Frolov A. V., Akhmetova L. A., Vaz-de-Mello F. Z. 2019. Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature. Journal of Natural History. 53 (11-12): 725-747.

DOI: 10.1080/00222933.2019.1606953

 

Contribution to the knowledge of Aegidinus Arrow (Coleoptera: Scarabaeidae: Orphninae): new species and comments on the classification and nomenclature

 

Frolov, A.V.1,2, L.A. Akhmetovaand Fernando Z. Vaz-de-Mello2

1Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St.-Petersburg, 199034 Russia. Email: frolov@scarabaeoidea.com

2Universidade Federal de Mato Grosso, Instituto de Biociências, Departamento de Biologia e Zoologia, Av. Fernando Corrêa da Costa, 2367, Boa Esperança, 78060-900 Cuiabá, MT, Brazil.

Abstract

Two new species of the Neotropical Orphninae genus Aegidinus Arrow are described: Aegidinus noriegai sp. nov. and Aegidinus colbyae sp. nov. The lectotype of Orphnus candezei Preudhomme de Borre, 1886, is designated. Aegidium steinheili Harold, 1880, considered a synonym of Aegidinus guianensis (Westwood, 1846) is revalidated as Aegidinus steinheili (Harold). The following new synonymies are established: Aegidinus oreibates Colby, 2009 = Aegidinus guianensis (Westwood, 1846), syn. nov.; Aegidinus crypticus Colby, 2009 = Aegidinus guianensis (Westwood, 1846), syn. nov.; Aegidinus sunidigea Colby, 2009 = Aegidinus candezei (Preudhomme de Borre, 1886), syn. nov.; Aegidinus unicus Colby, 2009 = Aegidinus cornutus Colby, 2009, syn. nov. The male of Aegidinus simulatus Colby, 2009, is described. An updated diagnostic key to the males of Aegidinus is provided. The taxonomic and diagnostic characters as well as phylogenetic relationships of Aegidinus are discussed.

Introduction

In the Neotropics, the scarab beetles of the subfamily Orphninae are represented by the endemic tribe Aegidiini Paulian and comprise five genera and 47 species, two thirds of which were described in the past decade  (Colby 2009, Frolov et al. 2015, Frolov and Vaz-de-Mello 2015, Frolov et al. 2017b, Frolov et al. 2017a, Rojkoff and Frolov 2017). Aegidinus Arrow is the second most speciose genus of the Aegidiini after Aegidium Westwood which comprises 22 species (Frolov et al. 2017a).

Aegidinus was recently revised by Colby (2009) who described 12 new species in addition to the three known by then. We have re-examined the types of the species described by Westwood, Harold, Preudhomme de Borre, and Colby, and the results suggested us to revise some of the taxonomic decisions of the previous authors and propose new synonymies. The purpose of this research is also to describe two new species and the previously unknown male of Aegidinus simulatus Colby.

Material and methods

The material used in this work is housed in the collections of the following organizations (curators in brackets):

BGCO            Bruce Gill collection, deposited in the Canadian National Collection, Ottawa, Canada  (Bruce Gill)

BMNH           Natural History Museum, London, United Kingdom (Maxwell Barclay)

CEMT             Entomology Sector of Zoological Collection, Biosciences Institute, Federal University of Mato Grosso, Cuiabá, Brazil (Fernando Vaz-de-Mello)

CMN               Canadian Museum of Nature, Ottawa, Canada (François Génier)

IRSNB                       Belgian Royal Institute of Natural Sciences, Bruxelles, Belgium (Alain Drumont)

MNHN           National Museum of Natural History, Paris, France (Olivier Montreuil)

OUMNH        Oxford University Museum, Oxford, United Kingdom (Darren Mann)

RMNHL         Royal Museum of Natural History Naturalis, Leiden, Netherlands (Hans Huijbregts, Jan Krikken)

UNMSM         Natural History Museum, National University Mayor de San Marcos, Lima, Peru (Angélico Asenjo, Luis Figueroa)

ZIN                 Zoological Institute, Russian Academy of Sciences, Saint-Petersburg, Russia (Andrey Frolov)

Morphological terminology follows Frolov (2012) and Frolov et al. (2016). Labels of the type specimens are cited verbatim and separated by a slash and our comments are in square brackets. The holotypes of the new species are supplied with red labels “HOLOTYPUS Aegidinus [species name] Frolov, Akhmetova & Vaz-de-Mello 2018”; the paratypes are supplied with yellow labels “PARATYPUS Aegidinus [species name] Frolov, Akhmetova & Vaz-de-Mello 2018”. Specimens, digital images and locality maps preparation follow Frolov et al. (2017a).

Genus Aegidinus Arrow, 1904

Type species: Aegidium guianensis Westwood, designated by Paulian (1984).

Diagnosis

Body small to mid-sized (length 6 to 12 mm), reddish brown to dark-brown. Mandibles subsymmetrical, with long processes on the outer sides. Clypeus with tubercle or horn on anterior margin medially in males, without horn in females. Pronotum variably excavated medially in males, convex to depressed medially in females; anterior margin of pronotum in males with a tubercle or horn medially. Propleurae with carinae separating anterolateral areas from basal area. Scutellum narrowly rounded apically, about 1/15 length of elytra. Elytra convex, with marked humeral umbones and striae marked with elongated punctures. Wings fully developed. Metepisternon triangular, its posterior angle rounded to triangular and situated in distinct concavity of epipleuron. Protibiae with 3 strong outer teeth in both sexes and with a smaller medial tooth in majority of males. Each procoxa with two cavities. Phallobase tube-shaped with strongly sclerotized ventral side but without differentiation of ventral and dorsal sclerites. Parameres relatively short, with complex shape, apices without setae. Gonocoxites separated into 2 more or less distinct parts.

Aegidinus is distinct among the orphnines in having the following unique characters: mandibles with processes on the outer sides; shape of the gonocoxites is diverse and species specific at least in majority of species; procoxae with two cavities each; parameres are asymmetric in two species; phallobase with protruding ventroapical plate in some species. From other Aegidiini, Aegidinus also differs in the absence of a hole connecting mesocoxal cavities, and in having more or less distinct transverse carina on metatibiae.

Diagnostic characters of species

Most Aegidinus species are similar externally and, despite some interspecific variation of the body sculpture and head and protoracic armature in some species, reliable identification is only possible by the examining on the shape of the male genitalia. External female genitalia (gonocoxites) are also highly specific although their usefulness in the distinguishing closely related species requires further evaluation (see Discussion below).

