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Synopsis of the pelidnotine scarabs (Coleoptera, Scarabaeidae, Rutelinae, Rutelini) and annotated catalog of the species and subspecies
expand article infoMatthew R. Moore, Mary L. Jameson§, Beulah H. Garner|, Cédric Audibert, Andrew B. T. Smith#, Matthias Seidel¤
‡ University of Florida, Gainesville, United States of America
§ Wichita State University, Wichita, United States of America
| Natural History Museum, London, United Kingdom
¶ Musée des Confluences, Lyon, France
# Canadian Museum of Nature, Ottawa, Canada
¤ University of Prague, Prague, Czech Republic
Open Access

Abstract

The pelidnotine scarabs (Scarabaeidae: Rutelinae: Rutelini) are a speciose, paraphyletic assemblage of beetles that includes spectacular metallic species (“jewel scarabs”) as well as species that are ecologically important as herbivores, pollinators, and bioindicators. These beetles suffer from a complicated nomenclatural history, due primarily to 20th century taxonomic and nomenclatural errors. We review the taxonomic history of the pelidnotine scarabs, present a provisional key to genera with overviews of all genera, and synthesize a catalog of all taxa with synonyms, distributional data, type specimen information, and 107 images of exemplar species. As a result of our research, the pelidnotine leaf chafers (a paraphyletic group) include 27 (26 extant and 1 extinct) genera and 420 valid species and subspecies (419 extant and 1 extinct). Our research makes biodiversity research on this group tractable and accessible, thus setting the stage for future studies that address evolutionary and ecological trends. Based on our research, 1 new species is described, 1 new generic synonym and 12 new species synonyms are proposed, 11 new lectotypes and 1 new neotype are designated, many new or revised nomenclatural combinations, and many unavailable names are presented. The following taxonomic changes are made:

New generic synonym: The genus Heteropelidnota Ohaus, 1912 is a new junior synonym of Pelidnota MacLeay, 1819.

New species synonyms: Plusiotis adelaida pavonacea Casey, 1915 is a syn. n. of Chrysina adelaida (Hope, 1841); Odontognathus gounellei Ohaus, 1908 is a revised synonym of Pelidnota ebenina (Blanchard, 1842); Pelidnota francoisgenieri Moore & Jameson, 2013 is a syn. n. of Pelidnota punctata (Linnaeus, 1758); Pelidnota genieri Soula, 2009 is a syn. n. of Pelidnota punctata (Linnaeus, 1758); Pelidnota lutea (Olivier, 1758) is a revised synonym of Pelidnota punctata (Linnaeus, 1758); Pelidnota (Pelidnota) texensis Casey, 1915 is a revised synonym of Pelidnota punctata (Linnaeus, 1758); Pelidnota (Strigidia) zikani (Ohaus, 1922) is a revised synonym of Pelidnota tibialis tibialis Burmeister, 1844; Pelidnota ludovici Ohaus, 1905 is a syn. n. of Pelidnota burmeisteri tricolor Nonfried, 1894; Rutela fulvipennis Germar, 1824 is syn. n. of Pelidnota cuprea (Germar, 1824); Pelidnota pulchella blanda Burmeister, 1844 is a syn. n. of Pelidnota pulchella pulchella (Kirby, 1819); Pelidnota pulchella scapularis Burmeister, 1844 is a syn. n. of Pelidnota pulchella pulchella (Kirby, 1819); Pelidnota xanthogramma Perty, 1830 is a syn. n. of Pelidnota pulchella pulchella (Kirby, 1819).

New or revised statuses: Pelidnota fabricelavalettei Soula, 2009, revised status, is considered a species; Pelidnota rioensis Soula, 2009, stat. n., is considered a species; Pelidnota semiaurata semiaurata Burmeister, 1844, stat. rev., is considered a subspecies.

New or comb. rev. and revised status: Plusiotis guaymi Curoe, 2001 is formally transferred to the genus Chrysina (C. guaymi (Curoe, 2001), comb. n.); Plusiotis transvolcanica Morón & Nogueira, 2016 is transferred to the genus Chrysina (C. transvolcanica (Morón & Nogueira, 2016), comb. n.). Heteropelidnota kuhnti Ohaus, 1912 is transferred to the genus Pelidnota (P. kuhnti (Ohaus, 1912), comb. n.); Odontognathus riedeli Ohaus, 1905 is considered a subspecies of Pelidnota rubripennis Burmeister, 1844 (Pelidnota rubripennis riedeli (Ohaus, 1905), revised status and comb. rev.); Pelidnota (Strigidia) acutipennis (F. Bates, 1904) is transferred to the genus Sorocha (Sorocha acutipennis (F. Bates, 1904), comb. rev.); Pelidnota (Odontognathus) nadiae Martínez, 1978 is transferred to the genus Sorocha (Sorocha nadiae (Martínez, 1978), comb. rev.); Pelidnota (Ganonota) plicipennis Ohaus, 1934 is transferred to the genus Sorocha (Sorocha plicipennis (Ohaus, 1934), comb. rev.); Pelidnota similis Ohaus, 1908 is transferred to the genus Sorocha (Sorocha similis (Ohaus, 1908), comb. rev.); Pelidnota (Ganonota) yungana Ohaus, 1934 is transferred to Sorocha (Sorocha yungana (Ohaus, 1934), comb. rev.); Pelidnota malyi Soula, 2010: 58, revised status; Xenopelidnota anomala porioni Chalumeau, 1985, revised subspecies status.

To stabilize the classification of the group, a neotype is designated for the following species: Pelidnota thiliezi Soula, 2009. Lectotypes are designated for the following names (given in their original combinations): Pelidnota brevicollis Casey, 1915, Pelidnota brevis Casey, 1915, Pelidnota debiliceps Casey, 1915, Pelidnota hudsonica Casey, 1915, Pelidnota oblonga Casey, 1915, Pelidnota pallidipes Casey, 1915, Pelidnota ponderella Casey, 1915, Pelidnota strenua Casey, 1915, Pelidnota tarsalis Casey, 1915, Pelidnota texensis Casey, 1915, and Scarabaeus punctatus Linnaeus, 1758.

The following published infrasubspecific names are unavailable per ICZN Article 45.6.1: Pelidnota (Odontognathus) cuprea var. coerulea Ohaus, 1913; Pelidnota (Odontognathus) cuprea var. rufoviolacea Ohaus, 1913; Pelidnota (Odontognathus) cuprea var. nigrocoerulea Ohaus, 1913; Pelidnota pulchella var. fulvopunctata Ohaus, 1913; Pelidnota pulchella var. sellata Ohaus, 1913; Pelidnota pulchella var. reducta Ohaus, 1913; Pelidnota unicolor var. infuscata Ohaus, 1913.

The following published species name is unavailable per ICZN Article 11.5: Neopatatra synonyma Moore & Jameson, 2013.

The following published species name is unavailable per application of ICZN Article 16.1: Parhoplognathus rubripennis Soula, 2008.

The following published species name is unavailable per application of ICZN Article 16.4.1: Strigidia testaceovirens argentinica Soula, 2006, Pelidnota (Strigidia) testaceovirens argentinica (Soula, 2006), and Pelidnota testaceovirens argentinica (Soula, 2006).

The following published species names are unavailable per application of ICZN Article 16.4.2: Homonyx digennaroi Soula, 2010; Homonyx lecourti Soula, 2010; Homonyx mulliei Soula, 2010; Homonyx simoensi Soula, 2010; Homonyx wagneri Soula, 2010; Homonyx zovii Demez & Soula, 2011; Pelidnota arnaudi Soula, 2009; Pelidnota brusteli Soula, 2010; Pelidnota chalcothorax septentrionalis Soula, 2009; Pelidnota degallieri Soula, 2010; Pelidnota lavalettei Soula, 2008; Pelidnota lavalettei Soula, 2009; Pelidnota dieteri Soula, 2011; Strigidia gracilis decaensi Soula, 2008, Pelidnota (Strigidia) gracilis decaensi (Soula, 2008), and Pelidnota gracilis decaensi (Soula, 2008); Pelidnota halleri Demez & Soula, 2011; Pelidnota injantepalominoi Demez & Soula, 2011; Pelidnota kucerai Soula, 2009; Pelidnota malyi Soula, 2010: 36-37; Pelidnota mezai Soula, 2009; Pelidnota polita darienensis Soula, 2009; Pelidnota polita orozcoi Soula, 2009; Pelidnota polita pittieri Soula, 2009; Pelidnota punctulata decolombia Soula, 2009; Pelidnota punctulata venezolana Soula, 2009; Pelidnota raingeardi Soula, 2009; Pelidnota schneideri Soula, 2010; Pelidnota simoensi Soula, 2009; Pelidnota unicolor subandina Soula, 2009; Sorocha carloti Demez & Soula, 2011; Sorocha castroi Soula, 2008; Sorocha fravali Soula, 2011; Sorocha jeanmaurettei Demez & Soula, 2011; Sorocha yelamosi Soula, 2011; Xenopelidnota bolivari Soula, 2009; Xenopelidnota pittieri pittieri Soula, 2009.

Due to unavailability of the name Pseudogeniates cordobaensis Soula 2009, we describe the species as intentionally new (Pseudogeniates cordobaensis Moore, Jameson, Garner, Audibert, Smith, and Seidel, sp. n.).

Keywords

leaf chafers, jewel beetles, New World, taxonomy

Introduction

The pelidnotine leaf chafers (Rutelinae: Rutelini) include the brilliantly metallic jewel scarabs (Chrysina spp.; e.g., Fig. 10), large, showy species that are used in ornamentation and jewelry (e.g., Chrysophora chrysochlora [Latreille]; Fig. 13), and species that exhibit dramatic sexual dimorphism (e.g., the bulging and dilated hind legs of male Pelidnota burmeisteri burmeisteri Burmeister; Fig. 56). The intensely lustrous, metallic colors of Chrysina Kirby species have been studied for their rare, cuticular reflection of circularly polarized light (Sharma et al. 2009, Pye 2010). Further studies have demonstrated that this may reduce predation by allowing for communication between conspecifics while remaining cryptic to avoid detection by predators (Brady and Cummings 2010). Ecologically, the leaf chafers have been proposed as valuable bioindicators of high-quality forest (Morón et al. 1997). The group is named for their leaf herbivory tendencies as adults, yet some species may also serve as pollinators of flowering plants (e.g., Pelidnota sumptuosa Vigors visits the flowers of Rourea induta Planch. and Stryphnodendron adstringens (Mart.) Coville in Brazil) (Gottsberger and Silberbauer-Gottsberger 2006). The extant pelidnotine leaf chafers are entirely distributed in the New World and include endemic genera such Pseudogeniates Ohaus (endemic to Argentina) and Homothermon Ohaus (endemic to Paulista center of endemism in Brazil), as well as widespread genera such as Pelidnota MacLeay (distributed from Canada to central Argentina).

Pelidnotine leaf chafers are a poorly studied group with a great need for systematics research. The lack of a taxonomic and phylogenetic framework remains an impediment to the circumscription of natural, monophyletic groups within the Rutelini. Lacking this essential foundation, we cannot understand the evolution of characters such as circular polarization of light in the cuticle of these beetles, the broad context of ecological services such as pollination that the species may provide, and we cannot reconstruct biogeographic patterns nor predict future distributional changes of genera and species of Rutelini.

The objective of this paper is to provide a foundation for understanding the taxonomy of 27 (26 extant and 1 extinct) genera and over 400 species of pelidnotine beetles, assist in stabilizing the classification and nomenclature of the genera, enable identification of genera, and provide a foundation for continued biodiversity research on leaf chafer scarabs. This work synthesizes the taxonomic and biodiversity literature for the pelidnotine scarabs, also encapsulating work that assisted in clarifying the nomenclature for the group (Moore and Jameson 2013, Moore et al. 2014). For the purposes of this research, we refer to this paraphyletic assemblage of taxa as the “pelidnotine scarabs,” and it is our aim that this work will set the stage for future research that addresses broad trends and patterns within the ruteline scarabs.

Legacy and history. Fredrick Bates (1829–1903, collection at BMNH), the younger brother of the well-known tropical biologist and coleopterist Henry Walter Bates (see O’Hara 1995), also had a love for taxonomy and entomology. He conducted research on the Heteromera (Tenebrionoidea) and the pelidnotine scarab beetles. Fredrick Bates died in 1903, and his single work on scarabs was published posthumously with the aid of Gilbert Arrow (The Natural History Museum). In the introduction, Arrow stated: “this revision of a difficult group of beetles represents many months of constant and strenuous investigation, continued to within a very few days of my friend’s death” (Arrow in Bates 1904: 249). Bates had intended the work to be more comprehensive, but his health did not allow further research. Upon Bates’s request, Arrow finished the work. Arrow stated that he confined himself to editorial functions with the exceptions of a few additions and modifications. Some of these modifications are clear within the text as evidenced by brackets and Arrow’s initials. For example, the commentary and diagnosis for the genus Mecopelidnota F. Bates and M. arrowi F. Bates were clearly added by Arrow.

Friedrich Ohaus (1864–1946, collection at ZMHB) was a student of coleopterist Edgar von Harold and a practicing medical doctor, which allowed him to travel to South America as a ship’s doctor. Ohaus provided the most comprehensive body of literature on world Rutelinae, and he developed the classification of the subfamily Rutelinae that is still used today. His work synthesized the body of knowledge on this highly biodiverse group, providing catalogs of species and their distributions, keys to higher-level groups, natural history, illustrations, and interpretation of characters. Ohaus’s classification of subtribes and genera is largely artificial, but this was a reflection of the state of systematics at the time. The Genera Insectorum on the Rutelini (Ohaus 1934b) was delayed for more than 20 years before publication. Instead of waiting for this larger, comprehensive catalog to be completed, Ohaus (1915b) published his concepts on some subtribal taxa within Rutelini and included descriptions of genera. In this work, he formalized the use of the subtribe Pelidnotina (as “Pelidnotinorum”) (Ohaus 1915b).

Johann W. Machatschke (1912–1975, collection at NHMB) continued Ohaus’s work, completing the Genera Insectorum volumes on Orthochilous Rutelinae (Anoplognathini, Adoretini, Anatistini) as well as Anomalini. He was the curator of the Coleoptera at Deutsche Entomologische Institut (DEI) in Berlin and later at the Museum G. Frey in Tutzing near Munich.

Marc Soula (1947–2012, collection at CCECL) was a mathematics teacher and naturalist who lived in Massat, France. He traveled broadly to South America and Thailand where he collected Rutelinae. At the Muséum National d’Histoire Naturelle in Paris, he began his life-long work on Rutelinae. Soula, in the fashion of Sciences Nat volumes, created guides to assorted rutelines, particularly the larger and showier groups. These volumes, which were published in parts, provide a preliminary effort to understand the diversity in the group. The benefit of these guides is that they provide color images of most species (dorsal habitus and often male genitalia) and species names that, in most cases, were verified by type examination. However, unlike monographic revisions, these guides suffer many shortfalls. They are based on a very limited number of specimens (often holotype specimens only), lack generalized distributional data and, prior to his death in 2012, most holotypes of species named by Soula were unavailable for general study because they were deposited in his personal collection. Soula’s works were written in an unusual style for scientific work. In effect, they were a “stream of consciousness” and lacked synthesis, analysis, and did not make meaningful interpretations or comparisons of characters. Rather than synthesizing his body of work, he published his work in disjunct parts. Sprinkled throughout the volumes (sometimes in red font, sometimes in bold font) he provided corrections to previous volumes such as amended taxonomic decisions, new combinations, new synonymies, and new distributions. These notes are very difficult to track and contributed to Soula’s numerous errors (synonyms, homonyms, lapsus calami, unavailable names, and transcriptional errors) (Moore and Jameson 2013, Moore et al. 2014). Soula’s guides provided an outlet for description of many new genera, species, and subspecies, but lacked unified species- or generic-level concepts. Additionally, Soula’s ruteline volumes were not peer-reviewed, were not widely available, and were expensive (thus reducing access). Because the volumes were not peer-reviewed, the data in them were not subjected to the objective scrutiny of other experts on phytophagous scarabs or agreed upon through scientific consensus. The volumes were not well edited, and they suffer from many misspellings (e.g., localities and scientific names), language that is not concise, and omissions (e.g., in the index and catalog). In addition, the lack of peer review and proper scientific editing for Soula’s volumes left numerous published names unavailable when the zoological rules of nomenclature (ICZN 1999) are applied due to various shortcomings and rule violations in the descriptions. Soula’s large collection of Rutelinae now resides in the Musée des Confluences, Lyon, France where it is databased, curated, and accessible for biodiversity research.

Higher-level nomenclature. Many of Soula’s descriptions of new genera within the Rutelini lack information regarding higher-level classification (e.g., Patatra Soula, Pachacama Soula, and Homeochlorota Soula were not clearly assigned to a subtribe of Rutelini at the time of their description). Because his work was published in parts, they included a mix of many genera from formerly accepted subtribes (Pelidnotina, Anticheirina) or accepted subtribes (Areodina, Lasiocalina), and they were not organized in a systematic manner. Thus, Soula’s tribal and subtribal classification within the Rutelinae was not clear. Soula recognized that his classification was not based on monophyletic groups (“La plupart des taxons supragénériques, n’étant pas monophylétiques...” [Soula 2011: 3]), but he maintained this classification pending further phylogenetic research. At the same time, however, he abandoned the subtribe Lasiocalina (Soula 2006) based on “dissimilarity” of the two genera included by Ohaus (1934b), but he failed to reclassify taxa in the group. Later, Soula (2011) revalidated the subtribe Lasiocalina without discussion. Soula’s classification (Soula 2011) omitted Rutelinae tribes (Alvarengiini and Adoretini) and included subtribes that are no longer accepted (e.g., Anticheirina, Pelidnotina) (Smith 2006, Bouchard et al. 2011). Additionally, Soula’s (2011) classification contradicted information in previous publications including the classification of Minilasiocala Soula (=Microogenius Gutiérrez) in the lasiocaline scarabs versus the pelidnotine scarabs (Soula 2006) and the classification of Pseudochlorota Ohaus and Lasiocala Blanchard as both pelidnotines and lasiocalines (Soula 2011). Two genera that were formerly included in the subtribe Pelidnotina were omitted by Soula (2011): Oogenius Solier and Eremophygus Ohaus. Because Soula provided no characters or justification for his higher classification and existing phylogenetic evidence demonstrates that Pelidnotina and Anticheirina are not monophyletic groups (Jameson 1998), we follow the classification of Bouchard et al. (2011) which lists these subtribes in synonymy under Rutelini.

Nomenclature. Pelidnota was placed on the Official List of Generic Names in Zoology (ICZN 2003) and included in the subtribe Rutelina (tribe Rutelini) by Bouchard et al. (2011). Although the subtribe Rutelina was hypothesized to be paraphyletic (Jameson 1998), it is the name-bearer for higher-level taxa (Rutelinae, Rutelini). The name has nomenclatural priority over the names Chasmodiidae Burmeister, Chrysophoridae Burmeister, Macraspididae Burmeister, Pelidnotidae Burmeister, Antichirides Lacordaire, Plusiotina Bates, and Fruhstorferina Ohaus (Bouchard et al. 2011).

The type species of Pelidnota is Scarabaeus punctatus Linnaeus, 1758. To ensure nomenclatural stability, the name was conserved due to homonymy with Scarabaeus punctatus Villers, 1789 (the dynastine scarab Pentodon bidens punctatus [Villers]) (ICZN 1999, Krell et al. 2012, Moore and Jameson 2013).

The name “Pelidnota” (from which the subtribe Pelidnotina takes its name) is derived from the blackish markings (“pelidnos” or “pelios” = Greek for black; “nota”=Latin for markings) that are common on the elytra of North American Pelidnota species.

Life history and biology. Immature life stages are known for only a handful of the pelidnotine genera including Homonyx Guérin-Méneville (Morelli 1996), Chrysina (Ritcher 1966, Morón 1976, 1985, Morón and Deloya 1991), Chrysophora Dejean (Pardo-Locarno and Morón 2007), and Pelidnota (Ritcher 1945, 1966, Morón 1976, Morón and Deloya 2002, Rodriguez et al. 2012, Garcia et al. 2013). Based on life history studies, life cycles are one to two years in duration (Ritcher 1966, Morón 1976, Morelli 1996). Larvae are sapro-xylophagous (Morón 1991) and feed on dry, rotten wood (Pelidnota virescens Burmeister; Morón and Deloya 2002), hollow trunks and tree stumps (P. punctata (Linnaeus) [Hoffman 1926]; Epichalcoplethis velutipes Arrow [Chalumeau 1985]), organic matter in the soil, and rotten roots (Morón 1991). One species, P. filippiniae Soula, is a significant defoliator and high numbers could contribute to plantation damage (Lunz et al. 2011).

Human cultural uses. The beauty, large size, and ease of collecting of many pelidnotine leaf chafers have promoted the cultural use of many species. For example, in Ecuador and Peru, the Jivaro and Sequoia Indians use the brilliant, metallic green elytra, pronota, or entire bodies of Chrysophora chrysochlora (Latreille) to make necklaces and headdresses (Ratcliffe 2006, Ratcliffe et al. 2015). In Guatemala, local people developed a cottage industry for tourists creating pendants, bracelets, and bola ties using local species of Chrysina (Woodruff 2009). The Yanomami people of Venezuela and Brazil extract and eat the larvae of Pelidnota sp. (known as “Makoia”) from logs in their gardens (Paoletti et al. 2000, Paoletti and Dufour 2005). Many attractive pelidnotine chafers are used in natural, artistic displays, including those of designer and photographer Christopher Marley (Marley 2008).

Fossil pelidnotines. Fossil organisms provide important information on ancestral character states, habitats, ecosystems, and adaptations. The only known leaf chafer fossil sets the minimum age of the subfamily Rutelinae at 50-42 mya (Krell 2006). The pelidnotine-like Pelidnotites atavus Cockerell is from the Eocene of England in the Bartonian Bagshot Beds of Bournemouth (Cockerell 1920, Carpenter 1992). Cockerell (1920) described the fossil belonging to the Rutelini and “in the vicinity of Pelidnota and Cotalpa” (Cockerell 1920: 463). This fossil should be examined to place the taxon within the Rutelini and to hypothesize sister-group relationships. No pelidnotine relatives are currently distributed in England or the Old World. Thus, this fossil revealed distributional patterns quite different from the current range of the pelidnotine Rutelini.

Identification of pelidnotine scarabs. Keys to the genera of “Pelidnotina” were created by F. Bates (1904) and Ohaus (1934b), and these provided a weak foundation for future work in the group. Bates’s (1904) posthumous work was based almost exclusively upon specimens available to him at the Natural History Museum (BMNH). Because this collection did not contain all described taxa in Pelidnotina, the revision and key were incomplete. Keys to genera and species groups that were provided by Ohaus (1934b) are not adequate for reliable identification of pelidnotine scarabs. Many of Ohaus’s (1934b) couplets are, in our estimation unclear, asymmetrical, and based on characters that vary widely across species and genera of Pelidnotina. Soula’s (2006) key to pelidnotine genera was based on Ohaus’s keys. Soula’s (2006) key omitted five genera that were newly described or elevated to generic status by Soula (Chipita Soula, Epichalcoplethis Burmeister, Pachacama, Patatra, and Sorocha Soula) and included genera that previously had been transferred out of Rutelini or synonymized (Pelidnotopsis Ohaus, Peltonotus Burmeister, and Plusiotis Burmeister) (see Moore and Jameson 2013). To date, there is no reliable, comprehensive key that facilitates accurate and repeatable identification of pelidnotine genera by non-experts. We think that consistent generic- and species-level identification of pelidnotine scarabs is not possible at this time. This lack of basic information is a great impediment to biodiversity and ecological research on pelidnotines

.

Classification and phylogeny. The leaf chafers (Rutelini) are members of the phytophagous scarab beetle clade (Melolonthinae, Cetoniinae, Dynastinae, Rutelinae, and a few minor subfamilies), a group that has been widely accepted as monophyletic for about 150 years and corroborated by molecular and morphological phylogenetic studies (Smith et al. 2006, Ahrens et al. 2011, McKenna et al. 2014). At the tribal- and subtribal-level morphological phylogenetic analyses demonstrated the inadequacies of the Rutelini classification sensu Machatschke (1972) (Jameson 1998). Based on this research, several subtribes (e.g., Pelidnotina, Anticheirina, and Rutelina) in the Rutelini were not monophyletic and, in fact, they were uninformative and logically inconsistent (Jameson 1998). Paraphyly of the ruteline subtribes has been a matter of discussion for well over a century. In his work on the pelidnotine scarabs, F. Bates (1904) noted the “close relationship” of Pelidnota and Rutela Latreille, and this idea was corroborated based on phylogenetic analyses of Rutelinae (Jameson 1998, Ahrens et al. 2011). Jameson (1998) provided the basis for elimination of the subtribe Pelidnotina and concluded that many genera were poorly characterized and not based on synapomorphic or autapomorphic characters.

Relationships among the pelidnotine scarabs need to be addressed by phylogenetic analyses. Pelidnotine genera, especially Pelidnota and the generic-level synonyms of Pelidnota (e.g., Strigidia Burmeister), should be re-structured into monophyletic groups with clear hypothesized synapomorphies. Based on phylogenetic analyses (Jameson 1998), Pelidnota and several related genera form a grade of taxa that are currently treated as genera or subgenera. Because of the poor understanding of the group and the lack of synapomorphies delimiting genera, past workers elevated distinctive species to the rank of genus, thus creating many monotypic genera (e.g., Pelidnotopsis [=Chrysina] and Ectinoplectron Ohaus). Hardy (1975) did not discuss relationships among the subgenera of Pelidnota, although he noted that the classification and subgeneric concept as proposed by Ohaus were in need of study. In our estimation, several of the current pelidnotine genera are probably not valid and Pelidnota, in particular, is likely to include several distinct, monophyletic groups (i.e., at the generic-level).

Research on specific pelidnotine genera has led to classification changes that affected the composition of pelidnotine scarabs. For example, Plusiotis and Chrysina were historically separate genera. As new species were described, our understanding of characters that circumscribe these groups was broadened and, as stated by Morón and Howden (1992: 208), the “characters that have been used to separate Plusiotis and Chrysina form a non-concordant mosaic.” Based on molecular and morphological data, Hawks (2001) synonymized Plusiotis as well as Pelidnotopsis within the genus Chrysina. Additionally, revisionary research on the genus Peltonotus, which was considered a member of the pelidnotine scarabs, provided sound evidence that the genus Peltonotus is closely related to members of the subfamily Dynastinae rather than the Rutelinae (Jameson and Wada 2004, 2009, Smith et al. 2006, Jameson and Drumont 2013). In spite of phylogenetic evidence that the subtribe Pelidnotina was paraphyletic, Soula (2006, 2008, 2009) maintained the subtribe, maintained the generic composition of the subtribe by including genera that had been transferred or synonymized, and he refrained from re-characterizing the group in any way.

We reiterate that Pelidnotina, as historically defined, is a paraphyletic grouping of disparate genera and species, and it should not be considered a valid taxon. We consider the subtribe a synonym of the subtribe Rutelina (Jameson 1998, Bouchard et al. 2011). We refer to “pelidnotine scarabs” in order to: 1) synthesize information on a group of genera that was chaotically treated by Soula, 2) incorporate genera new to the subfamily Rutelinae and previously of uncertain tribal placement (Peruquime Mondaca and Valencia and Neogutierrezia Martínez), 3) provide a mechanism for generic identification (in the form of a provisional dichotomous key), and 4) set the stage for future research that addresses broader trends within the Rutelini scarabs.

Materials and methods

Specimens and taxonomic material. Specimens examined for this study were provided by many institutions and private collections. We include information on type specimens to provide a foundation for continued research in the leaf chafers. Type specimens are international standards for scientific names (Knapp et al. 2004) and are tied to species hypotheses. The type specimen provides the nomenclatural stability that assures that the name reflects the described species and is linked through history in the literature. Acronyms for loaning institutions follow Evenhuis (2016).

