identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
5B62C6B2BE0D4F372BAF31F54566C5AB.text	5B62C6B2BE0D4F372BAF31F54566C5AB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Acrobolbia Ohaus 1912	<div><p>Acrobolbia Ohaus, 1912</p><p>Type species.</p><p>Acrobolbia macrophylla Ohaus, 1912, by monotypy.</p><p>Valid taxa.</p><p>One species.</p><p>The northern South American genus Acrobolbia is known from Peru, Ecuador, and possibly Venezuela (Ohaus 1912, Machatschke 1972, Jameson et al. 2002) (Fig. 51). Acrobolbia has a complicated classification history. Ohaus (1912) described A. macrophylla based upon a single male specimen collected in Peru. Ohaus (1912) compared Acrobolbia to Cyclocephala, but he ultimately classified the genus in the subtribe Areodina ( Rutelinae: Rutelini). Ohaus (1918) later transferred the genus into its own subtribe, Acrobolbiina, within Rutelini . Acrobolbia triangularis was the second species to be described into the genus, but this species was later treated as a synonym and a “variant” of A. macrophylla (Benderitter 1922, Ohaus 1934a, b).</p><p>Based on the elongated antennal club of the male in Acrobolbia, the genus was transferred into the ruteline subtribe Oryctomorphina (Dechambre and Ponchel 1999). Most recently, Acrobolbia was reviewed and transferred into Cyclocephalini by Jameson et al. (2002). Acrobolbia is hypothesized to be related to Ancognatha based upon characters of the clypeus, mentum, pronotum, prosternal process, protarsus, and mandibles (Jameson 1998, Jameson et al. 2002). Specimens of Acrobolbia are rare in collections, and almost nothing is known of their biology (Jameson et al. 2002). Acrobolbia macrophylla adults are attracted to lights at night, though specimens do not land or rest at light traps (Jameson et al. 2002). Specimens have been collected from 400-1,200 m in elevation (Jameson et al. 2002). The immature stages are undescribed and unknown.</p><p>Acrobolbia species can be recognized by the following combination of characters: 1) dorsal coloration varying from all black with variable reddish brown margins of the elytra and elytral suture, or with the elytra partially testaceous; 2) body not anteroposteriorly compressed or dorsoventrally flattened; 3) clypeal apex acuminate in dorsal view; 4) frontoclypeal suture distinct, but incomplete medially; 5) mandibles long, sickle-shaped, with pointed apex; 6) mandibular molar area with rows of circular micropunctures; 7) apical margin of mentum weakly emarginate to nearly straight; 8) galea of maxilla reduced to small, rectangular mound in dorsal view; 9) galea on inner surface with teeth greatly reduced to peg-like projections at the middle and apex; 10) galea on inner surface lacking teeth at base; 11) males with antennal club (segments 8-10) elongated, nearly twice as long as antennomeres 1-7; 12) pronotum with broadly incomplete beaded basal margin; 13) males and females with 3 protibial teeth, basal tooth reduced, removed from the apical 2 teeth, and oriented laterally; 14) protibial spur straight to weakly deflexed; 15) males with inner protarsal claw enlarged and narrowly cleft at apex; 16) mesocoxae touching, nearly contiguous; 17) meso- and metatibiae with distal, divided carinae; 18) metacoxae with lateral edge perpendicular to ventral surface; 19) anterior edge of hindwing distal to apical hinge lacking setae and with produced, membranous border; 20) vein RA with 2 rows of pegs extending distally nearly to margin of apical hinge.</p></div>	https://treatment.plazi.org/id/5B62C6B2BE0D4F372BAF31F54566C5AB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
57846B736B8B5C8904BA3ACCB623C260.text	57846B736B8B5C8904BA3ACCB623C260.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ancognatha Erichson 1847	<div><p>Ancognatha Erichson, 1847</p><p>Type species.</p><p>Ancognatha scarabaeoides Erichson, subsequent designation by Casey 1915: 111.</p><p>Valid taxa.</p><p>22 species.</p><p>The 22 species of Ancognatha are distributed from the southwestern United States south to Argentina (Fig. 52). The species diversity in the genus is concentrated in north and western South America and in Mexico, west of the Isthmus of Tehuantepec. Biological information on Ancognatha species is lacking, and almost nothing is known about the natural history of adults. In Meso- and Central America, Ancognatha species are associated with premontane, lower montane, and montane tropical forests with some species being collected at elevations from 2,000 to 3,500 m above sea level (Ratcliffe 2003, Ratcliffe and Cave 2006, Ratcliffe et al. 2013). This pattern also holds in South America. Several Ancognatha species have been recorded from elevations over 4,000 m in Peru and northern Chile (Mondaca 2016, Figueroa and Ratcliffe 2016). Some South American Ancognatha species can be very large for the tribe. For example, A. matilei Dechambre from Colombia is up to the 36 mm long (Dechambre 2000). Adults are attracted to lights at night.</p><p>Larvae are described for four Ancognatha species (Ritcher 1966, Ramírez-Salinas et al. 2004, Vallejo and Morón 2008, Neita-Moreno and Morón 2008). South American larval descriptions are largely based on material collected in agroecosystems, and thus the natural ecology of Ancognatha immatures is poorly known. Mondaca (2016) reported the larvae of A. aymara Mondaca feeding on grass roots high in the altiplano steppe of northern Chile.</p><p>Ancognatha species can be recognized by the following combination of characters: 1) dorsal coloration variable, from all or partially black or testaceous, to light brown with variable dark maculae; 2) body convex and not strongly anteroposteriorly or dorsoventrally compressed; 3) clypeal apex rounded to parabolic, never truncate or emarginate; 4) frontoclypeal suture incomplete medially; 5) males with anterolateral margin of the mandibles without teeth; 6) mandibular apices narrow and elongated, recurved dorsally; 7) mandibular molar area with rows of circular micropunctures; 8) apical margin of mentum narrowly and deeply emarginated; 9) galea of maxilla reduced to a roughly quadrate process; 10) galea of the maxilla on inner surface lacking well-developed teeth, teeth when present and visible greatly reduced into spine-like projec tions; 11) males and females with 3 protibial teeth, basal tooth slightly removed from the more apical 2 teeth, and oriented laterally; 12) protibial spur straight to weakly deflexed; 13) males with inner protarsal claw enlarged and narrowly cleft at apex; 14) mesocoxae narrowly separated and touching; 15) meso- and metatibiae with distal, transverse carinae; 16) metacoxae with lateral edge perpendicular to ventral surface; 17) anterior edge of hindwing distal to apical hinge lacking setae and with produced, membranous border; 18) vein RA with single row of pegs extending distally nearly to margin of apical hinge; 19) elytral margin membranous.</p><p>The relationship of Ancognatha species to other cyclocephaline genera has not been evaluated. Acrobolbia may be related to Ancognatha based on characters of the clypeus, mentum, pronotum, prosternal process, protarsus, and mandibles (Jameson 1998, Jameson et al. 2002). Surutu also shares some intriguing characters with Ancognatha, which may be indicative of a close relationship between these two genera. For example, Ancognatha and Surutu species all have a rounded to parabolic clypeal apex and a narrowly, but deeply, emarginated apex of the mentum. Surutu species have a anteriorly projecting tooth at the apex of the labrum, and this is also shared in some Ancognatha species.</p></div>	https://treatment.plazi.org/id/57846B736B8B5C8904BA3ACCB623C260	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
