116 urn:lsid:arphahub.com:pub:619a5b3a-5ec8-5ff7-b0b1-5070a7c17694 urn:lsid:zoobank.org:pub:70C65CC0-001D-487B-A05D-B86A205B9582 Contributions to Entomology CTE 0005-805X 2511-6428 Senckenberg Gesellschaft für Naturforschung 10.3897/contrib.entomol.74.e126799 126799 Research Article Scarabaeoidea Trogidae Taxonomy An illustrated catalogue of South American species of Omorgus Erichson, 1847 (Coleoptera, Trogidae, Omorginae) including a neotype designation and taxonomical changes da Costa-Silva Vinícius silvavinicius92@gmail.com https://orcid.org/0000-0002-4556-3793 1 2 Strümpher Werner P. https://orcid.org/0000-0002-7047-2666 3 Barclay Maxwell V. L. https://orcid.org/0000-0003-4989-2014 4 Vaz-de-Mello Fernando Z. https://orcid.org/0000-0001-9697-320X 2 Department of Zoology & Entomology, University of Pretoria, Private Bag X20, Hatfield, 0028, South Africa Universidade Federal de Mato Grosso (UFMT) Cuiabá Brazil Laboratório de Scarabaeoidologia, Instituto de Biociências, Universidade Federal de Mato Grosso (UFMT), Cuiabá, Mato Grosso, Brazil University of Pretoria Hatfield South Africa Ditsong National Museum of Natural History, PO Box 413, Pretoria 0001, South Africa Ditsong National Museum of Natural History Pretoria South Africa Department of Life Sciences, Natural History Museum, London SW7 5BD, UK Natural History Museum London, SW7 5BD United Kingdom

Corresponding author: Vinícius da Costa-Silva (silvavinicius92@gmail.com)

Academic editor: Vinicius S. Ferreira

 2024 29 07 2024 74 1 81 101 B2861E35-EA80-53AF-A4AD-1BA2816CBD9C 339AB68C-DCB1-435F-B88E-2C42D0A7B05E 04 05 2024 11 07 2024 Vinícius da Costa-Silva, Werner P. Strümpher, Maxwell V. L. Barclay, Fernando Z. Vaz-de-Mello This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. http://zoobank.org/339AB68C-DCB1-435F-B88E-2C42D0A7B05E

An illustrated catalogue of South American species of the genus Omorgus Erichson is presented. Based on this study we propose the following taxonomic changes: Omorgus (Omorgus) capillaceus Scholtz, 1990, syn. nov. and Omorgus (O.) lucidus Pittino, 2010, syn. nov. are proposed as junior subjective synonyms of Omorgus (O.) fuliginosus (Robinson, 1941) and Omorgus (O.) ciliatus (Blanchard, 1847), respectively. Lastly, we designated a male neotype for the ubiquitous New World species Trox suberosus Fabricius, 1775 to replace the missing type.

 Keratin feeding Omorgini Scarabaeoidea South America taxonomy type specimens Introduction

Omorgus Erichson, 1847 (Scarabaeoidea, Trogidae) is a diverse and widespread genus of Trogidae MacLeay, 1819 with c. 150 valid names (Scholtz 1986a; Strümpher et al. 2016; Zidek 2017). After the morphological phylogeny study conducted by Scholtz (1986a) and later supported by Strümpher et al. (2014 – molecular phylogeny), three subgenera are recognized within Omorgus: Afromorgus Scholtz, 1986a with 57 species distributed throughout the sub-Saharan Africa, Oriental, and Palaearctic regions (Kalawate and Strümpher 2024); the monotypic Haroldomorgus Scholtz, 1986a [Omorgus (H.) batesi Harold, 1872] endemic to South America; and the nominotypical Omorgus with 94 species reported from the Neotropical, Nearctic, Palearctic, Oriental and Australasian regions (Huchet and Costa-Silva 2018; Costa-Silva et al. 2021; Strümpher and Kalawate 2023). For a more comprehensive taxonomical overview of the genus, see Strümpher et al. (2016).

From South America, 17 species of Omorgus are recognized after the studies carried out by Vaurie (1962), Scholtz (1990), Pittino (1987, 2010), and Huchet and Costa-Silva (2018), where redescriptions, taxonomic changes, and identification keys for the species can be found. More recently, Costa-Silva et al. (2021) have addressed a modern revision of Brazilian species of Omorgus, presenting for the first time a high-resolution image of all name-bearing types (with the exception of Omorgus suberosus – see results) of the group.

Thus, our main goal is to present an illustrated catalogue of those name-bearing type specimens of South American species of Omorgus whose images have never been published before. Our results, as well as the publication by Smith (2017) and Costa-Silva et al. (2021), include all South American Omorgus types available for consultation. This study will provide researchers around the World who are interested in the South American Omorgus the opportunity to examine the type specimens through high-definition photography, subsidizing the taxonomic and nomenclatural stability of the names studied in the future. Additionally, after studying all type specimens of Omorgus from South America, we propose here Omorgus (O.) lucidus Pittino, 2010 as a junior subjective synonym of Omorgus (O.) ciliatus (Blanchard, 1847), and Omorgus (O.) capillaceus Scholtz, 1990 as a junior subjective synonym of Omorgus (O.) fuliginosus (Robinson, 1941). Also, in order to fix the problem regarding the missing type specimen of Trox suberosus Fabricius, 1775 from Joseph Banks collection (see Hielkema and Hielkema (2019) and Costa-Silva et al. (2021)), a neotype is designated in accordance with Art. 75.3 of the International Code of Zoological Nomenclature (ICZN 1999).

 Material

A total of 899 specimens of Omorgus (including types and non-type material) from the institutions listed below were examined. The curators are presented in brackets.

CEMT Coleção Entomológica de Mato Grosso Eurides Furtado, Cuiabá, Mato Grosso, Brazil (Fernando Z. Vaz-de-Mello)

CMNCCanadian Museum of Nature, Ottawa, Canada (Robert Anderson and François Génier)

CMNCHCarnegie Museum of Natural History, Pittsburgh, Pennsylvania, United States (Robert Androw)

CNCCanadian National Collection of Insectsy, Ottawa, Canada (Patrice Bouchard)

CVMD Collection of Víctor Manuel Diéguez [Private Collection], Peñalolén, Santiago, Chile (Víctor Manuel Diéguez).

DZUPMuseu de Entomologia ‘Pe. Jesus Santiago Moure’, Universidade Federal do Paraná, Curitiba, Paraná, Brazil (Cibele Stramare Ribeiro-Costa).

IADIZA Instituto Argentino de Investigaciones de Zonas Áridas, Mendoza, Mendoza, Argentina (Gustavo Flores and Sergio Roig).

IFMLInstituto Fundación Miguel Lillo, Tucumán, Argentina (Emília Perez).

LACMLos Angeles County Museum of Natural History, Los Angeles, California, United States (Brian Brown)

MACNMuseo Argentino de Ciencias Naturales ‘Bernardino Rivadavia’, Buenos Aires, Argentina (Pablo Mulieri).

MCZCMuseum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, United States (Rachel Hawkins)

MNHNMuséum national d’Histoire naturelle, Paris, France (Olivier Montreuil and Antoine Mantilleri)

MZSPMuseu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (Sônia Casari).

NHMBNaturhistorisches Museum, Basel, Switzerland (Christoph Germann)

NHMUKNatural History Museum, London, United Kingdom (Maxwell Barclay)

RBINSRoyal Belgian Institute of Natural Sciences, Brussels, Belgium (Alain Drumont and Wouter Dekoninck)

RPMI Collection of Riccardo Pittino [Private Collection], Milan, Italy (Riccardo Pittino).

ZMUK Zoologisches Museum, Universität Kiel, Kiel, Germany (Michael Kuhlmann)

 Methods

Label data for primary type specimens are transcribed ipsis litteris in quotation marks (“…”). Forward slashes “/” were used to indicate a new line of text on the same label. Comments and additional information are provided in square brackets “[…]”. All handwritten information is underlined. The non-type material follows the format present by Costa-Silva et al. (2021): COUNTRY (in capital letters); followed by Department, province of state name (in bold letters); city or specific locality (when mentioned in the label); metres above sea level (when mentioned), geographical coordinates (when mentioned), date, name of the collector(s), number of examined specimens, gender and depository. The countries, first-order political divisions and cities and/or specific localities are given in alphabetical order.

All images taken by the authors were made using a Leica model m205C stereomicroscope with MC190 HD image capture system. Photographs that were not taken by the authors are indicated in the captions of the respective images with the respective copyright of the author and the museum.

 Results Systematics

Order Coleoptera

Family Trogidae MacLeay, 1819

Subfamily Omorginae Nikolajev, 2005

Tribe Omorgini Nikolajev, 2005

Animalia Coleoptera Trogidae 6373E2A7-5A8A-5F2C-8F3D-F4F8CF4C1AB0 Omorgus Erichson, 1847 Note.

For morphological description and diagnosis, see Vaurie (1962 – as “species group suberosus”), Scholtz (1990), and Strümpher et al. (2016).

 Type species.

Trox suberosus Fabricius, 1775 (by subsequent designation, Lacordaire, 1856).

 Identification key.

The identification keys of Vaurie (1962) and Scholtz (1990) can be used to identify South American species of Omorgus, while Costa-Silva et al. (2021) can be used for Brazilian species.

 Catalogue of South American species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Omorgus">Omorgus</tp:taxon-name-part></tp:taxon-name></italic> Animalia Coleoptera Trogidae AAE41682-797B-5571-9EB0-5CFA4076CFA2 Omorgus (Haroldomorgus) batesi (Harold, 1872) Trox batesi Harold, 1872: 126 (original description); Arrow 1912: 54 (catalogue); Blackwelder 1944: 218 (catalogue); Vaurie 1962: 154 (redescription). Trox (Omorgus) batesi : Scholtz 1982: 8 (catalogue). Omorgus (Haroldomorgus) batesi : Scholtz 1986a: 362 (phylogenetics); Scholtz 1990: 1418 (redescription); Zidek 2013: 7 (catalogue); Strümpher et al. 2014: 549 (systematics); Strümpher et al. 2016: 62 (systematics); Zidek 2017: 98 (catalogue); Hielkema and Hielkema 2019: 10 (checklist for the Guianas); Costa-Silva et al. 2021: 1997 (review of Brazilian species). Omorgus batesi : Deloya 2005: 122 (checklist). Type specimen examined.

Holotype (♂ MNHN). See Costa-Silva et al. (2021: 1997) for high-quality images of the type specimen. Type locality: “Amazonestrom” [Amazonas River].

 Geographic distribution.

Argentina, Bolivia, Brazil and Paraguay (Zidek 2017; Costa-Silva et al. 2021).

 Non-type examined material

(four specimens additionally to Scholtz (1990) and Costa-Silva et al. (2021)). BRAZIL – Amazonas • 1; Parana da Eva; 8 Nov. 1969; J.M. & B.A. Campbell leg.; CNC. – Mato Grosso • 1; Juina; 1 Oct. 2022; K.R. Ferreira leg; CEMT • 1; Santa Cruz do Xingu, Faz. da Ilha; 9°45'41"S, 52°15'41"W; UV light trap; 11–17 Jan. 2024; D.F.Rodrigues & T.I.Vitor-da-Silva leg.; CEMT. – Rio de Janeiro • 1♀; Magé, 70 km N [north] of Rio de Janeiro; 15 Oct.–10 Nov. 1999; W. Grosser leg.; NHMUK.

 Note.

The monotypic subgenus Haroldomorgus Scholtz, 1986a was established to accommodate Trox batesi Harold, 1872, following a morphological phylogenetic analysis aimed at elucidating the suprageneric relationships within Trogidae (refer to Scholtz, 1986a for details). One of the most distinctive features of Omorgus (Haroldomorgus) species is a complete absence of tubercles or ridges on the pronotum and elytra. Additionally, this species exhibits an oval-shaped scutellum that is not constricted at the base, which distinguishes it from the typical modern concept of Omorgus species. Following a morphological phylogenetic analysis conducted by some of the authors (VCS, WPS, and FZVM – Costa-Silva et al. in press), the genus Omorgus was recovered to be paraphyletic, restoring Haroldomorgus as a sister group to Omorgus + Polynoncus. In the first and only molecular phylogeny of Trogidae performed to date, by Strümpher et al. (2014), no specimens of Omorgus (Haroldomorgus) batesi were used in the analysis. However, we have now obtained fresh specimens to revisit the molecular analysis and finally test the Scholtz’s (1986a) hypothesis and the position of Haroldomorgus within Trogidae.

 Animalia Coleoptera Trogidae A7BB9E96-A795-5872-96D1-BFBFE26C500D Omorgus (Omorgus) badeni (Harold, 1872) Trox badeni Harold, 1872: 83 (original description); Arrow 1912: 54 (catalogue); Vaurie 1962: 147 (as syn. of Omorgus borrei); Scholtz 1982: 9 (as syn. of Omorgus borrei). Omorgus (Omorgus) badeni : Scholtz 1990: 1416 (revalidated, redescribed and lectotype designated); Zidek 2013: 7 (checklist); Zidek 2017: 98 (checklist); Costa-Silva et al. 2021: 2001 (review of Brazilian species). Omorgus badeni : Morrone 2001: 57 (mention, biogeography); Deloya 2005: 122 (checklist); Morrone 2014: 57 (mention, biogeography). Type specimen examined.

Lectotype (RBINS). See Costa-Silva et al. (2021: 2001) for high quality images of the type specimen. Type locality: “Ceará, Brazil”.

 Geographic distribution.

Brazil. For details of distribution, see Costa-Silva et al. (2021). Morrone (2001, 2014) report this species from “Northern Colombia and northwestern (sic!)”, without specific locality. Although plausible, so far, no specimen of Omorgus (O.) badeni from Colombia has been examined by the authors.

