Annotated catalog and bibliography of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae, Cyclocephalini)

Matthew R. Moore; Ronald D. Cave; Marc A. Branham

doi:10.3897/zookeys.745.23685

Catalogue

Annotated catalog and bibliography of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae, Cyclocephalini)

Matthew R. Moore^‡, Ronald D. Cave^§, Marc A. Branham^‡

‡ University of Florida, Gainesville, United States of America

§ University of Florida, Fort Pierce, United States of America

Corresponding author: Matthew R. Moore ( cyclocephala@gmail.com )

Academic editor: Andrey Frolov

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.

Citation: Moore MR, Cave RD, Branham MA (2018) Annotated catalog and bibliography of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae, Cyclocephalini). ZooKeys 745: 101-378. https://doi.org/10.3897/zookeys.745.23685

ZooBank: urn:lsid:zoobank.org:pub:8785DC6B-C2A2-44FD-94B6-243EB07C717F

Abstract

Cyclocephaline scarab beetles represent the second largest tribe of the subfamily Dynastinae, and the group includes the most speciose genus of dynastines, Cyclocephala. The period following publication of Sebő Endrődi’s The Dynastinae of the World has seen a huge increase in research interest on cyclocephalines, and much of this research has not been synthesized. The objective of this catalog and bibliography is to compile an exhaustive list of taxa in Cyclocephalini. This paper provides an updated foundation for understanding the taxonomy and classification of 14 genera and over 500 species in the tribe. It discusses the history of cataloging dynastine species, clarifies issues surrounding the neotype designations in Endrődi’s revision of Cyclocephalini, synthesizes all published distribution data for cyclocephaline species, and increases accessibility to the voluminous literature on the group by providing an easily searchable bibliography for each species. We propose the nomen novum Cyclocephala rogerpauli, new replacement name, for C. nigra Dechambre.

Keywords

masked chafers, rhinoceros beetles, catalog, bibliography

Introduction

The Cyclocephalini, a group first defined by French naturalist Francis de Laporte de Castelnau in 1840, represents the second largest tribe of the subfamily Dynastinae. Cyclocephala Dejean, the type genus of Cyclocephalini, is the most speciose dynastine genus and comprises over 350 species-group taxa as of 2017. The last comprehensive, synoptic treatment of the tribe was The Dynastinae of the World (Endrődi 1985a). Endrődi’s foundational book revolutionized the study of the subfamily and paved the way for a veritable explosion of new research into dynastines. This influence is most apparent in the scientific literature covering cyclocephaline scarab beetles. The post-Endrődi era of cyclocephaline research has been marked by ever diversifying interests and approaches to the group. Papers on cyclocephalines now span all modern entomological disciplines from taxonomy, evolutionary biology, ecology, ethology, agronomics, and physiology.

Figure 1.

Cumulative number of described cyclocephaline species-group taxa by decade. Species description accumulation was based on the compiled catalog. The synonymy curve also includes names that were homonyms and later replaced.

Ever-growing numbers of cyclocephaline species have, at times, engendered lighthearted dismay among researchers of the group. For example, the Costa Rican C. unamas Ratcliffe (Spanish “una mas”) was named after the overwhelming feeling one gets after the discovery of yet another new Cyclocephala species, epitomized by the species name C. nodanotherwon Ratcliffe. Over 170 new cyclocephaline species-group taxa have been described since 1985, and this has created challenges for species identification in several genera. While the most intense period of new species descriptions has probably passed (Fig. 1), many new South American taxa are likely to be discovered, especially in the genera Cyclocephala and Stenocrates Burmeister (Ratcliffe 2015).

Starting in the mid-1970s, a growing body of research covering the floral ecology of cyclocephalines began to develop. Many faunistic studies in Mesoamerica (especially Mexico) and South America have reported a great deal of cyclocephaline locality data that has yet to be synthesized. Additionally, researchers in the United States and South America have greatly expanded the agronomic literature on the tribe since publication of The Dynastinae of the World. This hugely expanded literature for the tribe has not been adequately synthesized and is unwieldy and inaccessible as a result. The objective of this catalog is to: 1) provide an updated foundation for understanding the taxonomic history of 14 genera and over 500 species of cyclocephaline scarab beetles; 2) identify destabilizing issues in the classification and nomenclature of the genera and species; 3) create an easily searchable bibliography to further promote research on these beetles; and 4) synthesize known distribution data for all species in the tribe.

Brief history of cyclocephalines in catalogs, checklists, and bibliographies

Coleopterists have a long history of compiling species catalogs at a global or regional scale. Prior to the Information Age, these catalogs were invaluable resources for the entomological community because they served to organize biodiversity research. Cataloging the diversity of cyclocephaline species began in Germany with the fourth volume of Catalogus Coleopterorum Hucusque Descriptorum Synonymicus et Systematicus (Harold 1869b). This catalog included information on over 120 valid species in the group and provided a brief list of citations for each taxon along with locality information. French entomologists Louis Chevrolat, Albert Fauvel, Auguste Sallé, and Edmond Fleutriaux provided early lists of cyclocephaline diversity in French Guiana (Fauvel 1861), Cuba (Chevrolat 1865), and Guadeloupe (Fleutiaux and Sallé 1889). Gilbert Arrow (1937b) published a comprehensive catalog of Dynastinae in the Coleoptorum Catalogus series. Arrow’s catalog featured an updated classification of the subfamily and the tribe Cyclocephalini. His concept of Cyclocephalini was broader than that of later workers, and he included several genera in the tribe that would be subsequently included in Oryctoderini. This new catalog also updated the bibliographic information for each species (Arrow 1937b). Burgeon (1947) followed up Arrow’s work and cataloged dynastine species of the Democratic Republic of Congo and provided an image and redescription of the African cyclocephaline Ruteloryctes morio (Fabricius). Friedrich Ohaus and Johann Machatschke published several catalogs of Rutelinae in the mid-20^th century in the Coleopterorum Catalogus and Genera Insectorum series, which at the time included information about the cyclocephaline genera Peltonotus Burmeister and Acrobolbia Ohaus (Ohaus 1918, 1934b, Machatschke 1972, 1974). Milan Krajcik (2005, 2012) published exhaustive checklists of world Dynastinae and Scarabaeoidea in his Annima.X series.

Many North American Coleopterists created catalogs and checklists that included cyclocephalines. Frederick Melsheimer, Samuel Haldeman, and John LeConte compiled the first catalog of Coleoptera of the United States (Melsheimer et al. 1853). George Horn’s (1894) list of Coleoptera of Baja California included three species of Cyclocephala. W. S. Blatchley (1910, 1930) cataloged the Coleoptera of Indiana and created a checklist of scarabs of Florida. Charles Leng, Andrew Mutchler, and Richard Blackwelder compiled enormous catalogs and checklists of beetles, which included cyclocephalines, throughout the New World, including the West Indies (Leng and Mutchler 1914, 1917, Leng 1920, Blackwelder 1939, 1944, 1948). George Wolcott (1923, 1936) assembled annotated checklists of the insects of Puerto Rico and later provided interesting observations of the natural history of Puerto Rican scarabs in other works. Alan Hardy (1991) created a very detailed annotated catalog of Rutelinaeand Dynastinae of North America. Andrew Smith created a checklist of all scarabaeoid beetles of the Nearctic Realm (Smith 2009). Stewart Peck, along with collaborators, has cataloged Coleoptera from parts of the West Indies (Peck 2009, 2010, 2016, Peck et al. 2002). Relatively modern checklists or reviews of scarabaeoids have been produced for Canada (McNamara 1991, Bousquet et al. 2013, Ratcliffe and Cave 2017) and portions of the United States including Florida (Woodruff 1973, Peck and Thomas 1998), Maryland (Staines 1984), Nebraska (Ratcliffe 1991, Ratcliffe and Paulsen 2008), South Carolina (Harpootlian 2001), and Texas (Riley and Wolfe 2003).

Checklists of the economically injurious insects of Honduras (Passoa 1983), Nicaragua (Maes and Robleto 1988), Colombia (Posada Ochoa), French Guiana (Remillet 1988), and Suriname (Van Dinther 1960) report unique and fascinating records of cyclocephalines causing damage in agroecosystems. Cyclocephaline floral association data were compiled for the family Araceae (Gibernau 2003, 2011) and for the beetle tribe (Moore and Jameson 2013). Relatively recent checklists focusing on mainland Neotropical scarabaeoid or dynastine taxa have been produced for parts of Mexico (Deloya et al. 2014a, 2016), Nicaragua (Maes 1987, Maes et al. 1997), Panama (Ratcliffe 2002a), Colombia (Restrepo et al. 2003, Gasca-Álvarez and Amat-García 2010), Peru (Ratcliffe et al. 2015), and French Guiana (Ponchel 2011). Regional checklists of Dynastinae were published for Costa Rica and Panama (Ratcliffe 2003), Honduras, Nicaragua, and El Salvador (Ratcliffe and Cave 2006), Mexico, Guatemala, and Belize (Ratcliffe et al. 2013), the West Indies (Ratcliffe and Cave 2015), and the United States (Ratcliffe and Cave 2017). An updated catalog of Stenocrates was provided by Ratcliffe (2015), and Dupuis (2017) added details about Stenocrates in French Guiana. An updated checklist of the Cyclocephala of Colombia was provided by Gasca-Álvarez and Deloya (2016). Cyclocephala is the only genus in the tribe for which a tailored bibliography has been produced (Pike et al. 1976).

Materials and methods

All available literature was reviewed for compiling this catalog and bibliography. References to a cyclocephaline genus only were not included in the list of references. Cited references must have used a trackable specific epithet to have been included. The institutional and collection acronyms used throughout the catalog follow Evenhuis (2016) when possible.

CAS California Academy of Sciences, San Francisco, California, USA

CERPE Coleção Entomológica da Universidade Federal Rural de Pernambuco, Recife, Brazil

CMNC Canadian Museum of Nature, Ottawa, Ontario, Canada

CNC Canadian National Collection of Insects, Ontario, Ottawa, Ontario, Canada

BMNH The Natural History Museum, London, United Kingdom

FSCA Division of Plant Industry, Florida State Collection of Arthropods, Gainesville, Florida, USA

FDPC Fabien Dupuis Collection, Saint-Chamond, France

FUJI Masayuki Fujioka Collection, Tokyo, Japan

HNHM Hungarian Natural History Museum, Budapest, Hungary

ICN Universidad Nacional de Colombia, Insituto de Ciencias Naturales de la Universidad Nacional, Bogotá, Colombia

IEE Institute of Ecology and Evolution, Russian Academy of Sciences, Moscow, Russia

IEXA Colección Entomológica, Instituto de Ecología, A.C., Xalapa, México

IMQC Insectarium de Montreal, Montreal, Québec, Canada

INPA Instituto Nacional de Pesquisas da Amazonia, Colecão Sistemática da Entomologia, Manaus, Amazonas, Brazil

IREC Institut de Recherches Entomologique de la Caribe, Pointe-a-Pitre, Guadeloupe (also known as Centre de Recherches Agronomiques Antilles Guyana, Duclos, Petit-Bourg [CRAAG])

JPVC J. Pierre Voirin Collection, Le Luc, France

LEMQ Ste. Anne de Bellevue, McGill University, Lyman Entomological Museum, Québec, Canada

MACN Museo Argentina de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Argentina

MCMC Museo de Historia Natural de la Ciudad de Mexico, Distrito Federal, Mexico

MCZ Harvard University, Museum of Comparative Zoology, Cambridge, Massachusetts, USA

MIZA Museo del Instituto de Zoología Agrícola, Maracay, Venezuela

MLUH Zentralmagazin Naturwissenschaftlicher Sammlungen, Martin-Luther Universität Halle-Wittenberg, Halle, Germany

MNCR Museo Nacional de Costa Rica, San José, Costa Rica

MNHN Muséum National d’Histoire Naturelle, Paris, France

MNNC Coleccion Nacional de Insectos, Museo Nacional de Historia Natural, Santiago, Chile

MUSENUV Universidad de Valle, Museo de Entomología, Cali, Colombia

MUSM Museo de Historia Natural de la Universidad Nacional Mayor de San Marco, Lima, Peru

MTD Museum für Tierkunde, Dresden, Germany

MXAL Miguel Ángel Morón Collection, Xalapa, Mexico

MZSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil.

NHMB Naturhistorisches Museum, Basel, Switzerland

NHRS Naturhistoriska riksmuseet, Stockholm, Sweden

NMPC National Museum (Natural History), Prague, Czech Republic

NSMT National Science Museum (Natural History), Tokyo, Japan

QSBG Queen Sirikit Botanic Garden, Chiang Mai, Thailand

RIEB Research Institute of Evolutionary Biology, Tokyo, Japan

RPDC Roger-Paul Dechambre Collection, Paris, France

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany

UCDC UCDC

UNSM University of Nebraska State Museum, Lincoln, Nebraska, USA

USNM National Museum of Natural History, Washington, District of Columbia, USA

UUZM Uppsala University, Uppsala, Sweden

UVGC Universidad del Valle de Guatemala, Colección de Artrópodos, Guatemala City, Guatemala

WADA Kaoru Wada Collection, Tokyo, Japan

ZMH Zoologiska Museum, University of Helsinki, Helsinki, Finland

ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany

ZMUC University of Copenhagen, Zoological Museum, Copenhagen, Denmark

ZMUH Universität von Hamburg, Zoologisches Institut und Zoologisches Museum, Hamburg, Germany

ZMUK Universität Kiel, Zoologisches Museum, Kiel, Germany

ZSMC Zoologische Staatssammlung des Bayerischen Staates, Munich, Germany

A special note must be made about the type specimen housing institution reporting herein. For older literature, the remarks on type depositories relied on Endrődi’s (1966) explanations. However, many private collections (or portions of collection holdings) have changed hands in the intervening period. This is most relevant for the Antonio Martínez Collection, Sebő Endrődi Collection, Henry and Anne Howden Collection, Frey Collection, and for some Fabrician types. These collections contain a significant number of primary type material for cyclocephaline species. Holotypes deposited in the Martínez Collection should be at MACN. Holotypes and invalid neotypes deposited in the Endrődi Collection should be at HNHM. Holotypes deposited in the Howden collection should be at CMNC. Holotypes deposited in the Frey Collection should be at NHMB. Fabrician lectotypes designated by Endrődi (1966) were originally at ZMUK, but they should now be at ZMUC.

Most of Endrődi’s (1966) lectotypes were clearly designated. However, the types of many species (especially Arrow and Bates types at BMNH and Casey types at USNM) were not clearly discussed. The original descriptions of these species were not always explicit about the number of specimens in a type series. Endrődi (1966) does not clarify these cases and simply listed that a “Type” was at an institution. Herein, these “Type” specimens were not speculated to be holotypes by monotypy or as parts of a syntype series. Future workers who further examine the original descriptions and the type material will have to make those judgements.

As noted by a few authors, Endrődi’s cyclocephaline neotype designations are invalid on several grounds (Dechambre 1991b, Ratcliffe and Hoffman 2011). Endrődi’s (1966) revision of Cyclocephalini was published after the recently adopted 1964 version of the International Code of Zoological Nomenclature (ICZN 1964). ICZN (1964) Article 75c had six conditions that must have been met by Endrődi for his neotype designations to be valid: 1) a statement of characters for differentiating the taxon for which the neotype was designated (or a reference to such a statement); 2) data and description sufficient so that the neotype can be recognized; 3) explanation for believing that all of the original type material is lost or destroyed and the steps that were taken to determine this was the case; 4) explanation of why the neotype specimen is considered consistent with the original-type material; 5) explanation that the neotype came for as near as possible to the original type-locality; and 6) a statement that the neotype is immediately, or upon publication, the property of a recognized scientific or educational institution that maintains a research collection.

Every one of Endrődi’s (1966) twenty neotype designations variably violates the conditions of ICZN (1964) Article 75c. Endrődi possibly satisfied condition (1) of Article 75c in some cases because his neotype designations were accompanied by detailed descriptions (but not necessarily explicitly stated to be descriptions of the neotype). Condition (2) was violated in every case. For example, he did not describe the labels of any of his neotype specimens, hampering recognition of the neotypes. He sometimes also omitted an explicit statement about the sex of the neotype (though they are presumably male). Usually, Endrődi only made vague statements about his search for type materials, amounting to the fact that he did not find a type. This violated condition (3). He typically did not report where exactly he had searched for the material that he did not find. For example, after his description of Cyclocephala villosa Blanchard, Endrődi only stated, “Die Type war trotz sorgfältiger Nachforschung nicht aufzufinden, darum designierte Ich mein einziges Exemplar als Neotype ♂” [“The type was not found despite a careful search, so I designated my only specimen as a neotype ♂”]. In contrast, he went into detail about his search (with the help of Bengt-Olof Landin) for Linnean type material of C. amazona amazona.

Endrődi never described why he thought his neotypes were consistent with the original type material, nor did he explicitly mention type locality (violating conditions 4 and 5). He was obviously aware of the concept of type locality as evidenced by his decisions when designating neotypes. For example, he designated a neotype of C. castanea (Olivier) from “Surinam”, which is the type locality of this species based on the original description (Olivier 1789). However, he offered no explanation of these concepts. Lastly, all of Endrődi’s neotypes were deposited in his personal collection (now at HNHM). Dechambre (1991b) recognized that deposition in Endrődi’s collection did not satisfy condition (6). Endrődi’s invalid neotypes are listed below, in their original name combinations, and are noted in the catalog. In some cases, Dechambre (1991b) discovered syntypes of these species at MNHN and designated lectotypes. Those lectotypes are noted here where applicable.