5E6B8BC3BB8C5B646B85600F2BB84EDD.text	5E6B8BC3BB8C5B646B85600F2BB84EDD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Arriguttia Martinez 1960	<div><p>Arriguttia Martinez, 1960</p><p>Type species.</p><p>Cyclocephala brevissima Arrow, 1911, by monotypy.</p><p>Valid taxa.</p><p>Two species.</p><p>Arriguttia contains two South American species known only from the Brazilian Amazon, Guyana, and French Guiana (Arrow 1911, 1937b, Blackwelder 1944, Martínez 1960a, 1968a, Endrődi 1966, 1985a, Ponchel 2006, 2011, 2015) (Fig. 53). Very little is known about the biology of Arriguttia species. Arriguttia brevissima (Arrow) feeds within the inflorescences of Victoria sp. in Brazil ( Martínez 1968a). In French Guiana, A. brevissima was found in the spathes of an unidentified terrestrial aroid ( Araceae) (Ponchel 2006, 2015). In Brazilian cerrado habitat, A. brevissima are floral visitors of Annona coriacea Mart. and are likely late-season, secondary pollinators of this species (Costa et al. 2017). Specimens of A. brevissima have been collected at lights at night ( Martínez 1968a). The immature stages are undescribed and unknown.</p><p>Arriguttia was compared to Surutu in the original description of the genus ( Martínez 1960a). This is possibly confusing for identification purposes. Arriguttia shares many more characters with Cyclocephala and Augoderia than with Surutu . Arriguttia species can be recognized by the following combination of characters: 1) dorsal coloration varying from all black or with variable dark, reddish coloration on the elytra; 2) body convex and anteroposteriorly compressed, creating a relatively round gestalt; 3) clypeus quadrate in dorsal view, with sides nearly parallel, and the apex distinctly reflexed upwards (most obvious in lateral view); 4) frontoclypeal suture complete medially; 5) males with anterolateral margin of the mandibles weakly toothed; 6) mandibular molar area with rows of circular micropunctures; 7) apical margin of mentum weakly emarginated; 8) galea of the maxilla on inner surface with 3 fused basal teeth, a free median tooth, and 2 fused apical teeth (3-1-2 arrangement); 9) pronotum with broadly incomplete beaded basal margin; 10) males and females with 3 protibial teeth, basal tooth reduced, removed from the more apical 2 teeth, and oriented anteriorly; 11) protibial spur straight to weakly deflexed; 12) males with inner protarsal claw enlarged and narrowly cleft at apex; 13) mesocoxae widely separated; 14) meso- and metatibiae with distal, transverse carinae; 15) metacoxae with lateral edge perpendicular to ventral surface; 16) anterior edge of hindwing distal to apical hinge lacking setae and with produced, membranous border; 17) vein RA with 2 rows of pegs extending distally nearly to margin of apical hinge.</p><p>The relationships of Arriguttia to other cyclocephaline genera have not been clearly discussed in the literature. Martínez (1968a) stated that Arriguttia should be "placed next to" Surutu, but he did not offer any character justifications for this hypothesis. Endrődi (1966) considered Arriguttia to be a “primitive” cyclocephaline based on his poorly justified character analysis. Arriguttia shares hindwing characters (two rows of pegs on vein RA and a membrane on the leading edge of the hindwing distal to the apical hinge) with Augoderia, Aspidolea, and Cyclocephala . The form of the maxilla (3-1-2 teeth arrangement), the mandibular form (males with anterolateral margin weakly toothed and the molar area with rows of circular micropunctures), the incomplete bead on the basal margin of the pronotum, and the shape and arrangement of the protibial teeth are shared among Arriguttia, Augoderia, and some Cyclocephala (especially species like C. sexpunctata Laporte and species formerly placed in Stigmalia Casey). Future analyses should focus on comparing characters in these Cyclocephala species-groups and genera to Arriguttia, rather than Surutu .</p></div>	https://treatment.plazi.org/id/5E6B8BC3BB8C5B646B85600F2BB84EDD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
D8927393A343DAB9D5EB78C18D028AD6.text	D8927393A343DAB9D5EB78C18D028AD6.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Aspidolea Bates 1888	<div><p>Aspidolea Bates, 1888</p><p>Type species.</p><p>Aspidolea singularis Bates, 1888: 296-297, by monotypy.</p><p>Valid taxa.</p><p>26 species.</p><p>Aspidolea contains 26 species ranging from northern Mexico south through South America (Fig. 54) ( Endrődi 1966, 1985a, Ratcliffe 2003, Ratcliffe and Cave 2006, Ratcliffe et al. 2013). The genus includes both widespread and narrowly distributed species. Most Aspidolea (22 of 26 species) are known only from a few South American localities. In contrast, A. fuliginea and A. singularis occur from Mexico south to Argentina and Ecuador, respectively. Bates (1888) described Aspidolea based upon the "elongate and robust" yet toothless maxillary galea found in the type species A. singularis . Bates (1888) noted a similar reduction in maxillary teeth in " Cyclocephala fuliginea Burmeister" and Ancognatha species. Aspidolea contained only A. singularis for over 30 years until Höhne (1922a, b, c) recircumscribed the genus and placed many new species into the group.</p><p>Höhne (1922a) offered an expanded diagnosis of Aspidolea using characters of the clypeus (sides parallel at base with apical margin perpendicular to the sides), maxilla (toothless and with penicillate setae at the apex), and dorsum (yellow to brownish coloration and generally lacking maculae) to distinguish the genus. Cyclocephala clypeata Burmeister and C. laticeps Harold were transferred into Aspidolea along with ten new species described by Höhne (1922a). The new genus Paraspidolea was erected to contain species similar to Aspidolea, but with at least two small teeth present at the apex of the galea ( Höhne 1922a). Six new species were included in Paraspidolea along with the Burmeister species C. fuliginea ( Höhne 1922a, b). The subgenus Aspidolea (Aspidolites) was erected to contain the species A. atricollis Höhne ( Höhne 1923c). The homonym Aspidolites Höhne was replaced with Aspidolella (Prell 1936). Aspidolea atricollis is conspecific with C. histrionica Burmeister ( Endrődi 1966), and the subgenus Aspidolella is considered a synonym of Cyclocephala . Paraspidolea was also synonymized within Aspidolea ( Endrődi 1966).</p><p>The last major contribution to the knowledge of Aspidolea was provided by Dechambre (1992). Dechambre (1992) described three new Aspidolea species, which he included in the " Aspidolea helleri species-group" along with A. helleri ( Höhne) and A. chalumeaui Endrődi . These species were placed into the " helleri species-group" based on the bidentate form of the protibial margin in males. This male protibial character is shared with species formerly included in Mimeoma and some Cyclocephala species (like C. amazona) (see Moore et al. 2015). The dorsal coloration of the " helleri species-group", especially the elongated, triangular maculae found along the elytral suture, is like that found in some former Mimeoma species (especially Cyclocephala acuta Arrow and C. englemani (Ratcliffe)). These characters suggest that Aspidolea may not be monophyletic as presently defined.</p><p>There is little available biological data for Aspidolea species. Aspidolea adults seem to be readily attracted to lights at night and can occasionally be collected in large numbers (Ratcliffe and Cave 2006, Touroult et al. 2010, Grossi et al. 2011). Floral association data for Aspidolea are mostly lacking. Aspidolea fuliginea were collected in male- and female-phase inflorescences of Oenocarpus bataua Mart. ( Arecaceae) in Colombia, though they were only sporadically encountered ( Núñez-Avellaneda and Rojas-Robles 2008). In French Guiana, A. quadrata Endrődi was collected from the inflorescence of Montrichardia arborescens (L.) Schott ( Araceae) (Gibernau et al. 2003, Ponchel 2006). Neita-Moreno et al. (2007) described the larva and pupa of A. singularis . Larvae of A. singularis were collected from soil beneath cultivated cassava ( Manihot esculenta Crantz; Euphorbiaceae) in Colombia (Neita-Moreno et al. 2007).