Species composition and distribution

Aegidinus comprises 14 species including two new described in this work. Members of the genus occur in South America, mostly in the Amazon and Guianan moist forest regions to Yungas in the west, and in the Trinidad island (Fig. 7).

Key to Aegidinus species (males)

1.       Parameres separated into dorsomedial and ventrolateral lobes (Figs. 1H, I, 2E, F, 3B–D).......... 2

-        Parameres not separated into dorsomedial and ventrolateral lobes (Figs. 6C–E) Aegidinus cornutus

2.       Phallobase with ventroapical plate (Figs. 1G, 3C, 4D) ..............................................................  3

-        Phallobase without ventroapical plate (Figs. 2D, 4H, 5D) ......................................................... 6

3.       Parameres symmetrical ............................................................................................................... 4

-        Parameres asymmetrical ............................................................................................................. 5

4.       Ventrolateral lobe of paramere with subapical tooth (Colby 2009: Fig. 55) Aegidinus howdenorum

-        Ventrolateral lobe of paramere without subapical tooth (Fig. 1H).............. Aegidinus guianensis

5.       Parameres longer, more asymmetrical (Figs. 4B, C); ventroapical plate of phallobase longer than wide (Fig. 4D); protibia without medioapical tooth .................................. Aegidinus noriegai sp. nov.

-        Parameres shorter, less asymmetrical (Figs. 3B, D); ventroapical plate of phallobase wider than long (Fig. 3C); protibia with medioapical tooth ............................................................. Aegidinus candezei

6.       Mediobasal margins of dorsomedial lobes of parameres feebly sclerotized, membranous (Fig, 2E); protibia with medioapical tooth ..............................................................................................................  7

-        Mediobasal margins of dorsomedial lobes of parameres strongly sclerotized (4G, 5E); protibia without medioapical tooth ......................................................................................................................  9

7.       Ventrolateral lobes of parameres long and slender (in lateral view), reasonably longer than dorsomedial lobes (Fig. 2F)................................................................................................................. steinheili

-        Ventrolateral lobes of parameres triangular and obtuse in lateral view, not longer than dorsomedial lobes .................................................................................................................................................... 8

8.       Ventrolateral lobes of parameres as long as dorsomedial lobes......................... Aegidinus petrovi

-        Ventrolateral lobes of parameres reasonably shorter than dorsomedial lobes Aegidinus teamscaraborum

9.       Dorsal sides of parameres strongly overlapping and separated by slit (Fig. 4F, arrowed) Aegidinus simulatus

-        Dorsal sides of parameres less overlapping and not separated by slit (Fig. 5F)........................ 10

10.     Dorsal processes of parameres carina-shaped (Fig. 5E, arrowed)  Aegidinus colbyae sp. nov.

-        Dorsal processes of parameres tooth or spur-shaped ...........................................................  11

11.     Dorsal processes of parameres long, spur-shaped ..................................... Aegidinus brasiliensis

-        Dorsal processes of parameres short, tooth-shaped  .......................................  Aegidinus howeae

Aegidinus guianensis (Westwood, 1845)

(Figures 1A–I)

Aegidium guianense Westwood, 1845: 175.

= Aegidinus oreibates Colby, 2009: 26, syn. nov.

= Aegidinus crypticus Colby, 2009: 20, syn. nov.

Type locality. Guyana, Upper Takutu-Upper Essequibo.

Type material examined

Ae. guianensis: holotype (Fig. 1A), female at OUMNH labelled “Guiana / Aegidium ? Guianense Westw. / TYPE WESTWOOD Trans. Ent. Soc L.1846.P.175 T. 12. 8. 5. Coll. Hope Oxon. / TYPE COL: 488 Aegidium guianensis Westw. HOPE DEPT. OXFORD / W / WB”.

Ae. oreibates: holotype (Fig. 1F), male at CMN labelled “TRINIDAD: ST. GEORGE 8 km N. Arima, Simla Res. Sta., 260 m, 14-24.VI.1993 S. & J. Peck, lower mont. rainforest, f.i.t., 93-48 / Aegidinus oreibates Colby HOLOTYPE”. Paratypes: GUYANA. Potaro-Siparuni: one female at CMN labelled “Guyana: Iwokrama F. R. 4°40ʹ19N 58°41ʹ04W 100-200 m, V-VI.2001 R.Brooks & Z.Falin in FIT / Aegidinus oreibates J. Colby PARATYPE”. TRINIDAD AND TOBAGO. One female at IRSNB labelled “♀ / Coll. R. I. Sc. N. B. Caparo Trinidad A. Heyne, Berlin-Wilm. / Aegidinus oreibates Colby PARATYPE / Paratype / Aegidinus guianense Westw ♀ E. Benderitter, det”. Arima: one male at CMN labelled “TRINIDAD, 16 km N Arima Andrews Trace, 620m up. montane rainforest FIT, 7-24.VI.1993 S&J Peck, 93-14 / PARATYPE Aegidinus oreibates J. Colby”; one male at CMN labelled “TRINIDAD, 8 km N Arima Simla Res. Sta. 260m 24.VI-8.VII.93, lower montane rainforest FIT S&J Peck, 93-57 / Aegidinus oreibates J. Colby PARATYPE”; one female at CMN labelled “TRINIDAD: ST. GEORGE 8 km N. Arima, Simla Res. Sta., 260 m, 14-24.VI.1993 S. & J. Peck, lower mont. rainforest, f.i.t., 93-48 / Aegidinus oreibates Colby ALLOTYPE”; one male and two females at CMN labelled “TRINIDAD: ST. GEORGE 8 km N. Arima, Simla Res. Sta., 260 m, 14-24.VI.1993 S. & J. Peck, lower mont. rainforest, f.i.t., 93-48 / Aegidinus oreibates Colby PARATYPE”; two males and one female at CMN labelled “TRINIDAD, 8 km N Arima Simla Res. Sta. 260m 6-14.VI.93 trop. forest FIT S&J Peck, 93-9 / PARATYPE Aegidinus oreibates J. Colby”. Couva-Tabaquite-Talparo: one male at CMN labelled “TRINIDAD, 11 km SE Arima, Arena For. Res. 13-22.VI.93, 80m rainforest FIT S&J Peck, 93-30 / PARATYPE Aegidinus oreibates J. Colby”; one male at CMN labelled “TRINIDAD, 13 km S Arima 2km N Talparo, Quesnell Farm, rainforest FIT 12-22.VI.93, 50m S&J Peck, 93-28 / PARATYPE Aegidinus oreibates J. Colby”. VENEZUELA. Bolívar: one female at CMN labelled “VENEZUELA:BOLIVAR Guri, 14.VI.1996 H.&A. Howden Dry forest L+ / Aegidinus oreibates J. Colby PARATYPE”.

