pitfall trap, L .D . Audino leg . (2 CVMD); Barão de Cotegipe , May 1966, S . Giacomel leg . (1 DZUP); Pelotas , 15 March 1951, Blezanko leg . (2 CVMD); Pelotas , 18 January 1950, C .M . Biezanko leg . (1♂ BMNH); Pelotas , 31 August 1951, C .M . Biezanko leg . (1 BMNH); Porto Alegre, April 1928, no collector (1 MGAP); Rio Grande, no date or collector (1 BMNH); São Francisco de Paula , February 1944, no collector (1 MGAP), without specific locality, date or collector (1 MNHN) . Santa Catarina, Bom Jardim da Serra, Mantiqueira , December 2001, C . Arioli leg . (1♂ CEMT); Florianópolis, São João do Rio Vermelho, 800 m, April 1958, no collector (1 DZUP); Joinville , November 1950, Dirings leg . (2 MZSP); Mafra , 800 m, April 1961, no collector (1 DZUP); Theresópolis colony [currently Águas Mornas], 1888, H . Fruhstorfer leg . (1 MNHN); Urubici, São Joaquim National Park , 1685 m, 28.158°S, 49.626°W, 13 January 2016, meat, Atlantic Forest, P .G . da Silva leg . (1♂ CEMT); Urubici, São Joaquim National Park , 1738 m, 28.117°S, 49.500°W, 14 January 2016, meat, Atlantic Forest, P .G . da Silva leg . (1♀ EMAC); Urubici, São Joaquim National Park , 1751 m, 28.120°S, 49.495°W, 14 January 2016, pitfall trap with meat, Atlantic Forest, P .G . da Silva leg . (2♀ CEMT); without specific locality, date or collector (6 BMNH) . São Paulo, Campos do Jordão , January 1952, M .A.V.A leg. (1 MZSP); São Paulo, Brooklin , 16 August 1959, Reichardt leg . (2 MZSP); São Paulo, Ipiranga, no date or collector (5 MZSP); São Paulo , Jabaquara , January 1959, B . Pohl leg . (2 MZSP); São Paulo, no date or collector (3 MZSP); São Paulo, Serra da Cantareira , March 1952, Dirings leg . (1 MZSP). Without specific locality, Paraná River, no date or collector (1 MNHN); without specific locality, date or collector (1♂ and 2 unsexed MNHN); ‘ Rio’ [sic!], no date or collector (1 MNHN) . CHILE: Valparaíso (V Region), without specific locality, June 1836, C . Gaudichaud-Beaupré [ Exp. La Bonite ] leg. (1 MNHN) . Without specific locality, date or collector (1 BMNH); without specific locality, date or collector (1 MNHN) . PARAGUAY: without specific locality, date or collector (1 MACN) . URUGUAY: Camelones, Camelones, Estación Ferrocaril , 8 March 2002, Gandini leg . (3 CEMT); Puntas del Arenal no date or collector (1 MZSP) . Cerro Largo, Bañado de Medina , 13–20 December 2011, pitfall trap (human + pig dung as bait), R .M . Moraes leg. (1 CEMT). Colônia, La Estanzuela National Institute for Agricultural Research , 12 September 1995, J .R . Verdú leg . (1♂ and 1 unsexed CEMT) . Florida, Cerro Colorado , 1911, P . Serre leg . (1 MNHN). Maldonado, Bella Vista , November 1980, W .D . Rovera leg. (2 HNHM). Montevideo, Rocha , La Esmeralda, 2 July 1977, A . Witeck leg . (1 SAMC); Rocha, Rocha, no date or collector (1 MACN); Villa Colon , May 1886, Oldfield Thomas leg . (1♀ BMNH); without specific locality or date, C . Darwin [Beagle voyage] leg. (3 BMNH) ; without specific locality, 1820, no collector (1 MNHN); without specific locality, 1861, Birabent leg. (4 MNHN); without specific locality, 1863, A . Brunel leg . (4 MNHN); without specific locality, 1912, P . Serre leg . (2 MNHN); without specific locality, date or collector (3♂ and 1 unsexed BMNH; 1 MACN; 6 MNHN; 2 MZSP); without specific locality, January 1976, W .D . Rovera leg. (1 HNHM); without specific locality or date, A . Beyer leg . (1 MACN); Paysandú, Paysandú, no date or collector (3 MNHN) . Soriano, without specific locality or date, T . Havers leg . (1 BMNH). Without specific locality, date or collector (3 MNHN) .

Remarks. The year of description of Trox pilularius is historically dated as 1824 according to the title page of Germar’s work entitled ‘ Insectorum species novae aut minus cognitae, descriptionibus illustratae. Volumen primum. Coleoptera . Cum Tab. aen. II’. (ie Harold 1872; Arrow 1912; Blackwelder 1944; Vaurie 1962; Scholtz 1982, 1990; Zidek 2013, 2017; Smith 2017; Costa-Silva and Diéguez 2020). However, as first mentioned by Ersch (1828, p. 1166) and followed by Bousquet (2016, p. 211), the correct date of publication of Germar’s work is 1823.

Figures

Figure 57. Lectotype of Trox pilularius Germar, 1823 (now Polynoncus pilularius): habitus in a, dorsal and b, lateral view; c, labels. Scale bar: 1 mm.

Figure 58. Distribution map of Polynoncus pilularius (red circles), P. sallei (green triangles), and P. tenebrosus (blue squares) in South America. The records with a black dot in the centre indicate the type locality of the respective species.