Homocopris torulosus

Homocopris torulosus (Eschscholtz, 1822) Figs 9–10, 21–22, 29, 35, 38, 41, 45, 47, 49, 54, 61, 67 Copris torulosa Eschscholtz, 1822: 29 (original description). Copris valdiviana Philippi, 1859: 661 (original description). Pinotus torulosus var. minor Luederwaldt, 1925: 68 (original description) (uncertain status, see remarks below). Pinotus Dahli Landin, 1955: 5 (original description) syn. nov. Copris (Homocopris) torulosa – Burmeister 1846: [3] (mention). Copris torulosa – Solier 1851: 60 (monograph). Copris torulosus – Redtenbacher 1868: 55 (distribution). Pinotus torulosus – Harold 1869a: 133 (monograph); 1869b: 1011 (catalogue). — Waterhouse 1891: 361 (comment). — Bruch 1911: 187 (catalogue). — Gillet 1911: 62 (catalogue). — Joseph 1929: 31 (nidification). — Gutiérrez 1940: 277 (monograph). — Pereira 1941: 267 (checklist). — Blackwelder 1944: 208 (checklist). — Lange 19476: 313 (misidentification). [Copris] valdivianus – Harold 1869b: 1011 (synonymy). — Heyne 1900: 62 (synonym). — Gillet 1911: 62 (synonym). Copris valdivianus – Harold 1869a: 133 (synonym). — Reed 1876: 281 (synonym). — Bruch 1911: 187 (synonym). [Pinotus] torulosus – Reed 1876: 281 (catalogue). — Heyne 1900: 62 (diagnosis, distribution). Pinotus valdivianus – Waterhouse 1891: 361 (comment). Pinotus (P.) torulosus var. valdivianus – Luederwaldt 1929: 649 (monograph). Pinotus torulosus var. Minor – Luederwaldt 1929:649 (monograph).— Gutiérrez1940: 280 (monograph). — Blackwelder 1944: 208 (checklist). — Mondaca 2023: 223 (synonym). Pinotus (P.) torulosus – Luederwaldt 1929: 650 (monograph). Pinotus torulosus var. valdivianus – Gutiérrez 1940: 279 (monograph). — Blackwelder 1944: 208 (checklist). Pinotus torulosus minor – Lange 1947: 313 (biology, distribution). Dichotomius (D.) torulosus – Martínez 1959: 87 (catalogue). Dichotomius torulosus – Halffter & Matthews 1966: 7 (biology). — Halffter 1977: 242 (nidification). — Halffter & Edmonds 1982: 89 (biology). — Zunino 1983: 541 (classification). — Klemperer 1983: 61 (ecology). — Cabrera Walsh & Gandolfo 1996: 586 (nidification). [Dichotomius (D.) torulosus] var. minor – Vaz-de-Mello 2000: 193 (checklist). [Dichotomius (D.) torulosus] var. valdivianus – Vaz-de-Mello 2000: 193 (checklist). Homocopris torulosus – Vaz-de-Mello et al. 2010: 192 (new combination). — Jerez 2015: 1 (biology). — Ratcliffe et al. 2015: 197 (checklist). — González et al. 2015: 53 (ecology). — Tarasov & Génier 2015: 21 (phylogeny). — Tarasov & Dimitrov 2016: 7 (phylogeny). — Rebolledo et al. 2017: 27 (faunistic). — Tello et al. 2017: 61 (fossil); 2021: 139 (distribution). — Mondaca 2023: 223 (checklist, distribution). Copris valdiviana – Mondaca 2023: 223 (synonymy). Pinotus dahli – Mondaca 2023: 223 (synonym). Differential diagnosis Dorsal ocular width subequal to one-sixth interocular distance; lateral pronotal edge angular on anterior fourth in dorsal view; width of ellipse formed by lateral pronotal carina and edge approximately one third the elliptical length; posterior pronotal surface chagrined; elytral interstriae punctation visible at 10× magnification; metasternal depression entire, extending into anterior portion of median metasternal lobe. ♂ pronotal ridge with three sets of distinct bilaterally paired tubercles in large individuals; paramere apex acutely tipped; FLP endophallite