Additional material examined

VENEZUELA. Bolívar: Guri, dry forest, 14.VI.1996, H. Howden and A. Howden leg., one female (CEMT); Guri, wet forest, 17.VI.1996, B.D. Gill leg., one female (BGCO). TRINIDAD AND TOBAGO. 11 km SE Arima, Arena Forest Reserve, 80 m, rainforest, FIT, 13-22.VI.1993, S. Peck, J. Peck leg., one male (BGCO); 8 km N Arima, Simla Research Station, 260 m, lower mountane rainforest FIT, 6-14.VI.1993, S. Peck, J. Peck leg., one male (BGCO); 16 km N Arima, Andrews Trace, 620 m, upper montane rainforest, FIT, 24.VI-7.VII.1993, S. Peck, J. Peck leg., one male (BGCO); 8 km N Arima, Simla Research Station, 260 m, lower montane rainforest, FIT, 24.VI-8.VII.1993, S. Peck, J. Peck leg., one female (BGCO). GUYANA. Potaro-Siparuni: Pataro-Siparuni Iwokara Rain Forest, 70 m, human faeces baited, 13.IV–31.V.2009, S. Phelps leg., one female (OUMNH). SURINAME. Road to Amotopo, km 39, unbaited pitfalls, secondary forest, 29.V–2.VI.1981, J. Huijbregts leg., one female (RMNHL); Sipaliwini W-bank Suriname River opposite Pingpe 4°02ʹ30ʺN 55°27ʹ00ʺW, FIT in forest, 25.I-1.II.2012, M.A. Hielkema leg., one female (RMNHL); road to Amotopo, km 34, primary forest, pitfalls baited with human dung, 28.V-1.VI.1981, J. Huijbregts leg., one female (RMNHL). FRENCH GUIANA. Mont Grand Matoury, 24.I.2013, SEAG leg., two females (ZIN); 7.I.2013, SEAG leg., one male (ZIN); Gourdonville, 1905, E. Le Moult leg., one male and one female (MNHN). BRAZIL. Roraima: Cariri, Vila Napoleão, 4.V.1972, J. Celso leg., one male (CEMT); 10.IV.1972, J. Celso leg., one female (CEMT). Pará: Belém, one female (MNHN).

Diagnosis

Aegidinus guianensis is most similar to Ae. howdenorum but differs from it in having the ventrolateral lobe of the parameres without a subapical tooth (Fig. 1H). From other Aegidinus species it differs in having the phallobase with a ventroapical plate and symmetrical parameres (Figs. 1G, I).

Figure 1. Aegidinus guianensis, holotype, female (a, b, d), female, paratype of Ae. oreibates (c), female from Villa Napoleão, Brazil (e), male, holotype of Ae. oreibates (f), male, paratype of Ae. oreibates (g–i). Habitus in dorsal view (a, c, f), labels (b), female external genitalia (d, e), aedeagus in lateral (h) and ventral view (g), parameres in dorsal view (i).
Figure 1. Aegidinus guianensis, holotype, female (a, b, d), female, paratype of Ae. oreibates (c), female from Villa Napoleão, Brazil (e), male, holotype of Ae. oreibates (f), male, paratype of Ae. oreibates (g–i). Habitus in dorsal view (a, c, f), labels (b), female external genitalia (d, e), aedeagus in lateral (h) and ventral view (g), parameres in dorsal view (i).

Remarks

Paulian (1984) and Colby (2009) drew male characters of Ae. guianensis from the type of Ae. steinheili, which was considered as a synonym of Ae. guianensis after Arrow (1904). Colby (2009) erroneously considered the female specimen with the label “Cayenne”, housed in BMNH, as the holotype of Ae. guianensis and drew the female characters from it. This specimen in our opinion belongs to Ae. howeae Colby or Ae. brasiliensis, but it lacks the abdomen and genitalia. Ae. guianensis was described from British Guiana (nowadays Guyana) and the holotype of it is housed in OUMNH (Figs. 1A, B). It agrees with the original description of the species and its mouthparts were dissected and illustrated by Westwood (1845: plate XII, Fig. 5). The specimen was likely collected by Robert Schomburgk in Pirara on the banks of Lake Amucu (Darren Mann, pers. comm.).

By the shape of the female genitalia and external morphological characters, the type of Ae. guianensis (Figs. 1A, D) cannot be distinguished from Ae. oreibates Colby (Figs. 1C, E), described from a long series mostly from the Guiana Shield and the island of Trinidad. For this reason, we propose that Aegidinus oreibates Colby is a synonym of Aegidium guianense Westwood. Because this long series includes both sexes, the male characters of Ae. guianensis can be reliably based on the males from this series.

Ae. crypticus Colby was described from Guyana from a single male specimen. The difference between Ae. oreibates and Ae. crypticus is, according to the author, that in the later species the median lobes of the parameres are longer and thinner and the shape of the ventral plate of phallobase at the apex is much rounder (Colby 2009). However, the detailed photographs of the parameres provided in the original descriptions (Colby 2009: Figs. 46–49, 59–60) show that these characters are not diagnostic: the median lobes of the parameres are of about the same length and width and the ventral plates are equally rounded. The differences between the photographs are due to the slightly different angle of view and because the illustrated parameres of Ae. oreibates are opened while those of Ae. crypticus are close to each other. Therefore, we propose that Aegidinus crypticus Colby is also a synonym of Aegidium guianense Westwood.

Aegidinus steinheili (Harold, 1880), distinct species

(Figures 2A–G)

Aegidium steinheili Harold, 1880: 42.

Aegidinus guianensis nec Westwood, 1845: 175, auct.: Arrow 1904: 739, 1912: 31; Paulian 1984, 98; Colby 2009: 20.

Type locality. Colombia, Antioquia, San Carlos.

Type material examined

Lectotype. Male (Fig. 2A) at MNHN labelled “S. Carlos / Columbia Aegidinus Steinheili Harold / Col. Benderitter ae. guianense / Aegidinus guganesis (Westw.) R. Paulian det. / Aegidinus steinheili (Har.) R. Paulian det. / Museum Paris / Museum Paris Coll. M. Pic / HOLOTYPE / Aegidinus guianensis”. The labels of this specimen do not agree completely with the data provided by Paulian (1984), but we believe that this is the specimen that Paulian intended to designate as the lectotype. There are cases that Paulian did not properly label his material with type labels (Olivier Montreuil, Antoine Mantilleri, pers. comm.). However, this specimen is the only one bearing Paulian’s handwritten identification labels and it definitely belongs to the type series from San Carlos.