BIOG Biodiversity Institute of Ontario, University of Guelph, Guelph, Ontario, Canada (Renee Labbee)

BMNH The Natural History Museum, London, United Kingdom (Max Barclay, Beulah Garner)

CAS California Academy of Sciences, San Francisco, California, USA (Norman Penny)

CCECL Musée des Confluences, Lyon, France (Cédric Audibert)

CMNC Canadian Museum of Nature Collection, Ottawa, Canada (Robert Anderson, François Génier)

CNC Canadian National Collection of Insects, Arachnids, and Nematodes, Ottawa, Canada (Pat Bouchard)

CNIN Colección Nacional de Insectos, Instituto de Biología, Universidad Nacional Autónoma de México (UNAM), México D. F., México (Harry Alperowitz, Santiago Zaragoza Caballero)

DBPC Denis Bouchard Personal Collection, Autouillet, France

DEPC David Ebrard Personal Collection, Velars sur Ouche, France

DJCC Daniel Curoe Personal Collection, Palo Alto, California, USA

EAPZ Escuela Agrícola Panamericana, Tegucigalpa, Honduras (Ron Cave, Jesús Orozco)

EMEC Essig Museum of Entomology, University of California, Berkeley, California, USA (Cheryl Barr, Peter Oboyski)

FMNH Field Museum of Natural History, Chicago, Illinois, USA (Alfred Newton, Crystal Maier)

FSCA Florida State Collection of Arthropods, Gainesville, Florida, USA (Paul Skelley)

HNHM Hungarian Natural History Museum, Budapest, Hungary (Ottó Merkl)

IAZA Instituto Argentino de Investigaciones de las Zonas Áridas, Mendoza, Argentina (Adriana Marvaldi)

IEXA Colección Entomológica, Instituto de Ecología, A.C., Xalapa, México (Miguel Ángel Morón)

IFML Instituto Fundación Miguel Lillo, Tucumán, Argentina (Dominga Carolina Berta)

IRSNB Institute Royal des Sciences Naturelles de Belgique, Brussels (Alain Drumont)

INBC Instituto Nacional de Biodiversidad, San José, Costa Rica (Ángel Solís)

INPA National Institute for Amazonian Research, Manaus, Brazil (Marcio Luiz de Oliveira)

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] (Girard Chovet, Fortuné Chalumeau)

JEMC José Mondaca E. Personal Collection, Peñaflor, Chile

JPBC Jean-Pierre Beraud Personal Collection, Cuernavaca, Morelos, México

LACM Los Angeles County Museum of Natural History, Los Angeles, California, USA (Brian Brown, Weiping Xie)

MACN Museo Argentino de Ciencias Naturales, Buenos Aires, Argentina (Arturo Roig)

MCZ Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA (Brian Farrell, Naomi Pierce)

MHNN Muséum d’Histoire Naturelle, Geneva, Switzerland (Peter Schwendinger)

MHNP Museo de Historia Natural, Universidad Nacional de San Antonio Abad, Cusco, Perú (Percy Yangue Yucra)

MIUP Museo de Invertebrados “G.B. Fairchild”, Universidad de Panamá, Panamá (Diomedes Quintero Arias)

MIZA Museo del Instituto de Zoología Agrícola, Maracay, Venezuela (José Clavijo)

MLJC Mary Liz Jameson Personal Collection, Wichita, Kansas, USA

MLPA Museo de la Plata, Universidad Nacional de la Plata, La Plata, Argentina (Analía Lanteri, Nora Cabrera)

MLUH Zentralmagazin Naturwissenschaftlicher Sammlungen, Martin-Luther-Universität Halle-Wittenberg, Halle, Germany (Karla Scheider, Joachim Händel)

MNHN Muséum National d’Histoire Naturelle, Paris, France (Olivier Montreuil)

MNNC Coleccion Nacional de Insectos, Museo Nacional de Historia Natural, Santiago, Chile (Mario Elgueta)

MNCR Museo Nacional de Costa Rica, San José, Costa Rica (formerly INBC, Ángel Solís)

MSPC Matthias Seidel Personal Collection, Prague, Czech Republic

MTD Museum für Tierkunde, Dresden, Germany (Klaus-Dieter Klass, Olaf Jäger)

MXAL Miguel Ángel Morón Collection, Xalapa, México

NHMB Naturhistorisches Museum, Basel, Switzerland (Daniel H. Burckhardt)

NMPC Department of Entomology, National Museum (Natural History), Prague, Czech Republic (Jiří Hájek)

OUMNH University Museum of Natural History, Oxford, United Kingdom (Darren Mann, Amoret Spooner)

PAPC Patrick Arnaud Personal Collection, Saintry sur Seine, France

PMNH Peabody Museum of Natural History, Yale University, New Haven, Connecticut, USA (Leonard Munstermann)

PVGH Pedro Vidal Personal Collection, Santiago, Chile

SDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (Lothar Zerche, Konstantin Nadein)

STRI Smithsonian Tropical Research Institute, Balboa, Panama (Annette Aiello)

UAEH Universidad Autónoma del Estado Hidalgo, Pachuca, Hidalgo, México (Juan Marquez Luna)

UAG Escuela de Biología de la Universidad Autónoma de Guadalajara, México (Jose Luis Navarette)

UCCC Museo de Zoología, Universidad de Concepción, Concepción, Chile (Jorge Artigas)

UCRC Entomology Research Museum, Department of Entomology, University of California, Riverside, California, USA (Doug Yanega)

UFRJ Museu Nacional, São Cristóvão, Rio de Janeiro, Brazil (Miguel Monné, Marcela Monné)

UVGC Colección de Artrópodos, Universidad del Valle de Guatemala, Guatemala City, Guatemala (Jack Schuster, Enio Cano)

UNSM University of Nebraska State Museum, Lincoln, Nebraska, USA (Brett Ratcliffe, M. J. Paulsen)

USNM U.S. National Museum, Washington, D.C. (currently housed at the University of Nebraska State Museum for off-site enhancement) (Brett Ratcliffe, Floyd Shockley)

UUZM Zoological Institute, Uppsala University, Uppsala, Sweden (Hans Mejlon)

WBWC William B. Warner Personal Collection, Chandler, Arizona, USA

WSU Maurice T. James Entomological Collection, Washington State University, Pullman, Washington, USA (Richard Zack)

ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany (Manfred Uhlig, Joachim Willers, Johannes Frisch)

ZSMC Zoologische Staatssammlung des Bayerischen Staates, Munich, Germany (Martin Baehr)

Images and terminology . Digital images of type specimens were taken over a 10-year period and were captured using several imaging applications including the Leica Application Suite V3.8. Images were edited in Adobe Photoshop CS2 (background removed, contrast manipulated). Figure legends for type specimens provide the valid, accepted name (as in the catalog) and the original combination of the species. We provide images of type specimen labels, but we largely defer from designating specimens as lectotypes. In our view, this is incumbent upon future revisionary scientists who will observe the best practices of systematics (ICZN 2003) and properly assign type status to specimens based on thorough review of all literature. Morphological terminology follows Hardy (1975) and Jameson (1998). Characters and specimens were observed with 6.3–50.0x magnification and fiber-optic illumination.

Literature reviewed. In compiling this work, we reviewed all available literature including major catalogs (Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, 1974, Smith 2009, Krajcik 2008, 2012, 2013). Some nomenclatural decisions were made by Krajcik (2012), but these were not indicated with “new synonymy” or “new combination,” and the rationale for these changes was not provided. For example, Krajcik did not accept Soula’s concepts of Epichalcoplethis, Chalcoplethis Burmeister, and Sorocha. Instead, he synonymized all names under Pelidnota. Also, Krajcik’s acceptance of names was not homogenous. He included some of Soula’s species and subspecies, but not all of them. For example, from the same publication (Soula 2009), Krajcik included Pelidnota estebanabadiei Soula (2009: 34), but he did not include Pelidnota equatoriana Soula (2009: 32). Krajcik included all of the following names: Pelidnota bondili Soula (2006: 10), Pelidnota castroi Soula (2008), and P. belti boyacaensis Soula (2006: 73), but he did not include any of the following names: Pseudogeniates cordobaensis Soula (2009), Pelidnota brusteli Soula (2010a), P. ohausi piurensis Soula (2006: 22), and P. sanctidomini caliensis Soula (2006: 79). Lacking his rationale, we did not follow Krajcik’s classification for the pelidnotine chafers. We discuss Krajcik’s (2008, 2012, 2013) catalogs to synthesize all available information for pelidnotine scarabs. However, we are agnostic about Krajcik’s listing of generic-, specific-, and subspecific-level synonyms amongst pelidnotine scarab taxa. Krajcik’s catalogs, in our opinion, are useful for tracking data on the proliferation of names in the hyper-diverse Scarabaeoidea, but should not be used to inform classifications. In our generic overviews and annotated catalog, we were forced in most cases to follow Soula’s generic- and species-level classifications (inadequate and uninformative though they are) because they are valid until addressed in broader systematic and revisionary works.

Soula’s descriptions of new taxa were often vague about the number, sex, and associated label data of type specimens. To rectify this lack of basic information, we report the verbatim label data of every pelidnotine scarab type specimen deposited in the Soula collection now housed at CCECL and CMNC. Due to the number of taxa that Soula named and described, both collections are rich in Rutelini type specimens. Among the pelidnotine scarabs, Soula’s material contains over 80 primary types (holotypes and neotypes) and nearly 700 secondary types (allotypes, paratypes, and paralectotypes) that are now all curated at CCECL. Additionally, examination of the CCECL collection revealed Soula specimens with type labels that had not been validly described. These “manuscript names” and the associated specimen data are listed in the appropriate genera (see “Annotated Catalog” below) as in litteris, and they are unavailable names. In a few instances, it appears that Soula omitted paratype data from the original published description or added paratype specimens to a type series after publication of a species description. For example, he added 17 “paratypes” collected in 2011 to Pseudochlorota peruana lecourti (described by Soula in 2005). Soula clearly knew that this violated nomenclatural rules, because he stated: “Répétons que de nombreux “cotypes” de cette Collection ne sont pas de “bons” types car ils ont été désignés et étiquetés après la description originale” (Soula 2005). We report these specimens as “invalid types” or “probable paratypes.” Soula also mislabeled type specimens (holotypes and paratypes), and we noted and corrected these mistakes when possible by referencing Soula’s original species descriptions.

During our study of the Soula collection at CCECL, we were able to return some primary type material to the institutions that had loaned specimens to Marc Soula (Garner and Audibert 2015). This type material had remained in his collection following his death. Additionally, we were able to gather information about what happened to the Soula collection after his passing, but before it was properly and legally accessioned by CCECL. Before CCECL acquired the Soula collection, it was briefly in control of his family. Unfortunately, the scientific value of some specimens was not recognized initially by the family and it seems likely that some material from the Soula collection or loaned specimens (possibly primary types) of an unknown species or several species (described only as a Pelidnota-like species with large legs [possibly some species similar to Pelidnota burmeisteri Burmeister or Chrysina species]) were lost in a sale to an antique collector (pers. comm. from Patrick Arnaud, June 2014) (Garner and Audibert 2015). Future systematists will have to deal with the uncertainty surrounding these possibly lost type specimens. Fortunately, it seems to us that this was an isolated and limited incident. We stress here that this is a cautionary tale that highlights the importance of properly maintaining loan records, providing temporary institutional labels on loaned specimens, and tracking the fate of personal collections that contain type material, regardless of taxonomic group.

Annotated Catalog. We list the author and date of the description of the species and genera, type species of genera (indicated with an asterisk), subspecies and forms, and transfer of species to other genera. This catalog builds on the work of Ohaus (1918, 1934b) and Machatschke (1972, 1974) with additions by Hardy (1975), Soula (1998, 2002a, 2002b, 2003, 2006, 2008, 2009, 2010a, 2010b, 2011), Krajcik (2008, 2012, 2013), and other authors. References to original descriptions of all species and genera are provided. Entries for species in the catalog provide: 1) the valid species name, author, date, and abbreviated citation, 2) original spelling and combination (if applicable), misspellings, new combinations, and invalid names in chronological order, 3) synonyms and the reference in which the synonym was designated, 4) general distribution data including the country (in capital letters) and states/provinces/departments/communes when they are known. Distributions are based on the literature and on specimens that we examined.

Rules of zoological nomenclature. Numerous nomenclatural changes within the pelidnotine scarabs are necessary due to misspellings, invalid type designations, and unavailability of infrasubspecific names (Moore and Jameson 2013, Moore et al. 2014). We follow the International Code of Zoological Nomenclature (ICZN 1999) as a means of stabilizing the taxonomy and classification of the pelidnotine scarabs.

Infrasubspecific names such as varieties and forms were widely used by authors such as Friedrich Ohaus and occasionally Marc Soula. These names are used to indicate unique color variants. Many of these names were treated as forms (forma; infrasubspecific entities) in catalogs (Machatschke 1972, 1974) as well as in works by Soula. According to ICZN Article 45.6.4: “A name is subspecific if first published before 1961 and its author expressly used one of the terms “variety” or “form” (including use of the terms “var.”, “forma”, “v.” and “f.”), unless its author also expressly gave it infrasubspecific rank, or the content of the work unambiguously reveals that the name was proposed for an infrasubspecific entity, in which case it is infrasubspecific” (see Lingafelter and Nearns 2013). Thus, named entities need to be interpreted within the context of the publication to discern if a name was unambiguously infrasubspecific. That is, if the author described or discussed both subspecies and varieties within a work, then it is clear that varieties can be treated as infrasubspecific, thus making the name unavailable unless further action was taken to correct the names prior to 1985 (ICZN Art. 45.6.4.1).

Species for which Soula designated type material but for which specimens are missing and presumed lost resulted in our neotype designations. The International Code of Zoological Nomenclature (ICZN 1999) requires that a neotype “is validly designated when there is an exceptional need and only when that need is stated expressly” (75.3). To be validly designated, Article 75.3.7 (ICZN 1999) requires a statement regarding the accessibility of the specimen. Upon publication, the specimen must be “the property of a recognized scientific or educational institution, cited by name.” Thus, some neotypes were invalidly designated by Soula. Designation of some neotype specimens was necessary for names proposed by Soula.

The lack of synthesis and attention to detail in Soula’s works resulted in some names that were not validly described (see Moore and Jameson 2013, Moore et al. 2014). For all new species-group names described after 1999, the holotype and the type depository must be explicitly stated for the name to be deemed available (ICZN Art. 16.4). Because Soula (2006, 2008, 2009, 2010a, 2011) did not explicitly state the location of holotypes for several species, these names are unavailable.

For groups that have dramatic sexual dimorphism, some taxonomists refer to the “alloréférent” or the “neallotype” specimen for the first specimen of the opposite sex that is described in a publication subsequent to the original description (Dechambre 2001). Unlike name-bearing type specimens (e.g., holotype, lectotype, neotype), these specimens have no formal nomenclatural status (Hawksworth 2010). Soula frequently made use of the term “alloréférent” in his collection and his published works. We stress that these specimens are not name-bearing type specimens.

Poor editing and many misspellings compromise the scientific value of Soula’s works (e.g., scientific names, localities, descriptive characters, figure legends, indices, and identification keys). These errors pose problems because they can be propagated by future researchers. And, in some cases, the error confuses or obscures Soula’s intended species name. We include these misspellings to limit confusion and promote future research.

Type specimens and lectotype designation. For purposes of nomenclatural stability, we designate lectotypes for some species (ICZN Art. 74). In these cases, a specimen was selected among a group of syntype specimens or cotype specimens. During this research (initiated by MLJ in the late 1990s) many type series were studied, lectotype labels added, and specimens returned to museum collections. However, when research on the group became intractable due to Soula’s concurrent work on the group, these lectotypes were not published. Soula also designated lectotypes. In some cases, he removed previous lectotype labels and he changed the collection depository. In other cases, we have reason to reject Soula’s attempted lectotypification. For example, Soula designated paralectotypes without first designating a lectotype (see P. laevissima Burmeister in Soula 2009), he stated that the type series consists of only a holotype but he provided an image of a lectotype (see Homothermon praemorsus (Burmeister) in Soula 2008) and, for a species named by Sharp in 1877, he designated a holotype and paratype (rather than a lectotype and paralectotype) (see P. prolixa Sharp in Soula 2009). All of these cases gave us reason to be cautious of Soula’s lectotypification. Thus, in instances where we have verified label data, we refer to types specifically (lectotype, paralectotype, holotype). In other instances, however, we refer to type specimens as “types,” and we leave lectotype designation to future systematists. We provide images of many type specimens, but it is not our purpose to nomenclaturally fix or designate types with these images.

Concepts for genera and species. In this work, we do not generally assess the validity of species, subspecies, or genera. In our view, this is best conducted as part of comprehensive, revisionary studies. Instead, we provide a taxonomic and nomenclatural framework for future research. Although we do not name new species within this work, we adhere to the phylogenetic species concept (Wheeler and Platnick 2000) in our interpretations: “A species is the smallest aggregation of (sexual) populations or (asexual) lineages diagnosable by a unique combination of character states.” New nomenclatural acts in this work, such as new synonyms and new homonyms, are based on examination of type specimens and in accordance with the rules of zoological nomenclature (International Code of Zoological Nomenclature, 1999).

Hardy’s standards for species circumscription provide a solid basis for ruteline systematics. Hardy’s (1975) classic work on Pelidnota from North and Central America provided the foundation for our knowledge of pelidnotine species as well as a rigorous foundation for interpretation of intraspecific variation. Hardy’s species concepts within the genus Pelidnota have endured for over four decades. Hardy (1975) considered species to be “variable entities” and he allowed for intraspecific variation in coloration, maculae, the degree of posterior coxal corner production, and even the form of male genitalia. Hardy allowed intraspecific variation in parameres in P. punctata (Hardy 1975; Figs 3436) and P. costaricensis H. W. Bates (Hardy 1975; Figs 38, 39) just to name a few. Because all specimens of all populations of species cannot realistically be studied, actual distributions of characters must be theorized based on available specimens. Species-level lineages are hypothetically circumscribed and these hypotheses should be testable (i.e., subject to the consideration of additional data). Only by examining a large number of specimens and seeing continuity between populations was Hardy (1975) able to conclude that the observed variation was intraspecific rather than interspecific.

In our view, Hardy’s (1975) method of study provided a good test of historical species concepts in Pelidnota as additional specimens and populations were discovered and species boundaries could be critically evaluated (see Wheeler 2004). Critical evaluation of species boundaries is an important (but largely undiscussed) concept within pelidnotine leaf chafer taxonomy due to the elaborate male genital morphology (considered diagnostic for identification) present in many species. Pelidnotine scarab species, and Rutelini more generally, often have asymmetric parameres and ventral genital plates originating from the phallobase (e.g., in the genera Homothermon, Xenopelidnota F. Bates, and many Pelidnota). The cuticular generation of these asymmetric structures is certainly a complex and highly sensitive developmental process that we think gives rise to a great deal of intraspecific variation in male genital characters. Many historical Pelidnota workers, outside of Hardy (1975), have interpreted this type of genital shape variation to be interspecific and subsequently split species where we may have lumped them into broader, variable species.

In contrast to Hardy’s (1975) and our species concept, Soula’s species concept (1998, 2002a, b, 2003, 2006, 2008, 2009, 2010a, b, 2011) did not allow for intraspecific variation. In Soula’s works, slight differences in color, punctation, or form of male parameres equated to different species or subspecies. In fact, slight differences in male parameres could be attributed to minor deviation in the manner in which parameres were viewed. For example, Soula’s description of S. purpurea esperitosantensis was based on two male specimens from Espírito Santo, Brazil (Soula 2006). Soula’s line drawings of the male parameres are extremely similar to the nominate subspecies, and Soula remarked that parameres of both were “slightly different.” Both the nominate species and subspecies were known to Soula from fewer than five specimens in total, and only the nominate form of the species was included in Soula’s key (2006: 9-12).

In the pelidnotine scarabs, Soula (2006, 2008, 2009, 2010a, 2011) described over 150 new species and subspecies and ultimately classified approximately 100 new species and subspecies in Pelidnota. Quantifying the number of specimens that Soula’s pelidnotine species-group names were based upon illustrates the lack of intraspecific variation incorporated into his species concept. Forty-four percent (67 of 152) of Soula’s new species and subspecies of pelidnotine scarabs were described from two (minimum number for both sexes to be described) or one specimen. Approximately 33% (50 of 152) of Soula’s new pelidnotine names were based on descriptions of a single, male holotype specimen. In total, 41% (63 of 152) of Soula’s new pelidnotine names were based upon descriptions of only one sex. Soula’s species and subspecies concepts were almost singularly reliant on slight differences in male paramere morphology and/or broadly separated populations. These concepts, when paired with the limited number of specimens of some genera to which he had access, led to his inability to reliably diagnose species when either of these conditions was violated. For example, Soula stated that he could not diagnose the females of Sorocha Soula species when they are even narrowly sympatric. For example, for diagnosis of “S. yelamosi” Soula stated (2011: 82), “Là encore la femelle est à capturer et à repérer. Plusieurs espèces semblent cohabiter et il ne sera pas facile d’appareiller les couples”. In sum, adequate characters were not provided to circumscribe many of Soula’s species-level hypotheses. Soula’s guides provided the outlet for description of many new genera, species, and subspecies, but an adequate concept that guided his hypotheses was lacking. In our view, Soula’s species and subspecies concepts cast doubt on the validity of many of his taxa.

DNA barcode analysis for Pelidnota punctata. Cytochrome Oxidase 1 (CO1) DNA data were used to address genetic variation in Pelidnota punctata across the distributional range of the species. Using the Barcode of Life Database (BOLD: http://www.boldsystems.org), CO1 data were gathered for P. punctata (13 specimens) and 10 other species of Pelidnota (38 specimens). The distance model used the Kimura 2 parameter with a neighbor-joining tree building method in BOLD. Nodes are labeled by species name, BOLD ID number, and country and state/province where the specimen was collected (Fig. 4).

Overviews of genera. Biological and natural history data in the “Generic Overviews” were synthesized from the literature, specimens, and specimen labels from many institutions. Overviews do not summarize all literature and all specimens. Instead, they highlight: 1) potential complications such as paraphyly and nomenclatural issues, 2) potential synapomorphic characters and discuss possible sister-group relationships, 3) basic distribution and habitat affiliations, and 4) known larvae and natural history information.

Overview of the pelidnotine genera

Diagnosis. Pelidnotine scarabs are members of the tribe Rutelini (for a key to tribes of Rutelinae, see Jameson 1990, 2005). Characters that diagnose the Rutelini pelidnotine scarabs include: obvious membranous border on the elytra lacking (versus membranous border present at the elytral apex as in Anomalini); frontoclypeal suture obsolete at middle (versus complete as in the ruteline subtribes Heterosternina and Areodina); labrum that is horizontally produced with respect to the clypeus (versus vertically produced as in Geniatini, Anoplognathini, Anatistini, and Adoretini) and sinuate at apex; protarsomeres that are subcylindrical and lack ventral setose pads (versus dorsoventrally flattened and densely setose ventrally as in Geniatini); protibial spur apical (versus subapical as in Anomalini); and terminal spiracle positioned in pleural suture (versus not positioned in pleural suture as in Anomalini).

Males and females are generally separated based on the inner protarsal claw that is wider than the outer claw and may or may not possess a small, inner tubercle. Protarsal claws of the females, in comparison, are more similar in width and lack a small, inner tubercle.

Identification key. We provide a provisional key to the pelidnotine scarabs that should be used with caution. First, the subtribe Pelidnotina is paraphyletic and users should not be misled into thinking that the key circumscribes a natural group. Second, some genera are also very likely paraphyletic, thus causing complications for circumscription and identification. Third, the key will not be useful for both males and females for some genera (e.g., Mesomerodon Ohaus, Hoplopelidnota F. Bates) due to use of sexually dimorphic characters. Fourth, owing to likely paraphyly, the genera Microogenius and Eremophygus could not be separated in the key. Fifth, two genera are keyed twice (Chalcoplethis, Epichalcoplethis). These complications in key construction are indicators of the complexity of the group and need for further systematics studies.

Key to the genera of pelidnotine scarabs (Coleoptera, Scarabaeidae, Rutelinae, Rutelini)

Males: inner protarsal claw wider than the outer claw; may or may not possess a small, inner tubercle; sternites usually concave. Females: protarsal claws similar in width; lack a small, inner tubercle; sternites usually convex.

1 Claws on all legs simple and of similar size; protarsal claw (male) lacking apical or subapical tubercle, lacking apical split Neogutierrezia Martínez
Claws on all legs with the inner claw different than the outer claw (wider or split apically); protarsal claw (male) wider than outer claw, with or without small, inner tubercle, and with or without apical incision 2
2 Labrum and clypeus fused anteriorly Peruquime Mondaca and Valencia
Labrum and clypeus not fused anteriorly, free 3
3 Lateral edge of mandible lobe-like and flattened, without reflexed teeth (e.g., Fig. 1A) 4
Lateral edge of mandible not flattened, with 1 or 2 reflexed teeth (Fig. 1B, C, E, F) 8
4 Apex of labrum extends beyond clypeal apex, visible from dorsal view 5
Apex of labrum does not extend beyond clypeal apex, not visible from dorsal view 6
5 Metatarsomere 4 at apex with 4-6 spinules, medial spinules thickened Oogenius Solier
Metatarsomere 4 at apex with 4-6 spinules, medial spinules seta-like (not thickened) Microogenius Gutiérrez and Eremophygus Ohaus
6 Clypeus with apex quadrate or subquadrate (Fig. 1D); apex emarginated 7
Clypeus with apex rounded, parabolic, or trapezoidal; apex with or without emargination (Fig. 1A, B, C, E, F) 8
7 Lateral edge of protibia with 2 teeth Chipita Soula
Lateral edge of protibia with 3 teeth Parhoplognathus Ohaus
8 Apex of elytra in males with acute, spiniform projections (Fig. 3C, D) 9
Apex of elytra in males rounded 10
9 Males without acute process on posterior margin of mesofemur. Females with 2 deep emarginations near the apex of the terminal sternite; pygidial disc with a concavity. Dorsal color metallic green Hoplopelidnota F. Bates
Males with acute process on posterior margin of mesofemur (Fig. 2E). Females lacking emarginations at apex of terminal sternite, instead apex is rounded; pygidial disc convex. Dorsal color testaceous or light-brown (with or without weak metallic-green reflections) Mesomerodon Ohaus
10 Pronotum with apical bead obsolete or lacking medially (Fig. 1A) Chrysina Kirby
Pronotum with apical bead complete medially (Fig. 1B, C, D, F) 11
11 Males with metatibia enlarged, curved, produced posteriorly at apex Chrysophora Dejean
Males without metatibia enlarged, curved, produced posteriorly at apex 12
12 Metatarsomeres 1–5 longer than metatibia Chalcoplethis Burmeister
Metatarsomeres 1–5 subequal to metatibia 13
13 Metatibia somewhat laterally flattened (Fig. 2A) Epichalcoplethis F. Bates
Metatibia not laterally flattened (Fig. 2B) 14
14 Prosternal projection (between procoxae) produced to level of procoxae 15
Prosternal projection (between procoxae) shorter, not produced to level of procoxae Xenopelidnota F. Bates
15 Base of metatibia with semicircular notch (Fig. 2C) Mecopelidnota F. Bates
Base of metatibia lacking semicircular notch, straight (Fig. 2D) 16
16 Apex of metatibia straight and with numerous spinules Ectinoplectron Ohaus
Apex of metatibia not straight (biemarginate or with external apex produced), with 0–8 spinules 17
17 Metatibia lacking produced external apex, lacking apical spinules 18
Metatibia with external apex produced posteriorly and with apical spinules 26
18 Disc of frons with weak V-shaped depression (Fig. 1E) Sorocha Soula
Disc of frons planar, smooth, lacking a V-shaped depression (Fig. 1A, B, C, D, F) 19
19 Metatibia laterally flattened 20
Metatibia not laterally flattened 21
20 Metatarsomeres 1-5 subequal to metatibia Epichalcoplethis F. Bates
Metatarsomeres 1-5 longer than metatibia Chalcoplethis Burmeister
21 Elytral shoulder rounded (not flat in ventral view), lacking bead Homothermon Ohaus
Elytral shoulder flat in ventral view, with bead 22
22 Mesometasternal keel surpassing mesocoxae (Fig. 3B) Pelidnota MacLeay
Mesometasternal keel not surpassing mesocoxae (Fig. 3A) 23
23 Lateral edge of mandible with two reflexed teeth (Fig. 1B, E, F) 24
Lateral edge of mandible with one reflexed tooth (Fig. 1C) 25
24 Metatarsomere 3 with apical setae (externally) of unequal length and width; color castaneous to black Homonyx Guérin-Méneville
Metatarsomere 3 with apical setae (externally) of equal length and width; color metallic green Catoclastus Solier
25 Fifth meso- and metatarsomeres without internal teeth, tarsomeres simple Pseudogeniates Ohaus
Fifth meso- and metatarsomeres with one or two internal teeth (may be rounded) Parhomonyx Ohaus
26 Protibia with 2 external teeth Pachacama Soula
Protibia with 3 external teeth 27
27 Labrum with apex bilobed Patatra Soula
Labrum with apex projecting anteriorly at middle, not bilobed Homeochlorota Soula
Figure 1.

Clypeal shape varying from rounded, parabolic, trapezoidal, subquadrate (A–D), and emarginate (E–F). Lateral edge of mandibles with no reflexed teeth (lacking teeth in A, but the mandible is reflexed rather than flattened in A, two reflexed teeth (B, E, F), or one reflexed tooth (C)). Apical bead of pronotum varying from obsolete (A) to complete medially (B, C, D, F). Disc of frons with V-shaped depression (E) or frons planar, smooth, lacking V-shaped depression (A–D, F) A Chrysina beyeri Skinner B Epichalcoplethis velutipes velutipes (Arrow) C Parhomonyx fuscoaeneus (Ohaus) D Chipita mexicana (Ohaus) E Sorocha sp. F Homonyx elongatus (Blanchard).

Figure 2.

Characters of the mesofemora and metatibiae in pelidnotine genera. A Epichalcoplethis aciculata (F. Bates), metatibia somewhat flattened (dorsal view) B Pelidnota virescens, metatibia not flattened (dorsal view) C Mecopelidnota sp., metatibia at base with a semicircular notch D Pseudogeniates cordobaensis Moore et al., metatibia simple at base E Mesomerodon gilletti Soula male, acute production of posterior margin of mesotibia (ventral view).

Figure 3.

Characters of the thorax (ventral view) and elytral apex (dorsal and lateral views). Mesosternal keel not surpassing base of mesocoxae (A) or keel surpassing base of mesocoxae (B). Acute, spiniform projections at apex of elytra (C, D). A Homonyx elongatus B Pelidnota dobleri Frey C Mesomerodon gilletti, dorsal view D Mesomerodon gilletti, lateral view.

Catoclastus Solier, 1851

Fig. 5

Type species

Catoclastus chevrolatii Solier, 1851.

Species

3 species; length 14–23 mm.

Three species are included in this genus and are distributed in western Peru. Species are elongate-oval, metallic green with dark red appendages, and similar in overall appearance to species of Mecopelidnota and Homonyx. Soula (2010a) apparently overlooked C. rabinovichi Martínez, a species that is known only from the male holotype from Cusco, Peru. Species in the genus are characterized by having all claws simple; male protarsal claw with inner tubercle; bidentate mandibles; pronotum with bead complete apically, laterally and basally; elytral base with dimple lateral of scutellum; elytral epipleuron shelf-like (not rounded); fifth meso- and metatarsomeres lacking internomedial tooth; apex of the metatibia with weak corbel and with four to five spinules apically (biemarginate in Homonyx); mesosternal keel not surpassing mesocoxae; and metasternum with longitudinal groove (not paired as in Hoplopelidnota). Sister-group relationships of the genus require analysis. Specimens have been collected from 2000 to 3500 m elevation in the months of April and June. Specimens are rare in collections, and larvae are not described.

Chalcoplethis Burmeister, 1844

Fig. 6

Type species

Chrysophora kirbii Gray, 1832.

Species

2 subspecies; length 22–27 mm.

As circumscribed by Soula (2006), the genus Chalcoplethis includes only C. kirbii kirbii Gray and C. kirbii misionesensis Soula. Whereas F. Bates (1904), Ohaus (1934b), and Hardy (1975) considered Pelidnota (Chalcoplethis) to include a broad group of Pelidnota species with metallic green, rugose elytra, Soula considered Chalcoplethis as unique and monotypic. Soula (2006) also considered Epichalcoplethis to be separate and distinct from Chalcoplethis. It is clear that species of Chalcoplethis and Epichalcoplethis share a number of characters (form of the male genitalia, pronotal bead obsolete apicomedially, lack of spinules at apex of metatibia, well-developed prosternal process, and mesometasternal keel surpassing the mesocoxae). Relationships of these three genera need to be studied and placed within the broader context of ruteline genera.

Chalcoplethis kirbii is diagnosed by its metallic green color, striate elytra, elytral epipleuron shelf-like (not rounded), pronotum with bead incomplete apically (complete laterally and basally), metatibia of the male that is strongly compressed (less so in females) and lacking apical spinules, meso- and metatarsomere 5 lacking an internomedial tooth; mandibles that are bidentate externally, prosternal process well developed, and mesometasternal keel surpassing the mesocoxae. The species is distributed in the Atlantic Coastal Forest of Brazil from Bahia in the north to Rio Grande do Sul in the south. Larvae are not described.

Chipita Soula, 2008

Fig. 7

Type species

Byrsopolis mexicana Ohaus, 1905.

Species

1 species; length 14–18 mm.

The monotypic genus Chipita was proposed by Soula (2008) for Chipita mexicana (formerly Parhoplognathus mexicanus), which is known from Sinaloa, Guerrero, Jalisco, Nayarit, and Oaxaca states in Mexico. Following Ohaus’s (1934b) classification of the genus Parhoplognathus, Soula (2008) created this monotypic genus. Several characters provide sufficient rationale for the genus: form of the thorax (broadest at base versus broadest at the middle in Parhoplognathus), elytra (striate versus not striate in Parhoplognathus), mesosternum not produced anteriorly (produced or not produced anteriorly in Parhoplognathus), and protibia with 2 external teeth (with 3 external teeth in Parhoplognathus). The taxon shares several similarities with species in the genus Platyrutela Bates (an anticheirine scarab), thus requiring examination within a phylogenetic framework.