</p><p>Aspidolea species can be recognized by the following combination of characters: 1) dorsal coloration highly variable, with or without black or brown maculae on the pronotum and elytra; 2) body not anteroposteriorly compressed or dorsoventrally flattened; 3) clypeus robust and broad, with sides more or less parallel at base, appearing quadrate in dorsal view; 4) frontoclypeal suture complete medially; 5) males with anterolateral margin of the mandibles weakly toothed (in A. fuliginea) or not; 6) mandibular molar area with rows of circular micropunctures; 7) apical margin of mentum broadly and deeply (nearly to level of labial palp insertion) emarginated; 8) galea of maxilla dorsoventrally flattened; 9) dentition of galea of maxilla variable, inner surface of galea lacking teeth or with reduced teeth (2 small, yet obvious teeth at the apex with 1 greatly reduced tooth at the base, presence or absence of medial teeth varies among species, teeth often obscured by dense setae); 10) apex of galea with dense brush of penicillate setae; 11) pronotum with broadly incomplete or complete beaded basal margin; 12) males with 2 or 3 protibial teeth, females with 3 protibial teeth, when 3 teeth are present, basal tooth reduced, removed from the more apical 2 teeth, and oriented laterally; 13) protibial spur straight to weakly deflexed or strongly deflexed; 14) males with inner protarsal claw enlarged and entire (not cleft with a small ramus) or narrowly cleft at apex; 15) mesocoxae widely separated; 16) meso- and metatibiae with distal, transverse carinae; 17) metacoxae with lateral edge acutely angled with respect to ventral surface; 18) anterior edge of hindwing distal to apical hinge lacking setae and with produced, membranous border; 19) vein RA with 2 rows of pegs extending distally nearly to margin of apical hinge.</p></div>	https://treatment.plazi.org/id/D8927393A343DAB9D5EB78C18D028AD6	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
66E7806F106DD08D25FB1870EC85CF9E.text	66E7806F106DD08D25FB1870EC85CF9E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Augoderia Burmeister 1847	<div><p>Augoderia Burmeister, 1847</p><p>Type species.</p><p>Augoderia nitidula Burmeister, 1847: 34, by monotypy.</p><p>Valid taxa.</p><p>Five species and subspecies.</p><p>The five species and subspecies of Augoderia are distributed in Argentina, Bolivia, Brazil, French Guiana, Peru, and Venezuela (Burmeister 1847, Harold 1869b, Arrow 1937b, Blackwelder 1944, Guimarães 1944, Martínez 1966, Gibbs et al. 1977, Endrődi 1966, 1967a, 1981, 1985a, Riehs 2005, Ronqui and Lopes 2006, Ponchel 2009, Grossi et al. 2011, Ratcliffe et al. 2015) (Fig. 55). Augoderia species are similar to some Cyclocephala in overall appearance, although three taxa ( A. giuglarisi Ponchel, A. nitidula nitidula, and A. nitidula yungana Martínez) are notable for their metallic, mother-of-pearl luster that reflects circularly polarized light, a cuticular trait that is rare in Dynastinae ( Endrődi 1967a, 1981, Ponchel 2009, Pye 2010). The biology of Augoderia species is completely unknown. Gibbs et al. (1977) reported A. nitidula as a floral visitor of Magnolia ovata, but this beetle was likely a misidentified Cyclocephala species (see Gottsberger et al. 2012, Moore and Jameson 2013). The immature stages are undescribed. Adults are attracted to lights at night (Riehs 2005, Ronqui and Lopes 2006, Grossi et al. 2011).</p><p>Augoderia, though maintained as a valid genus since Burmeister (1847), is poorly defined and diagnosed in the literature. The irregularly spaced punctures of the elytra and the mother-of-pearl sheen of some taxa are the only characters historically used to separate Augoderia from Cyclocephala . Thus, the genus has no clearly hypothesized synapomorphic characters. For example, many characters used to diagnose Augoderia in Endrődi’s (1985a) Dynastinae of the World are all variably present in Cyclocephala, Arriguttia, and Aspidolea species: 1) body short, convex; 2) dorsal coloration yellow, with dark maculae, and with or without metallic reflections; 3) mandibles of males with small anterolateral tooth, lacking in females; 4) frontoclypeal suture complete; 5) 10-segmented antennae with a short club in both sexes; 6) large eyes; 7) males with thickened protarsi; and 8) protibia tridentate in both sexes.</p><p>The following combination of characters can be used to recognize Augoderia species: 1) dorsal coloration yellowish or light brown, with or without elytral maculae, with or without metallic, mother-of-pearl sheen; 2) body not anteroposteriorly compressed or dorsoventrally flattened; 3) clypeal apex evenly rounded in dorsal view; 4) frons mesad of eyes with long, erect setae; 5) frontoclypeal suture complete; 6) males with anterolateral margin of mandibles weakly toothed; 7) mandibular molar area with rows of circular micropunctures; 8) apical margin of mentum weakly emarginated; 9) galea of the maxilla on inner surface with 3 fused basal teeth, a free median tooth, and 2 fused apical teeth (3-1-2 arrangement); 10) pronotum at base with incomplete or complete marginal bead; 11) pronotum on anterolateral portions with long, erect setae; 12) males and females with 3 protibial teeth, basal tooth reduced, removed from the apical 2 teeth, and oriented anteriorly; 13) protibial spur straight to weakly deflexed; 14) males with inner protarsal claw enlarged and narrowly cleft at apex; 15) mesocoxae widely separated; 16) metatibiae without distal, transverse carinae; 17) metacoxae with lateral edge perpendicular to ventral surface; 18) anterior edge of hindwing distal to apical hinge lacking setae and with produced, membranous border; 19) vein RA with 2 rows of pegs extending distally nearly to margin of apical hinge.</p></div>	https://treatment.plazi.org/id/66E7806F106DD08D25FB1870EC85CF9E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
4D3B7EDEB303C44E1FFCB0928B63E267.text	4D3B7EDEB303C44E1FFCB0928B63E267.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chalepides Casey 1915	<div><p>Chalepides Casey, 1915</p><p>Type species.</p><p>Parachalepus (Chalepides) eucephalus Casey, 1915, by original designation.</p><p>Valid taxa.</p><p>15 species.</p><p>The nomenclatural history of Chalepides was complicated by a case of homonymy. Chalepides was originally proposed as a subgenus of Parachalepus (Casey 1915). Parachalepus Casey, 1915 is a homonym of Parachalepus Baly, 1885 ( Coleoptera: Chrysomelidae) (Prell 1936, Arrow 1937a). To rectify this problem, Chalepides was elevated to the status of genus and comprised the seven species originally included in Parachalepus (Casey 1915, Prell 1936, Arrow 1937a). Parachalepus was proposed based on abdominal characters. Parachalepus included Dyscinetus -like species with a rigid fusion of the propygidium and the pygidium (Casey 1915). The subgenus Parachalepus (Chalepides) was proposed for species with a dramatic reduction of the pygidium in addition to propygidial/pygidial fusion (Casey 1915). Chalepides has been recognized as a valid genus by subsequent authors and was recently revised (Arrow 1937a, b, Endrődi 1966, 1985a, Joly and Escalona 2002).</p><p>The 15 species of Chalepides are distributed across South America and the West Indies ( Martínez 1978b, Endrődi 1966, 1973a, 1985a, Joly and Escalona 2002, Riehs 2005, Ratcliffe and Cave 2015) (Fig. 56). Species of Chalepides described by Prokofiev (2012) require a special discussion. Chalepides euhirtus Prokofiev and C. unduavicus Prokofiev were described based on specimens from Peru and Bolivia (Prokofiev 2012), and the Peruvian data would represent a new country record for Chalepides . However, both species were placed into the wrong genus, based on the original descriptions and images of the holotypes. The holotype of C. euhirtus appears to be a female specimen of A. fuliginea (Prokofiev 2012). Chalepides unduavicus was later synonymized under A. scarabaeoides and was also considered an infrasubspecific ( “ab.”) entity (Prokofiev 2013, 2014). The discussion below covering the biology and genus-level recognition of Chalepides will exclude information on the misclassified species C. euhirtus and C. unduavicus .