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

Pinotus torulosus var. minor Luederwaldt: no type material examined.

Type locality

17 km W Angol (800 m), Malleco, Chile.

Type material

Neotype of Copris torulosus (present designation)

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°15′30″ S, 71°46′0″ W]; 12 Jan. 1951; Ross and Michelbacher; CAS • 7 ♂♂, 7 ♀♀; 11 km W of Angol; [37°49′30″ S, 72°47′0″ W]; 1000 m; 9 Dec. 1984 – 16 Feb. 1985; S. and J. Peck; boggy mixed forest remnant, carrion trap; CMNC • 1 ♂, 1 ♀; 15 km W of Victoria; [38°15′ S, 72°31′ W]; 200 m; 28–31 Dec. 1976; S. Peck; dung; BDGC • 6 ♂♂, 11 ♀♀; same locality as for preceding; 28–31 Dec. 1976; S. Peck; carrion; CMNC • 10 ♂♂, 27 ♀♀; same locality as for preceding; 28–31 Dec. 1976; S. Peck; dung; CMNC • 1 ♂; same locality as for preceding; 28–31 Dec. 1976; S. Peck; carrion; CNC • 21 ♂♂, 17 ♀♀; same locality as for preceding; 28–31 Dec. 1976; S. Peck; dung; CNC • 2 ♀♀; same locality as for preceding; 28 Dec. 1976; H.F. Howden; CNC • 1 ♀ (alloreferent of Copris torulosus), 5 ♂♂, 8 ♀♀; 17 km W of Angol; [37°48′ S, 72°51′30″ W]; 800 m; 8 Dec. 1984 – 16 Feb. 1985; S. and J. Peck; mixed Nothofagus, flight interception trap; CMNC • 2 ♀♀; 20 km E of Manzanar; [38°26′30″ S, 71°31′0″ W]; 1100 m; 19–25 Dec. 1976; S. Peck; dung; CNC • 1 ♂; 20 km E of Temuco; [38°46′ S, 72°22′ W]; 8 Jan. 1951; Ross and Michelbacher; CAS • 1 ♂, 2 ♀♀; 21 km NE of Pucón, Lago Caburga; [39°11′ S, 71°49′ W]; 600 m; 15 Dec. 1984 – 10 Feb. 1985, S. and J. Peck; mixed forest remnant, flight interception trap; CMNC • 2 ♂♂, 1 ♀; 22 km E of Temuco; [38°40′ S, 72°15′30″ W]; Jun.–Jul. 1951; M.G. Smith; CAS • 1 ♂; same locality as for preceding; 2 Jan. 1951; Ross and Michelbacher; CAS • 1 ♂, 1 ♀; 30 km NE of Villarrica; [39°8′47″ S, 72°3′32″ W]; Jan. 1965; L. Peña; CAS • 1 ♀; same locality as for preceding; 16–31 Dec. 1964; L. Peña; CAS • 1 ♀; 4 km W of Victoria; [38°14′ S, 72°24′30″ W]; 300 m; 26–27 Dec. 1976; H.F. Howden; CNC • 3 ♂♂, 6 ♀♀; 40 km W of Angol, Parque Nacional Nahuelbuta; [37°49’30″ S, 73°0′30″ W]; 1200– 1500 m; 9 Dec. 1984 – 17 Feb. 1985; S. & J. Peck; flight interception trap; CMNC • 1 ♀; 6 km W of Curacautín; [38°25′ S, 71°58′ W]; 750 m; 12 Dec. 1984 – 16 Feb. 1985; S. and J. Peck; grazed Nothofagus remnant, carrion trap; CMNC • 1 ♀; 9 km S of Pucón, Parque Nacional Volcán Villarrica; [39°22′ S, 71°57′30″ W]; 900 m; 15 Dec. 1984 – 10 Feb. 1985; S. and J. Peck; Nothofagus grove on ash, carrion trap; CMNC • 1 ♀; along road near Río Picoiquén, Cotoco, Parque Nacional Nahuelbuta; [37°45.812′ S, 72°59.314′ W]; 980 m; 15 Feb. 2004; A. Smith, M. Paulsen and J. Mondaca; CMNC • 1 ♀; Boroa Central, SE Nueva Toltén; [39°16′ S, 73°8′ W]; 22 Jan. 2004; M. Snižek; NMPC • 1 ♀; Collipulli; [37°57′ S, 72°26′ W]; 15 Feb. 1967; Mendez; CNC • 1 ♀; Crest of Sierra Nahuelbuta, W of Angol; [37°50′ S, 72°52′ W]; 1200 m; 13 Jan. 1951; [anonymous]; CAS • 2 ♀♀; Curacautín; [38°26′ S, 71°53′ W]; 20 Jul. 1950; [anonymous]; CAS • 3 ♀♀; same locality as for preceding; 7 Apr. 1971; P. Ramírez; CMNC • 3 ♂♂, 8 ♀♀; Flor del Lago, 15 km NE of Villarrica; [39°10′ S, 72°10′ W]; 300 m; 14 Dec. 1984 – 10 Feb. 1985; S. and J. Peck; Nothofagus forest, carrion trap; CMNC • 6 ♂♂, 13 ♀♀; same locality as for preceding; 14 Dec. 1984 – 10 Feb. 1985; S. and J. Peck; Nothofagus forest, flight interception trap; CMNC • 1 ♀; Manzanar; [38°28.071′ S, 71°41.873′ W]; 800 m; 1–3 Feb. 2005; S. Bilý; NMPC • 8 ♂♂, 7 ♀♀; Monumento Natural Contulmo, Purén; [38°1′ S, 73°10′ W]; 350 m; 11 Dec. 1984 – 13 Feb. 1985; S. and J. Peck; mixed evergreen forest, flight interception trap; CMNC • 1 ♀; Parque Nacional Nahuelbuta; [37°48′ S, 73°0′ W]; 1200 m; 9 Sep. 1966; E.I. Schlinger and E.I. Erwin; Nothofagus and Araucaria association; CAS • 1 ♀; Pucón; [39°16′ S, 71°58′ W]; 9–16 Nov. 1989; S. A. Marshall; dung trap near lake; CMNC • 1 ♀; Quepe; [38°52′ S, 72°37′ W]; 31 Oct. 1965; T. Cekalovic; CNC • 2 ♀♀; S Melipeuco; [38°52′ S, 71°41′30″ W]; 23 Jan. 2004; M. Snižek; NMPC • 4 ♂♂, 5 ♀♀; Salto de la Princesa, 20 km E of Curacautín; [38°28′30″ S, 71°40′30″ W]; 1000 m; 12 Dec. 1984 – 16 Feb. 1985; S. and J. Peck; Nothofagus forest, flight interception trap; CMNC • 3 ♂♂, 2 ♀♀; Selva Obscura; [38°22′ S, 72°10′ W]; 6 Oct. 1967; C. Oyarzo; CNC • 2 ♂♂, 2 ♀♀; Sendero Lemu Mau, Monumento Natural Contulmo; [38°74′ S, 73°11.13′ W]; 410 m; 8–24 Dec. 2002; Thayer and Newton; (FMHD #2002-062); Nothofagus obliqua -Eucryphia cordifolia with fern and bamboo understory, carrion trap, octopus; CMNC • 1 ♀; Temuco; [38°45′ S, 72°36′ W]; Jan. 1906; [anonymous]; CAS • 1 ♂; same locality as for preceding; Jan. 2004; [anonymous]; CMD • 1 ♂; same locality as for preceding; Feb. 2002; [anonymous]; CMD • 4 ♂♂, 4 ♀♀; same locality as for preceding; 16 Nov. 1926; [anonymous]; CAS • 1 ♀; Termas del Río Blanco, Curacautín; [39°6′30″ S, 71°37′0″ W]; 3 Feb. 2004; M. Snižek; NMPC • 1 ♂; Victoria; [38°14′ S, 72°21′ W]; [no date]; [anonymous]; MNHN • 1 ♀; same locality as for preceding; [no date]; F. Schneider; NMPC.