Chalepus luridus Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966). Joly and Escalona (2002a) listed the housing institution as MLUH.

Cyclocephala concolor Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype ♀ at MNHN (Dechambre 1991b).

Cyclocephala fulvipennis Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype ♀ at MNHN (Dechambre 1991b).

Cyclocephala gregaria Heyne & Taschenberg: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966).

Cyclocephala nigricollis Burmeister: invalid ♂ neotype at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype ♂ at MNHN (Dechambre 1991b).

Cyclocephala occipitalis Fairmaire: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966).

Cyclocephala octopunctata Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype ♀ at MNHN (Dechambre 1991b).

Cyclocephala putrida Burmeister: invalid neotype at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype ♀ of C. putrida at MNHN (Dechambre 1991b).

Cyclocephala rubescens Bates: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966).

Cyclocephala signaticollis Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype ♀ at MNHN (Dechambre 1991b).

Cyclocephala subsignata Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype ♀ at MNHN (Dechambre 1991b).

Cyclocephala tetrica Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966).

Cyclocephala villosa Blanchard: invalid ♂ neotype at MNHN (Endrődi Collection) (Endrődi 1966).

Cyclocephala villosa Burmeister: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966).

Melolontha castanea Olivier: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966).

Melolontha immaculata Olivier: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966). Chalumeau and Gruner (1977) stated this neotype was at MNHN.

Melolontha picipes Olivier: invalid neotype ♂ at HNHM (Endrődi Collection) (Endrődi 1966).

Melolontha rustica Olivier: invalid neotype at HNHM (Endrődi Collection) (Endrődi 1966).

Melolontha undata Olivier: invalid ♂ neotype at HNHM (Endrődi Collection) (Endrődi 1966).

Scarabaeus amazonus Linnaeus: invalid neotype at HNHM (Endrődi Collection) (Endrődi 1966).

How to use this catalog

Entries for genera follow a format that tracks the history of genera as either valid or invalid through time. The generic-level entries also include the type species of the genus, a list of references that contain identification keys for the genus and/or its species, and the number of valid species and subspecies in the genus.

The very first line, in bold, is the current valid name of the taxon. Underneath that, the taxonomic history of the name is presented in chronological order. These histories can be extremely complicated and difficult to track. The types of changes included here are: 1) changes in generic classification; 2) changes in subgeneric classification; 3) names that the current valid name may have been synonymized under for a period; 4) revalidation of names as either species or subspecies; 5) changes in status that include movement between specific, subspecific, and infrasubspecific categories. These changes, outlined above, are cited using an abbreviated reference, with pagination, and what actions that author took.

Entries for synonyms are indented and labeled “syn.” in bold. The taxonomic histories of synonyms can be just as complicated as valid names and those details are provided here in the same format as the taxonomic history of the valid name. Information about the primary types, as far as could be ascertained, is provided beneath the taxonomic histories. These include citations for that information and the primary type repository. A generalized distribution from the literature is provided below the type information. Country records are in all capital letters and state/department/commune-level records are given after the country. Lastly, a bibliography of each species is provided in rough chronological order and sorted by author.

Remarks are also given that clarify some data about a taxon where it is applicable. Remarks given here generally relate to conflicting distribution data in the literature or potential minor taxonomic and nomenclatural issues. Because so little is known of cyclocephaline relationships, the catalog is presented in alphabetic order by genus and then species-group taxa, instead of systematic order.

Annotated catalog and bibliography of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae, Cyclocephalini)

Tribe Cyclocephalini Laporte, 1840

Cyclocephalites Laporte, 1840: 124 [original usage].

Cyclocephalidae [Burmeister, 1847: 21].

Cyclocephalides [Lacordaire, 1856: 393].

Cyclocephalini [LeConte, 1862: 143].

Cyclocephalinae [Bates, 1888: 296].

Type genus

Cyclocephala Dejean, 1821.

Genus Acrobolbia Ohaus, 1912

Acrobolbia Ohaus, 1912: 316 [original usage].

Type species

Acrobolbia macrophylla Ohaus, 1912, by monotypy.

Keys

Jameson et al. 2002.

Valid taxa

1 species.

Acrobolbia macrophylla Ohaus, 1912

Acrobolbia macrophylla Ohaus, 1912: 317–318 [original combination].

syn.Acrobolbia triangularis Benderitter, 1922: 147 [original combination].

Acrobolbia macrophylla Ohaus [synonymy by Ohaus 1934a: 14].

Types

Holotype ♂ of A. macrophylla at ZMHB (Jameson et al. 2002). Neotype ♂ of A. triangularis at UNSM (Jameson et al. 2002).

Distribution

ECUADOR: Napo, Pastaza. PERU: Huanuco, Madre de Dios. VENEZUELA.

References

Lucas 1918a, Benderitter 1922, Ohaus 1912, 1918, 1934a, b, Anonymous 1940, Machatschke 1972, Jameson 1998, Dechambre and Ponchel 1999, Jameson et al. 2002, Breeschoten et al. 2013.

Genus Ancognatha Erichson, 1847

Ancognatha Erichson, 1847a: 97 [original usage].

Cyclocephala Dejean [synonymy by Lacordaire 1856: 398].

Ancognatha Erichson [revalidated genus status by Bates 1888: 297].

syn.Barotheus Bates, 1891: 30–31 [original usage]. Type species: Barotheus andinus Bates, 1891, by monotypy.

Ancognatha Erichson [synonymy by Endrődi 1966: 365].

syn.Lissodon Paulian, 1954: 1154 [original usage]. Type species: Lissodon argodi Paulian, 1954, by monotypy.

Ancognatha Erichson [synonymy by Endrődi 1969b: 38].

syn.Pseudoancognatha Otoya, 1945: 275 [original usage]. Proposed as a subgenus. Type species: Ancognatha nigriventris Otoya, 1945, by original designation. Ancognatha Erichson [synonymy by Martínez 1965a: 64].

Type species

Ancognatha scarabaeoides Erichson, subsequent designation by Casey 1915: 111.

Keys

Saylor 1945 (USA), Arnett 1968 (USA), Endrődi 1966, 1985a, Figueroa and Ratcliffe 2016 (Peru), Morón and Deloya 1991 (Mexico, Durango), Morón-Ríos and Morón 2001 (Mexico, Chiapas), Ratcliffe et al. 2013 (Mexico), Neita-Moreno and Morón 2008 (larvae), Vallejo and Morón 2008 (larvae), Pardo-Locarno et al. 2006, Gasca-Álvarez and Amat-García 2010 (Colombia), Mondaca 2011 (Chile), Villalobos-Moreno et al. 2017 (Colombia), Ratcliffe and Cave 2017 (USA and Canada).

Valid taxa

22 species.

Ancognatha atacazo (Kirsch, 1885)

Cyclocephala atacazo Kirsch, 1885: 223 [original combination].

Ancognatha atacazo (Kirsch) [new combination by Endrődi 1966: 370].

Types

Lectotype ♀ at MTD (Endrődi 1966).

Distribution

COSTA RICA: Cartago, San José. COLOMBIA: Quindío, Tolima, Valle del Cauca. ECUADOR: Pichincha.

References

Kirsch 1885, Bertkau 1886, Arrow 1937b, Blackwelder 1944, Pike et al. 1976, Endrődi 1966, 1967a, 1985a, Ratcliffe 2003, Pardo-Locarno et al. 2006, Krajcik 2005, 2012.

Ancognatha aymara Mondaca, 2016

Ancognatha aymara Mondaca, 2016: 60–63 [original combination].

Types

Holotype ♂ at MNNC (Mondaca 2016).

Distribution

CHILE: Arica y Parinacota.

References

Mondaca 2016.

Remarks

Specimens of A. lutea reported from Chile (e.g., see Gutiérrez 1950, Mondaca 2011, Ferrú and Elgueta 2011) were later determined to be A. aymara (Mondaca 2016).

Ancognatha castanea Erichson, 1847

Ancognatha castanea Erichson, 1847a: 98 [original combination].

Cyclocephala castanea (Erichson) [new combination by Lacordaire 1856: 398, 399].

Barotheus castaneus (Erichson) [new combination by Arrow 1911: 169].

Ancognatha castanea Erichson [revised combination by Endrődi 1966: 365].

syn.Barotheus andinus Bates, 1891: 31 [original combination].

Barotheus castaneus (Erichson) [synonymy by Arrow 1911: 169].

syn.Lissodon argodi Paulian, 1954: 1154–1155 [original combination].

Ancognatha castanea Erichson [synonymy by Endrődi 1969b: 38].

Types

Lectotype ♀ of A. castanea at ZMHB (Endrődi 1966). Type of B. andinus at BMNH (Endrődi 1966). Holotype ♂ at MNHN (Paulian 1954).

Distribution

COLOMBIA: Nariño. ECUADOR: Chimborazo, Napo, Pichincha. PERU: Ayacucho, Cuzco, Lima.

References

Erichson 1847a, Lacordaire 1856, Marschall 1857, Harold 1869b, Bates 1891, Bertkau 1892, Arrow 1911, 1937b, Blackwelder 1944, Paulian 1954, Endrődi 1966, 1969b, 1974, 1985a, Onore 1997, 2005, Pardo-Locarno et al. 2006, Krajcik 2005, 2012, Breeschoten et al. 2013, Ratcliffe et al. 2015, Dossey et al. 2016, Figueroa and Ratcliffe 2016, Mitasuhashi 2016.

Remarks

Lacordaire’s (1856) rejection of Ancognatha created a case of homonymy between the names C. castanea (Erichson) and C. castanea (Olivier). Cyclocephala peruana was proposed as a replacement for the junior homonym C. castanea (Erichson) (Harold 1869a). Subsequent authors did not use this replacement name (e.g., Arrow 1911, 1937b), with Endrődi (1966) stating that it was an “incorrect” new name.

Ancognatha corcuerai Figueroa & Ratcliffe, 2016

Ancognatha corcuerai Figueroa & Ratcliffe 2016: 65–67 [original combination].

Types

Holotype ♂ at MUSM (Figueroa and Ratcliffe 2016).

Distribution

PERU: Cajamarca.

References

Figueroa and Ratcliffe 2016.

Ancognatha erythrodera (Blanchard, 1846)

Cyclocephala erythrodera Blanchard, 1846: 191 [original combination].

Endrődi 1967a, 1985a, Dechambre 2000, Pardo-Locarno et al. 2006, Krajcik 2005, 2012.

Ancognatha humeralis (Burmeister, 1847)

Cyclocephala humeralis Burmeister, 1847: 40 [original combination].

Ancognatha humeralis (Burmeister) [new combination by Arrow 1914: 274].

syn.Cyclocephala longiceps Kirsch, 1870: 354–355 (paginated incorrectly as 370–371) [original combination].

syn.Ancognatha aequata Bates, 1888: 297 [original combination].

Ancognatha manca (LeConte) [synonymy by Arrow 1911: 169].

syn.Ancognatha durangoana Casey, 1915: 125 [original combination].

Ancognatha manca (LeConte) [synonymy by Saylor 1945: 125].

syn.Ancognatha laevigata Bates, 1888: 297–298 [original combination].

Ancognatha manca (LeConte) [synonymy by Saylor 1945: 379].

syn.Ancognatha perspicua Casey, 1915: 126 [original combination].

Ancognatha manca (LeConte) [synonymy by Arrow 1937b: 6].

syn.Ancognatha zuniella Casey, 1915: 127 [original combination].

Ancognatha manca (LeConte) [synonymy by Arrow 1937b: 6].

Types

Type of C. manca at MCZ (Endrődi 1966). Types of A. aequata and A. laevigata at BMNH (Endrődi 1966). Types of A. zuniella, A. perspicua, and A. durangoana at USNM (Endrődi 1966).

Distribution

Cyclocephala ustulata Burmeister, 1847: 39 [original combination].

Ancognatha ustulata (Burmeister) [new combination by Bates 1888: 297].

COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Puntarenas, San José. PANAMA: Chiriquí.

References

Ratcliffe 1992d, 2002a, 2003, Krajcik 2005, 2012.

Ancognatha vulgaris Arrow, 1911

Ancognatha vulgaris Arrow, 1911: 169–170 [original combination].

syn.Ancognatha (Pseudoancognatha) nigriventris Otoya, 1945: 275–282 [original combination].

Ancognatha vulgaris Arrow [synonymy by Martínez 1965a: 64].

Types

Type of A. vulgaris at BMNH (Endrődi 1966). Holotype ♂ of A. nigriventris at ICN (Otoya 1945).

Distribution

BOLIVIA. BRAZIL: Amazonas. COLOMBIA: Antioquia, Boyacá, Cauca, Cundinamarca, Huila, Magdalena, Meta, Nariño, Norte de Santander, Quindío, Risaralda, Santander, Tolima. COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Limón, Puntarenas, San José. ECUADOR. PANAMA: Bocas del Toro, Chiriquí, Coclé, Panamá, Veraguas. PERU: Ayacucho, Cajamarca, Cuzco, Huánuco, La Libertad, Lima, Loreto, Pasco, Piura, San Martín, Ucayali. VENEZUELA: Mérida.

References

Arrow 1911, 1937b, Blackwelder 1944, Otoya 1945, Gutiérrez 1950, Martínez 1965a, Howden and Campbell 1974, Endrődi 1966, 1985a, Ruiz and Pamalpa 1990, Montoya et al. 1994, Restrepo et al. 2003, Ratcliffe 2002a, 2003, Onore 1997, 2005, Pardo-Locarno et al. 2005a, Útima and Vallejo 2008, Gasca-Álvarez and Amat-García 2010, Krajcik 2005, 2012, Moore 2012, Breeschoten et al. 2013, López-García et al. 2015, Ratcliffe et al. 2015, Dossey et al. 2016, Figueroa and Ratcliffe 2016, Mitasuhashi 2016, Villalobos-Moreno et al. 2016, 2017.

Remarks

Some authors attributed the name A. humeralis to Bates (1888) and subsequently treated this taxon as a synonym of A. vulgaris (e.g., Arrow 1937b, Endrődi 1966, 1985a, Krajcik 2005, 2012). Bates (1888) clearly attributed the name A. humeralis to Burmeister (1847), and his notes on this species should not be considered a description of a new species.

The identity and species status of A. nigriventris is ambiguous and needs clarification. Ancognatha nigriventris was described from male and female specimens collected in the Colombian departments of Meta and Santander (Otoya 1945). The species was placed in a new subgenus based on its relatively well-developed maxillary teeth (Otoya 1945). Gutiérrez (1950) discussed the subgenus and Martínez (1965a) implied that the species was a synonym of A. vulgaris. Endrődi (1966) remarked that he had not seen the type series, or any specimens, of A. nigriventris and treated the species as valid. Endrődi (1985a) did not further treat A. nigriventris. Some subsequent papers have cited the species from Colombia (Ruiz and Pamalpa 1990, Montoya et al. 1994), while others have ignored it (e.g., Restrepo et al. 2003 and Gasca-Álvarez and Amat-García 2010). Ancognatha nigriventris is reported from the Colombian states of Meta, Nariño, and Santander (Otoya 1945, Ruiz and Pamalpa 1990, Montoya et al. 1994).

Genus Arriguttia Martínez, 1960

Arriguttia Martínez, 1960a: 97–98 [original usage].

Type species

Cyclocephala brevissima Arrow, 1911, by monotypy.

Keys

Endrődi 1966, 1985a, Ratcliffe 1985, Jameson et al. 2002.

Valid taxa

2 species.

Arriguttia bolivari Martínez, 1968

Arriguttia bolivari Martínez, 1968a: 185–188 [original combination].

Types

Holotype ♂ at MACN (Antonio Martínez Collection) (Martínez 1968a).

Distribution

BRAZIL: Amazonas.

References

Martínez 1968a, Krajcik 2005, 2012.

Arriguttia brevissima (Arrow, 1911)

PERU: Pasco. VENEZUELA: Carabobo.

References

Erichson 1847a, Burmeister 1847, Harold 1869a, Arrow 1937b, Pike et al. 1976, Endrődi 1966, 1985a, Krajcik 2005, 2012, Ratcliffe et al. 2015.

Aspidolea lindae Ratcliffe, 1977

Aspidolea lindae Ratcliffe, 1977: 429–430 [original combination].

Types

Holotype ♂ at UNSM (Ratcliffe 1977).

Distribution

COLOMBIA: Amazonas. PERU.

References

Ratcliffe 1977, Dupuis 1999, Restrepo-Giraldo et al. 2003, Krajcik 2005, 2012, Otavo et al. 2013, Ratcliffe et al. 2015.

Aspidolea mimethes (Höhne, 1922)

Paraspidolea mimethes Höhne, 1922a: 93–94 [original combination].

Aspidolea mimethes (Höhne) [new combination by Endrődi 1966: 342, 353–354].

Types

Male type was not found. A female type is at MTD (Endrődi 1966).

Distribution

PERU: Pasco.

References

Höhne 1922a, Arrow 1937b, Blackwelder 1944, Endrődi 1966, 1985a, Krajcik 2005, 2012, Ratcliffe et al. 2015.

Aspidolea notaticollis Höhne, 1922

Aspidolea notaticollis Höhne, 1922a: 86 [original combination].

syn.Aspidolea bigutticollis Höhne, 1922a: 87 [original combination].