</p><p>Relatively little is known about the biology and natural history of Chalepides species. It is unclear, based on available data, if Chalepides species are floral visitors. Mannerheim (1829) reported that C. dilatatus (Mannerheim) was collected in flowers without further detail. Valla and Cirino (1972) reported a single specimen of an unidentified Chalepides species from the inflorescence of a Victoria cruziana A.D. Orb. Chalepides barbatus adults and larvae are associated with sugar cane fields in Puerto Rico (Wolcott 1923, 1948). In Puerto Rico, adult C. barbatus are prey for the invasive cane toad R. marina (Wolcott 1937, 1948). Like Dyscinetus, Chalepides species may have some semi-aquatic habits. Chalepides luridus (Burmeister) and C. alliaceus (Burmeister) have been collected along the edges of river banks ( Endrődi 1973a). Chalepides barbatus reportedly attacks the invasive, aquatic weed water hyacinth ( Eichhornia crassipes [Mart.] Solms [ Pontederiaceae]) in Uruguay (Silveira Guido 1965, Perkins 1974, Buckingham and Bennett 1989). Chalepides species are attracted to lights at night (Kusui 1992, Riehs 2005, Albuquerque et al. 2016).</p><p>Chalepides species can be recognized by the following combination of characters: 1) dorsal coloration yellowish brown, dark brown, or almost black with greenish reflections in some species; 2) body convex, not strongly anteroposteriorly compressed or dorsoventrally flattened; 3) clypeus trapezoidal with apex truncate in dorsal view; 4) frontoclypeal suture complete or narrowly incomplete medially; 5) males with anterolateral margin of the mandibles lacking weak tooth; 6) mandibular molar area with rows of circular micropunctures; 7) mandibular molar area on proximal margin with 2 semicircular depressed pits; 8) galea of maxilla on inner surface with 2 fused basal teeth, 2 free medial teeth, and 2 fused apical teeth (2-2-2 arrangement); 9) pronotum with broadly incomplete beaded basal margin; 10) males and females with 3 protibial teeth on lateral margin, basal tooth not greatly reduced, only slightly removed from apical 2 teeth, and oriented laterally; 11) protibial spur straight to weakly deflexed; 12) males with inner protarsal claw enlarged and entire at apex, not cleft; 13) mesocoxae not widely separated, nearly touching; 14) metacoxae on lateral edge with transverse, depressed sulcus; 15) metacoxae with lateral edge perpendicular to ventral surface; 16) meso- and metatibiae with distal, transverse carinae; 17) anterior edge of hindwing distal to apical hinge with erect setae and lacking produced, membranous border; 18) vein RA with single row of pegs proximal to the apical hinge; 19) propygidium expanded, propygidium and pygidium fused, pygidium with long, dense setae.</p></div>	https://treatment.plazi.org/id/4D3B7EDEB303C44E1FFCB0928B63E267	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
04F3F6313AEA7F6A98F2061640B983DD.text	04F3F6313AEA7F6A98F2061640B983DD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cyclocephala Dejean 1821	<div><p>Cyclocephala Dejean, 1821</p><p>Type species.</p><p>Scarabaeus amazonus Linnaeus, 1767: 551, subsequent designation by Casey (1915).</p><p>Valid taxa.</p><p>359 species and subspecies.</p><p>The speciose genus Cyclocephala contains over 350 taxa distributed throughout the Nearctic and Neotropical realms (Fig. 57). Cyclocephala contains the only adventive species in Cyclocephalini, with C. pasadenae and C. signaticollis established in Hawaii and Australia, respectively (Carne 1956, Jameson et al. 2009). The greatest number of Cyclocephala species is found in northern South America, but many endemic species occur in Meso- and Central America. Some Cyclocephala species are extremely geographically widespread. For example, C. lunulata occurs from the southwestern United States south to Argentina. In contrast, there are also cases of endemism in mainland species of the genus. The pollination mutualist C. jalapensis occurs only in a narrow band of habitat in eastern Mexico (Veracruz, Puebla, Oaxaca, Querétaro, and Hidalgo states) where its host plant Magnolia schiedeana Schltl. is found (Dieringer and Delgado 1994, Dieringer and Espinosa 1994).</p><p>Cyclocephala is a difficult genus to diagnose due to its species richness, diversity of forms, and probable non-monophyly. Many of the character descriptions below are complicated by these factors. Cyclocephala species can be recognized by the following combination of characters: 1) dorsal coloration highly variable; unicolored black, green, or light brown, pronotum in some species cherry-red, light brown species often have complex maculae patterns of the pronotum and elytra; 2) body not anteroposteriorly compressed or dorsoventrally flattened; 3) clypeal apex variable; evenly rounded, parabolic, acute, emarginate, triemarginate, or nearly straight; 4) frons mesad of eyes with or without long, erect setae; 5) frontoclypeal suture complete or incomplete medially; 6) males with anterolateral margin of mandibles weakly toothed or not; 7) mandibular molar area with rows of circular micropunctures either present or absent; 8) apical margin of mentum weakly emarginated or broadly and deeply emarginated; 9) galea of the maxilla well-developed [with or without teeth] or reduced into a rounded process; 10) galea of the maxilla dorsoventrally flattened or not; 10) galea of maxilla on inner surface variable (not all character states are given here); with 3 fused basal teeth, a free median tooth, and 2 fused apical teeth (3-1-2 arrangement) (in C. amazona -like species and former Mimeoma, the galea are flattened and the basal tooth is compressed and rotated, giving the appearance of being bidentate with the third tooth shifted dorsally); with 2 fused basal tooth and 2 fused apical teeth (2-0-2 arrangement); with 2 fused basal teeth, 1 middle tooth, and 2 fused apical teeth (2-1-2 arrangement); 11) pronotum at base with incomplete or complete marginal bead; 12) pronotum on anterolateral portions with or without long, erect setae; 13) males with 2 or 3 protibial teeth, females always with 3; 14) protibial spur straight to weakly deflexed or strongly decurved; 15) males with inner protarsal claw enlarged and narrowly cleft at apex or entire at apex; 16) mesocoxae widely separated or nearly touching, contiguous; 17) metatibiae with or without distal, transverse carinae; 18) metacoxae with lateral edge perpendicular to ventral surface or with lateral edge angled underneath the ventral surface; 19) anterior edge of hindwing distal to apical hinge lacking setae and with produced, membranous border or lacking membranous border and with decumbent setae ( C. cribrata species-group); 20) vein RA with 2 rows of pegs extending distally nearly to margin of apical hinge.</p></div>	https://treatment.plazi.org/id/04F3F6313AEA7F6A98F2061640B983DD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
77E1BB001FC725CE151A09B47286947C.text	77E1BB001FC725CE151A09B47286947C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Dyscinetus Harold 1869	<div><p>Dyscinetus Harold, 1869</p><p>Type species.</p><p>Melolontha geminata Fabricius, 1801, by monotypy.</p><p>Valid taxa.</p><p>21 species.</p><p>The genus Dyscinetus comprises 21 species distributed from North America south to Argentina and the West Indies (Fig. 58). Smooth, large, and mostly black Dyscinetus species superficially resemble hydrophilid beetles. Dyscinetus is generally not considered an aquatic or semiaquatic genus. However, some species in the genus have an intriguing association with moist, mucky soils and aquatic plants. Dyscinetus rugifrons and another Dyscinetus sp. attack water hyacinth in Uruguay (Silveira Guido 1965, Bennett and Zwolfer 1968, Perkins 1974). Dyscinetus morator also attacks water hyacinth in Florida (Perkins 1974, Buckingham and Bennett 1989). These species are considered scavengers and enhancers of damage started by other arthropods on water hyacinth, though they are known to attack healthy tissues (Perkins 1974, Buckingham and Bennett 1989). Feeding damage on water hyacinth occurs inside petioles, crowns, petiole bases, and submerged roots (Perkins 1974, Buckingham and Bennett 1989).</p><p>Experiments indicated that D. morator can survive submerged in water for up to 36 hours (Buckingham and Bennett 1989). The mechanism allowing for this prolonged submersion is unclear. Air bubbles are visible along the elytral margin and on the metathorax in submerged individuals (Buckingham and Bennett 1989). Dyscinetus laevipunctatus Bates was also observed submerged in association with water hyacinth in Mexico ( García-Rivera and Contreras-Ramos 2015). Unlike many other genera in the group, Dyscinetus adults are not known to visit flowers. A Brazilian Dyscinetus species was reportedly attracted to the floral odors of Annona sp., although these beetles were not encountered in any inflorescences (Gottsberger 1989). This is the only mention of Dyscinetus floral attraction in the literature.