 Animalia Coleoptera Trogidae 91E0E208-6F9B-525B-9E65-64F2976E63D1 Omorgus (Omorgus) borrei (Harold, 1872) Trox borrei Harold, 1872: 84 (original description); Bruch 1911: 193 (checklist); Arrow 1912: 54 (catalogue); Blackwelder 1944: 218 (catalogue); Vaurie 1962: 147 (redescription). Trox (Omorgus) borrei : Scholtz 1982: 9 (catalogue). Omorgus (Omorgus) borrei : Scholtz 1986a: 361 (phylogenetics); Scholtz 1990: 1417 (redescription); Zidek 2013: 8 (checklist); Zidek 2017: 98 (checklist); Costa-Silva et al. 2021: 2002 (review of Brazilian species, lectotype designation). Omorgus borrei : Deloya 2005: 122 (checklist, as “borreri”); Morrone 2014: 78 (mention, biogeography). Type specimen examined.

Lectotype (RBINS). See Costa-Silva et al. (2021: 2002) for high-quality images of the type specimen. Type locality: “Montevideo, Uruguay”.

 Geographic distribution.

Argentina, Bolivia, Brazil, Paraguay and Uruguay (Vaurie 1962; Scholtz 1990; Costa-Silva et al. 2021).

 Non-type examined material

(23 specimens additionally to Costa-Silva et al. 2021). ARGENTINA – Chaco • 1♂ and 1 unsexed; Charata; Dec. 1995; S. Bolle leg.; NHMUK • 1; Pampa del Infierno; Set. 1982, A. Martinez leg.; CMNC. – Currientes • 1♀; Bella Vista, Bords du Parana; Dec. 1903, E.R. Wagner leg.;MNHN • 1; 15 Mar. 1827–20 Apr. 1828, d’Orbigny leg.; MNHN. – Misiones • 1; 1932; K.J. Hayward leg. NHMUK. – Santa Fe • 1; Villa Ana; Nov. 1925, K.J. Hayward leg.; NHMUK. – Santiago del Estero • 1♂ and 7 unsexed; El Pinto; Dec. 1956; CNC • 1♀; Icaño, Edges of Rio Salado; Dec. 1910, E.R. Wagner leg.; MNHN.

BOLIVIA • 1; Jansen leg.; NHMUK.

BRAZIL • 1; Matto Grosso (sic!); 1886; P. Germain leg.; MNHN.

PARAGUAY – Presidente Hayes • 2; Nanawa, “Paraguayan Chaco”; Mar. 1927; G.S. Carter leg.; NHMUK.

URUGUAY – Artigas • 1; 20 km SE Artigas; 30°31'S, 56°22'W; 27 Dec. 2002, S. & J. Peck leg.; CMNC. – Montevideo • 1; Montevideo; MNHN. – Without specific locality • 1♀; MNHN.

 Animalia Coleoptera Trogidae 444630C0-E5CC-5052-B7AF-0E8D509751C1 Omorgus (Omorgus) candezei (Harold, 1872) Fig. 1 Trox candezei Harold, 1872: 113 (original description); Burmeister 1876: 259 (as syn. of Trox ciliatus); Bruch 1911: 193 (checklist – as syn. of Trox ciliatus); Arrow 1912: 55 (catalogue – as syn. of Trox ciliatus); Blackwelder 1944: 218 (checklist – as syn. of Trox ciliatus); Vaurie 1962: 153 (redescription). Trox (Omorgus) candezei : Scholtz 1982: 9 (catalogue). Omorgus candezei : Scholtz 1990: 1406 (redescription); Diéguez and Gómez 2004: 94 (checklist); Deloya 2005: 122 (checklist); Gómez 2008: 514 (key and notes for Argentinean species); Zidek 2013: 8 (checklist);Zidek 2017: 99 (checklist); Smith 2017: 84 (notes, distribution). Type specimen examined.

Holotype, by monotypy (RBINS – Fig. 1). First label [red, typeset]: “Coll. R. I. Sc. N. B. / Argentine / Pampas / ex coll. Candeze”. Second label [white]: “det. Harold 1872 / [yellow card, Edgar von Harold`s handwriting] Tr. Candezei / Harold typ.”. Third label [red, typeset]: “TYPE”. Fourth label [white aged with green border]: “Candezei / Har. / Pampas / Ch.”. Fifth label [white, printed]: “Omorgus candezei / (Harold) / det. Scholtz 1988” (Fig. 1C). Type locality: “die Pampas von Südamerika” [the Pampas of South America].

10.3897/contrib.entomol.74.e126799.figure1 D15C0638-8132-54BD-98A1-02782EF2351F

Holotype of Trox candezei Harold, 1872 (now Omorgus candezei). A. Habitus in dorsal view; B. Habitus in lateral view; C. Labels. Photos by Jonathan Brecko (RBINS). Length of specimen: 10.5 mm.

https://binary.pensoft.net/fig/1103863 Note.

According to Harold (1872: 114), the holotype was found in a factory in Verviers (Belgium) in a consignment of sheep’s wool originally from the Pampas region (Argentina).

 Geographic distribution.

Argentina (Vaurie 1962; Scholtz 1990; Smith 2017).

 Non-type examined material

(15 specimens). ARGENTINA – Buenos Aires • 1; C. Bruch leg.; MACN. – Córdoba • 1; [Villa] Yacanto, A. Stevenin leg.; MACN • 1; Calamuchita, “El Sauco”; Apr. 1959, M.J. Viana leg.; CVMD. – Mendoza • 1; Santa Rosa, Reserva de Biosfera Ñacuñan; 22–25 Mar. 2010; G. Arriagada & R. Garcez leg.; CVMD • 1; Santa Rosa, Reserva de Biosfera Ñacuñan; 11–13 Dec. 2002; V.M. Diéguez leg.; IADIZA. – Río Negro • 1; Río Colorado; Dec. 1963; Bachmann leg.; CMNC. – Río Negro • 1; Village de Patagones [=Viedma]; Set. 1829; D’Orbigny leg.; MNHN. – San Luis • 2; 18 km S Ariona; 18–26 Jan. 1982; H. & A. Howden leg.; CMNC • 1; Arizona; Fev. 1980; M.J. Viana leg.; CVMD • 1; Quines; 24 Nov. 1995; Flores & Roig leg.; CVMD • 1; San Geronimo; Fev. 1914; M.J. Viana leg.; CVMD. – Santa Fe • 1; Weiser leg.; MACN. – Santiago del Estero • 1; Lago Mayo; 22 Mar.–22 May 1957, Golbach leg.; IFML. – Tucumán • 1; A. Breyer leg.; MACN.

 Animalia Coleoptera Trogidae E586ADE0-317A-596E-AB87-8FF0D85C27AE Omorgus (Omorgus) ciliatus (Blanchard, 1847) Fig. 3 Trox ciliatus Blanchard, 1847: 190 (original description); Harold 1869: 1088 (catalogue); Harold 1872: 112 (redescription); Bruch 1911: 193 (checklist); Arrow 1912: 54 (catalogue); Blackwelder 1944: 218 (catalogue – as “ciliata”); Vaurie 1962: 152 (redescription). Trox (Lagopelus) ciliatus : Burmeister 1876: 258, 265 (key and diagnosis); Preudhomme de Borre 1886: 59 (key and comments). Trox (Omorgus) ciliatus : Scholtz 1982: 9 (catalogue). Omorgus ciliatus : Scholtz 1990: 1405 (redescription); Diéguez and Gómez 2004: 94 (checklist); Deloya 2005: 122 (checklist); Gómez 2008: 514 (key for Argentinean species); Zidek 2013: 8 (checklist); Zidek 2017: 99 (checklist); Smith 2017: 85 (lectotype designation). Omorgus lucidus Pittino, 2010 (new synonym); Zidek 2013: 12 (checklist); Zidek 2017: 103 (checklist). Type specimen examined.

Lectotype (MNHN). See Smith (2017: 2085) for high-quality images of the type specimen.

 Type locality:

“village the Patagones” [=Viedna, Río Negro, Argentina].

 Type specimen

of Omorgus (Omorgus) lucidus Pittino, 2010 (new synonym) examined. Holotype, by monotypy (♀ RPMI – Fig. 2). First label [white, Riccardo Pittino’s handwriting]: “ARGENTINA / Buenos Ayres / ex Coll. Winkler”. Second label [white with red border, Riccardo Pittino’s handwriting]: “Omorgus ♀ / lucidus n. sp. / HOLOTYPUS / Det. R. Pittino 1980” (Fig. 2C). Type locality: “Buenos Ayres” [=Buenos Aires], Argentina.

10.3897/contrib.entomol.74.e126799.figure2 AA8FD2F4-3E77-5F62-9999-A40E66815405

Holotype of Omorgus lucidus Pittino, 2010. A. Habitus in dorsal view; B. Habitus in lateral view; C. Labels. Photos by Alberto Ballerio (Brescia). Length of specimen: 13.8 mm.

https://binary.pensoft.net/fig/1103864 Remarks.

Omorgus lucidus Pittino, 2010 was described to accommodate a single female specimen closely resembling Omorgus ciliatus (Blanchard, 1847: 190), from “Buenos Ayres” (Argentina) (see Pittino 2010: 80). According to the author, both species are distinguished by “several minor characters” that include the form of clypeal edge, punctures of the forebody and elytral sculpture (which is visible in Omorgus ciliatus and without any trace of tubercle in Omorgus lucidus, according to the author). However, Trogidae species tend to show great intraspecific morphological variation (Costa-Silva, pers. communication) and we deem that Omorgus lucidus morphology falls within the intraspecific variability of O. ciliatus for the reasons specified below.

The diagnostic characteristics cited by Pittino (2010) to separate O. lucidus from O. ciliatus, include the shape of clypeal edge, punctures of the forebody and mainly the elytral sculpture. These features are dubious and, in our opinion, do not allow to confirm the validity of this species based on only a single female specimen, considering the morphological variation in Trogidae.

During an investigation of many South American and European collections, more than 170 specimens of proper O. ciliatus (according to the description provided by Scholtz 1990: 1405) were examined, being 25 males dissected to study. All specimens were studied considering the external and internal morphology, beyond the photographs of both type-species of O. lucidus (Fig. 2) and O. ciliatus (see Smith 2017: 2085).

In conclusion, we found a great variability in O. ciliatus for morphological characters used by Pittino (2010) as diagnostic for O. lucidus, as for example, the elytral sculpture (Fig. 3A–F). According to the author, the elytra of O. lucidus are “scarcely convex with no tubercles” and, after our study, we agree with the author’s statement. However, this character state is also found in a few specimens of O. ciliatus, as shown in Fig. 3A–C, whose aedeagus was identical to the one of the more typical O. ciliatus (Fig. 3G–I). Therefore, we deem that Omorgus (Omorgus) lucidus Pittino, 2010 to be a junior subjective synonym of Omorgus (O.) ciliatus (Blanchard, 1847), that presents a wide distribution in Argentina.

10.3897/contrib.entomol.74.e126799.figure3 E5B9A723-CCA2-5B62-AB97-18820E5AA29C

Details of elytra, dorsal habitus and aedeagus of Omorgus ciliatus (Blanchard, 1847): A. Elytral tubercles present and visible (from Catamarca, Argentina); B. Poorly visible (Salta, Argentina) and C. Elytral tubercles absent (Neuquén). Habitus in dorsal view (D–F) of Omorgus ciliatus specimens and their respective aedeagus (G–I) showing no appreciable variation. Scale bars: 1 mm.

https://binary.pensoft.net/fig/1103865 Geographic distribution.

Argentina and Bolivia [?]. We have examined only one specimen at NHMUK, which bears a small label inscribed ‘Bolivia’ and another label with the accession number ‘67-45’. This number corresponds to a data entry in the museum’s book, which refers to a donation made by Edward Wesley Janson (1822–1891) around 1867. This is probably the same specimen previously studied by Vaurie (1962). However, we treat this single record of Omorgus ciliatus from Bolivia with caution until new and reliable data becomes available.

