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Subfamily Melolonthinae Our diagnosis of the Melolonthinae below is based largely on Evans (2002). Diagnosis. Scarabaeidae. Adults 3���60 mm in length. Dorsal surface glabrous, setose or scaled; sometimes setae or scales forming distinct patches or lines; color testaceous to reddish brown or black, occasionally with metallic blue, green, coppery, or brassy luster. Head usually unarmed, never with frontal extensions or horns. Eyes only partially divided by ocular canthus. Antennae with 7���10 antennomeres, rarely more, and insertions not visible from above; lamellate club usually consisting of 3���7 antennomeres; lamellae glabrous or with only a few setae, often longer in males. Mandibles sclerotized, usually well developed, and concealed in dorsal view. Transverse, narrowed, or conical labrum free or fused beneath clypeus, sometimes extending beyond or fused to apical clypeal margin. Thorax with pronotal disc more-or-less convex, never concave or with raised carinae; scutellum exposed. Elytra with lateral margins more-or-less straight behind humerus, with mesepimeron usually covered by elytral bases. Metathoracic wings fully developed or reduced. Metepimeron rectangular or triangular. Legs with procoxae transverse or conical; protibiae usually with apical spurs; mesotibiae and metatibiae usually with a pair of apical spurs located below or set on either side of tarsal articulation, or with upper spur adjacent to tarsal articulation; opposing mesotarsal and metatarsal claws equal in thickness (except Hoplia, which have a single, enlarged metatarsal claw; see comments below on female Acoma) and length and simple, cleft, bifid, toothed, or pectinate. Abdomen often concave medially in males, flat or slightly convex in females; sternites imbricate or smooth, narrowed medially or not, with sutures visible or completely effaced medially; sternite and propygidium fused or not with or without trace of suture; sixth sternite partially or completely retracted beneath fifth; pygidium exposed; 7 or fewer pairs of abdominal spiracles scarcely or not divergent posteriorly, with 1���3 posterior abdominal spiracles located in sternites or tergites. Male genitalia without sclerotized median lobe. Evans & Smith (2009) considered the Nearctic genera Acoma, Chaunocolus, and Chnaunanthus as incertae sedis, while following the traditional placement of both Phobetus and Warwickia in tribe Tanyproctini. The tribal placement of all these genera is reconsidered below. During the course of our investigations, all worldwide Melolonthinae tribes (see Table 1 for a complete list) were studied and we found no compelling morphological characters indicating that Acoma, Chaunocolus, Chnaunanthus, Phobetus, or Warwickia belong to any of them. Excluded taxa. Oncerinae new status and Podolasiinae new status. Evans (2002, 2003), Smith (2006), and Bouchard et al. (2011) all considered Oncerini and Podolasiini as tribes within the Melolonthinae. With regards to Oncerini, we now follow Saylor (1938), who considered both Oncerus LeConte, 1856 and Nefoncerus Saylor, 1938 in the subfamily Oncerinae on the basis of the placement of all abdominal spiracles within the pleural membrane. Ritcher (1969) confirmed Saylor���s findings for the placement of abdominal spiracles in Oncerus, but apparently did not examine the more rarely collected Nefoncerus. In addition to the placement of their abdominal spiracles, both of these monotypic genera are characterized by having hidden mandibles, labrum fused and coplanar with the clypeus, small and moderately setose lamellate clubs, and abdominal sutures that are distinct or not. Howden (1997) erected the tribe Podolasiini for Podolasia Harold, 1869 and his new genus, Podostena Howden, 1997, and stated that its position within the Melolonthinae is ���problematical��� and ���remains debatable since some characters are not typical for the subfamily���. Ritcher (1969) found Podolasia to have seven pairs of functional spiracles and a vestigial eighth pair, all