</p><p>Dyscinetus species can be recognized by the following combination of characters: 1) dorsal coloration dark piceous to black; 2) body convex, not strongly anteroposteriorly compressed or dorsoventrally flattened; 3) clypeus trapezoidal with apex truncate in dorsal view; 4) frontoclypeal suture complete medially; 5) males with anterolateral margin of the mandibles lacking weak tooth; 6) mandibular molar area with rows of circular micropunctures; 7) mandibular molar area on proximal margin with 2 semicircular depressed pits; 8) galea of maxilla on inner surface with 2 fused basal teeth, 2 free medial teeth, and 2 fused apical teeth (2-2-2 arrangement); 9) pronotum with broadly incomplete beaded basal margin; 10) males and females with 3 protibial teeth on lateral margin, basal tooth not greatly reduced, only slightly removed from the more apical 2 teeth, and oriented laterally; 11) protibial spur straight to weakly deflexed; 12) males with inner protarsal claw enlarged and narrowly cleft at apex; 13) mesocoxae not widely separated, nearly touching; 14) metacoxae on lateral edge with transverse, depressed sulcus; 15) metacoxae with lateral edge perpendicular to ventral surface; 16) meso- and metatibiae with distal, transverse carinae; 17) anterior edge of hindwing distal to apical hinge with erect setae and lacking produced, membranous border; 18) vein RA with single row of pegs proximal to apical hinge; 19) propygidium not expanded, with propygidium and pygidium not fused.</p></div>	https://treatment.plazi.org/id/77E1BB001FC725CE151A09B47286947C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
BDB2A805D58ED5DC1C82B5B366EE731F.text	BDB2A805D58ED5DC1C82B5B366EE731F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Erioscelis Burmeister 1847	<div><p>Erioscelis Burmeister, 1847</p><p>Type species.</p><p>Apogonia emarginata Mannerheim, 1829, by monotypy.</p><p>Valid taxa.</p><p>Five species.</p><p>The five species of Erioscelis are distributed in South America north to Nicaragua (Fig. 59). Erioscelis species are remarkable among cyclocephalines for their well-characterized floral visitation syndromes. Erioscelis species are associated with nocturnally blooming genera in the family Araceae . Three Erioscelis species have been reported from the spathes of Dieffenbachia, Philodendron Schott, Syngonium Schott, Montrichardia Crueg., and possibly Xanthosoma Schott (Schrottky 1910, Gottsberger and Amaral 1984, Young 1986, Grayum 1996, Croat 1997, Morón 1997, Beath 1998, 1999, Gibernau et al. 2003). While the association between Erioscelis species and aroid flowers is firmly established, there is little evidence of species- or genus-level specificity in this pollination mutualism. For example, Erioscelis columbica Endrődi has been collected from the spathes of nine different Philodendron species in Heredia, Costa Rica (Grayum 1996, Croat 1997, Morón 1997, Moore and Jameson 2013). Based on feeding damage to Philodendron inflorescences by Erioscelis, it was hypothesized that this genus may be an interloper on the cyclocephaline/aroid mutualism (Goldwasser 1987). Other observations seem to indicate that Erioscelis species are part of this mutualism.</p><p>The descriptions of Erioscelis spp. visitation of Dieffenbachia and Philodendron inflorescences are some the most detailed available for Cyclocephalini . In Costa Rica, E. columbica is a pollinator of Dieffenbachia nitidipetiolata Croat &amp; Grayum (Young 1986, 1988a, 1988b, 1990). Erioscelis columbica arrive at receptive female-phase inflorescences during nightfall, where they feed on staminodia and mate (Young 1986). The beetles exit the spathe after 24 hours when the spadix is in the male-phase and shedding pollen (Young 1986). Erioscelis columbica are covered in sticky pollen grains while exiting the spathe, and they may also feed on some of the pollen (Young 1986). Erioscelis proba (Sharp) displays similar behavior in the inflorescences of two other Dieffenbachia species in French Guiana (Gibernau 2015a).</p><p>Observational and experimental evidence suggests that Erioscelis emarginata (Mannerheim) prefers to feed upon sterile staminate flowers on the spadix in two Philodendron species (Maldonado et al. 2015). Furthermore, analyses of nutritional and defensive compound (calcium oxalate) content of sterile and fertile flowers in these Philodendron species suggested that sterile staminate flowers have lower amounts of defensive compounds (Maldonado et al. 2015). Erioscelis species are seemingly attracted to the strong floral scents that are volatilized during thermogenesis and receptivity of the staminate flowers in these aroids. The dynamics of floral scent attraction are mostly unexplored for Erioscelis . In the case of Philodendron adamantium Mart. ex Schott, a single dominant flower scent compound (Dihydro-β-ionone) extracted from this species was sufficient to attract E. emarginata to scent traps (Pereira et al. 2014).</p><p>Erioscelis was first revised by Saylor (1946) and again by Endrődi (1966, 1985a). These works provide a strong foundation for species-level identification, but characters that separate Erioscelis from other cyclocephalines are largely undiscussed. For example, Saylor (1946) commented, "When compared with such species as Cyclocephala (Stigmalia) mafaffa Burmeister, or C. (Aclinidia) castanea (Fabricius), the only character definitely to separate Erioscelis is the unenlarged front tarsal claws of both sexes". Unique protibial (2 teeth on the lateral margin in both sexes, subapical position of reduced protibial spur) and abdominal (bisinuate margin of 6th abdominal sternite, terminal spiracle not positioned on pleural suture) characters of Erioscelis emarginata also complicate recognition of the genus and may be reasons to doubt the monophyly of the group. These characters (except for the bisinuate margin of 6th abdominal sternite) are associated with Anomalini ( Rutelinae) and are absent in all other members of Erioscelis and Cyclocephalini more broadly. Sister-relationships of Erioscelis have not been hypothesized and the immature stages are unknown for the genus.</p><p>Erioscelis species can be recognized by the following combination of characters: 1) dorsal coloration castaneous, rufocastaneous, or piceous; 2) body not dorsoventrally flattened nor anteroposteriorly compressed; 3) clypeal apex truncate, weakly emarginate, or deeply emarginate in dorsal view; 4) frontoclypeal suture complete medially; 5) apical margin of mentum shallowly emarginate; 6) anterolateral margin of mandible lacking tooth; 7) mandibular molar area with rows of circular micropunctures; 8) galea of maxilla not dorsoventrally flattened; 9) galea of maxilla on inner surface with 6 teeth in 2-2-2 arrangement (each pair shares a base); 10) pronotum with apical bead complete medially; 11) basal bead of pronotum incomplete medially; 12) anterior membrane of pronotum straight at middle, not projected anteriorly; 13) anterior membrane of the pronotum extending laterally to apicolateral margins of the pronotum; 14) protibia with 2 or 3 lateral teeth in both sexes; 15) when protibia tridentate, basal tooth not greatly reduced, only slightly removed from the apical 2 teeth, and oriented laterally; 16) protibial spur subapical or apically positioned; 17) protibial spur straight to weakly reflexed; 18) males and females with protarsal claws simple, not enlarged; 19) males and females with inner protarsal claws with apex entire, not cleft; 20) mesocoxae not widely separated, nearly touching; 21) metacoxae with lateral edge perpendicular to ventral surface; 22) anterior edge of hindwing distal to apical hinge simple (lacking setae or membrane) or with row of long, erect setae extending along vein; 23) vein RA with double row of pegs proximal to apical hinge; 24) terminal abdominal spiracle situated on pleural suture or not.</p></div>	https://treatment.plazi.org/id/BDB2A805D58ED5DC1C82B5B366EE731F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
