Phanaeus furiosus Bates, 1887

Moctezuma, Victor, Halffter, Gonzalo & Lizardo, Viridiana, 2021, The Phanaeus tridens species group (Coleoptera: Scarabaeoidea): a dung beetle group with genital morphological stasis but a changing ecological niche, Acta Entomologica Musei Nationalis Pragae 61 (2), pp. 447-482 : 461-463

Phanaeus furiosus Bates, 1887

( Figs 19–21 , 27 , 38 , 50 , 61 )

Phanaeus furiosus Bates, 1887: 61 . Type locality: Mexico, Ventanas. Phanaeus furcosus Felsche, 1901: 155 . Unjustified emendation.

Type material examined. MEXICO. L ECTOTYPE (designated by EDMONDS 1994, examined from photographs; Fig. 21 ):♂, Ventanas ( BMNH: NHMUK 013903639).

Non-type material examined. MEXICO: AG UASCAL I ENTES: 1 ♀, Calvillo, Mesa El Roble ( IEXA). G UANA J UATO: 1 ♀, Cueramaro ( IEXA). J AL I SCO: 3 ♀♀, Agua del Obispo ( GHVM); 1 ♂ 4 ♀♀, Ajijic ( GHVM); 1 ♂ 4 ♀♀, Amatitlán ( IEXA); 1 ♀, Chapala ( IEXA); 2 ♂♂ 4 ♀♀, La Calera ( IEXA); 3 ♂♂ 11 ♀♀, Mezquitán ( IEXA); 3 ♂♂ 6 ♀♀, Ponctitlán ( IEXA); 1 ♀: San Miguel de Hidalgo ( IEXA); 10 ♂♂ 12 ♀♀, Trans. Carretera Guadalajara-Colotlán ( GHVM: 7 ♂♂ 7 ♀♀; VMPM: 3 ♂♂ 5 ♀♀); 1 ♀, Zapopan ( IEXA). M I CHOACÁN: 1 ♂, Coalcomán ( GHVM); 1 ♂, without specific locality ( GHVM). N AYAR I T: 1 ♂ 2 ♀♀, El Venado ( IEXA); 1 ♂ 5 ♀♀, Tepic, La Cantera ( IEXA); 1 ♂ 1 ♀, San Blas ( IEXA: 1 ♀, GHVM: 1 ♂); 1 ♂, San José Mojarras, Santa María del Río ( IEXA) 3 ♂♂ 4 ♀♀, Villa Hidalgo ( IEXA); 1 ♂, without specific locality ( GHVM). S I NALOA: 4 ♂♂ 3 ♀♀, Copala ( GHVM); 1 ♀, Corte Alto ( IEXA); 1 ♂ 3 ♀♀, El Zapotillo ( IEXA); 1 ♀, Escuinapa ( GHVM); 1 ♂ 1 ♀, Loberas ( IEXA); 1 ♀, Mazatlán ( GHVM); 2 ♀♀, 48 km N Mazatlán ( IEXA). S ONORA: 2 ♂♂ 1 ♀, 4 km S San Nicolás ( IEXA). Z ACATECAS: 2 ♂♂ 1 ♀, La Manchada ( IEXA: 1 ♂; GHVM: 1 ♂ 1 ♀); 4 ♂♂ 3 ♀♀, San Lorenzo ( GHVM); 5 ♂♂ 14 ♀♀, San Lorenzo de los Mota ( GHVM). U NKNOWN LOCAL I TY: 1 ♂ ( GHVM).

Diagnosis. Metallic green ( Figs 19, 21 ), blue ( Fig. 61 ) or red species ( Fig. 20 ). Sides of pronotal disc granulate ( Figs 19–21 , 50 ). Pronotal disc rugose to lightly granulorugose ( Figs 19–21 , 50 ). Posteromedial process of pronotum produced into thick denticle, short and strongly emarginate apically ( Figs 38 , 50 ). Anteromedial portion of pronotal disc with variable number of denticles and tubercles ( Fig. 50 ). Anterolateral margins of pronotal disc with strongly developed tubercles forming complete ridge ( Fig. 50 ). Posterolateral angles of pronotum fairly short, much shorter that posteromedial process ( Figs 38 , 50 ). Elytral striae scabriculous, distinctly impressed, superficially punctate ( Figs 19–21 ). Elytral interstriae scabriculous, smooth, superficially punctate, convex ( Figs 19–21 ).

Variation. Minor male. Similar to major males, except for reduction of secondary sexual characters (i.e., cephalic horn, pronotal processes and posterolateral angles). Female. Similar to male, except for head showing trituberculate carina; pronotal sculpture granulate to rugose posteriorly; pronotum with anteromedial black macula, and anteromedial carina followed by posterior concavity ( Fig. 61 ).

Three typical colour phases are showed by this species (namely P. furiosus , Figs 19–21 , 50 , 61 ), but it is frequent to found combinations between these colours. EDMONDS (1994) suggested that the colouration of P. furiosus varies geographically.According to him, the green phase ( Figs 19, 21 ) is fixed in the majority of the populations from Sonora, Sinaloa, Nayarit, Jalisco, and Michoacán. On the other hand, the blue phase ( Fig. 61 ) is scarce in Sonora, Sinaloa, Nayarit and western Jalisco; but it becomes common in northern Jalisco and Michoacán. The specimens revised in the present study do not support the geographic variation of colour proposed by EDMONDS (1994). The red chromatic phase ( Fig. 20 ) is fairly common in Jalisco, Zacatecas and Nayarit, but it was omitted by him. Moreover, red ( Fig. 20 ) and green specimens ( Figs 19, 21 ) are found in similar proportions in the aforementioned Mexican states. Additionally, there are some populations of the Pacific slope of Mexico where the blue phase ( Fig. 61 ) almost completely dominates. Nevertheless, it is important to highlight that a geographic cline for colour variation is not recognized nor proposed by us.

