Phanaeus daphnis Harold, 1863

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 : 455-457

Phanaeus daphnis Harold, 1863

( Figs 11–13 , 35 , 46 , 57 )

Phanaeus daphnis Harold, 1863: 165 . Type locality: Mexico.

Type material examined. MEXICO: Lൾർඍඈඍඒඉൾ (designated by Aඋ- ඇൺඎൽ 1982a, examined from photographs; Fig. 12 ): J, without specific locality ( MNHN: MNHN EC10558). Pൺඋൺඅൾർඍඈඍඒඉൾ (examined from photographs): J, without specific locality ( MNHN: MNHN EC10559). Non-type material examined. MEXICO: Eඌඍൺൽඈ ൽൾ Mඣඑංർඈ: 1 J 7 ♀♀, Chalma ( GHVM: 1 J 6 ♀♀; VMPM: 1 ♀); 11 ♀♀, Zona arqueológica Malinalco ( GHVM: 10 ♀♀; VMPM: 1 ♀). Gඎൾඋඋൾඋඈ: 2 ♀♀, Acahuizotla ( IEXA); 1♀, Coapango ( VMPM); 5JJ 2♀♀, Hueyecantenango, Chilapa ( GHVM); 11JJ 30♀♀, Mochitlán,Acahuizotla ( IEXA); 2 JJ, Taxco ( GHVM:1J; VMPM: 1J); 1♀, 2 mi. W Taxco ( GHVM); 2 JJ 1 ♀, Tetipac ( IEXA). Mඈඋൾඅඈඌ: 3 JJ, Alpuyeca ( GHVM); 3 JJ 2 ♀♀, Cuautla ( GHVM); 8 JJ 12 ♀♀, Cuernavaca ( IEXA: 5 JJ 8 ♀♀; GHVM: 3 JJ 4 ♀♀); 1 ♀, 9 mi. S Cuernavaca ( GHVM); 2 JJ 1 ♀, Huautla ( IEXA); 1 J, Jojutla, Cerro del Higuerón ( IEXA); 1 J 1 ♀, Oaxtepec ( GHVM); 1J, Palmiras, Cuernavaca ( GHVM); 1 ♀, Progreso ( GHVM); 2 JJ 1 ♀, Puente de Ixtla ( GHVM: 1 J; VMPM: 1 J 1 ♀); 1 J, Real del Puente ( GHVM); 26 JJ 7 ♀♀, Tepoztlán ( GHVM: 21 JJ 6 ♀♀; VMPM: 5 JJ 1 ♀); 1 ♀, Tequesquitengo ( GHVM); 3 JJ 5 ♀♀, Xochitepec ( GHVM: 3 JJ 4 ♀♀, VMPM: 1 ♀). Oൺඑൺർൺ: 1 J, Oaxaca ( IEXA). Pඎൾൻඅൺ: 2 ♀♀, 9 mi. N Amatitlán ( GHVM); 1 J, Matamoros ( VMPM). UඇKඇඈඐඇ අඈർൺඅංඍඒ: 2 JJ ( GHVM).

Diagnosis. Typically metallic green ( Figs 10, 12 , 46 , 57 ), rarely deep blue-green ( Fig. 11 ). Sides of pronotal disc granulate anteriorly, becoming punctate with effaced granules posteriorly ( Figs 10–11 , 46 ). Pronotal disc coarsely rugose ( Figs 10–11 , 46 ). Posteromedial process of pronotum produced into fairly short denticle, not bifurcated apically ( Figs 35 , 46 ). Anteromedial portion of pronotal disc with line of denticles or carina ( Fig. 46 ). Anterolateral margins of pronotal disc without ridge or line of tubercles ( Fig. 46 ). Posterolateral angles of pronotum much longer than posteromedial process of pronotum ( Figs 35 , 46 ). Elytral striae scabriculous, distinctly impressed, superficially punctate ( Figs 10–12 ). Elytral interstriae scabriculous, superficially punctate, flat; I, III, V, VI and VII frequently with lightly roughened integument ( Figs 10–12 ).

Variation. Minor male. Similar to major males, except for reduction of secondary sexual characters (i.e., cephalic horn, pronotal processes and posterolateral angles). Particularly, posterolateral angles of pronotum are reduced into a couple of pronotal keels, while posteromedial process is obsolete. Female. Similar to male, except for head showing trituberculate carina; pronotal sculpture granulate, becoming coarsely punctate posteriorly; pronotum with anteromedial black macula, and anteromedial trapezoidal carina, followed by posterior concavity ( Fig. 57 ).

Comments. Eൽආඈඇൽඌ (1994) and Eൽආඈඇൽඌ & ZටൽൾK (2012) proposed that P. coeruleus ( Figs 13, 47 ) and P. herbeus ( Figs 16–18 , 37 , 49 , 59 ) were junior subjective synonyms of P. daphnis ( Figs 11–13 , 35 , 46 , 57 ). In turn, Aඋඇൺඎൽ (2002) considered both subspecies of P. daphnis . According to the findings of the present study, both P. coeruleus and P. herbeus merit full species status. Major males of P. daphnis never show the posteromedial process of pronotum produced into a distinctly elongate denticle ( Figs 35 , 46 ), while those of P. coeruleus ( Fig. 47 ) and P. herbeus ( Figs 18 , 37 , 49 ) do.

Less than 3% of the examined specimens of P. daphnis are deep blue-green ( Fig. 11 ), but their colouration is not similar to that of P. coeruleus ( Figs 13, 47 ). This fact likely led Eൽආඈඇൽඌ (1994) and Eൽආඈඇൽඌ & ZටൽൾK (2012) to consider P. coeruleus and P. daphnis synonyms. Nevertheless, the deep blue-green chromatic phase of P. daphnis ( Fig. 11 ) never shows the distinctly smooth elytral interstriae seen in P. coeruleus ( Fig. 13). The typical green specimens of P. daphnis ( Figs 10, 12 , 46 , 57 ) and P. herbeus ( Figs 16–17 , 59 ) are similar in colour. Nevertheless, P. daphnis never shows a green-red chromatic phase like in P. herbeus ( Fig. 49 ). Phanaeus daphnis and P. herbeus are sympatric in an area between Estado de México and Morelos ( Fig. 65 ).

Distribution. Mexico: Estado de México, Guerrero, Morelos, Puebla, Oaxaca ( Fig. 65 ).

Figures

Figs 10–12. Phanaeus daphnis Harold, 1863. 10 – male green phase; 11 – male deep blue-green phase; 12 – lectotype and labels (by Christophe Rivier, MNHN). 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 16–18. Phanaeus herbeus Bates, 1887, stat. rev. 16 – male green phase; 17 – holotype and labels (by Mario Cupello, UFPR); 18 – P. tricornis Olsoufieff, 1924, junior subjective synonymy, redrawn from Oඅඌඈඎൿංൾൿൿ (1924). Scale bar = 1.0 mm.

Fig. 65. Predicted distribution of P. daphnis Harold, 1863, P. substriolatus Balthasar, 1939 and P. herbeus Bates, 1887.