Dichotomius (selenocopris), 1846

Nunes, Rafael V. & Vaz-de-Mello, Fernando Z., 2019, Taxonomic revision of Dichotomius (Cephagonus) Luederwaldt 1929 and the taxonomic status of remaining Dichotomius Hope 1838 subgenera (Coleoptera: Scarabaeidae: Scarabaeinae: Dichotomiini), Journal of Natural History 53 (37), pp. 2231-2351 : 2240-2244

Dichotomius (selenocopris) bicuspis ( Germar 1824)

( Figure 12 (a))

Copris bicuspis ı Germar (1824) ı p. 101 (description)

Copris (selenocopris) bicuspis ı Burmeister (1846)ı p. 87

Pinotus bicuspis ı Harold (1869) ı p. 1009

Pinotus rotundigena ı Felsche (1901) ı p. 52 new synonym ( Figure 3 (b))

Pinotus (Cephagonus) ascanius ı Luederwaldt 1929 ı pp. 115 – 116

Pinotus (Cephagonus) bicuspis rotundigena ı Luederwaldt 1929 ı p. 117 – 118

Pinotus (Cephagonus) ascanius piceus ı Luederwaldt 1930 ı pp. 119 new synonym ( Figure 3 (c)) Pinotus (Cephagonus) ascanius piceus var. rotundiceps ı Luederwaldt 1930 ı p. 119 (invalid name)

Pinotus (Cephagonus) ascanius ı Luederwaldt 1931 ı pp. 357

Pinotus (Cephagonus) ascanius piceus ı Luederwaldt 1931 ı p. 357

Pinotus (Cephagonus) ascanius aceratus ı Pereira 1944 new synonym

Pinotus ascanius ı Blackwelder 1944 ı p. 206

Pinotus ascanius piceus ı Blackwelder 1944 ı p. 206

Pinotus ascanius rotundiceps ı Blackwelder (1944) ı p. 206

Dichotomius (selenocopris) bicuspis ı Martínez 1951 ı p. 142

Dichotomius (selenocopris) bicuspis ı Vaz-de-Mello 2000 ı p.193

Dichotomius (selenocopris) ascanius aceratus ı Vaz-de-Melloı 2000ı p. 193

Dichotomius (Luederwaldtinia) guaranii Gandini & Aguiları 2009 ı p. 155 (description) new synonym

Diagnosis. D. (s.) bicuspis is separated from other species in the subgenus by the following combined characters: male´s cephalic horn with conical apex; larger male´s pronotal disc four-lobed (as in larger males of larger species in ascanius , fissus and quadraticeps groups in Cephagonus and speciosus group in selenocopris); ocellate punctures present on anteriorı posterior pronotal angles and pronotal posterior margin (as in sericeus and batesi species group of selenocopris) and female´s sixth ventrite three times larger than fifth ventrite medially (as geminatus group in selenocopris; pygidium swollenı strongly convex (as in females of Cephagonus and groupo inachus of selenocopris).

● HOLOTYPE of Copris bicuspis (male) ( Figure 12 (a)): Type Germar {whiteı handwritten}/ 7 {whiteı handwritten}/ bicuspis m. {blue with black marginsı handwritten}/Brasilia P. bicuspis Germ. {white with red marginsı handwritten}/Ex Musaeo E. Harold {white with black marginsı printed} [NMHU];

● HOLOTYPE of Pinotus rotundigena (male): Mirandaı XI-94 – II-95 (Andeer) {white with black marginsı printed}/Typus {white with black marginsı handwritten in red colour}/Coll. C. Felsche Kauf 10ı 1910 {green with black marginı printed}/ rotundigena Felsche C. Brasilien {white with purple marginsı handwritten}/Museum für Tierkunde Dresden {whiteı printed}/ HOLOTYPE Pinotus rotundigena Felsc. {red with black marginsı partially handwritten} [SNSD];

● LECTOTYPE of Pinotus (Cephagonus) ascanius piceus (maleı here designated): São Pauloı Ipiriangaı x-1914 {white with black marginsı partially handwritten}/COTIPO {red with black marginsı printed}/ Pinotus ascanius Hr. Lued. Det. 14 {whiteı handwritten}/ Pinotus ascanius subspecies piceus Luederw. Det 29 {whiteı handwritten}/ 26861 {whiteı printed} [MZSP];

● HOLOTYPE of Pinotus (Cephagonus) ascanius aceratus : we have not fond the type material for this speciesı see this species remarks.