Chipita mexicana is diagnosed by the following characters: profemur produced anteriorly and widest at middle (autapomorphic for the genus); protibia with 2 external teeth (shared with Platyrutela); mandibular palp with deep, horizontal sulcus (shared with Platyrutela); clypeus quadrate and apex reflexed (shared with Platyrutela); clypeus greatly declivous with respect to plane of frons; pronotum broadest at base (shared with Platyrutela); elytra striate; elytral epipleuron rounded; claws simple on all legs (male and female; shared with Platyrutela); male protarsal and mesotarsal claws with inner, apical tubercle; meso- and metatarsomere 5 with internomedial tooth; apex of metatibia with short spines (versus long setae in Platyrutela); color gray or castaneous with or without metallic green sheen.

Adult C. mexicana inhabit tropical deciduous and sub-deciduous forests at elevations between sea level and 200 m (Morón et al. 1997). Adults are temporally distributed between June and November, and are attracted to lights at night (Morón et al. 1997). The larvae of C. mexicana are undescribed and their biology is unknown. Male specimens of Chipita mexicana are rare in collections, and may be indicative of unusual natural history.

Chrysina Kirby, 1828

Figs 1A, 8, 9, 10, 11, 12

Type species

Chrysina peruviana Kirby, 1828.

Species

113 species; length 19–40 mm.

Species in the genus Chrysina are commonly known as the “jewel scarabs” for their spectacular metallic and iridescent coloration and large size. Species range from metallic green, pink, purple, gold, and silver, and their elytra may be adorned with metallic gold or silver pin stripes or polka dots. The males of some species have enlarged metafemora (e.g., Chrysina macropus [Francillon]). Morón (1990) reviewed the 73 species of Chrysina (then referred to as Plusiotis, Chrysina, and Pelidnotopsis). Since that time, an additional 40 species have been described, and no updated revision or monograph is available for the group. The following characters serve to diagnose species in the genus: clypeal apex rounded, with or without emargination; all claws simple; male protarsal claw with or without inner tubercle; mandibles rounded externally; pronotum with bead incomplete apically and basally (complete laterally) (Chrysophora with bead complete on all margins); elytral epipleuron shelf-like (not rounded); fifth meso- and metatarsomeres with internomedial tooth; metatarsi shorter than tibia (longer than tibia in Chrysophora and Chalcoplethis); apex of the metatibia with or without corbel; meso- and metatarsomere 5 with internomedial tooth; mesosternal keel surpassing mesocoxae.

The genera Plusiotis and Chrysina were historically separate genera. Morón and Howden (1992) noted an apparent grade of characters within the taxa. Based on molecular and morphological data, Hawks (2001) synonymized Plusiotis as well as Pelidnotopsis with Chrysina. Soula (2008) resurrected the genus Pelidnotopsis, asserting that the genus was “closer” to Pelidnota than to Chrysina. Moore and Jameson (2013) again synonymized Pelidnotopsis within Chrysina. In an effort to develop identification tools for species of conservation importance, Moron and Noguiera (2016) advocated for the use of both Plusiotis and Chrysina. Although they acknowledge that several species possess “transitional characters”, they argue that the evidence for synonymy of Plusiotis was based on unpublished data (Hawks 2001). Characters, they assert, clearly differentiate the two genera, but they do not provide a list of these characters nor a diagnosis for each genus. In our view, the transitional characters provide support for one clade, thus we advise the unity of these genera into the senior name, Chrysina. An analysis in preparation by Morón will elucidate the relationships of the genera (Morón and Noguiera 2016).

Species in the genus are distributed from the southwestern United States to Ecuador with the greatest diversity of species occurring between 1000–2000 m elevation (Morón 1991). Many species have narrow habitat requirements and are negatively impacted by unfaltering deforestation that serves to reduce and isolate populations, thus placing species at risk (Morón and Nogueira 2016). Species are found in primary forests (pine, juniper, and pine-oak) between 50-3800 m. Species feed on the foliage (adults) or rotting logs (larvae) of various trees including species in the genera Abies, Alnus, Arbutus, Heliocarpus, Juglans, Juniperus, Liquidambar, Pinus, Platanus, Quercus, and Turpinia (Morón 1991). Representative larvae have been described in the genus (Ritcher 1966, Morón 1976, 1985). Adults are frequently attracted to lights, and larvae live in rotten logs.

Chrysophora Dejean, 1821

Fig. 13

Type species

Melolontha chrysochlora Latreille, 1812.

Species

1 species; length 25–42 mm.

The dazzling, metallic green Chrysophora chrysochlora is a distinctive species and the only member of its genus. The large size, conspicuously rugose elytra, and elongate legs of the male are distinguishing characteristics. Additional characters include the metatibia of the male that is prolonged and acuminate at the apex, the 5th tarsomere with an internal tooth (all legs), the mandibles that are broadly rounded externally, the pronotum with a complete bead, and the mesosternum that is not appreciably produced beyond mesometasternal suture. Research is needed to examine sister-group relationships of this monotypic genus.

The species is distributed in Colombia, Ecuador, and Peru where the Jivaro, Shuar, and Sequoia Indians use the elytra, pronota, legs, or entire body for adornment (Ratcliffe 2006, Ratcliffe et al. 2015, Le Tirant and Limoges 2016). The species is primarily found in dry and humid tropical forest between 180-550 m elevation (Pardo-Locarno and Morón 2007), although the species is recorded between 500-1000 m elevation (Morón 1990). The species is associated with Buddleja L. (Scrophulariaceae), Gynerium sagittatum (Aubl.) Beauvois (Poaceae; arrow cane, wild cane) (both Ohaus 1934b), Senna reticulata (Willd.) H. S. Irwin and Barneby (Leguminosae), and Leucaena leucocephala (Lam.) de Wit (Leguminosae) (both Pardo-Locarno and Morón 2007). Adults feed on the leaves of G. sagittatum during the day (Ohaus 1934b) from February to May (Morón 1990), and they fly at twilight or at night (Ohaus 1934b). Larvae and pupae are described and share several characters with Pelidnota larvae and pupae (Pardo-Locarno and Morón 2007).

Ectinoplectron Ohaus, 1915

Fig. 14

Type species

Homonyx oryctoides Ohaus, 1905.

Species

1 species; length 21–23 mm.

This monotypic genus is endemic to northwestern Mexico. Adults have a rufous dorsal coloration without metallic reflections, and are similar to Pelidnota (Pelidnota) in overall appearance. Adults in the genus Ectinoplectron are diagnosed by the disc of the prosternal peg that is weakly concave with reflexed margins (an autapomorph). Additional diagnostic characters include: lateral edge of mandibles with two reflexed teeth; apex of metatibia straight (not biemarginate) and lacking spinules or setae; meso- and metatarsomere 5 lacking an internomedial tooth; mesosternum not appreciably produced beyond the mesometasternal suture; pronotum with bead complete apically, basally, and laterally; lateral edge of protibia with three rounded teeth; and, apex of clypeus subtrapezoidal to subtriangular.

Ectinoplectron oryctoides is known from Pacific coastal states of Mexico (Durango, Jalisco, Michoacán, Nayarit, Sinaloa and Sonora), northern Chihuahua (Lugo et al. 2011), and western Durango (Machatschke 1972, Hardy 1975, Morón 1990) where it occupies tropical deciduous forests of oak and pine (Morón et al. 1997). Temporal distribution is from late June to September (Morón et al. 1997). Individuals of E. oryctoides occur from sea level to 2000 m elevation (Hardy 1975, Morón et al. 1997), are attracted to lights at night, and tend to fly near dusk (Morón et al. 1997). Larvae of E. oryctoides are undescribed.

Epichalcoplethis F. Bates 1904

Figs 2A, 15, 16, 17, 18, 19

Type species

Pelidnota velutipes Arrow, 1900.

Species

16 species and subspecies; length 15–19 mm.

Previously considered a subgenus of Pelidnota, Epichalcoplethis was circumscribed by Soula (2006) as distinct from the monotypic genus Chalcoplethis and composed of 16 species and subspecies. Chalcoplethis kirbii shares many characters with species of Epichalcoplethis including form of the male genitalia, pronotal bead which is obsolete apicomedially, lack of spinules at apex of metatibia, well-developed prosternal process, and mesometasternal keel surpassing the mesocoxae. Sister-group relationships require examination.

Species in the genus Epichalcoplethis can be diagnosed, in part, based on the following characters: metatibia weakly compressed (strongly compressed in C. kirbii) and apex lacking spinules; meso- and metatarsomere 5 lacking internomedial tooth; punctate-striate elytra; elytral epipleuron shelf-like (not rounded); pronotum with bead incomplete apically (complete laterally and basally); mandibles that are bidentate externally; prosternal process well-developed; and, mesometasternal keel surpassing the mesocoxae. Epichalcoplethis chamaeleon (Herbst) differs from other species in the genus based on the form of the male parameres, form of the metatibia in the male (not compressed laterally and apex with a well-developed corbel). For many years, this large and conspicuous species was misidentified as Pelidnota rostrata Burmeister.

Species in the genus are distributed from Guatemala and Belize, St. Vincent and the Grenadines, Trinidad and Tobago, and south to Argentina, Uruguay, and Paraguay. In Grenada, E. velutipes is common in the temperate zone from April to May (Chalumeau 1985). Although the larvae are not described for this species, Chalumeau (1985) noted that he obtained larvae from the decaying trunks of mango trees.

Eremophygus Ohaus, 1910

Figs 20, 21, 22, 23, 24

Type species

Eremophygus philippii Ohaus, 1910.

Species

6 species; length 14–15 mm.

Rarity of specimens in collections as well as possible paraphyly with the genera Oogenius, Microogenius, Peruquime, and Lasiocala hampers our understanding of the biodiversity of this group. Species in the genus Eremophygus are distributed in the altiplano of Bolivia, Argentina, Peru, and Chile. Gutiérrez described two species in the genus (Gutiérrez 1951, 1952), discussed the genus (Gutiérrez 1949, 1950, 1951, 1952), and provided the most recent key to species (Gutiérrez 1952), yet he did not discuss the group’s relationships or context within the Rutelinae.

Some species in the genus lack the independently movable claws that are diagnostic of Rutelinae (that is, the apex of meso- and metatarsomere 5 lack a longitudinal slit, a character suite shared with cyclocephaline rhinoceros beetles [Dynastinae: Cyclocephalini]). One species, Eremophygus pereirai Martínez (from Jujuy, Argentina), was transferred to the dynastine tribe Cyclocephalini and the genus Cyclocephala by Martínez (1975b) who compared the toothless maxillary galea of E. pereirai to the similar maxilla in Cyclocephala zischkai Martínez from Bolivia (Martínez 1960, 1965). Endrödi (1977) agreed with the tribal transfer and also compared E. pereirai to C. zischkai, considering these species distinctive enough so that, together, they could warrant subgeneric status within Cyclocephala. Eremophygus pereirai (as C. pereirai) was later included in the key to world Dynastinae and Cyclocephalini (Endrödi 1985). Krajcik (2012) included E. pereirai under Cyclocephala following Endrödi. Cyclocephala zischkai and C. pereirai have male parameres that are formed from two, laterally articulated plates, a character associated with Cyclocephalini and not Rutelini (male parameres are fused into a single plate that is not laterally articulating). This represents another example of genera historically considered to be part of Rutelini (e.g., Peltonotus and Acrobolbia Ohaus) that were later transferred to Cyclocephalini. This highlights the need for phylogenetic analyses including Eremophygus to broadly sample taxa from Cyclocephalini and Rutelini to resolve the tribal, and thus the subfamilial, placement of this genus.

Diagnostic characters have greatly diminished reliability because of overlap with Lasiocala, Oogenius, Peruquime, Microogenius, and Cyclocephala and should be used with great caution: dorsal surface often with long, tawny setae; apex of labrum extends beyond clypeal apex, visible in dorsal view; antenna 9- or 10-segmented (9-segmented according to Mondaca and Valencia [2016]); lateral edge of mandibles rounded and without reflexed teeth; apex of clypeus varies from rounded to subtrapezoidal; pronotum with apical bead complete medially, laterally, and basally; lateral edge of protibia with three rounded teeth; apex of fourth metatarsomere lacking spiniform attenuation; base of metatibia nearly straight, lacking distinct notch; apex of meso- and metatibia with many spinules, and; mesosternum not appreciably produced beyond the mesometasternal suture. In some species (e.g., E. lasiocalinus Ohaus), the protarsal claw is enlarged and deeply split; the meso- and metatarsal claws may be deeply split or simple; the unguitractor plate of meso- and metatarsus is subcylindrical with 2 or 3 setae; and the apex of tarsomere 5 (meso- and metatarsus) with 2 weak, longitudinal slits at apex (a character that is not shared by most other Rutelini; instead it is more common in the Melolonthinae and Dynastinae). Larvae, natural history, and sister-group relationships are not known.

Homeochlorota Soula, 2006

Fig. 25

Type species

Pseudochlorota chiriquina Ohaus, 1905.

Species

1 species; length 18–20 mm.

The monotypic genus Homeochlorota is rarely encountered in collections and is narrowly distributed in Costa Rica and Panama. As the generic name implies, the genus shares similarities with the genus Chlorota (an anticheirine leaf chafer) including the form of the metatibia (with emargination at apex and with external apex posteriorly produced), form of the claws (widely toothed), and metamesosternal peg that is produced ventrally. In general appearance, it could be confused with Chlorota flavicollis Bates. Analyses should closely examine relationships with Chlorota and other anticheirine leaf chafers in combination with lasiocaline and pelidnotine chafers.

The ruteline genera Pseudochlorota and Lasiocala comprise the subtribe Lasiocalina (Ohaus 1934b). Soula (2006) abandoned the subtribe because “it clearly is not monophyletic” (translated from French) (Soula 2006: 144), then reinstituted it without reason (Soula 2011). Soula (2006) observed that the species of Pseudochlorota possessed “some similarities” as well as many characters that separate them. On this basis, Soula (2006) transferred Pseudochlorota chiriquina into the genus Homeochlorota, creating this monotypic genus.

The taxon is characterized by the following features: pronotum with apical bead lacking or obsolete medially; mesosternum not appreciably produced beyond metamesosternal suture; metamesosternal peg produced ventrally; lateral edge of mandible with one reflexed tooth; labrum extends beyond apex of the clypeus; apical margin of the labrum arcuate and with a small tooth at the middle; frontoclypeal suture obsolete; metatibia with emargination at apex and with external apex posteriorly produced, and; larger claw on all legs widely cleft (shared with Lasiocala). Natural history and larvae are not known, and sister-group relationships have not been examined.

Homonyx Guérin-Méneville, 1839

Figs 1F, 3A, 26, 27, 28, 29, 30, 31, 32, 33

Type species

Homonyx cupreus Guérin-Méneville, 1839.

Species

14 species and subspecies; length 12–19 mm.

Species in the genus Homonyx are elongate, parallel-sided, subcylindrical, and dark-colored beetles. They strongly resemble the allied genus Parhomonyx but can be separated based on the form of the mandibles (bidentate in Homonyx and broadly rounded with one apical tooth in Parhomonyx), the apex of the metatibia (with many spinules in Parhomonyx and biemarginate in Homonyx), and the feathery fringe of setae at the apex of the elytra (exposed in Parhomonyx; hidden in Homonyx). These genera share additional characters: prosternal process short (well-developed in H. planicostatus); mesosternum not produced beyond the mesometasternal suture; pronotum with bead complete apically, laterally, and basally; claws simple; lateral set of setae on apical edge of 3rd metatarsomere of unequal length and width (versus equal in length and width in Catoclastus).

Species in the genus are distributed in Argentina, Brazil, Bolivia, Ecuador, Uruguay, and Peru. Soula (2010a) provides the most current treatment of species in the genus, but did not include a key for identification. Larvae, sister-group relationships and natural history are poorly known for species in the genus.

Homothermon Ohaus, 1898

Figs 34, 35

Type species

Homothermon bugre Ohaus, 1898.

Species

4 species; length 9–19 mm.

The genus Homothermon includes four uncommon species that are distributed in the Paulista center of endemism in Brazil and Argentina (Rio de Janeiro in the north to Santa Catarina, Rio Grande do Sul, and Misiones in the south) (Müller 1973). Species in the genus are characterized by greatly enlarged metatibia in the male, claws on all legs simple (both male and female), male with a medial tubercle on the protarsal claw, pronotal basal bead incomplete anterior to the scutellum, elytral margin without a bead (=rounded), scutellum that is nearly twice as wide as long, clypeus semi-circular or subtrapezoidal, apex of mandibles bidentate, and parameres with a well-developed ventral plate.

Classification and nomenclatural history of members of the genus has been confused. Ohaus (1898) postulated that the genus was closely related to Thyridium Burmeister and placed it in the subtribe Anticheirina. Bates (1904) omitted the genus Homothermon (even though it was described six years prior to his revision) and also overlooked Homothermon praemorsus (Burmeister) (then classified as Strigidia praemorsus) even though he treated Strigidia as a valid genus. In the Coleopterorum Catalogus (Ohaus 1898, 1918), the genus was placed in the subtribe Pelidnotina. Soula (2008) commented that the genus “approaches” Pelidnota. Based on the elytral margin that lacks a bead, it is possible that the genus may be allied with Plesiorutela Jameson (Rutelini). Future research should examine the relationships of Homothermon, placing it within a broad context of Rutelini.

Homothermon serrano Ohaus and H. bugre are apparently sympatric. Based on our examination of specimens (including type specimens), the two species differ only in color but are conspecific in all other respects. Soula (2008) maintained H. serrano and H. bugre as separate species, and he stated that these are “good examples of two populations where the aedeagus is very similar, however the populations represent two good species that are sympatric” (Soula 2008: 31; translated from French). Lacking additional character evidence, we think that this is open to interpretation.

Natural history for the genus is unknown. Homothermon serrano is known from the forested mountains near Theresopolis, Santa Catarina, Brazil (Ohaus 1898).

Hoplopelidnota F. Bates, 1904

Figs 36, 37

Type species

Hoplopelidnota candezei F. Bates, 1904.

Species

1 species; length 19–24 mm.

The monotypic genus Hoplopelidnota is rarely found in collections. Similar to species of Chalcoplethis and Catoclastus, it possesses metallic green, rugose elytra. Prior to this work, females were not associated with males. The elytral callus of the male possesses a well-developed spine (shared with the pelidnotine genus Mesomerodon; lacking in females of both Mesomerodon and Hoplopelidnota). In addition to the spinose elytra, several unusual characters serve to diagnose the genus: fringe of setae produced beyond apex of elytra; metatibial apex straight (lacking a corbel); mesosternum produced beyond the mesometasternal suture; metasternum with two parallel, longitudinal furrows; pygidium of female with a well-developed horizontal ridge and weak discal concavity; terminal sternite in the female with two deep emarginations on either side of the apex.

The genus includes one species, Hoplopelidnota metallica (Laporte), which has a turbulent nomenclatural history (see “Annotated Catalog”; Moore and Jameson 2013). No analyses have examined the relationships of the genus to other rutelines. Hoplopelidnota metallica is distributed in northern South America, and we provide country records for Brazil, Guyana, and Venezuela.

Mecopelidnota F. Bates, 1904

Figs 2C, 38

Type species

Mecopelidnota arrowi F. Bates, 1904.

Species

8 species; length 17–26 mm.

Species in the genus Mecopelidnota are distinctive for their dark metallic green coloration, large size, elongate body form, and emargination at the base of the metatibia in the male. As currently constituted, the genus includes eight species (Soula 2008), but this may be an over-estimate. Members are distributed both on the east and west sides of the Andes in Colombia, Ecuador, and Peru. The record for M. cylindrica from Guatemala is questionable (Monzón 1996).

The form of the male metatibia (base with an emargination) serves as a synapomorph for the group. Based on our analyses of external morphological characters, the genus includes two lineages: one to the west side of the Andes and one on the east of the Andes. Species on the west of the Andes (M. arrowi, M. cylindrica (Waterhouse), M. marxi Soula, and M. obscura [Taschenberg]) share the form of the male parameres (with enlarged “thumb” in lateral view) and greatly enlarged female gonocoxites. Species on the east side of the Andes (M. witti Ohaus, M. gerardi Soula, M. mezai Soula, and M. dewynteri Soula) share the form of the male parameres (lacking the enlarged “thumb” in lateral view) and reduced female gonocoxites. Both lineages exhibit north-south clinal variation in the form of the male parameres, and Soula (2008: 23) also alludes to this “transitional” variation in species on either side of the Andes.

Species in the genus are recorded from less than 10 m elevation (M. cylindrica and M. obscura; Paucar-Cabrera 2005) to 2700 m (M. obscura). Ohaus (1908b) recorded M. arrowi in the flowers of yellow Mimosa sp. (Leguminosae) in Guayaquil, Ecuador, during the rainy season. In Ecuador, species were collected January to April in tropical regions (Paucar-Cabrera 2005). Larvae and sister-group relationships are not known.

Mesomerodon Ohaus, 1905

Figs 2E, 3C, D, 39, 40

Type species

Mesomerodon spinipenne Ohaus, 1905.

Species

2 species; length 17–24 mm.

The genus Mesomerodon includes two species that are distributed in Colombia, Ecuador, Peru, and Bolivia (Machatschke 1972, Soula 2008). Members are sexually dimorphic, with males having an acute, spiniform processes on the apical callus of the elytra (shared with Hoplopelidnota) and an acute process on the posterior margin of the mesofemur. Species in the genus are testaceous in color, ovate-shaped, and similar in overall gestalt to species of Pelidnota (Pelidnota). The genus is diagnosed by the following additional characters: lateral edge of mandibles without reflexed teeth; lateral edge of protibia with three teeth; pronotum with apical bead complete medially, laterally and basally; mesosternum produced beyond mesometasternal suture; male parameres with a well-developed ventral plate.

The biology of Mesomerodon species is unknown and larvae are not described; sister-group relationships have not been examined. Adults are collected at lights at elevations between 300–750 m.

Microogenius Gutiérrez, 1951

Figs 41, 42, 43

Type species

Oogenius martinezi Gutiérrez 1951.

Species

4 species; length 10–13 mm.

The classification and nomenclatural history of this genus are quite complicated due to two impediments: lack of robust circumscription of ruteline groups and access to literature. Historically, the genus Microogenius was considered a member of the subtribe Lasiocalina and closely related to Lasiocala Blanchard (Martínez 1974) as well as a member of the subtribe Pelidnotina closely related to Eremophygus and Oogenius (Ohaus 1934b). Based on similarities, the two subtribes were combined (Martínez 1974), but this publication was effectively lost until Soula (2006) noted Martínez’s synonymy of Oogenius (Microogenius) and created the new genus Minilasiocala Soula. However, based on the Principle of Priority, Microogenius should be considered the valid name (Moore and Jameson 2013). Similar problems circumscribing ruteline groups led Soula to initially consider the taxon a lasiocaline scarab (Soula 2006) and later to consider it a pelidnotine scarab (Soula 2011). Clearly, phylogenetic and revisionary research must examine relationships of the South American genera Microogenius, Oogenius, Eremophygus, and Lasiocala.

Although the validity of the genus requires evaluation, a few characters can be used with caution for diagnosis: apex of labrum extends beyond clypeal apex and visible in dorsal view (shared with Eremophygus and Oogenius); metatarsomere 4 at apex with 4-6 long setae that are subequal in length and thickness; mandible on external margin rounded (shared with Eremophygus); pronotal basal bead complete (shared with Eremophygus); terminal tergite of female rounded at apex (shared with Eremophygus).

As currently composed, species in the genus are distributed in the altiplano of Bolivia and Argentina. Larvae, natural history, and sister group relationships are not known.

Neogutierrezia Martínez, 1953

Fig. 44

Type species

Neogutierrezia mirabilis Martínez, 1953.

Species

10 species; length 6–9 mm.

Similar to Peruquime, the genus Neogutierrezia is a difficult-to-place taxon with affinities to both Melolonthinae and Rutelinae. Molecular and morphological phylogenetic analyses provided strong evidence that the genus is closely related to members of the Rutelinae, thus Neogutierrezia was transferred from Melolonthinae to Rutelinae (Ocampo et al. 2010). The recent discovery of Peruquime and comparison with Neogutierrezia and Eremophygus establishes an association with pelidnotine chafers, thus our rationale for including the genus herein.

The genus Neogutierrezia is endemic to the Monte biogeographic province in Argentina (Mendoza, Río Negro, Neuquén, Chubut), a shrub steppe region and that coincides with the distribution of Larrea spp., Bulnesia spp., and Plectocarpha spp. (all Zygophyllaceae) (Ocampo et al. 2010). The genus is hypothesized to be a relictual ruteline group that evolved and adapted in situ to the extreme arid conditions of the desert sand dunes (Ocampo et al. 2010).

The genus is diagnosed by the following characters: antennal club longer than stem, and club 3- or 4-segmented (3-segmented in Peruquime); labrum kidney-shaped; pygidial apex “recumbent towards metacoxae” in males; parameres with dorsal and ventral plates fused. Other characters include: frontoclypeal suture complete or obsolete at middle; pronotal apical bead obsolete at middle, complete laterally and basally; and all claws simple.

Species are associated with sandy habitats (sea shores, dunes), and females of one species (N. araucana Martínez) are known to be flightless, probably living underground and only coming to the surface to mate (Martínez 1973). Adults have been collected at light (UV and kerosene lamp) and with un-baited pitfall traps. An identification key to species is available (Ocampo et al. 2010). Larvae are not known.

Oogenius Solier, 1851

Figs 45, 46, 47, 48, 49

Type species

Oogenius virens Solier, 1851.

Species

7 species; length 12–23 mm.

Species in the genus Oogenius are egg-shaped (from which the generic name was derived) and distributed in Chile and Argentina. Based on prevailing usage of the name, Mondaca (2005) corrected the spelling of the genus from “Oogeneius” to Oogenius and provided a catalog of included species. Oogenius was later revised by Mondaca (2016), and this work provided a comprehensive key to species, a distribution map, high-quality images of diagnostic characters, and additional biological information.

Circumscription of the genus and phylogenetic analyses that include Microogenius, Eremophygus, and Lasiocala are necessary to better understand the composition of the genus and sister group relationships. Although Soula (2006) treated the genera Lasiocala and Microogenius (or Minilasiocala by Soula 2006), he omitted the genera Oogenius and Eremophygus. Species in these groups possess a broad overlap in characters and many species are quite rare in collections.

The genus Oogenius can provisionally be identified based on the following characters: pronotum with basal bead obsolete or complete medially, complete laterally and apically; clypeus broadly rounded apically, reflexed; labrum produced beyond apex of clypeus; mandibles broadly rounded externally; inner claw enlarged and weakly split in male; unguitractor plate subcylindrical; 5th meso- and metatarsomeres lacking medial tooth; mesosternum not appreciably produced beyond mesometasternal suture; and ventral surface densely setose.

The immature stages of Oogenius have not been described, but Mondaca (2016) reported that larvae feed on roots and decaying plant matter in three species: O. castilloi Martínez and Peña, O. chilensis Ohaus, and O. virens Solier.

Pachacama Soula, 2006

Fig. 50

Type species

Pachacama ocampoi Soula, 2006.

Species

2 subspecies; length 15–17 mm.

As noted by Soula (2006; translated from French) in his description of this unusual genus, “cladistics or molecular analysis is needed more than ever.” Soula (2006) included this lustrous, dark green chafer in the pelidnotine scarabs, and he noted characters that it shared with Minilasiocala (now a junior synonym of Microogenius) and Chrysophora. Phylogenetic analyses are needed to address sister-group relationships of this taxon. Pachacama ocampoi possesses unusual autapomorphs (prosternum produced anteriorly, mesosternum posterior to prosternal peg with transverse fold), and it is possible that it is more closely related to some anticheirine scarabs.

Pachacama can be diagnosed based on the following characters: dorsal surface smooth (lacking striae, obvious punctures or rugosity); clypeus elongate with parabolic apex (subequal in length and width); external margin of mandible bisinuate with apical tooth reflexed; pronotum with apical bead incomplete at middle (bead complete medially and basally); protibia with 2 external teeth; apex of metatibia produced on external margin; metatarsus 1 short (half the length of metatarsus 2); metacoxal corner produced, acute; mesosternum appreciably produced beyond metamesosternal suture; protarsal claws of male with internal claw enlarged, split (female split); meso- and metatarsal claws simple in male (widely split in female); 5th tarsomeres (all legs) with medial tooth; uncus subcylindrical, tapering at apex.

Pachacama ocampoi is endemic to Ecuador where it is recorded between 500 to 1650 m elevation in the provinces of Cañar and Pichincha. Natural history and larvae are unknown.

Parhomonyx Ohaus, 1915

Figs 1C, 51

Type species

Homonyx fuscoaeneus Ohaus, 1905.

Species

1 species; length 17–22 mm.

The monotypic genus Parhomonyx is endemic to northern Argentina. Ohaus (1905) described P. fuscoaeneus in the genus Homonyx and in conjunction with another unusual pelidnotine, E. oryctoides (originally Homonyx orcytoides). According to Ohaus (1915b), the genera Homonyx and Parhomonyx were closely related, and differences in the mandibular form (apex with two teeth in Homonyx versus apex rounded in Parhomonyx) provide character support for both genera (=lineages). Species in the pelidnotine genera Parhomonyx, Homonyx, and Pseudogeniates are distributed primarily in the southern half of South America. Ohaus (1915b) considered Parhomonyx to be an “intermediate stage” that “led Homonyx to Pseudogeniates” (Ohaus 1915b: 258), and that characters such as coloration, clypeus, mouthparts, elytra, metatibia, and antennae indicated a progression of forms (Jameson and Ocampo 2012). Additional research should examine sister-group relationships of the taxon.

Parhomonyx fuscoaeneus is castaneous-bronze in color and is diagnostic for its rounded mandibular apex with apical tooth (shared with Pseudogeniates; bidentate in Homonyx); metatibial apex with many spinules (biemarginate in Homonyx); elytral apex with a fringe of setae (shared with Pseudogeniates; hidden in Homonyx); protibia lacking weak, basal notch; all claws simple; fifth meso- and metatarsomeres with one or two internal teeth (lacking in Pseudogeniates, shared with Homonyx); lateral set of setae on apical edge of 3rd metatarsomere of equal length and width (versus unequal length and width in Pseudogeniates); pronotum with bead complete apically, laterally, and basally; prosternal process short; mesosternal peg lacking (shared with Homonyx and Pseudogeniates); elytra striate (shared with Pseudogeniates and Homonyx); and body form elongate and parallel-sided (shared with Homonyx and Pseudogeniates). Larvae are not described. Label data indicate that specimens are collected at blacklight.

Parhoplognathus Ohaus, 1915

Fig. 52

Type species

Areoda maculata Gory, 1833.

Species

4 species; length 12–16 mm.

On first glance, the Brazilian Atlantic Coastal forest endemic genus Parhoplognathus appears similar to areodine leaf chafers such as Areoda MacLeay or Byrsopolis Burmeister due to their strongly convex form (in lateral view) and the apex of the metatibia that possesses many spinules. However, whereas areodine chafers possess a complete frontoclypeal suture, species in the genus Parhoplognathus have an obsolete frontoclypeal suture.

Ohaus (1934b) considered Chipita mexicana, to be a member of the genus Parhoplognathus, but several morphological characters (in addition to the disjunct distribution) provide rationale for the monotypic genus Chipita. The genera Parhoplognathus, Chipita, and Platyrutela (an anticheirine leaf chafer) share several similarities, and phylogenetic analyses could examine this overlap.

Species in the genus Parhoplognathus are diagnosed by the following characters: pronotum with apical bead obsolete or lacking medially; clypeal apex quadrate, reflexed, with or without emargination; external edge of protibia with 3 teeth; all claws simple (shared with Chipita and Platyrutela); mandibular palp with horizontal/longitudinal sulcus.