Aspidolea notaticollis Höhne [synonymy by Endrődi 1966: 339, 354].

syn.Aspidolea tibialis Höhne, 1922a: 85–86 [original combination].

Endrődi 1980, 1985a, Gibernau et al. 2003, Touroult et al. 2010, Krajcik 2005, 2012, Dupuis 2014, Ponchel 2006, 2015.

Aspidolea singularis Bates, 1888

Aspidolea singularis Bates, 1888: 296–297 [original combination].

syn.Aspidolea cevallosi Martínez, 1975a: 307–313 [original combination].

Aspidolea singularis Bates [synonymy by Endrődi 1985a: 155].

syn.Aspidolea similis Höhne, 1922a: 82–83 [original combination].

Aspidolea singularis ab. similis (Höhne) [new infrasubspecific status by Endrődi 1966: 356].

Aspidolea singularis Bates [synonymy by Ratcliffe 2002a: 27].

syn.Aspidolea texana Höhne, 1922a: 84 [original combination].

Aspidolea singularis Bates [synonymy by Endrődi 1966: 356].

Types

Genus Chalepides Casey, 1915

Parachalepus (Chalepides) Casey, 1915: 176–177 [original usage]. Proposed as a subgenus.

Chalepides Casey [new genus status by Prell 1936: 151; see also Arrow 1937a: 36].

syn.Parachalepus (Parachalepus) Casey, 1915: 175–176 [original usage]. Parachalepus Casey is a junior homonym of Parachalepus Baly, 1885.

Type species

Parachalepus (Chalepides) eucephalus Casey, 1915, by original designation.

Keys

Casey 1915, Endrődi 1966, 1985a, Jameson et al. 2002, Joly and Escalona 2002a, Gasca-Álvarez and Amat-García 2010 (Colombia).

Valid taxa

15 species.

Chalepides alliaceus (Burmeister, 1847)

Chalepus alliaceus Burmeister, 1847: 77 [original combination].

Dyscinetus alliaceus (Burmeister) [new combination by Harold 1869a: 123].

Parachalepus (Chalepides) alliaceus (Burmeister) [new combination and new subgeneric classification by Casey 1915: 176].

Chalepides alliaceus (Burmeister) [new combination by Prell 1936: 151].

Types

Lectotype ♂ at MLUH (Endrődi 1966, Joly and Escalona 2002a).

Distribution

BOLIVIA: Beni, Santa Cruz. BRAZIL: Mato Grosso, Mato Grosso do Sul, Paraná, Rio de Janeiro.

References

Burmeister 1847, Harold 1869a, Ohaus 1909, Casey 1915, Prell 1936, Arrow 1937b, Anonymous 1940, Blackwelder 1944, Endrődi 1966, 1973a, 1985a, Joly and Escalona 2002a, Riehs 2005, Krajcik 2005, 2012.

Chalepides anomalus Martínez, 1978

Chalepides anomalus Martínez, 1978b: 17–19 [original combination].

Types

Holotype ♂ at MACN (Antonio Martínez Collection) (Martínez 1978b, Joly and Escalona 2002a).

Distribution

ARGENTINA: Corrientes, Entre Ríos. URUGUAY: Artigas.

References

Martínez 1978b, Endrődi 1985a, Joly and Escalona 2002a, Krajcik 2005, 2012.

Chalepides barbatus (Fabricius, 1787)

Scarabaeus barbatus Fabricius, 1787: 10 [original combination].

Melolontha barbata (Fabricius) [new combination by Fabricius 1801: 167].

Chalepus barbatus (Fabricius) [new combination by Burmeister 1847: 77].

Dyscinetus barbatus (Fabricius) [new combination by Harold 1869a: 123].

Parachalepus (Parachalepus) barbatus (Fabricius) [new combination and new subgeneric classification by Casey 1915: 175].

Chalepides barbatus (Fabricius) [new combination by Prell 1936: 151].

syn.Chalepides hydrophiloides argentinus Prell, 1937c: 9 [original combination].

Chalepides barbatus argentinus Prell [new subspecific status by Endrődi 1966: 403].

Chalepides barbatus (Fabricius) [synonymy by Ratcliffe and Cave 2015: 71].

Types

Lectotype ♀ of S. barbatus deposited at ZMUK, now housed at ZMUC (Endrődi 1966). Lectotype ♀ of C. hydrophiloides argentinus at ZMHB (Endrődi 1966).

Distribution

Arrow 1937a, b, Prell 1937a, c, Blackwelder 1944, Roze 1955, Martínez 1978b, Endrődi 1966, 1985a, Joly and Escalona 2002a, Restrepo-Giraldo et al. 2003, Gasca-Álvarez and Amat-García 2010, Ponchel 2011, Krajcik 2005, 2012.

Chalepides dilatatus (Mannerheim, 1829)

Apogonia dilatata Mannerheim, 1829: 55–56 [original combination].

Chalepus dilatatus (Mannerheim) [new combination by Burmeister 1847: 77].

Dyscinetus dilatatus (Mannerheim) [new combination by Harold 1869a: 123].

Chalepides dilatatus (Mannerheim) [new combination by Arrow 1937b: 18].

Types

Lectotype ♀ at ZMH (Endrődi 1966).

Distribution

BRAZIL: Minas Gerais, São Paulo.

References

Mannerheim 1829, Burmeister 1847, Harold 1869a, Arrow 1937b, Endrődi 1966, 1985a, Joly and Escalona 2002a, Krajcik 2005, 2012.

Chalepides eucephalus (Casey, 1915)

Parachalepus (Chalepides) eucephalus Casey, 1915: 176–177 [original combination].

Chalepides eucephalus (Casey) [new combination by Prell 1936: 151].

Types

Holotype ♀ at USNM (Endrődi 1966, Joly and Escalona 2002a).

Distribution

BRAZIL: Espírito Santo. PARAGUAY: Alto Paraná.

References

Casey 1915, Prell 1936, Endrődi 1966, 1981, 1985a, Joly and Escalona 2002a, Krajcik 2005, 2012.

Chalepides euhirtus Prokofiev, 2012

Chalepides euhirtus Prokofiev, 2012: 1–2 [original combination].

Types

Holotype ♀ at IEE (Prokofiev 2012).

Distribution

Peru: Junín.

References

Prokofiev 2012.

Remarks

Based on the original description and figures of the holotype, C. euhirtus was described from a misidentified female specimen of Aspidolea fuliginea.

Chalepides fuliginosus (Burmeister, 1847)

Chalepus fuliginosus Burmeister, 1847: 78 [original combination].

Dyscinetus fuliginosus (Burmeister) [new combination by Harold 1869a: 123].

Parachalepus (Chalepides) fuliginosus (Burmeister) [new combination and new subgeneric classification by Casey 1915: 176].

Chalepides fuliginosus (Burmeister) [new combination by Prell 1936: 151].

Types

Lectotype ♀ at MLUH (Endrődi 1966).

Distribution

ARGENTINA: Misiones. BOLIVIA: Santa Cruz. BRAZIL: Espírito Santo, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, São Paulo, Santa Catarina. CHILE: Santiago. URUGUAY: Artigas.

References

Burmeister 1847, Harold 1869a, Casey 1915, Prell 1936, Arrow 1937b, Blackwelder 1944, Endrődi 1966, 1985a, Joly and Escalona 2002a, Grossi et al. 2011, Krajcik 2005, 2012.

Remarks

Chalepides fuliginosus was collected at lights at night in Wakayama, Japan, but a population has not established there (Kusui 1992).

Chalepides howdenorum Joly & Escalona, 2002

Chalepides howdenorum Joly & Escalona, 2002a: 42, 43, 57–59 [original combination].

Types

Holotype ♂ at CNC (Joly and Escalona 2002a).

Distribution

BOLIVIA: Beni.

References

Joly and Escalona 2002a, Krajcik 2005, 2012.

Chalepides hydrophiloides (Burmeister, 1847)

Chalepus hydrophiloides Burmeister, 1847: 77 [original combination].

Dyscinetus hydrophiloides (Burmeister) [new combination by Harold 1869a: 123].

Dyscinetus barbatus (Fabricius) [synonymy by Bates 1888: 313].

Parachalepus (Parachalepus) hydrophiloides (Burmeister) [revalidated status, new combination, and new subgeneric classification by Casey 1915: 175].

Chalepides hydrophiloides (Burmeister) [new combination by Prell 1936: 151].

Chalepides barbatus (Fabricius) [synonymy by Arrow 1937b: 18].

Chalepides barbatus hydrophiloides (Burmeister) [new subspecies status by Endrődi 1966: 403].

Chalepides hydrophiloides (Burmeister) [revalidated species status by Joly and Escalona 2002a: 41, 43, 49].

syn.Parachalepus (Parachalepus) rhomboidalis Casey, 1915: 175 [original combination].

Chalepides rhomboidalis (Casey) [new combination by Prell 1936: 151].

Chalepides barbatus (Fabricius) [synonymy by Arrow 1937b: 18].

Chalepides barbatus hydrophiloides (Burmeister) [synonymy by Endrődi 1966: 403].

syn.Chalepides acillioides Prell, 1937c: 8–9 [original combination].

Chalepides barbatus hydrophiloides (Burmeister) [synonymy by Endrődi 1966: 403].

Types

Lectotype ♀ of C. hydrophiloides at MLUH (Endrődi 1966). Type of C. rhomboidalis at USNM (Endrődi 1966). Lectotype ♂ of C. acillioides at ZMHB (Endrődi 1966).

Distribution

ARGENTINA: Chaco, Santa Fe. BOLIVIA. BRAZIL: Bahia, Espírito Santo, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. PARAGUAY: Distrito Capital. URUGUAY: Artigas, Canelones, Cerro Largo, Durazno, Florida, Maldonado, Montevideo, Rivera, Treinta y Tres.

References

Burmeister 1847, Reiche 1859, Harold 1869b, Steinheil 1874, Bates 1888, Tremoleras 1910, Casey 1915, Heikertinger 1919, Arrow 1937b, Prell 1936, 1937c, Endrődi 1966, 1969a, 1985a, Saenz and Morelli 1985, Joly and Escalona 2002a, Krajcik 2005, 2012.

Chalepides luridus (Burmeister, 1847)

Chalepus luridus Burmeister, 1847: 78 [original combination].

Joly and Escalona 2002a, Krajcik 2005, 2012.

Chalepides unicolor (Endrődi, 1963)

Cyclocephala unicolor Endrődi, 1963: 331 [original combination].

Chalepides unicolor (Endrődi) [new combination by Endrődi 1964: 466].

Chalepides luridus (Burmeister) [synonymy by Endrődi 1966: 408].

Chalepides unicolor (Endrődi) [revalidated species status by Joly and Escalona 2002a: 42, 43, 64].

Types

Holotype ♂ at ZSMC (Endrődi 1966).

Distribution

ARGENTINA: La Rioja. BOLIVIA: Beni, La Paz, Santa Cruz. BRAZIL: Amapá, Amazonas, São Paulo, Rondônia.

References

Pike et al. 1976, Endrődi 1963, 1964, 1966, 1985a, Joly and Escalona 2002a, Krajcik 2005, 2012.

Genus Cyclocephala Dejean, 1821

Cyclocephala Dejean, 1821: 57 [original useage].

syn.Aclinidia Casey, 1915: 113 [original usage]. Type species: Melolontha castanea Olivier, by original designation.

Cyclocephala (Aclinidia) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Endrődi 1966: 34].

syn.Albridarollia Bolívar y Pieltain, Jiménez-Asúa, & Martínez, 1963: 182 [original usage]. Type species: Albridarollia ocellata Bolívar y Pieltain, Jiménez-Asúa, & Martínez, by original designation.

Cyclocephala Dejean [synonymy by Endrődi 1966: 34].

syn.Aspidolella Prell, 1936: 374 [original usage]. Proposed as a subgenus, replacement name for the subgenus Aspidolites Höhne. Type species: Aspidolea (Aspidolites) atricollis Höhne, by monotypy.

Cyclocephala Dejean [synonymy by Endrődi 1966: 61].

syn.Diapatalia Casey, 1915: 111 [original usage]. Type species: Cyclocephala discicollis Arrow, by original designation.

Cyclocephala (Diapatalia) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Endrődi 1966: 33].

syn.Dichromina Casey, 1915: 112 [original usage]. Type species: Cyclocephala dimidiata Burmeister, by original designation.

Cyclocephala (Dichromina) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Saylor 1945: 380].

syn.Graphalia Casey, 1915: 159 [original usage]. Proposed as a subgenus of Ochrosidia. Type species: not yet designated.

Cyclocephala (Graphalia) Casey [new subgenus classification by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Endrődi 1966: 34].

syn.Halotosia Casey, 1915: 113 [original usage]. Type species: Cyclocephala fasciolata Bates, by original designation.

Cyclocephala (Halotosia) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Endrődi 1966: 62].

syn.Homochromina Casey, 1915: 111 [original usage]. Type species: Homochromina divisa Casey, by original designation.

Cyclocephala (Homochromina) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Endrődi 1966: 34].

syn.Isocoryna Casey, 1915: 136 [original usage]. Proposed as a subgenus. Type species: Cyclocephala (Isocoryna) jalapensis Casey, by monotypy.

Cyclocephala Dejean [synonymy by Endrődi 1966: 62].

syn.Mimeoma Casey, 1915: 111 [original usage]. Type species: Cyclocephala maculata Burmeister, by original designation.

Cyclocephala Dejean [synonymy by Moore et al. 2015: 898].

syn.Mononidia Casey, 1915: 110 [original usage]. Type species: Cyclocephala carbonaria Arrow, by original designation.

Cyclocephala (Mononidia) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Endrődi 1966: 33].

syn.Ochrosidia Casey, 1915: 112 [original usage]. Type species: Melolontha immaculata Olivier, by original designation.

Cyclocephala (Ochrosidia) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Saylor 1945: 380].

syn.Paraclinidia Martínez, 1965b: 13 [original usage]. Proposed as a subgenus. Type species: Cyclocephala (Paraclinidia) endrodii Martínez, by original designation.

Cyclocephala Dejean [synonymy by Endrődi 1966: 34].

syn.Plagiosalia Casey, 1915: 135 [original usage]. Proposed as a subgenus. Type species: not yet designated.

Cyclocephala Dejean [synonymy by Endrődi 1966: 62].

syn.Spilosota Casey, 1915: 112 [original usage]. Type species: Spilosota nubeculina Casey, by original designation.

Cyclocephala (Spilosota) Casey [new subgenus status by Arrow 1937b: 8].

Cyclocephala Dejean [synonymy by Saylor 1945: 380].

syn.Stigmalia Casey, 1915: 111 [original usage]. Type species: Cyclocephala mafaffa Burmeister, by original designation.

Cyclocephala (Stigmalia) Casey [new subgenus status by Arrow 1937b: 8].

Dechambre 1980, Endrődi 1985a, Krajcik 2005, 2012.

Cyclocephala amazona amazona (Linnaeus, 1767)

Scarabaeus amazonus Linnaeus, 1767: 551 [original combination].

Melolontha amazona (Linnaeus) [new combination by Schönherr 1817: 188].

Cyclocephala amazona (Linnaeus) [new combination by Burmeister 1847: 45].

Cyclocephala amazona amazona (Linnaeus) [new subspecies status by Endrődi 1966: 147].

syn.Cyclocephala (Cyclocephala) auriculata Casey, 1915: 141 [original combination].

Cyclocephala detecta Bates [synonymy by Arrow 1937b: 10].

syn.Cyclocephala detecta Bates, 1888: 300 [original combination].

Cyclocephala (Cyclocephala) detecta Bates [new subgeneric classification by Casey 1915: 141].

Cyclocephala detecta Bates [removal of subgeneric classification by Arrow 1937b: 8, 10].

Cyclocephala amazona (Linnaeus) [synonymy by Endrődi 1964: 466].

syn.Cyclocephala (Cyclocephala) beaumonti Casey, 1915: 140 [original combination].

Cyclocephala detecta Bates [synonymy by Arrow 1937b: 10].

syn.Cyclocephala signata var. inconstans Burmeister, 1847: 45 [original combination].

Cyclocephala (Cyclocephala) inconstans Burmeister [new subgeneric classification and new species status by Casey 1915: 140].

Cyclocephala inconstans Burmeister [removal of subgeneric classification by Arrow 1937b: 8, 11].

Cyclocephala amazona ab. inconstans Burmeister [revised infrasubspecific status by Endrődi 1966: 147].

syn.Melolontha pallens Fabricius, 1798: 132 [original combination].

Cyclocephala pallens (Fabricius) [new combination by Burmeister 1847: 46].

Cyclocephala amazona ab. pallens (Fabricius) [new subspecific status by Endrődi 1964: 466].

syn.Melolontha uncinata Illiger, 1802b: 49 [original combination].

Cyclocephala signata (Fabricius) [synonymy by Arrow 1937b: 16].

syn.Melolontha signata Fabricius, 1781: 39 [original combination].

Cyclocephala signata (Fabricius) [new combination by Burmeister 1847: 43].

Cyclocephala (Cyclocephala) signata (Fabricius) [new subgeneric classification by Casey 1915: 137].

Cyclocephala signata (Fabricius) [removal of subgeneric classification by Arrow 1937b: 8, 16].

Cyclocephala amazona signata (Fabricius) [new subspecies status by Endrődi 1964: 466].