 Non-type examined material

(171 specimens). ARGENTINA – Buenos Aires, • 1; Baía Blanca; C. Bruch leg.; MACN • 1; Carmen de Patagones; MACN • 1; Medanos; 11 Nov. 1946; Hayward y Willink leg.; IFML. – Catamarca • 2; 120 km N Belén; 26°59'44"S, 66°18'48"W; 3 Fev. 2009; V.M. Diéguez & G. Arriagada leg.; CVMD • 4; Campo Posuelo, 60 km S Punta de Balasto; 8 Fev. 1997; G. Arriagada leg.; CVMD • 2; Punta Balasto; 18 Jan. 1997; G. Arriagada leg.; CVMD • 2; Recreo; Dec. 1928; IFML • 1; María; 27 Fev. 1992; A. Ugarte leg.; NHMUK • 1; C. Bruch leg.; 1 MACN. – Chubut • 2; Biedma, Puerto Madryn, 4 km W costa; 42°47'8"S, 65°2'56"W; 11 Nov. 2011; G. Cheli leg.; IADIZA • 2; Estepa, Trelew; 10 Apr. 2010; D. Rojas Lanus leg.; CVMD • 1; Paleontology Park Bryn Gwyn; Apr.–May 2012; D. Rojas Lanus leg.; CVMD • 1; Península Valdes; 1 Mar. 1998; D. Rojas Lanus leg.; CVMD • 1; Peninsula Valdés, Playa Colombo; 12 Dec. 2001; D. Rojas Lanus leg.; CVMD • 1; Peninsula Valdés, Punta Pardelas; 12 Dec. 2011; D. Rojas Lanus leg.; CVMD • 1; Pto. Madryn, Playa El Doradillo; 23 Oct. 2011; D. Rojas Lanus leg.; CVMD. – Córdoba • 1; Río Cuarto, Las Albahacas; 2 Nov. 2001; R. Gómez leg. CVMD. – La Pampa • 1; Curacó, Puelches, Estância La Gracielita; 21 Oct. 2001; Bárbara Corró Molas leg.; IADIZA • 1; Curacó, Puelches, Estância La Gracielita; 26 Nov. 2002; Bárbara Corró Molas leg.; IADIZA • 1; MACN. – La Rioja • 1; Mascasin; Jan. 1959; CNC. – Mendoza • 6; 17 km E refugio Alvarado, Res. Laguna del Diamante; 35°15'22"S, 69°12'4"W; 19 Nov. 2012; G. Arriágada leg.; CVMD • 1; Desaguadero; 28–29 Fev. 2005; R. Barrera leg.; CVMD • 12; La Valle, Reserva de Telteca; 8–10 Dec. 2002; V.M. Diéguez leg.; CVMD • 1; La Valle, Reserva de Telteca; 9 Dec. 2002; R. Gómez leg. CVMD • 3; Lavalle, Telteca; 10 Dec. 2002; G. Flores leg.; IADIZA • 1; Lavalle, Telteca; 16–17 Dec. 2005; A. Marvaldi, R. Ruiz y G. Flores leg.; IADIZA • 3; Malargüe, Payunia; 35°40'47"S, 68°41'40"W; 6 Jan. 2016; F. Aballay leg.; IADIZA • 8; Malargüe, Payunia; 35°59'59"S, 68°52'55"W; 2 Dec. 2015; F. Aballay leg.; IADIZA • 2; Malargüe, Payunia, Mina Ethel; 36°0'4"S, 68°50'11"W; 4 Jan. 2016; F. Aballay leg.; IADIZA • 4; San Carlos, Laguna del Diamante; 31 Jan. 2016; F. Aballay & F. Jofré leg.; IADIZA • 36; San Carlos, Laguna del Diamante; 31 Dec. 2015; F. Aballay & F. Jofré leg.; IADIZA • 1; San Rafael, “El Sosneado”; MACN • 1; Santa Rosa, Ñacuñan; 15 Nov. 1996; G. Debandi leg.; IADIZA • 1; Santa Rosa, Ñacuñan; 16 Fev. 1982; S. Claver leg.; IADIZA • 1; Santa Rosa, Ñacuñan; 34°2'42"S, 67°54'33"W; 24 Oct. 2002; G. Debandi leg.; IADIZA • 1; Santa Rosa, Ñacuñan; Jan. 1976; S. Roig leg.; IADIZA • 1; Santa Rosa, Ñacuñan, Nov.–Dec. 1998; S. Roig leg.; IADIZA • 12; Santa Rosa, Reserva de Ñacuñan; 11–13 Dec. 2002; V.M. Diéguez leg.; IADIZA • 8; Santa Rosa, Reserva de Ñacuñan; 8–10 Dec. 2002; V.M. Diéguez leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; 10 Mar. 2014; G. Arriadaga leg.; CVMD • 2; Santa Rosa, Reserva Provincial Telteca; 9 Dec. 2002; R. Gómez leg.; IFML • 1; Santa Rosa, Reserva Provincial Telteca; 9 Dec. 2002; R. Gómez leg.; IADIZA • 1; MNHN • 1; Nov. 1946; Moyano leg.; IFML • 1; Dec. 1897; C. Bruch leg.; MACN. – Neuquén • 1; A. Breyer leg.; MACN • 1; Zapala; 12–22 Dec. 1946; Hayward y Willink leg.; IFML. – Río Negro • 1; Aguada Cecílio; MACN • 1; Río Colorado; 13–19 Nov. 1946; Willink leg.; IFML • 3; Río Colorado; Dec. 1990; CMNC • 3; A. Breyer leg.; MACN • 1; H. Richter leg.; MLPA • 1; Weiske leg.; MACN. – Salta • Anta, 33 km NE Joaquín Gonzales; 22–28 Nov. 1979; Willink leg.; IFML • 1; Cachi; MACN. – San Luis • 1; 30 Km NE San Luis; Dec. 2009, P. Wegner leg.; CEMT • 3; Desaguadero; 15–22 Jan. 1997; Jorge Jensen leg; CVMD • 1; Desaguadero, 8–19 Jan. 1996; E. Escobar leg.; CVMD • 1; San Jeronimo; Jan. 1983; CMNC • 1; Santa Rosa; Fev. 1998; Jaime Pizzaro leg.; IADIZA. – Santiago del Estero • 3; El Pinto; Nov. 1956; CNC • 1; La Banda; Oct. 1945; Briones leg.; CMNC. – Tucumán • 1; C. Bruch leg.; MACN.

BOLIVIA [?] – Without further locality • 1; NHMUK.

 Animalia Coleoptera Trogidae ADD0D40B-E883-59D1-A676-7E32E3629EC7 Omorgus (Omorgus) indigenus Scholtz, 1990 Fig. 4 Omorgus (Omorgus) indigenus Scholtz 1990: 1411 (original description); Zidek 2013: 11 (checklist); Zidek 2017: 102 (checklist). Omorgus indigenus : Morrone 2001: 63 (mention); Deloya 2005: 122 (checklist); Morrone 2014: 62 (mention, biogeography). Type specimens examined.

Holotype, by original designation (♂ CMNC). First label [white, typeset]: “ECU: Galap; Espanola / Bahia Manzanilla / 5–10.VI.85, S&J Peck / Prospis grove behind / beach, carrion traps”. Second label [white with black border, printed]: “[QR code] / Canadian Museum of / Musée canadien de la / NATURE / CMNCEN 00011596”. Third label [white with red border, Clarke Scholtz’s handwriting]: Omorgus / indigenous / C.H. Scholtz 1988 / HOLOTYPE”. Type locality: “Bahia Manzanilla, Española Is., Galapagos” [Ecuador].

10.3897/contrib.entomol.74.e126799.figure4 22753C35-15FF-5752-8358-36578E5355AB

Holotype of Omorgus indigenusScholtz 1990. A. Habitus in dorsal view; B. Habitus in lateral view; C. Aedeagus in dorsal view; D. Aedeagus in lateral view; E. Labels. Photos by François Génier (CMNC).

https://binary.pensoft.net/fig/1103866

Paratypes examined: ECUADOR – Galapagos • 15; Española Is., Bahia Manzanilla; Prosopis grove behind beach; carrion traps; 5–10 Jun. 1985; S. & J. Peck leg.; CMNC • 2; same collection data; NHMUK.

 Geographic distribution.

Endemic to Galapagos Islands [Ecuador].

 Non-type examined material

(24 specimens). ECUADOR – Galapagos • 3; Española Is., Bahia Manzanilla; 5–10 Jun. 1985; S. & J. Peck leg.; CMNC • 15; Galapagos, Española Is., Bahia Manzanilla; 25 Apr.–2 May 1985; S. Peck leg.; CMNC • 6; Galapagos, Española Is., Pta. Suarez; 2 Jun. 1992; S. Peck leg.; CMNC.

 Animalia Coleoptera Trogidae 5A39B49D-422A-51EC-89FA-77D9D59DB106 Omorgus (Omorgus) loxus (Vaurie, 1955) Trox loxus Vaurie, 1955: 58 (original description); Vaurie 1962: 147 (redescription); Ratcliffe 1978: 301 (new distribution records). Trox (Omorgus) loxus : Scholtz 1982: 11 (catalogue). Omorgus (Omorgus) loxus : Scholtz 1986a: 361 (phylogenetics); Scholtz 1990: 1416 (redescription), Moragues 2010: 77 (checklist – as “lauxus”); Zidek 2013: 12 (checklist); Zidek 2017: 103 (checklist); Hielkema and Hielkema 2019: 11 (checklist of the Guianas); Costa-Silva et al. 2021: 1999 (review of Brazilian species). Omorgus loxus : Deloya 2000: 67 (checklist); Diéguez and Gómez 2004: 94 (checklist); Deloya 2005: 122 (checklist); Gómez 2008: 515 (key for Argentinean species). Type specimen examined.

Holotype, by original designation (♀ MCZH). See Costa-Silva et al. (2021: 1999) for high-quality images of the type specimen. Type locality: “El Palmar, 16k.W. of Tetzonapa” [Mexico].

 Geographic distribution.

Argentina, Brazil, Colombia, Costa Rica, French Guiana, Mexico, Panama, and Paraguay (Vaurie 1955; Ratcliffe 1978; Scholtz 1990; Costa-Silva et al. 2021).

 Non-type examined material

(three specimens additionally to Costa-Silva et al. 2021). ARGENTINA – La Rioja • 1♀; Patquia; 1933; K.J. Hayward leg.; NHMUK • 1♀; Santiago del Estero, Bords du Rio Salado, D’Icano, Mistol paso; 1909; E.R. Wagner leg.; MNHN.

BRAZIL • 1; Matto Grosso (sic!); 1886; P. Germain leg.; MNHN.

 Animalia Coleoptera Trogidae A182B442-C86E-55DA-B2E2-411DCF6DE621 Omorgus (Omorgus) nocheles Scholtz, 1990 Fig. 5 Omorgus (Omorgus) nocheles Scholtz, 1990: 1412 (original description); Gómez 2008: 515 (key for Argentinean species); Zidek 2013: 14 (checklist); Zidek 2017: 105 (checklist); Smith 2017: 86 (notes). Omorgus nocheles : Deloya 2005: 122 (checklist). Type specimen examined.

Holotype, by original designation (♂ LACM). First label [white, typeset]: “El Bolson, Rio Negro, / ARGENTINA / Oct. 24–25, 1956 / Andor Kovacs”. Second label [white with red border, Clarke Scholtz’s handwriting]: “Omorgus / nocheles / 1988 / Scholtz / HOLOTYPE”. Third label [white, printed]: “[QR Code] / LACM ENT 160322” (Fig. 5E). Type locality: “Argentina, El Bolsón, Río Negro”.

10.3897/contrib.entomol.74.e126799.figure5 6FFC447E-F115-5C50-B1A4-1C2D0634977F

Holotype of Omorgus nocheles Scholtz, 1990. A. Habitus in dorsal view; B. Habitus in lateral view; C. Aedeagus in dorsal view; D. Aedeagus in lateral view; E. Labels. Photos by Brian Brown (LACM). Length of specimen: 15.5–16.5 mm.

https://binary.pensoft.net/fig/1103867 Geographic distribution.

Omorgus (O.) nocheles is known only from two specimens collected in 1956 in El Bolsón, Argentina (Scholtz 1990).

 Animalia Coleoptera Trogidae 7E86444E-7AE6-5F60-B172-EB037C1E731E Omorgus (Omorgus) pastillarius (Blanchard, 1847) Trox pastillarius Blanchard, 1847: 187 (original description); Harold 1869: 1089 (catalogue); Harold 1972: 50 (redescription); Bruch 1911: 193 (checklist); Blackwelder 1944: 218 (catalogue – as “pastillaria”); Vaurie 1962: 149 (redescription). Trox (Chesas) pastillarius : Burmeister 1876: 250, 264 (key and diagnosis); Preudhomme de Borre 1886: 59 (key and comments); Arrow 1912: 59 (catalogue). Trox (Omorgus) pastillarius : Scholtz 1982: 12 (catalogue). Omorgus pastillarius : Diéguez and Gómez 2004: 94 (checklist); Deloya 2005: 122 (checklist); Gómez 2008: 515 (key for Argentinean species). Omorgus (Omorgus) pastillarius : Scholtz 1990: 1404 (redescription). Type specimen examined.

Lectotype (MNHN). See Smith (2017: 87) for high-quality images of the type specimen. Type locality: “en Patagonie, […], en dehors de la baie de San-Blas [Buenos Aires, Argentina]].

 Geographic distribution.

Argentina, Bolivia, and Chile (Vaurie 1962; Scholtz 1990; Smith 2017). Here we present the new country record for Paraguay (National Park Cerro Corá).