F094999B3517A55A792B927FBFE083D5.text	F094999B3517A55A792B927FBFE083D5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Harposceles Burmeister 1847	<div><p>Harposceles Burmeister, 1847</p><p>Type species.</p><p>Harposceles paradoxus Burmeister, 1847: 35, by monotypy.</p><p>Valid taxa.</p><p>One species.</p><p>The monotypic genus Harposceles was erected for the species H. paradoxus . This striking, relatively large cyclocephaline occurs in lowland forests in Brazil, Ecuador, French Guiana, Peru, Suriname, and possibly Colombia (Burmeister 1847, Harold 1869b, Arrow 1937b, Blackwelder 1944, Endrődi 1966, 1985a, Endrődi and Dechambre 1976, Lachaume 1992, Couturier and Kahn 1992, Andreazze 2001, Andreazze and da Silva Motta 2002, Touroult et al. 2010, Ponchel 2011, Saltin and Ratcliffe 2012, Ratcliffe et al. 2015) (Fig. 60). Males display dramatic, and unique, characters of the protibia. Harposceles paradoxus males have elongated, arcuate protibia with the protibial spurs fused to the base of the tibia. Females are much less common than male specimens in collections, and males are readily attracted to lights at night, especially between midnight and 4 am (Andreazze 2001, Andreazze and da Silva Motta 2002, Touroult et al. 2010, Saltin and Ratcliffe 2012). The immature stages of H. paradoxus are associated with the palms Astrocaryum chonta Mart. and A. carnosum F. Kahn &amp; B. Millán ( Arecaceae) (Couturier and Kahn 1992). The larvae and pupae were found in the organic litter accumulated between leaf sheaths of A. carnosum (Couturier and Kahn 1992). The immature stages are undescribed.</p><p>Harposceles species can be recognized by the following combination of characters: 1) dorsal coloration dark piceous to black; 2) body dorsoventrally flattened; 3) clypeus rounded in dorsal view; 4) frontoclypeal suture incomplete medially; 5) apical margin of mentum truncate; 6) anterolateral margin of mandible lacking tooth; 7) mandibular molar area with surface lacking circular pits, with large, disorganized, canal-like invaginations; 8) galea of maxilla dorsoventrally flattened; 9) galea on inner surface at base with large, flattened, blade-like, tooth (less produced than in Surutu species); 10) galea on inner surface with 7 teeth in 2-1-1-1-2 arrangement from base to apex; 11) apical and basal beaded margins of pronotum complete at middle; 12) anterior membrane of the pronotum interrupted before lateral pronotal margins; 13) males with protrochanter with ventrally produced tooth; 14) protibia with 3 teeth in both sexes; 15) males with protibia elongated and arcuate; 16) protibial spur straight to weakly reflexed; 17) males with protibial spur fused to protibia, not articulated at its base; 18) males with inner protarsal claw thickened and not cleft at apex; 19) mesocoxae not widely separated, nearly touching; 20) metacoxae with lateral edge perpendicular to ventral surface; 21) apices of the meso- and metatibiae with a corbel; 22) anterior edge of hindwing distal to apical hinge lacking membranous border; 23) anterior edge of hindwing distal to apical hinge with decumbent setae surrounding vein and originating away from apical hinge; 24) vein RA with single row of pegs proximal to apical hinge.</p><p>The relationship of Harposceles to other cyclocephalines has not been elaborated upon in the literature. However, H. paradoxus shares some characters with Surutu that may be indicative of a close relationship between the two genera. The rounded shape of the clypeal apex in H. paradoxus is like the clypeal form in S. dytiscoides . The single row of RA pegs in H. paradoxus is shared between Ancognatha and Surutu, though Ancognatha species lack setae on the anterior edge of the hindwing distal to the apical hinge. The decumbent setae of the hindwing leading edge (distal to apical hinge) found in H. paradoxus is also found in Surutu species and the " Cyclocephala cribrata species group" (which included species previously placed in Mononidia and Surutoides) (Dechambre 1997). These groups also all share corbeled meso- and metatibial apices and entirely black coloration. Harposceles paradoxus shares other interesting characters with Surutu species. These shared characters include: 1) body strongly dorsoventrally flattened; 2) dorsoventrally flattened maxillary galea; 3) a seven-toothed maxillary galea in a 2-1-1-1-2 arrangement from the base to apex; 4) an incomplete frontoclypeal suture; and 5) the apical pronotal membrane interrupted before the lateral margins of the pronotum. The large basal tooth of the maxillary galea is much smaller and less produced in H. paradoxus than in Surutu species. Several male characters of H. paradoxus are autapomorphic in Cyclocephalini: 1) the protibial spur fusion to the protibial; 2) the arcuate, elongated protibia (seen also in some Dynastini); and 3) the ventrally produced protrochanter teeth.</p></div>	https://treatment.plazi.org/id/F094999B3517A55A792B927FBFE083D5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
421163B0212FD6D753C0C9BAD6E0E7A4.text	421163B0212FD6D753C0C9BAD6E0E7A4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Peltonotus Burmeister 1847	<div><p>Peltonotus Burmeister, 1847</p><p>Type species.</p><p>Peltonotus morio Burmeister, 1847: 75, by monotypy.</p><p>Valid taxa.</p><p>25 species.</p><p>Peltonotus species are distributed throughout Southeast Asia, southern China, and the eastern portion of the Indian Subcontinent (Fig. 61). Peltonotus is currently considered the sole Asian lineage of Cyclocephalini, though its subfamilial classification has been unstable. The genus is remarkable for its confounding combination of morphological and behavioral traits that blurred the lines between historical concepts of the subfamilies Dynastinae and Rutelinae . For example, the sexual dimorphism of the protarsi in Peltonotus species has long been compared to that found in Cyclocephala (e.g., see Burmeister 1847). In contrast, the labral morphology of Peltonotus species matches that found in Asian parastasiine and fruhstoferiine ( Rutelinae) scarabs (Arrow 1908, 1910). The floral feeding behavior of Peltonotus species on Araceae is also shared between cyclocephalines and Asian parastasiines, adding a further layer of intrigue to unresolved evolutionary relationships between the groups at the subfamilial- and tribal-level (e.g., see Moore and Jameson 2013, Kumano-Nomura and Yamaoka 2006, Kumano-Nomura and Yamaoka 2009, Tung et al. 2010, Hoe et al. 2011, 2016).</p><p>Peltonotus was described by Burmeister (1847), and he included it within the Chalepidae division of Cyclocephalidae . The classification of Peltonotus was stable until Arrow (1908, 1910) transferred the genus to Rutelinae based upon the exposed (in dorsal view, produced apically beyond the clypeus) and chitinized labrum. Arrow (1917) later erected the “division” Peltonotini for Peltonotus within his classification of Rutelinae . Ohaus (1918, 1934b) and Machatschke (1972) rejected Peltonotini and included Peltonotus in Pelidnotina ( Rutelini) in their catalogs of Rutelinae . Morphological phylogenetic analysis of Rutelina ( Rutelinae: Rutelini) suggested that Peltonotus were more closely related to Cyclocephalini than Rutelini (Jameson 1998). Subsequent works on the genus have treated Peltonotus as a member of Cyclocephalini (Jameson and Wada 2004, 2009, Jameson and Jákl 2010, Jameson and Drumont 2013).</p><p>Little is known about the biology and natural history of Peltonotus species. The immatures are undescribed. Adults are attracted to lights at night (Jameson and Wada 2004). Peltonotus malayensis Arrow was collected from the spathes of Epipremnum falcifolium Engl. ( Araceae), where males and females were observed mating and feeding (Jameson and Wada 2004). In Thailand, P. nasutus visit the large inflorescences of the terrestrial aroid Amorphophallus paeoniifolius (Dennst.) Nicolson, where adult beetles feed and mate (Grimm 2009). Peltonotus nasutus can be attracted to the inflorescences in high numbers (over 70 individuals) (Danell 2010).