A synopsis of the species in the genus was provided by Soula (2008), but identification key, natural history, and general distributional information were omitted. Natural history, larvae, and sister-group relationships have not been examined for any species in the genus.

Patatra Soula, 2008

Type species

Patatra mathani Soula, 2008.

Species

1 species; length 15.5 mm.

Patatra mathani is metallic green, the internal protarsal claw is widely toothed and other claws are simple, and the parameres share some similarity to species of Chlorota Burmeister or Pseudothyridium Soula (anticheirine scarabs). Soula (2008, 2009) described this monotypic genus based on one male specimen from Pará (Brazil), and he placed it in the tribe Rutelini. He created a case of double homonymy by describing the genus and species identically in two publications (Soula 2008, 2009, see Moore and Jameson 2013). He noted that the taxon possessed characters of both pelidnotines scarabs (complete pronotal basal bead) as well as anticheirine scarabs (scutellum wider than long in the middle, male medial tarsal claws split and other claws simple). The genus ultimately was classified in the pelidnotine scarabs (Soula 2011), but characters that supported this were not provided. The genus was not included in any generic keys, and Soula’s descriptions do not provide adequate characters for separation from other genera. Future research should address the classification and relationships of Patatra mathani. Larvae and natural history for the species are not known.

Pelidnota MacLeay, 1819

Figs 2B, 3B, 53, 54, 55, 56, 57, 58, 59, 60, 61, 62, 63, 64, 65, 66, 67, 68, 69, 70, 71, 72, 73, 74, 75, 76, 77, 78, 79, 80, 81, 82, 83, 84, 85, 86, 87, 88, 89, 90, 91, 92, 93, 94, 95, 96, 97

Type species

Scarabaeus punctatus Linnaeus, 1758.

Species

195 species and subspecies; length 11–37 mm.

From southeastern Canada to Argentina and the Caribbean, members of the genus Pelidnota are obvious members of the entomofauna with diverse forms (some with enlarged metafemora such as P. burmeisteri), diverse colors (from metallic silver in P. teocuitlamayatli Delgado-Castillo, Deloya, and Morón to shiny red and blue in P. rubripennis riedeli [Ohaus]), and diverse maculations (striped green and tan in P. liturella [Kirby] or colorfully spotted in P. xanthospila [Germar]). Their large size, abundance, and beauty make them fairly recognizable. Some species are recognized as pests: P. filippiniae Soula, which defoliates plantations (Lunz et al. 2011) and P. punctata that feeds on leaves in vineyards (Ratcliffe and Paulsen 2008). Complete life cycle and representative larvae are described (Ritcher 1945, 1966, Morón 1976, Morón and Deloya 2002, Rodriguez et al. 2012, Garcia et al. 2013).

Hardy (1975) revised the genus Pelidnota from North and Central America and provided a key to species. He geographically restricted his revision to North and Central American species due to the large size of the group. The work stabilized the classification of North and Central American taxa and provided the only method of accurately identifying species in this region. He did not discuss relationships among the subgenera of Pelidnota, although he noted that the classification and subgeneric concept (as proposed by Ohaus) were in need of study. Subsequent to Hardy’s revision (Hardy 1975), many new species of Pelidnota have been described. Keys to the Mexican species (Delgado et al. 1988) and Costa Rican species (Solís and Morón 1994) of Pelidnota are available. Soula (2006, 2008, 2009, 2010a, 2010b, 2011) described 104 species and subspecies and provided difficult-to-use keys to many species.

Research on Pelidnota and its allies was initiated by one of us in the 1990s (MLJ). This research, however, became intractable when Soula began describing many pelidnotine taxa and depositing type specimens in his private collection where they were not accessible to other scientists. Additionally, Soula created many new species for North American morphotypes of P. punctata (see “Pelidnota punctata (Linnaeus) species hypothesis and synonyms” below). A comprehensive revision of the genus and its allies is needed, including identification resources for all species.

Molecular and morphological phylogenetic analyses are necessary to unravel the evolutionary and ecological patterns within this interesting group. For over a century, taxonomy and nomenclature of the genus has been mired with several genus-level nomenclatural and classification conflicts (F. Bates 1904, Ohaus 1918, 1934b, Machatschke 1970, 1972, 1974, Krajcik 2008, Özdikmen 2009, Soula 2006, 2008, 2009, 2010a, 2011) (see Moore and Jameson 2013). Whereas the taxonomy and composition of Pelidnota (Pelidnota) is stable (ICZN 2003) and fairly homogeneous, other genus-level names are much less stable, the composition unknown, and identification is problematic (including Chalcoplethis, Epichalcoplethis, Strigidia, Odontognathus, Ganonota Ohaus). It is possible that Pelidnota sensu lato includes several natural groups (=genera), but to truly unravel the group, an unabridged systematic revision (taxonomy, phylogeny) must be undertaken and the group must be examined within a broad context of the Rutelini.

Due to possible paraphyly, diagnosis of the genus is difficult. For most species of Pelidnota, the pronotal basal bead is complete (obsolete in some); external margin of the mandible is bidentate; mesosternum with a transverse suture that separates the metasternum; prosternal projection more or less prominent and beaded; scutellum as wide as long; mesosternal projection not well-developed, not strongly produced anteriorly; elytral shoulder with a bead; metatrochanters sometimes protruding; claws simple in both sexes; male protarsal claw with or without inner tubercle; metatibia simple, gradually widening from base or corbeled.

Ohaus (1912) described the genus Heteropelidnota based on one, unusual male specimen (Ohaus 1934b; Plate 2, Fig. 11) (Fig. 72). The color and form of the specimen (the holotype of P. kuhnti [Ohaus] and the only known representative of the taxon) (Fig. 72) was compared with individuals of P. aeruginosa var. citripennis (valid name P. semiaurata citripennis) (Ohaus 1912). Examination of this specimen reveals that it is an aberrant, teratological specimen (see discussion of P. kuhnti in “Annotated Catalog”). In Ohaus’ (1934b) discussion of the genus Heteropelidnota, he compared the genus with Hoplopelidnota and Xenopelidnota, both of which possess a dense row of setae near the ventral apex of the elytra. Ohaus (1934b) stated that Hoplopelidnota and Xenopelidnota differ from Heteropelidnota based on the bidentate mandible and produced mesometasternal peg. It should be noted that the dense row of setae on the ventral side of the elytra is observed within many rutelines, but the position (subapically, anteapically, apically) and the density of setae varies widely. The function of this character is unknown (possibly functioning in flight or preventing water loss) and should be investigated. Ohaus included a new species in the genus, P. cribrata (Ohaus), and he transferred Pelidnota rostrata Burmeister (Ohaus 1918) to the genus. Martínez (1967) described P. ustarani (Martínez), also including it in the genus. After examination of the species included in the genus and based on lack of sufficient “collective” characters that support the genus, Soula (2008) transferred P. cribrata, P. ustarani, and P. rostrata to the genus Pelidnota (Soula 2008). However, he retained H. kuhnti in the genus based on its many “singularities.” Indeed, Soula (2008) also seemed to imply that H. kuhnti was a member of the genus Pelidnota. Herein, we consider Heteropelidnota a new junior synonym of Pelidnota. Lacking certainty of the species association due to the extreme deformities, we retain the species name and transfer the species to the genus Pelidnota.

Peruquime Mondaca & Valencia, 2016

Fig. 98

Type species

Peruquime arequipensis Mondaca & Valencia, 2016.

Species

1 species; length 8.3–10.5 mm.

Peruquime arequipensis is a small, setose scarab that inhabits high elevation (3,800–4,000 m), arid regions in southern Peru. The monotypic taxon possesses several unusual characters that are not typically observed in the Rutelinae: labrum projects anteriorly beyond the clypeal apex and fused to the clypeus (similar to some Melolonthinae: Tanyproctini or “pachydemine” scarabs), labrum horizontally produced with respect to the clypeus, antennal club is greatly enlarged (Mondaca and Valencia 2016). The taxon was classified in the tribe Rutelini based on the independently movable claws and laterally flattened unguitractor plate. The taxon was compared with Eremophygus, but it differs based on the pyriform mentum (form oval in Eremophygus) and antenna with 10 segments and enlarged club (antenna 9- or 10-segmented and lacking enlarged club in Eremophygus). It was postulated that Peruquime, together with Neogutierrezia, possess convergent characters that allow for adaptions to arid habitats (Mondaca and Valencia 2016).

Peruquime arequipensis is endemic to the Puna biogeographic region of the Andes, an area known for high endemism. Adult Peruquime arequipensis are diurnal and are active during the rainy season where they were collected in traps (flight intercept, pan, and pitfall). Larvae and sister-group relationships are not known.

Pseudogeniates Ohaus, 1910

Fig. 99

Type species

Pseudogeniates richterianus Ohaus, 1910.

Species

3 species; length 12–19 mm.

The genus Pseudogeniates is endemic to Argentina, and species are associated with arid habitats in the Chaco, Pampa, Espinal, and Monte ecoregions (Jameson and Ocampo 2012). The genus includes three species that are poorly represented in collections. Ohaus (1910a) puzzled over the first specimens that he studied in the genus, and originally thought that they represented teratological abnormalities due to the unusual form of the clypeus and mouthparts that resemble species in the Geniatini and Anoplognathini (both Rutelinae). As the genus name implies, members resemble species in the genus Geniates (Geniatini), but they are easily diagnosed by the feathery fringe of setae on the ventral edge of the elytra, the mesosternal peg that is lacking, claws on all legs that are simple (lacking inner tubercle), the incomplete frontoclypeal suture, the maxillae that lack teeth, and the mandibular apex that has only one, recurved tooth (Jameson and Ocampo 2012).

Species in the genus are reviewed and an identification key is available (Jameson and Ocampo 2012). Natural history is poorly known, and the immature stages have not been described. Adults have been collected at lights from December to February at elevations ranging from 500–750 m.

Sorocha Soula, 2006

Figs 1E, 100, 101, 102, 103, 104, 105

Type species

Pelidnota acutipennis F. Bates, 1904.

Species

16 species and subspecies; length 16–19 mm.

Soula (2006) described the genus Sorocha for a homogeneous group of species that Ohaus (1934b) had placed in the “P. pulchella group” based on the smooth, shiny elytra that lack markings, and distribution in the Andean highlands. Based on overall gestalt, species in the genus Sorocha are similar to species in the genus Pseudochlorota Ohaus (Lasiocalina), but they differ in the following respects: larger claw on all legs (simple in Sorocha; widely split in Pseudochlorota); unguitractor plate (flat and with two apical setae in Sorocha; subcylindrical and with two or more setae in Pseudochlorota); pronotal basolateral corner (quadrate in Sorocha; rounded in Pseudochlorota); and apex of the metatibia (lacking spinules in Sorocha; possessing spinules in Pseudochlorota).

This taxon requires phylogenetic analysis because we think some species are probably more appropriately placed in Pelidnota. Sorocha can be characterized, in part, by the following characters: disc of the frons with a V-shaped depressed region (shared with Pseudochlorota); all claws simple; male protarsal claw with inner tubercle; bidentate mandibles; pronotum with bead complete or incomplete apically (complete laterally and basally); elytral base with a median “dimple” lateral of scutellum; elytral epipleuron shelf-like (not rounded); protibia with basal external tooth slightly removed from apical teeth; clypeal length shorter than length of frons; eyes large; apex of the metatibia biemarginate and lacking apical spinules; meso- and metatarsomere 5 lacking internomedial tooth; mesometasternal keel not surpassing mesocoxae; metasternum with dense pilosity.

Species in the genus are superficially similar, and identification is hampered due to lack of a key. Females cannot currently be identified due to similarity among species. Species in the genus are distributed at high elevations from Colombia and Venezuela to Ecuador, Bolivia, and Peru. Larvae are not known. Soula (2006) stated that species in the genus are not readily attracted to lights at night.

Xenopelidnota F. Bates, 1904

Figs 106, 107

Type species

Plusiotis anomala Burmeister, 1844.

Species

3 species and subspecies; length 19–27 mm.

Species in the genus Xenopelidnota resemble castaneous-colored Pelidnota, but the taxon is easily diagnosed by its dark-brown color and parabolic clypeus. The apices of the mandibles are quite variable (weakly bidentate, unidentate, rounded), perhaps due to wear and age. Additional characters that diagnose the genus are as follows: claws simple; male protarsal claw with inner tubercle; pronotum with bead complete apically, laterally and basally; elytral epipleuron shelf-like (not rounded); elytral apex with dense, short tawny setae; fifth meso- and metatarsomeres lacking internomedial tooth; apex of metatibia expanded, straight (lacking corbel or emarginations), and with many spinules; prosternal keel short (not produced to level of procoxae); and mesosternum not appreciably produced beyond mesometasternal suture.

Species in the genus are distributed in northern South America (Colombia, Venezuela, Trinidad, St. Vincent and the Grenadines). As typical of rutelines in this region, species are externally quite similar but male parameres possess a great deal of variability. Phylogenomic analyses of the Xenopelidnota lineage may reveal a greater understanding of the biogeography of the region. Larvae, natural history, and sister-group relationships of the group are not known.

Pelidnota punctata (Linnaeus) species hypothesis and synonyms

Pelidnota punctata (Linnaeus) is a widespread species in North America occurring from Ontario and Quebec to Florida west to South Dakota and Texas. The host plant of this species is grape (Vitis Linnaeus; Vitaceae) foliage and fruit and the larvae develop in rotting stumps and logs of various deciduous trees.

The taxonomic history of this species dates back to the very beginning of zoological binomial nomenclature with a brief description by Linnaeus (1758) (as Scarabaeus punctatus), followed by more extensive (and quite sufficient) descriptions in later editions of Systema Naturae (Linnaeus 1764, 1767). The locality was erroneously given as “India” (Linnaeus 1758, 1764), but was later corrected to “Carolina” (Linnaeus 1767). The photograph of the Linnaeus lectotype specimen leaves no doubt about the identity of this species (Fig. 79). The lectotype is formerly of the Ludovicae Ulricae collection, which is now housed in the Zoological Institute in Uppsala University, Sweden (UUZM).

Melolontha lutea Olivier, 1789 was later described, but it has since been recognized that Olivier (1789) was describing the lighter colored and non-spotted southeastern United States version of this same species (see Hardy 1975 for a detailed discussion). Casey (1915) has the unfortunate notoriety of describing a further 10 synonyms of P. punctata based largely on intraspecific color variations in this species. Hardy (1974, 1975) synonymized all of these Casey names during the course of his taxonomic revision of the genus Pelidnota. Hardy (1975) gave a good account of the color variation of P. punctata and detailed a north-south cline of variation. We have observed that specimens from Canada and the northern United States always have dark legs and clearly defined spots (six on the elytra and two on the pronotum) while the legs and spots can be much lighter in specimens from the southern United States. We have observed that many specimens from Florida and Texas have light legs and little to no trace of spots on the pronotum or elytra. Considering the scope of the variation, even within smaller regions, we postulate that larval diet/nutrition, environmental conditions during development, and the length of time spent in the larval stage can have a significant impact on color patterns along with genetics for this particular species. Ritcher (1966) stated that in Lexington, Kentucky, P. punctata pass the winter in the larval stage and appear to have a two-year life cycle. It is possible that the life cycle of this species is accelerated in the southern part of the range. Pelidnota punctata may have an extended life cycle in the northern part of the range in response to decreased temperature and more extreme seasonal climate fluctuation.

Pelidnota genieri Soula was described as a purported species endemic to Ottawa, Ontario, Canada (Soula 2009). Soula’s (2009) description was based on color patterns and trivial structural characters without any detailed comparisons with P. punctata specimens from other parts of Canada and North America. We studied the holotype, allotype, and ten paratypes (five males, five females) from Soula’s type series and have concluded that the color patterns observed are well within the range of variation observed from specimens of P. punctata from other parts of Ontario, Quebec, and across the eastern half of the United States (over 500 specimens were examined). In fact, the holotype, allotype, and two paratypes were from a larger series of 51 specimens from the same collecting event (Ottawa, ONT. / 5. VIII.1971 / J.E.H. Martin) all in the Canadian Museum of Nature Collection. Having seen only four of the 51 specimens from this collecting event, Soula was unaware that the color variation observed in this series alone undermined many of the characters used to justify his new species (varying shades of dorsal coloration, different sizes of dark spots, different amounts of metallic green reflections around and between the eyes). Therefore, we are placing Pelidnota genieri in synonymy with Pelidnota punctata (syn. n.).

Soula’s (2009) motivation for describing such an obvious synonymy is unclear, but the quality of his work is highly suspect in our opinions after using his publications and examining material identified and described by him in the Canadian Museum of Nature collection and CCECL. Since Marc Soula’s death in 2012, his taxonomic work has come under increased scrutiny and criticism for poor quality (e.g., Moore and Jameson 2013, Garner and Audibert 2015). A case in point relevant to Pelidnota genieri was the fact that Soula (2006) had previously described Strigidia genieri Soula, 2006, and then he transferred this species to Pelidnota (in Soula 2009), creating a secondary homonym with the Pelidnota genieri in the very same paper where the former name was described! Moore and Jameson (2013) fixed this homonymy problem by erecting Pelidnota francoisgenieri Moore & Jameson, 2013 as a replacement name for Pelidnota genieri Soula, 2009 (not Pelidnota genieri [Soula, 2006]). As an objective synonym of Pelidnota genieri, Pelidnota francoisgenieri is also here placed in synonym with Pelidnota punctata (syn. n.).

Soula (2009) also re-validated the names Pelidnota lutea (Olivier) and Pelidnota texensis Casey from synonymy with Pelidnota punctata for the Florida and Texas populations, respectively. He discussed some morphological differences between these populations and pointed out some variations in the male genitalia but also remarked that a DNA analysis would be necessary to determine the classification of this group of species. While we acknowledge that there are some morphological differences, we do not see stable and consistent differences between the populations or forms of P. punctata enough to warrant splitting this species at this time. DNA barcoding data is available for specimens from Ontario, Florida, Arkansas, and Texas (Fig. 4). Based on the lack of consistent morphological differences and the virtually identical CO1 barcoding data for specimens from the northern and southern extremes of the distribution, we hereby reinstate the junior synonymy of both Pelidnota lutea and Pelidnota texensis with Pelidnota punctata.

Figure 4.

Neighbor-joining tree for individuals of P. punctata across the species’ distribution based on CO1 data. Between-species divergence is typically above 10% (e.g., P. punctulata and P. strigosa), whereas within species divergence is typically less than 1% (e.g., P. punctata and P. lugubris).

In the genus Pelidnota, analysis of the CO1 barcode data (Fig. 4) shows that between-species divergence is typically more than 10% (e.g., P. punctulata and P. strigosa). In contrast, the CO1 barcode region typically shows less than 1% divergence within species (e.g., P. lugubris), even when the individuals are separated by more than 200 miles (Sonora, Mexico to Arizona, USA). Similarly, individuals of P. punctata that were collected from across the species’s range (Florida, USA to Ontario, Canada) showed less than 1% CO1 divergence (mean: 0.24 % infraspecific divergence; max: 0.77 % infraspecific divergence). Individuals of Pelidnota punctata from Ontario, Canada (in the northern extent of the distribution) possess dark metallic green elytral spots, legs, and venter. Some individuals from Florida, USA (in the southern extent of the distribution) possess reduced elytral spots or no elytral spots and possess tan legs and venter. The CO1 data demonstrate that these phenotypic differences are not strong characters for species cohesion and provide support that all morphotypes of P. punctata are, in fact, conspecific (see “Annotated Catalog” for synthetic taxonomic information of P. punctata and synonyms).

Annotated Catalog of the Pelidnotine Scarabs (Coleoptera: Scarabaeidae: Rutelinae: Rutelini)

Tribe RUTELINI MacLeay, Subtribe RUTELINA MacLeayGroup Pelidnotine scarabs (paraphyletic)

27 genera (26 extant and 1 extinct) and 420 species and subspecies (419 extant and 1 extinct).

CATOCLASTUS Solier, 1851

Catoclastus Solier, 1851: 95–96.

Type species

Catoclastus chevrolatii Solier, 1851: 96-97, by monotypy.

Gender

Masculine.

Species

3 species.

Catoclastus chevrolatii Solier, 1851

Catoclastus chevrolatii Solier, 1851: 96–97 [original combination].

Catoclastus chevrolati Solier [incorrect subsequent spelling by Harold 1869b: 1226].

Distribution

PERU: Ayacucho (Ohaus 1918, 1934b, 1952, Blackwelder 1944, Machatschke 1972, Ratcliffe et al. 2015).

Types

Soula (2010a: 4) designated the male neotype of Catoclastus chevrolatii at MNHN.

Remarks

Solier (1851) described C. chevrolatii from “various parts of Chile” (translated from Spanish) and subsequent authors continued to cite these data (Harold 1869b, Reed 1876, Philippi 1887, Ohaus 1910c, 1918, 1934b, 1952, Blackwelder 1944, Machatschke 1972, Krajcik 2008, Soula 2010a). We have not examined any Catoclastus specimens from Chile and we consider these data erroneous. Catoclastus chevrolatii is currently known only from Peru.

Catoclastus jaumesi Soula, 2010

Catoclastus jaumesi Soula, 2010a: 6 [original combination].

Distribution

PERU (Soula 2010a, Ratcliffe et al. 2015).

Types

The following specimens are deposited at CCECL. 1 ♂ Holotype (Fig. 5A, B, C), 1 ♀ allotype (Fig. 5D, E), 8 ♂ paratypes, 4 ♀ paratypes: “Matucama; Pérou 2000m; II/2002//Holotype Catoclastus jaumesi S. 2010 Soula” (47031020); “Matucama; Pérou 2000m; II/2002//Allotype Catoclastus jaumesi S. 2010 Soula” (47031021); Eight paratypes with identical label data: “Matucama; Pérou 2000m; II/2002//Paratype Catoclastus jaumesi S. 2010 Soula” (47031022 to 47031028, exch58); “Chancho Moy Peru Kirsch//Paratype Catoclastus jaumesi S. 2010 Soula” (47031029); Three paratypes with identical label data: “Pérou coll. – SOULA//Paratype Catoclastus jaumesi S. 2010 Soula” (47031030 to 47031032). Genitalia card-mounted underneath the male holotype and five male paratypes. Box 4618690 SOULA.

Catoclastus rabinovichi Martínez, 1971

Catoclastus rabinovichi Martínez, 1971: 79–81[original combination].

Distribution

PERU: Cusco (Martínez 1971, Ratcliffe et al. 2015).

CHALCOPLETHIS Burmeister, 1844

Chalcoplethis Burmeister, 1844: 410.

Pelidnota (Chalcoplethis) Burmeister [new subgenus status by Ohaus 1915b: 258–259].

Chalcoplethis Burmeister [revised genus status by Soula 2006: 98-99].

Type species

Chrysophora kirbii Gray, 1832: 516, by monotypy.

Gender

Feminine.

Species

2 subspecies.

Remarks

Krajcik (2012, 2013) considered Chalcoplethis to be a junior synonym of Pelidnota.

Figure 5.

Catoclastus jaumesi Soula holotype male and allotype female from CCECL. A Dorsal habitus, holotype B Specimen labels, holotype C Male genitalia, dorsal view, holotype D Dorsal habitus, allotype E Specimen labels, allotype.

Chalcoplethis kirbii kirbii (Gray, 1832)

Chrysophora kirbii Gray, 1832: 516 [original combination].

Chalcoplethis kirbii (Gray) [new combination by Burmeister 1844: 410–411].

Chalcoplethis kirbyi (Gray) [incorrect subsequent spelling by Harold 1869b: 1224].

Pelidnota (Chalcoplethis) kirbyi (Gray) [new subgeneric combination by Ohaus 1918: 29].

Chalcoplethis kirbyi (Gray) [revised combination by Soula 2006: 99–100].

Distribution

BRAZIL: Bahia, Paraná, Espírito Santo, Rio Grande do Sul (Gray 1832, Burmeister 1844, Blanchard 1851, Harold 1869b, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Hardy 1975, Soula 2006, Krajcik 2008). COSTA RICA (Hardy 1975). PARAGUAY: Cororó (HNMB).

Types

1 ♂ holotype of Chrysophora kirbii at BMNH (Soula 2006).

Figure 6.

Chalcoplethis kirbii misionesensis Soula holotype male from CCECL. A Dorsal habitus B Lateral habitus C Specimen labels D Male genitalia, lateral view E Male genitalia, dorsal view.

Chalcoplethis kirbii misionesensis Soula, 2010

Chalcoplethis kirbyi misionesensis Soula, 2010a: 46–47 [original combination].

Chalcoplethis kirbii misionesensis Soula [see suggested correct spelling by Moore and Jameson 2013: 383].

Distribution

ARGENTINA: Misiones (Soula 2010a).

Types

The following specimens are deposited at CCECL. 1 ♂ holotype (Fig. 6), 1 ♀ allotype, 2 ♂ paratypes, 2 ♀ paratypes: “Pozo Azul, Misiones Argentine, XII/2009//Holotype Chalcoplethis kirbyi misionesensis Soula 2010” (47030871); “Pozo Azul, Misiones Argentine, XII/2009//Allotype Chalcoplethis kirbyi misionesensis Soula 2010” (47030872); Three paratypes with identical label data: “Pozo Azul, Misiones Argentine, XII/2009//Paratype Chalcoplethis kirbyi misionesensis Soula 2010” (47030873, 47030956 and 47030957); “Misiones Arg. M. SOULA det. 19//Paratype Chalcoplethis kirbyi misionesensis Soula 2010” (47030874). Genitalia are card-mounted underneath the male holotype and the male paratype specimens. Box 4618649 SOULA and 4616345 PORION.

Chipita Soula, 2008

Chipita Soula, 2008: 10.

Type species

Byrsopolis mexicana Ohaus 1905: 324, by monotypy.

Gender

Feminine.

Species

1 species.

Chipita mexicana (Ohaus, 1905)

Byrsopolis mexicana Ohaus, 1905: 324–325 [original combination].

Parhoplognathus mexicanus (Ohaus) [new combination by Ohaus 1915b: 257].

Chipita mexicana (Ohaus) [new combination by Soula 2008: 10].

Distribution

MEXICO: Sinaloa (FSCA), Guerrero, Jalisco, Nayarit, Oaxaca (Ohaus 1905, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón et al. 1988, Rodríguez-Palafox and Corona 2002, Soula 2008, Deloya et al. 2014).

Types

1 ♀ lectotype and 1 paralectotype of Byrsopolis mexicana at ZMHB (Soula 2006). An exemplar specimen is figured (Fig. 7).

Figure 7.

Chipita mexicana (Ohaus) female specimen from FSCA. A Dorsal habitus B Lateral habitus.

CHRYSINA Kirby, 1828

Chrysina Kirby, 1828: 522.

synonym. Plusiotis Burmeister, 1844

Plusiotis Burmeister, 1844: 417. [Type species. Pelidnota victorina Hope, 1841, by original designation].

Chrysina Kirby [syn. by Hawks 2001: 2].

synonym. Plusiotina Casey, 1915

Plusiotina Casey, 1915: 84. [Type species. Plusiotina aeruginis Casey, 1915 by subsequent designation (Hawks 2001: 2) (= Chrysina lecontei [Horn, 1882])].

Plusiotis Burmeister [syn. by Ohaus 1934b: 59].

synonym. Pelidnotopsis Ohaus, 1915b

Pelidnotopsis Ohaus, 1915b: 257. [Type species. Pelidnota plusiotina Ohaus, 1912, by monotypy].

Chrysina Kirby [syn. by Hawks 2001: 2].

Pelidnotopsis Ohaus [revised genus status by Soula 2010b: 11].

Chrysina Kirby [syn. by Moore and Jameson 2013: 381].

Type species

Chrysina peruviana Kirby 1828: 523, by monotypy.

Gender

Feminine.

Species

113 species.

Remarks

Casey (1915) did not designate a type species for the genus Plusiotina. No authors addressed this until Machatschke (1972) subsequently designated Plusiotis woodi Horn as the type of Plusiotina. This type designation was invalid based on ICZN Article 67.2 that states “A nominal species is only eligible to be fixed as the type species of a nominal genus or subgenus if it is an originally included nominal species”. Plusiotis woodi Horn was not an originally included nominal species of Plusiotina and was thus invalidly designated as the type species of Plusiotina. Hawks (2001) corrected this by subsequently designating Plusiotina aeruginis Casey (= Chrysina lecontei [Horn, 1882]), an originally included nominal species of Plusiotina, as the type species of Plusiotina.

Chrysina adelaida (Hope, 1841)

Pelidnota adelaida Hope, 1841: 147 [original combination].

Plusiotis adelaida (Hope) [new combination by Burmeister 1844: 421].

Plusiotis adelaidae (Hope) [incorrect subsequent spelling by Nonfried 1891: 302].

Chrysina adelaida (Hope) [new combination by Hawks 2001: 7].

synonym. Pelidnota ornatissima Sturm, 1843

Pelidnota ornatissima Sturm, 1843: 341-342 [original combination].

Plusiotis adelaida (Hope) [syn. by Burmeister 1844: 421].

synonym. Plusiotis adelaida pavonacea Casey, 1915

Plusiotis adelaida pavonacea Casey, 1915: 84.

Chrysina adelaida (Hope) [syn. n.].

Distribution

MEXICO: Chiapas, Chihuahua, Coahuila, Colima, Durango, Guerrero, Hidalgo, Jalisco, México, Michoacán, Morelos, Oaxaca, Puebla, Queretaro, San Luis Potosí, Tamaulipas, Tlaxcala, Veracruz (Hope 1841, Sturm 1843, Burmeister 1844, Lucas 1865, Harold 1869b, Boucard 1875, H. W. Bates 1888, Nonfried 1891, Casey 1915, Ohaus 1918, 1934b, Blackwelder 1944, Cazier 1951, Carrillo et al. 1966, Machatschke 1972, Morón 1985, 1990, 1991, 1994, Morón and Zaragoza 1976, Morón and Deloya 1991, Ratcliffe et al. 1992, Deloya et al. 1993, Lobo and Morón 1993, García-Montiel et al. 2003, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Márquez and Sierra-Martínez 2008, Muñoz-Hernández et al. 2008, Morón and Márquez 2012, Márquez et al. 2013, Deloya et al. 2014).

Remarks

Hawks (2006) provided an online “Checklist of Chrysina species” wherein he considered C. adelaida pavonacea Casey to be a synonym of C. adelaida. Casey (1915) proposed this subspecies for individuals from Guerrero, Mexico, which differed from the nominative form based on “feebly convex intervals”, “very shallow clypeal sinuation”, and slight differences in color. We agree that this subspecies is conspecific with the nominative form. Because the on-line checklist is not considered to be formally published for nomenclatural purposes, we synonymize this subspecies herein. Chrysina adelaida (=C. ornatissima) was reported from “San Gerónimo”, Guatemala (possibly San Jerónimo, Baja Verapaz) (H. W. Bates 1888, Ohaus 1918, 1934b, Blackwelder 1944). There have been no further published collection records for C. adelaida in Guatemala and these data need to be re-evaluated.

Chrysina adolphi Chevrolat, 1859

Chrysina adolphi Chevrolat, 1859: 481 [original combination].

Chrysina macropus var. adolphi Chevrolat [new infrasubspecific status by H. W. Bates 1888: 285].

Chrysina macropus adolphi Chevrolat [new subspecific status by Machatschke 1972: 17].