located in the pleural membrane. Lacroix (2007) considered the Podolasiinae (Podolasiini), along with the Palaearctic Chasmatopterinae (Chasmatopterini) to be in the Chasmatopteridae without discussion or justification. As mentioned previously, this presumed relationship is untenable given that Chasmatopterus Dejean, 1821 and Podolasia have very different abdominal spiracle configurations. The lack of any posterior abdominal spiracles in the tergites or sternites, and the small, nearly globose, and densely setose lamellate antennal clubs, combined with the cephalic armature and genital capsule characters found in recently described Podolasia (see Smith & Paulsen 2017), and an undescribed Podostena species (Evans, personal observation) from Mexico precludes the inclusion of Podolasiini within the Melolothinae, thus requiring its elevation to the subfamily rank of Podolasiinae. The phylogenetic position of this new subfamily with relation to the Melolonthinae needs further study and it will likely occupy a branch basal to the phytophagous scarab clade (see Smith et al. 2006)., Published as part of Evans, Arthur V. & Smith, Andrew B. T., 2020, On the tribal classification of the Nearctic Melolonthinae (Coleoptera: Scarabaeidae), with descriptions of new species of Acoma Casey, 1889, pp. 51-77 in Zootaxa 4748 (1) on pages 52-54, DOI: 10.11646/zootaxa.4748.1.3, http://zenodo.org/record/3697578, {"references":["Evans, A. V. (2002) Melolonthinae. In: Arnett, R. H., Thomas, M. C., Skelley, P. E. & Frank, J. H. (Eds.), American Beetles. Vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, Florida, pp. 51 - 60.","Bouchard, P., Bousquet, Y., Davies, A. E., Alonso-Zarazaga, M. A., Lawrence, J. F., Lyal, C. H. C., Newton, A. F., Reid, C. A. M., Schmitt, M., Slipinski, S. A. & Smith, A. B. T. (2011) Family-group names in Coleoptera (Insecta). ZooKeys, 88, 1 - 972. https: // doi. org / 10.3897 / zookeys. 88.807","Lobl, I. & Lobl, D. (Eds.) (2016) Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea-Scirtoidea-Dascilloidea-Buprestoidea-Byrrhoidea. Revised and Updated Edition. Brill, Leiden, 983 pp.","Smith, A. B. T. & Evans, A. V. (2018) Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America. Zootaxa, 4471 (1), 279 - 308. https: // doi. org / 10.11646 / zootaxa. 4471.2.3","Lacordaire, J. T. (1855) Histoire naturelle des insectes. Genera des Coleopteres, or expose methodique et criticque de tous genres proposes jusquici dans cet odre d'insectes. Tome Troiseme. Contenant les familles des Pecticornes et Lamellicornes. Librarie Encyclopedique de Roret, Paris, 594 pp. https: // doi. org / 10.5962 / bhl. title. 67686","Evans, A. V. & Smith, A. B. T. (2009) Checklist of the New World Melolonthinae, Version 3. Available from: www. museum. unl. edu / research / entomology / SSSA / nwmelos. htm (accessed 14 July 2019)","Evans, A. V. (2003) A checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211 (1), 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Saylor, L. W. (1938) Revision of the subfamily Oncerinae with description of a new genus (Coleoptera: Scarabaeidae). Proceedings of the Entomological Society of Washington, 40, 99 - 103.","LeConte, J. L. (1856) Synopsis of the Melolonthidae of the United States. Journal of the Academy of Natural Sciences of Philadelphia, Series 2, 3, 225 - 288.","Ritcher, P. O. (1969) Spiracles of adult Scarabaeoidea (Coleoptera) and their phylogenetic significance. I. The abdominal spiracles. Annals of the Entomological Society of America, 62, 869 - 880. https: // doi. org / 10.1093 / aesa / 62.4.869","Howden, H. F. (1997) Podolasiini Howden, new tribe, and a revision of the included genera, Podolasia Harold and Podostena Howden, new genus (Coleoptera: Scarabaeidae: Melolonthinae). The Coleopterists Bulletin, 51, 223 - 255.","von Harold, E. (1869) Scarabaeidae. In: Gemminger, M. & Harold, E. (Eds.), Catalogus Coleopterorum. bucusque descriptorum, synonymicus et systematics, autotribus. Tome IV. Scarabaeidae. E. H. Gummi, Munich, pp. 976 - 1346.","Lacroix, M. (2007) Pachydeminae