● HOLOTYPE of Dichotomius guaranii : not examinedı picture and description in the original paper and geographical locality considered by us to be enough to consider the present synonymy. [MNHNP].

Non type material. ARGENTINA: Misiones: San Pedro. xi-2012. P Wagner [4 ƋƋ at CEMT]. BOLIVIA: Santa Cruz: Chiquitos. ENE Santiago. 18³20 ʹ 10 ” Sı 59 ³35 ʹ 0 ” W. 622 m. Bosque chiquitano. Hum. faec. xi-2008 WD Edmonds & T Vidaurre [2 ♀♀ at CEMT]. BRASIL: Bahia: Barreiras. x-xii-1991. Luz. [3 ƋƋ at CEMT]ı same municipality but ix- 1991ı JC Zanúncio (lgt) [1 ♀ at CEMT]; Distrito Federal: Brasília. FAL-Unbı x-xi-1985 [2 ƋƋı 1 ♀ at CEMT]ı same locality but iv-1986 [1 Ƌ at CEMT]; same municipality but Recor IBGEı 9³15 ’ Sı 47 ³51 ʹ W. 1150 m. Cerrado s. str. FIT. 9-xii-1998. MS Milhomem [3 ƋƋı 4 ♀♀ at CEMT]; same municipality but Brazlândia. I-1994. E Grossi [1 ♀ at CEMT]; same municipatily but Parque Nacional de Brasíliaı 15 ³43 ʹ 24 ” S 47³59 ʹ 53 ” Wı pitfall. 8-iii-2015. MCV Rocha [1 ♀ at CEMT]; same municipality but Planaltina. Embrapa CPA C. 15³36 ʹ 24 ” Sı 47 ³42 ʹ 45 ” W. 1025 mosl. Cerradão. FIT. CM Oliveira [1m #ı 1 ♀ at CEMT]; Espírito Santo: Reserva Nat. Valeı porco. 7-9-ix-2012. RC Lima [1 Ƌ at CEMT]; Goiás: Aragarças. Aeródromo. 7-x- 1989ı A Bello [3 Ƌ at CEMT]; Joanápolis. Chácara São José. 21-ii-2015 [1 ♀ at CEMT]; Mineirosı Parque Nacional das Emas. 17³54 ʹ 20 ” Sı 52³59 ʹ 04 ” Wı 837 m. 15-iii-2011. Luz. MF Souza [1 Ƌı 3 ♀♀ at CEMT]; Mato Grosso: Barão de Melgaço [1 Ƌ at CEMT]; Barra do Garças. 15-xi-1988 [1 ♀ at CEMT]; same municipality but Cema. 20-iv-1991. MA Serrano [1 Ƌ at CEMT]; Campo Verde. Rio Engano. 14-x-2012. Peña & Peña [5 ƋƋı 1 ♀ at CEMT]; same municipality but Fazenda Caverá. 11-x-2011. A Biava [1 ♀ at CEMT]; Chapada dos Guimarães. 2012. WO Souza [1 ♀ at CEMT]; Cuiabáı Santa Isabel. 19-i-2012. M Alexandre [1 ♀ at CEMT]; same municipality but Jardim Presidente. x-2011. MMR Castro [1 ♀ at CEMT]; Paratininga. 10-15- xi-2011. O Neto [1 Ƌ at CEMT]; Primavera do Leste. Fazenda Ilha Grande. 15³50 ’ Sı 54 ³13 ʹ W. ii-2013. AS Tissiani [2 ƋƋı 2 ♀♀ at CEMT]; Santo Antônio de Leverger. São Vicente da Serra. 15³49 ʹ 42 ” Sı 55 ³25 ʹ 11 ” W. 15-xii-2010. AS Tissiani & FZ Vaz-de-Mello [1 Ƌı 1 ♀ at CEMT]; Tangará da Serra. Campus UNEMAT. 10-xi-2011. P Souza [1 ♀ at CEMT]; same locality but Sitio Clemente. 14³39 ʹ 46 ” Sı 57 ³49 ʹ 35 ” W. 368 m. FIT. RJ Silva [1 Ƌ at CEMT]; Várzea Grande. Bom Sucesso. 20-x-2011. S Simm [3 ♀♀ at CEMT]; same municipality but Praia Grande. 15-xi-2011. D Barbosa [1 Ƌ at CEMT]; Vila Bela da Santissima Trindade. 7-22- ix-1997. M Calleffo [1 ♀ at CEMT]; Mato Grosso do Sul: Bonito. Fazenda Brasil Bonito. 