Mannerheim 1829, Burmeister 1847, Harold 1869b, Casey 1915, Luederwaldt 1916, Höhne 1923b, Arrow 1937b, Blackwelder 1944, Martínez 1975b, Pike et al. 1976, Dechambre 1980, Endrődi 1963, 1964, 1966, 1985a, Ramírez 1992, Lunau 2002, Restrepo et al. 2003, Gottsberger and Silberbauer-Gottsberger 1988, 2006, Silberbauer-Gottsberger et al. 2001, 2003, Cavalcante et al. 2009, Krajcik 2005, 2012, Maia et al. 2012, Breeschoten et al. 2013, Paulino-Neto 2014, Gasca-Álvarez and Deloya 2016, Gottsberger 1986, 1988, 1989, 1992, 1999, 2016, Costa et al. 2017.

Remarks

A single no-data specimen of C. atricapilla was reported from Mexico (Endrődi 1966), but this species is South American (Ratcliffe et al. 2013).

Cyclocephala atriceps (Casey, 1915)

Homochromina atriceps Casey, 1915: 164 [original combination].

Bates 1888, Casey 1915, Arrow 1937b, Blackwelder 1944, Pike et al. 1976, Beach 1982, Endrődi 1966, 1975b, 1985a, Young 1986, 1988a, 1990, Maes 1987, 1994, Ratcliffe 1992a, 2002a, 2003, Restrepo et al. 2003, Ratcliffe and Cave 2006, Neita-Moreno 2011, Krajcik 2005, 2012, Moore and Jameson 2013, Gasca-Álvarez and Deloya 2016.

Cyclocephala aulustjaorum Hielkema, 2017

Cyclocephala pubescens brevis Höhne, 1923b: 373 [original combination].

Cyclocephala pubescens var. brevis Höhne [new infrasubspecific status by Arrow 1937b: 15].

Cyclocephala brevis Höhne [new species status by Ratcliffe 2002a: 28].

Cyclocephala aulustjaorum Hielkema [new replacement name by Hielkema 2017: 8].

syn.Cyclocephala pubescens Burmeister, 1847: 68–69 [original combination].

Dechambre 1995, Krajcik 2005, 2012.

Cyclocephala boliviana Dechambre, 1992

Cyclocephala borealis Arrow, 1911

Cyclocephala borealis Arrow, 1911: 172 [original combination, new replacement name for Cyclocephala villosa Burmeister).

syn.Cyclocephala villosaBurmeister 1847: 54 [original combination, homonym of Cyclocephala villosa Blanchard].

Cyclocephala angularis (Knoch) [synonymy by Harold 1869b: 1241].

Cyclocephala carbonaria Arrow, 1911: 173–174 [original combination].

Mononidia carbonaria (Arrow) [new combination by Casey 1915: 114–115].

Cyclocephala carbonaria Arrow [revised combination by Arrow 1937b: 8, 9].

Mononidia carbonaria (Arrow) [revised combination by Bolívar y Pieltain et al. 1963: 185].

Cyclocephala carbonaria Arrow [revised combination by Endrődi 1966: 164].

syn.Mononidia carbonaria punctulata Prell, 1934: 162 [original combination].

Cyclocephala carbonaria var. punctulata (Prell) [new combination and new infrasubspecific status by Arrow 1937b: 9].

Cyclocephala carbonaria ab. punctulata (Prell) [revised infrasubspecific status by Endrődi 1964: 466].

Cyclocephala carbonaria Arrow [synonymy by Endrődi 1966: 164].

syn.Mononidia carbonaria trachypyga Prell, 1934: 162 [original combination].

Cyclocephala carbonaria Arrow [synonymy by Arrow 1937b: 9].

syn.Cyclocephala howdeni Endrődi, 1967b: 83–84 [original combination].

Cyclocephala carbonaria Arrow [synonymy by Ratcliffe 2003: 99].

Types

Type of C. carbonaria at BMNH (Endrődi 1966). Holotype ♂ of C. howdeni at ZMHB (Endrődi 1967b). Location of the Prell types was not reported by Endrődi (1966).

Distribution

BELIZE: Cayo, Stann Creek. BOLIVIA. COLOMBIA: Boyacá, Chocó, Córdoba, Cundinamarca, Santander, Valle del Cauca. COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Limón, Puntarenas. ECUADOR: Guayas, Santo Domingo de los Tsáchilas. GUATEMALA: Alta Verapaz, Huehuetenango, Izabal, Petén, Zacapa. HONDURAS: Atlántida, Colón, Cortés, Olancho, Santa Bárbara, Yoro. MEXICO: Chiapas, Puebla. NICARAGUA: Chontales, Jinotega, RAA Norte, Río San Juan. PANAMA: Bocas del Toro, Chiriquí, Coclé, Colón, Former Canal Zone, Panamá, San Blas. VENEZUELA: Amazonas, Bolívar.

References

Casey 1915, Prell 1934, Arrow 1911, 1937b, Blackwelder 1944, Bolívar y Pieltain et al. 1963, Pike et al. 1976, Endrődi 1964, 1966, 1967b, 1985a, Maes 1987, 1994, Joly 1995a, Dechambre 1991a, 1997, Ratcliffe and Morón 1997, Beath 1998, Ratcliffe 2002a, 2003, Restrepo et al. 2003, Neita-Moreno et al. 2006, Ratcliffe and Cave 2006, Pacheco-F. et al. 2008, Neita-Moreno 2011, Krajcik 2005, 2012, Breeschoten et al. 2013, Ratcliffe and Cave 2013, García-López et al. 2013, Ratcliffe et al. 2013, Yepes-Rodriguez et al. 2013, López-García et al. 2015, Gasca-Álvarez and Deloya 2016.

Cyclocephala cardini Chapin, 1935

References

Ratcliffe and Cave 2009, Krajcik 2012, Ratcliffe et al. 2013.

Cyclocephala castanea (Olivier, 1789)

Melolontha castanea Olivier, 1789: 79 [original combination].

Cyclocephala castanea (Olivier) [new combination by Hope 1837: 40].

Aclinidia castanea (Olivier) [new combination by Casey 1915: 113, 165].

Cyclocephala castanea (Olivier) [revised combination by Arrow 1937b: 8, 9].

syn.Melolontha elongata Olivier, 1789: 23–24 [original combination].

Cyclocephala castanea (Olivier) [synonymy by Burmeister 1847: 49].

syn.Cyclocephala latipes Laporte, 1840: 125 [original combination].

Cyclocephala castanea (Olivier) [synonymy by Burmeister 1847: 49].

syn.Melolontha valida Schönherr, 1817: 187 [original combination].

Cyclocephala cerea Burmeister [revalidated species status by Ratcliffe and Cave 2015: 75, 83].

syn.Cyclocephala sororia Bates, 1888: 303 [original combination].

Cyclocephala cerea Burmeister [synonymy by Ratcliffe and Cave 2015: 83].

Cyclocephala complanata Burmeister, 1847: 48–49 [original combination].

Cyclocephala (Plagiosalia) complanata Burmeister [new subgeneric classification by Casey 1915: 135].

Cyclocephala complanata Burmeister [removal of subgeneric classification by Arrow 1937b: 8, 9].

syn.Cyclocephala (Plagiosalia) emacerata Casey, 1915: 136 [original combination].

Gruner 1971, Pike et al. 1976, Endrődi 1966, 1985a, Dechambre 1992, Maes et al. 1997, Ratcliffe 1992a, b, 2002a, 2003, Restrepo et al. 2003, Ratcliffe and Cave 2006, Krajcik 2005, 2012, García-López et al. 2013, Dupuis 2014, Ratcliffe et al. 2013, 2015, Gasca-Álvarez and Deloya 2016.

Cyclocephala conspicua Sharp, 1877

Cyclocephala conspicua Sharp, 1877: 135 [original combination].

Stigmalia conspicua (Sharp) [new combination by Casey 1915: 123].

Cyclocephala conspicua Sharp [revised combination by Arrow 1937b: 8, 9].

syn.Cyclocephala conspicua gregaroides Dechambre, 1992: 71 [original combination].

Holotype ♂ in the Yannig Ponchel Collection (Ponchel 2009).

Distribution

FRENCH GUIANA: Maripasoula.

References

Ponchel 2009, 2011, Krajcik 2012.

Cyclocephala deceptor (Casey, 1915)

Stigmalia deceptor Casey, 1915: 117–118 [original combination].

Cyclocephala mafaffa (Burmeister) [synonymy by Arrow 1937b: 12].

Cyclocephala deceptor (Casey) [revalidated species status by Ratcliffe and Delgado 1990: 43–45].

syn.Stigmalia cuernavacana Casey, 1915: 116-117 [original combination].

Cyclocephala mafaffa Burmeister [synonymy by Arrow 1937b: 12].

Cyclocephala mafaffa ab. cuernavacana (Casey) [new infrasubspecific status by Endrődi 1966: 247].

Cyclocephala deceptor (Casey) [synonymy by Ratcliffe and Delgado-Castillo 1990: 43].

syn.Stigmalia deficiens Casey, 1915: 117 [original combination].

Cyclocephala mafaffa Burmeister [synonymy by Arrow 1937b: 12].

Cyclocephala deceptor (Casey) [synonymy by Ratcliffe and Delgado-Castillo 1990: 43].

syn.Stigmalia fallaciosa Casey, 1915: 117 [original combination].

Cyclocephala mafaffa Burmeister [synonymy by Arrow 1937b: 12].

Cyclocephala mafaffa ab. fallaciosa (Casey) [new infrasubspecific status by Endrődi 1966: 247].

Cyclocephala deceptor (Casey) [synonymy by Ratcliffe and Delgado-Castillo 1990: 43].

Types

These Casey types are at USNM (Endrődi 1966).

Distribution

BELIZE: Cayo. EL SALVADOR: Ahuachapán, La Libertad, La Paz, Morazán, San Salvador, San Vicente, Santa Ana, Sonsonate. GUATEMALA: Baja Verapaz, Chimaltenango, Chiquimula, El Progreso, Escuintla, Guatemala, Huehuetenango, Izabal, Jalapa, Jutiapa, Petén, Quiché, Sacatepéquez, San Marcos, Santa Rosa, Suchitepéquez, Zacapa. HONDURAS: Choluteca, Comayagua, Copán, Cortés, El Paraíso, Francisco Morazán, La Paz, Olancho. MEXICO: Aguascalientes, Chiapas, Colima, Durango, Estado de México, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, Sinaloa, Sonora, Tamaulipas, Veracruz. NICARAGUA: Estelí, Matagalpa.

References

Casey 1915, Arrow 1937b, Blackwelder 1944, Endrődi 1966, 1985a, Ratcliffe and Delgado-Castillo 1990, Ratcliffe and Cave 2006, Krajcik 2005, 2012, Ratcliffe et al. 2013, Deloya et al. 2014a, 2016, Romero-López and Morón 2017.

Cyclocephala dechambrei Dupuis, 2018

Cyclocephala dilatata (Prell, 1934)

Mononidia dilatata Prell, 1934: 162 [original combination].

Casey 1915, Arrow 1902, 1937b, Blackwelder 1944, Gruner 1971, Pike et al. 1976, Endrődi 1966, 1985a, Ratcliffe 2003, Núñez-Avellaneda and Neita 2009, Krajcik 2005, 2012, Núñez-Avellaneda 2014.

Remarks

Specimens of C. discicollis were reported from Nayarit, Mexico (Endrődi 1966, 1985a). Deloya et al. (2014) reported C. discicollis from Guerrero, Jalisco, and Nayarit. Navarrete-Heredia et al. (2001) reported C. discicollis from Jalisco. Major faunistic studies did not record any specimens from Mexico (Ratcliffe et al. 2013).

Cyclocephala discolor (Herbst, 1790)

Melolontha discolor Herbst, 1790: 73 [original combination].

Cyclocephala discolor (Herbst) [new combination by Burmeister 1847: 45–46].

syn.Cyclocephala andina Bréthes, 1905: 331–332 [original combination].

Cyclocephala discolor andina Bréthes [new subspecies status by Endrődi 1966: 185].

Cyclocephala discolor (Herbst) [synonymy by Ratcliffe 2003: 118].

syn.Cyclocephala aurantiaca Prell, 1937b: 496 [original combination].

Cyclocephala discolor ab. aurantiaca Prell [new infrasubspecific status by Endrődi 1966: 185].

Cyclocephala discolor (Herbst) [synonymy by Ratcliffe 2003: 118].

syn.Melolontha unciata Schönherr, 1817: 189 [original combination].

Cyclocephala discolor (Herbst) [synonymy by Burmeister 1847: 46].

Types

Lectotype ♀ of C. discolor at ZMHB (Endrődi 1966). Type of C. aurantiaca at ZMHB (Endrődi 1966). Endrődi (1966) apparently did not examine the type of C. andina but wrote that the type was at “Mus. Buenos Aires”, possibly referring to MACN.

Distribution

ARGENTINA: Salta, Tucumán. BELIZE: Cayo, Stann Creek. BOLIVIA: Beni, Cochabamba, La Paz. BRAZIL: Amazonas, Minas Gerais. COLOMBIA: Amazonas, Antioquia, Cesar, Chocó, Cundinamarca, Meta, Valle del Cauca. COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Limón, Puntarenas. ECUADOR: Los Ríos, Morona-Santiago, Napo, Pastaza. FRENCH GUIANA: Cayenne. GUATEMALA: Alta Verapaz, Izabal. GUYANA. HONDURAS: Atlántida, Colón, Comayagua, Cortés, El Paraíso, Francisco Morazán, Gracias a Dios, La Paz, Olancho, Yoro. MEXICO: Chiapas, Colima, Jalisco, Michoacán, Nayarit, Oaxaca, San Luis Potosí. NICARAGUA: Jinotega, RAA Norte, Río San Juan. PANAMA: Bocas del Toro, Chiriquí, Coclé, Colón, Darien, Former Canal Zone, Panamá. PERU: Cusco, Huánuco, Loreto, San Martín. SURINAME. TRINIDAD AND TOBAGO: Trinidad. VENEZUELA: Aragua, Capital District, Monagas.

References

Herbst 1790, Schönherr 1817, Burmeister 1847, Erichson 1848a, Senoner 1864, Harold 1869b, Bréthes 1905, Seidlitz 1905, Arrow 1937b, Prell 1937b, Blackwelder 1944, Martínez 1954, Pike et al. 1976, Endrődi 1963, 1966, 1969b, 1985a, Maes et al. 1997, Ratcliffe and Morón 1997, Ervik et al. 1999, Morón-Ríos and Morón 2001, Navarrete-Heredia et al. 2001, Ratcliffe 2002a, 2003, Restrepo et al. 2003, Ponchel 2006, Ratcliffe and Cave 2006, Pacheco-F. et al. 2008, Núñez-Avellaneda and Rojas-Robles 2008, Núñez-Avellaneda and Neita 2009, Neita-Moreno 2011, Krajcik 2005, 2012, Breeschoten et al. 2013, Ratcliffe et al. 2013, 2015, Deloya et al. 2016, Gasca-Álvarez and Deloya 2016, Romero-López and Morón 2017.

Remarks

Endrődi (1966) reported C. discolor from Haiti with no further details. This record is either spurious or erroneous (Ratcliffe and Cave 2015).

Cyclocephala dispar (Herbst, 1790)

Melolontha dispar Herbst, 1790: 65–66 [original combination].

Cyclocephala dispar (Herbst) [new combination by Reiche 1859: 7].

syn.Cyclocephala dorsalis Burmeister, 1847: 64 [original combination].

Mimeoma englemani Ratcliffe, 1977: 430–432 [original combination].

Cyclocephala englemani (Ratcliffe) [new combination by Moore et al. 2015: 898].

Types

Holotype ♂ at UNSM (Ratcliffe 1977).

Distribution

PANAMA: Darien, Former Canal Zone, Panamá.

References

Ratcliffe 1977, 2002a, 2003, Endrődi 1985a, Krajcik 2005, 2012, Moore et al. 2015.

Cyclocephala enigma Ratcliffe, 2003

Cyclocephala enigma Ratcliffe, 2003: 60, 70, 121–123 [original combination].

Types

Holotype ♂ at MNCR (originally deposited at INBio) (Ratcliffe 2003).

Distribution

COSTA RICA: Guanacaste.

References

Ratcliffe 2003, Krajcik 2005, 2012.

Cyclocephala epistomalis Bates, 1888

Cyclocephala epistomalis Bates, 1888: 303–304 [original combination].

Homochromina epistomalis (Bates) [new combination by Casey 1915: 165].

Cyclocephala epistomalis Bates [revised combination by Arrow 1937b: 8, 10].

Cyclocephala frontalis Chevrolat, 1844: 90–91 [original combination].

Cyclocephala cubana Chapin [synonymy by Endrődi 1966: 179].

Cyclocephala gravis Bates, 1888: 308 [original combination].

Cyclocephala hirta LeConte, 1861: 346 [original combination].

Spilosota hirta (LeConte) [new combination by Casey 1915: 132].

Cyclocephala (Spilosota) hirta LeConte [new subgeneric classification by Saylor 1937: 68].

Cyclocephala hirta LeConte [removal of subgeneric classification by Arrow 1937b: 8, 11].

syn.Spilosota inconspicua Casey, 1915: 133 [original combination].

Cyclocephala inconspicua (Casey) [new combination by Arrow 1937b: 8, 11].