Paralectotypes. COLOMBIA. Antioquia: one female at MNHN labelled “San Carlos / G. J. Arrow Vidit 1903. / Ex. Musæo E. Steinheil / MNHN”; two males at MNHN labelled “San Carlos / G. J. Arrow Vidit 1903. / sp. nov. / Ex. Musæo E. Steinheil / Museum Paris / Aegidinus candezei (Preudhomme de Borre) det. J.B.Colby 2008 / MNHN”; one male at MNHN labelled “Nare (Wallis) / G. J. Arrow Vidit 1903. / Ex. Musæo E. Steinheil / Museum Paris / MNHN 03-2005 ADSmith / MNHN / Aegidinus guianensis (Westwood) det. J.B.Colby 2008”; one female at BMNH labelled “S. Carlos / Aegidinus guianensis, Westw. Agrees with type (♀) / Ex. Musæo E. Steinheil / ex Coll. Oberthur. 1904–175. / Cotype / BMNH / Aegidinus guianensis Westwood INVALID TYPE det. J.B.Colby 2007”; one female at MNHN labelled “San Carlos / G. J. Arrow Vidit 1903. / Ex. Musæo E. Steinheil / Brit. Mus. Accepit. / MNHN”; one female at MNHN labelled “San Carlos / G. J. Arrow Vidit 1903. / Ex. Musæo E. Steinheil / MNHN”; one female at MNHN labelled “San Carlos / G. J. Arrow Vidit 1903. / Ex. Musæo E. Steinheil / Museum Paris / MNHN 03-2005 ADSmith / MNHN / Aegidinus guianensis (Westwood) det. J.B.Colby 2008”; one male at MNHN labelled “San Carlos / G. J. Arrow Vidit 1903. / Steinheili Harold / Ex. Musæo E. Steinheil / Brit. Mus. Accepit. / Museum Paris / MNHN / Aegidinus guianensis (Westwood) det. J.B.Colby 2008”; one male at BMNH labelled “San Carlos / Steinheili Har. […] / Ex. Musæo E. Steinheil / ex Coll. Oberthur. 1904–175. / Cotype / BMHN / Aegidinus guianensis Westwood INVALID TYPE det. J.B.Colby 2007”.

Additional material examined

COLOMBIA. One male at MNHN, Canoas [not traced locality]. One male at IRSNB labelled “Coll. R. I. Sc. N. B. Bresil Steinheil. ex coll. Candèze / Aegidium Alatum Cast. Br. / Aegidinus guianensis (Westwood) det. J.B. Colby 2008”; a mislabelled specimen (see below) most probably collected in San Carlos.

Diagnosis

Aegidinus steinheili is most similar to Ae. petrovi Colby and Ae. teamscaraborum Colby but differs from them by having ventrolateral lobes of parameres long and slender (in lateral view), reasonably longer than dorsomedial lobes (Figs. 2 D–F). The female has a punctation of the body similar to that of the male (Fig. 2C); the external female genitalia as in Fig. 2G.

Figure 2. Aegidinus steinheili, lectotype, male (a, d–f), male with erroneous label ‘Bresil Steinheil. . .’. (b), female, paralectotype (c, g). Habitus in dorsal view (a–c), female external genitalia (g), aedeagus in lateral (f) and ventral view (d), parameres in dorsal view (e).
Figure 2. Aegidinus steinheili, lectotype, male (a, d–f), male with erroneous label ‘Bresil Steinheil. . .’. (b), female, paralectotype (c, g). Habitus in dorsal view (a–c), female external genitalia (g), aedeagus in lateral (f) and ventral view (d), parameres in dorsal view (e).

Remarks

Arrow (1904) considered Ae. steinheili (Harold, 1880) as a synonym of Ae. guianensis (Westwood, 1845) although he didn’t study the genitalia of the specimens. Paulian (1984) and Colby (2009) followed Arrow apparently because they didn’t study the type of Ae. guianensis and thus misinterpreted this species. Examination of the female type of Ae. guianensis and conspecific males (see above) showed that Ae. steinheili is distinct from it and all other Aegidinus species.

Paulian (1984) studied most of the Aegidinus specimens from San Carlos and one from Nare and, based on the external characters, identified them as two species, Ae. steinheili (=Ae. guianensis sensu Paulian) and Ae. candezei. He designated the larger male from San Carlos with the denser punctation of the body as the lectotype of Ae. steinheili (Fig. 2A). We dissected all specimens of this series and found that characters of the male and female genitalia are congruent with the characters of the external morphology. In this material we found two male morphotypes and two female morphotypes and we agree with Paulian that they belong to the two species mentioned above.

In the collection of IRSNB there is a male specimen of this species with the labels “Coll. R.I.Sc.N.B. Bresil Steinheil. ex coll. Candèze / Aegidium Alatum Cast. Br. / Aegidinus guianensis (Westwood) det. J.B.Colby 2008”. It was among a small series of Aegidiellus alatus Laporte de Castelnau having the same printed labels “Coll. R.I.Sc.N.B. Bresil […]” but varying additional data. These labels were put by IRSNB technical staff under previously unlabelled specimens that were in the collection drawer next to the bottom label “Aegidiellus alatus”. The record of Ae. steinheili from Brazil (Colby 2009, as Ae. guianensis) is apparently based on this mislabelled specimen which is very similar to the larger males of the type series (Fig. 2B).

Aegidinus candezei (Preudhomme de Borre, 1886)

(Figures 3A–E, G, H, J, K)

Orphnus candezei Preudhomme de Borre, 1886: 116.

Aegidinus candezei (Preudhomme de Borre); Arrow, 1912: 31; Schmidt, 1913: 76; Paulian 184: 89; Colby 2009: 17.

Aegidinus sunidigea Colby, 2009, syn. nov.

Type locality. Colombia, Chocó.

Type material examined

Lectotype (designated here, Figs. 3A, E), male at IRSNB labelled “Coll. R. I. Sc. N. B. Colombie ex coll Candeze Choco Wallis / det. P. de Borre 1886 Orphnus Candezei ♂ Type / Holotype / Lectotypus Orphnus candezei P.d.B. Frolov des. 2018”.

Paralectotype (Figs. 3F, I, L), female at IRSNB labelled “Coll. R. I. Sc. N. B. Colombie: Choco ex Wallis ex coll. Candeze / det. P. de Borre 1886 Orphnus Candezei ♀ Type / Allotype / Aegidinus candezei (Preudhomme de Borre) det. J.B.Colby 2009 / Paralectotypus Orphnus candezei P.d.B. Frolov des. 2018”.