21³6 ’ Sı 56 ³38 ʹ W. Fezes humanas. FO Roque [5 ƋƋı 1 ♀ at CEMT]; same municipality but Fazenda Morro Alto. 21³02 ʹ 25 ” Sı 56 ³37 ʹ 47 ” W. xi-2009. FO Roque [1 Ƌ at CEMT]; Campo Grande. Embrapa CNPG C. Pastagem Brachiaria. Fezes Bovinas (Nelore). 9-xi-1983. WW Koller [1 Ƌ at CEMT]; same municipalityı locality and habitat but Luz UV. ii-1987. A Gomes [1 ♀ at CEMT]; Campo Grande. x-1947. A Maller [3 ƋƋı 1 ♀ at CEMT]; same municipality but FIT. iii-1986. Bianchin [2 ƋƋ at CEMT]. Dourados. x-2010. VR Grando [1 ♀ at CEMT]; same municipality but 14-xi-2010. SM Ramos [1 Ƌ at CEMT]; same municipality but Campus UFG. 26-xi-2012. FZ Vaz-de-Mello [2 ƋƋ at CEMT]; same municipality but 5-xi- 2005. M Milo [1 Ƌ at CEMT]; Três Lagoas. Horto Rio Verde. LUZ UV. 19-xi-1993. CH Fletchmann [1 Ƌı 1 ♀ at CEMT]; Minas Gerais: Barroso. 8-ix-2007. MA Nascimento [1 Ƌı 1 ♀ at CEMT]; Belo Horizonte. PE Rola-Moça. 19³50 ’ Sı 43 ³50 ʹ W. 900 m. R Durães [1 ♀ at CEMT]; Buritis. Fazenda Querência. Luz. 3-xii-2012. CM Oliveira. [5 ƋƋı 10 ♀♀ at CEMT]; Caraça. xi-1993. A Bello [1 Ƌ at CEMT]; Carmo da Cachoeira. 21³22 ʹ 47 ” Sı 45 ³5 ʹ 33 ” W. Frutos Annona crassiflora. 19-iii-2008. DL Bandeira de Melo [32 ♀♀ at CEMT]; Caxambú. xii-1990. A Bello [4 ♀♀ at CEMT]; same locality but iii-1994. A Bello [1 ♀ at CEMT]; Conceição dos Ouros. Mata da Chapada Rio Sapucaí. ii-2003. GP Almeida-Neto [1 ♀ at CEMT]; Cordisburgo. Fazenda Pontinhas. ii-1996. FZ Vaz-de-Mello [2 ♀♀ at CEMT]; same municipality but Lagoa Bonita. i-2011. FZ Vaz-de-Mello [1 ♀ at CEMT]; Divinópolis. xi-1998. R Campos [2 ♀♀ at CEMT]; Ipatinga. xi-1992. E Grossi [1 ♀ at CEMT]; Itumirim. xii-2012. F França [1 Ƌ at CEMT]; Lavras. 17-ix-1997 [1 ♀ at CEMT]; same municipality but iii-1998. JNC Louzada [1 ♀ at CEMT]; same municipality but 25.iv.2000. H Carlos [1 ♀ at CEMT]; same municipality but 22.ii.2002. RF Melo [1 Ƌ at CEMT]; same municipality but 21³19 ’ Sı 44 ³59 ʹ W. 20.i-2008. MR Rocha & DHT Takahashi [17 ♀♀ at CEMT]; same municipality but Campus UFLA. 27-iii-2008 [1 Ƌ at CEMT]; same municipality and locality but Luz. 11. xii.2011. FZ Vaz-de-Mello [6 ♀♀ at CEMT]; Mariana. 7-xii-1998. JL Moreno & T Mestre. [1 Ƌ at CEMT]; Montes Claros. i-2002. GLD Leite. [1 ♀ at CEMT]; Prados. 21³03 ʹ 26 ” Sı 44 ³05 ʹ 52 ” W. 1094 mosl. Baço bovino. 17-ii-2002. L Vieira [1 Ƌı 1 ♀ at CEMT]; Poços de Caldas. ii-2000. C Lopes-Andrade [1 ♀ at CEMT]; São Lourenço. 20-i-1993. J Celso [1 Ƌ at CEMT]; São Roque de Minas. PN Canastra Jaguaré. 20³15 ʹ 17 ” Sı 46 ³25 ʹ 11 ” W. Campo Limpo. Manual. MF Souza [1 Ƌı 6 ♀♀ at CEMT]; São Tomé das Letras. xi-1991. J Celso. [1 ♀ at CEMT]; Serrinha. 