Chrysina macropus (Francillon) [syn. by Morón 1990: 54].

Chrysina adolphi Chevrolat [revised species status by Hawks 2001: 2].

Distribution

MEXICO: Guerrero, Oaxaca (Chevrolat 1859, Harold 1869b, Boucard 1875, H. W. Bates 1888, Ohaus 1918, Blackwelder 1944, Machatschke 1972, Hawks 2001, Thomas et al. 2006, Krajcik 2008).

Types

1 ♀ lectotype of Chrysina adolphi at BMNH (Hawks 2001).

Chrysina aenigmatica (Morón, 1990)

Plusiotis aenigmatica Morón 1990: 29 [original combination].

Chrysina aenigmatica (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: México, Morelos (Morón 1990, Deloya et al. 1993, Krajcik 2008).

Types

1 ♂ holotype, 1 ♀ allotype and 3 paratypes of Plusiotis aenigmatica at MXAL (Morón 1990); 1 paratype at MNHN (Morón 1990); 1 paratype at BMNH (Morón 1990); 2 paratypes at ZMHB (Morón 1990); 2 paratypes at CNC (Morón 1990); 2 paratypes at IEXA (Morón 1990).

Chrysina alfredolaui (Hawks, 1995)

Plusiotis alfredolaui Hawks, 1995: 273–275 [original combination].

Chrysina alfredolaui (Hawks) [new combination by Hawks 2001: 8].

Distribution

GUATEMALA (Thomas et al. 2006, Monzón 2010). MEXICO: Veracruz (Hawks 1995, Krajcik 2008, Thomas et al. 2006).

Types

1 ♂ holotype of Plusiotis alfredolaui at CAS (Hawks 1995); 1 ♀ allotype at EMEC (Hawks 1995).

Chrysina alphabarrerai (Morón, 1981)

Plusiotis alphabarrerai Morón, 1981: 57-63 [original combination].

Chrysina alphabarrerai (Morón) [new combination by Hawks 2001: 8].

Distribution

MEXICO: Veracruz (Morón 1981, 1990, Lobo and Morón 1993, Krajcik 2008, Thomas et al. 2006).

Types

1 ♂ holotype and 1 ♀ allotype at MXAL (Morón 1981).

Chrysina arellanoi Monzón, 2012

Chrysina arellanoi Monzón, 2012: 1–4 [original combination].

Distribution

MEXICO: Oaxaca (Monzón 2012, Thomas et al. 2013, Morón and Nogueira 2016).

Types

1 ♂ holotype and 1 ♀ allotype at CNIN (UNAM) (Monzón 2012).

Remarks

Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis arellanoi. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina arellanoi.

Chrysina argenteola (H. W. Bates, 1888)

Plusiotis argenteola H. W. Bates, 1888: 277 [original combination].

Chrysina argenteola (H. W. Bates) [new combination by Hawks 2001: 8].

Distribution

COLOMBIA: Antioquia, Cauca, Chocó, Nariño, Putumayo, Valle del Cauca (H. W. Bates 1888, Nonfried 1892, Ohaus 1903, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Arnaud 1994, Restrepo et al. 2003, Neita-Moreno et al. 2006, Neita-Moreno 2011, Thomas et al. 2006). ECUADOR: Bolívar, Cotopaxi, Esmeraldas, Pichincha (Ohaus 1903, 1908b; 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Arnaud 1994, Paucar-Cabrera 2005, Thomas et al. 2006, Krajcik 2008, Camacho Cárdenas 2015). PERU: Junín, Lima (Ohaus 1903, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Ratcliffe et al. 2015).

Types

1 ♂ neotype at MNHN (Arnaud 1994).

Chrysina aurigans (Rothschild & Jordan, 1894)

Plusiotis aurigans Rothschild & Jordan, 1894: 504–505 [original combination].

Chrysina aurigans (Rothschild and Jordan) [new combination by Hawks 2001: 8].

synonym. Plusiotis keithi Linell, 1895

Plusiotis keithi Linell, 1895: 1–2 [original combination].

Plusiotis aurigans Rothschild and Jordan [syn. by Ohaus 1918: 18].

Distribution

COSTA RICA: Alajuela, Cartago, San José (Rothschild and Jordan 1894, Linell 1895, Ohaus 1918, 1934b, Blackwelder 1944, 1952, Machatschke 1972, Morón 1990, Thomas et al. 2006, Krajcik 2008, García-López et al. 2013). PANAMA (Thomas et al. 2006).

Chrysina aurilisternum Pérez-Flores, Villagomez, & Galindo, 2016

Chrysina aurilisternum Pérez-Flores, Villagomez, & Galindo, 2016: 607–610 [original combination].

Distribution

MEXICO: Guanajuato (Pérez-Flores et al. 2016).

Types

1 ♂ holotype, 1 ♀ allotype and 17 paratypes at CNIN (UNAM) (Pérez-Flores et al. 2016).

Chrysina auripes Gray, 1832

Chrysina auripes Gray, 1832: 517 [original combination].

Plusiotis auripes (Gray) [new combination by Burmeister 1844: 419].

Chrysina auripes Gray [revised combination and revised application by Hawks 2001: 3].

synonym. Pelidnota auripes Hope, 1841

Pelidnota auripes Hope 1841: 147 [original combination].

Plusiotis auripes (Gray) [syn. by Burmeister 1844: 419].

Chrysina auripes Gray [syn. by Hawks 2001: 3].

synonym. Plusiotis chalchihuitli Morón, 1990

Plusiotis chalchihuitli Morón, 1990: 16, 36-37 [original combination].

Chrysina auripes Gray [syn. by Hawks 2001].

Distribution

MEXICO: Nuevo León, Oaxaca, San Luis Potosi, Tamaulipas (Gray 1832, Laporte 1840, Burmeister 1844, Blanchard 1851, Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Hawks 2001, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ lectotype at OUMNH (Hawks 2001).

Remarks

Krajcik (2012, 2013) erroneously listed C. chalchihuitli as a valid name.

Chrysina aurofoveata (Morón, 1981)

Plusiotis aurofoveata Morón, 1981: 50–57 [original combination].

Chrysina aurofoveata (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Hidalgo, Puebla (Morón 1981, 1990, 1993, 1994, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Muñoz-Hernández et al. 2008, Márquez et al. 2013).

Types

1 ♂ holotype, 1 ♀ allotype and paratypes at MXAL (Morón 1981).

Chrysina auropunctata Ohaus, 1913

Plusiotis auropunctata Ohaus, 1913: 491 [original combination].

Chrysina auropunctata (Ohaus) [new combination by Hawks 2001: 7].

Distribution

GUATEMALA: San Marcos (Morón 1990, Monzón 1995, Monzón et al. 1999, Thomas et al. 2006). MEXICO: Chiapas (Ohaus 1913, 1918, 1934b, Machatschke 1972, Morón 1990, 1991, Thomas 1993, Monzón 1995, Thomas et al. 2006, Krajcik 2008).

Chrysina aurora (Boucard, 1875)

Plusiotis aurora Boucard, 1875: 119 [original combination].

Chrysina aurora (Boucard) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: Alajuela, San José (Morón 1990, Thomas et al. 2007, García-López et al. 2013). NICARAGUA: Chontales (Maes 1987). PANAMA: Chiriquí (H. W. Bates 1888, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Maes 1987, Morón 1990, Ratcliffe 2002, Krajcik 2008, Thomas et al. 2007).

Chrysina badeni (Boucard, 1878)

Plusiotis badeni Boucard, 1878: 298–295 [original combination].

Chrysina badeni (Boucard) [new combination by Hawks 2001: 4].

Distribution

MEXICO: Hidalgo, Puebla, San Luis Potosí, Veracruz (Boucard 1878, H. W. Bates 1888, Nonfried 1892, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, 1993, 1994, Ratcliffe et al. 1992, Cano and Morón 1994, Morón et al. 1997, Hawks 2001, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Morón and Márquez 2012, Márquez et al. 2013).

Types

1 ♀ lectotype at ZMHB (Hawks 2001).

Chrysina baileyana Monzón, 2010

Chrysina baileyana Monzón, 2010: 7–10 [original combination].

Distribution

GUATEMALA: Huehuetenango (Monzón 2010).

Types

1 ♂ holotype and 1 ♀ allotype at UVGC (Monzón 2010); paratypes at UVGC, FSCA and WSU (Monzón 2010). 2 paratypes at MSPC (Fig. 8).

Figure 8.

Chrysina baileyana Monzón paratype male from MSPC. A Dorsal habitus B Lateral habitus C Specimen labels and male genitalia D Male genitalia, lateral view E Male parameres, dorsal view.

Chrysina batesi (Boucard, 1875)

Plusiotis batesi Boucard, 1875: 119–120 [original combination].

Chrysina batesi (Boucard) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: Cartago, San José (Boucard 1875, 1878, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Curoe 2001, Thomas et al. 2006, Krajcik 2008). PANAMA: Chiriquí (Morón 1990, Curoe 2001, Ratcliffe 2002, Thomas et al. 2006).

Chrysina beckeri H. W. Bates, 1889

Chrysina beckeri H. W. Bates, 1889: 411 [original combination].

Distribution

MEXICO: Durango (H. W. Bates 1889, Ohaus 1918, 1934b, Blackwelder 1944, Cazier 1951, Machatschke 1972, Morón 1990, Krajcik 2008, Thomas et al. 2007).

Types

Holotype of Chrysina beckeri at MNHN.

Chrysina benesi Pokorný & Curoe, 2012

Chrysina benesi Pokorný & Curoe, 2012: 111–116 [original combination].

Distribution

MEXICO: Chiapas (Pokorný and Curoe 2012).

Types

1 ♂ holotype (Fig. 9) and 1 ♀ allotype at NMPC (Pokorný and Curoe 2012); 1 paratype at BMNH; additional paratypes at DJCC, MXAL, and other private collections (Pokorný and Curoe 2012).

Figure 9.

Chrysina benesi Pokorný and Curoe holotype male from NMPC. A Dorsal habitus B Lateral habitus C Specimen labels and male genitalia D Male genitalia, lateral view E Male parameres, caudal view.

Chrysina beraudi (Warner, Hawks, & Bruyea, 1992)

Plusiotis beraudi Warner, Hawks, & Bruyea, 1992: 99–100 [original combination].

Chrysina beraudi (Warner, Hawks, and Bruyea) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: San José (Warner et al. 1992, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype at CAS (Warner et al. 1992).

Chrysina beyeri (Skinner, 1905)

Plusiotis beyeri Skinner, 1905: 289-290 [original combination].

Chrysina beyeri (Skinner) [new combination by Hawks 2001: 7].

synonym. Plusiotis ampliata Casey, 1915

Plusiotis ampliata Casey, 1915: 82 [original combination].

Plusiotis beyeri Skinner [syn. by Cazier 1951: 5].

synonym. Plusiotis beyeri ocularis Casey, 1915

Plusiotis beyeri ocularis Casey, 1915: 83 [original combination].

Plusiotis beyeri Skinner [syn. by Cazier 1951: 4].

Distribution

MEXICO: Chihuahua, Sinaloa, Sonora (Coolidge 1911, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Thomas et al. 2006, Lugo et al. 2011). USA: Arizona (Skinner 1905, Biederman 1907, Coolidge 1911, Casey 1915, Ohaus 1918, 1934b, Leng 1920, Blackwelder 1939, 1944, Cazier 1951, Machatschke 1972, Gibson and Carrillo 1959, Morón 1990, 1991, Hardy 1991, Thomas et al. 2006, Krajcik 2008).

Chrysina blackalleri Monzón & García, 2011

Chrysina blackalleri Monzón & García, 2011: 1–4 [original combination]

Distribution

MEXICO: Oaxaca (Monzón and García 2011, Thomas et al. 2012).

Types

1 ♂ holotype and 1 ♀ allotype at CNIN (UNAM) (Monzón and García 2011).

Chrysina boucardi (Sallé, 1878)

Plusiotis boucardi Sallé, 1878: 21 [original combination].

Chrysina boucardi (Sallé) [new combination by Hawks 2001: 8].

synonym. Plusiotis magnificus Arrow, 1919

Plusiotis magnificus Arrow, 1919: 380 [original combination].

Plusiotis boucardi Arrow [syn. by Morón 1990: 32].

Distribution

COSTA RICA: Puntarenas, San José (Boucard 1878, H. W. Bates 1888, Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Curoe 1999, Thomas et al. 2006, Krajcik 2008). PANAMA: Chiriquí (Arrow 1919, Ohaus 1934b, Morón 1990, 1991, Curoe 1999, Ratcliffe 2002, Krajcik 2008).

Chrysina brevis (Rothschild & Jordan, 1894)

Plusiotis brevis Rothschild & Jordan, 1894: 507 [original combination].

Chrysina brevis (Rothschild and Jordan) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Durango, Sinaloa (Rothschild and Jordan 1894, Ohaus 1918, 1934b, Blackwelder 1944, Cazier 1951, Machatschke 1972, Morón 1990, Morón et al. 1997, Krajcik 2008).

Chrysina bruyeai (Hawks, 1999)

Plusiotis bruyeai Hawks, 1999: 22–24 [original combination].

Chrysina bruyeai (Hawks) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Alajuela, Cartago, Guanacaste, Heredia (Hawks 1999, Thomas et al. 2006, Krajcik 2008, García-López et al. 2013). HONDURAS: El Paraiso, Olancho (Hawks 1999, Thomas et al. 2006). NICARAGUA: Zelaya (Hawks 1999, Thomas et al. 2006).

Types

1 ♂ holotype, 1 ♀ allotype and 31 paratypes at MNCR (Hawks 1999); 5 paratypes at MXAL (Hawks 1999); 1 paratype at BMNH (Natural History Museum 2014). 1 ♂ paratype at CMNC.

Chrysina cavei Hawks & Bruyea, 1999

Chrysina cavei Hawks & Bruyea, 1999: 16–18 [original combination].

Distribution

HONDURAS: Olancho, Yoro (Hawks and Bruyea 1999, Thomas et al. 2006, Krajcik 2008).

Types

4 paratypes at BMNH (Natural History Museum 2014, BHG pers. obs. Aug. 2016); 3 ♂ paratypes at CMNC.

Chrysina centralis (Morón, 1990)

Plusiotis centralis Morón 1990: 20 [original combination].

Chrysina centralis (Morón 1990) [new combination by Hawks 2001: 7].

Distribution

GUATEMALA: Quetzaltenango, San Marcos (Morón 1990, 1991; Blackaller-Bages and Delgado 1994, Krajcik 2008, Thomas et al. 2010, Monzón 2010, Morón and Nogueira 2016).

Types

1 ♂ holotype at MXAL (Morón 1990).

Remarks

Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis centralis. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina centralis.

Chrysina chalcothea (H. W. Bates, 1888)

Plusiotis chalcothea H. W. Bates, 1888: 284 [original combination].

Chrysina chalcothea (H. W. Bates) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Cartago, San José (H. W. Bates 1888, Nonfried 1892, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ lectotype at BMNH (Natural History Museum 2014).

Chrysina chloreis (H. W. Bates, 1888)

Plusiotis chloreis H. W. Bates, 1888: 282 [original combination].

Chrysina chloreis (H. W. Bates) [new combination by Hawks 2001: 8].

Distribution

MEXICO: Chiapas, Oaxaca, Veracruz (H. W. Bates 1888, Nonfried 1892, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1981, 1990, Lobo and Morón 1993, Thomas 1993, Krajcik 2008).

Chrysina chrysargyrea (Sallé, 1874)

Pelidnota chrysargyrea Sallé, 1874: 362 [original combination].

Plusiotis chrysargyrea (Sallé) [new combination by Boucard 1875: 120].

Chrysina chrysargyrea (Sallé) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: Puntarenas, San José (Sallé 1874, Boucard 1875, 1878, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas et al. 2006, Krajcik 2008). PANAMA: Chiriquí (H. W. Bates 1888, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Ratcliffe 2002, Thomas et al. 2006).

Chrysina chrysopedila (H. W. Bates, 1888)

Plusiotis aurora var. chrysopedila H. W. Bates, 1888: 277 [original combination].

Plusiotis chrysopedila H.W. Bates [new species status by Ohaus 1912: 307].

Chrysina chrysopedila (H. W. Bates) [new combination by Hawks 2001: 4].

Distribution

COSTA RICA (Morón 1990, Thomas et al. 2006). NICARAGUA: Chontales (H. W. Bates 1888, Nonfried 1892, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Maes 1987, Morón 1990, Thomas et al. 2006). PANAMA: Chiriquí (H. W. Bates 1888, Nonfried 1892, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Maes 1987, Morón 1990, Hawks 2001, Ratcliffe 2002, Thomas et al. 2006, Krajcik 2008).

Types

1 ♀ lectotype and 8 paralectotypes of Plusiotis aurora chrysopedila at BMNH (Hawks 2001); 1 paralectotype at CNC (Hawks 2001).

Chrysina citlaltepetlamayatli (Blackaller-Bages & Delgado, 1994)

Plusiotis citlaltepetlamayatli Blackaller-Bages & Delgado, 1994: 79–83 [original combination].

Chrysina citlaltepetlamayatli (Blackaller-Bages and Delgado) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Querétaro, Veracruz (Blackaller-Bages and Delgado 1994, Krajcik 2008, Thomas et al. 2007, Morón and Nogueira 2016).

Types

1 ♂ holotype and 1 ♀ allotype at MXAL (declaration by authors of final deposition at CNIN [UNAM]) (Blackaller-Bages and Delgado 1994); paratypes at CAS and MXAL (Blackaller-Bages and Delgado 1994).

Remarks

Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis citlaltepetlamayatli. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina citlaltepetlamayatli.

Chrysina clypealis (Rothschild & Jordan, 1894)

Plusiotis clypealis Rothschild & Jordan, 1894: 505–506 [original combination].

Chrysina clypealis (Rothschild and Jordan) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: Cartago, Limón (Rothschild and Jordan 1894, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Hawks 1999, Krajcik 2008, Thomas et al. 2007).

Chrysina colima (Morón, 1992)

Plusiotis colima Morón, 1992: 60-62 [original combination].

Chrysina colima (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Colima, Jalisco (Morón 1992, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype, 1 ♀ allotype and paratypes at MXAL (Morón 1992); 2 paratypes at ZMHB (Morón 1992).

Chrysina confusa (Ohaus, 1913)

Plusiotis confusa Ohaus, 1913: 487–488 [original combination].

Chrysina confusa (Ohaus) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA (Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Krajcik 2008).

Chrysina costata (Blanchard, 1851)

Plusiotis costata Blanchard, 1851: 210 [original combination].

Plusiotis psittacina var. costata Blanchard [new infrasubspecific status by Nonfried 1891: 304].

Plusiotis costata Blanchard [revised species status by Ohaus 1918: 16].

Chrysina costata (Blanchard) [new combination by Hawks 2001: 8].

Distribution

MEXICO: México, Oaxaca, Puebla, Veracruz (Blanchard 1851, Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Carrillo et al. 1966, Machatschke 1972, Morón 1981, 1990, 2010, Ratcliffe et al. 1992, Thomas et al. 2006, Krajcik 2008, Muñoz-Hernández et al. 2008, Delgado-Castillo et al. 2012).

Types

Holotype of Chrysina costata at MNHN.

Chrysina crassimargo (Rothschild & Jordan, 1894)

Plusiotis crassimargo Rothschild & Jordan, 1894: 506 [original combination].

Chrysina crassimargo (Rothschild and Jordan) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Colima, Guerrero, Jalisco, México, Michoacán (Rothschild and Jordan 1894, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, García-Montiel et al. 2003, Thomas et al. 2006, Krajcik 2008, Deloya et al. 2014).

Chrysina cunninghami (Curoe, 1999)

Plusiotis cunninghami Curoe, 1999: 1–4 [original combination].

Chrysina cunninghami (Curoe) [new combination by Hawks 2001: 8].

Distribution

PANAMA: Bocas del Toro (Curoe 1999, Krajcik 2008, Thomas et al. 2007).

Types

1 ♂ holotype at MIUP (Curoe 1999); 1 ♀ allotype at UNSM (Curoe 1999); paratypes at MXAL (Curoe 1999); 1 ♂ paratype at MSPC (Fig. 10).

Figure 10.

Plusiotis cunninghami (Curoe) (valid name Chrysina cunninghami [Curoe]) paratype male from MSPC. A Dorsal habitus B Lateral habitus C Specimen labels and male genitalia D Male genitalia, lateral view E Male parameres, caudal view.

Chrysina cupreomarginata (F. Bates, 1904)

Plusiotis cupreomarginata F. Bates, 1904: 272 [original combination].

Chrysina cupreomarginata (F. Bates) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Cartago, San José (F. Bates 1904, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ lectotype and 1 ♀ paralectotype at BMNH (Natural History Museum 2014, BHG pers. obs. Aug. 2016).

Chrysina curoei (Warner, LeBlanc, Hawks, & Bruyea, 1992)

Plusiotis curoei Warner, LeBlanc, Hawks, & Bruyea, 1992: 96–99 [original combination].

Chrysina curoei (Warner, LeBlanc, Hawks, and Bruyea) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: San José (Warner et al. 1992, Krajcik 2008, Thomas et al. 2007).

Types

1 ♂ holotype at CAS (Warner et al. 1992); 1 ♀ allotype at JPBC (Warner et al. 1992).

Chrysina cusuquensis (Curoe, 1994)

Plusiotis cusuquensis Curoe, 1994: 35–37, 38 [original combination].

Chrysina cusuquensis (Curoe) [new combination by Hawks 2001: 7].

Distribution

HONDURAS: Cortés (Curoe 1994, Krajcik 2008, Thomas et al. 2009, Jocque et al. 2013).

Types

1 ♂ holotype and 1 ♀ allotype at CAS (Curoe 1994); paratypes at EAPZ, CNIN (UNAM), BMNH, MNHN, ZMHB and MXAL (Curoe 1994). The following specimen is deposited at CCECL. 1 ♂ paratype: “HONDURAS: CUSUCO EL CANTIL 1840 m 1-10/VII/94 luz Hg BOSQUE SECUNDARIO PINO LATIFOLIADO D. CUROE COL.//PARATIPO Plusiotis cusuquensis Curoe” (47030024). Genitalia card-mounted underneath specimen. Box 4618644 SOULA.

Chrysina dianae (Ratcliffe & Taylor, 1992)

Plusiotis dianae Ratcliffe & Taylor, 1992: 62–63 [original combination].

Chrysina dianae (Ratcliffe and Taylor) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Veracruz, Oaxaca (Ratcliffe et al. 1992, Cano and Morón 1994, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype at MXAL (Ratcliffe et al. 1992); 1 ♀ allotype at UNSM (Ratcliffe et al. 1992); 2 paratypes at ZMHB (Ratcliffe et al. 1992).

Chrysina difficilis (Morón, 1990)

Plusiotis difficilis Morón, 1990: 19–20 [original combination].

Chrysina difficilis (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Hidalgo, Querétaro, Tlaxcala (Morón 1990, Blackaller-Bages and Delgado 1994, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Márquez et al. 2013, Morón and Nogueira 2016).

Types

1 ♀ holotype and 1 paratype at MXAL (Morón 1990).

Remarks

Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis difficilis. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina difficilis.

Chrysina diversa (Ohaus, 1912)

Plusiotis diversa Ohaus, 1912: 306–307 [original combination].

Chrysina diversa (Ohaus) [new combination by Hawks 2001: 7].

synonym. Chalcochlamys nobilis Ohaus, 1935

Chalcochlamys nobilis Ohaus, 1935: 125 [original combination].

Chrysina diversa (Ohaus) [syn. Jameson and Ratcliffe 2011: 39].

Distribution

BELIZE: Cayo (Gillett 2009). GUATEMALA: Alta Verapaz, Quiche (Cano and Morón 1994, Thomas et al. 2006). MEXICO: Chiapas, Oaxaca, Tabasco, Veracruz (Ohaus 1912, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Morón et al. 1985, 1997, Thomas 1993, Lobo and Morón 1993, Cano and Morón 1994, Thomas et al. 2006, Krajcik 2008).

Chrysina donthomasi Monzón & García, 2011

Chrysina donthomasi Monzón & García, 2011: 5–8 [original combination].

Distribution

MEXICO: Nuevo León (Monzón and García 2011, Thomas et al. 2012).

Types

1 ♂ holotype and 1 ♀ allotype at CNIN (UNAM) (Monzón and García 2011).

Chrysina dzidorhum (Arnaud, 1994)

Plusiotis dzidorhum Arnaud, 1994: 36–37 [original combination].

Chrysina dzidorhum (Arnaud) [new combination by Hawks 2001: 8].

Distribution

ECUADOR: Cañar, Pichincha (Arnaud 1994, Paucar-Cabrera 2005, Krajcik 2008).

Types

1 ♂ holotype and 1 ♀ allotype at PAPC (Arnaud 1994); paratypes at BMNH, MNHN, UFRJ and MXAL (Arnaud 1994); 1 ♂ paratype at CMNC.

Chrysina ericsmithi (Monzón & Cano, 1999)

Plusiotis ericsmithi Monzón & Cano, 1999: 213–214 [original combination].

Chrysina ericsmithi (Monzón and Cano) [new combination by Hawks 2001: 2].

Distribution

GUATEMALA: Izabal (Monzón and Cano 1999, Krajcik 2008, Thomas et al. 2007).

Types

1 ♂ holotype and 1 ♀ allotype at UVGC (Monzón and Cano 1999); paratypes at UVGC, FSCA, MXAL and UNSM (Monzón and Cano 1999).

Chrysina erubescens H. W. Bates, 1889

Chrysina erubescens H. W. Bates, 1889: 411 [original combination].

Distribution

MEXICO: Chihuahua, Durango, Nayarit, Sinaloa (H.W. Bates 1889, Ohaus 1918, 1934b, Blackwelder 1944, Cazier 1951, Machatschke 1972, Morón 1990, Morón and Deloya 1991, Lobo and Morón 1993, Thomas et al. 2006, Krajcik 2008).

Types

Holotype of Chrysina erubescens at MNHN.

Chrysina expansa (Ohaus, 1913)

Plusiotis expansa Ohaus, 1913: 489–490 [original combination].

Chrysina expansa (Ohaus) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Guerrero, Oaxaca (Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas et al. 2006, Krajcik 2008).

Chrysina eyai Curoe, 2012

Chrysina eyai Curoe, 2012: 9–15 [original combination].

Distribution

PANAMA: Darien (Curoe 2012, Thomas et al. 2014).

Types

1 ♂ holotype at EMEC (Curoe 2012).

Chrysina flohri (Ohaus, 1905)

Plusiotis flohri Ohaus, 1905: 321 [original combination].

Chrysina flohri (Ohaus) [new combination by Hawks 2001: 8].

Distribution

MEXICO: Durango, Nayarit (Ohaus 1905, 1918, 1934b, Blackwelder 1944, Cazier 1951, Machatschke 1972, Morón 1990, Morón et al. 1997, 1998, Thomas et al. 2006, Krajcik 2008).

Chrysina gaellae Ebrard & Soula, 2010

Chrysina gaellae Ebrard & Soula, 2010: 7–9 [original combination].

synonym. Chrysina hawksi Monzón, 2010

Chrysina hawksi Monzón, 2010: 4-7 [original combination].

Chrysina gaellae Ebrard and Soula [syn. by Soula 2011: 83].

Distribution

GUATEMALA: Baja Verapáz, Huehuetenango, Zacapa (Ebrard and Soula 2010, Monzón 2010, Thomas et al. 2016, Morón and Nogueira 2016). MEXICO: Chiapas (Monzón 2010, Thomas et al. 2016, Morón and Nogueira 2016).

Types

1 ♂ holotype of Chrysina gaellae at DEPC (Soula and Ebrard 2010).

Remarks

Possibly with ill intentions, the names Plusiotis hawksi (Monzón 2010 [Oct. 15]) and C. gaellae (Ebrard and Soula 2010 [Oct. 1]) were published the same month (see discussion in Soula 2011: 83). Based on the date of publication provided in Soula (2011), C. gaellae is the valid name. Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis hawksi. It is possible that they overlooked the description of C. gaellae (Ebrard and Soula 2010) and/or the synonymy by Soula (2011). Because Morón and Nogueira (2016) do not revalidate the genus Plusiotis, we use the currently valid genus Chrysina.

Chrysina gaitalica Curoe & Hawks, 2012

Chrysina gaitalica Curoe & Hawks, 2012: 9–15 [original combination in Curoe 2012).

Distribution

PANAMA: Coclé (Curoe 2012).

Types

1 ♂ holotype at UCRC (Curoe 2012).

Chrysina giesberti Monzón, 2010

Chrysina giesberti Monzón, 2010: 1 [original combination].

Distribution

GUATEMALA: Huehuetenango, Quiché (Monzón 2010). MEXICO: Veracruz (Monzón 2010).

Types

1 ♂ holotype and 1 ♀ allotype at UVGC (Monzón 2010); paratypes at UVGC, FSCA, WSU (Monzón 2010); 2 paratypes at MSPC (Fig. 11).

Figure 11.

Chrysina giesberti Monzón paratype male from MSPC. A Dorsal habitus B Lateral habitus C Specimen labels and male genitalia D Male genitalia, lateral view E Male parameres, caudal view.

Chrysina gloriosa (LeConte, 1854)

Plusiotis gloriosa LeConte, 1854: 221–222 [original combination].

Chrysina gloriosa (LeConte) [new combination by Hawks 2001: 8].

Distribution

MEXICO: Chihuahua, Coahuila, Sonora (Coolidge 1911, Ohaus 1918, 1934b, Blackwelder 1944, Cazier 1951, Machatschke 1972, Morón 1990, Hardy 1991, Morón et al. 1997, Thomas et al. 2006). USA: Arizona, New Mexico, Texas (Boucard 1875, Nonfried 1891, Skinner 1911, Casey 1915, Ohaus 1918, 1934b, Leng 1920, Blackwelder 1939, 1944, Cazier 1951, Gibson and Carrillo 1959, Machatschke 1972, Morón 1990, 1991, Hardy 1991, Thomas et al. 2006, Krajcik 2008).

Chrysina gorda Delgado, 2003

Chrysina gorda Delgado, 2003: 319–321 [original combination].

Distribution

MEXICO: Hidalgo, Querétaro, Veracruz (Delgado 2003, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Márquez et al. 2013).

Types

1 ♂ holotype and 1 ♀ allotype at IEXA (Delgado 2003); paratypes at UAEH (Delgado 2003).

Chrysina guatemalensis (Monzón, Cano, & Bailey, 1999)

Plusiotis guatemalensis Monzón, Cano, & Bailey, 1999: 183–184 [original combination].

Chrysina guatemalensis (Monzón, Cano, and Bailey) [new combination by Hawks 2001: 2].

Distribution

GUATEMALA: San Marcos (Monzón et al. 1999, Thomas et al. 2006, Krajcik 2008, Monzón 2010).

Types

1 ♂ holotype and 1 ♀ allotype at UVGC (Monzón et al. 1999); paratypes at UVGC, FSCA, and UNSM (Monzón et al. 1999).

Chrysina guaymi (Curoe, 2001)

Plusiotis guaymi Curoe, 2001: 46–49 [original combination].

Chrysina guaymi (Curoe) [comb. n.].

Distribution

PANAMA: Chiriquí (Curoe 2001; Thomas et al. 2009).

