Phanaeus malyi Arnaud, 2002a

Moctezuma, Victor & Halffter, Gonzalo, 2021, Taxonomic revision of the Phanaeus endymion species group (Coleoptera: Scarabaeidae), with the descriptions of five new species, European Journal of Taxonomy 747, pp. 1-71 : 26-29

Phanaeus malyi Arnaud, 2002a

Figs 1I , 2E , 7 , 16–17 , 18J , 19J

Phanaeus pyrois malyi Arnaud, 2002a: 4 .

Phanaeus pyrois – Bates 1887: 58 (in part, as a variety of P. pyrois ). — Kohlmann et al. 2018: 78 (in part).

Phanaeus blanchardi (error) – Olsoufieff 1924: 92 (not P. blanchardi Harold, 1871: 114 , in part). — Vulcano & Pereira 1967: 575 (in part). — Martínez & Pereira 1967: 68 (in part, as synonym of P. funereus ). — Edmonds 1972: 830, fig. 256 (in part); 1994: 3, 8, 45–46 (in part, as synonym of P. pyrois ). — Arnaud 1982: 116 (in part, as synonym of P. funereus ). — Krajcik 2006: 152 (as synonym of P. pyrois ). — Edmonds & Zídek 2012: 5, 13 (in part, as synonym of P. pyrois ). — Chamorro et al. 2019: 220–221 (in part, as synonym of P. pyrois ).

Phanaeus pyrois malyi – Arnaud 2002b: 97 (as subspecies of P. pyrois ). — Solís & Kohlmann 2012: 1, 9–10 (as subspecies of P. pyrois ). — Edmonds & Zídek 2012: 1, 8, 13 (as synonym of P. pyrois ). — Krajcik 2006: 152 (as subspecies of P. pyrois ). — Kohlmann et al. 2018: 78.

Phanaeus (Notiophanaeus) pyrois malyi – Arnaud 2002b: 96 (as subspecies of P. pyrois ).

Phanaeus malyi – Solís & Kohlmann 2012: 1, 7, 31, fig. 1. — Edmonds & Zídek 2012: 3, 6 (as synonym of P. pyrois ). — Kohlmann et al. 2018: 67–69 , 78–80, 82, 88–89, figs 8, S1b. — Gillett & Toussaint 2020: fig. 4.

Phanaeus (Notiophanaeus) malyi – Kohlmann et al. 2018: 78 .

Non Phanaeus olsoufieffi (error) – Balthasar 1939: 242 (in part). — Edmonds 1994: 8, 45–46 (in part, as synonym of P. pyrois ). — Arnaud 2002b: 96 (in part). — Krajcik 2006: 152 (in part, as synonym of P. pyrois ). — Edmonds & Zídek 2012: 3, 5–6 (in part, as synonym of P. pyrois ).

Non Phanaeus bothrus (error) – Blackwelder 1944: 209 (in part). — Martínez & Pereira 1967: 68 (in part, as synonym of P. funereus ). — Edmonds 1994: 8, 45 (in part, as synonym of P. pyrois ). — Arnaud 2002b: 97 (in part, as synonym of P. olsoufieffi ). — Krajcik 2006: 152 (in part, as synonym of P. pyrois ). — Edmonds & Zídek 2012: 3, 5 (in part, as synonym of P. pyrois ).

Non Phanaeus (Phanaeus) blanchardi (error) – Martínez & Pereira 1967: 68 (in part, as synonym of P. funereus ).

Non Phanaeus (Phanaeus) olsoufieffi (error) – Martínez & Pereira 1967: 68 (in part, as synonym of P. funereus ).

Non Phanaeus (Notiophanaeus) pyrois olsoufieffi (error) – Arnaud 2002b: 96 (in part).

Non Phanaeus pyrois olsoufieffi (error) – Arnaud 2002b: 98; 2018: 4 (in part).