Description

Male neotype (Figs 9, 21, 29, 35, 38, 47, 49)

MEASUREMENTS. Body length 13.9 mm.

HEAD. Clypeus bidentate with broad, rounded teeth. Anteroventral clypeal tooth broad. Clypeal and genal surface rugose, clypeogenal suture demarcated by shallow notch. Posterior genal angle obtuse. Frontal surface reduced, smooth. Dorsal ocular width subequal to one-sixth interocular distance. Interocular surface medially smooth. Cephalic horn emerging centrally from clypeofrontal surface, tapering apically, straight, 2.9 mm in length.

PROTHORAX. Anterior pronotal edge medially indented. Lateral pronotal edge angular on anterior fourth in dorsal view. Anterolateral pronotal lobe surface with irregular transverse rugulae. Anterior pronotal declivity surface (Fig. 49) with transverse irregular, fused punctures. Pronotal ridge with three sets of distinct bilaterally paired tubercles. Posterior pronotal surface chagrined. Prosternal apex acutely tipped.

ELYTRA (Fig. 47). Glossy overall. Striae 1–6 widest anteriorly; narrowing posteriorly. Third and fourth striae posteriorly joined. Eighth stria absent throughout. Ninth stria effaced along anterior quarter of elytral length. Interstriae convex, minutely punctate.