Additional material examined

COLOMBIA. Boyacá: Otanche, 5.V.1995, Arriágada leg., five females (CEMT). Santander: Puerto Araújo, Los Manantiales Ranch, V.2000, J. Noriega leg., one male (CEMT); Puerto Araújo, Las Marias, 130 m, V.2000, J. Noriega leg., one female (CEMT). Tolima: Mariquita, Municipal forest, 680 m, IV.2000, J. Noriega leg., one male (CEMT). Antioquia: Medellín, forest, pitfall, IV.1998, J. Noriega leg., one male (CEMT).

Diagnosis

Ae. candezei is most similar to Ae. noriegai sp. nov. but can be separated from it by the shorter and less asymmetrical parameres, ventroapical plate of the phallobase wider than long (Figs. 3B–D), and protibia with medioapical tooth.

Figure 3. Aegidinus spp. Aegidinus candezei, lectotype, male (a, e), male (b–d), female (h, k), female, holotype of Ae. sunidigea (g, j); Aegidinus sp., paralectotype of Ae. candezei (f, i, l). Habitus in dorsal view (a, g, h, i); labels (e, f); female external genitalia (j–l); aedeagus in lateral (b) and ventral view (c); parameres in dorsal view (d).
Figure 3. Aegidinus spp. Aegidinus candezei, lectotype, male (a, e), male (b–d), female (h, k), female, holotype of Ae. sunidigea (g, j); Aegidinus sp., paralectotype of Ae. candezei (f, i, l). Habitus in dorsal view (a, g, h, i); labels (e, f); female external genitalia (j–l); aedeagus in lateral (b) and ventral view (c); parameres in dorsal view (d).

Remarks

Aegidinus candezei was described with no explicit information about the number of the examined specimens although it was indicated that both sexes were studied. There are two specimens (Figs. 3A, I) in the IRSNB collection which agree with the original description and might be the only specimens studied by Preudhomme de Borre. These specimens should be considered syntypes since no lectotype had been designated so far. The male specimen bears a red printed label “Holotype” (Fig. 3E) and the female specimen a label “Allotype” (Fig. 3F). These labels were put by the IRSNB technical staff in 1980–2000 (Alain Drumont, pers. comm.). Paulian (1984) mentioned the “holotype” apparently based on these labels. The identity and conspecificity of the two specimens were not questioned by Paulian (1984) and Colby (2009) and the latter author drew the female characters of Ae. candezei from the female syntype. However, we think that the two syntypes are not conspecific. The body punctation and the shape of the genitalia of the female syntype (Figs. 3I, L) do not agree with the female specimens from San Carlos (Figs. 3H, K) belonging to the series of both sexes with trustable locality data. Only the male syntype bears the original locality label “Choco Wallis”. The locality label of the female syntype was added by IRSNB technical staff later and put under an unlabelled specimen. Although it seems probable that the both specimens originate from the Candeze’s collection and might have been collected by Wallis, the locality of the female syntype is unclear since Wallis made extensive expeditions throughout South America (Veitch 1906).

Aegidinus sunidigea Colby was described from a female (Figs. 3G, J) from San Carlos. As discussed above, the series from San Carlos includes two species represented by both sexes, so the female characters of Ae. candezei can be inferred from these specimens with sparse punctation, including the type specimen of Ae. sunidigea. To ensure stability of the nomenclature, we here designate the male syntype (Fig. 3A) as the lectotype of Orphnus candezei andpropose that Aegidinus sunidigea isa synonym of Orphnus candezei.

Aegidinus noriegai Frolov, Akhmetova & Vaz-de-Mello sp. nov.

(Figures 4A–D)

Type locality. Colombia, Antioquia, Medellín.

Type material examined

Holotype. Male (Fig. 4A) at CEMT labelled “COLOMBIA: Antioquia. Porce. Picardia V-1998. pastizal, pitfall. J. Noriega”.

Diagnosis

Ae. noriegai sp. nov. is most similar to Ae. candezei but can be separated from it by the longer and more asymmetrical parameres, ventroapical plate of the phallobase longer than wide (Figs. 4B–D), and the protibia without medioapical tooth.

Description

Male (Figs. 4A). Body length 8.5 mm. Colour uniformly blackish brown. Anterior margin of frontoclypeus with a horn rounded apically.

Pronotum with widely rounded lateral margins, narrower than elytra, 1.6 times wider than length. Posterior angles widely rounded. Anterior margin bordered, border interrupted medially, with feeble gibbosity. Base of pronotum not bordered, with a few large rounded punctures laterally and a few small medially. Pronotal disc feebly excavated anteromedially, with 2 gibbosities in centre. Pronotum punctate with a few large rounded punctures laterally and anteromedially and with minute, feebly visible punctures throughout.

Elytra almost as long as wide, widest medially and rounded apically, with humeral and apical humps. First elytral stria as continuous line, connected basally with undulate line from scutellum to humeral hump. Other striae marked with rows of sparse punctures; punctures somewhat V- and comma-shaped on basal part of elytra becoming smaller toward apices.

Protibia without medioapical tooth.

Aedeagus relatively short and wide, ratio phallobase length/parameres width 1.8 (Fig. 4B). Ventroapical plate of phallobase narrow and long, 1/3 as wide as phallobase and longer than wide (Fig. 4D). Parameres strongly asymmetrical (Figs. 4C, D): right paramere with relatively long medial and lateral processes, medial process relatively straight; left paramere with vestigial lateral process; medial process curved outwards, with small tooth near apex.

Figure 4. Aegidinus spp. Ae. noriegai sp. nov., holotype (a–d), Ae. simulatus, male (e–h). Habitus in dorsal view (a, e); aedeagus in lateral (b, f) and ventral view (d, h); parameres in dorsal view (c, g).
Figure 4. Aegidinus spp. Ae. noriegai sp. nov., holotype (a–d), Ae. simulatus, male (e–h). Habitus in dorsal view (a, e); aedeagus in lateral (b, f) and ventral view (d, h); parameres in dorsal view (c, g).

Distribution

The species is known from a single locality in Antioquia, Colombian Andes (Fig. 7).

Etymology

The new species is named after Jorge Noriega (University of Los Andes, Bogotá, Colombia).

Aegidinus colbyae Frolov, Akhmetova & Vaz-de-Mello sp. nov.

(Figures 5A–F)

Type locality. Peru, Loreto, Iquitos, Sachamama.

Type material examined

Holotype (Fig. 5A, E, D, F), male at CEMT labelled “PERU: LO: IQUITOS SACHAMAMA 100m 22-III-02 CUROE ROTTEN FISH”.