Diagnosis

Easily diagnosed species by the bright black colour with red-green sheen on frontolateral angles of pronotum ( Fig. 2E ); elytral striae fine, smooth, impressed basally as distinct fossae ( Fig. 7 ). The rest of black species within the P. endymion species group ( P. funereus , P. olsoufieffi , P. panamensis sp. nov.) are differentiated from P. malyi by the elytral striae not strongly impressed basally as distinct fossae and the shape of the endophallite copulatrix ( Fig. 1 ).

Type material

Holotype (not studied)

COSTA RICA • ♂, Arnaud 2002a: 3; Puntarenas, Carara National Park, Estación Quebrada Bonita ; originally deposited at INBIO; MNCR.

Non-type material revised (10 ♂♂, 6 ♀♀)

COLOMBIA • 1 ♂; unknown locality; “[Illegible data]/ Ex-mus ӕo D. Sharp 1890/ Museum Paris ex Coll. R. Oberthur / PARALECTOTYPE/ PARALECTOTYPE Phanaeus (Phanaeus) blanchardi Olsoufieff, 1924 / MNHN EC10569 ”; MNHN .

COSTA RICA – Puntarenas • 1 ♂; “ San Luis, San Luis Vly. VII-29–1996. Coll. Louis La Pierre ”; VMC • 1 ♀; “ 6km S. San Vito. 27-IV / 7-V 1967. 100m. D.F. Viers col. human feces? ”; TAMU • 1 ♂; “ S.Vito , Las Cruces. July 1982. B.Gill. 1200 m ”; IEXA • 1 ♂, 1 ♀; same collection data as for preceding; TAMU • 1 ♂, 1 ♀; “ S.Vito , Las Cruces. 17 VIII-12 IX 1982. B.Gill. 1200 m ”; TAMU • 1 ♂; same collection data as for preceding; UVGC • 1 ♀; “ S.Vito , Las Cruces. 15–18 VIII 1982. B.Gill. 1200 m ”; VMC • 1 ♂; “ Rincón de Osa. 3-X-69. G. Halffter y P. Reyes C., col. Selva tropical lluviosa. Cebo excremento. Día”; GHC • 1 ♀; “ Rincón de Osa. 5-X-69. G. Halffter y P. Reyes C., col. Selva tropical lluviosa. Cebo excremento. Día”; GHC • 1 ♀; “ Rincón de Osa. 30-IX-69. G. Halffter y P. Reyes C., col. Selva tropical lluviosa. Cebo excremento. Día (11–17 hrs.)”; VMC • 1 ♂; “ Rincón de Osa. 22–5–1965. Col. P. Kazan ”; GHC .

PANAMA – Chiriquí • 1 ♂; “ Cerro Hornito 15 km NE Gualaca. 17–21 VI 1982. B.Gill. 1200 m.”; TAMU • 1 ♂; “ Cerro Pelota 4 km N Sta.Clara. 9–18 VIII 1982. B.Gill. 1500 m.”; VMC .

Type locality

Costa Rica, Puntarenas, Carara National Park, Estación Quebrada Bonita.

Redescription

Major male

HEAD. Completely bright black. Clypeus bidentate, with roughened sculpture. Genae with roughened sculpture. Cephalic horn curved posteriorly over pronotum ( Figs 2E , 7A ).

PRONOTUM. Keel absent in the middle of anterior pronotal margin. Disc triangular, flat; with two weakly developed, elongate tubercles on anterior portion. Triangle bright black, smooth, scabriculous, impunctate. Sides bright black, becoming bright metallic red-green; scabriculous, with smooth sculpture, almost effaced punctures. Lateral lines of triangle straight. Posterolateral angles short, widened. Lateral

fossae distinctly impressed. Basal fossae distinctly to superficially impressed. Posterior margin with distinctly to superficially impressed punctures ( Figs 2E , 7A ).