Types

1 ♂ holotype and 1 ♀ allotype at MIUP (Curoe 2001); paratypes at STRI, MNCR, and MXAL (Curoe 2001).

Remarks

The genus Plusiotis was synonymized with Chrysina in 2001 (Hawks 2001), the same year that Curoe described a new species from Panama (Curoe 2001). In Hawks’ (2006) online “Checklist of Chrysina species”, Plusiotis guaymi Curoe is listed as the new combination Chrysina guaymi. Because the online checklist is not considered to be formally published for nomenclatural purposes, we formalize this new combination herein.

Chrysina halffteri (Morón, 1990)

Plusiotis halffteri Morón, 1990: 28 [original combination].

Chrysina halffteri (Morón) [new combination by Hawks 2001: 7].

Distribution

GUATEMALA: Huehuetenango (Thomas et al. 2006). MEXICO: Chiapas (Morón 1990, 1991, Thomas 1993, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype and 1 ♀ allotype MXAL (Morón 1990).

Chrysina howdenorum (Morón, 1990)

Plusiotis howdenorum Morón, 1990: 31–32 [original combination].

Chrysina howdenorum (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Oaxaca (Morón 1990, Krajcik 2008, Thomas et al. 2007).

Types

The following specimens are deposited at CMNC. 1 ♂ holotype, 1 ♀ allotype, and 1 ♂ paratype: “7000’, 32mi. S. Valle Nacional Oax. Mex. V.21-24, 1971 H. Howden//H. & A. Howden Collection//HOLOTIPO//Plusiotishowdenorum Morón M. A. Morón, det. 1987//[barcode matrix] Canadian Museum of Musée canadien de la NATURE CMNEN 00011917”, allotype with identical collecting data label and database number CMNEN 00011918; 1 paratype at MXAL (Morón 1990).

Chrysina intermedia (Ohaus, 1913)

Plusiotis intermedia Ohaus, 1913: 488–489 [original combination].

Chrysina intermedia (Ohaus) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Oaxaca (Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Krajcik 2008, Thomas et al. 2006).

Chrysina karschi (Nonfried, 1891)

Plusiotis karschi Nonfried, 1891: 306 [original combination].

Chrysina karschi (Nonfried) [new combination by Ohaus 1912: 308].

Distribution

GUATEMALA: Baja Verapaz, Zacapa (Monzón 1995, Young 1999, Thomas et al. 2006). HONDURAS: Cortés (Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Monzón 1995, Thomas et al. 2006, Krajcik 2008, Jocque et al. 2013).

Chrysina lacordairei (Boucard, 1875)

Plusiotis lacordairei Boucard, 1875: 122 [original combination].

Chrysina lacordairei (Boucard) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Guerrero, Oaxaca (Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Cano and Morón 1994, Morón et al. 1997, Thomas et al. 2006, Krajcik 2008, Deloya et al. 2014).

Chrysina laniventris (Sturm, 1843)

Pelidnota laniventris Sturm, 1843: 339–340 [original combination].

Plusiotis laniventris (Sturm) [new combination by Burmeister 1844: 420].

Chrysina laniventris (Sturm) [new combination by Hawks 2001: 7].

synonym. Pelidnota latipennis Sturm, 1843

Pelidnota latipennis Sturm, 1843: 338-339 [original combination].

Pelidnota laniventris Sturm [syn. by Burmeister 1844: 420].

synonym. Plusiotis mnizechii Boucard, 1875

Plusiotis mnizechii Boucard, 1875: 124 [original combination].

Plusiotis mniszechii Boucard [incorrect subsequent spelling by Nonfried 1891: 304].

Plusiotis mniszechi Boucard [incorrect subsequent spelling by Ohaus 1918: 16].

Plusiotis laniventris Sturm [syn. by Morón 1990: 23].

Distribution

MEXICO: Distrito Federal, Guerrero, Hidalgo, Jalisco, México, Michoacán, Morelos, Veracruz (Sturm 1843, Burmeister 1844, Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón and Zaragoza 1976, Morón 1990, Deloya et al. 1993, Morón et al. 1997, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Deloya et al. 2014).

Types

1 ♂ lectotype of Pelidnota laniventris at BMNH (Natural History Museum 2014).

Remarks

Boucard (1875) described Plusiotis mnizechii from Mexico and named this species for “Count Georges de Mnizech” of the Polish noble Mniszech family. Nonfried (1891) spelled the name P. mniszechii and Ohaus (1918) further shortened this to P. mniszechi. Ohaus’s (1918) spelling of P. mniszechi Boucard was used for this valid species by all subsequent authors until it was synonymized under P. laniventris (Sturm) (Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990). We think the original spelling of Boucard (1875) as P. mnizechii should be considered correct as there is no evidence of printing error or lapsus calami per ICZN Article 32.5. Indeed, Boucard (1878) referred to this species again as P. mnizechii in a figure plate legend. The spelling of this synonym has created some confusion in the literature (see “Remarks” under Chrysina macropus (Francillon)). If the misspelled name were to be given revalidated species status, it would be at risk of homonymy with Chrysina macropus var. mniszechi H. W. Bates. We stress that correct spelling of this synonym is Plusiotis mnizechii Boucard.

Chrysina lecontei (Horn, 1882)

Plusiotis lecontei Horn, 1882: 120–121 [original combination].

Plusiotina lecontei (Horn) [new combination by Casey 1915: 87].

Plusiotis lecontei Horn [revised combination by Ohaus 1918: 17].

Chrysina lecontei (Horn) [new combination by Hawks 2001: 7].

synonym. Plusiotina aeruginis Casey, 1915

Plusiotina aeruginis Casey, 1915: 85 [original combination].

Plusiotis aeruginis (Casey) [new combination by Ohaus 1934b: 62].

Plusiotis lecontei Horn [syn. by Cazier 1951: 7].

synonym. Plusiotina angusta Casey, 1915

Plusiotina angusta Casey, 1915: 86 [original combination].

Plusiotis angustata (Casey) [new combination and incorrect subsequent spelling by Ohaus 1934b: 62].

Plusiotis lecontei Horn [syn. by Cazier 1951: 7].

synonym. Plusiotina sonorica Casey, 1915

Plusiotina sonorica Casey, 1915: 87 [original combination].

Plusiotis sonorica (Casey) [new combination by Ohaus 1934b: 63].

Plusiotis lecontei Horn [syn. by Cazier 1951: 7].

synonym. Plusiotina subenodis Casey, 1915

Plusiotina subenodis Casey, 1915: 86 [original combination].

Plusiotis subenodis (Casey) [new combination by Ohaus 1934b: 63].

Plusiotis lecontei Horn [syn. by Cazier 1951: 7].

Distribution

MEXICO: Chihuahua, Durango, Sinaloa, Sonora (H. W. Bates 1888, Coolidge 1911, Casey 1915, Ohaus 1918, 1934b, Blackwelder 1939, 1944, Cazier 1951, Machatschke 1972, Morón 1990, Hardy 1991, Blackaller-Bages and Delgado 1994, Thomas et al. 2006, Morón and Nogueira 2016). USA: Arizona, New Mexico (Horn 1882, H. W. Bates 1888, Nonfried 1891, Casey 1915, Leng 1920, Blackwelder 1939, Ohaus 1918, 1934b, Cazier 1951, Gibson and Carrillo 1959, Hardy 1991, Morón 1990, 1991, Morón and Deloya 1991, Blackaller-Bages and Delgado 1994, Thomas et al. 2006, Krajcik 2008, Morón and Nogueira 2016).

Remarks

Krajcik (2008) listed Plusiotis angustata Machatschke (1972) in synonymy with Chrysina lecontei (Horn). This is misleading as P. angustata is not a validly described species but rather is a subsequent misspelling by Ohaus (1934b) of P. angusta Casey. Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis lecontei. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina lecontei.

Chrysina limbata (Rothschild & Jordan, 1894)

Plusiotis limbata Rothschild & Jordan, 1894: 505 [original combination].

Chrysina limbata (Rothschild and Jordan) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: Cartago, Limón, San José (Morón 1990, Thomas et al. 2006, Krajcik 2008, García-López et al. 2013). PANAMA: Chiriquí (Morón 1990, Ratcliffe 2002, Thomas et al. 2006).

Chrysina luteomarginata (Ohaus, 1913)

Plusiotis luteomarginata Ohaus, 1913: 492–493 [original combination].

Chrysina luteomarginata (Ohaus) [new combination by Hawks 2001: 4].

Distribution

COSTA RICA: Alajuela, Cartago (Morón 1990, Monzón 1995, Young 2002, Thomas et al. 2006, García-López et al. 2013). GUATEMALA: Izabal (Monzón 1995, Young 2002). HONDURAS: Cortés (Thomas et al. 2006). NICARAGUA (Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Maes 1987, Morón 1990, Monzón 1995, Hawks 2001, Young 2002, Krajcik 2008). PANAMA: Chiriquí, Veraguas (Morón 1990, Monzón 1995, Ratcliffe 2002, Young 2002, Thomas et al. 2006).

Types

1 ♂ lectotype at ZMHB (Hawks 2001).

Chrysina macropus (Francillon, 1795)

Scarabaeus macropus Francillon, 1795: 1–4 [original combination].

Trichius macropus (Francillon) [new combination by Gyllenhaal 1817: 262].

Chrysina macropus (Francillon) [new combination by Burmeister 1844: 416].

synonym. Chrysina mexicana Gray, 1832

Chrysina mexicana Gray, 1832: 516–517 [original combination].

Chrysina macropus (Francillon) [syn. by Burmeister 1844: 416].

synonym. Chrysina macropus mniszechi H. W. Bates, 1888

Chrysina macropus var. mniszechi H. W. Bates, 1888: 285 [original combination].

Chrysina macropus mniszechi H. W. Bates [new subspecific status by Machatschke 1972: 17].

Chrysina macropus (Francillon) [syn. by Morón 1990: 54].

Distribution

MEXICO: Guerrero, Hidalgo, Oaxaca, Puebla, Querétaro, San Luis Potosí, Veracruz (Burmeister 1844, Gistel 1850, Blanchard 1851, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Gibson and Carrillo 1959, Carrillo et al. 1966, Machatschke 1972, Morón 1985, 1990, 1993, 1994, Morón et al. 1997, Hawks 2001, Thomas et al. 2006, Krajcik 2008, Pacheco Flores et al. 2006, Delgado-Castillo and Márquez 2006, Márquez 2008, Muñoz-Hernández et al. 2008, Delgado-Castillo et al. 2012, Márquez et al. 2013, Deloya et al. 2014).

Remarks

The name “Chrysina henrybatesi” is cited as a synonym of C. macropus on the website checklist of Chrysina species (Hawks 2006). As discussed by Moore and Jameson (2013), this name was proposed in a manuscript version of Hawks (2001), and it was removed before publication. Apparently the manuscript version of the published paper (Hawks 2001) was used for the website checklist. This name is a nomen nudum and should not be listed as a synonym of C. macropus.

The names C. macropus mniszechi H. W. Bates and C. mnizechii Boucard have created some confusion (see “Remarks” under C. laniventris). The spelling of these names differs, but revalidation of species status would risk homonymy.

Chrysina magnistriata (Morón, 1990)

Plusiotis magnistriata Morón 1990: 32 [original combination].

Chrysina magnistriata (Morón) [new combination by Hawks 2001: 7].

Distribution

PANAMA: Chiriquí (Morón 1990, 1991, Ratcliffe 2002, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype and 1 ♀ allotype at MNHN (Morón 1990).

Chrysina marginata (Waterhouse, 1871)

Plusiotis marginata Waterhouse, 1871: 5–6 [original combination].

Chrysina marginata (Waterhouse) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Cartago, Limón, Puntarenas (Morón 1990, Cano and Morón 1994, Hawks 1999, Thomas et al. 2006, Krajcik 2008). PANAMA: Chiriquí (Waterhouse 1871, Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Cano and Morón 1994, Hawks 1999, Ratcliffe 2002, Thomas et al. 2006).

Types

1 ♀ holotype at BMNH (Natural History Museum 2014).

Chrysina miguelangeli Nogueira & Curoe, 2012

Chrysina miguelangeli Nogueira & Curoe, 2012: 2–5 [original combination].

Distribution

MEXICO: Oaxaca (Nogueira and Curoe 2012, Thomas et al. 2014).

Types

1 ♂ holotype and 1 ♀ allotype at MXAL (Nogueira and Curoe 2012); paratypes at MXAL, CNIN (UNAM), UAG and IEXA (Nogueira and Curoe 2012).

Chrysina misteca (Morón, 1990)

Plusiotis misteca Morón, 1990: 37 [original combination].

Chrysina misteca (Morón) [new combination by Hawks 2001: 8].

Distribution

MEXICO: Oaxaca (Morón 1990, Krajcik 2008, Thomas et al. 2010).

Types

The following specimens are deposited at CMNC. 1 ♂ holotype: “MEXICO: Oaxaca Disto. de Yautepec Juquila Mixes VI. 1973 W. Miller//H. & A. Howden Collection//HOLOTIPO//Plusiotismisteca Morón M. A. Morón, det. 1989//[barcode matrix] Canadian Museum of Musée canadien de la NATURE CMNEN 00011919”. 2 paratypes at MXAL (Morón 1990).

Chrysina modesta (Sturm, 1843)

Pelidnota modesta Sturm, 1843: 338 [original combination].

Chrysina macropus var. modesta (Sturm) [new combination and new infrasubspecific status by H. W. Bates 1888: 285].

Chrysina modesta (Sturm) [revised species status by Ohaus 1912: 308–309].

Distribution

MEXICO: México, Michoacán (Sturm 1843, H. W. Bates 1888, Ohaus 1912, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas et al. 2006, Krajcik 2008).

Chrysina moroni (Curoe & Beraud, 1994)

Plusiotis moroni Curoe & Beraud, 1994: 31–33 [original combination].

Chrysina moroni (Curoe and Beraud) [new combination by Hawks 2001: 7].

Distribution

GUATEMALA: San Marcos (Thomas et al. 2006). MEXICO: Chiapas (Morón et al. 1997, Morón-Ríos and Morón 2001, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype and 1 ♀ allotype at CNIN (UNAM) (Curoe and Beraud 1994); paratypes at UVGC and MXAL (Curoe and Beraud 1994).

Chrysina nogueirai (Morón, 1992)

Plusiotis nogueirai Morón, 1992: 62-66 [original combination].

Chrysina nogueirai (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Aguascalientes, Jalisco (Morón 1992, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype, 1 ♀ allotype and paratypes at MXAL (Morón 1992); paratypes at ZMHB (Morón 1992).

Chrysina ofidiodontophallica Curoe, 2011

Chrysina ofidiodontophallica Curoe, 2011: 2–4 [original combination].

Distribution

PANAMA: Darien (Curoe 2011).

Types

1 ♂ holotype at MIUP (Curoe 2011); 1 paratype at MXAL (Curoe 2011).

Chrysina ohausi (Franz, 1928)

Plusiotis ohausi Franz, 1928: 3–5 [original combination].

Chrysina ohausi (Franz) [new combination by Hawks 2001: 8].

Distribution

PANAMA (Franz 1928, Ohaus 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Curoe 2001, Ratcliffe 2002, Krajcik 2008).

Remarks

Chrysina ohausi (Franz) was known only from a single holotype female described from “Panama” and the holotype is apparently lost (Morón 1990; Curoe 2001). It is possible that C. ohausi (Franz) is a junior synonym of C. batesi (Boucard) based on the description of C. ohausi (Franz) and comparisons with other Chrysina species from Panama (Curoe 2001).

Chrysina optima (H. W. Bates, 1888)

Plusiotis optima H. W. Bates, 1888: 279 [original combination].

Chrysina optima (H. W. Bates) [new combination by Hawks 2001: 8].

synonym. Plusiotis melior Rothschild & Jordan, 1894

Plusiotis melior Rothschild & Jordan, 1894: 506 [original combination].

Plusiotis optima var. melior Rothschild and Jordan [new infrasubspecific status by Ohaus 1934b: 64].

Plusiotis optima melior (Rothschild and Jordan) [new subspecific status by Machatschke 1972: 16].

Plusiotis optima H. W. Bates [syn. by Morón 1990: 47].

Distribution

COSTA RICA: Cartago (H. W. Bates 1888, Nonfried 1892, Rothschild and Jordan 1894, H. W. Bates 1888, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas et al. 2006, Krajcik 2008). PANAMA: Chiriquí (Morón 1990, Ratcliffe 2002, Thomas et al. 2006).

Types

1 ♂ holotype of Plusiotis optima at BMNH (Natural History Museum 2014).

Chrysina oreicola (Morón, 1992)

Plusiotis oreicola Morón, 1992: 72–73 [original combination].

Chrysina oreicola (Morón) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: Limón (Morón 1992, Krajcik 2008, Thomas et al. 2013). PANAMA: Chiriquí (Ratcliffe 2002).

Types

1 ♂ holotype at MNCR (Morón 1992); 1 ♀ allotype at MXAL (Morón 1992).

Remarks

Hawks (2001), in reference to species groups in the genus Chrysina, listed C. oreicola as “incertae sedis” (meaning that he was not able to assign the species to a species group). In this usage, Hawks (2001) was not suggesting that the validity of the species was in question.

Chrysina orizabae (H. W. Bates, 1889)

Plusiotis orizabae H. W. Bates, 1889: 410 [original combination].

Chrysina orizabae (H. W. Bates) [new combination by Hawks 2001: 5].

synonym. Plusiotis alticola H. W. Bates, 1889

Plusiotis alticola H. W. Bates, 1889: 409–410 [original combination].

Chrysina orizabae (H. W. Bates) [syn. by Hawks 2001: 5].

Distribution

MEXICO: Colima, Distrito Federal, Hidalgo, Jalisco, México, Morelos, Puebla, Tlaxcala, Veracruz (H. W. Bates 1889, Nonfried 1892, Casey 1915, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Blackaller-Bages and Delgado 1994, Morón et al. 1997, Hawks 2001, Thomas et al. 2006, Krajcik 2008, Morón and Márquez 2012, Márquez et al. 2013, Morón and Nogueira 2016).

Types

1 ♀ holotype of Plusiotis orizabae at BMNH (Fig. 12); 1 ♂ holotype of Pelidnota alticola at BMNH.

Remarks

Krajcik (2012, 2013) erroneously listed C. alticola as a valid name. Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis orizabae. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina orizabae.

Figure 12.

Plusiotis orizabae H. W. Bates (valid name Chrysina orizabae [H. W. Bates]) holotype from BMNH. A Dorsal habitus B Lateral habitus C Specimen labels.

Chrysina pastori (Curoe, 1994)

Plusiotis pastori Curoe, 1994: 37–39 [original combination].

Chrysina pastori (Curoe) [new combination by Hawks 2001: 8].

Distribution

HONDURAS: Cortés (Curoe 1994, Krajcik 2008, Thomas et al. 2007, Jocque et al. 2013).

Types

1 ♂ holotype and 1 ♀ allotype at CAS (Curoe 1994); paratypes at EAPZ, CNIN (UNAM) and MXAL (Curoe 1994).

Chrysina pehlkei (Ohaus, 1930)

Plusiotis pehlkei Ohaus, 1930b: 265–266 [original combination].

Chrysina pehlkei (Ohaus) [new combination by Hawks 2001: 7].

Distribution

GUATEMALA: Chimaltenango, El Quiché, Quetzaltenango, Sacatepéquez, San Marcos, Sololá (Ohaus 1930b, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Blackaller-Bages and Delgado 1994, Thomas et al. 2006, Krajcik 2008, Morón and Nogueira 2016).

Remarks

Chrysina pehlkei was reported from Chiapas and Oaxaca, Mexico (Thomas 1993, Morón et al. 1997), but these data refer to C. rutelidedundeei (pers. comm. from Don Thomas, Aug. 2016). Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis pehlkei. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina pehlkei.

Chrysina peruviana Kirby, 1828

Chrysina peruviana Kirby 1828: 523 [original combination].

Chrysina macropus (Francillon) [syn. by Burmeister 1844: 416].

Chrysina peruviana Kirby [revised species status by Hawks 2001: 2].

synonym. Pelidnota aeruginosa Sturm, 1843

Pelidnota aeruginosa Sturm, 1843: 337 [original combination].

Chrysina amoena (Sturm) [syn. by Burmeister 1844: 417].

synonym. Plusiotis hoegei Boucard, 1895

Plusiotis högei Boucard, 1895: 4–5 [original combination].

Chrysina hoegei Boucard [emendation by Machatschke 1972: 17].

Chrysina amoena (Sturm) [syn. by Ohaus 1912: 308].

synonym. Pelidnota amoena Sturm, 1843

Pelidnota amoena Sturm, 1843: 337–338 [original combination].

Chrysina amoena (Sturm) [new combination by Burmeister 1844: 417].

Chrysina peruviana Kirby [syn. by Hawks 2001: 2].

Distribution

MEXICO: Hidalgo, Puebla, Querétaro, San Luis Potosí, Veracruz (Sturm 1843, Burmeister 1844, H. W. Bates 1888, Ohaus 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Morón et al. 1997, Hawks 2001, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Muñoz-Hernández et al. 2008, Márquez 2008, Márquez and Sierra-Martínez 2009, Márquez et al. 2013).

Types

1 ♀ holotype at OUMNH (Hawks 2001).

Remarks

Krajcik (2012, 2013) erroneously listed C. amoena as a valid name and omitted the name C. peruviana. Chrysina peruviana was reported from Guanajuato, Mexico (Morón and Márquez 2012), but these data refer to C. aurilisternum (Pérez-Flores et al. 2016).

Chrysina plusiotina (Ohaus, 1912)

Pelidnota plusiotina Ohaus, 1912: 304–305 [original combination].

Pelidnotopsis plusiotina (Ohaus) [new combination by Ohaus 1915b: 257].

Chrysina plusiotina (Ohaus) [new combination by Hawks 2001:4].

Pelidnotopsis plusiotina (Ohaus) [revised combination by Soula 2010b: 11].

Chrysina plusiotina (Ohaus) [revised combination by Moore and Jameson 2013: 381].

Distribution

MEXICO: Coahuila, Nuevo León (Ohaus 1912, 1915b, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Hawks 2001, Thomas et al. 2006, Krajcik 2008, Soula 2010b).

Types

1 ♀ lectotype and 1 paralectotype at ZMHB (Hawks 2001).

Remarks

Krajcik (2012, 2013), perhaps erroneously, considered Pelidnotopis plusiotina as valid.

Chrysina prasina (Boucard, 1878)

Plusiotis prasina Boucard, 1878: 295 [original combination].

Chrysina prasina (Boucard) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Guerrero, Hidalgo, Oaxaca, Puebla, Veracruz (Boucard 1878, H. W. Bates 1888, Nonfried 1892, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1993, 1994, Ratcliffe et al. 1992, Morón et al. 1997, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Márquez et al. 2013).

Chrysina prototelica (Morón & Howden, 1992)

Plusiotis prototelica Morón & Howden, 1992: 205–209 [original combination].

Chrysina prototelica (Morón and Howden) [new combination by Hawks 2001: 7].

Distribution

GUATEMALA: Baja Verapáz, Guatemala, Sacatepéquez (Morón 1991, Morón and Howden 1992, Monzón 1995, Krajcik 2008, Thomas et al. 2007).

Types

The following specimens are deposited at CMNC. 1 ♂ holotype: “GUATML. B. Verapaz 5 km S Sn Jeronimo May 24-31 1989 4500’ J. E. Wappes//No match at BM 1990//SEM//HOLOTIPO//Plusiotisprototelica Morón-Howden M. A. Morón, det. 1990//[barcode matrix] Canadian Museum of Musée canadien de la NATURE CMNEN 00000040”, 4 ♂ and 5 ♀ paratypes at CMNC. 1 ♀ allotype at MXAL (Morón and Howden 1992); paratypes at UVGC and MXAL (Morón and Howden 1992).

Chrysina psittacina (Sturm, 1843)

Pelidnota psittacina Sturm, 1843: 340 [original combination].

Plusiotis auripes (Gray) [syn. by Burmeister 1844: 419].

Plusiotis psittacina (Sturm) [revised species status by Boucard 1875: 123].

Chrysina psittacina (Sturm) [new combination and revised application by Hawks 2001: 3].

synonym. Plusiotis amalia Burmeister, 1844

Plusiotis amalia Burmeister, 1844: 422 [original combination].

Plusiotis laeta Sturm [syn. by Boucard 1875: 121].

Plusiotis psittacina Sturm [syn. by H. W. Bates 1888: 281].

synonym. Pelidnota laeta Sturm, 1843

Pelidnota laeta Sturm, 1843: 341 [original combination].

Plusiotis psittacina (Sturm) [syn. by Boucard 1878: 294].

Plusiotis adelaidae (Hope) [syn. by Nonfried 1891: 302].

Plusiotis psittacina var. laeta (Sturm) [syn. by Ohaus 1918: 17].

Distribution

MEXICO: Chiapas, Hidalgo, Nuevo León, San Luis Potosi, Tamaulipas (Sturm 1843, Burmeister 1844, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1981, 1990, Thomas 1993, Hawks 2001, Thomas et al. 2006, Krajcik 2008, Márquez et al. 2013).

Types

1 ♀ lectotype of Pelidnota psittacina at BMNH (Hawks 2001).

Remarks

In reference to species groups, Hawks (2001) listed C. amalia as “incertae sedis” (that is, Hawks was not able to assign the species to a species group). In his usage of “incertae sedis”, Hawks (2001) was not suggesting that the validity of the species was in question. H. W. Bates (1888) noted that P. amalia had uncertain locality data but synonymized the species under Plusiotis psittacina. Morón (1990) maintained the synonymy. We cautiously list C. amalia here in synonymy with Chrysina psittacina until the validity of the species is reevaluated.

Chrysina purpurata (Morón, 1990)

Plusiotis purpurata Morón, 1990: 20–21 [original combination].

Chrysina purpurata (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Guerrero (Morón 1990, Blackaller-Bages and Delgado 1994, Thomas et al. 2006, Krajcik 2008, Morón and Nogueira 2016).

Types

1 ♂ holotype and 1 ♀ allotype at MXAL (Morón 1990); 1 paratype at CNIN (UNAM) (Morón 1990); 1 paratype at ZMHB (Morón 1990).

Remarks

Morón and Nogueira (2016) considered the valid name for this species to be Plusiotis purpurata. Lacking a clearly articulated and evidence-based rationale for this nomenclatural change, we use the name Chrysina purpurata.

Chrysina purulhensis (Warner & Monzón, 1993)

Plusiotis purulhensis Warner & Monzón, 1993: 211–213 [original combination].

Chrysina purulhensis (Warner & Monzón, 1993) [new combination Hawks 2001: 2].

Distribution

BELIZE: Cayo (Warner and Monzón 1993, Thomas et al. 2006). GUATEMALA: Alta Verapáz, Huehuetenango, Quiché (Warner and Monzón 1993, Thomas et al. 2006, Krajcik 2008, Monzón 2010).

Types

1 ♂ holotype at FSCA (Warner and Monzón 1993); 1 ♀ allotype at USNM (Warner and Monzón 1993); paratypes at FSCA and MXAL (Warner and Monzón 1993).

Chrysina quetzalcoatli (Morón, 1990)

Plusiotis quetzalcoatli Morón, 1990: 22 [original combination].

Chrysina quetzalcoatli (Morón) [new combination by Hawks 2001: 7].

Distribution

GUATEMALA: Alta Verapaz, El Quiché, Huehuetenango, Jutiapa, Sacatepéquez, San Marcos (Morón 1990, 1991, Thomas et al. 2006, Jocque et al. 2013). HONDURAS: Comayagua, Cortés (Morón 1991, Thomas et al. 2006, Schlein 2011, Jocque et al. 2013). MEXICO: Chiapas (Morón 1990, 1991, Thomas 1993, Morón-Ríos and Morón 2001, Thomas et al. 2006, Krajcik 2008, Jocque et al. 2013).

Types

1 ♂ holotype, 1 ♀ allotype and 5 paratypes at MXAL (Morón 1990); 1 paratype at MNHN (Morón 1990); 2 paratypes at BMNH (Morón 1990, and BHG pers. obs. Aug. 2016); 2 paratypes at ZMHB (Morón 1990).

Chrysina quiche (Morón, 1990)

Plusiotis quiche Morón, 1990: 41 [original combination].

Chrysina quiche (Morón) [new combination by Hawks 2001: 8].

Distribution

GUATEMALA: Quetzaltenango, San Marco, Zacapa (Morón 1990, Monzón 1995, Cano and Morón 1994, Monzón et al. 1999, Krajcik 2008, Thomas et al. 2009). MEXICO: Chiapas (Morón-Ríos and Morón 2001).

Types

1 ♂ holotype at MXAL (Morón 1990).

Chrysina ratcliffei (Morón, 1990)

Plusiotis ratcliffei Morón, 1990: 44–45 [original combination].

Chrysina ratcliffei (Morón) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Limón, Puntarenas (Morón 1990, Hawks 1999, Thomas et al. 2006). PANAMA: Former Canal Zone, Colón, Darien, Panama (Morón 1990, 1991, Cano and Morón 1994, Hawks 1999, Ratcliffe 2002, Thomas et al. 2006, Krajcik 2008).

Types

The following specimen is deposited at CMNC. 1 ♂ holotype “Panamá: Canal Zone Barro Colorado Is. 9°10'N 79°50'W//5. vi. 1977 H. A. Hespenheide//H. & A. Howden Collection//HOLOTIPO//Plusiotisratcliffei Morón M. A. Morón, det. 1988//[barcode matrix] Canadian Museum of Musée canadien de la NATURE CMNEN 00011920”; 1 ♂ paratype at CMNC (Morón 1990).

Chrysina resplendens (Boucard, 1875)

Plusiotis resplendens Boucard, 1875: 119 [original combination].

Chrysina resplendens (Boucard) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Puntarenas, San José (Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas et al. 2006, Krajcik 2008). PANAMA: Chiriquí (H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Ratcliffe 2002, Thomas et al. 2006).

Chrysina rodriguezi (Boucard, 1878)

Plusiotis rodriguezi Boucard, 1878: 295 [original combination].

Chrysina rodriguezi (Boucard) [new combination by Hawks 2001: 8].

Distribution

GUATEMALA: Alta Verapaz, Baja Verapaz, Guatemala, Huehuetenango, Quetzaltenango, Quiché (Boucard 1878, Dohrn 1883, H. W. Bates 1888, Nonfried 1892, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Cano and Morón 1994, Thomas et al. 2006, Krajcik 2008, Monzón 2010). MEXICO: Chiapas, Guerrero (H. W. Bates 1888, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Thomas 1993, Thomas et al. 2006).

Chrysina rutelidedundeei Soula, 2012

Chrysina rutelidedundeei Soula, 2012: 5–6 [original combination].

Distribution

MEXICO: Chiapas, Oaxaca (Thomas 1993, Morón et al. 1997, Soula 2012, Thomas et al. 2014).

Types

The following specimens are deposited at CCECL. 1 ♂ holotype, 1 ♀ allotype: “Mexique Chiapas San Cristobal VII. 2011//Holotype 2012 Chrysina ebrardi S. Soula//Chrysina rutelidedundeei M. Soula det. 2012 Holotype ♂” (47030025); “Mexique Chiapas San Cristobal VII. 2011// Chrysina rutelidedundeei M. Soula det. 2012 Allotype ♀” (47030026). Genitalia card-mounted underneath the male holotype and female allotype. Chrysina ebrardi is a manuscript name and an invalid label on the male holotype. Box 4618645 SOULA.