elongate, bent, with internal angle adjacent to sclerotized extension. Name-bearing type data Copris torulosus Eschscholtz, neotype ♂, here designated (CMNC) (Fig. 54): “ CHILE: Malleco | 17kmWAngol, 800m | 8.XII.84 - 16.II.85 | S & JPeck, FIT | mixed Nothofagus ”; “WORLD | SCARAB. | DATABASE | WSD00035423” [barcode label]; “Southern Neotropical Scarabs | database # AS2599338 | Homocopris torulosus | (Eschscholtz, 1822) ♂ | DET: A.B.T.Smith 2005“ [with black border]; “N E O T Y P E ♂ | Copris | torulosus | Eschscholtz, 1822 | des. Génier & Darling, 2021” [on red card with black border.]. Pinotus torulosus var. minor Luederwaldt: no type material examined. Pinotus dahli Landin, allotype ♂ (MZLU) (Fig. 57): “ Chile ” [handwritten]; “Allotypus” [red card with black border]; “ Pinotus Dahli m. | allotype ♂ | B. O. Landin det” [partly handwritten]; “Photo 2018 | by MZLU”; “MZLU | Type no | 2775:2” [partly red]; “MZLU | 2018 | 001” [green paper]; “ZML.2005 | 202” [green paper]; “WORLD | SCARAB. | DATABASE | WSD00035268” [barcode label]; “Southern Neotropical Scarabs | database # AS2607242 | Homocopris torulosus | (Eschscholtz, 1822) ♂ | DET: A.B.T.Smith 2006”; “ Homocopris ♂ | torulosus | (Eschscholtz, 1822) | det. Génier & Darling, 2018”. Type locality 17 km W Angol (800 m), Malleco, Chile. Type material Neotype of Copris torulosus (present designation) CHILE • ♂ (Fig. 54); “ CHILE: Malleco | 17kmWAngol, 800m | 8.XII.84 - 16.II.85 | S & JPeck, FIT | mixed Nothofagus ”; “WORLD | SCARAB. | DATABASE | WSD00035423” [barcode label]; “Southern Neotropical Scarabs | database # AS2599338 | Homocopris torulosus | (Eschscholtz, 1822) ♂ | DET: A.B.T.Smith 2005“ [with black border]; “N E O T Y P E ♂ | Copris | torulosus | Eschscholtz, 1822 | des. Génier & Darling, 2021” [on red card with black border]; CMNC. Allotype of Pinotus dahli CHILE • ♂ (Fig. 57); “Chile” [handwritten]; “Allotypus” [red card with black border]; “ Pinotus Dahli M. | allotype ♂ | B. O. Landin det” [partly handwritten]; “Photo 2018 | by MZLU”; “MZLU | Type no | 2775:2” [partly red]; “MZLU | 2018 | 001” [green paper]; “ ZML.2005 | 202” [green paper]; “WORLD | SCARAB. | DATABASE | WSD00035268” [barcode label]; “Southern Neotropical Scarabs | database # AS2607242 | Homocopris torulosus | (Eschscholtz, 1822) ♂ | DET: A.B.T.Smith 2006”; “ Homocopris ♂ | torulosus | (Eschscholtz, 1822) | det. Génier & Darling, 2018”; MZLU. Paratype de Pinotus dahli CHILE • 1 ♀; same locality as for allotype; [no date]; [anonymous]; MZLU. Other material examined (163 ♂♂, 206 ♀♀) ARGENTINA – Río Negro • 1 ♂; El Bolsón; [41°58′ S, 71°32′ W]; Nov. 1967; [anonymous]; CPFA • 1 ♀; same locality as for preceding; [no date]; A. Kovács; NMPC • 1 ♀; same locality as for preceding; 1956; A. Kovács; NMPC • 1 ♂; same locality as for preceding; Dec. 1956; [anonymous]; NMPC. CHILE • 6 ♂♂, 1 ♀; [unspecified locality]; 1907; M.J. Rivera; MNHN • 1 ♂; same locality as for preceding; [no date]; M.J. Rivera; CMNC. – Región de Los Lagos • 1 ♂, 2 ♀♀; 11 km NW of Castro, Isla Chiloé; [42°24′ S, 73°51′ W]; 200 m; 27 Dec. 1984 – 1 Feb. 1985; S. and J. Peck; ravine second growth forest, carrion trap; CMNC • 1 ♂, 9 ♀♀; 13 km W of Entre Lagos; [40°37′30″ S, 72°44′0″ W]; 200 m; 21 Dec. 1984 – 8 Feb. 1985; S. and J. Peck; forest remnant, carrion trap; CMNC • 2 ♀♀; 34 km E of Puerto Montt, Lago