ELYTRA. Striae fine, smooth, completely bright black, with superficially impressed to effaced punctures, scabriculous, impressed basally as distinct fossae. Interstriae bright black, with smooth surface, scabriculous, impunctate or with almost effaced punctures. Sutural margin without apical tooth ( Figs 7A ).

PROTIBIAE. Quadridentate with apical spine.

TERGITE VIII. Bright metallic red-green, scabriculous sculpture; with rough, superficially impressed punctures. Basal margin with setae variable in size.

GENITALIA. Right and left lobes of endophallite copulatrix similar in size. Right lobe obtusely triangular in shape, rounded superiorly. Left lobe obtusely lobed. Central ridge less developed than central column ( Fig. 1I ).

Minor male

Like the major male, except for the reduction of the secondary sexual characters (i.e., cephalic horn, pronotal triangle and tubercles, and pronotal posterolateral angles).

Female

Similar to the male, except for the head showing a cephalic trituberculate carina, with nearly aligned, rounded or carinate, weakly developed tubercles; middle tubercle slightly more developed than lateral tubercles; frons with superficially impressed punctures; pronotum with almost effaced punctures; pronotal process trituberculate, lacking concavity; middle pronotal tubercle more developed, slightly more projected posteriorly than lateral tubercles; all tubercles rounded in shape; posterior pronotal midline completely effaced to almost effaced ( Fig. 7B ).

Variation

Mean length 17.8 mm (12.8–20.5 mm). Colour variation was not found for P. malyi .

Distribution

Southern Pacific costal area, Costa Rica, Panama and Colombia ( Figs 16–17 ). The specimens revised by us represent the first accurate records of P. malyi from Panama. Apparently, the distributions of P. malyi and P. panamensis sp. nov. show a significant area of sympatry and both species may be collected in the same locality (e.g., Cerro Hornito, Panama). Nevertheless, P. malyi and P. panamensis sp. nov. are confidently identified by the diagnosis provided herein and putative hybrid specimens were not found by us. The MNHN EC10569 specimen represents the first record for P. malyi in Colombia. Nevertheless, the illegible label data prevents us to provide an accurate locality ( Fig. 7C ). The extent of the distribution of P. malyi in Colombia needs to be confirmed by future research.

Remarks

Phanaeus malyi was considered by previous authors as a synonym of P. pyrois ( Edmonds & Zídek 2012) . Nevertheless, clear differences in the external and genital morphology of P. malyi were found by us. These differences support the full species status suggested by Solís & Kohlmann (2012) and Kohlmann et al. (2018). When revising the type material housed at MNHM, a Colombian minor male specimen of P. malyi was found to be labelled as a paralectotype of P. blanchardi ( Fig. 7C , MNHN EC10569).

Figures

Fig. 1. Endophallite copulatrix of the Phanaeus endymion species group (except for P. arletteae Arnaud, 2018). A. Phanaeus bravoensis Moctezuma, Sánchez-Huerta & Halffter, 2017 (IEXA). B. Phanaeus chiapanecus sp. nov. (TAMU). C. Phanaeus edmondsi Moctezuma, Deloya & Halffter, 2019 (IEXA). D. Phanaeus endymion Harold, 1863 (VMC). E. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). F. Phanaeus halffterorum Edmonds, 1979 (TAMU). G. Phanaeus huichol Moctezuma, Sánchez- Huerta & Halffter, 2017 (IEXA). H. Phanaeus jackenioi sp. nov. (UVGC). I. Phanaeus malyi Arnaud, 2002 (VMC). J. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). K. Phanaeus pacificus sp. nov. (UVGC). L. Phanaeus panamensis sp. nov. (TAMU). M. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). N. Phanaeus pyrois Bates, 1887 (VMC).O. Phanaeus rzedowskii sp. nov. (TAMU).P. Phanaeus zapotecus Edmonds, 2006 (VMC). Q. Phanaeus zoque Moctezuma & Halffter, 2017 (IEXA).