 Non-type examined material

(352 specimens). ARGENTINA – Buenos Aires • 3; Bahía Blanca; 6 Set.– 3 Oct. 1832; C. Darwin leg.; NHMUK • 1; Estancia Barran, 30 km SO Viila Lais; Nov. 1946; CMNC • 16–30 Set. 1968; Willink, Terán y Stange leg.;IFML. – Catamarca • 1; 10 km S Andalgalá; May 2004; Pio Brizuela leg.; IADIZA • 1; 6 km Santa María; 6 Dec. 1968; A. Willink y Stange leg.; IFML • 1; Belen, La Cienega; 1926; Weister Wolters leg.; MACN • 1; Belén, Loma Negra; Jan. 1927; IFML • 1; Belén; IFML • 1; Catamarca; Nov. 1983; L. Peña leg.; CMNC • 1; Los Nacimientos; 27°12'S, 66°40'W; 15 Oct. 1997; S. Roig leg.; IADIZA • 1; Punta de Balasto; 18 Jan. 1997; G. Arriágada leg.; CVMD • 1; Recreo; Dec. 1920; IFML • 2; Santa María; 10–17 Dec. 1995; G. Arriágada leg.; CVMD • 1; Santa María; 27 Fev. 1992; NHMUK • 1; C. Bruch leg.; MACN • 1; 11 Dec. 1896; C. Bruch leg.; MACN • 1; MACN • 1; MLPA. – Chubut • Valdés, San Pablo Médano; 42°41'43"S, 64°10'44"W; 23 Aug. 2008; G. Cheli leg.; IADIZA • 2; MACN • 1; May 1905; C. Bruch leg.; MACN. – Córdoba • 1; 24 Oct. 1987; Gonir leg.; MACN • 1; Argüello, Dec. 1956–Jan. 1957; A. Espinal leg.; IFML • 1; Calamuchita, Belgrano; Oct. 1971; CMNC • 1; Pocho Chancaní; 19 Set.–23 Oct. 1993; L. Acosta leg.; IADIZA • 1; Rio Cuarto, Las Albahacas; 17 Nov. 2001; CVMD • 1; MACN • 3; MNHN. – La Pampa • 4; Chical Có, Reserva Provincial La Humada, Puesto Los Toldos; 29 Set.–24 Oct. 2002; N. Gouts leg.; IADIZA • 1; Curacó, Puelches, Estancia La Gracielita; 18 Set. 2002; Martínez leg.; IADIZA • 1; Curacó, Puelches, Estancia La Gracielita, 18 Set. 2002; Bárbara Corró Molas leg.; IADIZA • 14; Curacó, Puelches, Estancia La Gracielita; 21 Dec. 2001; Bárbara Corró Molas leg.; IADIZA • 1; Curacó, Puelches, Estancia La Gracielita; 26 Nov. 2001; Bárbara Corró Molas leg.; IADIZA • 2; Curacó, Puelches, Estancia La Gracielita; 6 Apr. 2000; Bárbara Corró Molas leg.; IADIZA • 1; La Humada, Puesto Los Toldos; 3 Oct. 2002; Maceda leg.; IADIZA • 5; Lihué Calel, Lihué Calel Parque Nacional; 17 Aug. 1996; Pessino leg.; IADIZA • 1; Lihué Calel, Lihué Calel Parque Nacional; 21 Dec. 1992; Bárbara Corró Molas leg.; IADIZA • 3; Lihué Calel, Lihué Calel Parque Nacional; 23 Mar. 1994; Pessino leg.; IADIZA • Reserva Provincial Pichi Mahuida, Salto Anderson; 22 Nov. 2003; Maceda leg.; IADIZA. – La Rioja • 1; Chamical, “las viscacheras”; Fev. 1989; Zunino, Barbero & Luzzatto leg.; CVMD • 1; Mascasin; Jan. 1960; M. Viana leg.; CVMD • 1; Parque Talampaya; 10 Jan. 1997; CVMD • 1; Patquia; Mar. 1948; Bremer leg.; CMNC • 1; Patquia; K.J. Hayward leg.; NHMUK • 1; Talampaya; 15–16 Jan. 1997; IADIZA. – Mendoza • 2; 17 km E refugio Alvarado; 36°15'22"S, 69°12'4"W; 19 Nov. 2012; G. Arriágada leg.; CVMD • 1; El Carrizal; Set. 1974; S. Roig leg.; IADIZA • 1; General Alvear, RP 188; 3 Jan. 2019; S. Roig & R. Carrara leg.; IADIZA • 2; La Cruceta; Jan. 1983; A. Crimi leg.; IADIZA • 6; La Paz, 14 km S Maquinista Levet; 2–14 Jan. 2019; S. Roig & R. Carrara leg.; IADIZA • 1; La Paz, 51 km S de La Paz; 14 Jan. 2015; S. Roig, R. Carrara leg.; IADIZA • 1; La Tosca; 17 Jun. 1976; A. Roig leg; IADIZA • 3; Las Catitas; 10 Dec. 1951; IADIZA • 3; Lavalle, Reserva Telteca; 32°22'59"S, 68°3'14"W; 12 Apr. 2008; L. Muñoz leg.; IADIZA • 2; Lavalle, Telteca; 10 Oct.–3 Dec. 1996; R. Gonzáles leg.; IADIZA • 1; Lavalle, Telteca; 12 May–17 Jun. 1995; Flores & Roig leg.; IADIZA • 2; Lavalle, Telteca; 1–24 Nov. 1995; G. Flores, S. Roig leg.; IADIZA • 1; Lavalle, Telteca; 13 Dec. 1994–3 Fev. 1995; G. Flores leg.; IADIZA • 2; Lavalle, Telteca; 25 Set.–31 Oct. 1995; Flores & Roig leg.; IADIZA • 2; Lavalle, Telteca; 26 Mar.–2 May 1996; Flores & Roig leg.; IADIZA • 1; Lavalle, Telteca; 3 Fev.–14 Mar. 1995; G. Flores leg.; IADIZA • 2; Lavalle, Telteca; 3 Dec. 1996–6 Jan. 1997; Flores & Roig leg; IADIZA • 1; Lavalle, Telteca, 6 Nov.–3 Dec. 1996; Flores & Roig leg.; IADIZA • 1; Malargüe, 5 km E [East] Mina Ethel, Puesto Poso de Agua; 6 Jan. 2016; R. Carrara & G. Flores leg.; IADIZA • 1; Malargüe, 5 km E Mina Ethel, Puesto Poso da Agua; 6 Jan. 2016; R. Carrara & G. Flores leg.; IADIZA • 3; Malargüe, Agua Escondida; Dec. 1975; S. Roig leg.; IADIZA • 1; Malargüe, La Senillosa, Payunia; 20 Jan. 1998; A. Atencio leg.; IADIZA • 4; Malargüe, Ñacuñan; Dec. 1975; S. Roig leg.; IADIZA • 2; Reserva Ecologica Ñacuñan; Dec. 1990; Zunino, Barbero Valdinazzi leg.; CEMT • 2; Reserva Ñacuñán; Dec. 1975; Arturo Roig leg.; CVMD • 1; San Carlos, Laguna del Diamante; 31 Dec. 2015; F. Aballay & F. Jofré leg.; IADIZA • 1; San Rafael, 20 km N Ruta 146; 14 Jan. 2019; S. Roig, R. Carrara leg.; IADIZA • 1; San Rafael, 22 km W 25 de Mayo; 17 Dec. 1998; Flores & Roig leg.; IADIZA • 6; San Rafael, 30 km S Nihuil; 15 Jan. 2019; S. Roig & R. Carrara leg.; IADIZA • 1; San Rafael; MNHN • 1; San Rafael, RP 190, 32 km S Punta de Agua; 4 Jan. 2019; S. Roig & R. Carrara leg.; IADIZA • 1; San Rafael, Solar del Nihuil, 20 km S El Nihuil; 35°10'S, 68°41'W; 24 Fev. 2006; R. Carrara leg.; IADIZA • 2; Santa Rosa, Divisadero; 11 Mar. 2005; A. Scollo leg.; IADIZA • 1; Santa Rosa, El Divijadero; IADIZA • 21; Santa Rosa, Ñacuñan; 12 Dec. 1998; S. Lagos leg.; IADIZA • 2; Santa Rosa, Ñacuñan; 15 Dec. 1983; Videla Puig leg.; IADIZA • 11; Santa Rosa, Ñacuñan; 15 Dec. 1996; S. Lagos leg.; IADIZA • 5; Santa Rosa, Ñacuñan; 16 Fev. 1982; S. Claver leg.; IADIZA • 3; Santa Rosa, Ñacuñan; 16 Fev. 1996; G. Debandi leg.; IADIZA • 10; Santa Rosa, Ñacuñan; 18 Jun.–16 Jul. 1998; S. Lagos leg.; IADIZA • 2; Santa Rosa, Ñacuñan; 20 Apr. 1998; S. Roig leg.; IADIZA • 10; Santa Rosa, Ñacuñan; 20 Nov. 1992; S. Roig leg.; IADIZA • 3; Santa Rosa, Ñacuñan; 24 Apr. 1998; S. Lagos leg.; IADIZA • 1; Santa Rosa, Ñacuñan; 29 Dec. 1997–7 Fev. 1998; S. Lagos leg.; IADIZA • 1; Santa Rosa, Ñacuñan; 4 Fev. 1982; S. Claver leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; 1 Dec. 1981; S. Claver leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; 11–13 Dec. 2002; V.M. Diéguez leg.; CVMD • 1; Santa Rosa, Reserva Ñacuñan; 12 Dec. 1998; S. Lagos leg.; IADIZA • 3; Santa Rosa, Reserva Ñacuñan; 12 Dec. 1998; S. Lagos leg.; CVMD • 2; Santa Rosa, Reserva Ñacuñan; 18 Jul.–16 Aug. 1998; S. Lagos leg.; CVMD • 1; Santa Rosa, Reserva Ñacuñan; 20 Oct.–22 Nov. 1997; S. Lagos leg.; CVMD • 3; Santa Rosa, Reserva Ñacuñan; 24 Set. 1995; C. Campos leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; 26 Mar. 1984; Ruig Videla leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; 27 Jan. 1994; M. Lacomi leg.; IADIZA • 9; Santa Rosa, Reserva Ñacuñan; 3 Dec. 1981; S. Claver leg.; IADIZA • 5; Santa Rosa, Reserva Ñacuñan; 7 Fev. 1998; S. Lagos leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; 9 Nov. 1996; IADIZA • 2; Santa Rosa, Reserva Ñacuñan; Fev. 1974; A. Roig leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; Mar. 1976; F. Roig leg.; IADIZA • 1; Santa Rosa, Reserva Ñacuñan; Apr. 1976; S. Roig leg.; IADIZA • 1; Telteca; 15 Dec. 1994–3 Fev. 1995; Flores & Roig leg.; CVMD • 1; C. Bruch leg.; MACN • 1; MNHN • 13; MLPA • 1; MACN. – Misiones • 2; 1932; K.J. Hayward leg. NHMUK. – Neuquén • 2; Añelo; 24 Nov. 1976; O. de Ferraris leg.; IADIZA • 1; Confluencia, Estación Challacó; C. Bruch leg.; MACN • 2; Plaza Huincul; 29 Oct. 1970; M. Gentili leg. IADIZA • 1; Ramon M. Castro; 25 Set. 1971; M. Gentili leg.; IADIZA • 1; Santa Rosa, Sierra Auca Mahuida; 22 Mar. 2002; Bernardo Parisek leg.; CVMD • 1; Villa La Angostura, Cerro Bayo; 15 Oct. 1982; M. Gentili leg.; IADIZA • 1; C. Bruch leg.; MACN • 5; 1907; Dr. Adolf Lendl leg.; MNHN • 2; Zapala, Cerro Mesa, 20 km E Zapala (Ruta 16); 8 Dec. 2017; E. Stevani, G. Flores, R. Carrara, F. Aballay leg.; IADIZA. – Río Negro • 1; Pringles; Mar. 1895; Koehler leg.; MLPA • 1; Río Colorado; 13–19 Nov. 1946; Willink leg.; IFML • 1; Villa Regina; Oct. 1961; DZUP • 4; Village de Patagones [= Viedma], “salinas d’Andrés Paz”; Mar.–Apr. 1829; A. d’Orbigny leg.; MNHN. – La Rioja • Castro Barros; 18 Fev. 1939; Biraben-Scott leg.; MLPA • 1; MLPA. – Salta • 1; Cafayate (Yacochuya); 6 Apr. 1988; Stange y Willink leg.; IFML • 1; Cafayate, Tolombóm; 9 Nov. 1995; G. Flores leg.; IADIZA • 1; Merán; Mar. 1929; IFML • 1; Piedra del Molino; Dec. 1990; CMNC • 1; Ruiz de los Llanos; Fev. 1947; R. Golbach leg.; IFML. – San Luis • 1; 18 km S Arionza; 18–26 Jan. 1982; H & A Howden leg; CMNC • 3; Belgrano, Fundo El Molle; 15–20 Jan. 2007; José Gómez leg.; CVMD • 2; Belgrano, Fundo El Molle; 20–28 Jan. 2013; G. Arriágada leg.; CEMT • 2; Belgrano, Fundo El Molle; 33°2'22"S, 66°30'47"W; 10–15 Jan. 2009; José Gómez leg.; CVMD • 3; Belgrano, San Isidro; Jan. 2007; E. Abadie leg.; CEMT • 4; Desaguadero; 15 Jan. 1998; Daniel Velasquez leg.; CVMD • 1; Desaguadero; 4–7 Jan. 2008: V.M. Diéguez leg.; CVMD • 4; Jarilla; Dec. 1974; S. Roig leg.; IADIZA • 1; San Jeronimo; Nov. 1974; S. Roig leg; IADIZA • 6; Suyuque; MLPA. – Santiago del Estero • 3; Choya; Apr. 1968; M. Viana leg.; CVMD • 1; Estación Puni-Tajo; 11 Dec. 1939; Biraben - Bezzi leg.; MLPA • 5; Icano, Rio Salado; 1910; E.R. Wagner leg.; MNHN • 1; La Banda; Oct. 1945; Briones-Prosen leg.; CMNC • 1; Lago Muyo; 29 Mar.–12 Apr. 1957; R. Golbach leg.; IFML • 1; Mal Paso; Oct. 1946; Ruiz Huidobro leg.; IFML • 2; Monte Quemado; MACN • 1; Río Salado; Wagner leg.; MLPA • 2; MNHN • 2; Jan. 1952; IADIZA. – Tucumán • 1; Cruz Alta, La Soledad, Cañete; 27 Jan. 1966; Bucher leg.; IFML • 1; MNHN.

BOLIVIA • 3; NHMUK.

CHILE • 8; NHMUK.

PARAGUAY – Amambay • 1; P.N. [National Park] Cerro Corá; 30 Mar. 2003; R. Garcés leg.; CVMD.

 Animalia Coleoptera Trogidae F2920855-5F63-5C0D-B1C5-5BEC367E3C94 Omorgus (Omorgus) persuberosus (Vaurie, 1962) Trox persuberorus Vaurie, 1962: 145 (original description). Trox (Omorgus) persuberosus : Scholtz 1982: 12 (catalogue). Omorgus persuberosus : Deloya 2005: 122 (checklist); Ratcliffe 2015: 189 (checklist from Peru); Casari et al. 2024: 629 (image). Omorgus (Omorgus) persuberosus : Scholtz 1986a: 361 (phylogenetics); Scholtz 1990: 1411 (redescription); Zidek 2013: 15 (catalogue); Zidek 2017: 106 (catalogue); Smith 2017: 87 (distribution data); Costa-Silva et al. 2021: 2005 (review of Brazilian species). Type specimens examined.

Holotype, by original designation (♂ MZSP). See Costa-Silva et al. (2021: 2003) for high-quality images of the type specimen. Type locality: “Ypiranga, São Paulo” [Brazil].

Paratypes. Forty specimens from several localities in different institutions (see Vaurie 1962: 146).

 Geographic distribution.

Argentina, Bolivia, Brazil, Paraguay, Peru, and Uruguay (Vaurie 1962; Scholtz 1990; Costa-Silva et al. 2021).

