Phanaeus zapotecus Edmonds, 2006

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 : 47-50

Phanaeus zapotecus Edmonds, 2006

Figs 1P , 2L , 14–15 , 18Q , 19Q

Phanaeus zapotecus Edmonds, 2006: 31–32 , 35–37, figs 1, 3, 5, 7.

Phanaeus dionysius Kohlmann et al., 2018: 67–70 , 76, 78, 82, 88–89, figs 1b, 3a–b, d, 4–7. Syn. nov.

Phanaeus zapotecus – Edmonds & Zídek 2012: 1, 7–8. — Moctezuma & Halffter 2017: 52, 54–55, fig. 23. — Moctezuma et al. 2017: 114, 130, 132; 2019: 253.

Phanaeus (Notiophanaeus) zapotecus – Edmonds & Zídek 2012: 3, 13, figs 139, 141, 143, 148–151.

Phanaeus (Notiophanaeus) dionysius – Kohlmann et al. 2018: 68 , 70.

Non Phanaeus endymion – Edmonds 1994: 44 (referred to as “ Oaxaca ” population).

Non Phanaeus (Notiophanaeus) “ Oaxaca ” endymion – Edmonds 1994: fig. 221.

Type material of Phanaeus zapotecus revised (3 ♂♂, 5 ♀♀)

Holotype (not studied, temporarily lost)

MEXICO – Oaxaca • ♂, Edmonds 2006: 32; 8 km south of San Miguel Sola de Vega; IEXA ?

Paratypes

MEXICO – Oaxaca • 1 ♂, 1 ♀; “ 8 km Sola de Vega, 1850 m, 7–17 ix 05, Pine oak forest, D. Curoe col. Mushroom-baited trap ”; CEMT • 1 ♂, 1 ♀; same collection data as for preceding; TAMU • 1 ♂, 2 ♀♀; same collection data as for preceding; VMC • 1 ♀; “ 8 km Sola de Vega, 1850 m, 4–9 vii 05, Pine oak forest, D. Curoe col. Mushroom-baited trap ”; TAMU .

Type material of Phanaeus dionysius revised (2 ♂♂, 2 ♀♀)

Holotype

MEXICO – Oaxaca • “ La mesita San Pablo Etla , 23–VI–17, coprotrampa, x- 96 °44’18.91’’O, y- 17°9’54.36’’N, bosque de Encino, 1976 m, Arriaga A. & Arenas A. Col. ”; IEXA.

Paratypes

MEXICO – Oaxaca • 2 ♀♀; same collection data as for holotype; IEXA • 1 ♂; “ La mesita San Pablo Etla, 14–VII–17, coprotrampa, x- 96 °44’53.55’’O, y- 17°9’53.55’’N, bosque de Encino, 1954 m, Arriaga A. & Arenas A. Col. ”; IEXA .

Non-type material revised (1 ♂)

MEXICO – Oaxaca • 1 ♂; “ San Pablo Etla , Reserva San Pablo Etla , 20/X/2016, C.D. 17°10’1.23’’N, 96°44’14.25’’O, 1994 m, Alfosina Arriaga Col.”; VMC .

Type locality

Mexico, Oaxaca, 8 km south of San Miguel Sola de Vega.

Phanaeus dionysius

Kohlmann et al. (2018) recently described P. dionysius based upon specimens from San Pablo Etla, Oaxaca, Mexico and suggested that P. zapotecus and P. dionysius were sister taxa. The following character combination helps to separate both species according to Kohlmann et al. (2018): P. dionysius has long and slender pronotal posterolateral angles, whereas P. zapotecus has short and rounded posterolateral angles. The basal border of the tergite VIII in P. dionysius forms a small indentation at its middle, whereas it runs completely straight in P. zapotecus . Additionally, the apex of the parameres of P. dionysius is more projected, than that from P. zapotecus . Moreover, the middle sinuation of the parameres in lateral view is much more pronounced in P. dionysius . Nevertheless, we consider that the description and diagnosis of P. dionysius exemplify some of the frequent problems occurring in taxonomical work ( Komarek & Beutel 2006) as follows.

Insufficient study of types

Kohlmann et al. (2018) did not indicate how many specimens of P. zapotecus they revised, but they commented that the paratypes housed at CNIN were checked by them. Taking into account this information and the original description of P. zapotecus ( Edmonds 2006) , we assume that Kohlmann et al. (2018) revised two paratypes at most.

Insufficient assessment of the range of character variation

Kohlmann et al. (2018) were not able to adequately assess the morphological variation of P. zapotecus , as a consequence of the reduced number of paratypes studied by them. The pronotal posterolateral angles were found by us to be long and slender in some paratype major males of P. zapotecus ( Fig. 14A ). The basal border of the tergite VIII is highly variable: four paratypes of P. zapotecus revised by us show the small indentation at the middle of the basal border of the tergite VIII ( Fig. 14D ), whereas it runs completely straight in two paratypes. The small indentation is present in the basal border of the tergite VIII of three specimens of P. dionysius revised by us, whereas it runs completely straight in two paratypes ( Fig. 14E ).