Chapo; [41°25′ S, 72°35′ W]; 300 m; 24 Dec. 1984 – 2 Feb. 1985; S. and J. Peck; second growth Nothofagus, carrion trap; CMNC • 1 ♂; Ahori Alto; 21–23 Feb. 1988; 70 m; L. Masner; CNC • 1 ♀; Repucura forest trail, Anticura, Parque Nacional Puyehue; [40°40′ S, 72°10′30″ W]; 500 m; 19 Dec. 1984 – 6 Feb. 1985; S. and J. Peck; forest, carrion trap; CMNC • 1 ♂; Terao, 10 km S of Chonchi, Isla Chiloe; [42°43′1″ S, 73°38′47″ W]; 45 m; 12–13 Feb. 2005; [anonymous]; yellow pan trap; CMNC • 1 ♂; [unspecified locality]; 30 Jan. 2007; [anonymous]; CMD • 1 ♀; same locality as for preceding; 10 Feb. 2007; [anonymous]; CMD. – Región de Los Ríos • 3 ♂♂, 3 ♀♀; 34 km WNW of La Union; [40°13′45″ S, 73°21′35″ W]; 700 m; 17 Dec. 1984 – 7 Feb. 1985; S. and J. Peck; mixed evergreen forest, carrion trap; CMNC • 3 ♂♂; Corral; [39°53′ S, 73°26′ W]; 14 Oct. 1913; R.H. Beck; CAS • 1 ♂; same locality as for preceding; 16 Oct. 1913; R.H. Beck; CAS • 1 ♂, 1 ♀; same locality as for preceding; 20 Oct. 1913; R.H. Beck; CAS • 1 ♂; Panguipulli; [39°38′20″ S, 72°20′30″ W]; Jan. 1952: [anonymous]; CMNC • 1 ♂; same locality as for preceding; 1927; [anonymous]; CAS • 2 ♂♂, 3 ♀♀; Valdivia; [39°50′ S, 73°13′ W]; [no date]; [anonymous]; MNHN. – Región del Biobío • 1 ♀; 4 km SE of Recinto; [36°52′ S, 71°36′ W]; 700 m; 17 Dec. 1976; H.F. Howden; CMNC • 2 ♀♀; 40 km E of San Carlos; [36°33′ S, 71°34′ W]; 24 Dec. 1950; Ross and Michelbacher; CAS • 5 ♂♂, 4 ♀♀; 60 km SE od Chillan, Thermas Road; [36°54′ S, 71°33′ W]; 1300 m; 7 Dec. 1984 – 19 Feb. 1985; S. and J. Peck; beech forest, flight interception trap; CMNC • 1 ♂; Arauco; [37°15′ S, 73°19′ W]; 15 Apr. 1965; Sanfeliu; CMNC • 1 ♀; Concepción; [36°48′ S, 73°1′ W]; [no date]; [anonymous]; CAS • 1 ♂, 1 ♀; Contulmo; [38°1′ S, 73°14′ W]; 28 Mar. 1940; D.S. Bullock; CAS • 1 ♀; same locality as for preceding; 7 Jan. 1967; Sanfeliu; CNC • 1 ♂; same locality as for preceding; 10 Feb. 1967; T. Cekalovic; CNC • 7 ♂♂, 4 ♀♀; same locality as for preceding; 10 Mar. 1969; Sanfeliu; CNC • 7 ♂♂, 1 ♀; same locality as for preceding; 20 Dec. 1969; T. Cekalovic; CNC • 1 ♀; Hulpencillo; [36°47′ S, 73°5′30″ W]; 20 Mar. 1964; T. Cekalovic; CNC • 2 ♂♂, 2 ♀♀; Invernada; [36°52′30″ S, 71°34′30″ W]; Mar. 1970; L.E. Peña; PMOC • 1 ♂, 1 ♀; Las Trancas; [36°54′ S, 71°40′ W]; 1 Feb. 1974; L.E. Peña; PMOC • 1 ♂, 1 ♀; Las Trancas, Curico; [36°54′ S, 71°30′ W]; 1 Mar. 1979; L.E. Peña; CMNC • 1 ♂; Los Castaños, Chillán, El Carmen; [36°37′30″ S, 72°5′0″ W]; 21 Nov. 2003; M. Snižek; NMPC • 1 ♂; Mulchén; [37°43′ S, 72°14′ W]; Jan. 1935; Gutiérrez; CMNC • 2 ♂♂; same locality as for preceding; Dec. 1935; Gutiérrez; CMNC • 3 ♂♂, 1 ♀; Recinto; [36°48′ S, 71°44′ W]; 1 Mar. 1971; [anonymous]; CPFA • 2 ♂♂, 2 ♀♀; Talcahuano; [36°45′ S, 73°6′ W]; 1841; Jacquinot; MNHN. – Región del Maule • 1 ♂; Fundo Malcho, Cordillera Parral; [36°19′ S, 71°25′ W]; Dec.1956; M. Rivera; CNC • 2 ♂♂, 2 ♀♀; same locality as for preceding; May 1957; [anonymous]; CPFA • 1 ♀; same locality as for preceding; 1 May 1957; [anonymous]; CPFA • 1 ♂; same locality as for preceding; 11–20 Nov. 1964; M. Rivera; CNC • 2 ♀♀; Romehual, Cordillera Parral; 5–10 Nov. 1960; [anonymous]; CPFA • 1 ♂, 3 ♀♀; Vilches Alto, Talca; [35°36′ S, 71°4′ W]; Dec. 1973; J. Valencia; CMNC. – Región La Araucanía • 2 ♂♂; 10 mi NE of Pucón; [39°

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