3. iii.2007. DHT Takahashi [1 Ƌı 8 ♀♀ at CEMT]; Turvolândia. ii-1997. FO Micherell [1 ♀ at CEMT]; Três Marias. x-1990 JC Zanuncio [1 Ƌ at CEMT]; Uberaba. x-1995. FZ Vaz-de-Mello [1 ♀ at CEMT]; Viçosa. x-xi-1981. F Fiuza [2 ƋƋı 7 ♀♀ at CEMT]; same municipality but 4-x-1988. DF Avilés [2 ♀♀ at CEMT]; same municipality but 13-x-1988. A Mayhe [1 ♀ at CEMT]; same municipality but 5.vi.1990. P Jusselino [1 ♀ at CEMT]; same municipality but x-1992. JNC Louzada [5 ♀♀ at CEMT]; same municipality but Mata do Paraíso. Luz UV. xi- 1992. F Fiuza [1 ♀ at CEMT]; same municipality but Mata do Paraíso. 20³48 ʹ 08 ” Sı 42³51 ʹ 31 ”. xii-1984. Luz. F Fiuza [1 Ƌ at CEMT]; same municipality but 30.viii.1994. JNC Louzada [1 ♀ at CEMT]; same municipality but Campus UFV. 18-xi-1995. FZ Vaz-de-Mello [1 ♀ at CEMT]; same municipality and locality but xi-1996. FZ Vaz-de-Mello [1 ♀ at CEMT]; same municipality and locality but x-1997. FZ Vaz-de-Mello [2 ♀♀ at CEMT]; Paraná: Balsa Nova. São Luiz do Purunã. Fazenda Monjolo. 25³45 ’ Sı 50 ³10 ʹ W. Pitfall. i-2007. P Lowenberg-Neto [1 Ƌ at CEMT]; Cornélio Procópio. PE Mata São Francisco. 23³9 ʹ 20 ” Sı 50 ³34 ʹ 20 ” W. Pitfall. N Cipola [1 Ƌ at CEMT]; Londrina. 11-xii-1998. IM Medri [2 ♀♀ at CEMT]; same municipality but Mata dos Godoy. 11-xii-1998. J Lopes [2 ƋƋ at CEMT]; Quatro Barras. Engenho Lacerda. 25³20 ʹ 02 ” Sı 48 ³54 ʹ 07 ” W. 873 mosl. Luz. A Specht [1 ♀ at CEMT]; Tibagi. Cânion Guartelá. 10-xi-2007. P Grossi [1 Ƌı 2 ♀♀ at CEMT; Rio de Janeiro: Itaguaí. i-1996. SL Pinto [2 ƋƋ at CEMT]; Nova Friburgo. Campo Coelho. xii-1997. A Penna [1 Ƌ at CEMT]; same municipality but i-1999. P Grossi [1 Ƌ at CEMT]; Rio de Janeiro. Pedra do Rio. 17-x-1992. A Bello [1 ♀ at CEMT]; same data but xii-1993. A Bello [1 Ƌı 3 ♀♀ at CEMT]; same data but ii-2010. A Bello [1 Ƌ at CEMT]; same municipality but Sumaré. 1-xii-1989. A Bello [1 Ƌı 1 ♀ at CEMT]; Teresópolis. PN Serra dos Orgãos. 20-i-1981. A Bello [2 ♀♀ at CEMT]; Rio Grande do Sul: colourado. 12-xi-1993. D Link [2 ƋƋı 2 ♀♀ at CEMT]; Derrubadas. PE Turvo. pitfall. 31-x-2003 [2 ♀♀ at CEMT]; Passo Fundo. 720 m. Luz. xii-1998. Salvadori [1 Ƌı 1 ♀ at CEMT]; Santa Maria. ii-2011. D Link [1 ♀ at CEMT]; Santa Catarina: Anchieta. 26³29 ’ Sı 58 ³18 ʹ W. Fezes humanas. xii-2014 [1 Ƌ at CEMT]; Campos Novos. 27³23 ’ Sı 51 ³12 ʹ W. Pitfall fezes humanas. ii-2010. MIM Hernandéz & G Viegas [1 Ƌ at CEMT]; same data but ii-2014 RC Campos [3 ƋƋı 6 ♀♀ at CEMT]; São Paulo: Cerqueira César. xii-1992. J Carlos [1 ♀ at CEMT]; Guaratinguetá. Rodovia Dutra km 270ı Clube dos 500. 25-27-xii-1994. A Mendes [1 Ƌ at CEMT]; Guarulhos. Cumbica. 2-xi-1968. A Bello [1 ♀ at CEMT]; Piracicaba. 540 m. Campus Esalq. 8-xi-1967. E Berti-Filho [1 Ƌı 1 ♀ at CEMT]; São Paulo. i-1969. A Bello [1 ♀ at CEMT]. PARAGUAY: Guairá: Melgarejo. 10-x-1994. Dreschel [1 Ƌ at CEMT]. Caaguasu: Capiata. 20-xi-2001. C Aguilar [1 ♀ at CEMT];