 Animalia Coleoptera Trogidae BAC76C22-2D50-5D2B-A371-75E418D5B1F8 Omorgus (Omorgus) spatulatus (Vaurie, 1962) Fig. 6 Trox spatulatus Vaurie, 1962: 151 (original description). Trox (Omorgus) spatulatus : Scholtz 1982: 13 (catalogue). Omorgus spatulatus : Deloya 2005: 122 (checklist). Omorgus (Omorgus) spatulatus : Scholtz 1990: 1404 (redescription); Zidek 2013: 16 (checklist); Zidek 2017: 108 (checklist). Type specimen examined.

Holotype, by original designation (♂ NHMB). First label [white aged, typeset]: “Provincia / Buenos Aires”. Second label [white aged, unknown handwriting]: “II [?]”. Third label [white, typeset]: “Museum Frey / Tutzing”. Fourth label [red, Patricia Vaurie’s handwriting]: “TYPE / Trox / spatulatus / Vaurie” (Fig. 6D). Type locality: “Provincia Buenos Aires” [Argentina].

10.3897/contrib.entomol.74.e126799.figure6 AF797C0B-73C4-58B5-8838-205A3572B212

Holotype of Trox spatulatus Vaurie, 1962 (now Omorgus spatulatus). A. Habitus in dorsal view; B. Habitus in lateral view; C. Spiculum gastrale and aedeagus in dorsal view; D. Labels. Photos by Christoph Germann (NHMB). Length of specimen: 10 mm.

https://binary.pensoft.net/fig/1103868 Geographic distribution.

Argentina (Vaurie 1962; Scholtz 1990).

 Non-type examined material

(215 specimens). ARGENTINA – Buenos Aires • 1; La Plata; MLPA • 12; Quéquen Puerto; 7 Mar. 1928; M.D. Jurado leg.; MACN • 1; Quéquen Puerto; G. Pellerano leg.; MACN • 1; Quéquen; Fev. 1937; CEMT • 1; Quéquen; Fev. 1937; DZUP • 12; A. Breyer leg.; MACN • 2; C. Bruch leg. MACN • 175; MACN. – Córdoba • 10; Miramar; MLPA.

 Animalia Coleoptera Trogidae 7227468B-AD2B-59A4-88DB-92E016E0915D Omorgus (Omorgus) suberosus (Fabricius, 1775) Fig. 7 Trox suberosus Fabricius, 1775: 31 (original description); Fabricius 1781: 34 (diagnosis); Fabricius 1787: 18 (diagnosis); Olivier 1789: 6 (diagnosis); Gmelin 1790: 1586 (diagnosis); Herbst 1790: 29 (description); Fabricius 1792: 87 (diagnosis); Fabricius 1801: 111 (diagnosis); Illiger 1802: 332 (catalogue); Schönherr 1806: 118 (catalogue); Castelnau 1840:107 (as ‘tuberosus’ – misspelling); Sturm 1843: 112 (checklist); Blanchard 1847: 190 (catalogue); Lacordaire 1856: 151 (type designation of Omorgus) Harold 1869: 1090 (checklist); Harold 1872: 119 (redescription); Horn 1874: 5 (diagnosis, comments); Burmeister 1876: 257 (diagnosis); Berg 1881: 99 (checklist); Arrow 1903: 516 (notes about fauna of the St. Vincent Island); Bruch 1911: 194 (checklist); Leng 1920: 253 (catalogue); Mutchler 1925: 238 (catalogue of Galapagos Islands); Blatchley 1928: 64 (records for Florida); Leng 1928: 422 (catalogue of New York); Sim 1934: 11 (larvae description); Denier 1936: 205 (natural history); Hayward 1936: 217 (feeding habit); Blackwelder 1944: 219 (catalogue – as ‘suberosa’); Van Dyke 1953: 123 (distribution data); Haaf 1954: 739 (catalogue of Australia); Vaurie 1955: 60 (redescription); Howden and Vaurie 1957: 4 (pronotum and aedeagus drawing); Ritcher 1958: 325 (biology); Vaurie 1962: 144 (redescription); Landin 1963: 4 (record from Cape Verde Is.); Zimsen 1964: 38 (catalogue); Ritcher 1966: 73 (biology); Hatch 1971: 464 (key to the Pacific Northwest beetles); Chalumeau and Gruner 1974: 787 (catalogue of French Antilles); Paulian 1981: 4 (larvae morphology); Young and Hamm 1985: 93 (feeding experiment); Young 2006: 271 (biology); Bouchard et al. 2024: 372 (mentioned as type species). Trox (Omorgus) suberosus : Burmeister 1876: 257 (redescription); Arrow 1912: 62 (catalogue); Scholtz 1982: 13 (systematics). Omorgus suberosus : Erichson 1847: 111 (new combination); Baker 1968: 42 (larvae’s description); Chalumeau 1977: 231 (corrigenda); Scholtz 1986a: 361 (phylogenetics); Scholtz 1986b: 54 (distribution from Australia); Scholtz 1990: 1407 (redescription); Ratcliffe 1991: 157 (redescription); Baraud 1992: 24 (diagnosis); Browne et al. 1993: 199 (phylogeny); Muñoz-Batet and Lopez-Colon 1995: 279 (record from Czech Republic); Páramo 1997: 29–31 (distribution); Costa et al. 1988: 109 (larvae description and illustration); Deloya 2000: 69 (checklist); Ratcliffe 2002: 8 (checklist from Panama); Rosano-Hernandes and Deloya 2002: 32: (natural history); Morón 2003: 408 (checklist); Deloya 2003: 132 (diagnosis and distribution); Diéguez and Gómez 2004: 94 (checklist); Deloya 2005: 122 (checklist); Nikolajev 2005: 322 (larvae characteristics); Löbl and Smetana 2006: 79 (catalogue); Lopes et al. 2007: Mora-Aguilar and Montes de Oca 2009: 575 (distribution); Philips 2009: 4 (association with mites); Krell 2010: 4 (checklist); Carvajal et al. 2011: 161 (checklist); Zidek 2013: 17 (checklist); Verdugo 2014: 212 (mention); Ratcliffe 2015: 189 (checklist from Peru); Ziani et al. 2015: 3 (distribution); Baena et al. 2015: 2, 4 (feeding behaviour); Pittino and Bezdĕk 2016: 54 (checklist); Strümpher et al. 2016: 57 (phylogeny); Zidek 2017: 109 (checklist); Smith 2017: 87 (notes, distribution); Cortez et al. 2017: 4 (natural history); Huchet and Costa-Silva 2018: 565 (new distribution records from South America); Hielkema and Hielkema 2019 (checklist of the Guianas); Miquel 2019: 184, 186 (distribution); Giraldo-Mendoza 2021: 64 (checklist from Peru); Costa-Silva et al. 2021: 2008 (review of Brazilian species). Omorgus (Omorgus) suberosus : Gianizella and Prado 1999: 749–751 (in poultry houses); Lopes et al. 2007: 29–31 (in poultry houses); Strümpher and Kalawate 2023: 518 (catalogue of the Oriental and Palearctic species); Pablo-Cea et al. 2023: 16 (catalogue of El Salvador). Synonyms.

For a comprehensive list, see Pittino and Bezdĕk (2016: 54–55) and Smith (2017: 88).

 Type specimen examined.

Neotype, here designated (♂ NHMUK – Fig. 7). First label [white, printed]: “BRASIL: Rio de Janeiro, / Cabo Frio, I-1991, / F.Z. Vaz-de-Mello”. Second label [red, Vinicius Costa-Silva’s handwriting]: “NEOTYPE / Trox suberosus / Fabricius, 1775 / des. V. Costa-Silva, 2022”. Third label [white with black border, Vinicius Costa-Silva’s handwriting]: “Omorgus / suberosus / Det. V. Costa-Silva, 2022”. Fourth label [white, printed]: “[QR Code] / WORLD / TROGIDAE / DATABASE / WTD0000386” (Fig. 7E). Type locality: “Brasil, Rio de Janeiro, Cabo Frio”.

10.3897/contrib.entomol.74.e126799.figure7 A73C3B1E-6953-5B32-A19B-CF823ACB4560

Trox suberosus Fabricius, 1775 (now Omorgus suberosus), Neotype, here designated (♂ NHMUK): A. Habitus in dorsal view; B. Habitus in lateral view; C. Aedeagus in dorsal view; D. Aedeagus in lateral view; E. Labels. Scale bars: 1 mm.

https://binary.pensoft.net/fig/1103869 Geographic distribution.

Widespread (see below). For details of distribution, see Huchet and Costa-Silva (2018) and Costa-Silva et al. (2021).

Neotype designation of Trox suberosus Fabricius, 1775.Trox suberosus was originally described in 1775 by the Danish entomologist Johan Christian Fabricius (1745–1808) based on an unspecified number of specimens collected from “Brasilia” (Latin spelling meaning Brazil) that he had examined in the collection of Sir Joseph Banks (“Mus. Dom. Banks”). The collection is housed currently in The Natural History Museum, London, UK (Fabricius 1775; Zimsen 1964). Seventy years later, Erichson (1847) described the genus Omorgus, transferring Trox suberosus to the new genus. Even after the proposal of Erichson (1847) and the subsequent designation of Omorgus suberosus as type species of the genus Omorgus by Lacordaire (1856), the species was historically cited as in its original combination by several authors (i.e., Harold 1869; Harold 1872; Burmeister 1876; Arrow 1903; Bruch 1911; Arrow 1912; Leng 1920; Mutchler 1925; Leng 1928; Denier 1936; Hayward 1936; Blackwelder 1944; Van Dyke 1953; Haaf 1954; Vaurie 1955; Ritcher 1958; Vaurie 1962; Zimsen 1964; Ritcher 1966; Hatch 1971; Chalumeau and Gruner 1974; Scholtz 1982). The species remained in the original combination until the systematic study carried out by Scholtz (1986a), where the author proposed Omorgus as a genus based on several synapomorphies (see Scholtz 1986a for more details). Scholtz’s proposal (1986a), which was adopted by recent authors, was supported by the molecular phylogeny conducted by Strümpher et al. (2014).

Omorgus suberosus is native to the New World and is widely distributed throughout South, Central and North America (from Canada to southern Argentina) (Vaurie 1962; Scholtz 1990; Huchet and Costa-Silva 2018; Costa-Silva et al. 2021). In South America Omorgus suberosus is present in all countries except Chile (see Diéguez 2008). It has also been recorded from the Galapagos Archipelago (Ecuador) (Vaurie 1962; Scholtz 1990).

The ubiquitous New World species has also been recorded from other parts of the world (outside of the New World) such as: Australia, Europe (from Belgium, Czech Republic, Spain), North Africa (from Morocco), Southeast Asia (from the Philippines), and on oceanic islands in the Pacific (Fiji, New Caledonia) and Atlantic (Cape Verde, Canary Islands) (Vaurie 1955, 1962; Scholtz 1986b; Scholtz 1990; Batet and López-Colón 1995; Páramo 1997; Ziani et al. 2015; Pittino and Bezdĕk 2016; Huchet and Costa-Silva 2018; Costa-Silva et al. 2021). The human hand may have contributed to the dispersal of Omorgus suberosus to almost all continents through human migrations, and international transport of products mainly via shipping. Harold (1872) mentioned specimens of Trox suberosus found in a wool shipment imported from Argentina in a factory in Verviers (Belgium). With the fast and silent widespread introductions, plus the high degree of intraspecific variability (i.e., colour, size, the shape of elytral tubercles; see Harold 1872) of T. suberosus (originally described from Brazil), a lot of new species names were described for several countries/continents when in fact, they are just morphotypes of the introduced T. suberosus. For the list of synonyms, see Smith (2017) and Zidek (2017).

Omorgus suberosus is also known to be highly opportunistic, and in the absence of carcasses will exploit virtually any other source of keratin (or chitin) present in their environment. This species has been recorded feeding on eggs, dung and chicken feathers in poultry farms in Brazil (Gianizella and Prado 1999; Lopes et al. 2007), locust eggs in Argentina (Baker 1986), as a “potential predator” of the eggs of the turtles in Mexico, Costa Rica and the Galapagos Islands (Rosano-Hernández and Deloya 2002; Baena et al. 2015), and iguana eggs also in Galapagos (Allgower 1979; Rosano-Hernández and Deloya 2002), being considered a risk factor for the survival of these (and other) species considered as vulnerable according to the International Union for the Conservation of Nature (IUCN) (see Baena et al. 2015).

On the compilation of Fabricius types, Zimsen (1964) presented a list mentioning two type specimens of Trox suberosus: one from Joseph Banks’ collection (housed in the British Museum of Natural History - NHMUK) and another from the Zoologisches Museum, Kiel University (ZMUK). Both specimens were meticulously studied by us; however, as first noted by Vaurie (1962), the type specimen of Trox suberosus lodged in Banks’ collection (NHMUK) corresponds to Trox monachus (now Omorgus monachus – Fig. 8) described by Johann Friedrich Wilhelm Herbst in 1790, 15 years after Trox suberosus description. In the description, Fabricius (1775) mentioned “elytris striatis” and later “Elytra minus rugosa […]”, but none of these statements correspond to the specimen standing as “type” of Trox suberosus in Banks’ collection. This statement/description of “elytris striatis” was repeated in all of Fabricius’ subsequent publications that mentioned T. suberosus (Fabricius 1781, 1787, 1792 and 1801; Fig. 9). According to Harold (1872), Trox monachus presents “tuberculata elytrorum omnia, etiam suturalia, rotundam omnino tomentosa” and Vaurie (1962) presents records of this species only from the USA (Florida, Nebraska, Kansas, Oklahoma and Texas); the type of Trox suberosus is from Brazil, probably Rio de Janeiro (see Papavero 1971). The other specimen mentioned by Zimsen (1964), from ZMUK (Fig. 10), was studied through detailed photographs kindly provided to us by Dr. Michael Kuhlmann (ZMUK). With these high-resolution images from various angles, it was possible to confirm, without any doubt, that the specimen from ZMUK mentioned as a Trox suberosus’ syntype by Zimsen (1964) is a specimen that belongs to the modern concept of the genus Trox, and not Omorgus. The identification to species level was not conclusive. However, considering that the species described by Fabricius comes from the Banks Collection, there is no reason (or evidence) to consider the specimen housed in ZMUK as part of the type series. The label with Fabricius’ handwriting saying “suberosus” on the specimen from ZMUK was probably put on later and corresponds to either a misidentification of the spe­cies or an inadvertent misspelling (between “sabulosus” and “suberosus”) of a specimen from his own collection.