Paratype (Fig. 5B, C), female at CEMT labelled “PERU: LO: IQUITOS SACHAMAMA 100m 22-III-02 CUROE ROTTEN FISH”.

Diagnosis

Ae. colbyae sp. nov. is most similar to Ae. brasiliensis Arrowand Ae. howeae Colby but can be separated from them by the dorsal processes of the parameres being carina-shaped (Fig. 5E, arrowed), while this process is tooth or spur-shaped in Ae. brasiliensis and Ae. howeae.

Description

Male (Fig. 5A). Body length 10.8 mm. Colour uniformly blackish brown. Anterior margin of frontoclypeus with a long horn rounded apically.

Pronotum with widely rounded lateral margins, almost as wide as elytra, 1.5 times wider than length. Posterior angles widely rounded. Anterior margin bordered, border narrowly interrupted medially, with transversal, bimodal gibbosity. Base of pronotum not bordered, with a row of rounded punctures. Pronotal disc feebly excavated anteromedially, with 2 gibbosities in centre. Pronotum punctate with a few large rounded punctures laterally and anteromedially and with minute, feebly visible punctures throughout.

Elytra almost as long as wide, widest medially and tapering apically, with humeral and apical humps. First elytral stria as continuous line, connected basally with undulate line from scutellum to humeral hump. Other striae marked with rows of sparse punctures; punctures V-shaped near base, smaller and irregulary elongate on disc.

Protibia without medioapical tooth.

Aedeagus relatively long and slender, ratio phallobase length/parameres width 2.8 (Fig. 5F). Ventroapical plate of phallobase absent (Fig. 5D). Parameres symmetrical, mediobasal margins of dorsomedial lobes strongly sclerotized (5E), dorsal processes of parameres carina-shaped (Fig. 5E, arrowed).

Figure 5. Aegidinus colbyae, holotype male (a, e, d, f), paratype female (b, c). Habitus in dorsal view (a, b); female external genitalia (c); aedeagus in lateral (f) and ventral view (d); parameres in dorsal view (e).
Figure 5. Aegidinus colbyae, holotype male (a, e, d, f), paratype female (b, c). Habitus in dorsal view (a, b); female external genitalia (c); aedeagus in lateral (f) and ventral view (d); parameres in dorsal view (e).

Female paratype (Fig. 5B) differs from male in having relatively wider elytra, pronotum and head without excavations and armature, and in having a protibial spur. Body length 8.5 mm. External female genitalia as in Fig. 5C.

Distribution

The species is known from a single locality in Peru: Loreto, Iquitos, Sachamama (Fig. 7).

Etymology

The new species is named after Julia Colby (Peggy Notebaert Nature Museum, Chicago, U.S.A.).

Aegidinus cornutus Colby, 2009

(Figures 6A–F)

Aegidinus cornutus Colby, 2009: 18.

= Aegidinus unicus Colby, 2009: 37, syn. nov.

Type locality. Brazil, Mato Grosso, Sinop.

Type material examined

Aegidinus cornutus: Holotype. Male at CMN labelled “SINOP 12°31ʹS, 55°37ʹW BR 163 km 500 a 600 Mato Grosso, BRASIL 350 m IX.1974 Alvarenga & Roppa col. / Coleção M. Alvarenga / H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidinus cornutus Colby HOLOTYPE”.

Paratype. Male at CMN labelled “Oct. 976 BRASIL Minas Gerais Sinópolis Alvarenga - leg. Coll. Martinez / H. & A. HOWDEN COLLECTION ex. A. Martinez coll. / Aegidinus cornutus Colby PARATYPE”.

Additional material examined

BRAZIL. Mato Grosso: Sinop, 12°31ʹS, 55°37ʹW, BR 163 km 500–600, 350 m, IX.1974, Alvarenga, Roppa leg., two males and one female (CEMT). Pará: Redençao, Pinkaiti - Aik, X.1999, P.Y. Scheffler leg., 18 males and nine females (CEMT); Redençao, Pinkaiti - Aik, FIT, 2.XI.1998, P.Y. Scheffler leg., one male (CEMT).

Diagnosis

Ae. cornutus can be easily distinguished by the parameres not separated into dorsomedial and ventrolateral lobes (Figs. 6C–E). External female genitalia as in Fig. 6F.

Figure 6. Aegidinus cornutus, male (a, c, d, e), female (b, f). Habitus in dorsal view (a, b); female external genitalia (f); aedeagus in lateral (c) and ventral view (e); parameres in dorsal view (d).
Figure 6. Aegidinus cornutus, male (a, c, d, e), female (b, f). Habitus in dorsal view (a, b); female external genitalia (f); aedeagus in lateral (c) and ventral view (e); parameres in dorsal view (d).

Remarks

Aegidinus cornutus and Ae. unicus were both described from Sinop, Mato Grosso, Brazil, from a male and a female, respectively. Additional specimens of both sexes with the same locality labels as the holotype and a rather long series of the both sexes from Redençao (Figs. 6A, B) show that the both names belong to the same species. Therefore, a new synonymy is here proposed.

The label of the paratype of Ae. cornutus is wrong. There is no locality “Sinópolis” in Minas Gerais and Sinop has been probably misinterpreted as an abbreviation; if in the original data the state was noted as MGr or MG, it could be confused with Minas Gerais (instead of Mato Grosso). M. Alvarenga has been in Sinop (Mato Grosso), then a just founded town, more than once in the 70’s and he was proud of his extensive collecting done there (personal comments to FZVM in several occasions, 1992-2008).

Aegidinus venezuelensis Colby, 2009

Aegidinus venezuelensis Colby, 2009: 38.

Type locality. Venezuela, Distrito Capital, Caracas.

Type material examined

Holotype. Female at MNHN labelled “Vénézuéla Caracas / EX MUSEO N VAN DE POLL / MUSÉUM PARIS 1936 COLL. A. BOUCOMONT / Aegidinus brasiliensis R. Paulian det. Arr. / Aegidinus venezuelensis Colby HOLOTYPE”.

Remarks

The type of this species cannot be reliably separated from Ae. guianensis and Ae. howdenorum females. Distribution suggests that it may belong rather to Ae. howdenorum but additional material is needed. It should be noted that Ae. howdenorum is known from a pair of specimens; the third female from “Colombia” was tentatively identified as Ae. howdenorum and cannot be separated from females of Ae. guianensis. Considering similarity of the two species—guianensis and howdenorum differ only in a slightly different shape of the parameres, the difference smaller than in other Aegidinus species, — additional material is also needed to clarify if this difference is not due to the geographic variability among populations of one species.