Cyclocephala (Spilosota) hirta LeConte [synonymy by Saylor 1937: 68].

syn.Spilosota magister Casey, 1915: 132 [original combination].

Cyclocephala magister (Casey) [new combination by Arrow 1937b: 8, 12].

Cyclocephala (Spilosota) hirta LeConte [synonymy by Saylor 1937: 68].

syn.Spilosota nubeculina Casey, 1915: 131 [original combination].

Cyclocephala nubeculina (Casey) [new combination by Arrow 1937b: 8, 13].

Cyclocephala hirta LeConte [synonymy by Saylor 1945: 384].

syn.Spilosota pallidissima Casey, 1915: 133 [original combination].

Cyclocephala pallidissima (Casey) [new combination by Arrow 1937b: 8, 14].

Cyclocephala (Spilosota) hirta LeConte [synonymy by Saylor 1937: 68].

Types

Endrődi (1966) was uncertain about the housing institutions for the types of C. hirta and its synonyms. He thought they were all at USNM.

Distribution

MEXICO: Aguascalientes, Baja California, Chihuahua, Coahuila, Durango, Estado de México, Sonora. USA: Arizona, California, Colorado, Iowa, Kansas, Nebraska, Nevada, New Mexico, Oklahoma, South Dakota, Texas, Utah.

References

LeConte 1861, Gerstaecker 1862, Harold 1869b, Horn 1871, Crotch 1873, Henshaw 1885, Fall 1901, Casey 1915, Leng 1920, Dawson 1922, Moore 1937, Arrow 1937b, Saylor 1936, 1937, 1945, 1948, Blackwelder 1939, 1944, Bohart 1947, Blackwelder and Blackwelder 1948, Howden and Endrődi 1966, Hatch 1971, Kirk and Balsbaugh 1975, Ritcher and Baker 1974, Pike et al. 1976, 1977, Endrődi 1966, 1969b, 1985a, Hardy 1991, Ratcliffe 1991, Stahly and Klein 1992, Tanada and Kaya 1993, Thurston et al. 1993, 1994, Kaya et al. 1992, 1993, 1995, Cranshaw and Ward 1996, Poole and Gentili 1996, Ratcliffe and Morón 1997, Converse and Grewal 1998, Koppenhöfer and Kaya 1996, 1997, 1998, Theunis 1998, Bauernfeind 2001, Stock et al. 2001, Koppenhöfer et al. 1999, 2000a, b, c, 2002a, b, 2007, 2012, 2013, Riley and Wolfe 2003, Yildrim and Hoy 2003, Ansari et al. 2004, Cranshaw 2004, Mottern et al. 2004, Koppenhöfer and Grewal 2005, Grewal et al. 2005, Jaramillo et al. 2005, Jackson and Klein 2006, Oestergaard et al. 2006, Stuart et al. 2006, Koppenhöfer 2008, Koppenhöfer and Fuzy 2003b, c, 2008b, Ratcliffe and Paulsen 2008, Morales-Rodriguez and Peck 2009, Smith 2003, 2009, Bélair et al. 2010, Holmstrup et al. 2010, Negrisoli et al. 2010, Krajcik 2005, 2012, Perera et al. 2012, Breeschoten et al. 2013, de Coninck et al. 2013, Ratcliffe et al. 2013, Hussaini 2014, Li and Bouwer 2014, Christians et al. 2016, Gyawaly et al. 2016, Del Valle et al. 2017, Wu et al. 2017, Ratcliffe and Cave 2017.

Cyclocephala hirta pilosicollis Saylor, 1936

Spilosota hirta pilosicollis Saylor, 1936: 2 [original combination].

Cyclocephala (Spilosota) hirta pilosicollis Saylor [new subgeneric classification by Saylor 1937: 69].

Cyclocephala pilosicollis (Saylor) [removal of subgeneric classification and new species status by Arrow 1937b: 8].

Cyclocephala hirta LeConte [synonymy by Ratcliffe et al. 2013: 165].

Cyclocephala hirta pilosicollis Saylor [revalidated subspecies status by Ratcliffe and Cave 2017: 67].

Types

Type at USNM (Endrődi 1966).

Distribution

USA: California.

References

Saylor 1936, 1937, 1945, Blackwelder and Blackwelder 1948, Howden and Endrődi 1966, Pike et al. 1976, Endrődi 1966, 1985a, Hardy 1991, Poole and Gentili 1996, Krajcik 2005, 2012, Ratcliffe et al. 2013, Ratcliffe and Cave 2017.

Cyclocephala histrionica Burmeister, 1847

Cyclocephala histrionica Burmeister, 1847: 41 [original combination].

Paraspidolea histrionica (Burmeister) [new combination by Arrow 1937b: 7].

Cyclocephala histrionica Burmeister [revised combination by Endrődi 1966: 77, 127, 217–218].

Melolontha ferruginea Fabricius, 1801: 170 [original combination].

Cyclocephala ferruginea (Fabricius) [new combination by Burmeister 1847: 58].

Cyclocephala immaculata (Olivier) [synonymy by Chalumeau and Gruner 1977: 582].

Cyclocephala immaculata ferruginea (Fabricius) [revalidated subspecies status by Endrődi 1985a: 101].

syn.Melolontha nigriceps Gyllenhal, 1817a: 188–189 [original combination].

Cyclocephala ferruginea (Fabricius) [synonymy by Burmeister 1847: 58].

Types

Lectotype ♀ of M. ferruginea deposited at ZMUK, now housed at ZMUC (Endrődi 1966). The type of M. nigriceps was unknown to Endrődi (1966).

Distribution

FRENCH GUIANA: Cayenne.

References

Fabricius 1801, Gyllenhal 1817a, Schönherr 1817, Dejean 1821, 1833, 1836b, Sturm 1843, Burmeister 1847, Harold 1869b, Arrow 1937b, Blackwelder 1944, Zimsen 1964, Pike et al. 1976, Chalumeau and Gruner 1977, Dechambre 1980, Endrődi 1966, 1985a, Ponchel 2011, Krajcik 2005, 2012, Ratcliffe and Cave 2015.

Cyclocephala immaculata immaculata (Olivier, 1789)

Melolontha immaculata Olivier, 1789: 29 [original combination].

Cyclocephala immaculata (Olivier) [new combination by Dejean 1821: 57].

Ochrosidia (Ochrosidia) immaculata (Olivier) [new combination and new subgeneric classification by Casey 1915: 112, 142].

References

Pike et al. 1976, Endrődi 1964, 1966, 1985a, Young 1986, 1987, 1988a, 1990, Croat 1997, Ratcliffe 1992a, 2002a, 2003, Garcia-Robledo et al. 2004, 2005, Ratcliffe and Cave 2006, Krajcik 2005, 2012, García-López et al. 2013, Ratcliffe et al. 2013, 2015, Gasca-Álvarez and Deloya 2016.

Cyclocephala kechua (Martínez, 1957)

Cyclocephala longula LeConte [revised combination and removal of subgeneric classification by Arrow 1937b: 8, 12].

syn.Ochrosidia (Ochrosidia) abrupta Casey, 1915: 152 [original combination].

Cyclocephala abrupta (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8].

Cyclocephala (Spilosota) abrupta (Casey) [new subgeneric classification by Saylor 1937: 69].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

syn.Ochrosidia (Ochrosidia) ambiens Casey, 1915: 155 [original combination].

Cyclocephala ambiens (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

syn.Ochrosidia (Ochrosidia) rustica Casey, 1915: 157 [original combination].

Cyclocephala californica Arrow, 1937b: 9 [original combination, new replacement name for Cyclocephala rustica (Casey)].

Cyclocephala (Spilosota) abrupta (Casey) [synonymy by Saylor 1937: 69].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

syn.Ochrosidia (Ochrosidia) marcida Casey, 1915: 155 [original combination].

Cyclocephala marcida (Casey) [new combination and removal of subgeneric classification by Arrow 1937b, 8, 12].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

syn.Ochrosidia (Ochrosidia) modulata Casey, 1915: 154 [original combination].

Cyclocephala modulata (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 13].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

syn.Ochrosidia (Ochrosidia) obesula Casey, 1915: 156 [original combination].

Cyclocephala obesula (Casey) [new combination by Arrow 1937b: 8, 13].

Cyclocephala (Spilosota) abrupta (Casey) [synonymy by Saylor 1937: 69].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

syn.Ochrosidia (Ochrosidia) oblongula Casey, 1915: 156 [original combination].

Cyclocephala oblongula (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 13].

Cyclocephala (Spilosota) abrupta (Case) [synonymy by Saylor 1937: 69].

syn.Ochrosidia (Ochrosidia) phasma Casey, 1915: 153 [original combination].

Cyclocephala phasma (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 14].

Cyclocephala (Spilosota) abrupta (Casey) [synonymy by Saylor 1937: 69].

syn.Ochrosidia (Ochrosidia) prona Casey, 1915: 157 [original combination].

Cyclocephala prona (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 14].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

syn.Ochrosidia (Ochrosidia) reflexa Casey, 1915: 153 [original combination].

Cyclocephala reflexa (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 15].

Cyclocephala (Spilosota) abrupta (Casey) [synonymy by Saylor 1937: 69].

syn.Ochrosidia (Ochrosidia) rugulifrons Casey, 1915: 154 [original combination].

Cyclocephala rugulifrons (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 15].

Cyclocephala longula LeConte [synonymy by Saylor 1945: 384].

Types

Type of C. longula at MCZ (Endrődi 1966). Types of the Casey synonyms are at USNM (Endrődi 1966).

Distribution

CANADA: British Columbia. MEXICO: Baja California, Baja California Sur, Chihuahua, Sonora. UNITED STATES: Arizona, California, Colorado, Idaho, Illinois, Kansas, Montana, Nebraska, Nevada, New Mexico, Oklahoma, Oregon, South Dakota, Texas, Utah, Washington, Wyoming.

References

LeConte 1863, Gerstaecker 1865, Crotch 1873, Henshaw 1885, Horn 1894, Wickham 1896, Fall 1901, Casey 1915, Leng 1920, Dawson 1922, Arrow 1937b, Moore 1937, Blackwelder 1939, 1944, Riegel 1942b, Saylor 1937, 1945, 1948, Blackwelder and Blackwelder 1948, Allred and Beck 1965, Ritcher 1966, Howden and Endrődi 1966, Arnett 1968, Pike et al. 1976, Endrődi 1966, 1969b, 1985a, Hatch 1971, Kirk and Balsbaugh 1975, Bechtel et al. 1983, Hardy 1991, McNamara 1991, Ratcliffe 1991, Poole and Gentili 1996, Ratcliffe and Morón 1997, Bauernfeind et al. 1999, Bauernfeind 2001, Riley and Wolfe 2003, Ratcliffe and Paulsen 2008, Smith 2003, 2009, Krajcik 2005, 2012, Haviland and Hernandez 2012, Bousquet et al. 2013, Breeschoten et al. 2013, Ratcliffe et al. 2013, Ratcliffe and Cave 2017.

Remarks

Cyclocephala longula was reported from Wisconsin and Florida (USA) (Endrődi 1966, 1985a). These records are likely based on misidentifications (Ratcliffe and Paulsen 2008). Additionally, C. longula was reported from Nicaragua (Boaco and Managua) (Maes 1987, 1994). The validity of these records is uncertain as faunistic studies focused on Nicaragua did not report additional specimens of C. longula (Ratcliffe and Cave 2006).

Cyclocephala lunulata Burmeister, 1847

Cyclocephala lunulata Burmeister, 1847: 62 [original combination].

Ochrosidia (Graphalia) lunulata (Burmeister) [new combination and new subgeneric classification by Casey 1915: 159].

Cyclocephala lunulata Burmeister [revised combination and removal of subgeneric classification by Arrow 1937b: 8, 12].

syn.Cyclocephala nubeculosa Burmeister, 1847: 63 [original combination].

Cyclocephala lunulata Burmeister [synonymy by Endrődi 1964: 466].

syn.Ochrosidia (Graphalia) oblita Casey, 1915: 159 [original combination].

Cyclocephala oblita (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 13].

Cyclocephala lunulata Burmeister [synonymy by Endrődi 1964: 466].

Types

Lectotype ♀ of C. lunulata at MLUH (Endrődi 1966). Lectotype ♂ of C. nubeculosa at MLUH (Endrődi 1966). Type of O. oblita at USNM (Endrődi 1966).

Distribution

ARGENTINA: Buenos Aires, Tucumán. BELIZE: Cayo, Orange Walk, Stann Creek, Toledo. BOLIVIA: Beni, Cochabamba, La Paz, Santa Cruz. BRAZIL: Acre, Amazonas, Bahia, Espírito Santo, Goiás, Minas Gerais, Paraná, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. COLOMBIA: Antioquia, Bolívar, Boyacá, Caldas, Caquetá, Cauca, Cesar, Chocó, Córdoba, Cundinamarca, Huila, La Guajira, Meta, Nariño, Norte de Santander, Putumayo, Quindío, Risaralda, Santander, Tolima, Valle del Cauca. COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Limón, Puntarenas, San José. ECUADOR: Bolívar, Cañar, Chimborazo, El Oro, Guayas, Loja, Morona-Santiago. EL SALVADOR: Ahuachapán, Cabañas, La Libertad, La Unión, Morazán, San Miguel, San Salvador, Santa Ana, Usulután. FRENCH GUIANA: Cayenne, Sinnamary, St.-Laurent du Maroni. GUATEMALA: Alta Verapaz, Baja Verapaz, Chiquimula, El Progreso, Escuintla, Guatemala, Huehuetenango, Izabal, Jalapa, Jutiapa, Petén, Quetzaltenango, Quiché, Talhuleu, Sacatepéquez, San Marcos, Santa Rosa, Suchitepéquez, Zacapa. GUYANA: Cuyuni-Mazaruni, East Berbice-Corentyne. HONDURAS: Atlántida, Choluteca, Comayagua, Copán, Cortés, El Paraíso, Francisco Morazán, Gracias a Dios, Intibucá, Isla de la Bahía, Lempira, Ocotepeque, Olancho, Santa Bárbara, Valle, Yoro. MEXICO: Aguascalientes, Chiapas, Colima, Baja California Sur, Campeche, Chiapas, Coahuila, Distrito Federal, Estado de México, Guanajuato, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, Quintana Roo, San Luis Potosí, Sinaloa, Sonora, Tabasco, Tamaulipas, Veracruz, Yucatán, Zacatecas. NICARAGUA: Carazo, Chinandega, Chontales, Estelí, Jinotega, León, Managua, Masaya, Matagalpa, RAA Norte, RAA Sur, Río San Juan. PANAMA: Bocas del Toro, Chiriquí, Coclé, Colón, Darien, Former Canal Zone, Veraguas. PARAGUAY: Alto Paraná, Amambay. PERU: Cusco, Junín, Lima, Madre de Dios, Pasco. TRINIDAD AND TOBAGO: Trinidad. UNITED STATES: Arizona, Texas. VENEZUELA: Aragua, Capital District, Carabobo, Mérida.

References

Burmeister 1847, Reiche 1859, Harold 1869b, Bates 1888, Casey 1915, Bodkin 1919, Arrow 1937b, Blackwelder 1944, Guimarães 1944, Howden 1955, Carrillo-S. et al. 1966, Gruner 1971, Pike et al. 1976, Dechambre 1979a, Pava-O. et al. 1983, King 1984, Endrődi 1963, 1964, 1966, 1969a, b, 1985a, Maes and Tellez Robleto 1988, Hardy 1991, Pérez Dominguez 1991, Thomas 1993, Lobo and Morón 1993, Deloya and Morón 1994, Maes 1987, 1992, 1994, Morón 1994, Poole and Gentili 1996, Abraca and Quesada 1997, Ratcliffe and Morón 1997, Sanchez Soto 1997, Aragón-Garcia and Morón 2000, Marín Jarillo 2001, Navarrete-Heredia et al. 2001, Andreazze and da Silva Motta 2002, Ratcliffe 2002a, 2003, Restrepo et al. 2003, Pardo-Locarno et al. 2005a, Díaz Mederos et al. 2006, Neita-Moreno et al. 2006, Ratcliffe and Cave 2006, Múñoz-Hernández et al. 2008, Pacheco-F. et al. 2006, 2008, Útima and Vallejo 2008, García et al. 2009, Smith 2003, 2009, Stechauner-Rohringer and Pardo-Locarno 2010, Neita-Moreno 2011, Yanes-Gómez and Morón 2010, Ponchel 2011, Aragón-García et al. 2012, Krajcik 2005, 2012, Lugo et al. 2012, Breeschoten et al. 2013, García-López et al. 2013, Pardo-Locarno 2013, Rivera-Gasperín et al. 2013, Rivera-Gasperín and Morón 2013, Yepes-Rodriguez et al. 2013, Morón et al. 1985, 1988, 1998, 2014, Ratcliffe et al. 2013, 2015, López-García et al. 2015, Deloya et al. 1993, 2014a, b, 2016, Gasca-Álvarez and Deloya 2016, Ratcliffe and Cave 2017.

Remarks

Endrődi (1966) reported a single C. lunulata specimen, without details, from New Mexico (USA). Major faunistic studies did not find further specimens from New Mexico (Ratcliffe and Cave 2017).

Cyclocephala lurida coahuilae Bates, 1888

Cyclocephala coahuilae Bates, 1888: 302 [original combination].

Ochrosidia (Ochrosidia) coahuilae (Bates) [new combination and new subgeneric classification by Casey 1915: 151].