Redescription (larger males). Colour: blackı shiny. Head: clypeo-frontal and genal surfaces smooth. Clypeo-genal angleı if presentı slightly obtuse – then emarginations present laterally to each clypeal tooth. Interocular space bearing few ocellate punctures posterior to cephalic horn. Cephalic horn with rounded apex. Pronotum: anterior portion bearing sparse ocellate punctures. Anterior angles acuteı bearing few ocellate punctures. Anterior pronotal excavation smooth. Pronotal disc with four aligned lobes. Pronotal disc smoothı shiny. Posterior margin bearing two rows of coarse ocellate punctures and a third row ill-defined of finer punctures. Metasternum: scattered setaeı half the length of those on hypomera. Elytra: striae deeply impressedı ocellate punctures separated by their diameter. Interstriae convexı shinyı bearing fine punctures (viewed under 20x magnification). Abdomen: ocellate punctures along the anterior margin of each ventrite interrupted mediallyı except for 2nd ventriteı where it is continuous. Sixth ventrite with basal sulcus interrupted in the medial portion. Pygidium: smoothı lacking distinct punctures (viewed under 30x magnification)ı swollen. Aedeagus: parameraı dorsally having longitudinal excavations taking 2/3 of paramera length; apexes acuteı convergent. Laterallyı abrupt declivity towards apex; anterior border of longitudinal excavation carinate. Ventrallyı base widely excavated. Subgenital plate with slight basal emarginationı rounded at apex.