</p><p>Peltonotus species can be recognized by the following combination of characters: 1) dorsal coloration brown to black with variable presence of maculae; 2) body convex, not dorsoventrally flattened; 3) clypeal apex rounded to straight in dorsal view; 4) frontoclypeal suture incomplete medially; 5) apical margin of mentum variably shaped with weak emargination; 6) anterolateral margin of mandible lacking tooth; 7) mandibular molar area with rows of circular micropunctures; 8) galea of maxilla not strongly dorsoventrally flattened; 9) galea of the maxilla on inner surface with 3 fused basal teeth, a free median tooth, and 2 fused apical teeth (3-1-2 arrangement); 10) galea with articulated medial tooth; 11) labrum extending apically beyond clypeal apex (obvious in dorsal view); 12) apical and basal margins of pronotum with beaded margin complete or incomplete at middle; 13) protibia of males with 2 or 3 teeth, females with 3 teeth; 14) protibial spur straight to weakly reflexed; 15) males with inner protarsal claw thickened and not cleft at apex (nib variably present or absent); 16) mesocoxae not widely separated, nearly touching; 17) metacoxae with lateral edge perpendicular to ventral surface; 18) anterior edge of hindwing distal to apical hinge lacking membranous border; 19) anterior edge of hindwing distal to apical hinge with row of long setae extending from apical hinge along length of the costal vein; 20) vein RA with single row of pegs proximal to apical hinge.</p></div>	https://treatment.plazi.org/id/421163B0212FD6D753C0C9BAD6E0E7A4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
9E44E2A8C7B92A00690D7E84C2EA0B48.text	9E44E2A8C7B92A00690D7E84C2EA0B48.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ruteloryctes Arrow 1908	<div><p>Ruteloryctes Arrow, 1908</p><p>Types species.</p><p>Ruteloryctes tristis Arrow, 1908: 336, by monotypy.</p><p>Valid taxa.</p><p>Two species.</p><p>The two species of Ruteloryctes are distributed in the Guinea-Congo lowland rainforests of West and Central Africa. Ruteloryctes specimens have been collected in Angola, Benin, Cameroon, Chad, Côte d’Ivoire, Democratic Republic of the Congo, Guinea, Guinea-Bissau, Nigeria, Senegal, Sierra Leone, and The Gambia (Burgeon 1947, Paulian 1954, Endrődi 1960, 1966, 1985a, Krell et al. 2003, Hirthe and Porembski 2003, Ervik and Knudsen 2003) (Fig. 62). Ruteloryctes morio is a pollinator of nocturnally blooming Nymphaea lotus L., and this floral association has been reported from Côte d’Ivoire, Senegal, and Nigeria (Fabricius 1798, Krell et al. 2003, Hirthe and Porembski 2003, Ervik and Knudsen 2003). The immature stages of Ruteloryctes are undescribed.</p><p>Ruteloryctes species can be recognized by the following combination of characters: 1) dorsal coloration black to dark brown; 2) body convex, not strongly anteroposteriorly compressed or dorsoventrally flattened; 3) clypeal apex truncate or rounded in dorsal view; 4) frontoclypeal suture incomplete medially; 5) males with anterolateral margin of the mandibles lacking weak tooth; 6) mandibular molar area with rows of circular micropunctures; 7) apex of mentum weakly emarginated at middle; 8) galea of maxilla on inner surface with 3 fused basal teeth, a free median tooth, and 2 fused apical teeth (3-1-2 arrangement); 9) pronotum with broadly incomplete beaded basal margin; 10) males and females with 3 protibial teeth on lateral margin, basal tooth not greatly reduced, slightly removed from apical 2 teeth, and oriented laterally; 11) protibial spur straight to weakly deflexed; 12) males with inner protarsal claw enlarged and narrowly cleft at apex; 13) mesocoxae not widely separated, nearly touching; 14) meso- and metatibiae with distal, transverse carinae; 15) metacoxae with lateral edge perpendicular to ventral surface; 16) anterior edge of hindwing distal to apical hinge lacking setae and with produced, membranous border; 17) vein RA with single row of pegs proximal to apical hinge.</p><p>The original description of Ruteloryctes compared the genus to New World Dyscinetus species, and it was hypothesized to have "strayed across the Atlantic" (Arrow 1908). Endrődi (1966) thought that Ruteloryctes was one of the most “primitive” cyclocephaline genera. The 3-1-2 arrangement of the teeth on the maxillary galea in Ruteloryctes is most similar to Arriguttia, Augoderia, and many Cyclocephala species. The membranous border of the hindwing present in Ruteloryctes is also shared with Arriguttia, Acrobolbia, Ancognatha, Aspidolea, and Cyclocephala . However, the single row of pegs present on the hindwing RA vein in Ruteloryctes is present in Ancognatha, Surutu, Harposceles, Stenocrates, Dyscinetus, Erioscelis, and Chalepides .</p></div>	https://treatment.plazi.org/id/9E44E2A8C7B92A00690D7E84C2EA0B48	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
4FA25544FD25CE9F1A025B5C5C36D2E3.text	4FA25544FD25CE9F1A025B5C5C36D2E3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Stenocrates Burmeister 1847	<div><p>Stenocrates Burmeister, 1847</p><p>Type species.</p><p>Scarabaeus laborator Fabricius, subsequent designation by Casey 1915: 114.</p><p>Valid taxa.</p><p>52 species and subspecies.</p><p>The enigmatic genus Stenocrates comprises 52 taxa distributed from Mexico south throughout South America (except Chile) and Jamaica (Fig. 63). Species diversity in the group is highest in the tropical forests of Brazil, especially the northern and western states of Amazonas, Pará, Acre, and Rondônia . Many Stenocrates species are also known from eastern Brazil, especially Bahia, Espírito Santo, São Paulo, and Santa Catarina. Stenocrates species are problematic to identify due to conserved external morphology among species, making the group, “… possibly the most difficult genus of Dynastinae in the Americas with which to work" (Ratcliffe and Cave 2015). Male paramere morphology is diagnostic for species-level identification in the genus, and females not associated with males at the time of collection cannot be reliably identified with existing literature. Nothing is known about the natural history and biology of Stenocrates species. Adults can be collected at lights at night ( Endrődi 1969a, Ratcliffe and Cave 2006, Ratcliffe 2014, 2015). Immature stages are undescribed for the genus.</p><p>Stenocrates was erected by Burmeister (1847) for species that he considered highly similar to the historical concept of Chalepus, except for the lack of dimorphic protarsi. Burmeister (1847) included 4 species in Stenocrates and speculated that Melolontha rufipennis Fabricius could also be a member of the genus. Descriptions of new species of Stenocrates were slow to accumulate in the 19th and early 20th century. Kirsch (1870) described the sixth Stenocrates species from Colombia. Bates (1888) examined S. laborator specimens from Mexico and noted that the simple protarsi of the males and dorsoventrally flattened tibiae separated diagnosed Stenocrates within Cyclocephalini . Stenocrates was compared to Euetheola by Bates (1888) stating that the form of the mandibles and the proximal tarsomeres served to separate these genera. Arrow (1911, 1913) added two new species to Stenocrates, but he did not offer a diagnosis for the genus or make meaningful character comparisons for the genus. Stenocrates was revised by Endrődi (1966, 1985a), and many new species have been described since that work, which have not been incorporated into a comprehensive identification key.</p><p>Stenocrates species can be recognized by the following combination of characters: 1) dorsal coloration black or dark brown and without maculae; 2) body convex, not strongly anteroposteriorly compressed or dorsoventrally flattened; 3) clypeus trapezoidal with apex truncate in dorsal view; 4) frontoclypeal suture complete medially; 5) males with anterolateral margin of the mandibles lacking weak tooth; 6) mandibular molar area with rows of circular micropunctures; 7) mandibular molar area on proximal margin without semicircular depressed pits; 8) galea of maxilla on inner surface with 2 fused basal teeth, 2 fused medial teeth, and 2 fused apical teeth (2-2-2 arrangement); 9) pronotum with broadly incomplete beaded basal margin; 10) pronotum with narrowly incomplete beaded apical margin; 11) males and females with 3 protibial teeth on lateral margin, basal tooth not greatly reduced, only slightly removed from apical 2 teeth, and oriented laterally; 12) protibial spur straight to weakly deflexed; 13) males and females with protarsal claws simple (not cleft) and not enlarged; 14) mesocoxae not widely separated, nearly touching; 15) metacoxae on lateral edge without transverse, depressed sulcus; 16) metacoxae with lateral edge perpendicular to ventral surface; 17) meso- and metatibiae with distal, transverse carinae; 18) meso- and metatibiae dorsoventrally flattened and laterally expanded; 19) anterior edge of hindwing distal to apical hinge with erect setae and lacking produced, membranous border; 20) vein RA with single row of pegs proximal to apical hinge; 21) propygidium not expanded, propygidium and pygidium not rigidly fused.</p></div>	https://treatment.plazi.org/id/4FA25544FD25CE9F1A025B5C5C36D2E3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