10.3897/contrib.entomol.74.e126799.figure8 AFC42BBA-AC71-5A81-9664-05E1875B2E1F

Supposed syntype of Trox suberosus Fabricius, 1775 (=Trox monachus Herbst, 1790) from Joseph Banks collection (NHMUK): A. Habitus in dorsal view; B. Patricia Vaurie’s handwritten note about the type specimen; C. Type labels. Scale bar: 1 mm. Photo by Keita Matsumoto (NHMUK).

https://binary.pensoft.net/fig/1103870 10.3897/contrib.entomol.74.e126799.figure9 80D2BB4C-0B57-5D92-9684-6438F5A88861

Trox suberosus’ descriptions provided by Johan Christian Fabricius in his publications of A. 1775; B. 1781; C. 1787; D. 1792; and E. 1801.

https://binary.pensoft.net/fig/1103871 10.3897/contrib.entomol.74.e126799.figure10 2B1E1B15-226A-5E31-B6CC-CC7D6566DB2D

Alleged syntype of Trox suberosus Fabricius, 1775 (=Trox sp.) from Fabricius collection (ZMUK): A. Habitus in dorsal view; B. Habitus in fronto-lateral view; C. Type labels. Photo by Michael Kuhlmann (ZMUK).

https://binary.pensoft.net/fig/1103872

Since Vaurie (1962), the name-bearing type specimen of Trox suberosus (mainly that from NHMUK) has been cited as “problematic” (i.e., Vaurie 1962; Hielkema and Hielkema 2019; Costa-Silva et al. 2021), because the specimen does not correspond to the original description. In the case here presented, we believe that all specimens widely recognized and identified as Trox suberosus were based on “common sense” and on a universally recognized concept, but not based on a proper name-bearing type. As presented here and well-documented in the literature (see Vaurie 1955), both Omorgus suberosus and O. monachus are morphologically well-defined, and universally recognized as different species, and should be treated as such. Due to its non-compliance with the description and the stated type locality, we here propose that the specimen housed in the Banks collection under the name suberosus Fabricius is a specimen of the North American Trox monachus which has been mistakenly substituted with the original material of Trox suberosus from Brazil, and that the original type material of Trox suberosus has been lost. Such a substitution is particularly credible in the Joseph Banks collection, since the specimens do not have locality labels or determination labels, and the handwritten name labels, and labels indicating ‘type’, are pinned onto the drawers and not attached to the specimens. Hence a misplaced specimen would not be easy to recognize as such except by comparison with the original description.

In order to establish the single name-bearing type specimen of the widespread Trox suberosus, we propose the designation of a neotype. Our proposed neotype specimen (Fig. 7) morphologically match with the original description provided by Fabricius (1775; Fig. 9A) and fulfils the qualifying conditions of ICZN Article 75.3 (ICZN 1999). To avoid possible future confusion and to maintain the type locality as consistent as possible, and to comply with ICZN Article 75.3.6, we here select as the neotype a specimen from a similar locality as the presumed lost specimen collected by Joseph Banks during his voyage across the globe on the Endeavour (1768‒1771) (Zimsen 1964; Papavero 1971). According to Papavero (1971), “Brasilia, Dom. Banks of Fabricius” refers to the collection made by Sir. Joseph Banks in Ilha Rasa, off the coast near the Guanabara Bay (Rio de Janeiro) on December 7th, 1768 (for details, see Banks 1896; also see Cupello et al. 2023 for additional information about Joseph Bank collection). For detailed diagnoses and redescriptions of Omorgus suberosus, see Harold (1872), Vaurie (1962) and Scholtz (1990); and for additional high-quality images see also Huchet and Costa-Silva (2018) and Costa-Silva et al. (2021).

 Animalia Coleoptera Trogidae 15C23DA4-41D3-58C1-A43F-FCB09D3875C4 Omorgus (Omorgus) triestinae Pittino, 1987 Omorgus triestinae Pittino, 1987: 378; Scholtz 1990: 1407 (as new synonym of Omorgus suberosus), Zidek 2013: 17 (catalogue – as synonym of Omorgus suberosus); Smith 2017: 88 (as synonym of Omorgus suberosus); Zidek 2017: 109 (catalogue – as synonym of Omorgus suberosus); Huchet and Costa-Silva 2018: 559 (reinstated as a valid name); Costa-Silva et al. 2021: 2016 (review of Brazilian species). Type specimens examined.

Holotype, by original designation (♂ MNHN). See Costa-Silva et al. (2021: 2015) for high quality images of the type specimen. Type locality: “Brésil, Minas-Geraes” [Brazil].

Paratypes. Same data label of the holotype (11 MNHN).

 Geographic distribution.

Bolivia, Brazil, and Paraguay (Huchet and Costa-Silva 2018; Costa-Silva et al. 2021).

 <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Omorgus">Omorgus</tp:taxon-name-part> (<tp:taxon-name-part taxon-name-part-type="subgenus" reg="Omorgus">Omorgus</tp:taxon-name-part>) <tp:taxon-name-part taxon-name-part-type="species" reg="capillaceus">capillaceus</tp:taxon-name-part></tp:taxon-name> Scholtz, 1990 a new junior synonym of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Omorgus">Omorgus</tp:taxon-name-part> (<tp:taxon-name-part taxon-name-part-type="subgenus" reg="Omorgus">Omorgus</tp:taxon-name-part>) <tp:taxon-name-part taxon-name-part-type="species" reg="fuliginosus">fuliginosus</tp:taxon-name-part></tp:taxon-name> (Robinson, 1941). Animalia Coleoptera Trogidae 5EABDD67-70CF-585E-881D-66A15D4697CB Omorgus (Omorgus) fuliginosus (Robinson, 1941) Fig. 11 Trox (Omorgus) fuliginosus Robison, 1941: 134 (original description); Blackwelder and Blackwelder 1948: 31 (checklist). Trox fuliginosus : Vaurie 1955: 66 (redescription); Vaurie 1958: 45 (distribution); Blackwelder 1973: 35 (checklist); Ratcliffe 1978: 301 (distribution). Omorgus fuliginosus : Baker 1968: 43 (larvae description); Deloya 1992: 3 (natural history); Deloya 1996: 43 (natural history); Morón et al. 1998: 91 (distribution); Deloya 2000: 66, 73 (checklist); Smith 2003: 6 (checklist); Deloya 2003: 130 (diagnosis and distribution); Deloya 2005: 122, 124, 128 (key); Mora-Aguilar and Montes de Oca 2009: 575 (distribution). Omorgus (Omorgus) fuliginosus : Zidek 2013: 10 (checklist); Zidek 2017: 101 (checklist); Pablo-Cea et al. 2023: 16 (catalogue of El Salvador). Omorgus (Omorgus) capillaceus Scholtz, 1990: 1413 (new synonym): Deloya 2005: 122 (checklist); Zidek 2013: 8 (checklist); Zidek 2017: 99 (checklist). Type specimen examined.

Holotype, by original designation (♂ USNM – Fig. 11). First label [white aged, typeset]: “New Braunfels / IV.10 ’02 Tex”. Second label [white aged, typeset]: “H. Mittendorf / Collector”. Third label [red, printed]: “Type No / 55468 / USNM”. Fourth label [red, Mark Robinson’s handwriting]: “HOLOTYPE / Trox / fuliginosus / Mark Robinson”. Sixth label [white, printed]: “USNMENT / [QR Code] / 01474112” (Fig. 11C). Type locality: “New Braunfels” [Texas, United States].

10.3897/contrib.entomol.74.e126799.figure11 501E6441-5C5E-55B5-ACF7-6A376FA85A20

Holotype of Trox fuliginosus Robinson, 1941 (now Omorgus fuliginosus). A. Habitus in dorsal view; B. Habitus in lateral view; C. Labels. Photos by Robert Finn (USNM). Length of specimen: 14–15 mm.

https://binary.pensoft.net/fig/1103873 Type specimen

of Omorgus (Omorgus) capillaceus Scholtz, 1990 (new synonym) examined. Holotype, by original designation (♀ CMNCH – Fig. 12). First label [white aged, unknown’s handwriting]: “Botoga / Colombia / S. Amer.”. Second label [white, typeset]: “Carn. Mus. / Acc. 2275”. Third label [white, printed]: “[QR code] / CMNCH-IZ / 724,596”. Fourth label [white with red border, Clark Scholtz`s handwriting]: “Omorgus / capillaceus / C.H. Scholtz 1988 / HOLOTYPE” (Fig. 12C). Type locality: “Bogota, Colombia”.

10.3897/contrib.entomol.74.e126799.figure12 C4BA82C9-09BB-5DA5-904B-9D35C6AC0731

Holotype of Omorgus capillaceus Scholtz, 1990. A. Habitus in dorsal view; B. Habitus in lateral view; C. Labels. Photos by Vanessa Verdecia (CMNCH).

https://binary.pensoft.net/fig/1103874 Remarks.

Omorgus (Omorgus) capillaceus was one of the species described by Scholtz in 1990 during his monographic revision of the Trogidae of South America. This species was identified in the collection of the Carnegie Museum of Natural History (CMNCH) and described based on a single female specimen, distinguished by its morphological differences from other South American species. Scholtz noted that O. capillaceus was similar to O. monachus (Herbst, 1790) and O. fuliginosus (Robinson, 1941), both of which are North American species. However, he highlighted a diagnostic feature that distinguishes O. capillaceus from other similar-looking species: the absence of a velutinous covering on the elytral disc, which is present in its relatives.

Upon reviewing the holotype of O. capillaceus, it became evident that the specimen is merely a worn variation of O. fuliginosus. While we concur with Scholtz’s observation of the glabrous elytral disc, this feature alone is insufficient for distinguishing Trogidae species without considering male genitalia. Trogidae specimens often exhibit significant intraspecific variation, and it is not uncommon to encounter specimens that are partially or entirely glabrous (or worn) in collections (V. Costa-Silva, personal observation).

Another factor that may have led Scholtz to describe a new species is the specimen’s locality: Bogota, Colombia. The holotype of O. capillaceus was associated with the label number “2275” (Fig. 12C), which corresponds to specimen from the Henry Klages Collection, acquired by the CMNCH on May 28, 1903 (Robert Androw, personal communication to VCS). However, as documented by Nearns and Androw (2013) and Costa-Silva et al. (2024), the Henry Klages collection has been noted for containing numerous mislabelled Nearctic and Neotropical specimens, including some cases of misidentification within the Trogidae (see Costa-Silva et al. 2024 for examples). Omorgus fuliginosus is known from the United States, Mexico, Guatemala, El Salvador, and Costa Rica (Vaurie 1955; Ratcliffe 1978; Deloya 2000), making its record in Colombia questionable, but not impossible.

Therefore, we conclude that O. capillaceus should be considered a junior subjective synonym of O. fuliginosus, having been described based on a worn and mislabelled specimen from Bogotá, Colombia.

 Geographic distribution.

Costa Rica, El Salvador, Guatemala, Mexico, and United States (Vaurie 1955; Ratcliffe 1978; Deloya 2000).

 Non-type examined material

(9 specimens). MEXICO – Veracruz • 5; Carr. Actopan, Passando la Desv. Idolos; 30 Aug. 1994; L. Arellano & R. Sanchez leg.; CEMT • 1; Carr. Alto Lucero, Km 2 Desviación la Concepción; 18 Oct. 1994; L. Arellano & R. Sanchez leg.; CEMT • 1; Carretera Xalapa-Alto Lucera, 1 km de Espinal; 31 Aug. 1994; L. Arellano & R. Sanchez leg.; CEMT • 1; Catemaco, Parq. de la Flora y Fauna Silvestre tropical; 29 Apr. 1990; F. Capistran leg.; CEMT • 1; Plan de Hidalgo, Mpio de Papantla; 19 Mar. 1997; R. Sanchéz & M.E. Favila leg.; CEMT.

 Author contributions

Vinícius da Costa-Silva: Conceptualization, Methodology, Investigation, Resources, Data Curation, Writing - Original draft, Writing - Review and Editing, Visualization, Supervision, Project administration. Werner P. Strümpher: Methodology, Investigation, Resources, Writing - Review and Editing, Supervision. Maxwell V.L. Barclay: Methodology, Investigation, Writing - Review and Editing, Visualization. Fernando Z. Vaz-de-Mello: Investigation, Resources, Writing - Review and Editing, Visualization, Supervision.

 Acknowledgements

The authors thanks Michal Geiser for his help with the literature and catalogues about NHMUK specimens. We also wish to thank Michael Kuhlmann and Ulrich Irmler (ZMUK), Keita Matsumoto (NHMUK), Robert Androw and Vanessa Verdecia (CMNCH), François Génier (CMNC), Jonathan Brecko (RBINS), Brian Brown (LACM), Alberto Ballerio (Brescia), Robert Finn (USNM), Christoph Germann (NHMB) for providing further information and/or photographs of the types housed on their respective institution. VCS also thanks the Ernst Mayer Grants (Harvard University) and Stewart and Jarmila Peck Visiting Scientist Awards in Entomology (Canadian Museum of Nature) for the financial support to visit the Muséum national d’histoire Naturelle (Paris, France), The British Museum of Nature (London, UK), and the Canadian Museum of Nature (Ottawa, Canada). VCS thanks the Post-Doctoral Fellowship Programme from University of Pretoria and Prof. Catherine Sole (University of Pretoria, South Africa) for the provision of all laboratory facilities. We also extend thanks to Alberto Ballerio (Brescia) and Gimo M. Daniel (The National Museum, South Africa) for providing helpful comments and suggestions that significantly improved the manuscript.