Aegidinus simulatus Colby, 2009

(Figures 4E–H)

Aegidinus simulatus Colby, 2009: 32.

Material examined

PERU. Loreto: Curaray River, Gaseoducto Barrett, 25.II.2008, W. Yawarcani leg., two males and three females (UNMSM). COLOMBIA. Meta: San Martín, Tocancipa Ranch, 520 m, forest, pitfall, IV.2006, J. Noriega leg., one male (CEMT).

Diagnosis

Aegidinus simulatus is similar to Ae. brasiliensis, Ae. howeae and Ae. colbyae sp. nov. in having mediobasal margins of the dorsomedial lobes of the parameres strongly sclerotized but can be separated from them by having the dorsal sides of the parameres strongly overlapping and separated by a slit (Fig. 4F, arrowed).

Description of male

Body length 9.5-10.0 mm. Colour uniformly blackish brown. Anterior margin of frontoclypeus with a horn rounded apically (Fig. 4E).

Pronotum with widely rounded lateral margins, narrower than elytra, 1.6 times wider than length. Posterior angles widely rounded. Anterior margin bordered, border narrowly interrupted medially, with short somewhat bimodal tubercle. Base of pronotum not bordered, with obsolete punctures laterally. Pronotal disc feebly excavated anteromedially, with 2 gibbosities in centre. Pronotum punctate with a few large rounded punctures laterally and transversely elongated punctures anteromedially and with minute, feebly visible punctures throughout.

Elytra almost as long as wide, widest medially and rounded apically, with humeral and apical humps. First elytral stria as continuous line not connected basally with undulate line from scutellum to humeral hump. Other striae marked with rows of sparse elongate punctures.

Protibia without medioapical tooth.

Aedeagus relatively long and slender, ratio phallobase length/parameres width 2.6 (Fig. 4F). Ventroapical plate of phallobase absent (Fig. 4H). Parameres symmetrical, mediobasal margins of dorsomedial lobes strongly sclerotized (4G), dorsal sides of the parameres strongly overlapping and separated by a slit (Fig. 4F, arrowed).

Distribution

The species was described from Aliñahui (Ecuador, Napo Province) and Gigante (Huila, Colombia). The new records are from Curaray River (Peru, Loreto) and San Martín (Colombia, Meta) (Fig. 7).

Figure 7. Aegidinus spp. Locality records map.
Figure 7. Aegidinus spp. Locality records map.

Discussion

Species descriptions

Some nominal species of Aegidinus were described from males, other from females, yet other from both sexes (Arrow 1904, Paulian 1984, Colby 2009). However, the conspecificy of the sexes was not always verified in the last case. We think that the species delimitation and description methodology should be consistent in the Orphninae studies and thus the species should be described from the males only. Identification of the females should be based at least on a series of both sexes collected together, if the more reliable methods (e.g. in copula collecting) are not possible. We have examined a few series of specimens collected in one locality and thus we were able to assess conspecificy of the males and females of some species; it was not yet possible for all nominal species, though.

Taxonomic and diagnostic characters

Aegidinus is unique among the Orphninae in having a large variation of the shape of the gonocoxites (Colby 2009). However, whereas the usefulness of the male genitalia in the classification of the Scarabaeoidea in general and Orphninae in particular is proved by the majority of the taxonomic works, utilization of the female genitalia requires further research. Data about utilization of the female genitalia in the Scarabaeidae at species level are scarce. Özgül-Siemund and Ahrens (2015) showed that the female genitalia can provide species-level taxonomic characters in Sericinae, but these characters pertain to the secondary sclerotisations in the internal genitalia (ductus bursae) rather than to the external genitalia. Similar structures are absent in the Orphninae. Detailed examination of the female genitalia of a reasonable number of Aegidinus specimens dissected by a standardized procedure (Frolov, Akhmetova and Vaz-de-Mello 2017a) confirmed species-specific differences in most species but also showed that the differences between similar species are difficult to interpret unequivocally though these species can be rather reliably distinguished by the shape of the male genitalia. Therefore, we consider the shape of the external female genitalia in Aegidinus as a species-group or subgenus level character rather than species level character. As in the other Scarabaeoidea, description of the species from females may result in an unstable classification.

One of the supposedly informative Aegidinus female genital structures, vaginal palps (stili), proved non-reliable since they are subject to both intraspecific and interspecific variation. We found Aegidinus specimens of different species having partly segmented coxites as well as those having asymmetrical genitalia with only one coxite having a distinct stylus. Some species comprise specimens with stili and without. The mechanism of the development of such aberrations is probably the same as involved in the symphysocery: premature termination of the segmentation during ontogenesis. Symphysocery, the incomplete segmentation of the antennomers, is found in different groups of scarab beetles and is thought to be highly homoplastic (Krell 1992, Ahrens 2006, Frolov and Akhmetova 2016). Stili are known in many higher groups of the Scarabaeoidea (Dupuis 2005) but there is no apparent taxonomic pattern. Stili are also found in the majority of the Orphninae taxa but are subject to reasonable variability, both intraspecific and interspecific (Frolov’s unpublished data). Therefore, we consider segmentation of the coxites (presence or absence of the distinct stili) as a character of low phylogenetic, taxonomic and diagnostic value and thus do not use it in the taxa delimitation or identification.

Phylogenetic relationships of Aegidinus

 Aegidinus was placed in the tribe Aegidiini by Paulian (1984) and this placement was supported by Colby (2009) and Frolov (2012). Paulian (1984) did not discuss the phylogenetic relationships within the Aegidiini. Colby (2009) provided strict consensus trees of the two phylogenetic analyses one of which resulted in the polytomy of the Orphninae genera and another suggested the following relationships within the Aegidiini: ((Aegidinus + Aegidium) + Aegidiellus) + Paraegidium); however bootstrap support for all clades was low. Frolov (2012) suggested that (Aegidinus + Aegidiellus + Paraegidium) represent a lineage with unresolved relationships, which is sister to Aegidium.

Phylogenetic relationships within the Aegidiini should be further investigated. Frolov (2012) provisionally placed in this tribe the monotypical genus Stenosternus Karsch from São Tomé Island (Guinea Gulf), based on the shape of metepisternum, a keel separating basal and anterolateral parts of propleura, shape of the mandibles, and protibia with a medial tooth. Later examination of the additional material (Frolov and Akhmetova 2015) did not support the placement of Stenosternus in the Aegidiini. In the Frolov’s (2012) analysis, the clade (Aegidinus + Aegidiellus + Paraegidium) was supported by the single symplesiomorphy, the absence of the protibial process in males. However, in the present work we showed that the process is present in some Aegidinus species.