Comments. ARNAUD (2002) suggested that P. furiosus included two subspecies, the nominotypic one ( Figs 19–21 , 27 , 38 , 50 , 61 ) and P. furiosus pseudofurcosus ( Figs 22–23 , 39 , 51 , 62 ). EDMONDS (1994) and EDMONDS & ZÍDEK (2012) treated P. furiosus as a monotypic species instead, regarding the populations that ARNAUD (2002) had classified in P. furiosus pseudofurcosus as an odd disjunct population of P. tridens tridens . But despite this classification, EDMONDS & ZÍDEK (2012) confusingly cited P. pseudofurcosus as a junior subjective synonym of P. tridens instead of P. furiosus ( Figs 1–5 , 32 , 43 , 55 , 63 ); this was likely a lapse. Our conclusion, however, is that P. pseudofurcosus is neither a subspecies of P. furiosus nor a synonym of either P. furiosus or P. tridens , but a fully independent species (see comments below). Phanaeus furiosus , in turn, is indeed a monotypic species (i.e., a taxonomically “homogeneous” species that does not comprise different subspecies according to ZACHOS 2016) as suggested by EDMONDS (1994) and EDMONDS & ZÍDEK (2012). In spite of its several chromatic phases ( Figs 19–21 , 50 , 61 ), P. furiosus is a fairly distinctive species that has the posteromedial process of pronotum in major males produced into a thick denticle, short and strongly emarginate apically ( Figs 38 , 50 ).

Distribution. Mexico: Durango, Guanajuato, northern to central Jalisco, Michoacán, Nayarit, Sinaloa, Sonora and Zacatecas. This is the most widely distributed species within the P. tridens species group ( Fig. 66 ).

Figures

Figs 19–21. Phanaeus furiosus Bates, 1887. 19 – male green phase; 20 – male red phase; 21 – lectotype and labels (by Mario Cupello, UFPR). Scale bar = 1.0 mm.

Figs 27–28. Phanaeus victoriae Moctezuma sp. nov. 27 – holotype male; 28 – paratype female. Scale bar = 1.0 mm.

Figs 32–42. Posterior view of pronotum of major male. 32 – P. tridens Castelnau, 1840; 33 – P. moroni Arnaud, 2001, stat. rev.; 34 – P. bal- thasari Arnaud, 2001, stat. rev.; 35 – P. daphnis Harold, 1863; 36 – P. substriolatus Balthasar, 1939, stat. rev.; 37 – P.herbeus Bates, 1887, stat. rev.; 38 – P.furiosus Bates, 1887; 39 – P.pseudofurcosus Balthasar, 1939, stat. rev.; 40 – P. nimrod Harold, 1863; 41 – P. victoriae Moctezuma sp. nov.; 42 – P. eximius Bates, 1887. Scale bar = 1.0 mm.

Figs 43–54. Lateral view of pronotum of major male. 43 – P. tridens Castelnau, 1840; 44 – P. moroni Arnaud, 2001, stat. rev.; 45 – P. balthasari Arnaud, 2001; 46 – P. daphnis Harold, 1863; 47 – P. coeruleus Bates, 1887, stat. rev. (holotype, by Keita Matsumoto, BMNH); 48 – P. substriolatus Balthasar, 1939, stat. rev.; 49 – P. herbeus Bates, 1887, stat. rev. (green-red phase); 50 – P. furiosus Bates, 1887 (green phase); 51 – P. pseudofurcosus Balthasar, 1939, stat. rev.; 52 – P. nimrod Harold, 1863; 53 – P. victoriae Moctezuma sp. nov. (holotype); 54 – P. eximius Bates, 1887. Scale bar = 1.0 mm.

Figs 55–62. Pronotum of female. 55 – P. tridens Castelnau, 1840; 56 – P. balthasari Arnaud, 2001, stat. rev.; 57 – P. daphnis Harold, 1863; 58 – P. substriolatus Balthasar, 1939, stat. rev. (dark blue-black phase); 59 – P.herbeus Bates, 1887, stat. rev. (green phase); 60 – P. nimrod Harold, 1863 (blue-green phase); 61 – P. furiosus Bates, 1887 (dark blue phase); 62 – P. pseudofurcosus Balthasar, 1939, stat. rev. Scale bar = 1.0 mm.

Figs 22–23.Phanaeus pseudofurcosus Balthasar, 1939, stat.rev.22 – male green phase;23 – holotype and labels (by Jiří Hájek,NMPC).Scale bar = 1.0 mm.

Figs 1–5. Phanaeus tridens Castelnau, 1840. 1 – male green phase; 2 – male green-red phase; 3 – neotype and labels, present designation (by Christophe Rivier, MNHN); 4 – P. frankenbergeri junior subjective synonymy, holotype and labels (by Jiří Hájek, NMPC); 5 – Eൽආඈඇൽඌ’ (1994) neotype of P. tridens and labels (by Simon Hinkley, NMVA). Scale bar = 1.0 mm.

Fig.63. Phallobase, parameres and endophallite copulatrix of the P.tridens species group. Scale bar = 1.0 mm.

Fig. 66. Predicted distribution of P. furiosus Bates, 1887 and P. pseudofurcosus Balthasar, 1939.