Fig. 2. Lateral view of males within the Phanaeus endymion species group. A. Phanaeus chiapanecus sp. nov. (TAMU). B. Phanaeus endymion Harold, 1863 (VMC). C. Phanaeus funereus Balthasar, 1939 stat. rev. (VMC). D. Phanaeus jackenioi sp. nov. (UVGC). E. Phanaeus malyi Arnaud, 2002 (VMC). F. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. (VMC). G. Phanaeus pacificus sp. nov. (UVGC). H. Phanaeus panamensis sp. nov. (TAMU). I. Phanaeus porioni Arnaud, 2001 stat. rev. (VMC). J. Phanaeus pyrois Bates, 1887, green morph (VMC). K. Phanaeus rzedowskii sp. nov. (TAMU). L. Phanaeus zapotecus Edmonds, 2006 (VMC).

Fig. 7. Phanaeus malyi Arnaud, 2002.A. ♂ (TAMU). B. ♀ (TAMU). C. Colombian specimen previously labelled as a paralectotype of P. blanchardi Olsoufieff, 1924 (by Christophe Rivier, Muséum national d’histoire naturelle, Paris, France) (MNHN).

Fig. 16. Distribution of the Phanaeus endymion species group in Central America (in part).

Fig. 17. Distribution of the Phanaeus endymion species group in South America. Phanaeus malyi Arnaud, 2001 was reported from Colombia without accurate locality. Phanaeus arletteae Arnaud, 2018 was modified from Arnaud (2018).

Fig. 18. Pronotum of females within the Phanaeus endymion species group. A. Phanaeus arletteae Arnaud, 2018 (modified from Arnaud 2018). B. Phanaeus bravoensis Moctezuma, Sánchez-Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta).C. Phanaeus chiapanecus sp. nov. D. Phanaeus edmondsi Moctezuma, Deloya & Halffter, 2019 (by Jose Luis Sánchez-Huerta). E. Phanaeus endymion Harold, 1863. F. Phanaeus funereus Balthasar, 1939 stat. rev. G. Phanaeus halffterorum Edmonds, 1979 (by Jose Luis Sánchez-Huerta).H. Phanaeus huichol Moctezuma, Sánchez-Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta). I. Phanaeus jackenioi sp. nov. J. Phanaeus malyi Arnaud, 2002. K. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. L. Phanaeus pacificus sp. nov. M. Phanaeus panamensis sp. nov. N. Phanaeus porioni Arnaud, 2001 stat. rev. O. Phanaeus pyrois Bates, 1887. P. Phanaeus rzedowskii sp. nov. Q. Phanaeus zapotecus Edmonds, 2006. R. Phanaeus zoque Moctezuma & Halffter, 2017.

Fig. 19. Elytral integument of species within the Phanaeus endymion species group. A. Phanaeus arletteae Arnaud, 2018 (modified from Arnaud 2018). B. Phanaeus bravoensis Moctezuma, Sánchez- Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta). C. Phanaeus chiapanecus sp. nov. D. Phanaeus edmondsi Moctezuma, Deloya & Halffter, 2019 (by Jose Luis Sánchez-Huerta). E. Phanaeus endymion Harold, 1863. F. Phanaeus funereus Balthasar, 1939 stat. rev. G. Phanaeus halffterorum Edmonds, 1979 (by Jose Luis Sánchez-Huerta). H. Phanaeus huichol Moctezuma, Sánchez-Huerta & Halffter, 2017 (by Jose Luis Sánchez-Huerta). I. Phanaeus jackenioi sp. nov. J. Phanaeus malyi Arnaud, 2002.K. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. L. Phanaeus pacificus sp. nov. M. Phanaeus panamensis sp. nov. N. Phanaeus porioni Arnaud, 2001 stat. rev. O. Phanaeus pyrois Bates, 1887. P. Phanaeus rzedowskii sp. nov. Q. Phanaeus zapotecus Edmonds, 2006. R. Phanaeus zoque Moctezuma & Halffter, 2017.