du monde (Coleoptera, Melolonthidae). Genera et catalogue. Collection Hannetons, Paris, 450 pp.","Smith, A. B. T. & Paulsen, M. J. (2017) Two new species of Podolasia Harold (Coleoptera: Scarabaeidae: Melolonthinae: Podolasiini) from Coahuila, Mexico. The Coleopterists Bulletin, 71, 532 - 536. https: // doi. org / 10.1649 / 0010 - 065 X- 71.3.532","Smith, A. B. T., Hawks, D. C. & Heraty, J. M. (2006) An overview of the classification and evolution of the major scarab beetle clades (Coleoptera: Scarabaeoidea) based on preliminary molecular analysis. Coleopterists Society Monograph, 5, 35 - 46. https: // doi. org / 10.1649 / 0010 - 065 X (2006) 60 [35: AOOTCA] 2.0. CO; 2"]}

Acoma chihuahuaensis Evans & Smith, new species Figs. 5���10 urn:lsid:zoobank.org:act: BD4FB72A-5BD1-4671-886A-C0E05D1BE391 Type material. Holotype male (CASC Type 19747), labeled ��� MEXICO: Chihuahua / 7.1 mi. W Concho / VII-23- 1982 / Fred G. Andrews / coll. blacklight // Dr. Alan R. Hardy / Donation to the / California Academy / of Sciences // CASENT / 8424526 // HOLOTYPE / Acoma / chihuahuaensis / Evans & Smith, 2019���. Paratypes, (9 males): same data as holotype, except with CASC database numbers CASENT 8424527��� 8424535. Paratypes are deposited in AVEC (1 paratype; CASENT 8424534), CASC (6 paratypes; CASENT 8424527��� 8424532), CMNC (1 paratype; CASENT 8424535), UNAM (1 paratype; CASENT 8424533). Diagnosis. Brown dorsally; anterior clypeal margin emarginate medially; antennae composed of 10 antennomeres, antennomeres 8���10 lamellate; protibia distinctly tridentate; elytra with disc glabrous and scattered setae along margins. Description. Holotype male (Figs. 5���10), length: 6.9 mm, width: 2.6 mm. BODY elongate, moderately convex, and uniformly brown dorsally, underside and appendages slightly lighter. HEAD (Fig. 8) rugosely punctate; supraorbital carinae becoming obsolete at base of clypeus. Clypeus subtrapezoidal with margins strongly reflexed, lateral margins arcuate basally, becoming straight before broadly concave anterior margin flanked by prominent obtuse angles. Ocular canthus slightly less than one-third of eye length in lateral view, with scattered pale setae each not quite as long as canthus. PRONOTUM slightly wider than long, broadest at middle, and less than half length of elytron; disc convex, glabrous, shiny, sparsely punctate with punctures separated by four or more of their diameters, with a narrow longitudinal impunctate line medially; margins with distinct bead, anterior margin bisinuate between obtuse anterior angles, with marginal bead relatively broad and raised; anterolateral margins straight, converging anteriorly, posterolateral margins slightly converging posteriorly with posterolateral angles broadly rounded. SCUTELLUM small, parabolic, about one-fifth basal width of elytron, finely and setigerously punctate basally, and glabrous and becoming impunctate apically. ELYTRON convex, with disc shiny, glabrous, and distinctly punctostriate with deep punctures separated by 1.5���3.0 X their diameters, punctures becoming confused at apical declivity; margins setigerous, setae on lateral margins becoming shorter apically. LEGS with tridentate protibia moderately emarginate between apical and middle teeth, and shallowly emarginate between middle and basal teeth. PYGIDIUM parabolic, shiny, finely shagreened, with scattered setigerous punctures, setae long. GENITALIA with parameres symmetrical, slender and slightly sinuate when viewed laterally, with membranous flange extending to apical fifth, and about as long as phallobasis (Figs. 9���10). Variation. Length: 5.9���6.9 mm, width: 2.2���2.6 mm. Color varies from light brown to brown, occasionally with elytra lighter, while the head and pronotum are slightly darker. Clypeus varies from subtrapezoidal to nearly parabolic with narrowly to broadly concave anterior margin flanked by prominent angles or not. Etymology. This species is named after the Mexican state of Chihuahua, where the type series was collected. Remarks. Acoma glabrata Cazier, 1953, A. nonglabrata