Remarks

The unavailable manuscript name Chrysina ebrardi appears on a label underneath the male holotype specimen of C. rutelidedundeei. The distributional data from Thomas (1993) and Morón et al. (1997) were associated with the name C. pehlkei, but these specimens are C. rutelidedundeei (pers. comm. from Don Thomas, Aug. 2016).

Chrysina sallaei (Boucard, 1875)

Plusiotis sallaei Boucard, 1875: 123–124 [original combination].

Plusiotis sallei Boucard [incorrect subsequent spelling by Nonfried 1891: 304].

Chrysina sallaei (Boucard) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Hidalgo, Puebla, Veracruz (Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1993, 1994, Ratcliffe et al. 1992, Morón et al. 1997, Thomas et al. 2006, Delgado-Castillo and Márquez 2006, Krajcik 2008, Márquez 2008, Muñoz-Hernández et al. 2008, Márquez et al. 2013).

Chrysina schusteri (Monzón, Cano, & Bailey, 1999)

Plusiotis schusteri Monzón, Cano, & Bailey, 1999: 183, 184–185 [original combination].

Chrysina schusteri (Monzón, Cano, and Bailey) [new combination by Hawks 2001: 2].

Distribution

GUATEMALA: San Marcos (Monzón et al. 1999, Krajcik 2008, Thomas et al. 2009).

Types

1 ♂ holotype and 1 ♀ allotype at UVGC (Monzón et al. 1999); paratypes at UVGC, FSCA and UNSM (Monzón et al. 1999).

Chrysina sirenicola (Solís & Morón, 1994)

Plusiotis sirenicola Solís & Morón, 1994: 31, 37–40 [original combination].

Chrysina sirenicola (Solís and Morón) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Puntarenas (Solís and Morón 1994, Hawks 1999, Krajcik 2008).

Types

1 ♂ holotype, 1 ♀ allotype and 4 paratypes at MNCR (Solís and Morón 1994); 1 ♂ paratype at CMNC; 3 paratypes at MXAL (Solís and Morón 1994); (Solís and Morón 1994); 2 paratypes at ZMHB (Solís and Morón 1994).

Chrysina spectabilis (Ratcliffe & Jameson, 1992)

Plusiotis spectabilis Ratcliffe & Jameson, 1992: 59–61 [original combination].

Chrysina spectabilis (Ratcliffe and Jameson) [new combination by Hawks 2001: 7].

Distribution

HONDURAS: Cortés (Soula 2006, Thomas et al. 2006, Jocque et al. 2013).

Types

1 ♀ holotype at FMNH (Ratcliffe et al. 1992).

Chrysina strasseni (Ohaus, 1924)

Plusiotis strasseni Ohaus, 1924: 185–186 [original combination].

Chrysina strasseni (Ohaus) [new combination by Hawks 2001: 8].

Distribution

GUATEMALA: Zacapa (Monzón 1995, Young 2002, Thomas et al. 2006, Jocque et al. 2013). HONDURAS: Cortés, Olancho, Yoro (Ohaus 1924, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Young 2002, Thomas et al. 2006, Krajcik 2008, Jocque et al. 2013).

Chrysina tapantina (Morón, 1992)

Plusiotis tapantina Morón, 1992: 70–72 [original combination].

Chrysina tapantina (Morón) [new combination by Hawks 2001: 7].

Distribution

COSTA RICA: Cartago (Morón 1992, Krajcik 2008, Thomas et al. 2013). PANAMA (Thomas et al. 2013).

Types

1 ♀ holotype at MNCR (Morón 1992).

Chrysina taylori (Morón, 1990)

Plusiotis taylori Morón, 1990: 31 [original combination].

Chrysina taylori (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Hidalgo, Veracruz (Morón 1990, 1994, Morón et al. 1997, Delgado-Castillo and Márquez 2006, Krajcik 2008, Thomas et al. 2007, Márquez 2008, Márquez et al. 2013).

Types

1 ♂ holotype, 1 ♀ allotype and 1 paratype at MXAL (Morón 1990); 1 paratype at ZMHB (Morón 1990).

Chrysina tecunumani (Cano & Morón, 1994)

Plusiotis tecunumani Cano & Morón, 1994: 2–8 [original combination].

Chrysina tecunumani (Cano and Morón) [new combination by Hawks 2001: 8].

Distribution

GUATEMALA: El Progreso, Izabal (Cano and Morón 1994, Krajcik 2008, Monzón 2010).

Types

1 ♂ holotype at UVGC (Cano and Morón 1994).

Chrysina terroni (Morón, 1990)

Plusiotis terroni Morón, 1990: 35 [original combination].

Chrysina terroni (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Hidalgo (Morón 1990, 1993, 1994, Delgado-Castillo and Márquez 2006, Krajcik 2008, Thomas et al. 2007, Márquez 2008, Márquez et al. 2013).

Types

1 ♂ holotype and 1 ♀ allotype at MXAL (Morón 1990).

Chrysina transvolcanica (Morón & Nogueira, 2016)

Plusiotis transvolcanica Morón & Nogueira, 2016: 13–15 [original combination].

Chrysina transvolcanica (Morón and Nogueira) [comb. n.].

Distribution

MEXICO: Jalisco, Méxcio, Michoacán, Morelos, Puebla, Tlaxcala (Morón and Nogueira 2016).

Types

1 ♂ holotype and 1 ♀ allotype and paratypes at MXAL (Morón and Nogueira 2016).

Remarks

The genus Plusiotis was synonymized with Chrysina (Hawks 2001). Morón and Nogueira (2016) continued using Plusiotis for reasons of practicality and lack of published molecular evidence in support of Hawks’s hypothesis. Because Morón and Nogueira (2016) did not revalidate the genus Plusiotis, we transfer Plusiotis transvolcanica to the currently valid genus Chrysina.

Chrysina tricolor (Ohaus, 1922)

Plusiotis tricolor Ohaus, 1922: 323 [original combination].

Chrysina tricolor (Ohaus) [new combination by Hawks 2001: 8].

Distribution

COSTA RICA: Cartago, San José (Ohaus 1922, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Cano and Morón 1994, Krajcik 2008, Thomas et al. 2007).

Chrysina triumphalis Morón, 1990

Chrysina triumphalis Morón, 1990: 54–55 [original combination].

Distribution

GUATEMALA: San Marcos (Young 2002, Thomas et al. 2006). MEXICO: Chiapas (Morón 1990, Thomas 1993, Morón-Ríos and Morón 2001, Young 2002, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype, 1 ♀ allotype and 1 paratype at MXAL (Morón 1990).

Chrysina tuerckheimi (Ohaus, 1913)

Plusiotis türckheimi Ohaus, 1913: 491–492 [original combination].

Plusiotis tuerckheimi Ohaus [justified emendation by Machatschke 1972: 14].

Chrysina turckheimi (Ohaus) [new combination and incorrect subsequent spelling by Hawks 2001: 8].

Distribution

GUATEMALA: San Marcos (Monzón et al. 1999, Thomas et al. 2006). MEXICO: Chiapas (Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas 1993, Cano and Morón 1994, Thomas et al. 2006, Krajcik 2008).

Chrysina veraguana (Ohaus, 1922)

Plusiotis veraguana Ohaus, 1922: 324 [original combination].

Chrysina veraguana (Ohaus) [new combination by Hawks 2001: 8].

Distribution

PANAMA: Veraguas (Ohaus 1922, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Cano and Morón 1994, Ratcliffe 2002, Krajcik 2008).

Remarks

In reference to species groups, Hawks (2001) listed C. veraguana as “incertae sedis” (that is, Hawks was not able to assign the species to a species group). In this usage, Hawks (2001) was not suggesting that the validity of the species was in question.

Chrysina victorina (Hope, 1841)

Pelidnota victorina Hope, 1841: 147 [original combination].

Plusiotis victorina (Hope) [new combination by Burmeister 1844: 418].

Chrysina victorina (Hope) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Chiapas, Oaxaca, Veracruz (Hope 1841, Burmeister 1844, Blanchard 1851, Boucard 1875, H. W. Bates 1888, Nonfried 1891, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, Thomas 1993, Morón et al. 1997, Thomas et al. 2006, Krajcik 2008).

Chrysina wolfi (Ohaus, 1912)

Plusiotis wolfi Ohaus, 1912: 305 [original combination].

Chrysina wolfi (Ohaus) [new combination by Hawks 2001: 8].

Distribution

ECUADOR: Manabí, Pichincha (Ohaus 1912, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Morón 1990, 1991, Arnaud 1994, Paucar-Cabrera 2005, Krajcik 2008).

Chrysina woodi (Horn, 1884)

Plusiotis woodi Horn, 1884: xxxi [original combination].

Plusiotis woodii Horn [incorrect subsequent spelling by Horn 1885: 124].

Chrysina woodii (Horn) [new combination by Hawks 2001: 8].

Chrysina woodi (Horn) [suggested correct spelling by Moore and Jameson 2013: 383].

Distribution

MEXICO: Chihuahua (H. W. Bates 1888, Blackwelder 1944, Cazier 1951, Hardy 1991, Thomas et al. 2006). USA: Texas (Horn 1884, 1885, H. W. Bates 1888, Nonfried 1892, Skinner 1911, Casey 1915, Ohaus 1918, 1934b, Leng 1920, Blackwelder 1939, 1944, Cazier 1951, Machatschke 1972, Morón 1990, 1991, Hardy 1991, Thomas et al. 2006, Krajcik 2008).

Chrysina xalixteca (Morón, 1992)

Plusiotis xalixteca Morón, 1992: 66–70 [original combination].

Chrysina xalixteca (Morón) [new combination by Hawks 2001: 8].

Distribution

MEXICO: Jalisco (Morón 1992, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype, 1 ♀ allotype and paratypes at MXAL (Morón 1992); paratypes at ZMHB (Morón 1992); paratypes at CNIN (UNAM) and ZMHB (Morón 1992).

Chrysina zapoteca (Morón, 1990)

Plusiotis zapoteca Morón, 1990: 39–40 [original combination].

Chrysina zapoteca (Morón) [new combination by Hawks 2001: 7].

Distribution

MEXICO: Oaxaca (Morón 1990, Cano and Morón 1994, Thomas et al. 2006, Krajcik 2008).

Types

1 ♂ holotype and 1 paratype at MXAL (Morón 1990). 1 ♀ allotype at CMNC: “20mi. S. Juchatengo 6000’, Oax. Rt. 131, Mex. V.27-30, 1971 H. F. Howden//H. & A. Howden Collection//ALLOTYPE//ALLOTIPO//Plusiotiszapoteca Morón M. A. Morón, det. 1988//[barcode matrix] Canadian Museum of Musée canadien de la NATURE CMNEN 00011921”.

CHRYSOPHORA Dejean, 1821

Chrysophora Dejean, 1821: 60.

Type species

Melolontha chrysochlora Latreille, 1812: 131, by monotypy.

Gender

Feminine.

Types

1 species (Fig. 13).

Figure 13.

Chrysophora chrysochlora (Latreille) male exemplar specimen from MSPC. A Dorsal habitus B Lateral habitus C Male genitalia, lateral view D Parameres, dorsal view.

Chrysophora chrysochlora (Latreille, 1812)

Melolontha chrysochlora Latreille, 1812: 131 [original combination].

Chrysophora chrysochlora (Latreille) [new combination by Dejean 1821: 60].

Distribution

COLOMBIA: Antioquia, Boyacá, Caquetá, Cauca, Cesar, Chocó, Cundinamarca, Huila, Meta, Nariño, Norte de Santander, Tolima, Valle del Cauca (Guérin-Méneville 1834, Gistel 1850, Nonfried 1891, Gibson and Carrillo 1959, Morón 1990, Restrepo et al. 2003, Neita-Moreno et al. 2006, Pardo-Locarno and Morón 2007, Neita-Moreno 2011, López-García et al. 2015). ECUADOR: Los Ríos (FSCA), Esmeraldas, Guayas, Loja, Morona-Santiago, Napo, Pastaza, Pichincha, Sucumbíos (Blackwelder 1944, Ohaus 1903, 1918, 1934b, 1952, Machatschke 1972, Morón 1990, Paucar-Cabrera 2005, López-García et al. 2015). PERU: Huánuco (FSCA), Junín (FSCA), San Martín (FSCA), Loreto (Latreille 1812, Germar 1815, LePeletier and Serville 1828, Laporte 1840, Blackwelder 1944, Ohaus 1934b, 1952, Machatschke 1972, Morón 1990, Krajcik 2008, Ratcliffe et al. 2015, López-García et al. 2015).

ECTINOPLECTRON Ohaus, 1915

Ectinoplectron Ohaus, 1915b: 257.

Type species

Homonyx oryctoides Ohaus, 1905: 314–315, by monotypy.

Gender

Neuter.

Species

1 species (Fig. 14).

Figure 14.

Ectinoplectron oryctoides (Ohaus) male exemplar specimen from MSPC. A Dorsal habitus B Lateral habitus C Male genitalia, lateral view D Parameres, dorsal view.

Ectinoplectron oryctoides (Ohaus, 1905)

Homonyx oryctoides Ohaus, 1905: 314–315 [original combination].

Ectinoplectron oryctoides (Ohaus) [new combination by Ohaus 1915b: 257].

synonym. Pelidnota howdeni Hardy, 1975

Pelidnota howdeni Hardy, 1975: 6, 14-15 [original combination].

Ectinoplectron oryctoides (Ohaus) [syn. by Morón et al. 1997: 26–27].

Distribution

MEXICO: Chihuahua, Durango, Jalisco, Michoacán, Nayarit, Sinaloa, Sonora (Ohaus 1905, 1915b, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Hardy 1975, Morón 1990, Morón et al. 1997, 1998, Bustos-Santana and Rivera-Cervantes 2003, Krajcik 2008, Soula 2009, Lugo et al. 2011, Morón and Márquez 2012, Zamora-Vuelvas et al. 2014).

Types

1 ♂ lectotype of Homonyx oryctoides at ZMHB (Soula 2009); 1 ♂ Pelidnota howdeni paratype at CMNC.

EPICHALCOPLETHIS F. Bates, 1904

Epichalcoplethis F. Bates, 1904: 253, 272–273.

Chalcoplethis Burmeister, 1844 [new synonym by Ohaus 1915b: 258].

Epichalcoplethis F. Bates [revised genus status by Soula 2006: 101].

Type species

Pelidnota velutipes Arrow, 1900: 179, by monotypy.

Gender

Feminine.

Species

16 species and subspecies.

Remarks

Krajcik (2012, 2013) considered Epichalcoplethis to be a junior synonym of Pelidnota.

Epichalcoplethis aciculata (F. Bates, 1904)

Pelidnota aciculata F. Bates, 1904: 254, 261 [original combination].

Pelidnota (Chalcoplethis) aciculata F. Bates [new subgeneric combination by Ohaus 1918: 29].

Epichalcoplethis aciculata (F. Bates) [new combination by Soula 2006: 106–107].

Distribution

BOLIVIA: Santa Cruz (WBWC). BRAZIL: Amazonas, Pará (INPA). FRENCH GUIANA: Cayenne, St.-Laurent du Maroni (F. Bates 1904, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Krajcik 2008, Soula 2006, 2010c).

Types

1 ♂ holotype at IRSNB (Soula 2006). An exemplar specimen is figured (Fig. 15).

Figure 15.

Epichalcoplethis aciculata (F. Bates), male specimen. A Dorsal habitus B Lateral habitus C Male genitalia, lateral view D Male genitalia, dorsal view.

Epichalcoplethis benjamini Bouchard & Soula, 2006

Epichalcoplethis benjamini Bouchard & Soula, 2006: 102, 107 [original combination].

Distribution

BOLIVIA: La Paz (Soula 2006). PERU (Soula 2006, Ratcliffe et al. 2015).

Types

The following specimens are deposited at CCECL. 1 ♂ holotype, 1 ♀ allotype, 1 ♂ paratype: “Coll. G. Lecourt Ixiamas 11/91 310 m. Bolivie//Holotype Epichalcoplethis benjamini 2006 S. Soula” (47030043); “Coll. G. Lecourt Ixiamas 11/91 310 m. Bolivie//Allotype Epichalcoplethis benjamini 2006 S. Soula” (47030044); “Coll. G. Lecourt Ixiamas 11/91 310 m. Bolivie//Paratype Epichalcoplethis benjamini 2006 S. Soula” (47030045). Genitalia card-mounted underneath male specimens. Box 4618648 SOULA.

Figure 16.

Epichalcoplethis gilletti Soula holotype male from BMNH. A Dorsal habitus B Lateral habitus C Specimen labels.

Epichalcoplethis blancoi Soula, 2006

Epichalcoplethis blancoi Soula, 2006: 102, 109 [original combination].

Distribution

VENEZUELA: Bolívar, Miranda (Soula 2006).

Types

The following specimens are deposited at CCECL. 1 ♀ holotype, 2 ♀ paratypes: “Camp. Minero Payapal Rio Yuruan//Exp. Instituto Zool. Agricola//Venezuela Bolivar//El Dorado 190 m 23-30-V-87//Holotype Epichalcoplethis blancoi S. 2006 Soula” (47030055); “En la luz//VENEZUELA: Bolívar Guri 200 m 27-vi- al 6-vii-1998 L. J. Joly; J. L. García; Y. Zavala//Paratype Epichalcoplethis blancoi S. 2006 Soula” (47030056); “VENEZUELA: Miranda Tacarigua de Manporal 10°22’32”N 66°12’10”W 23-v-1998 Col. O. Hernández S.//Paratype 2006 Epichalcoplethis blancoi S. Soula Soula” (47030057). This is the entire series and it is noted in Soula (2006) that they are from the MIZA Collection. Box 4618648 SOULA.

Epichalcoplethis chamaeleon (Herbst, 1789)

Scarabaeus chamaeleon Herbst, 1789: 247–248 [original combination].

Pelidnota chamaeleon (Herbst) [new combination by Burmeister 1844: 407].

Pelidnota ignita var. chamaeleon (Herbst) [new infrasubspecific status by F. Bates 1904: 259].

Pelidnota (Chalcoplethis) chamaeleon (Herbst) [new subgeneric combination and revised species status by Ohaus 1918: 29].

Epichalcoplethis chamaeleon (Herbst) [new combination by Soula 2006: 108].

synonym. Pelidnota equestris Laporte, 1840

Pelidnota equestris Laporte, 1840: 122 [original combination].

Pelidnota ignita (Olivier) [syn. by Burmeister 1844: 408].

Chalcoplethis chamaeleon (Herbst) [syn. by Ohaus 1918: 29].

Epichalcoplethis chamaeleon (Herbst) [syn. by Soula 2006: 108].

synonym. Cetonia ignita Olivier, 1789

Cetonia ignita Olivier, 1789: 69–70 [original combination].

Rutela ignita (Olivier) [new combination by Schönherr 1817: 150].

Pelidnota ignita (Olivier) [new combination by Burmeister 1844: 407].

Chalcoplethis chamaeleon var. ignita (Olivier) [new combination and new infrasubspecific status by Ohaus 1918: 29].

Chalcoplethis chamaeleon forma ignita (Olivier) [revised infrasubspecific status by Machatschke 1972: 32].

Epichalcoplethis chamaeleon (Herbst) [syn. by Soula 2006: 108].

Distribution

BRAZIL: Amazonas, Roraima (INPA) (Laporte 1840, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972). COLOMBIA: Meta (Burmeister 1844, Blanchard 1851, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Restrepo et al. 2003, Krajcik 2008). FRENCH GUIANA: Cayenne, St.-Laurent du Maroni (Olivier 1789, Burmeister 1844, Ohaus 1918, Machatschke 1972, Krajcik 2008, Soula 2010c). GUYANA: Demerara-Mahaica (MNRJ) (Ohaus 1918). SURINAME (Olivier 1789, Ohaus 1934b, Machatschke 1972). TRINIDAD AND TOBAGO: Trinidad (Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Soula 2006). VENEZUELA (Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Soula 2006).

Remarks

Voet (1769) is often cited as the author for this species. However, names in Voet’s Catalogus Systematicus Coleopterorum (1778 and subsequent editions) are not consistently binomial (ICZN, Art. 11.4). As such, all of Voet’s names are rejected (Löbl and Smetana 2011). Soula (2006) credited Herbst (1769) as the author of the species.

Epichalcoplethis gilletti Soula, 2010

Epichalcoplethis gilletti Soula, 2010a: 48–49 [original combination].

Distribution

ECUADOR: Pastaza (Soula 2010a). PERU (Soula 2010a, Ratcliffe et al. 2015).

Remarks

According to Soula (2010a), the type specimen of this species should be conserved in the Soula Collection (CCECL). We located this holotype specimen (Fig. 16) at BMNH with the following data: “sara / yagu [indecipherable word] / [blue line] 80.14 // [printed and handwritten] Ohaus determin. 1913 / Pelidnotaaciculata F. Bates / sbsp. nov. // [red label] [printed and handwritten] Holotype / Epichalcoplethis / gilletti Sou / 2010 Soula”.

Epichalcoplethis ledezmaae Bouchard & Soula, 2006

Epichalcoplethis ledezmaae Bouchard & Soula, 2006: 102, 104–105 [original combination].

Distribution

BOLIVIA: Santa Cruz (Soula 2006).

Types

The following specimen is deposited at CCECL. 1 ♂ holotype: “Rte de Camiri à Sta. Cruz Bol. coll. – SOULA //Holotype Epichalcoplethis ledezmaae S. 2006 Soula” (47030042). The genitalia are card-mounted underneath this male specimen. Box 4618648 SOULA.

Epichalcoplethis monzoni Soula, 2006

Epichalcoplethis monzoni Soula, 2006: 102, 112 [original combination].

Distribution

BELIZE: Cayo (Soula 2006). GUATEMALA: Izabal, Petén (Soula 2006).

Types

The holotype of Epichalcoplethis monzoni is deposited at UVGC (Soula 2006). The following specimens are deposited at CCECL. 11 ♂ paratypes, 2 ♀ paratypes: “Finca Firmeza, Sierra de Caral, Morales, Izabal, Guatemala, 450m, 20/V/2006//Paratype 2006 Epichalcoplethis monzoni S. Soula” (47030047 to 47030053); five paratypes with identical label data: “Finca Firmeza 20/V/2006 Sierra de Caral, 450 m Morales - Izabal, GUATEMALA José Monzon leg.//Paratype 2006 Epichalcoplethis monzoni S. Soula” (47030958 to 47030962); “Chiquibul Forest Reserve, Cayo, Belize, VI/2006//Paratype 2006 Epichalcoplethis monzoni S. Soula” (47030054). All male specimens with genitalia card-mounted. Box 4618648 SOULA and 4616345 PORION. The following specimen is deposited at CMNC: 1 ♂ paratype: “GUATEMALA. Izabal Morales. Finca Firmeza Sierra de Caral, 45 msnm 450m. 20 V 2006 José Monzón Coll. COLLECCION J. MONZON//Paratype 2006 Epichalcoplethis monzoni S. Soula”. The following paratype is deposited at BMNH: “Belize (Cayo) / Chiquibul Forest Reserve / Las Cuevas Research Station / 16°44'N 88°59'W / June 2006 / BMNH {E} 2006-141 / C. Gillet & J. Kitson // Paratype 2006 / Epichalcoplethis / monzoni S. / Soula”.

Epichalcoplethis navarropoloi Soula, 2011

Epichalcoplethis navarropoloi Soula, 2011: 73 [original combination].

Distribution

ECUADOR: Pastaza (Soula 2011).

Types

The holotype ♂ is deposited at the Malý collection (Soula 2011).

Epichalcoplethis porioni Soula, 2010

Epichalcoplethis porioni Soula, 2010a: 48 [original combination].

Distribution

HONDURAS: Lempira (Soula 2010a).

Types

The following specimen is deposited at CCECL. 1 ♂ holotype: “HONDURAS-LEMPIRA Montana de Celaque AOUT 1995 Thierry PORION Leg//Coll. TH. PORION//Holotype 2009 Epichalcoplethis porioni S. Soula” (47030955). Genitalia card-mounted underneath the male holotype. Box 4616345 PORION.

Epichalcoplethis richteri (Ohaus, 1910)

Pelidnota richteri Ohaus, 1910a: 186 [original combination].

Pelidnota (Chalcoplethis) richteri Ohaus [new subgeneric combination by Ohaus 1918: 29].

Epichalcoplethis richteri (Ohaus) [new combination by Soula 2006: 103].

Distribution

BRAZIL: Mato Gross do Sul (Ohaus 1910a, 1918, 1934b; Blackwelder 1944; Machatschke 1972; Krajcik 2008). PARAGUAY: Alto Paraguay (Ohaus 1910a; Soula 2006).

Types

1 type specimen of Pelidnota richteri at MLPA.

Epichalcoplethis santistebani Bouchard & Soula, 2006

Epichalcoplethis santistebani Bouchard & Soula, 2006: 102, 105 [original combination].

Distribution

PERU: Huánuco (Soula 2006, Ratcliffe et al. 2015).

Types

The following specimen is deposited at CCECL (Fig. 17). 1 ♂ holotype: “Huanuco Pérou VII/2000 M. SOULA det. 19//Holotype 2006 Epichalcoplethis santistebani Sou. Soula” (47030046). Genitalia card-mounted underneath specimen. Box 4618648 SOULA.

Figure 17.

Epichalcoplethis santistebani Bouchard and Soula holotype male from CCECL. A Dorsal habitus B Male genitalia, dorsal view C Specimen labels.

Epichalcoplethis sanctijacobi (Ohaus, 1905)

Pelidnota sanctijacobi Ohaus, 1905: 318 [original combination].

Pelidnota (Chalcoplethis) sanctijacobi Ohaus [new subgeneric combination by Ohaus 1918: 29].

Epichalcoplethis sanctijacobi (Ohaus) [new combination by Soula 2006: 103–104].

Distribution

ARGENTINA: Córdoba, Salta, Santiago del Estero, Tucumán (Ohaus 1905, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Soula 2006, Krajcik 2008).BRAZIL (Soula 2006). FRENCH GUIANA: Mana (Gruner 1971). PARAGUAY (Soula 2006). URUGUAY (Soula 2006).

Types

1 ♂ type specimen of Pelidnota sanctijacobi at ZMHB (Fig. 18); 1 type specimen at MLPA; 1 type specimen at SDEI; 1 ♂ paralectotype at NHMB (Soula 2006) (see “Type Specimens and Lectotype Designation” in Methods).

Figure 18.

Pelidnota sanctijacobi Ohaus (valid name Epichalcoplethis sanctijacobi [Ohaus]) type male from ZMHB. A Dorsal habitus B Lateral habitus C Specimen labels, mouthparts, and male genitalia D Male genitalia, lateral view E Male parameres, dorsal view.

Epichalcoplethis schiffleri Bouchard & Soula, 2006

Epichalcoplethis schiffleri Bouchard & Soula, 2006: 102, 107–108 [original combination].

Distribution

PERU: Loreto, Piura (Soula 2006, Ratcliffe et al. 2015).

Types

The following specimens are deposited at CCECL. 1 ♂ holotype, 4 ♂ paratypes: “Iquitos, Loreto Pérou; VIII/2003//Holotype 2006 Epichalcoplethis schiffleri S. Soula” (47030027); “Iquitos, Loreto Pérou; VIII/2003//Paratype 2005 Epichalcoplethis schiffleri S. Soula” (47030028); “Iquitos 100 m 9.03 Loreto/PERU//Paratype 2005 Epichalcoplethis schiffleri S. Soula” (47030030); “Yamamono River Iquitos (P) 6/88// Paratype 2005 Epichalcoplethis schiffleri S. Soula” (47030031); “Carbajal, Rio Itaya Piura, Pérou, IX/2005// Paratype 2005 Epichalcoplethis schiffleri S. Soula” (47030029). The Yamamono River locality does not appear in the Soula (2006) description. All 5 specimens have their genitalia card-mounted. Box 4618648 SOULA.

Epichalcoplethis seriatopunctata (Ohaus, 1912)

Pelidnota seriatopunctata Ohaus, 1912: 304 [original combination].

Pelidnota (Chalcoplethis) seriatopunctata Ohaus [new subgeneric combination by Ohaus 1918: 29].

Epichalcoplethis seriatopunctata (Ohaus) [new combination by Soula 2006: 102–103].

Distribution

BRAZIL (Ohaus 1912, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Soula 2006, Krajcik 2008).

Epichalcoplethis velutipes romeroi Soula, 2006

Epichalcoplethis velutipes romeroi Soula, 2006: 111–112 [original combination].

Distribution

VENEZUELA: Bolívar (Soula 2006).

Types

The following specimens are deposited at CCECL. 1 ♂ holotype (Fig. 19A, B, D), 1 ♂ invalid holotype, 1 ♀ allotype (Fig. 19C, E), 3 ♂ paratypes, 4 ♀ paratypes: “Rio Cauja Bolivar coll. – SOULA//Holotype 2005 Epichalcoplethis velutipes romeroi S. Soula” (47030032); Jabillal Rio Caura (Bolivar) coll – SOULA [obverse] 03/94 //Allotype 2005 Epichalcoplethis velutipes romeroi S. Soula” (47030033); “N. Venezuela S. Klages 1904//Not valid Holotype probable paratype det. M. L. Jameson 2014//Holotype Epichalcoplethis romeroi Sou. Soula det. 2005” (47030034); “Rio Cauja (Bolivar) coll. – SOULA[obverse] 07/87 Venez. // Pel. (Chalcoplethis) velutipes (Arrow)//Paratype Epichalcoplethis velutipes romeroi 2005 Soula” (47030035); “N. Venezuela S. Klages 1904//Pel. (Chalcoplethis) velutipes (Arrow)//Museum Paris ex Coll. R. Oberthur// Paratype Epichalcoplethis velutipes romeroi 2005 Soula” (47030036); “P. N. Henri Pittier Choroni ; Venezuela V-VI/2005//Pel. (Chalcoplethis) velutipes (Arrow)//Paratype Epichalcoplethis velutipes romeroi S. 2005 Soula” (47030037); “Camp. minero Payapal Rio Yuruan//Exp. Instituto Zool. Agricola//Venezuela Bolivar//El Dorado 190 m 23-30-V-87 //Paratype 2005 Chalcoplethis velutipes romeroi S. Soula” (47030038); “Rio Cauja (Bolivar) coll. – SOULA [obverse] 07/87 //Pel. (Chalcoplethis) velutipes (Arrow)//Paratype 2005 Epichalcoplethis velutipes romeroi S. Soula” (47030039); “Choroni 200 m V/1998 M. SOULA det. 19 [obverse] Aragua Venezuela (Chez Romero) // Paratype 2005 Epichalcoplethis velutipes romeroi S. Soula” (47030040); “VENEZUELA: Bolívar Guri 200 m 27-vi-al 6-vii-1998 L. J. Joly; J. L. García; Y. Zavala//Paratype 2005 Epichalcoplethis velutipes romeroi S. Soula” (47030041). Genitalia card-mounted underneath the male holotype, the invalid male holotype, three male paratypes, and one female paratype (47030036, 47030037, 47030041, and 47030039). Box 4618648 SOULA.

Remarks

The male holotype specimen labeled from “N. Venezuela” is in fact not the true holotype specimen according to Soula (2006). We labeled this specimen as a probable paratype.

Figure 19.