Cyclocephala coahuilae Bates [revised combination and removal of subgeneric classification by Arrow 1937b: 8, 9].

Cyclocephala immaculata coahuilae Bates [new subspecific status by Endrődi 1966: 219].

Cyclocephala lurida coahuilae Bates [revised subspecific status by Endrődi 1985a: 107].

Types

Type at BMNH (Endrődi 1966).

Distribution

MEXICO: Chihuahua, Coahuila, Durango, Estado de México, Guanajuato, Hidalgo, Jalisco, Nuevo León, Puebla, Querétaro, San Luis Potosí, Sonora, Veracruz.

References

Bates 1888, Casey 1915, Arrow 1937b, Blackwelder 1944, Pike et al. 1976, Endrődi 1966, 1985a, Ratcliffe and Morón 1997, Marín Jarillo 2001, Carrillo-Ruiz and Morón 2003, Díaz Mederos et al. 2006, Múñoz-Hernández et al. 2008, Smith 2003, 2009, Krajcik 2005, 2012, Morón and Márquez 2012, Ratcliffe et al. 2013, Ratcliffe and Cave 2017.

Remarks

Endrődi (1966, 1985a) reported C. lurida coahuilae from Costa Rica (Cartago), Colombia (no details), and Brazil (no details). This species has not been recorded again from Costa Rica, and these data are considered erroneous (Ratcliffe 2003). Cyclocephala lurida coahuilae is apparently known only from Mexico in Central America (Ratcliffe et al. 2013), and the South American records are possibly incorrect.

Cyclocephala lurida lurida Bland, 1863

Cyclocephala lurida Bland, 1863: 354 [original combination].

Spilosota lurida (Bland) [new combination by Casey 1915: 131].

Cyclocephala lurida Bland [revised combination by Arrow 1937b: 8, 12].

Cyclocephala hirta LeConte [synonymy by Saylor 1945: 384].

Cyclocephala lurida Bland [revalidated species status by Endrődi 1985a: 107].

syn.Ochrosidia (Ochrosidia) pagana Casey, 1915: 148 [original combination].

Cyclocephala pagana (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 14].

Cyclocephala immaculata (Olivier) [synonymy by Saylor 1945: 385].

Cyclocephala lurida lurida Bland [synonymy by Endrődi 1985a: 107].

syn.Ochrosidia (Ochrosidia) protenta Casey, 1915: 144 [original combination].

Cyclocephala protenta (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 14].

Cyclocephala immaculata (Olivier) [synonymy by Saylor 1945: 385].

Cyclocephala lurida lurida Bland [synonymy by Endrődi 1985a: 107].

syn.Ochrosidia (Ochrosidia) rufifrons Casey, 1915: 145 [original combination].

Cyclocephala rufifrons (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 15].

Cyclocephala immaculata (Olivier) [synonymy by Saylor 1945: 385].

Cyclocephala lurida lurida Bland [synonymy by Endrődi 1985a: 107].

syn.Ochrosidia (Ochrosidia) tenuicutis Casey, 1915: 146 [original combination].

Cyclocephala mafaffa grandis Burmeister [new subspecific status by Chalumeau 1982: 336].

The original spelling of this specific epithet is “melane” (Bates 1888). Subsequently, some authors have spelled the name as “melanae” without explanation (Endrődi 1966, 1985a, Ratcliffe 1992a, 2002a, 2003, Croat 1997, Moore and Jameson 2013).

Cyclocephala melanocephala (Fabricius, 1775)

Melolontha melanocephala Fabricius, 1775: 36 [original combination].

Cyclocephala melanocephala (Fabricius) [new combination by Burmeister 1847: 56–57].

Dichromina melanocephala (Fabricius) [new combination by Casey 1915: 160].

Cyclocephala melanocephala (Fabricius) [revised combination by Arrow 1937b: 8, 13].

syn.Chalepus leucophthalmus Fischer, 1823: 265 [original combination].

Dyscinetus leucophthalmus (Fischer) [new combination by Harold 1869a: 123].

Cyclocephala melanocephala (Fabricius) [synonymy by Arrow 1911: 172].

syn.Cyclocephala dimidiata Burmeister, 1847: 57 [original combination].

Dichromina dimidiata (Burmeister) [new combination by Casey 1915: 161].

Cyclocephala (Dichromina) dimidiata (Burmeister) [revised combination and new subgeneric classification by Saylor 1937: 70].

Cyclocephala dimidiata Burmeister [removal of subgeneric classification by Arrow 1937b: 8, 10].

Cyclocephala melanocephala (Fabricius) [synonymy by Endrődi 1964: 466].

syn.Cyclocephala elegans Horn, 1871: 337 [original combination].

Cyclocephala dimidiata Burmeister [synonymy by Horn 1875: 143].

Dichromina elegans (Horn) [revalidated species status and new combination by Casey 1915: 162].

Cyclocephala dimidiata Burmeister [synonymy by Saylor 1945: 382].

Cyclocephala melanocephala (Fabricius) [synonymy by Endrődi 1964: 466].

syn.Cyclocephala rubiginosa Burmeister, 1847: 59 [original combination].

Cyclocephala melanocephala rubiginosa Burmeister [new subspecific status by Chalumeau and Gruner 1977: 584].

Cyclocephala melanocephala (Fabricius) [synonymy by Ratcliffe and Cave 2015].

syn.Cyclocephala ventralis Erichson, 1847a: 97 [original combination].

Cyclocephala melanocephala (Fabricius) [synonymy by Endrődi 1964: 466].

syn.Dichromina ocularis Casey, 1915: 162 [original combination].

Cyclocephala ocularis (Casey) [new combination by Arrow 1937b: 8, 14].

References

Ratcliffe 2003, Krajcik 2005, 2012.

Cyclocephala mutata Harold, 1869

Cyclocephala mutata Harold, 1869a: 124 [original combination, new replacement name for Cyclocephala frontalis Burmeister, 1847].

Cyclocephala sanguinicollis ab. mutata Harold [new infrasubspecific status by Endrődi 1966: 301].

Cyclocephala mutata Harold [revalidated species status by Ratcliffe 2002a: 30].

syn.Cyclocephala frontalis Burmeister, 1847: 50 [original combination, homonym of Cyclocephala frontalisChevrolat 1844].

Cyclocephala mutata Harold [new replacement name by Harold 1869a: 124].

syn.Cyclocephala laevicauda Arrow, 1902: 138–139 [original combination].

Homochromina laevicauda (Arrow) [new combination by Casey 1915: 165].

Cyclocephala laevicauda Arrow [revised combination by Arrow 1937b: 8, 11].

Cyclocephala sanguinicollis ab. laevicauda Arrow [new infrasubspecific status by Endrődi 1966: 301].

Cyclocephala mutata Harold [synonymy by Ratcliffe 2002a: 30].

syn.Cyclocephala pseudoisabellina Endrődi, 1980: 38 [original combination].

COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Puntarenas, San José. PANAMA: Chiriquí.

References

Bates 1888, Arrow 1937b, Blackwelder 1944, Pike et al. 1976, Endrődi 1964, 1966, 1985a, Goldwasser 1987, 2000, Croat 1997, Ratcliffe 1992a, 2002a, 2003, García-Robledo et al. 2004, 2005, Krajcik 2005, 2012, Breeschoten et al. 2013, García-López et al. 2013.

Cyclocephala nigra (Endrődi, 1979)

Cyclocephala notata (Illiger, 1806)

Melolontha notata Illiger, 1806: 235–236 [original combination, new replacement name for Melolontha signata Olivier].

Cyclocephala notata (Illiger) [new combination by Burmeister 1847: 55].

syn.Cyclocephala insularis Laporte, 1840: 125 [original combination].

Lectotype ♂ at ZMHB (Endrődi 1966).

Distribution

BRAZIL: Goiás, Mato Grosso, Minas Gerais, São Paulo.

References

Höhne 1923b, Arrow 1937b, Blackwelder 1944, Pike et al. 1976, Endrődi 1966, 1985a, Gottsberger 1986, 1988, 1989, 1992, Gottsberger and Silberbauer-Gottsberger 1988, 2006, Krajcik 2005, 2012, Costa et al. 2017.

Cyclocephala olivieri Arrow, 1911

Cyclocephala olivieri Arrow, 1911: 171 [original combination, new replacement name for Cyclocephala nigrocephala Schönherr].

syn.Melolontha melanocephala Olivier, 1789: 42 [original combination, junior homonym of Melolontha melanocephala Fabricius].

Melolontha nigrocephala Schönherr, 1817: 190 [original combination, new replacement name for Melolontha melanocephala Olivier].

syn.Melolontha nigrocephala Schönherr, 1817: 190 [original combination, new replacement name for Melolontha melanocephala Olivier and junior homonym of Melolontha nigrocephala (DeGeer)].

Cyclocephala nigrocephala (Schönherr) [new combination by Burmeister 1847: 58].

Cyclocephala olivieri Arrow [new replacement name by Arrow 1911: 171].

Types

Type material for this species was not found, and Endrődi’s (1966) description was based upon Burmeister specimens.

Distribution

ARGENTINA: Formosa, Tucumán. BOLIVIA: Beni. BRAZIL: Pernambuco. COLOMBIA: Cundinamarca. PARAGUAY: Asunción. SURINAME. TRINIDAD AND TOBAGO: Trinidad. URUGUAY: Canelones, Florida, Maldonado, Montevideo, Paysandú, Rocha. VENEZUELA: Apure, Capital District, Bolívar.

References

Olivier 1789, Schönherr 1817, Burmeister 1847, Harold 1869b, Arrow 1911, 1937b, Blackwelder 1944, Pike et al. 1976, Dechambre and Endrődi 1984, Saenz and Morelli 1984, Dechambre and Endrődi 1984, Endrődi 1966, 1985a, Dechambre 1999, Restrepo et al. 2003, Krajcik 2005, 2012, Gasca-Álvarez and Deloya 2016.

Cyclocephala ovulum Bates, 1888

Cyclocephala ovulum Bates, 1888: 306–307 [original combination].

Ochrosidia (Ochrosidia) ovulum (Bates) [new combination and new subgeneric classification by Casey 1915: 158].

Seidlitz 1904, Arrow 1903, 1937b, Blackwelder 1944, Pike et al. 1976, Endrődi 1966, 1985a, Gottsberger 1986, 1989, Restrepo et al. 2003, Riehs 2005, 2006, Costa and Iannuzzi 2010, Grossi et al. 2011, Krajcik 2005, 2012, Breeschoten et al. 2013, Albuquerque et al. 2014, 2016, Ratcliffe et al. 2015, Gasca-Álvarez and Deloya 2016.

Remarks

Cyclocephala paraguayensis was recorded from Honduras (Endrődi 1966, 1985a). Further studies have not encountered C. paraguayensis in Honduras, and this species is probably restricted to South America (Ratcliffe and Cave 2006).

Cyclocephala parallela (Casey, 1915)

Ochrosidia (Ochrosidia) parallela Casey, 1915: 144 [original combination].

Cyclocephala parallela (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8 14].

Cyclocephala borealis Arrow [synonymy by Saylor 1945: 385].

Cyclocephala parallela (Casey) [revalidated species status by Endrődi 1966: 279].

Types

Types at USNM (Endrődi 1966).

Distribution

UNITED STATES: Florida, Georgia, North Carolina, South Carolina.

References

Casey 1915, Leng 1920, Arrow 1937b, Saylor 1945, Kirk 1970, Woodruff 1973, Pike et al. 1976, Reinert 1979, Gordon and Anderson 1981, Endrődi 1966, 1985a, Boucias et al. 1986, Cherry and Boucias 1989, Cherry et al. 1990, Hardy 1991, Raid and Cherry 1992, Cherry and Coale 1994, Stansly et al. 1994, Cherry and Klein 1997, Peck and Thomas 1998, Harpootlian 2001, Cappaert and Smitley 2002, Buss 2006, Jackson and Klein 2006, Frank 2008, Smith 2003, 2009, Krajcik 2005, 2012, Cherry 1984a, b, 1985, 2012, Breeschoten et al. 2013, Ratcliffe et al. 2013, Gyawaly et al. 2016, Ratcliffe and Cave 2017.

Remarks

Endrődi (1966, 1985a) reported a no-data specimen of C. parallela from Mexico. This record is likely based on a misidentification and C. parallela is restricted to the southeastern United States (Ratcliffe et al. 2013). The data points listed outside of Florida for C. parallela should be reevaluated, as some authors have hypothesized that this species is endemic to Florida (Peck and Thomas 1998, Harpootlian 2001).

Cyclocephala pardolocarnoi Dechambre, 1995

Cyclocephala pardolocarnoi Dechambre, 1995: 12–13 [original combination].

Types

Holotype ♂ at “Museo de Ciencias Naturales, Cali (Colombie)” (Dechambre 1995).

Distribution

COLOMBIA: Amazonas, Chocó, Quindío, Sucre, Tolima, Valle del Cauca. PANAMA: Bocas del Toro, Former Canal Zone, Panamá.

References

Dechambre 1995, Ratcliffe 2002a, 2003, Neita-Moreno et al. 2006, Neita-Moreno 2011, Krajcik 2005, 2012, Pardo-Locarno 2013, Gasca-Álvarez and Deloya 2016.

Cyclocephala pasadenae (Casey, 1915)

Ochrosidia (Ochrosidia) pasadenae Casey, 1915: 148 [original combination].

Cyclocephala pasadenae (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 14].

syn.Ochrosidia (Ochrosidia) arizonica Casey, 1915: 149 [original combination].

Cyclocephala arizonica (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8].

Cyclocephala pasadenae (Casey) [synonymy by Saylor 1945: 385].

syn.Ochrosidia (Ochrosidia) facilis Casey, 1915: 150 [original combination].

Cyclocephala facilis (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 10].

Cyclocephala pasadenae (Casey) [synonymy by Saylor 1945: 385].

syn.Ochrosidia (Ochrosidia) melinaCasey 1915: 149 [original combination].

Cyclocephala melina (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 13].

Cyclocephala pasadenae (Casey) [synonymy by Saylor 1945: 385].

syn.Ochrosidia (Ochrosidia) ovulata Casey, 1915: 151 [original combination].

Cyclocephala ovulata (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 14].

Cyclocephala pasadenae (Casey) [synonymy by Saylor 1945: 385].

syn.Ochrosidia (Ochrosidia) pusilla Casey, 1915: 150 [original combination].

Cyclocephala melina var. pusilla [new combination, removal of subgeneric classification, and new infrasubspecific status by Arrow 1937b: 8, 13].

Cyclocephala pasadenae (Casey) [synonymy by Saylor 1945: 385].

syn.Ochrosidia (Ochrosidia) validicepsCasey 1915: 148 [original combination].

Cyclocephala validiceps (Casey) [new combination and removal of subgeneric classification by Arrow 1937b: 8, 16].

Cyclocephala pasadenae (Casey) [synonymy by Saylor 1945: 385].

Types

The type material of all of these Casey species are at USNM (Endrődi 1966).

Distribution

MEXICO: Baja California, Baja California Sur, Chihuahua, Coahuila, Durango, Hidalgo, Jalisco, Nuevo León, San Luis Potosí, Sonora, Tamaulipas, Zacatecas. UNITED STATES: Arizona, Arkansas, California, Colorado, Hawaii, Iowa, Kansas, Missouri, Nebraska, Nevada, New Mexico, Oklahoma, South Dakota, Texas, Utah.

References

Casey 1915, Leng 1920, Arrow 1937b, Saylor 1945, 1948, Werner and Butler 1965, Ritcher 1944, 1966, Howden and Endrődi 1966, Martínez 1969, Pike et al. 1976, Endrődi 1966, 1985a, Miller 1985, Gerhardt and Stanghellini 1971, Stone 1986, Bueno et al. 1988, Hardy 1991, Ratcliffe 1991, Cokendolpher 1993, Blanco-Montero and Ward 1995, Merchant and Crocker 1996, Poole and Gentili 1996, Ratcliffe and Morón 1997, Koppenhöfer and Kaya 1997, 1998, Potter 1998, Bauernfeind et al. 1999, Bauernfeind 2001, Koppenhöfer and Fuzy 2003b, Riley and Wolfe 2003, Yildrim and Hoy 2003, Ansari et al. 2004, Cranshaw 2004, Rogers and Potter 2004b, Koppenhöfer et al. 1999, 2000a, 2002a, b, 2004, Koppenhöfer and Grewal 2005, Grewal et al. 2005, Blanco-Montero and Hernandez 1995, 2006, Ratcliffe and Paulsen 2008, Jameson et al. 2009, Morales-Rodriguez and Peck 2009, Smith 2003, 2009, Bélair et al. 2010, Holmstrup et al. 2010, Mashtoly et al. 2010, Negrisoli et al. 2010, Krajcik 2005, 2012, Perera et al. 2012, Breeschoten et al. 2013, de Coninck et al. 2013, Ratcliffe et al. 2013, Hussaini 2014, Li and Bouwer 2014, Christians et al. 2016, Gyawaly et al. 2016, Wu et al. 2017, Ratcliffe and Cave 2017.

Cyclocephala perconfusa Dechambre, 1992

Cyclocephala perconfusa Dechambre, 1992: 64–65 [original combination].

Types

Holotype ♂ at MNHN (Dechambre 1992).

Distribution

ECUADOR: Cañar, Cotopaxi, Pichincha, Santo Domingo de los Tsáchilas.