new species, and A. chihuahuaensis new species all have antennae with 10 antennomeres, of which 8���10 form the lamellate club. Acoma chihuahuaensis new species is readily distinguished among these species by its darker color, distinct elytral punctostriae with deep punctures, more rounded and less pronounced protibial teeth, and its occurrence in mainland Mexico., Published as part of Evans, Arthur V. & Smith, Andrew B. T., 2020, On the tribal classification of the Nearctic Melolonthinae (Coleoptera: Scarabaeidae), with descriptions of new species of Acoma Casey, 1889, pp. 51-77 in Zootaxa 4748 (1) on pages 56-58, DOI: 10.11646/zootaxa.4748.1.3, http://zenodo.org/record/3697578, {"references":["Cazier, M. A. (1953) Review of the scarab genus Acoma. American Museum Novitates, 1624, 1 - 13."]}

Acoma pararobusta Evans & Smith, new species Figs. 23���28 urn:lsid:zoobank.org:act: 9342C833-0F57-4CCB-9819-C38902BAE444 Type material. Holotype male (CASTYPE 19749), labeled ��� MEX.: Baja Calif. Sur / El Sargento, 29.VII. / 1971. At U.V. light / H.G. Real & R.E. Main // n. sp. ? / DET. / A.R. Hardy 1979 // Acoma / n. sp. 3 / det. W.B.Warner 2015 // CASENT / 8424521 // HOLOTYPE / Acoma / pararobusta / Evans & Smith, 2019���. Paratypes, (2 males): same data as holotype, except no determination labels, CASC database numbers CASENT 8424522���8424523. Paratypes are deposited in CASC (1 paratype; CASENT 8424523), CMNC (1 paratype; CASENT 8424522). Diagnosis. Rufopiceous or rufobrunneus dorsally; anterior clypeal margin shallowly emarginate or rounded medially; frontoclypeal suture tumid; antennae composed of 9 antennomeres, antennomeres 5���9 lamellate, subequal in length; protibia distinctly tridentate; elytral disc with scattered, pale setae. Description. Holotype male (Figs. 23���28), length: 9.0 mm, width: 4.0 mm. BODY dark brown, elongate, robust, moderately convex. HEAD (Fig. 26) with narrowly raised transverse impunctate area, frons and base of clypeus deeply and rugosely punctate, reflexed clypeal margins smoother, with deep punctures mostly separated by their own diameters; short and irregular supraorbital carina obsolete at bases of head and clypeus. Clypeus somewhat semicircular, lateral margins narrowly reflexed, apical margin more broadly reflexed and slightly concave. Ocular canthus thick and broadly rounded apically, about one-third length of eye when viewed laterally, densely setose with setae somewhat longer than canthus. PRONOTUM about half length of elytron, as long as wide; disc glabrous with moderately dense punctures usually separated by twice their diameters sometimes briefly confluent on either side of narrow longitudinal impunctate area medially, punctures become denser laterally, even more so anteriorly; anterolateral margins sinuate, abruptly converging anteriorly from middle, posterolateral margins subparallel, shallowly concave medially; lateral and posterior margins with narrow bead, lateral margins distinctly setose with setae long; anterior margin mostly concave between acute and produced anterior angles, slightly protruding medially, with broad bead; posterior angles broadly obtuse. SCUTELLUM broadly elliptical, basal half sparsely and setigerously punctate, setae coarse. ELYTRON shiny and setose, with deep, narrow striae on disc with setigerous punctures bearing recumbent setae becoming short and irregular on declivities. LEGS with tridentate protibia deeply emarginate between apical and middle teeth, shallowly emarginate between middle and basal teeth. PYGIDIUM broadly parabolic, rugosely punctate along sides, and setose with setae long, erect. GENITALIA with parameres symmetrical, slender and slightly sinuate when viewed laterally, with membranous flange extending to apical fourth, and not as long as phallobasis (Figs. 27���28). Variation. Length: 7.5���9.0 mm, width: 3.2���4.0 mm. Color varies from light to dark brown. Anterior clypeal margin ranges from slightly concave to nearly rounded. Etymology. This is named in recognition of the similarities with A. robusta Van Dyke, 