892A43C226717AA2B9E45E65AC972314.text	892A43C226717AA2B9E45E65AC972314.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Surutu Martinez 1955	<div><p>Surutu Martinez, 1955</p><p>Type species.</p><p>Surutu dytiscoides Martínez, 1955: 245-249, by monotypy.</p><p>Valid taxa.</p><p>Five species.</p><p>The five species of the South American genus Surutu are distributed in Colombia, Bolivia, and Brazil ( Martínez 1955, D’Andretta and Martínez 1956, Endrődi 1966, 1975a, 1985a, Ratcliffe 1981, Andreazze 2001, Otavo et al. 2013) (Fig. 64). These spectacular black species are truly the monsters of the Cyclocephalini, with some specimens of Surutu seabrai D’Andretta and Martínez measuring over 4 cm in length. Nothing is known about the biology of Surutu species. At least some species are attracted to lights at night (Ratcliffe 1981). The immature stages are undescribed for the genus as currently circumscribed.</p><p>Surutu species can be recognized by the following combination of characters: 1) dorsal coloration dark piceous to black; 2) body dorsoventrally flattened; 3) clypeus rounded to parabolic in dorsal view; 4) frontoclypeal suture incomplete medially; 5) apex of mentum narrowly and deeply emarginated (in S. dytiscoides and S. seabrai; other species unknown); 6) anterolateral margin of mandible lacking tooth; 7) galea of maxilla dorsoventrally flattened (in S. dytiscoides and S. seabrai; other species unknown); 8) galea on inner surface at base with large, flattened, blade-like, bifurcated tooth (in S. dytiscoides and S. seabrai; other species unknown); 9) galea on inner surface with 7 teeth in 2-1-1-1-2 arrangement from base to apex (in S. dytiscoides and S. seabrai; other species unknown); 10) apical and basal beaded margins of pronotum incomplete at middle (in S. dytiscoides and S. seabrai; other species unknown); 11) anterior membrane of the pronotum interrupted before lateral pronotal margins (in S. dytiscoides and S. seabrai; other species unknown); 12) protibia with 3 teeth in both sexes; 13) protibial spur straight to weakly reflexed; 14) males with protibial spur articulated at base, not fused to protibia; 15) males with inner protarsal claw thickened and narrowly cleft at apex (claw apex entire in S. fenni Ratcliffe and S. schulzei Endrődi); 16) mesocoxae not widely separated, nearly touching; 17), metacoxae with lateral edge perpendicular to ventral surface; 18) apices of the meso- and metatibiae with a corbel (in S. dytiscoides and S. seabrai; other species unknown); 19) anterior edge of hindwing distal to apical hinge lacking membranous border; 20) anterior edge of hindwing distal to apical hinge with decumbent setae surrounding the vein and originating away from the hinge; 21) vein RA with single row of pegs proximal to apical hinge.</p><p>Some characters of the head, mouthparts, and elytra of Surutu have been compared to Ancognatha, Cyclocephala, and Mimeoma ( Martínez 1955, D’Andretta and Martínez 1956). The parabolic and rounded clypeal apex in Surutu species is like the clypeal form in several Ancognatha species. Surutu dytiscoides and S. seabrai, at least, have a deeply emarginated apex of the mentum that is also shared with Ancognatha species. The single row of RA pegs is also shared between Ancognatha and Surutu, although Ancognatha species lack setae on the anterior edge of the hindwing distal to the apical hinge. Instead, Ancognatha have a hindwing membrane like that found in Cyclocephala, Augoderia, Arriguttia, Aspidolea, and Acrobolbia . The dramatic dilations and knobs on the elytral epipleuron of S. seabrai are similar to those found in some Ancognatha and Cyclocephala species.</p><p>The distinctive setae of the hindwings found in Surutu are also found in Harposceles and species of the " Cyclocephala cribrata species group" (which included species previously placed in the genera Mononidia and Surutoides) (Dechambre 1997). These groups also share corbeled meso- and metatibial apices and entirely black coloration. Harposceles paradoxus shares other interesting characters with Surutu species, suggestive of a close relationship between the two genera. These shared characters include: 1) body strongly dorsoventrally flattened; 2) dorsoventrally flattened maxillary galea; 3) a 7-toothed maxillary galea in a 2-1-1-1-2 arrangement from the base to apex; 4) an incomplete frontoclypeal suture; and 5) the apical pronotal membrane interrupted before the lateral pronotal margins.</p><p>Platyphileurus felscheanus Ohaus ( Dynastinae: Oryctini) warrants special discussion here. This species was described twice. Platyphileurus felscheanus was described from specimens collected from Santa Catarina, Brazil (Ohaus 1910). This new genus was compared to Phileurus Latreille and later included in the tribe Phileurini (Ohaus 1910, Arrow 1937b). Endrődi (1975) later described Surutu jelineki from Rio de Janeiro based on two female specimens. Comparison of the types of these species revealed that they are conspecific, with the name Platyphileurus felscheanus having priority over Surutu jelineki (Grossi et al. 2010).</p><p>The immatures of Platyphileurus felscheanus are associated with bromeliads (Grossi et al. 2010, Albertoni et al. 2014). Based on examination of larval, pupal, and adult characters, P. felscheanus was excluded from Phileurini and proposed to be a member of Oryctini (Albertoni et al. 2014). However, there are some intriguing adult character similarities between P. felscheanus and other Surutu species. For example, P. felscheanus is black, dorsoventrally flattened, and has dimorphic protarsal claw morphology (enlarged in males, simple in females) ( Endrődi 1975, Grossi et al. 2010, Albertoni et al. 2014). The apices of the metatibiae in P. felscheanus are "weakly dentate" (Albertoni et al. [2014]: figure 30). Alternatively, the outer edge of the metatibia figured in Albertoni et al. (2014) could be considered not to be "weakly dentate", but corbeled (outer edge produced beyond the inner edge of the tibial apex). This tibial character is found in Surutu, Harposceles, and in the " Cyclocephala cribrata species group". The venter of the meso- and metatarsi in P. felscheanus is covered with dense, reddish, flattened setae (Albertoni et al. 2014). Similar flattened, scale-like setae are also found on the venter of the meso- and metatarsi of S. seabrai and S. dytiscoides . Future analyses of the tribal placement of P. felscheanus should focus on adult character comparisons with Surutu species and H. paradoxus, especially characters of the mandibles, maxillary galea, tibiae, tarsi, parameres, and hind wings.</p></div>	https://treatment.plazi.org/id/892A43C226717AA2B9E45E65AC972314	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Moore, Matthew R.;Cave, Ronald D.;Branham, Marc A.	Moore, Matthew R., Cave, Ronald D., Branham, Marc A. (2018): Synopsis of the cyclocephaline scarab beetles (Coleoptera, Scarabaeidae, Dynastinae). ZooKeys 745: 1-99, DOI: http://dx.doi.org/10.3897/zookeys.745.23683, URL: http://dx.doi.org/10.3897/zookeys.745.23683
