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].

syn.Cyclocephala meinanderi Endrődi, 1964: 457–459 [original combination].

Cyclocephala gravis ab. meinanderi Endrődi [new infrasubspecific status by Endrődi 1967: 90].

Cyclocephala gravis Bates [synonymy by Maes 1994: 11].

Types

Type of C. gravis at BMNH (Endrődi 1966). Holotype ♂ of C. meinanderi at HNHM (Endrődi Collection) (Endrődi 1964).

Distribution

BELIZE: Cayo, Orange Walk, Stann Creek, Toledo. BOLIVIA: Cochabamba, Santa Cruz. BRAZIL: Bahia, Espírito Santo, Mato Grosso, Minas Gerais, Pernambuco, Rio de Janeiro. COLOMBIA: Antioquia, Boyacá, Caquetá, Chocó, Cundinamarca, Magdalena, Santander, Tolima. COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia, Limón, Puntarenas, San José. ECUADOR: Santo Domingo de los Tsáchilas. EL SALVADOR: Morazán. FRENCH GUIANA: Cayenne, St.-Élie, St.-Laurent du Maroni. GUATEMALA: Alta Verapaz, Huehuetenango, Izabal, Petén, Sololá, Zacapa. GUYANA: Upper Demerara-Berbice. HONDURAS: Atlántida, Choluteca, Cortés, El Paraíso, Francisco Morazán, Gracias a Dios, Olancho, Santa Bárbara, Yoro. MEXICO: Campeche, Chiapas, Guanajuato, Oaxaca, Quintana Roo, San Luis Potosí, Tabasco, Veracruz, Yucatán. NICARAGUA: Chontales, Jinotega, Matagalpa, RAA Norte, Río San Juan. PANAMA: Bocas del Toro, Chiriquí, Colón, Former Canal Zone, Panamá. PARAGUAY: Distrito Capital. SURINAME. VENEZUELA: Capital District, Guárico.

References

Bates 1888, Bodkin 1919, Arrow 1937b, Blackwelder 1944, Pike et al. 1976, Dechambre 1979a, 1980, Endrődi 1964, 1966, 1967, 1985a, Young 1986, 1988a, b, 1990, Thomas 1993, Lobo and Morón 1993, Maes 1987, 1994, Ramírez and Brito 1992, Croat 1997, Ratcliffe and Morón 1997, Beath 1998, 1999, Ratcliffe 1992b, 2002a, 2003, Restrepo et al. 2003, Reyes Novelo and Morón 2005, Ratcliffe and Cave 2006, Krajcik 2005, 2012, Neita-Moreno 2011, Ponchel 2011, García-López et al. 2013, Moore and Jameson 2013, Ratcliffe et al. 2013, López-García et al. 2015, Gasca-Álvarez and Deloya 2016.

Cyclocephala gregaria Heyne & Taschenberg, 1907

Cyclocephala gregaria Heyne & Taschenberg, 1907: 91–92 [original combination]

Stigmalia gregaria (Heyne & Taschenberg) [new combination by Casey 1915: 115, 122].

Cyclocephala gregaria Heyne & Taschenberg [revised combination by Arrow 1937b: 8, 11].

syn.Cyclocephala gregaria pallida Arrow, 1911: 172 [original combination].

Cyclocephala gregaria ab. pallida Arrow [new infrasubspecific status by Endrődi 1966: 210].

Types

Invalid neotype ♂ of C. gregaria at HNHM (Endrődi Collection) (Endrődi 1966). Endrődi (1966) did not examine the type of C. gregaria pallida, and it may be at BMNH.

Distribution

BOLIVIA: Santa Cruz. BRAZIL: Pará, São Paulo. COLOMBIA: Antioquia, Boyacá, Caldas, Cauca, Cundinamarca, Magdalena, Quindío, Risaralda, Santander, Tolima, Valle del Cauca. ECUADOR: Morona-Santiago. VENEZUELA: Mérida.

References

Heyne and Taschenberg 1907, Casey 1915, Arrow 1911, 1937b, Blackwelder 1944, Daniel 1945, Martínez 1968c, Pike et al. 1976, Endrődi 1966, 1985a, Montoya et al. 1994, Dechambre 1995, Restrepo et al. 2003, García-Robledo et al. 2004, 2005, Útima and Vallejo 2008, García-Robledo 2010, Krajcik 2005, 2012, Breeschoten et al. 2013, Yepes-Rodriguez et al. 2013, López-García et al. 2015, Gasca-Álvarez and Deloya 2016.