1928. Remarks. Acoma pararobusta new species is distinguished from all other species in the genus by the characters given in the diagnosis. In addition to the tumid frontoclypeal suture, it is distinguished from A. robusta by the coarse, rugose sculpturing on the frons and base of the clypeus, occasional confluent punctures on pronotal disc, and the somewhat semicircular clypeal margin with bases that are more or less continuous with the supraorbital carinae., Published as part of Evans, Arthur V. & Smith, Andrew B. T., 2020, On the tribal classification of the Nearctic Melolonthinae (Coleoptera: Scarabaeidae), with descriptions of new species of Acoma Casey, 1889, pp. 51-77 in Zootaxa 4748 (1) on pages 61-62, DOI: 10.11646/zootaxa.4748.1.3, http://zenodo.org/record/3697578, {"references":["Van Dyke, E. C. (1928) Notes and descriptions of new species of Scarabaeidae from western North America. The Pan-Pacific Entomologist, 4, 151 - 162."]}

Subfamily Melolonthinae Our diagnosis of the Melolonthinae below is based largely on Evans (2002). Diagnosis. Scarabaeidae. Adults 3–60 mm in length. Dorsal surface glabrous, setose or scaled; sometimes setae or scales forming distinct patches or lines; color testaceous to reddish brown or black, occasionally with metallic blue, green, coppery, or brassy luster. Head usually unarmed, never with frontal extensions or horns. Eyes only partially divided by ocular canthus. Antennae with 7–10 antennomeres, rarely more, and insertions not visible from above; lamellate club usually consisting of 3–7 antennomeres; lamellae glabrous or with only a few setae, often longer in males. Mandibles sclerotized, usually well developed, and concealed in dorsal view. Transverse, narrowed, or conical labrum free or fused beneath clypeus, sometimes extending beyond or fused to apical clypeal margin. Thorax with pronotal disc more-or-less convex, never concave or with raised carinae; scutellum exposed. Elytra with lateral margins more-or-less straight behind humerus, with mesepimeron usually covered by elytral bases. Metathoracic wings fully developed or reduced. Metepimeron rectangular or triangular. Legs with procoxae transverse or conical; protibiae usually with apical spurs; mesotibiae and metatibiae usually with a pair of apical spurs located below or set on either side of tarsal articulation, or with upper spur adjacent to tarsal articulation; opposing mesotarsal and metatarsal claws equal in thickness (except Hoplia, which have a single, enlarged metatarsal claw; see comments below on female Acoma) and length and simple, cleft, bifid, toothed, or pectinate. Abdomen often concave medially in males, flat or slightly convex in females; sternites imbricate or smooth, narrowed medially or not, with sutures visible or completely effaced medially; sternite and propygidium fused or not with or without trace of suture; sixth sternite partially or completely retracted beneath fifth; pygidium exposed; 7 or fewer pairs of abdominal spiracles scarcely or not divergent posteriorly, with 1–3 posterior abdominal spiracles located in sternites or tergites. Male genitalia without sclerotized median lobe. Evans & Smith (2009) considered the Nearctic genera Acoma, Chaunocolus, and Chnaunanthus as incertae sedis, while following the traditional placement of both Phobetus and Warwickia in tribe Tanyproctini. The tribal placement of all these genera is reconsidered below. During the course of our investigations, all worldwide Melolonthinae tribes (see Table 1 for a complete list) were studied and we found no compelling morphological characters indicating that Acoma, Chaunocolus, Chnaunanthus, Phobetus, or Warwickia belong to any of them. Excluded taxa. Oncerinae new status and Podolasiinae new status. Evans (2002, 2003), Smith (2006), and Bouchard et al. (2011) all considered Oncerini and Podolasiini as tribes within the Melolonthinae. With regards to Oncerini, we now follow Saylor (1938), who considered both Oncerus LeConte, 1856 and Nefoncerus Saylor, 1938 in the subfamily Oncerinae on the basis