Phanaeus olsoufieffi Balthasar, 1939

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 : 29-33

Phanaeus olsoufieffi Balthasar, 1939 stat. rev.

Figs 1J , 2F , 8 , 17 , 18K , 19K

Phanaeus blanchardi Olsoufieff, 1924: 92 (not P. blanchardi Harold, 1871: 114 ).

Phanaeus olsoufieffi Balthasar, 1939: 242 .

Phanaeus bothrus Blackwelder, 1944: 209 .

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

Phanaeus blanchardi – Vulcano & Pereira 1967: 575. — Martínez & Pereira 1967: 68 (as synonym of P. funereus ). — Edmonds 1972: 830, fig. 256; 1994: 3, 8, 45–46 (as synonym of P. pyrois ). — Arnaud 1982: 116 (as synonym of P. funereus ). — Krajcik 2006: 152 (as synonym of P. pyrois ). — Edmonds & Zídek 2012: 5, 13 (as synonym of P. pyrois ). — Chamorro et al. 2019: 220–221 (as synonym of P. pyrois ).

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

Phanaeus (Notiophanaeus) pyrois olsoufieffi – Arnaud 2002b: 96.

Phanaeus pyrois olsoufieffi – Arnaud 2002b: 98; 2018: 4, pl. 1, figs d–e.

Non Phanaeus pyrois (error). — Medina et al. 2001: 140 (in part). — Moctezuma & Halffter 2017: 55 (in part). — Moctezuma et al. 2017: 114, 130 (in part).

Diagnosis

This is the largest species within the P. endymion species group, frequently attaining 21–24 mm in length. Phanaeus olsoufieffi is diagnosed by the black colour with bright metallic red-green sheen; and elytral striae not strongly impressed basally ( Fig. 8 ). This species is easily separated from the closely related P. panamensis sp. nov. by the larger body size and endophallite copulatrix ( Fig. 1 ). Additionally, the major males of P. olsoufieffi are diagnosed by the distinctly developed keel in the middle of anterior pronotal margin; and posterolateral angles rounded, strongly developed, and projected laterally ( Figs 2F , 8A ).

Type material (studied from photographs, 2 ♂♂, 3 ♀♀)

Lectotype

COLOMBIA – Valle del Cauca • ♂, Arnaud 1982: 116 ( Fig. 8C ); “M. de Mathan 1898 / Ph. blanchardi ♂ Olsuf. det. G. Olsufiew. / Phanaeus apollinaris Muzo 1928 / Muséum Paris 1952 Coll R. Oberthur / P. blanchardi Ols. LECTOTYPE ♂ P. ARNAUD DET 1980/ Lectotype / Phanaeus pyrois Bates, Det. W. D. Edmonds ’83 / MNHN EC10566 ”; MNHN.

Paralectotypes

COLOMBIA – 1 ♂; “ExMus ӕo VAN LANSBERGE /Museum Paris ex. Coll. R. Oberthur / PARALECTOTYPE/ Phanaeus blanchardi Ols. PARALECTOTYPE ♂ P. ARNAUD DET 1981/ Phanaeus pyrois Bates / Det. W. D. Edmonds ‘83”/ MNHN EC10568 ”; MNHN. – Boyacá • 1 ♀; “ Muzo / Coll.E.Steinheil /MuseumParisex. Coll.R.Oberthur / PARALECTOTYPE/ PARALECTOTYPE Phanaeus (Phanaeus) blanchardi Olsoufieff, 1924 / MNHN EC10570 ”; MNHN EC10570 . – Valle del Cauca • 1 ♀; “ Santa Rosa entre S. Francisco & Carthago. Eujenio Garzon Aout 1878 ”; MNHN. – Unknown locality • 1 ♀; “(Illegible data) / Ex-mus ӕo D. Sharp 1890/ Ph. blanchardi ♀ Olsuf. det. G. Olsoufiew. / Museum Paris ex Coll. R. Oberthur / Phanaeus blanchardi ♀’s. PARALECTOTYPE ♀ P. ARNAUD DET 1981/ PARALECTOTYPE / Phanaeus pyrois Bates Det. W. D. Edmonds ’83/ MNHN EC10567 ”; MNHN .

Non-type material revised (24 ♂♂, 16 ♀♀)

COLOMBIA – Chocó • 3 ♂♂, 1 ♀; “ Q.Taparal B.San Juan . CNF. Nov 3–12–92. L.C. Pardo leg”; TAMU • 1 ♂, 1 ♀; “ Pacurita , 53m. 25-XI-01. 5°41’N, 76°40’W, bosque. Excrem. J.C. Neita col”; IEXA • 5 ♂♂, 5 ♀♀; same collection data as for preceding; TAMU • 1 ♂, 1 ♀; same collection data as for preceding; UVGC • 1 ♂; same collection data as for preceding; VMC • 1 ♀; “ Tutunendó , (20 km NE Quibdó), 60m, 26-XI-01. J.C. Neita col”; TAMU • 3 ♀♀; “ Unión Panamericana , 115m. 5°32’45N, 76°44’33’’W (No date) J.C. Neita col”; TAMU • 1 ♂, 1 ♀; “ Lloró, (3k km S Quibdó) 5°30’N, 76°33.5’W 90 m. J.C. Neita col (no date)”; TAMU • 2 ♂♂; “ Lloró , 90m, 20-II-03. 5°31’N, 76°33’W. Olaya & Mosquero; TAMU • 1 ♀; same collection data as for preceding; VMC. – Tolima • 1 ♂; “ Honda ”; GHC. – Valle del Cauca • 1 ♂; “Escalerete CF. Jul 19–21–91. L.C. Pardo leg”; TAMU • 1 ♂; same collection data as for preceding; VMC • 1 ♂, 1 ♀; “ B/Ventura B.Calima , C.F. 50 msnm. Col. ME Hitchcox ”; TAMU • 1 ♂, 1 ♀; “S30- 290. L.C. Pardo Locarno Leg.”; GHC • 2 ♂♂; same collection data as for preceding; VMC .