Epichalcoplethis velutipes romeroi Soula holotype male and allotype female from CCECL. A Dorsal habitus, holotype B Specimens labels, holotype C Specimen labels, allotype D Male genitalia dorsal view, holotype E Dorsal habitus, allotype.

Epichalcoplethis velutipes velutipes (Arrow, 1900)

Pelidnota velutipes Arrow, 1900: 179 [original combination].

Epichalcoplethis velutipes (Arrow) [new combination by F. Bates 1904: 253, 272–273].

Pelidnota (Chalcoplethis) velutipes Arrow [revised combination and new subgeneric combination by Ohaus 1918: 29].

Epichalcoplethis velutipes (Arrow) [new combination by Soula 2006: 109–111].

Distribution

GRENADA (Leng and Mutchler 1914, Machatschke 1972, Hardy 1975, Chalumeau 1985, Soula 2006, Krajcik 2008, Peck 2010, 2016). GUATEMALA: Petén (Hardy 1975, Chalumeau 1985, Soula 2006). HONDURAS: Atlántida (Hardy 1975, Chalumeau 1985, Soula 2006). MEXICO: Chiapas (Palacios-Rios et al. 1990, Thomas 1993, Soula 2006). ST. VINCENT AND THE GRENADINES: St. Vincent (Ohaus 1918, 1934b, Chalumeau 1985, Soula 2006, Peck 2010, 2016). TRINIDAD AND TOBAGO: Trinidad, Tobago (Ohaus 1918, 1934b, Chalumeau 1985, Peck et al. 2002, Peck 2016). VENEZUELA (Ohaus 1918, 1934b, Chalumeau 1985, Soula 2006, Peck 2010, 2016).

Types

1 ♂ type at at BMNH (Hardy 1975).

Eremophygus Ohaus, 1910

Eremophygus Ohaus, 1910c: 21–22.

synonym. Heterocallichloris Gutíerrez, 1951

Heterocallichloris Gutíerrez 1951: 112–114. [Type species. Heterocallichloris bicolor Gutiérrez, 1951 by original designation].

Platycoelia Dejean [syn. by Machatschke 1965: 55].

Eremophygus Ohaus [syn. by Smith and Jameson 2001: 105].

Type species

Eremophygus philippii Ohaus, 1910c: 22, by monotypy.

Gender

Masculine.

Species

6 species.

Eremophygus bicolor (Gutiérrez, 1951)

Heterocallichloris bicolor Gutiérrez, 1951: 112, 114 [original combination].

Platycoelia bicolor (Gutiérrez) [new combination by Machatschke 1965: 60].

Eremophygus bicolor (Gutiérrez) [new combination by Smith and Jameson 2001: 105].

Distribution

BOLIVIA (Smith and Jameson 2001).

Remarks

Heterocallichloris bicolor was originally described in the ruteline tribe Anoplognathini (subtribe Platycoeliina). As a result of a broad analysis of the Platycoeliina (Smith 2003), the species was transferred to the genus Eremophygus (Smith and Jameson 2001).

Eremophygus calvus Gutiérrez, 1952

Eremophygus calvus Gutiérrez, 1952: 223–224 [original combination].

Distribution

BOLIVIA: La Paz (Gutiérrez 1952, Machatschke 1972).

Types

Holotype ♀ of Eremophygus calvus Gutiérrez at UCCC (Fig. 20).

Figure 20.

Holotype female of Eremophygus calvus Gutiérrez from UCCC. A Dorsal habitus B Specimen labels.

Eremophygus lasiocalinus Ohaus, 1915

Eremophygus lasiocalinus Ohaus, 1915a: 76–77 [original combination].

Distribution

BOLIVIA: La Paz (Ohaus 1915a, 1918, 1934b, Blackwelder 1944, Gutiérrez 1949, 1950, Machatschke 1972, Ferrú and Elgueta 2011). CHILE: Arica and Parinacota (Gutiérrez 1949, 1950; Machatschke 1972, Ferrú and Elgueta 2011).

Types

Holotype ♂ of Eremophygus lasiocalinus at ZMHB (Fig. 21).

Figure 21.

Eremophygus lasiocalinus Ohaus holotype male from ZMHB. A Dorsal habitus B Lateral habitus C Specimen labels, mouthparts, and male genitalia D Male genitalia, lateral view E Parameres, dorsal view.

Eremophygus leo Gutiérrez, 1951

Eremophygus leo Gutiérrez, 1951: 106 [original combination].

Distribution

ARGENTINA: Jujuy (Gutiérrez 1951, Machatschke 1972).

Types

Holotype ♂ of Eremophygus leo Gutiérrez at UCCC (Fig. 22).

Figure 22.

Holotype male of Eremophygus leo Gutiérrez from UCCC. A Dorsal habitus B Specimen labels, mouthparts, and male genitalia.

Eremophygus pachyloides Ohaus, 1925

Eremophygus pachyloides Ohaus, 1925: 76 [original combination].

Distribution

BOLIVIA (Ohaus 1925, 1934b, Blackwelder 1944, Gutiérrez 1949, Machatschke 1972).

Types

Holotype ♀ of Eremophygus pachyloides at ZMHB (Fig. 23).

Figure 23.

Eremophygus pachyloides Ohaus holotype female from ZMHB. A Dorsal habitus B Lateral habitus C Specimen labels, mouthparts, and egg.

Eremophygus philippii Ohaus, 1910

Eremophygus philippii Ohaus, 1910c: 22 [original combination].

Distribution

CHILE: Arica and Parinacota; Tarapacá (Ohaus 1910c, 1918, 1934b, Blackwelder 1944, Gutiérrez 1949, 1950, Machatschke 1972). PERU (Ohaus 1952, Ratcliffe et al. 2015).

Types

Holotype ♂ of Eremophygus philippii at ZMHB (Fig. 24).

Figure 24.

Eremophygus philippii Ohaus holotype male from ZMHB. A Dorsal habitus B Lateral habitus C Parameres, dorsal view D Male genitalia, lateral view E Specimen labels.

HOMEOCHLOROTA Soula, 2006

Homeochlorota Soula, 2006: 148–149.

Type species

Pseudochlorota chiriquina Ohaus, 1905: 306-307, by monotypy.

Gender

Feminine.

Figure 25.

Homeochlorota chiriquina (Ohaus) male from DBPC. A Dorsal habitus B Lateral habitus C Male genitalia, lateral view D Male parameres, dorsal view.

Species

1 species.

Remarks

Krajcik (2012, 2013) considered Homeochlorota to be a synonym of Pseudochlorota. Because the rationale for this nomenclatural change was not provided, we use the name Homeochlorota.

Homeochlorota chiriquina (Ohaus, 1905)

Pseudochlorota chiriquina Ohaus, 1905: 306–307 [original combination].

Homeochlorota chiriquina (Ohaus) [new combination by Soula 2006: 149–150].

Distribution

COSTA RICA: Guanacaste (Soula 2006). PANAMA: Chiriquí (Ohaus 1905, 1918, 1934b, Machatschke 1972, Soula 2006, Krajcik 2008).

Types

Lectotype male of Pseudochlorota chiriquina at ZMHB labeled: “Panama, V.d. Chiriqui”; “typus!” (red label, typed); male genitalia card mounted; “Pseudochlorota chiriquina Ohaus” (red label, handwritten). Paralectotype female at ZMHB labeled as lectotype with mouthparts card mounted. An exemplar specimen is shown in Fig. 25.

Remarks

Krajcik (2012, 2013) considered the valid name for this species to be Pseudochlorota chiriquina. Lacking his rationale for this nomenclatural change, we use the name H. chiriquina.

HOMONYX Guérin-Méneville, 1839

Homonyx Guérin-Méneville, 1839: 299–300.

Type species

Homonyx cupreus Guérin-Méneville, 1839: 300, by monotypy.

Gender

Masculine.

Species

14 species and subspecies.

Homonyx argentinus Gutiérrez, 1952

Homonyx planicostatus argentinus Gutiérrez, 1952: 224, 225 [original combination].

Homonyx argentinus Gutiérrez [new species status by Soula 2010a: 16].

Distribution

ARGENTINA: Jujuy, Mendoza, Salta, Tucumán (Gutiérrez 1952, Machatschke 1972, Krajcik 2008, Soula 2010a).

Types

1 ♀ paratype at MNNC. 1 ♂ and 5 ♀ paratypes at CMNC. Gutiérrez (1952) stated the holotype male was deposited in his collection at UCCC.

Remarks

Krajcik (2012, 2013) considered H. argentinus to be a subspecies of H. planicostatus.

Homonyx chalceus bahianus Ohaus, 1913

Homonyx bahianus Ohaus, 1913: 495–496 [original combination].

Homonyx chalceus bahianus Ohaus [new species status by Soula 2010a: 14].

Distribution

BRAZIL: Bahia (Ohaus 1913, 1918, 1934b; Machatschke 1972, Krajcik 2008, Soula 2010a).

Types

1 ♂ lectotype and 1 paralectotype at ZMHB (Soula 2010a) (Fig. 26).

Remarks

Krajcik (2012, 2013) considered H. bahianus to be a valid species rather than a subspecies of H. chalceus.

Figure 26.

Homonyx bahianus Ohaus (valid name H. chalceus bahianus) type male (see “Type specimens and lectotype designation” in Methods) from ZMHB. A Dorsal habitus B Lateral habitus C Specimen labels, mouthparts, and genitalia D Male genitalia, lateral view E Parameres, caudal view.

Homonyx chalceus chalceus Blanchard, 1851

Homonyx chalceus Blanchard, 1851: 214 [original combination].

Distribution

ARGENTINA: Corrientes, Mendoza, Salta, San Luis (Blanchard 1851, Burmeister 1855, Steinheil 1874, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972, Krajcik 2008, Soula 2010a).

Types

1 ♂ holotype at MNHN (Soula 2010a). An exemplar specimen identified by Ohaus and compared with Blanchard’s type specimen is figured (Fig. 27).

Figure 27.

Homonyx chalceus Blanchard male (male specimen compared [by Ohaus] with Blanchard’s type deposited at MNHN). A Dorsal habitus B Lateral habitus C Male genitalia, lateral view D Specimen labels and male genitalia.

Homonyx cupreus Guérin-Méneville, 1839

Homonyx cupreus Guérin-Méneville, 1839: 300 [original combination].

Distribution

ARGENTINA: Corrientes, Salta (Burmeister 1844, Ohaus 1913, 1918, 1934b).

Remarks

Homonyx cupreus Guérin-Méneville was erroneously reported from the extreme southern Chilean Magallanes Province and later from the specific locality of Port Famine (modern Puerto del Hambre) (Solier 1851, Reed 1876, Philippi 1887, Ohaus 1910c, 1918, 1934b, Machatschke 1972, Krajcik 2008). This locality is dubius based on the distribution of other known Homonyx species, which have their diversity centered in Peru, Ecuador, Bolivia, and central Argentina. Further collecting in southern Chile and southern Argentina is needed to establish whether Homonyx species indeed occur there.

Homonyx demezi Soula, 2010

Homonyx demezi Soula, 2010a: 23 [original combination].

Distribution

BRAZIL: Mato Grosso (Soula 2010a).

Types

The following specimens are deposited at CCECL. 1 ♂ holotype, 1 ♀ allotype, 3 ♂ paratypes, 1 ♀ paratype: “Mato Grosso Brésil coll. – SOULA//Holotype 2010 Homonyx demezi S. Soula (47030999); “Matto Grosso Brésil coll. – SOULA//Allotype 2010 Homonyx demezi S. Soula (47031000); “Rosario Matto Grosso M. SOULA det 19 [obverse] 10/61//Paratype 2010 Homonyx demezi S. Soula (47031001); “Corumba Matt.//Paratype 2010 Homonyx demezi S. Soula (47031002); “Rosario Oeste Matto Grosso 01/72 coll. – Soula [obverse] Rosario Oeste//Paratype 2010 Homonyx demezi S. Soula (47031003); “Gob. de Los Andes//Paratype 2009 Homonyx demezi S. Soula (47031004). Genitalia card-mounted underneath the male holotype and the three male paratypes. Box 4618689 SOULA. The following specimen is deposited at CMNC. 1 ♀ paratype: “BRASIL Mato Grosso Rosario Oeste A. Maller-leg. Coll. Martínez Oct.-968// H. & A. HOWDEN COLLECTION ex. A. Martinez coll.//Paratype 2010 Homonyx demezi S. Soula”.

Homonyx elongatus Blanchard, 1842

Rutela elongata Blanchard, 1842: plate 11 [original combination].

Homonyx elongatus (Blanchard) [new combination by Blanchard 1851: 214].

Distribution

ARGENTINA (Blackwelder 1944). BOLIVIA: Pando (Blanchard 1851, Burmeister 1855, Ohaus 1918, 1934b, Blackwelder 1944, Machatschke 1972).

Types

1 ♀ holotype at MNHN (Soula 2010a).

Homonyx feyeri Ohaus, 1913

Homonyx feyeri Ohaus, 1913: 496–497 [original combination].

Distribution

ECUADOR: Morona-Santiago (Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Paucar-Cabrera 2005, Soula 2010a).

Types

1 ♂ holotype specimen of Homonyx feyeri Ohaus at ZMHB (Fig. 28). Soula states that ♂ holotype is at ZMHB (Soula 2010a).

Figure 28.

Homonyx feyeri Ohaus holotype male from ZMHB. A Dorsal habitus B Lateral habitus C Specimen labels, mouthparts, and male genitalia D Male genitalia, lateral view E Parameres, caudal view.

Homonyx fuscocupreus (Ohaus, 1913)

Homonyx chalceus var. fuscocupreus Ohaus, 1913: 494 [original combination].

Homonyx chalceus fuscocupreus Ohaus [new subspecific status by Machatschke 1972: 19].

Homonyx fuscocupreus Ohaus [new species status by Soula 2011: 73–74].

Distribution

ARGENTINA: Catamarca, Tucumán (Ohaus 1913, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Krajcik 2008, Soula 2010a, 2011).

Types

1 lectotype and 1 paralectotype at ZMHB (Soula 2010a). An exemplar specimen is shown in Figure 29.

Remarks

Ohaus (1913) described Homonyx chalceus ssp. uruguayanus, Homonyx chalceus ssp. santiagensis, and Homonyx chalceus var. fuscocupreus. Thus, in the context of this publication, it is unambiguous that Homonyx chalceus var. fuscocupreus is infrasubspecific and should be interpreted in this manner. Some publications have treated the taxon as a subspecies (Homonyx chalceus fuscocupreus) according to ICZN Article 45.6.4.1, thus making this species-group name available. Homonyx fuscocupreus was elevated to species status by Soula (2011). Krajcik (2012, 2013) considered H. fuscocupreus to be a subspecies of H. chalceus.

Figure 29.

Homonyx chalceus var. fuscocupreus Ohaus (valid name H. fuscocupreus) type male from ZMHB. A Dorsal habitus B Lateral habitus C Specimens labels D Parameres, caudal view.

Homonyx holligeri Soula, 2010

Homonyx holligeri Soula, 2010a: 19–20 [original combination].

Distribution

BOLIVIA: Santa Cruz (Soula 2010a).

Types

The following specimens are deposited at CCECL. 1 ♂ holotype, 1 ♂ paratype: “Coroïco à Caranavi 850 m 10/88//Holotype 2010 Homonyx holligeri S. Soula (47030993); “Bolivia-Dept. Santa Cruz-800 m 25.X.1960-Zischka//Paratype 2010 Homonyx holligeri S. Soula (47030994). Genitalia card-mounted underneath the male holotype and the male paratype. Box 4618689 SOULA.

Homonyx maurettei Soula, 2010

Homonyx maurettei Soula, 2010a: 18–19 [original combination].

Distribution

PERU: Piura (Soula 2010a, Ratcliffe et al. 2015).

Types

The following specimen is deposited at CCECL. 1 ♂ holotype: “Abra Porculla, Dt. Piura N-W Pérou; 1800m II/2007//Holotype Homonyx maurettei S. 2010 Soula (47030998). Genitalia card-mounted underneath the male holotype. Box 4618689 SOULA.

Homonyx peruanus Ohaus, 1913

Homonyx planicostatus peruanus Ohaus, 1913: 496 [original combination].

Homonyx elongatus peruanus Ohaus [revised subspecies status by Ohaus 1918: 21].

Homonyx planicostatus peruanus Ohaus [revised subspecies status by Ohaus 1934b: 73].

Homonyx elongatus peruanus Ohaus [revised subspecies status by Ohaus 1952: 2].

Homonyx peruanus Ohaus [new species status by Soula 2010a: 18].

Distribution

PERU (Blackwelder 1944, Ohaus 1913, 1934b, 1952, Machatschke 1972, Soula 2010a, Ratcliffe et al. 2015).

Types

1 ♀ syntype specimen of Homonyx planicostatus peruanus Ohaus at ZMHB (Fig. 30) (probably the ♀ holotype referred to by Soula [2010a]).

Remarks

Krajcik (2012, 2013) considered H. peruanus to be a subspecies of H. elongatus.

Figure 30.

Homonyx planicostatus peruanus Ohaus (valid name H. peruanus) syntype female from ZMHB. A Dorsal habitus B Lateral habitus C Specimen labels and mouthparts.

Homonyx planicostatus Blanchard, 1851

Homonyx planicostatus Blanchard, 1851: 214 [original combination].

Distribution

ARGENTINA: Mendoza, Tucumán (Ohaus 1913, 1934b, Blackwelder 1944, Gutiérrez 1952, Machatschke 1972). BOLIVIA (Blanchard 1851, Burmeister 1855, Ohaus 1913, 1934b, Blackwelder 1944, Gutiérrez 1952, Machatschke 1972, Krajcik 2008, Soula 2010a).

Types

1 ♀ syntype at MNHN (Soula 2010a). An exemplar specimen identified by Ohaus and compared with Blanchard’s type specimen is figured (Fig. 31).

Remarks

CCECL contains a H. planicostatus specimen labeled as a male ♂ alloréférent with the following data: “Vaccaguzman [arrow] Camiri coll. – SOULA [obverse] 1615m//Alloréférent ♂ de Homonyx planicostatus Bl. M. SOULA det 19 (47030995). Genitalia card-mounted underneath specimen. Box 4618689 SOULA.

Figure 31.

Homonyx planicostatus Blanchard (male specimen compared [by Ohaus] with Blanchard’s type from MNHN). A Dorsal habitus B Lateral habitus C Specimen labels and male genitalia D Male genitalia, lateral view E Male parameres, caudal view.

Homonyx santiagensis Ohaus, 1913

Homonyx chalceus santiagensis Ohaus, 1913: 494 [original combination].

Homonyx santiagensis Ohaus [new species status by Soula 2010a: 12].

Distribution

ARGENTINA: Córdoba, Jujuy, Santiago del Estero (Ohaus 1913, 1934b, Blackwelder 1944, Machatschke 1972, Krajcik 2008, Soula 2010a).

Types

1 ♂ lectotype and 1 paralectotype at ZMHB (Fig. 32) (Soula 2010a).

Remarks

Krajcik (2012, 2013) considered H. santiagensis to be a subspecies of H. chalceus.

Figure 32.

Homonyx chalceus santiagensis Ohaus (valid name H. santiagensis) type male (see “Type specimens and lectotype designation” in Methods) from ZMHB. A Dorsal habitus B Lateral habitus C Specimens labels and male genitalia D Male genitalia, lateral view E Parameres, caudal view.

Homonyx uruguayanus Ohaus, 1913

Homonyx chalceus uruguayanus Ohaus, 1913: 494 [original combination].

Homonyx uruguayensis Ohaus [new species status and incorrect subsequent spelling by Soula 2010a: 13].

Distribution

ARGENTINA: Córdoba, Entre Ríos (Soula 2010a). URUGUAY (Ohaus 1913, 1934b, Blackwelder 1944, Machatschke 1972, Krajcik 2008, Soula 2010a).

Types

1 ♂ syntype of Homonyx chalceus uruguayanus at ZMHB (called a holotype by Soula 2010a) (Fig. 33).

Remarks

Krajcik (2012, 2013) considered H. uruguayensis to be a subspecies of H. chalceus.

Figure 33.

Homonyx chalceus uruguayanus Ohaus (valid name H. uruguayanus) syntype male from ZMHB. A Dorsal habitus B Lateral habitus C Specimens labels and male genitalia D Male genitalia, lateral view E Parameres, caudal view.

Unavailable names in Homonyx(application of ICZN Article 16.4.2)

We consider the following names proposed by Soula in Homonyx as unavailable per ICZN Article 16.4.2 which states that fixation of holotype specimens for new names must be accompanied by the following information, “where the holotype or syntypes are extant specimens, by a statement of intent that they will be (or are) deposited in a collection and a statement indicating the name and location of that collection”. The names below were proposed by Soula (2010, 2011), but the descriptions did not state the intent to deposit the holotype specimens in a collection. By applying ICZN Article 16.4.2 herein, the following names are unavailable: Homonyx digennaroi Soula 2010, Homonyx lecourti Soula 2010, Homonyx mulliei Soula 2010, Homonyx simoensi Soula 2010, Homonyx wagneri Soula 2010, and Homonyx zovii Demez and Soula 2011. Below we report the complete taxonomic history of these names and the data from their invalid type specimens that are deposited at CCECL.

Homonyx digennaroi Soula, 2010 Unavailable, invalid name

Homonyx digennaroi Soula, 2010a: 19, 21-22 [original combination, unavailable, invalid name].

Distribution

BOLIVIA (Soula 2010a).

Types

The following invalid type specimens are deposited at CCECL. 1 invalid ♂ holotype: “Rte de Camiri à Sta Cruz Bol. coll. – SOULA//Holotype 2010 Homonyx digennaroi S. Soula (47031008). Genitalia card-mounted underneath the male holotype. Box 4618689 SOULA.

Homonyx lecourti Soula, 2010 Unavailable, invalid name

Homonyx lecourti Soula, 2010a: 19, 20 [original combination, unavailable, invalid name].

Distribution

BOLIVIA: La Paz (Soula 2010a).

Types

The following invalid type specimens are deposited at CCECL. 1 invalid ♂ holotype and 1 invalid ♀ allotype: “Yocumo 920 m 26/10/2000 M. SOULA det 19//Holotype 2010 Homonyx lecourti S. Soula (47031006); “Yocumo 920 m 26/X/2000 M. SOULA det 19//Allotype 2010 Homonyx lecourti S. Soula (47031007). Genitalia card-mounted underneath the male holotype. Box 4618689 SOULA.

Homonyx mulliei Soula, 2010 Unavailable, invalid name

Homonyx mulliei Soula, 2010a: 23, 24 [original combination, unavailable, invalid name].

Distribution

BOLIVIA (Soula 2010a).

Types

The following invalid type specimen is deposited at CCECL. 1 invalid ♂ holotype: “Camiri à Sta Cruz coll. – Soula//Holotype 2010 Homonyx mulliei S. Soula (47031005). Genitalia card-mounted underneath the male holotype. Box 4618689 SOULA.

Homonyx simoensi Soula, 2010 Unavailable, invalid name

Homonyx simoensi Soula, 2010a: 22, 23 [original combination, unavailable, invalid name].

Distribution

BOLIVIA (Soula 2010a).

Types

The following invalid type specimen is deposited at CCECL. 1 invalid ♂ holotype: “Camiri à Sta Cruz coll. – SOULA//Holotype 2010 Homonyx simoensi S. Soula (47031009). Genitalia card-mounted underneath the male holotype. Box 4618689 SOULA.

Homonyx wagneri Soula, 2010 Unavailable, invalid name

Homonyx wagneri Soula, 2010a: 23, 25 [original combination, unavailable, invalid name].

Distribution

ARGENTINA: Salta (Soula 2010a).

Types

The following invalid type specimen is deposited at CCECL. 1 invalid ♂ holotype: “Salta Argentine XI/2006 M. SOULA det 19//Holotype 2010 Homonyx wagneri S. Soula (47030997). Genitalia card-mounted underneath the male holotype. Box 4618689 SOULA.

Homonyx zovii Demez & Soula, 2011 Unavailable, invalid name

Homonyx zovii Demez & Soula, 2011: 74 [original combination, unavailable, invalid name].

Distribution

PERU: San Martín (Soula 2011, Ratcliffe et al. 2015).

Types

The following invalid type specimen is deposited at CCECL. 1 invalid ♂ holotype: “Janjui San Martin IX/2010 M. SOULA det 19//Holotype 2011 Homonyx zovii S. 2011 Soula (47030996). Genitalia card-mounted underneath the male holotype. Box 4618689 SOULA.

HOMOTHERMON Ohaus, 1898

Homothermon Ohaus, 1898: 59-60.

Type species

Homothermon bugre Ohaus, 1898: 60, original designation by Ohaus 1898: 59–60.

Gender

Neuter.

Species

4 species.

Homothermon bugre Ohaus, 1898

Homothermon bugre Ohaus, 1898: 60 [original combination].

Distribution

ARGENTINA: Misiones (Ohaus 1898, 1934b, Machatschke 1972, Krajcik 2008, Soula 2008). BRAZIL: Rio Grande do Sul, Santa Catarina (Ohaus 1898, 1918, 1934b, Machatschke 1972, Soula 2008).

Types

1 ♂ lectotype and paralectotypes at ZMHB (Soula 2008); 1 paralectotype at MNHN (Soula 2010a).

Homothermon drumonti Soula, 2008

Homothermon drumonti Soula, 2008: 33 [original combination].

Distribution

BRAZIL: São Paulo (Soula 2008).

Types

The following specimen is deposited at CCECL. 1 ♂ holotype: “Brasil OL. Guillot//Det Dr. Ohaus Homothermon paulista Ohaus//Holotype Homothermon drumonti S. 2007 Soula” (47031074). Genitalia card-mounted underneath the male holotype. Box 4618691 SOULA.

Homothermon praemorsus (Burmeister, 1855)

Odontognathus praemorsus Burmeister, 1855: 521 [original combination].

Homothermon praemorsus (Burmeister) [new combination by Ohaus 1918: 30].

synonym. Homothermon paulista Ohaus, 1898

Homothermon paulista Ohaus, 1898: 61 [original combination].

Homothermon praemorsus (Burmeister) [syn. by Ohaus 1918: 30].

Distribution

BRAZIL: São Paulo (Burmeister 1855, Ohaus 1898, 1918, 1934b, Machatschke 1972, Krajcik 2008, Soula 2008).

Types

1 ♀ syntype of Odontognathus praemorsus at ZMHB (Soula 2008). 1 ♂ syntype specimen of Homothermon paulista at ZMHB (Fig. 34); 1 syntype specimen of Homothermon paulista at SDEI. Soula (2008: 32) stated that he found the holotype at ZMHB, yet he provided an image of a lectotype specimen (see “Type Specimens and Lectotype Designation” in Methods).

Figure 34.

Homothermon paulista Ohaus (valid name H. praemorsus Burmeister) type male (see “Type specimens and lectotype designation” in Methods) from ZMHB. A Dorsal habitus B Lateral habitus C Specimens labels, mouthparts, and male genitalia D Male genitalia, lateral view.

Homothermon serrano Ohaus, 1898

Homothermon serrano Ohaus, 1898: 60 [original combination].

Distribution

ARGENTINA: Misiones (Soula 2008). BRAZIL: Rio Grande do Sul, Santa Catarina (Ohaus 1898, 1918, 1934b, Machatschke 1972, Krajcik 2008, Soula 2008).

Types

1 ♂ lectotype and 2 paralectotypes at ZMHB (Fig. 35); 1 paralectotype at NHMB (Soula 2008).

Figure 35.

Homothermon serrano Ohaus paralectotype male from ZMHB. A Dorsal habitus B Lateral habitus C Specimens labels and male genitalia D Male parameres, caudal view.

HOPLOPELIDNOTA F. Bates, 1904

Hoplopelidnota Bates, 1904: 253, 274–275.

Type species

Hoplopelidnota candezei F. Bates, 1904: 274–275, by monotypy.

Gender

Feminine.

Species

1 species.

Hoplopelidnota metallica (Laporte, 1840)

Pelidnota metallica Laporte, 1840: 122 [original combination].

Hoplopelidnota candezei F. Bates [syn. by Machatschke 1972: 12].

Hoplopelidnota metallica (Laporte) [revised species status by Soula 2008: 17].

synonym. Hoplopelidnota armata Ohaus, 1912

Hoplopelidnota armata Ohaus, 1912: 309 [original combination; sometimes erroneously referred to as H. armata F. Bates].

Hoplopelidnota metallica (Laporte) [syn. by Moore and Jameson 2013: 381].

synonym. Hoplopelidnota candezei F. Bates, 1904

Hoplopelidnota candezei F. Bates, 1904: 274–275 [original combination].

Hoplopelidnota metallica (Laporte) [syn. by Soula 2008: 17].

Distribution

BRAZIL: Territorio de Amapa (Serra Navia). FRENCH GUIANA: Cayenne (Laporte 1840, F. Bates 1904, Ohaus 1912, 1918, 1934b, Blackwelder 1944, Machatschke 1972, Soula 2008, 2010c). GUYANA: Essequibo River, Moraballi Creek. VENEZUELA: Amazonas (Rio Negro).

Types

The following specimen is deposited at CCECL. 1 invalid ♂ neotype (Fig. 36): “pk 23 p. de Belizon G. F. 8/91 coll. – SOULA [obverse] pk 23//Néotype 2007 Pelidnota metallica Lap. Soula det.//Hoplopelidnota metallica (Lap.) M. SOULA 19 2007” (47031033). Genitalia card-mounted underneath the invalid neotype. Box 4618690 SOULA.

Remarks

The classification of Hoplopelidnota metallica (Figs 36, 37) has been tumultuous (Moore and Jameson 2013). Laporte (1840) named P. metallica, clearly indicating the unusual form of the elytral apex. Bates’s (1904) description of Hoplopelidnota candezei overlooked the conspecific P. metallica. Hoplopelidnota candezei was based on a single male specimen that was labeled “Pelidnota armata” by Candèze. The name “H. armata”, however, had not been validly described and therefore was not available. However, Ohaus (1912) provided a description of the female, he used the name “Hoplopelidnota armata” (rather than H. candezei). This act made the name H. armata an available name and a junior synonym of H. metallica. Most recently, Krajcik (2012, 2013) considered H. candezei to be a subspecies of H. metallica.

Soula (2008: 17–18) attempted to designate a neotype specimen for Hoplopelidnota metallica. Soula stated that the neotype is in “Collection Soula”, but Article 75.3.7 (ICZN 1999) requires a statement that the “neotype is, or immediately upon publication has become, the property of a recognized scientific or educational institution, cited by name, that maintains a research collection, with proper facilities for preserving name-bearing types, and that makes them accessible for study”. Because Soula’s collection was private at the time of designation, Soula’s neotype is invalid.

Hoplopelidnota metallica is distributed in northern South America. Prior to this work, H. metallica was only recorded from French Guiana. In addition to French Guiana, we record the species from Guyana (Moraballi Creek, Essequibo River), Venezuela (Amazonas Dept., Rio Negro) and Brazil (Territorio de Amapa, Serra Navia). The species is rare in collections, and is apparently much more wide spread in northern South America than previous data would indicate. Specimens are recorded from 140 m elevation in March, April, July, August, and November.

Figure 36.

Hoplopelidnota metallica (Laporte) invalid neotype male from CCECL. A Dorsal habitus B Male parameres, caudal view C Specimen labels.

Figure 37.

Hoplopelidnota metallica (Laporte) male from <