of the placement of all abdominal spiracles within the pleural membrane. Ritcher (1969) confirmed Saylor’s findings for the placement of abdominal spiracles in Oncerus, but apparently did not examine the more rarely collected Nefoncerus. In addition to the placement of their abdominal spiracles, both of these monotypic genera are characterized by having hidden mandibles, labrum fused and coplanar with the clypeus, small and moderately setose lamellate clubs, and abdominal sutures that are distinct or not. Howden (1997) erected the tribe Podolasiini for Podolasia Harold, 1869 and his new genus, Podostena Howden, 1997, and stated that its position within the Melolonthinae is “problematical” and “remains debatable since some characters are not typical for the subfamily”. Ritcher (1969) found Podolasia to have seven pairs of functional spiracles and a vestigial eighth pair, all located in the pleural membrane. Lacroix (2007) considered the Podolasiinae (Podolasiini), along with the Palaearctic Chasmatopterinae (Chasmatopterini) to be in the Chasmatopteridae without discussion or justification. As mentioned previously, this presumed relationship is untenable given that Chasmatopterus Dejean, 1821 and Podolasia have very different abdominal spiracle configurations. The lack of any posterior abdominal spiracles in the tergites or sternites, and the small, nearly globose, and densely setose lamellate antennal clubs, combined with the cephalic armature and genital capsule characters found in recently described Podolasia (see Smith & Paulsen 2017), and an undescribed Podostena species (Evans, personal observation) from Mexico precludes the inclusion of Podolasiini within the Melolothinae, thus requiring its elevation to the subfamily rank of Podolasiinae. The phylogenetic position of this new subfamily with relation to the Melolonthinae needs further study and it will likely occupy a branch basal to the phytophagous scarab clade (see Smith et al. 2006).

Acoma nonglabrata Evans & Smith, new species Figs. 17���22 urn:lsid:zoobank.org:act: 8EBF5164-E12F-4C2B-BE80-21ACF3429289 Type material. Holotype male (CASCTYPE 19748), labeled ��� MEXICO: BajaCal. Norte / Guadalupe Canyon, E. / side of SierraJuarez / 18-VI-1973 S.C. / Williams & K.B. Blair / SCW # 311 (2) // ACOMA / OCHLERA / HOWDEN / OR N.SP. NEAR / det. W.B.Warner 2015 // CASENT / 8424501 // HOLOTYPE / Acoma / nonglabrata / Evans & Smith, 2019���. Paratypes (7 males): 5, same data as holotype, except no determination label, CASC database numbers CASENT 8424502���8424506; 2, ��� MEX. BajaCalif.Norte / 5.3km S Bajia de Los / Angeles VII.9���10.78 / 59029.6 / E. L. Sleeper ��� (CMNC) and ���BCN- 5.3 km S / BAJIA DE LOS / ANGELES / VII-9/10-1978 // 59029.6 / E. L. Sleeper ��� (CMNC). Paratypes are deposited in AVEC (1 paratype; CASENT 8424502), CASC (2 paratypes; CASENT 8424505��� 8424506), CMNC (1 paratype; CASENT 8424503), UNAM (1 paratype; CASENT 8424504). Diagnosis. Light reddish tan dorsally with head and pronotum darker; anterior clypeal margin emarginate medially; antennae composed of 10 antennomeres, antennomeres 8���10 lamellate; protibia distinctly tridentate; elytral disc with scattered setae. Description. Holotype male (Figs. 17���22), length: 6.5 mm, width: 3.5 mm. BODY testaceous, elongate, moderately convex. HEAD (Fig. 20) with impunctate vertex shiny and slightly depressed, frons and clypeus rugose, clypeus with large shallow punctures evident and separated by their own diameters; supraorbital carina obsolete at base of head and clypeus. Clypeus subtrapezoidal, lateral margins moderately reflexed, anterior margin broadly and shallowly concave with angles not prominent. Ocular canthus about one-quarter length of eye when viewed laterally, setose with setae longer than canthus. PRONOTUM slightly wider than long, broadest medially, convex, about one-third length of elytron; disc convex, glabrous, moderately punctate, most punctures separated by 1���2 diameters, with longitudinal impunctate area medially; lateral and posterior margins with narrow