ECUADOR – Esmeraldas • 2 ♂♂; “ Charco Vicente . Ex. Pitfall heces humanas. 16/05/01. J. Quito F. Añapa ”; TAMU .

Type locality

Colombia, Valle del Cauca.

Redescription

Major male

HEAD. Clypeus bidentate, black anteriorly, bright metallic red posteriorly, with green sheen; roughened sculpture. Genae bright metallic red, with green sheen; roughened sculpture. Front black. Cephalic horn black, curved posteriorly over pronotum ( Figs 2F , 8A ).

PRONOTUM. Carinate, distinctly developed keel in the middle of anterior pronotal margin. Disc triangular, flat, with two distinctly developed tubercles on anterior portion. Triangle completely dull black, with bright metallic red sheen; completely smooth, scabriculous, impunctate. Sides bright metallic red, with green sheen; smooth sculpture, scabriculous, with effaced to almost effaced punctures. Lateral lines of pronotal triangle straight. Posterolateral angles rounded, strongly developed, projected laterally. Lateral fossae distinctly impressed. Basal fossae obtusely oval, deeply impressed to effaced. Posterior margin impunctate, becoming occasionally bright metallic red ( Figs 2F , 8A ).

ELYTRA. Striae fine, smooth, dull black, with superficially impressed to effaced punctation, scabriculous, not strongly impressed basally. Interstriae dull black, smooth, scabriculous, impunctate. Sutural margin without apical tooth ( Fig. 8A ).

PROTIBIAE. Quadridentate with apical spine.

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

GENITALIA. Right lobe of endophallite copulatrix slightly more developed than left lobe. Right lobe obtusely triangular in shape; weakly developed, projected frontally. Left lobe concave superiorly, lobed inferiorly. Central ridge less developed than central column ( Fig. 1J ).

Minor male

Like the major male, except for the reduction of 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 weakly developed tubercles; carinate middle tubercle, slightly more frontally projected and more developed than lateral tubercles; frons with superficially impressed punctures; pronotal sculpture smooth, with almost effaced to effaced punctures; pronotum almost completely black, becoming posteriorly and laterally bright metallic red with green sheen; pronotal process trituberculate, lacking concavities; pronotal tubercles weakly developed, well-spaced, with middle tubercle more developed and posteriorly projected than lateral tubercles; posterior pronotal midline completely effaced ( Fig. 8B ).

Variation

Mean length 20.1 mm (15.9–23.9 mm). Minor males occasionally show the pronotal disc almost completely bright metallic red, with green sheen.

Distribution

Pacific Slope of the Andes, north-central Colombia and northern Ecuador ( Fig. 17 ). Previous authors reported P. olsoufieffi from Panama ( Arnaud 2002b; Kohlmann et al. 2018). Nevertheless, these authors confused P. olsoufieffi with P. panamensis sp. nov. The specimens revised herein are the first accurate records of P. olsoufieffi from Ecuador. The distributions of P. olsoufieffi and P. funereus show a significant area of sympatry. Nevertheless, we did not find any putative hybrid specimens.

Remarks

Phanaeus olsoufieffi was considered as a synonym of P. pyrois by previous authors ( Edmonds 1994; Edmonds & Zídek 2012). Nonetheless, a diagnosis and an updated key to separate P. olsoufieffi and closely related species are provided by us. As a consequence, P. olsoufieffi is confidently resurrected herein from previous synonymy and full species status is assigned to it. The lectotype of P. olsoufieffi is a minor male ( Fig. 8C ). Therefore, the redescription mainly relies on the type series and some specimens (n =8) collected from the type locality (Valle del Cauca, Colombia). When revising the type material deposited at MNHM, we were not able to find out the locality data for a Colombian female paralectotype of P. olsoufieffi (MNHN EC10567). Its locality data is probably indicated in an illegible label. Additionally, we found out that a paralectotype of P. olsoufieffi pertained to P. malyi ( Fig. 7C , MNHN EC10569). Consequently, we conclude that the type series of P. olsoufieffi lumped together two distinct species. Previous authors ( Arnaud 1982; 2002a, 2002b; Edmonds 1994; Edmonds & Zídek 2012) did not realize this fact that solves the controversy considering P. malyi as a junior subjective synonym of P. olsoufieffi or not ( Edmonds & Zídek 2012; Solís & Kohlmann 2012; Kohlmann et al. 2018).

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. 8. Phanaeus olsoufieffi Balthasar, 1939 stat. rev. A. ♂ (TAMU). B. ♀ (TAMU). C. Lectotype ♂ (by Christophe Rivier, Muséum national d’histoire naturelle, Paris, France) (MNHN).

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.

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).