References

Dechambre 1992, Ratcliffe 1992b, Krajcik 2005, 2012.

Cyclocephala pereirai (Martínez, 1960)

Cyclocephala picipes (Olivier, 1789)

Dupuis 2008, Krajcik 2012, Dupuis 2014, Gasca-Álvarez and Deloya 2016.

Cyclocephala robusta LeConte, 1863

Cyclocephala robusta LeConte, 1863: 79 [original combination].

Cyclocephala nigricollis Burmeister [synonymy by Horn 1871: 336].

Cyclocephala robusta LeConte [revalidated species status by Saylor 1945: 384].

Cyclocephala nigricollis Burmeister [synonymy by Endrődi 1964: 466].

Cyclocephala robusta LeConte [revalidated species status by Ratcliffe and Hoffman 2011].

syn.Cyclocephala nigricollis Burmeister, 1847: 54 [original combination].

Ochrosidia nigricollis (Burmeister) [new combination by Buchanan 1927: 167].

Cyclocephala robusta LeConte [synonymy by Saylor 1945: 384].

Cyclocephala nigricollis Burmeister [new status by Endrődi 1964: 466].

Cyclocephala nigricollis Burmeister [nomen dubium by Ratcliffe and Hoffman 2011: 136].

syn.Cyclocephala subvittata Brown, 1930: 5 [original combination].

Cyclocephala robusta LeConte [synonymy by Sanderson 1940: 380].

Arrow 1911, 1937b, Blackwelder 1944, Pike et al. 1976, Endrődi 1966, 1985a, Dupuis 2006, Ponchel 2006, 2011, Krajcik 2005, 2012, Otavo et al. 2013, Ratcliffe et al. 2015, Gasca-Álvarez and Deloya 2016.

Cyclocephala rustica (Olivier, 1789)

Melolontha rustica Olivier, 1789: 27 [original combination].

Cyclocephala rustica (Olivier) [new combination by Burmeister 1847: 70].

syn.Cyclocephala rustica municipalis Höhne, 1923b: 365 [original combination].

Cyclocephala rustica var. municipalis Höhne [new infrasubspecific status by Arrow 1937b: 15].

Cyclocephala rustica municipalis Höhne [revalidated subspecies status by Endrődi 1985a: 35].

Cyclocephala rustica (Olivier) [synonymy by Ratcliffe et al. 2013: 597].

Types

Invalid neotype of M. rustica at HNHM (Endrődi Collection) (Endrődi 1966). Lectotype of C. rustica municipalis at ZMHB (Endrődi 1966).

Distribution

BRAZIL: Amazonas, Bahia, Mato Grosso, Pará, São Paulo. COLOMBIA: Casanare, Cundinamarca, Meta. FRENCH GUIANA: Cayenne, Kourou, Régina. PERU. SURINAME. TRINDAD AND TOBAGO: Trinidad. VENEZUELA: Capital District, Carabobo.

References

Olivier 1789, Burmeister 1847, Harold 1869b, Höhne 1923b, Arrow 1937b, Blackwelder 1944, Pike et al. 1976, Endrődi 1963, 1966, 1985a, Pellmyr 1985, Ratcliffe and Morón 1997, Restrepo et al. 2003, Maia and Schlindwein 2006, Ponchel 2006, 2011, Krajcik 2005, 2012, Breeschoten et al. 2013, Moore and Jameson 2013, Gibernau 2015b, López-García et al. 2015, Ratcliffe et al. 2013, 2015, Gasca-Álvarez and Deloya 2016, Peck 2016.

Remarks

Cyclocephala rustica was reported from Veracruz, Mexico (Höhne 1923b, Endrődi 1966, 1985a, Ratcliffe and Morón 1997) and Guadeloupe (Olivier 1789, Endrődi 1966, 1985a). Cyclocephala rustica has not been recorded from these areas again and is likely a South American species (Ratcliffe et al. 2013, Ratcliffe and Cave 2015).

Cyclocephala saltini Ratcliffe, 2008

Cyclocephala saltini Ratcliffe, 2008: 238–240 [original combination].

Types

Holotype ♂ at UNSM (Ratcliffe 2008).

Distribution

PERU: Huánuco.

References

Ratcliffe 2008, Krajcik 2012, Ratcliffe et al. 2015.

Cyclocephala sanguinicollis Burmeister, 1847

Cyclocephala sanguinicollis Burmeister, 1847: 49–50 [original combination].

Homochromina sanguinicollis (Burmeister, 1847) [new combination by Casey 1915: 165].

Cyclocephala sanguinicollis Burmeister [revised combination by Arrow 1937b: 8, 15].

syn.Homochromina divisa Casey, 1915: 163 [original combination].

Cyclocephala divisa (Casey) [new combination by Arrow 1937b: 8, 10].

Cyclocephala sanguinicollis Burmeister [synonymy by Endrődi 1966: 301].

Cyclocephala sexpunctata Laporte, 1840: 125 [original combination].

syn.Cyclocephala pubescens Erichson, 1847a: 96 [original combination].

Cyclocephala sexpunctata Laporte [synonymy by Endrődi 1966: 306].

syn.Cyclocephala pubescens spermophila Ohaus, 1910: 671 [original combination].

Cyclocephala pubescens var. spermophila Ohaus [new infrasubspecific status by Arrow 1937b: 15].

Cyclocephala sexpunctata var. spermophila Ohaus [new infrasubspecific status by Endrődi 1966: 306].

Cyclocephala sexpunctata ab. spermophila Ohaus [revised infrasubspecific status by Endrődi 1985a: 82].

syn.Cyclocephala pubescens nigripes Höhne, 1923b: 372–373 [original combination].

Cyclocephala pubescens var. nigripes Höhne [new infrasubspecific status by Arrow 1937b: 15].

Cyclocephala sexpunctata ab. nigripes Höhne [revised infrasubspecific status by Endrődi 1966: 306].

syn.Cyclocephala lucida Burmeister, 1847: 67 [original combination].

Stigmalia lucida (Burmeister) [new combination by Casey 1915: 120].

Cyclocephala sexpunctata Laporte [synonymy by Endrődi 1964: 466].

Cyclocephala sexpunctata ab. lucida Burmeister [new infrasubspecific status by Endrődi 1966: 306].

syn.Stigmalia circulifer Casey, 1915: 121 [original combination].

Cyclocephala lucida Burmeister [synonymy by Arrow 1937b: 12].

Cyclocephala sexpunctata Laporte [synonymy by Endrődi 1966: 306].

syn.Stigmalia costaricana Casey, 1915: 121 [original combination].

Cyclocephala costaricana (Casey) [new combination by Arrow 1937b: 8, 9].

Cyclocephala sexpunctata Laporte [synonymy by Endrődi 1966: 306].

syn.Stigmalia discoidalis Casey, 1915: 120 [original combination].

Cyclocephala lucida var. discoidalis (Casey) [new combination and new infrasubspecific status by Arrow 1937b: 12].

Cyclocephala sexpunctata Laporte [synonymy by Endrődi 1966: 306].

Cyclocephala stictica Burmeister [removal of subgeneric classification by Arrow 1937b: 8, 16].

syn.Cyclocephala stictica bilineata Höhne, 1923b: 357 [original combination].

Cyclocephala stictica var. bilineata Höhne [new infrasubspecific status by Arrow 1937b: 16].

Cyclocephala stictica ab. bilineata Höhne [revised infrasubspecific status by Endrődi 1966: 314].

Cyclocephala stictica Burmeister [synonymy by Ratcliffe 2002a: 31].

syn.Cyclocephala microspila Bates, 1888: 301 [original combination].

Cyclocephala stictica Burmeister [synonymy by Endrődi 1966: 314].

syn.Cyclocephala sexnotata Burmeister, 1847: 67 [original combination].

Cyclocephala stictica Burmeister [synonymy by Endrődi 1964: 466].

Types

syn.Cyclocephala annamariae Dutrillaux & Chalumeau, 2013: 64–65 [original combination in Dutrillaux et al. 2013].

Cyclocephala tridentata Fabricius [synonymy by Ratcliffe and Cave 2015: 113].

syn.Cyclocephala tridentata dominicensis Cartwright & Chalumeau, 1978: 25 [original combination].

Cyclocephala dominicensis Cartwright & Chalumeau [new species status by Dutrillaux et al. 2013: 64].

Cyclocephala tridentata (Fabricius) [synonymy by Ratcliffe and Cave 2015: 113].

syn.Melolontha biliturata Gyllenhal, 1817b: 105–106 [original combination].

syn.Cyclocephala rubicunda Burmeister, 1847: 61–62 [original combination].

Cyclocephala undata (Olivier) [synonymy by Endrődi 1966: 325].

syn.Melolontha spilophthalma Herbst, 1790: 163–164 [original combination].

ARGENTINA: Buenos Aires, Neuquén, Río Negro.

References

Joly and Escalona 2002b, Krajcik 2005, 2012.

Dyscinetus dubius (Olivier, 1789)

Melolontha dubia Olivier, 1789: 32 [original combination].

Dyscinetus dubius (Olivier) [new combination by Bates 1891: 32].

Dyscinetus (Palechus) dubius (Olivier) [new subgeneric classification by Casey 1915: 174].

Dyscinetus dubius (Olivier) [removal of subgeneric classification by Arrow 1937b: 17].

syn.Geotrupes lugubris Quensel, 1806: 21 [original combination].

Melolontha geminata Fabricius [synonymy by Schönherr 1817: 187].

syn.Dyscinetus frater Bates, 1888: 312 [original combination].

Dyscinetus dubius (Olivier) [synonymy by Endrődi 1966: 387].

syn.Dyscinetus (Dyscinetus) obtusus Casey, 1915: 170 [original combination].

Dyscinetus frater Bates [synonymy by Chapin 1932: 295].

syn.Melolontha geminata Fabricius, 1801: 166–167 [original combination].

Chalepus geminatus (Fabricius) [new combination by MacLeay 1819: 149–150].

Cyclocephala geminata (Fabricius) [new combination by Laporte 1840: 124].

Chalepus geminatus (Fabricius) [revised combination by Burmeister 1847: 78–79].

Dyscinetus geminatus (Fabricius) [new combination by Harold 1869a: 123].

Dyscinetus dubius (Olivier) [synonymy by Prell 1937a: 187].

Types

Type material of M. dubia is apparently missing (Endrődi 1966). Lectotype ♂ of M. geminata deposited at ZMUK, now housed at ZMUC (Endrődi 1966). Endrődi (1966) did not find the type material of G. lugubris. Lectotype ♂ of D. frater at BMNH (Endrődi 1966). Type of D. obtusus at USNM (Endrődi 1966).

Distribution

ARGENTINA: Buenos Aires, Entre Ríos, Misiones. BELIZE: Cayo. BOLIVIA: Beni, Chuquisaca, Cochabamba, La Paz, Santa Cruz. BRAZIL: Acre, Amazonas, Bahia, Espírito Santo, Goiás, Mato Grosso, Pará, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Norte, Rio Grande do Sul, Rondônia, Santa Catarina, São Paulo. CHILE. COLOMBIA: Amazonas, Antioquia, Atlántico, Boyacá, Caquetá, Cauca, Casanare, Chocó, Córdoba, Cundinamarca, Huila, Meta, Risaralda, Santander, Tolima, Valle del Cauca. COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Limón, Puntarenas, San José. CUBA: Camagüey, Ciego de Ávila, Isla de la Juventud, Matanzas, Pinar del Río, Sancti Spíritus. ECUADOR: Guayas. EL SALVADOR: Ahuachapán, Cabañas, Cuscatlán, La Libertad, San Miguel, San Salvador, Santa Ana. FRENCH GUIANA: Cayenne, St.-Laurent du Maroni. GUATEMALA: Alta Verapaz, Baja Verapaz, Chiquimula, Escuintla, Guatemala, Huehuetenango, Izabal, Petén, Quetzaltenango, Retalhuleu, San Marcos, Santa Rosa, Suchitepéquez, Zacapa. GUYANA: Demerara-Mahaica, Mahaica-Berbice. HONDURAS: Atlántida, Choluteca, Mayagua, Cortés, El Paraíso, Francisco Morazán, Gracias a Dios, La Paz, Olancho. MEXICO: Campeche, Chiapas, Distrito Federal, Guerrero, Jalisco, Nayarit, Oaxaca, Quintana Roo, San Luis Potosí, Sinaloa, Tabasco, Tamaulipas, Tlaxcala, Veracruz. NICARAGUA: Carazo, Granada, Managua, Masaya, RAA Sur, Río San Juan, Rivas. PANAMA: Bocas del Toro, Panama Canal Zone, Chiriquí, Coclé, Colón, Darien, Panamá, San Blas. PARAGUAY: Central, Cordillera, Guaíra, Paraguarí. PERU. SURINAME: Paramaribo District. TRINIDAD AND TOBAGO: Trinidad. VENEZUELA: Amazonas, Apure, Aragua, Bolívar, Carabobo, Delta Amacuro, Falcón, Guárico, Mérida, Monagas, Portuguesa, Táchira, Trujillo, Zulia.

References

Olivier 1789, Fabricius 1801, Quensel 1806, Schönherr 1817, MacLeay 1819, Dejean 1821, 1833, 1836b, Latreille 1829, Laporte 1840, Sturm 1843, Burmeister 1847, Erichson 1848a, Reiche 1859, Fauvel 1861, Harold 1869a, Bates 1888, 1891, Ohaus 1900, Casey 1915, Cleare 1925, 1930, Chapin 1932, Squire 1932, 1933, Dash 1934, Prell 1937a, Arrow 1937b, Anonymous 1900, 1940, Blackwelder 1944, Guimarães 1944, Wiehe 1951, Figueroa-P. 1952, Lima 1953, Roze 1955, Gibson and Carrillo 1959, Van Dinther 1960, Zimsen 1964, Endrődi 1966, 1973a, 1985a, Dechambre 1979a, Morón 1979, Remillet 1988, Aguilera et al. 1993, Thomas 1993, Lobo and Morón 1993, Maes 1987, 1994, Ratcliffe and Morón 1997, Sanchez Soto 1998, Andreazze and Fonseca 1998, Ratcliffe 1986, 2002a, 2003, Restrepo-Giraldo et al. 2003, Pardo-Locarno et al. 1995, 2005a, Riehs 2005, Fernández García 2006, Ratcliffe and Cave 2006, Pacheco-F. et al. 2008, Marques and Gil-Santana 2009, Gasca-Álvarez and Amat-García 2010, Joly and Escalona 2010, Ponchel 2011, Neita-Moreno and Yepes 2011, Neita-Moreno 2011, Krajcik 2005, 2012, Breeschoten et al. 2013, Pardo-Locarno 2013, Otavo et al. 2013, García-López et al. 2013, Ratcliffe et al. 2013, Deloya et al. 2014a, Ratcliffe et al. 2013, 2015, Ratcliffe and Cave 2015, García-Atencia and Martínez-Hernández 2015, López-García et al. 2015, Albuquerque et al. 2016.

Dyscinetus dytiscoides Arrow, 1911

Dyscinetus dytiscoides Arrow, 1911: 168 [original combination].

Chalepides dytiscoides (Arrow) [new combination by Endrődi 1967a: 411].

Dyscinetus dytiscoides Arrow [revised combination by Joly and Escalona 2002a: 40].

Types

Chapin 1935a, Bruner et al. 1975, Endrődi 1966, 1985a, Fernández García 2006, Krajcik 2005, 2012, Ratcliffe and Cave 2015.

Dyscinetus morator (Fabricius, 1798)

Scarabaeus morator Fabricius, 1798: 24 [original combination].

Geotrupes morator (Fabricius) [new combination by Schönherr 1806: 22].

Heteronychus morator (Fabricius) [new combination by Burmeister 1847: 97].

Dyscinetus morator (Fabricius) [new combination by Arrow 1937b: 17].

syn.Dyscinetus (Dyscinetus) bitumorosus Casey, 1915: 171 [original combination].

Dyscinetus bitumorosus Casey [removal of subgeneric classification by Arrow 1937b: 17].

Dyscinetus morator (Fabricius) [synonymy by Endrődi 1966: 393].

syn.Dyscinetus (Dyscinetus) borealis Casey, 1915: 171 [original combination].

Dyscinetus morator (Fabricius) [synonymy by Arrow 1937b: 18].

syn.Dyscinetus (Dyscinetus) trachypygus discedens Casey, 1915: 171 [original combination].

Dyscinetus morator (Fabricius) [synonymy by Arrow 1937b: 18].

COLOMBIA: Nariño.

References

Ratcliffe 1986, Restrepo-Giraldo et al. 2003, Gasca-Álvarez and Amat-García 2010, Krajcik 2005, 2012.

Dyscinetus paradytis (Ponchel & Dechambre, 2003)

Chalepides paradytis Ponchel & Dechambre, 2003: 267 [original combination].

Dyscinetus paradytis (Ponchel & Dechambre) [new combination by Joly and Escalona 2010: 206, 212].

Types

Holotype ♂ at MNHN (Ponchel and Dechambre 2003).

Distribution

BOLIVIA: Beni. BRAZIL: Mato Grosso. COLOMBIA: Caquetá. PERU: Loreto. VENEZUELA: Amazonas, Bolívar, Delta Amacuro, Guárico.

References

Ponchel and Dechambre 2003, Joly and Escalona 2010, Krajcik 2005, 2012, Ratcliffe et al. 2015.