 References AllgowerK (1979) Effect of the Scarab beetle Trox suberosus on the hatching success of the east Pacific green turtle Chelonia mydas agassizi in the Galapagos Islands. Charles Darwin Research Station Annual Report 152–154. ArrowGJ (1903) On the laparostict Coleoptera of Granada and St. Vincent (W. Indies).Transactions of the Entomological Society of London1903: 509–520. https://doi.org/10.1111/j.1365-2311.1903.tb01144.x ArrowGJ (1912) Coleopterorum Catalogus auspiciis et auxilio W. Junk editus a S. Schenkling. Pars 43.Scarabaeidae: Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. Berlin: W. Junk, 66 pp. BaenaMLEscobarFHalffterGGarcía-ChávezJH (2015) Distribution and feeding behavior of Omorgus suberosus (Coleoptera: Trogidae) in Lepidochelys olivacea turtle nests. PLOS ONE 10(9): e0139538. https://doi.org/10.1371/journal.pone.0139538 BakerCW (1968) Larval taxonomy of the Troginae in North America with notes on biologies and histories (Coleoptera: Scarabaeidae).Bulletin of the United States National Museum279(279): 1–79. https://doi.org/10.5479/si.03629236.279.1 BanksJ (1896) Chapter II: Rio de Janeiro. In Hooker JD (Ed.) Journal of the Right Hon. Sir Joseph Banks during Captain Cook’s first voyage in H.M.S. Endeavour in 1768–71 to Tierra del Fuego, Otahite, New Zealand, Australia, the Dutch East Indies, etc. Mac Millan & Co.Ltd., London & N. York, 406 pp.[, 2 pls., 4 maps.] https://doi.org/10.5962/bhl.title.56795 BaraudJ (1992) Coléoptères Scarabaeoidea d’Europe.Féderation Française des Sociétés de Sciences Naturelles78: 1–856. https://doi.org/10.3406/linly.1992.11007 BatetJMLópez-ColónJI (1995) Primer registro centroeuropeo de Omorgus suberosus (Fabricius, 1775) (Coleoptera, Trogidae).Nouvelle Revue d’Entomologie12(4): 279. BergC (1881) Insectos. Informe oficial de la comision científica agregada al Estado Mayor General de la Expedicion al Río Negro (Patagonia) realizada en los meses de Abril, Mayo y Junio de 1879, bajo las órdenes del General D. Julio A. Roca (com 16 láminas). Entrega I. – Zoología (con 4 láminas). Buenos Aires: Ostwald y Martinez, [xxiv +] 168 [+ 1 (Errores tipográficos)] pp. [+ 4 pls.] BlackwelderRE (1944) Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Part 2. Scarabaeidae, Troginae.Bulletin of the United States National Museum185: 218–219. https://doi.org/10.5479/si.03629236.185.2 BlackwelderREBlackwelderRM (1948) Fifth supplement to the Leng Catalogue of Coleoptera of America, north of Mexico. John D. Sherman; Mt.Vernon, New York, 87 pp. BlanchardCÉ (1847) Insectes de l’Amérique Méridionale. In: d’OrbignyABlanchardCÉBrulléA (Eds) Voyage dans l’Amérique Méridionale.Vol. 6, part 2, livraison 90. Paris (France): P. Bertrand, 185–232. BlatchleyWS (1928) The Scarabaeidae of Floridae.The Florida Entomologist12(4): 63–65. https://doi.org/10.2307/3492997 BouchardPBousquetYDaviesAECaiC  (2024) On the nomenclatural status of type genera in Coleoptera (Insecta).ZooKeys1194: 1–981. https://doi.org/10.3897/zookeys.1194.106440 BousquetY (2016) Litterature Coleopterologica (1758–1900): a guide to selected books related to the taxonomy of Coleoptera with publication dates and notes.Zookeys583: 1–776. https://doi.org/10.3897/zookeys.583.7084 BruchC (1911) Catálogo sistemático de los Coleópteros de la República Argentina. Pars IV: Familias Lucanidae, Scarabaeidae (Coproini-Cetonini), Passalidae. Revista del Museo de La Plata, Tomo XVII, 181–225. https://doi.org/10.5962/bhl.title.8799 BrowneDJScholtzCHKukalova-PeckJ (1993) Phylogenetic significance of wing characters in the Trogidae (Coleoptera: Scarabaeoidea).African Entomology1(2): 195–206. BurmeisterHCC (1876) Die argentinischen Arten der Gattung Trox F.Stettiner Entomologische Zeitung37: 241–268. CarvajalVLVillamarínSCOrtegaAMA (2011) Escarabajos del Ecuador: Principales géneros. Instituto de ciências biológicas de la Escuela Politécnica Nacional, Série Entomología No. 1. Quito, Ecuador., [xviii +] 350 pp. CasariSABiffiGIdeS (2024) Coleoptera Linnaeus, 1758. In: RafaelJAMeloGARCarvalhoCJBCasariSConstantinoR (Eds) Insetos do Brasil: Diversidade e Taxonomia.2ª ed. Instituto Nacional de Pesquisas da Amazônia, Manaus, 575–698. https://doi.org/10.61818/56330464c31 Castelnau Comtede (1840) Histoire naturelle des animaux Articulés, Annélides, Crustacés, Arachnides, Myriapodes et Insectes. Histoire naturelle des Insectes Coléoptères. Tome deuxième. Paris (Duménil), 563 pp. https://doi.org/10.5962/bhl.title.47104 ChalumeauF (1977) Contribution à l’étude des Scarabaeoidea des Antilles (Corrigenda et Addenda aux Scarabaeoidea des Antilles françaises).Bulletin Mensual de la Société Linnéenne de Lyon8: 231–240. ChalumeauFGrunerL (1974) Scarabaeoidea des Antilles francaises. Annales de la Société Entomologique de France (n.s.)10(4): 781–819. https://doi.org/10.1080/21686351.1974.12278526 CortezVVerdúJROrtizAJHalffterG (2017) Identification and evaluation of semiochemicals for the biological control of the beetle Omorgus suberosus (F.) (Coleoptera: Trogidae), a facultative predator of eggs of the sea turtle Lepidochelys olivacea (Eschscholtz). PLOS ONE 12(2): e0172015. https://doi.org/10.1371/journal.pone.0172015 CostaCVaninSACasariSA (1998) Larvas de Coleoptera do Brasil. Museu de Zoologia da Universidade de São Paulo, São Paulo, 282 pp [+ 165 pls]. Costa-SilvaVStrümpherWPVaz-de-MelloFZ (2021) Review of the Brazilian species of Omorgus Erichson, 1847 (Coleoptera: Trogidae: Omorginae).Journal of Natural History54(31–32): 1993–2024. https://doi.org/10.1080/00222933.2020.1833999 Costa-SilvaVStrümpherWPThyssenPJVaz-de-MelloFZ (2024) Taxonomic revision of the South American genus Polynoncus Burmeister, 1876 (Coleoptera: Scarabaeoidea: Trogidae).Journal of Natural History58(1–4): 14–166. https://doi.org/10.1080/00222933.2023.2260060 Costa-SilvaVStrümpherWPThyssenPJVaz-de-MelloFZ (in press) Phylogenetic systematics of the keratin-feeding genus Polynoncus Burmeister, 1876 (Coleoptera: Scarabaeoidea: Trogidae). Zoological Journal of the Linnean Society. DeloyaC (1992) Necrophilous Scarabaeidae and Trogidae beetles of tropical deciduous forest in Tepexco, Puebla, Mexico.Acta Zoologica Mexicana52: 1–13. https://doi.org/10.21829/azm.1992.49521952 DeloyaC (1996) Los Macro-Coleopteros necrofilos de Tepoztlan, Morelos, Mexico (Scarabaeidae, Trogidae, Silphidae).Folia Entomologica Mexicana97: 39–54. DeloyaC (2000) Distribucion de la familia Trogidae en México (ColeopteraLamellicornia).Acta Zoológica Mexicana81: 63–76. https://doi.org/10.21829/azm.2000.81811871 DeloyaC (2003) Familia Trogidae. In: MorónMA (Ed.) Atlas de los escarabajos de México.Vol II. Barcelona, 125–133. DeloyaC (2005) Omorgus rodriguezae a new species from Mexico, and a key for the separation of the Central and North America species of the genus (Coleoptera: Trogidae). Folia Entomologica Mexicana 44(Supl. 1): 121–129. DenierP (1936) Estado natural de mis conocimientos acerca del “champi” (Trox suberosus F.). In: Ministerio de Agricultura de la Nación, (Ed.) Memoria de la Comisión Central de Investigaciones Sobre La Langosta (Argentina).República Argentina, Buenos Aires, 205–216. DiéguezVM (2008) Conocimiento actual de los Trogidae de Chile (Coleoptera: Scarabaeoidea).Revista Chilena de Entomología34: 11–28. DiéguezVMGómezRS (2004) Aporte al conocimiento de las Trogidae de la Argentina.Revista de la Sociedad Entomológica de Argentina63(1–2): 92–95. ErichsonWF (1847) Conspectus coleopterorum quae in Republica Peruana observata sunt.Archiv für Naturgeschichte13(1): 67–185. FabriciusJC (1775) Systema Entomologiae, sistens insectorvm Classes, Ordines, Genera, Species, adiectis Synonymis, Locis, Descriptionibvs, Observationibus. Kortii, Flensburg and Leipzig [“Flensburgi et Lipsiae”]; Havniae [=Copenhagen], p. [32] + 832 pp. https://doi.org/10.5962/bhl.title.36510 FabriciusJC (1781) Species insectorvm exhibentes eorvm differentias specificas, synonyma avctorvm, loca natalia, metamorphosin adiectis observationibvs, descriptionibvs. Carol. Ernest. Bohn, Hambvrgi et Kilonii [= Hamburg & Kiel], Tom. I, [viii +] 552 pp. https://doi.org/10.5962/bhl.title.36509 FabriciusJC (1787) Mantissa Insectorum Sistens Eorum Species Nuper Detectas Adiectis Characteribus, Genericis, Differentiis, Specificis, Emendationibus, Observationibus. 1.Proft, Hafniae, 382 pp. https://doi.org/10.5962/bhl.title.36471 FabriciusJC (1792) Entomologia Systematica Emendata et Aucta. Secundum Classes, Ordines, Genera, Species Adjectis Synonymis, Locis, Observationibus, Descriptionibus. 1.Proft, Hafniae, 538 pp. https://doi.org/10.5962/bhl.title.125869 FabriciusJC (1801) Systema Eleutheratorum Secundum Ordines, Genera, Species Adiectis Synonymis, Locis, Observationibus, Descriptionibus. 1.Impensis Bibliopolii Adademici Novi, Kiliae [= Kiel], 506 pp. GianizellaSLPradoAP (1999) Ocorrência e sazonalidade de Omorgus (Omorgus) suberosus (Fabr.) (Trogidae: Coleoptera) em esterco de aves poedeiras, em Monte Mor, SP.Anais da Sociedade Entomológica do Brasil28(4): 749–751. https://doi.org/10.1590/S0301-80591999000400019 Giraldo-MendozaAE (2021) A preliminary list of beetles (Insecta: Coleoptera) of forensic importance from Peru.Dugesiana28(2): 61–74. https://doi.org/10.32870/dugesiana.v28i2.7148 GmelinJF (1790) Caroli a Linné Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima tertia, aucta, reformata. Tom. I. Pars IV. Georg Emanuel Beer, Lipsiae [= Leipzig], 1517–2224. GómezRS (2008) Trogidae. In: ClapsLEDebandiGRoig-JuñentS (Eds) Biodiversidad de Artrópodos Argentinos II.Sociedad Entomológica Argentina, 509–518. HaafE (1954) Die Australischen arten der gattung Trox (Col. Scarab.). Entomologische Arbeiten aus dem Museum G.Frey5(2): 691–740. HaroldE (1869) Tomo IV. Scarabaeidae. In: GemmingerMHaroldE (Eds) Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus.Monachii: Sumptu E.H. Gummi, 979–1346. https://doi.org/10.5962/bhl.title.9089 HaroldE (1872) Monographie der Gattung Trox.Coleopterologische Hefte9: 1–192. HatchMH (1971) The beetles of the Pacific northwest. Part 5. Rhipiceroidea, Sternoxi, Phytophaga, Rhynchophora and Lamellicornia.Publications in Biology of the University of Washington16: 433–485. HaywardKJ (1936) Contribucion al conocimiento de la langosta Schistocerca paranensis Burm. y sus enemigas naturales.Memoria de la Comisión Central de Investigaciones Sobre La Langosta (Argentina)1934: 199–229. HerbstJFW (1790) [Hefte 1: 1–151, 6 pls.]. In: Herbst JFW (Ed.) 1790–1791. Natursystem aller bekannten in- und ausländischen Insekten, als eine Forsetzung der von Büffonschen Naturgeschichte. Der Käfer Dritter Theil. Berlin: Joachim Pauli; p. xiv + 325 pp. + pls. 21–34. D–E [publication date following Bousquet 2016]. HielkemaAJHielkemaMA (2019) An annotated checklist of the Scarabaeoidea (Insecta: Coleoptera) of the Guianas.Insecta Mundi732: 1–306. HornGH (1874) Revision of the species of Trox of the United States.Transactions of the American Entomological Society5: 1–12. https://doi.org/10.2307/25076284 HowdenHFVaurieP (1957) Two new species of Trox from Florida (Coleoptera, Scarabaeidae).American Museum Novitates1818: 1–6. HuchetJBCosta-SilvaV (2018) A new species of Polynoncus Burmeister, 1876 from Brazil (Coleoptera: Trogidae).Zootaxa4524(5): 553–566. https://doi.org/10.11646/zootaxa.4524.5.3 ICZN [International Commission on Zoological Nomenclature] (1999) International Code of Zoological Nomenclature. 