Kohlmann et al. (2018) illustrated the endophallite copulatrix of P. dionysius (available from https://zookeys.pensoft.net/article/23029/zoom/fig/13/) but they overlooked its comparison with P. zapotecus ( Fig. 1P ). We noticed that the morphology of both P. zapotecus ( Fig. 1P ) and P. dionysius is identical when we compared the endophallites under the microscope. Furthermore, the endophallite copulatrix of the holotype of P. dionysius has suffered deformation and abrasion, maybe as a result of the preparation methods (soaking for 5 min in a solution of 5% boiling KOH) and/or storage (dry preservation in a plastic microvial). This structure was not found by us to be stored in a microvial with glycerol as mentioned by Kohlmann et al. (2018). The morphology of the cephalic carina and the pronotal tubercles of the females of P. zapotecus ( Fig. 14B ) and P. dionysius (see https://zookeys.pensoft.net/article/23029/zoom/fig/14/) did not differ. Moreover, the body colour in the paratypes of P. zapotecus varied from jet black or bright dark blue to jet black with a green-blue sheen ( Fig. 14A–C ), falling within the variation of P. dionysius ( Kohlmann et al. 2018) .

Kohlmann et al. (2018) commented that a major male was designated as holotype. Nevertheless, we found out that the holotype of P. dionysius housed at IEXA is the minor male illustrated by Kohlmann et al. (2018) in the original description (available from https://zookeys.pensoft.net/article/23029/zoom/fig/15/). As a consequence, we assume that the original description of P. dionysius was not based on the holotype. We found that a paratype minor male of P. zapotecus ( Fig. 14C ) and the holotype of P. dionysius (https://zookeys.pensoft.net/article/23029/zoom/fig/15/) were morphologically almost identical.

Characters not suitable for a study at a given taxonomic rank

The revision of a majority of species within the P. endymion species group led us to conclude that the morphology of the aedeagus is not taxonomically informative to separate species, as suggested by previous authors ( Edmonds 1994; Price 1995; Arnaud 2002b; Moctezuma & Halffter 2017; Moctezuma et al. 2017, 2019).

In the light of the findings presented herein, we conclude that the description and diagnoses provided by Kohlmann et al. (2018) do not justify the splitting of P. dionysius and P. zapotecus in two different species. As a consequence, a new junior subjective synonymy is recognized herein: Phanaeus zapotecus Edmonds, 2006 = Phanaeus dionysius Kohlmann, Arriaga-Jiménez & Rös, 2018 syn. nov.

Distribution

Sierra Madre del Sur and Sierra Norte, central Oaxaca ( Fig. 15 ).

Remarks

Mean length 16.4 mm (14.8–18.5 mm). To the original description of this species, we add that the elytral striae are impressed basally as distinct fossae ( Fig. 14 ); right lobe of endophallite copulatrix more developed than left lobe; right lobe obtusely triangular in shape, strongly developed superiorly; left lobe strongly reduced, lobed, slightly concave inferiorly; central ridge more developed than central column ( Fig. 1P ). For the female, the head shows a cephalic trituberculate carina; with almost aligned, conical tubercles; middle tubercle slightly more developed than lateral tubercles; frons with almost effaced punctures; pronotal process trituberculate, with posterior concavity; rounded pronotal tubercles; with middle tubercle slightly more developed than lateral tubercles; posterior pronotal midline distinctly impressed ( Fig. 14B ).

Edmonds (2006) commented that the holotype and three paratypes (1 ♂, 2 ♀♀) of P. zapotecus were housed at IEXA. However, the type material was not found by us at IEXA in a recent search. Therefore, the holotype and these paratypes are considered temporarily lost. It is possible that the former collection manager of IEXA, Professor Miguel Angel Morón Ríos, knew the location of the holotype and paratypes of P. zapotecus , but, unfortunately, we were not able to confirm this assumption because of his recent death.

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. 14. A–D. Phanaeus zapotecus Edmonds, 2006. A. ♂ (TAMU). B. ♀ (TAMU). C. Minor ♂ (VMC). D. Tergite VIII (TAMU). E. Phanaeus dionysius Kohlmann, Arriaga-Jiménez & Rös, 2018 syn. nov., tergite VIII (IEXA).

Fig. 15. Distribution of the Phanaeus endymion species group in North and Central America (in part). Phanaeus bravoensis Moctezuma, Sánchez-Huerta & Halffter, 2017;P. edmondsi Moctezuma, Deloya & Halffter, 2019; P. halffterorum Edmonds, 1979; and P. huichol Moctezuma, Sánchez-Huerta & Halffter, 2017 by Jose Luis Sánchez-Huerta.

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.