bead; anterior margin moderately bisinuate between obtuse anterior angles, with broad bead; anterolateral margin slightly sinuate and converging anteriorly, posterolateral margin straight and converging slightly posteriorly, with posterior angles broadly angulate. SCUTELLUM parabolic, about one-quarter width of elytron, disc mostly finely shagreened. ELYTRON convex, shiny, finely shagreened, disc punctostriate with punctures becoming confused on apical and lateral declivities, setigerous punctures with long suberect setae. LEGS with tridentate protibia moderately emarginate between teeth. PYGIDIUM parabolic, shiny, finely shagreened, with scattered setigerous punctures, setae long. GENITALIA with parameres symmetrical, slender and slightly sinuate when viewed laterally, with membranous flange extending to apical fourth, and not as long as phallobasis (Figs. 21���22). Variation. The type series is consistent in length, color, and surface sculpturing, but does exhibit minor variation in the degree of concavity of the anterior clypeal margin and prominence of clypeal angles. Etymology. This specific epithet is in reference to the superficial resemblance to A. glabrata, which also occurs in Baja California. Remarks. With 10 antennomeres and the club consisting of three antennomeres, A. nonglabrata new species superficially resembles A. glabrata, but is readily distinguished by the setose elytral disc and more angulate lateral pronotal margins., Published as part of Evans, Arthur V. & Smith, Andrew B. T., 2020, On the tribal classification of the Nearctic Melolonthinae (Coleoptera: Scarabaeidae), with descriptions of new species of Acoma Casey, 1889, pp. 51-77 in Zootaxa 4748 (1) on pages 60-61, DOI: 10.11646/zootaxa.4748.1.3, http://zenodo.org/record/3697578

Athlia rustica Erichson, 1835 Figs. 48���53, 72. Original combination. Athlia rustica Erichson, 1835: 267 Type locality. ��� Chile ��� (Erichson 1835). Type series. Lectotype (Figs. 48���53) at ZMHB labeled a) ���25008��� (typeset), b) ���Type��� (typeset, orange label), c) ��� rustica / Er. / Chile Meyen.��� (handwritten, green label), d) ���SYNTYPUS / Athlia / rustica Erichson, 1835 / labelled by MNHUB 2009��� (typeset, orange label), e) ��� ATHLIA / RUSTICA / ERICHSON, 1835 / LECTOTYPE / A.B.T. SMITH��� (handwritten and typeset, red label). Lectotype here designated. Four paralectotypes at ZMHB labeled a) ���Type��� (typeset, orange label), b) ���Hist.-Coll. (Coleoptera) / Nr. 25008 / Athlia / rustica Erichs. / Chili, Meyen / Zool. Mus. Berlin��� (typeset), c) ���SYNTYPUS / Athlia / rustica Erichson, 1835 / labelled by MNHUB 2009��� (typeset, orange label), d) ��� ATHLIA / RUSTICA / ERICHSON, 1835 / PARALECTOTYPE / A.B.T. SMITH��� (handwritten and typeset, yellow label). A lectotype is designated to preserve the stability of the nomenclature by selecting a single specimen as the name-bearing type for this species. Since Erichson (1835) did not explicitly state how many specimens he examined, it is impossible to know the exact composition of the type series. Distribution. Fig. 72. Chile: IV Regi��n de Coquimbo, V Regi��n de Valpara��so, Regi��n Metropolitana de Santiago, VI Regi��n del Libertador General Bernardo O���Higgins, VII Regi��n del Maule, VIII Regi��n del Biob��o, IX Regi��n de la Araucan��a (based on specimens from AMNH, AVEC, BMNH, CASC, CMNC, CNCI, FMNH, FSCA, INHS, LEMQ, MNHN, MNNC, UMCE, UMSP, USNM). One 19th century specimen is labeled ���Valdivia��� (BMNH) but we consider this record erroneous. The record from IX Regi��n de la Araucan��a is based on a single collecting event from ��� 30 km NE Villarrica���, which might need further verification., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 294-295, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Erichson, W. F. (1835) Neue Sudamerikanische Kafergattungen aus der familie der Blatterhorner. Archiv fur Naturgeschichte, 1, 256 - 270."]}