Morphological variation. Males BL: 15 – 19 mmı PW: 7 – 10 mm. Smaller males do not have pronotal lobes and their cephalic horn is very reduced into a single tubercle or knob. Howeverı their genitalia and puncture pattern are similar to those of larger males. Females BL: 15 – 18 mmı PW: 7 – 9 mm. Distinguished from males as follows: Head: clypeo-frontal and genal surface transversely wrinkled. Cephalic horn lowı produced in a single tubercle. Pronotum: opaqueı lacking anterior lobes. Elytra: interstriae opaque. Abdomen: sixth ventrite 2ı5x wider than the fifth ventrite. Pygidium: strongly swollen.

Distribution. Argentinaı Boliviaı Brazil (central and south portion) and Paraguay ( Figure 5 ).

Remarks. Pinotus ascanius piceus Lectotype is the most developed male of the syntype seriesı usually larger males have more informative morphology than smaller males and females ( Luederwaldt 1929; Nunes & Vaz-de-Mello 2016). Luederwaldt (1929) ı Pereira (1944) and Martínez (1951) characterised D. bicuspis as having a conspicuous clypeo-genal angleı clearly in disagreement with our type specimen examination and Germar ’ s description of the species ( Germar 1824). This misinterpretation led both H. Lüderwaldt and F. s. Pereira to wrongly consider D. ascanius (and the subspeciesı forms and varieties described by them) lacking clypeo-genal angulation and D. bicuspis having it. Indeedı larger males of D. bicuspis may have a subtle clypeo-genal angulationı while smaller males of D. ascanius will almost lack this. Howeverı the sixth ventrite groove of D. bicuspis is interrupted mediallyı while in males of D. ascanius it is not. We are considering Pinotus (Cephagonus) ascanius aceratus as new synonym of D. bicuspis based on its description. The holotype described by Pereira (1942) does not have a cephalic hornı or it was very low. Howeverı this should not be enough to consider D. ascanius aceratus different from D. bicuspis because it has a similar clypeusı pronotum and aedeagus shapeı according to its description ( Pereira 1944). Dichotomius guaranii was described from a female and placed in the Dichotomius (Luederwaldtinia) nisus group. Howeverı clypeusı ventral hair (mainly metasternum and ventrites) and pygidium and sixth ventrite shape are evidence to place this species near D. bicuspis (Nunes & Vaz-de- Melloı 2013). The examination of the descriptions and photographies of the holotype in the original paper ( Gandini and Aguilar 2009) holotype led us to conclude that this species is indeed a new synonym of D. bicuspis .

Figures

Figure 3. (a) Holotype of Dichotomius (Selenocopris) inachoides (Felsche) (b) Dichotomius (Homocanthonides) smaragdinus (Perty) (c) Holotype of Dichotomius (Dichotomius) prietoi (Martínez).

Figure 5. (a) Ventral view of the open pygidium of a female of D. ascanius: sixth – sixth ventrite, int – intestine;vgn – approximately location of vagina aperture;spt – spermatecha indicated by white arrow.(b) General dorsal aspect of Cephagonus female genitalia, the elytra and abdominal superior ventrites were removed: pyg – pygidium; spt – spermatheca; spdc – spermatheca duct; spgl – spermatheca gland; bcpl – bursa copulatrix. (c) D. ascanius spermatheca. (d) D. ingens spermatheca. (e) D. fissiceps spermatheca.

Figure 12. (a) Copris bicuspis Germar 1824, holotype dorsal habitus. (b) Dorsal habitus of Dichotomius (S.) punctulatipennis (Luederwaldt).