The placement of Aegidinus in the Aegidiini (excluding Stenosternus but including the recently described Onorius Frolov et Vaz-de-Mello (2015) is supported by a few putative synapomorphies: a keel separating basal and anterolateral parts of propleura (such keel in Stenosternus can be a homoplastic or plesiomorphic state) (Frolov 2012), tube-shaped phallobase (Frolov 2013), and the stridulatory ridges distinctly curved posteriad (unpublished data of the authors). However, Aegidinus differs significantly from the other genera in a number of characters (see Diagnosis).

Acknowledgements

We are thankful to all curators of the collections who provided access to the specimens. We are especially thankful to Darren Mann (OUMNH) for the information about Robert Schomburgk’s collecting in British Guiana, Oliver Hillert (Schöneiche bei Berlin, Germany) for providing material and help with tracing localities in Venezuela, Auke Hielkema (Paramaribo, Suriname) and Hans Huijbregts (RMNHL) for the help in tracing localities in Suriname. We also thank Daniel Curoe, Jorge Noriega, Celso Godinho, Gerardo Arriágada, Pamela Scheffler and Moacir Alvarenga (in memoriam) for donating specimens used to this study and now belonging to CEMT. Two anonymous reviewers provided comments and corrections that improved the manuscript. This work was supported by the National Council for Scientific and Technological Development of the Ministry of Science, Technology, and Innovation of Brazil, CNPq (405697/2013-9, 440604/2015-0, 400681/2014-5, 306745/2016-0) and partly by Russian Foundation for Basic Research (grant number 19-04-00565 А). The study was completed in the framework of the Russian state research project АААА-А19-119020690101-6.

References cited

 

Ahrens D. 2006. The phylogeny of Sericini and their position within the Scarabaeidae based on morphological characters (Coleoptera: Scarabaeidae). Systematic Entomology. 31: 113-144.

Arrow GJ. 1904. XXVII. Sound-production in the Lamellicorn Beetles. Transactions of the Royal Entomological Society of London. 52: 709-750.

Arrow GJ. 1912. Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. In: Coleopterorum Catalogus Berlin: Junk, W.; p. 1–66.

Colby J. 2009. Monographic revision of the genus Aegidinus Arrow (1904) and generic phylogeny of the world Orphninae (Coleoptera: Scarabaeidae: Orphninae). Insecta Mundi. 76: 1–41.

Dupuis F. 2005. L’abdomen et les genitalia des femelles de coléoptères Scarabaeoidea (Insecta, Coleoptera). Zoosystema. 27: 733–823.

Frolov AV. 2012. Diagnosis, classification, and phylogenetic relationships of the orphnine scarab beetles (Coleoptera, Scarabaeidae: Orphninae). Entomological Review. 2012/10/01; 92: 782–797.

Frolov AV. 2013. Stenosternus Karsch, a possible link between Neotropical and Afrotropical Orphninae (Coleoptera, Scarabaeidae). Zookeys. 335: 33–46.

Frolov AV, Akhmetova LA. 2015. Rediscovery of the enigmatic Stenosternus costatus Karsch (Coleoptera: Scarabaeidae: Orphninae) from São Tomé island. Zootaxa. 4007: 440–444.

Frolov AV, Akhmetova LA. 2016. Revision of the subgenus Orphnus (Phornus) (Coleoptera, Scarabaeidae, Orphninae). European Journal of Taxonomy. 241: 1–20.

Frolov AV, Akhmetova LA, Vaz-de-Mello FZ. 2017a. Revision of the mainland species of the Neotropical genus Aegidium Westwood (Coleoptera: Scarabaeidae: Orphninae). Journal of Natural History. 51: 1035–1090.

Frolov AV, Akhmetova LA, Vaz-de-Mello FZ. 2017b. Revision of the South American genus Paraegidium (Coleoptera: Scarabaeidae: Orphninae). Journal of Natural History. 51: 995–1014.

Frolov AV, Grossi PC, Vaz-de-Mello FZ. 2015. A new species of the genus Aegidium  (Coleoptera, Scarabaeidae) from the Atlantic Forest ecoregion in South America. Zootaxa. 4007: 437–439.

Frolov AV, Vaz-de-Mello FZ. 2015. A new genus and species of Orphninae (Coleoptera: Scarabaeidae) associated with epiphytes in an Andean cloud forest in Ecuador. Zootaxa. 4007: 433–436.

Harold E. 1880. Verzeichniss der von E.Steinheil in Neu-Granada gesammlten coprophagen Lamellicornien. Stettiner entomologische Zeitung. 41: 13-46.

Krell F-T. 1992. Verschmelzung von Antennomeren (Symphysocerie) als Regenfall bei Temnorhynchus repandus Burmeister, 1847, sowie phylogenetische, taxonomische, faunistische und nomenklaturische Anmerkungen zu diversen Taxa dieser Gattung. Deutsche Entomologische Zeitschrift. 39: 295-367.

Özgül-Siemund A, Ahrens D. 2015. Taxonomic utility of female copulation organs in Sericini chafers (Coleoptera, Scarabaeidae), with special reference to asymmetry. Contributions to Zoology. 84: 167-178.

Paulian R. 1984. Les Orphnidae Americains (Coleopteres, Scarabaeoidea). Annales de la Société Entomologique de France (N S). 20: 65-92.

Preudhomme de Borre A. 1886. Liste des Lamellicornes Laparostictiques recueillis par feu Camille Van Volxem pendant son voyage au Brésil et a la Plata en 1872, suivie de la description de dix huit espèces nouvelles et un genre nouveau. Annales de la Societe entomologique de Belgique. 30: 103-120.

Rojkoff S, Frolov AV. 2017. Révision des espèces du genre Aegidium Westwood, 1845, des Petites Antilles (Coleoptera: Scarabaeidae, Orphninae, Aegidiini). Annales de la Société Entomologique de France. 52: 354–368.

Schmidt A. 1913. Coleoptera. Lamellicornia. Fam. Scarabaeidae. Subfam. Aegialinae, Chironinae, Dynamopinae, Hybosorinae, Idiostominae, Ochodaeinae, Orphninae. Bruxelles: V. Verteneuil & L. Desmet.

Veitch JH. 1906. Hortus Veitchii: A history of the rise and progress of the nurseries of Messrs. James Veitch and Sons, together with an account of the botanical collectors and hybridists employed by them and a list of the most remarkable of their introductions. J. Veitch and Sons, London. 684 p.

Westwood JO. 1845. On the lamellicorn beetles which possess exserted mandibles and labrum, and 10-jointed antennae. Transactions of the Entomological Society of London. 4: 155–180.