4th Edition. The International Trust for Zoological Nomenclature, London, [xxix +] 306 pp. IlligerK (1802) Olivier’s Entomologie oder Naturgeschichte der Insekten mit ihren Gattungsund ArtMerkmalen, ihrer Beschreibung und Synonymie. Käfer. Uebersetzt und mit Zusätzen und Anmerkungen durchgängig begleitet von Karl Illiger. Karl Reichard, Braunschweig, Zweiter Theil. Mit Kupfern. [iv +] 266 pp. KalawateASStrümpherWP (2024) Catalogue of species of the subgenus Afromorgus Scholtz, 1986 (Coleoptera: Trogidae: Omorginae) known from the Oriental and Palaearctic Regions, including a description of one new species from the Deccan Plateau in India.Zootaxa5437(1): 021–063. https://doi.org/10.11646/zootaxa.5437.1.3 KrellF-T (2010) Catalogue of Colorado scarab and stag beetles (Coleoptera: Scarabaeoidea), based on literature records. Denver Museum of Nature & Science (Technical Report), 87 pp. LacordaireJT (1856) Histoire naturelle des insectes. Genera des Coléoptères ou exposé méthodique et critique de tous les genres proposés jusqu’ici dans cet ordre d’insectes.Tome troisième contenant les familles des pectinicornes et lamellicornes, Roret, Paris, 594 pp. LandinB-O (1963) The Lamellicorn beetles of the Cape Verde Islands, with some biogeographical aspects.Societas Scientiarum Fennica (commentationes biologicae)26(2): 1–27. LengCW (1920) Catalogue of the Coleoptera of America, north of Mexico. John D. Sherman, Jr., Mount Vernon, New York, 470 pp. https://doi.org/10.5962/bhl.title.8777 LengCW (1928) Order Coleoptera. In: LeonardMD (Ed.) A list of the insects of New York.With a list of the spiders and certain other allied groups, Vol. 101. Cornell University Agricultural Experiment Station Memoir, New York, 1–1121. LöblISmetanaA (2006) Catalogue of Palaearctic Coleoptera (Vol.3) Apollo Books, Stenstrup, 690 pp. LopesWDZda CostaFHÇopesWCZBalieiroJCSoaresVEPradoAP (2007) Omorgus (Omorgus) suberosus (Fabricius) (Coleoptera: Trogidae) em esterco de galinhas poedeiras de São João da Boa Vista, SP, Brasil.Arquivos do Instituto Biológico74(3): 227–232. https://doi.org/10.1590/1808-1657v74p2272007 MacLeayWS (1819) Horae entomologicae: or essays on the annulose animals. Containing general observations on the geography, manners, and natural affinities of the Insects which compose the genus Scarabaeus of Linnaeus; to which are added a few incidental remarks on the genera Lucanus and Hister of the same author. With an appendix and plates. Vol.1, Pt. 1. London: S. Bagster, 160 pp. https://doi.org/10.5962/bhl.title.65809 MiquelME (2019) On the distribution of Omorgus suberosus (Coleoptera: Trogidae) and its presence in the Philippines.Entomofauna8: 183–191. Mora-AguilarEFMontes de OcaE (2009) Escarabajos necrófagos (Coleoptera: Scarabaeidae y Trogidae) de la región central baja de Veracruz, México.Acta Zoológica Mexicana (nueva serie)25(3): 569–588. https://doi.org/10.21829/azm.2009.253662 MoraguesG. (2010) Aperçu des Trox de Guyane (Coleoptera: Trogidae). Contribution à l’étude des Coléoptères de Guyane, 1: 76–78. [Supplément au Bulletin de liaison de l’ACOREP France “Le Coléoptériste”]. MorroneJJ (2001) Toward a cladistic model for the Caribbean subregion: delimitation of areas of endemism.Caldasia23(1): 43–76. MorroneJJ (2014) Biogeographical regionalisation of the Netropical region.Zootaxa3782(1): 1–110. https://doi.org/10.11646/zootaxa.3782.1.1 MorónMA (1998) Fauna de ColeopteraLamellicornia de la región de Tepic, Nayarit, Mexico.Acta Zoologica Mexicana75: 73–116https://doi.org/10.21829/azm.1998.75751708. MorónMA (2003) Los coleópteros Scarabaeoidea que habitan en los cafetales bajo sombra de México.Folia Entomológica Mexicana42(2): 397–414. Muñoz-BatetJLopez-ColonJI (1995) Primer registro centroeuropeo de Omorgus suberosus (Fabricius, 17759) (Coleoptera, Trogidae).Nouvelle Revue d’Entomologie12(4): 279. MutchlerAJ (1925) Coleoptera from the Williams Galapagos expeditions.Zoologica5(20): 219–240. https://doi.org/10.5962/p.190343 NikolajevGV (2005) Omorgini (Coleoptera, Scarabaeidae, Troginae) – a new tribe of scarab beetles.Euroasian Entomological Journal4: 321–322. [In Russian] OlivierGA (1789) Entomologie, ou histoire naturelle des insectes, avec leurs caractères génériques et spécifiques, leur description, leur synonymie, et leur figure enluminée. Coléoptères. Baudouin. Tome premier, [xx +] 433 pp. https://doi.org/10.5962/bhl.title.61905 Pablo-CeaJDCaveRDSerrano-PerazaFAAlvarado-LariosRDeloyaCSerrano-ChicasKAAlfaroEChinchilla-RodríguezACGirón-SegoviaDNoriegaJA (2023) Catálogo y atlas de distribución de los Scarabaeoidea (Insecta: Coleoptera) de El Salvador. Revista Mexicana de Biodiversidad 94: e945117. https://doi.org/10.22201/ib.20078706e.2023.94.5117 PapaveroN (1971) Essays on the history of Neotropical dipterology, with special reference to collectors (1750–1905). Vol. I + II. Museu de Zoologia, Universidade de São Paulo, São Paulo, [viii +] 446 pp. https://doi.org/10.5962/bhl.title.101715 PáramoPB (1997) El género Omorgus Erichson, 1847 en la Península Ibérica (Coleoptera, Scarabaeoidea: Trogidae).Boletín de la Sociedad Entomológica Aragonesa17: 29–31. PaulianR (1981) Insectes Coléoptères Trogidae et Hybosoridae.Faune de Madagascar56: 1–29. PhilipsJR (2009) The mite (Acarina) fauna of trogid beetles (Coleoptera: Trogidae).International Journal of Acarology35(1): 1–17. https://doi.org/10.1080/01647950802709843 PittinoR (1987) New ColeopteraTrogidae from South America (XXXII contribution to the knowledge of ColeopteraScarabaeoidea).Giornale Italiano di Entomologia3(17): 377–397. PittinoR (2010) A new genus and species of Trogidae MacLeay, 1819 from Madagascar, and a new species of Omorgus Erichson, 1847 from Argentina (Coleoptera, Trogidae).Kogane11: 75–83. PittinoRBezdĕkA (2016) Family Trogidae W.S. Macleay, 1819. In: LöblILöblD (Eds) Catalogue of Palaearctic Coleoptera.Volume 3. Scarabaeoidea—Scirtoidea—Dascilloidea—Buprestoidea—Byrrhoidea. Revised and updated edition. Brill, Leiden, The Netherlands, 53–58. Preudhomme de BorreFPCA (1886) Catalogue des Trogides décrits jusque’ à ce jour précédé d’ un synopsis de leurs genres et d’une esquisse de leur distribution géographique.Annales de la Société entomologique de Belgique30: 56–82. RatcliffeBC (1978) A new species of Trox from the Amazon Basin with new distributional records for Central American Trox (Coleoptera: Scarabaeidae).Acta Amazonica8(2): 299–302. https://doi.org/10.1590/1809-43921978082299 RatcliffeBC (1991) The scarab beetles of Nebraska.Bulletin of the University of Nebraska State Museum12: 1–333. RatcliffeBC (2002) A checklist of the Scarabaeoidea (Coleoptera) of Panama.Zootaxa32(1): 1–48. https://doi.org/10.11646/zootaxa.32.1.1 RatcliffeBCJamesonMLFigueroaLCaveRDPaulsenMJCanoEBBeza-BezaCJimenez-FerbansLReyes-CastilloP (2015) Beetles (Coleoptera) of Peru: a survey of the families. Scarabaeoidea.Journal of the Kansas Entomological Society88(2): 186–207. https://doi.org/10.2317/kent-88-02-186-207.1 RitcherOP (1958) Biology of Scarabaeidae.Annual Review of Entomology3: 311–334. https://doi.org/10.1146/annurev.en.03.010158.001523 RitcherOP (1966) White grubs and their allies: a study of North American Scarabaeoid larvae.Oregon State University Press, Corvallis, Oregon, 220 pp. RobinsonM (1941) A new species of Trox from Texas (Coleoptera: Scarabaeidae).Entomological News52: 134–135. Rosano-HernándezMCDeloyaC (2002) Interacción entre trogidos (Coleoptera: Trogidae) y tortugas marinas (Reptilia: Cheloniidae) en el Pacifico Mexicano.Acta zoológica Mexicana87: 29–46. https://doi.org/10.21829/azm.2002.87871799 ScholtzCH (1982) Catalogue of world Trogidae (Coleoptera: Scarabaeoidea).Entomology Memoir, Department of Agriculture and Fisheries Republic of South Africa54: 1–27. ScholtzCH (1986a) Phylogeny and systematics of the Trogidae (Coleoptera: Scarabaeoidea).Systematic Entomology11: 355–363. https://doi.org/10.1111/j.1365-3113.1986.tb00186.x ScholtzCH (1986b) Revision of the genus Trox Fabricius (Coleoptera: Trogidae) of the Australian region.Australian Journal of Zoology125: 1–99. https://doi.org/10.1071/AJZS125 ScholtzCH (1990) Revision of the Trogidae of South America (Coleoptera: Scarabaeoidea).Journal of Natural History24: 1391–1456. https://doi.org/10.1080/00222939000770841 SchönherrCJ (1806) Synonymia insectorum, oder: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum geordnet. Mit Berichtigungen und Anmerkungen, wie auch Beschreibungen neuer Arten und illuminirten Kupfern. Erster Band. Eleutherata oder Käfer. Heinr. A. Nordström, Stockholm. Erster Theil. Lethrus – Scolytes, [xxii +] 293 pp. https://doi.org/10.5962/bhl.title.66107 SmithABT (2003) Checklist of the Scarabaeoidea of the Nearctic Realm. Version 3.Electronically published, Lincoln, Nebraska, 74 pp. SmithABT (2017) Southern South American Trogidae (Coleoptera): verification of specimens and notes on geographic distribution.Revista Chilena de Entomología43: 81–103. StrümpherWPKalawateAS (2023) Omorgus (Omorgus) khandesh (Coleoptera: Scarabaeoidea: Trogidae), a new species from India, along with an annotated catalogue of the Oriental and Palearctic species of the subgenus Omorgus Erichson, 1847.Zootaxa5231(5): 501–522. https://doi.org/10.11646/zootaxa.5231.5.1 StrümpherWPSoleCLVilletMHScholtzCH (2014) Phylogeny of the family Trogidae (Coleoptera: Scarabaeoidea) inferred from mitochondrial and nuclear ribosomal DNA sequence data.Systematic Entomology39(3): 548–562. https://doi.org/10.1111/syen.12074 StrümpherWPVilletMHSoleCLScholtzCH (2016) Overview and revision of the extant genera and subgenera of Trogidae (Coleoptera: Scarabaeoidea).Insect Systematics & Evolution47(1): 53–82. https://doi.org/10.1163/1876312X-46052133 SturmJ (1843) Catalog der Kaefer-Sammlung von Jacob Sturm. Mit 6 ausgemalten Kupfertafeln. Nürnberg, [xii +] 386 pp. https://doi.org/10.5962/bhl.title.37837 Van DykeEC (1953) The Coleoptera of the Galapagos Islands.Occasional papers of the California Academy of Sciences22: 1–181. VaurieP (1955) A revision of the genus Trox in North America (Coleoptera: Scarabaeidae).The Bulletin of the American Museum of Natural History106: 1–89. VaurieP (1958) New distribution records of North American Trox (Coleoptera: Scarabaeidae).The Coleopterists Bulletin12: 43–46. https://doi.org/10.5962/p.372226 VaurieP (1962) A revision of the genus Trox in South America (Coleoptera: Scarabaeoidea).The Bulletin of the American Museum of Natural History124: 105–167. VerdugoA (2014) Morfología de los estadios inmaduros, biología y comportamiento de Trox cotodognanensis Compte, 1986 en Cádiz (Coleoptera: Scarabaeoidea: Trogidae).Revista gaditana de Entomología5(1): 211–224. YoungOP (2006) Survival and reproduction of Trox suberosus F. (Coleoptera: Trogidae) on insect cadavers, cow dung, and mushroom.Journal of Entomological Science41(3): 271–276. https://doi.org/10.18474/0749-8004-41.3.271 YoungOPHammJJ (1985) The effect of the consumption of NPV-infected dead fall armyworm larvae on the longevity of two species of scavenger beetles.Journal of Entomological Science20(1): 90–94. https://doi.org/10.18474/0749-8004-20.1.90 ZianiSBezděkABrancoTHillertOJáklSKrálDMantičMRößnerESehnalR (2015) New country records of Scarabaeoidea from the Palaearctic region (Coleoptera). Insecta Mundi.A Journal of World Insect Systematics409: 1–36. ZidekJ (2013) Checklist and bibliography of the Trogidae (Coleoptera: Scarabaeoidea).Insecta Mundi314: 1–38. ZidekJ (2017) Updated checklist and bibliography of family Trogidae (Coleoptera: Scarabaeoidea).Folia Heyrovskyana25(1): 93–127. ZimsenE (1964) The type material of I.C. Fabricius. Copenhagen: Munksgaard, 656 pp.