HINDWING (Fig. 41). Membrane edge between AP and AA sinuous. AP vein entire, converging with J.

VENTRITES. Meso-metasternal suture straight. Median lobe of metasternum (Fig. 45) with deep impression extending throughout the entire length. Lateral metasternal lobe sparsely setose. Fifth abdominal ventrite with dense medial punctation. Pygidium coarsely punctate.

PROTHORACIC LEGS. Anterior and posterior surfaces of profemur with sparse row of long dark setae. Ventral profemoral surface finely punctate. Protibia with three distinct teeth. Protibial forespur parallel-sided, curving downward apically.

MESOTHORACIC LEGS. Posterior surface of mesotrochanter with three dark setae. Anterior and posterior surfaces of mesofemur with sparse row of long dark setae. Outer surface of mesotibia serrate. Width at mesotibial apex one-third mesotibial length.

METATHORACIC LEGS. Posterior surface of metatrochanter with three dark setae. Anterior and posterior surfaces of metafemur with sparse row of long dark setae. Outer surface of metatibia serrate. Width at metatibial apex one-third metatibial length.

MALE GENITALIA. Aedeagus length: 4.1 mm. Medially paired sclerites of genital segment short (Fig. 38). Paramere (Fig. 29) with apex acutely tipped. FLP endophallite (Fig. 35) elongate, bent, with internal angle adjacent to sclerotized extension.

Female alloreferent specimen (Figs 10, 22)

Similar to male with the following exceptions: clypeal teeth separated by V-shaped notch. Clypeofrontal carina width one-third interocular distance. Entire anterior pronotal surface with irregular, fused rugosities. Anterior pronotal ridge low, followed by shallow concavity. Protibial forespur apex sharply tapered, curving inward.

Variation

Body length 13–20 mm. Small males with medially projecting clypeofrontal ridge instead of cephalic horn. Pronotal ridge in small males low, medially notched.

Distribution (Fig. 67)

Chile in Región de los Lagos, Región de Los Ríos, Región de Valparaíso, Región del Biobío, Región del Maule, Región la Araucanía, Región Metropolitana de Santiago) and Argentina (Río Negro). Recorded elevations vary from 45 m to 1500 m. Rebolledo et al. (2017) report an elevation range from 0 m to 2000 m.

Natural history

Examined specimens were collected in beech forest, boggy mixed forest remnants, forest remnants, grazed Nothofagus remnants, mixed evergreen forest, mixed forest remnants, mixed Nothofagus, Nothofagus and Araucaria Juss. association, Nothofagus forest, Nothofagus grove on ash, Nothofagus obliqua –Eucryphia cordifolia with fern and bamboo understorey, second growth ravine forest and second growth Nothofagus . Attracted to pitfall traps baited with unspecified carrion and octopus carrion, unspecified dung and flight interception traps. A single specimen collected in a yellow pan trap.

Remarks

Copris valdivianus Philippi, 1859 was described from an unspecified number of male and female syntypes. This name was synonymized by Harold (1869a) under Pinotus torulosus (= H. torulosus), which he also considered a senior synonym of Copris punctatissimus . Luederwaldt (1929) later treated C. valdivianus as a variety of P. torulosus . José Mondaca recently published a checklist of Scarabaeoidea of Chile (Mondaca 2023) in which C. valdivianus is synonymized with H. torulosus . This synonymy is based on a putative male syntype deposited in the Paris Museum. “It is very probable that the specimen from Paris is the syntype of C. valdiviana since the specimen studied by Philippi is not found in the MNNC- Santiago. It is also not in the Prague Museum which also has some types of Philippi. It is probable that the possible syntype reached the Paris Museum through Philibert Germain. The locality written on the original label agrees with what was indicated by Philippi in the original publication and the spelling printed on the original label also agrees with that of the author of the species, therefore this only known specimen closest to the syntype that we have available.” (José Mondaca pers. com.).

The type material of Pinotus dahli Landin, 1956 (holotype ♀, allotype ♂) was studied. We were unable to find any differences supporting the validity of this species. We therefore propose the following new synonymy, the second epithet being valid: Pinotus dahli Landin, 1956 = Homocopris torulosus (Eschscholtz, 1822) syn. nov.

We were unable to study type material of Pinotus torulosus var. minor Luederwaldt, 1925 . Gutiérrez (1940) state that two males and four females from Chile and a single male specimen from Espírito Santo, Brazil, are deposited in the MZSP. The syntype series is probably a mixed series of H. torulosus and H. grossiorum if properly labelled. We have requested images of these syntypes, and we should be able to confirm the synonymy with H. torulosus once we obtain them.