Dyscinetus picipes (Burmeister, 1847)

Chalepus picipes Burmeister, 1847: 79–80 [original combination].

Dyscinetus picipes (Burmeister) [new combination by Harold 1869a: 123].

Dyscinetus (Dyscinetus) picipes (Burmeister) [new subgeneric classification by Casey 1915: 169].

Dyscinetus picipes (Burmeister) [removal of subgeneric classification by Arrow 1937b: 18].

syn.Chalepus geminatus Jacquelin du Val, 1857: 54 [original combination].

Chalepus picipes Burmeister [synonymy by Chevrolat 1865: 31].

syn.Chalepus obsoletus LeConte, 1854: 222 [original combination].

Dyscinetus obsoletus (LeConte) [new combination by Harold 1869a: 123].

Dyscinetus picipes (Burmeister) [synonymy by Saylor 1945: 380].

syn.Dyscinetus (Dyscinetus) ebeninus Casey, 1915: 169 [original combination].

Dyscinetus picipes (Burmeister) [synonymy by Chapin 1932: 293].

Dyscinetus puncticauda Casey [synonymy by Arrow 1937b: 18].

Dyscinetus picipes (Burmeister) [synonymy by Blackwelder 1944: 253].

syn.Dyscinetus (Dyscinetus) laevissimus Casey, 1915: 167 [original combination].

Dyscinetus obsoletus (LeConte) [synonymy by Arrow 1937b: 18].

Dyscinetus picipes (Burmeister) [synonymy Saylor 1945: 380].

syn.Dyscinetus (Dyscinetus) obsidianus Casey, 1915: 168 [original combination].

Dyscinetus obsidianus Casey [removal of subgeneric classification by Arrow 1937b: 18].

Dyscinetus picipes (Burmeister) [synonymy by Endrődi 1966: 395].

syn.Dyscinetus (Dyscinetus) obsoletus gilanus Casey, 1915: 168 [original combination].

Dyscinetus obsoletus var. gilanus Casey [new infrasubspecific status by Blackwelder 1944: 253].

Dyscinetus picipes (Burmeister) [synonymy by Saylor 1945: 380].

syn.Dyscinetus picipes Bates, 1888: 312 [original combination, homonym of Dyscinetus picipes (Burmeister)].

Dyscinetus picipes (Burmeister) [synonymy by Chapin 1932: 293].

Dyscinetus punctipes Saylor, 1945: 380 [new replacement name for Dyscinetus picipes Bates].

syn.Dyscinetus picipes puertoricensis Chalumeau, 1982: 342 [original combination].

Dyscinetus picipes (Burmeister) [synonymy by Ratcliffe and Cave 2015: 132].

syn.Dyscinetus puncticauda Casey, 1909: 282–283 [original combination].

Dyscinetus picipes (Burmeister) [synonymy by Chapin 1932: 293].

syn.Dyscinetus (Dyscinetus) subquadratus Casey, 1915: 166 [original combination].

Dyscinetus picipes (Burmeister) [synonymy by Chapin 1932: 293].

Dyscinetus puncticauda Casey [synonymy by Arrow 1937b: 18].

Dyscinetus picipes (Burmeister) [synonymy by Blackwelder 1944: 253].

Types

Endrődi (1966) commented that he doubted the D. picipes specimens in the Burmeister Collection at MLUH were the types. Chalumeau (1982) later designated a Chalepus picipes Burmeister lectotype ♂ at MLUH. Endrődi (1966) did not examine the types of C. geminatus. Type of C. obsoletus at MCZ (Endrődi 1966). Type of D. picipes Bates at BMNH (Endrődi 1966). The types of the Casey synonyms are at USNM (Endrődi 1966).

Distribution

ANTIGUA AND BARBUDA: Barbuda. BAHAMAS: Great Inagua. CUBA: Artemisa, Camagüey, Ciego de Ávila, Cienfuegos, Granma, Guantánamo, Holguin, Isla de a Juventud, La Habana, Matanzas, Mayabeque, Pinar del Río, Sancti Spíritus, Santiago de Cuba, Villa Clara. DOMINICAN REPUBLIC: La Vega, San Pedro de Macorís, Santo Domingo. GUADELOUPE: Gourbeyre, Marie-Galante, Pointe-à-Pitre, Saint-Claude, Trois-Rivières. HAITI: Ouest. MARTINIQUE. MEXICO: Aguascalientes, Coahuila, Distrito Federal, Durango, Estado de México, Guerrero, Hidalgo, Jalisco, Michoacán, Nayarit, Nuevo León, Puebla, Querétaro, San Luis Potosí, Sinaloa, Tabasco, Tamaulipas, Veracruz. PUERTO RICO: Adjuntas, Aibonito, Añasco, Arecibo, Barceloneta, Bayamón, Cabo Rojo, Carolina, Cayey, Comerio, Dorado, Fajardo, Guánica, Hormigueros, Humacao, Lajas, Loiza, Mayagüez, Orocovis, Ponce, Río Grande, San Germán, San Juan, San Sebastián, Toa Baja, Vega Baja, Yabucoa, Yauco. UNITED STATES: Arizona, Colorado, Iowa, Kansas, Missouri, Montana, Nebraska, New Mexico, Oklahoma, South Dakota, Texas.

References

Sturm 1843, Burmeister 1847, LeConte 1854, Jacquelin du Val 1857, Harold 1869a, Chevrolat 1865, Crotch 1873, Stahl 1882, Henshaw 1885, Bates 1888, Fleutiaux and Sallé 1889, Gundlach 1891, Casey 1909, 1915, Leng and Mutchler 1914, 1917, Leng 1920, Dawson 1922, Stahl and Scaramuzza 1929, Chapin 1932, Arrow 1937b, Blackwelder 1944, Saylor 1945, Paulian 1947, Blackwelder and Blackwelder 1948, Wolcott 1936, 1937, 1948, Barry 1951, Gibson and Carrillo 1959, Van Dinther 1960, Ritcher and Baker 1974, Bruner et al. 1975, Gruner 1975, Kirk and Balsbaugh 1975, Morón and Zaragoza 1976, Chalumeau and Gruner 1977, Martínez 1978b, Chalumeau 1982, Endrődi 1966, 1985a, Maes 1987, 1994, Hardy 1991, Morón et al. 1996, Poole and Gentili 1996, Ratcliffe and Morón 1997, González et al. 1998, Dechambre 1999, Navarrete-Heredia et al. 2001, Ratcliffe 1991, 2002b, Riley and Wolfe 2003, Fernández García 2006, Ratcliffe and Paulsen 2008, Smith 2003, 2009, Krajcik 2005, 2012, Breeschoten et al. 2013, Lugo et al. 2013, Ratcliffe et al. 2013, Ratcliffe and Cave 2008, 2015, 2017.

Remarks

Dyscinetus picipes was reported from Nicaragua (Maes 1987, 1994), but major faunistic studies did not find additional specimens (Ratcliffe and Cave 2006).

Dyscinetus plicatus (Burmeister, 1847)

Chalepus plicatus Burmeister, 1847: 80–81 [original combination].

Dyscinetus plicatus (Burmeister) [new combination by Harold 1869a: 123].

Types

Endrődi (1966) did not find any type material of this species.

Distribution

BRAZIL: Mato Grosso, Paraná, São Paulo.

References

Burmeister 1847, Harold 1869a, Arrow 1937b, Blackwelder 1944, Endrődi 1966, 1985a, Gottsberger 1989, Grossi et al. 2011, Krajcik 2005, 2012.

Dyscinetus questeli Chalumeau, 1982

Dyscinetus questeli Chalumeau, 1982: 340–341 [original combination].

Types

Holotype ♂ of D. questeli at USNM (Chalumeau 1982).

Distribution

GUADELOUPE: Gourbeyre, Marie-Galante, Petit-Bourg, Sainte-Anne, Saint-Claude, Saint-Louis, Sainte-Rose, Trois-Rivières.

References

Chalumeau 1982, Krajcik 2005, 2012, Ratcliffe and Cave 2015, Peck 2016.

Dyscinetus rugifrons (Burmeister, 1847)

Chalepus rugifrons Burmeister, 1847: 80 [original combination].

Dyscinetus rugifrons (Burmeister) [new combination by Harold 1869a: 123].

syn.Chalepus planatus Burmeister, 1847: 80 [original combination].

Dyscinetus planatus (Burmeister) [new combination by Harold 1869a: 123].

Dyscinetus rugifrons (Burmeister) [synonymy by Endrődi 1966: 396].

syn.Euetheola sinyaevi Prokofiev, 2012: 8–10 [original combination].

Dyscinetus rugifrons (Burmeister) [synonymy by Prokofiev 2013: 131].

Types

Lectotype ♂ of C. rugifrons and lectotype ♂ of C. planatus both at MLUH (Endrődi 1966). Holotype ♂ of E. sinyaevi at IEE (Prokofiev 2012).

Distribution

ARGENTINA: Buenos Aires, Chaco, Córdoba, Entre Ríos, Formosa, Jujuy, Misiones, Santa Fe, Tucumán. BOLIVIA: Cochabamba, Santa Cruz. BRAZIL: Bahia, Espírito Santo, Minas Gerais, Goiás, Paraná, Pernambuco, Rio de Janeiro, Rio Grande do Sul, Santa Catarina, São Paulo. PARAGUAY: Guairá, Itapúa. URUGUAY: Canelones, Montevideo. VENEZUELA: Bolívar.

References

Burmeister 1847, Reiche 1859, Harold 1869a, Steinheil 1874, Ohaus 1900, 1909, Tremoleras 1910, Luederwaldt 1915, Bosq 1936, Arrow 1937b, Anonymous 1900, 1940, Blackwelder 1944, Guimarães 1944, Guérin 1953, Silveira Guido 1965, Perkins 1974, Vidal et al. 1979, Endrődi 1966, 1969a, 1985a, Dennis and Knutson 1988, Costilla 1991, Lachaume 1992, Dechambre 1999, Vincini et al. 2000, Mauco and Favero 2004, Riehs 2005, Soave et al. 2006, Vitorino et al. 2008, Marques and Gil-Santana 2009, Joly and Escalona 2002b, 2010, Berón and Favero 2010, Krajcik 2005, 2012, Breeschoten et al. 2013, Cherman et al. 2013, Prokofiev 2012, 2013, 2014, Albuquerque et al. 2016.

Dyscinetus sculptus Dupuis, 2006

Dyscinetus sculptus Dupuis, 2006: 310–312 [original combination].

Types

Holotype ♂ at FDPC (Dupuis 2006).

Distribution

BOLIVIA: Santa Cruz.

References

Dupuis 2006, Krajcik 2012.

Dyscinetus subsericeus (Burmeister, 1847)

Chalepus subsericeus Burmeister, 1847: 81 [original combination].

Dyscinetus subsericeus (Burmeister) [new combination by Harold 1869a: 123].

Types

Lectotype ♂ at MLUH (Endrődi 1966).

Distribution

BRAZIL.

References

Burmeister 1847, Harold 1869a, Arrow 1937b, Blackwelder 1944, Endrődi 1966, 1985a, Krajcik 2005, 2012.

Genus Erioscelis Burmeister, 1847

Erioscelis Burmeister, 1847: 72–73 [original usage].

Type species

Apogonia emarginata Mannerheim, 1829, by monotypy.

Keys

Saylor 1946, Endrődi 1966, 1985a, Ratcliffe 1985, Maes 1994 (Nicaragua), Jameson et al. 2002, Ratcliffe 2003 (Costa Rica and Panama), Ratcliffe and Cave 2006 (Honduras, Nicaragua, and El Salvador), Gasca-Álvarez and Amat-García 2010 (Colombia).

Valid taxa

5 species.

Erioscelis columbica Endrődi, 1966

Erioscelis columbica Endrődi, 1966: 410, 413–414 [original combination].

Types

Holotype ♂ at HNHM (Endrődi collection) (Endrődi 1966).

Distribution

COSTA RICA: Heredia, Limón. COLOMBIA: Chocó, Cundinamarca, Meta. NICARAGUA: Jinotega, RAA Norte. PANAMA: Bocas del Toro, Colón, Darien, Panamá.

References

Endrődi 1966, 1985a, Young 1986, 1988a, 1988b, 1990, Maes et al. 1997, Beath 1998, 1999, Grayum 1996, Croat 1997, Morón 1997, Ratcliffe 2002a, 2003, Ratcliffe and Cave 2006, Gasca-Álvarez and Amat-García 2010, Neita-Moreno 2011, Krajcik 2005, 2012, Moore 2012, Moore and Jameson 2013.

Erioscelis emarginata (Mannerheim, 1829)

Apogonia emarginata Mannerheim, 1829: 54–55 [original combination].

Erioscelis emarginata (Mannerheim) [new combination by Burmeister 1847: 73].

Types

Type at ZMH (Endrődi 1966).

Distribution

ARGENTINA: Misiones. BRAZIL: Minas Gerais, Pará, Paraná, Rio de Janeiro, Rio Grande do Sul, São Paulo, Santa Catarina. ECUADOR. FRENCH GUIANA: Cayenne. PARAGUAY: Concepción.

References

Mannerheim 1829, Dejean 1821, 1833, 1836b, Burmeister 1847, Harold 1869b, Schrottky 1908, 1910, Strand 1912, Luederwaldt 1915, 1916, Arrow 1937b, Ogloblin 1941, Blackwelder 1944, Saylor 1946, Martínez 1968a, Gruner 1971, Gottsberger and Amaral 1984, Endrődi 1966, 1985a, Gottsberger and Silberbauer-Gottsberger 1991, Weber et al. 2001, Riehs 2005, Weber 2008, Dötterl et al. 2012, Krajcik 2005, 2012, Breeschoten et al. 2013, Gottsberger et al. 2013, Moore and Jameson 2013, Maia et al. 2014, Pereira et al. 2014, Maldonado et al. 2015, Gottsberger 1986, 1990, 2016.

Erioscelis peruana Saylor, 1946

Erioscelis peruana Saylor, 1946: 63–65 [original combination].

Types

Holotype ♂ at CAS (Saylor Collection) (Endrődi 1966).

Distribution

PERU: Ucayali.

References

Saylor 1946, Endrődi 1966, 1985a, Krajcik 2005, 2012, Ratcliffe et al. 2015.

Erioscelis proba (Sharp, 1877)

Cyclocephala proba Sharp, 1877: 135–136 [original combination].

Erioscelis proba (Sharp) [new combination by Endrődi 1966: 410, 412].

syn.Dyscinetus curtus Kirsch, 1873: 345–346 [original combination].

Melolontha morio Fabricius, 1798: 131 [original combination].

Chalepus morio (Fabricius) [new combination by Hope 1837: 71].

Heteronychus morio (Fabricius) [new combination by Burmeister 1847: 95].

Ruteloryctes morio (Fabricius) [new combination by Prell 1937a: 188].

syn.Ruteloryctes tristis Arrow, 1908: 336 [original combination].

Ruteloryctes morio (Fabricius) [synonymy by Paulian 1954: 1123].

Stenocrates inelegans Arrow, 1913: 465–466 [original combination].

Geotrupes holomelanus Germar, 1824: 116–117 [original combination].

Dechambre and Hardy 2004, Krajcik 2012, Ratcliffe 2015.

Stenocrates laborator (Fabricius, 1775)

Scarabaeus laborator Fabricius: 18 [original combination].

Geotrupes laborator (Fabricius) [new combination by Eschscholtz 1822: 121].

Stenocrates laborator (Fabricius) [new combination by Burmeister 1847: 85].

syn.Geotrupes globator Thunberg, 1814: 400 [original combination].

Stenocrates laborator (Fabricius) [synonymy by Endrődi 1966: 430].

syn.Geotrupes thoracicus Eschscholtz, 1818: 453–454 [original combination].

Geotrupes laborator var. thoracicus Eschscholtz [new infrasubspecific status by Eschscholtz 1822: 121].

Stenocrates laborator var. thoracicus (Eschscholtz) [new combination by Arrow 1937b: 19].

Ratcliffe 2015.

Stenocrates minutus Endrődi, 1966

Stenocrates minutus Endrődi, 1966: 415, 432–433 [original combination].

Endrődi 1985a, Dechambre and Hardy 2004, Krajcik 2005, 2012, Ratcliffe 1978, 2015.

Stenocrates rufipennis (Fabricius, 1801)

Melolontha rufipennis Fabricius, 1801: 167 [original combination].

Abdullah M, Roohi RA (1969) The cockchafers and dung-rollers of Pakistan of the Desmonycinae, Euchirinae and Rutelinae (Peltonotini, Parastasiini and Adorrhinyptiini) along with the description of five new species of Anomala (Coleoptera: Scarabaeidae). Pakistan Journal of Science and Industrial Research 12: 121–126.

Abraca G, Quesada M (1997) Especies del complejo de jobotos (Phyllophaga spp., Anomala spp. y Cyclocephala spp.) asociadas a cultivos, en el valle Central y Pacífico seco de Costa Rica. Agronomía Mesoamericana 8: 44–53.

Abrahão J, Amante E (1970) Fungos e insetos causadores de tombamento de mudas de algodoeiro no ano agrícola 1969–70. O Biológico 36: 24–25.

Achinelly MF, Camino NB (2008) A new Nematoda (Thelastomatidae) parasite of Coleoptera larvae from Argentina. Helminthologia 45: 86–88. https://doi.org/10.2478/s11687-008-0016-1

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