Ulata Saylor, 1945 Ulata Saylor, 1945: 441. Type species Ulata argentina Saylor, 1945 by original designation. Redescription. Length 4.5���8.0 mm. Elongate, widest just behind middle of elytra; dorsal surface, ventral surface of thorax and posterior ventrite reddish brown, remainder of abdomen dark brown; surfaces shiny with scattered short, white, and appressed setae. Head: labrum wide, broadly emarginate, evenly and shallowly reflexed anteriorly, completely fused to and slightly below plane of clypeus exposing anterior clypeal margin; labroclypeal suture distinct and terminating in small, marginal notches. Mentum longer than wide, not emarginate in front, with sides somewhat straight, gradually converging posteriorly. Antennae with 8 antennomeres, club with 3 short, thick lamellae. Pronotum: somewhat convex, slightly wider than long, widest medially. Elytra: somewhat flat with four rows (including sutural interval) of slightly raised, irregular, dark, and glabrous patches. Metathoracic wings fully developed. Abdomen: without lateral carinae on ventrites. Legs: protibia with three apical teeth on outer margin, basal tooth weak, spur present and barely reaching base of first tarsomere. Claws symmetrical and incised with a short, bisetose onychium in between. Metatibial apices with spurs set in notch, upper spur set below tarsal articulation and reaching about half the length of first tarsomere. Male genitalia: symmetrical parameres simple in form, not split or with strong sinuations, and subequal in length to phallobase. Composition. The only species, U. argentina Saylor, is known from Argentina and Chile., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 296-297, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Saylor, L. W. (1945) Studies in the melolonthine scarab beetle genera of the American continents. No. IV. A new genus from Argentina. Revista de Entomologia, 16, 441 - 444."]}

Athlia freyi Martínez, 1974 Figs. 24–27, 67. Original combination. Athlia freyi Martínez, 1974: 347. Type locality. “ Brasil, estado de São Paulo, Serra da Bocaina, 1500 m ” (Martínez 1974). Type series. Holotype at Georg Frey Collection, Naturhistorisches Museum Basel, Switzerland (not examined). One paratype at CMNC was examined. Distribution. Fig. 67. Brazil: São Paulo (based on specimens from CMNC)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 287, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Martinez, A. (1974) Nuevas consideraciones sobre el Genero Athlia Erichson, 1835 (Col., Scarabaeidae, Sericinae). Entomologische Arbeiten aus dem Museum G. Frey, 25, 343 - 353."]}

Apteroathlia translucida Smith & Evans, new species Figs. 6–10, 66. http://zoobank.org/urn:lsid:zoobank.org:act:D4F8FA09-1E22-41A0-91F1-CD2945005552 Type locality. Cerro, Montevideo, Uruguay. Type series. One male holotype, 14 male paratypes, and 14 female paratypes. Holotype male (Figs. 6–10) at CMNC labeled a) “Montevideo / Cerro / 1. X. 33 ” (handwritten), b) “H. & A. HOWDEN / COLLECTION / ex. A.Martínez coll.” (typeset with black border), c) “ Apterops / translucidus / (Cand. in litt.)” (handwritten). One paratype male at USNM labeled a) “ Bs As / Azul / V.956. Daguerre ” (handwritten), b) “ ARGENTINA / 1968 Colln. / J.Daguerre ” (typeset over red, white, and green background), c) “ Apterops / translucidus / (in lit.) Cand. / Vialle det” (handwritten). One paratype male at CMNC and one parartype female at USNM labeled a) “ Bs As / Azul / V.956. Daguerre ” (handwritten), b) “ ARGENTINA / 1968 Colln. / J.Daguerre ” (typeset over red, white, and green background). Four male paratypes at CMNC (2) and USMN (2) and six female paratypes at CMNC (3) and USNM (3) labeled a) “ ARGENTINA / Buenos Aires Prov. / Azul; V.1956 / J. Daguerre” (typeset). One male paratype at USNM labeled a) “ Montevideo / Cerro / 12 IX 34 ” (handwritten), b) “Apterops / translucidus / (Chand. i. litt)” (handwritten). One paratype male at CMNC labeled a) “ Montevideo / Cerro / 1. X. 33 ” (handwritten), b) “H. & A. HOWDEN / COLLECTION / ex. A.Martínez coll.” (typeset with black border). Four male and two female paratypes at CMNC labeled a) “R. O. del / Uruguay ” (typeset on faded green paper), b) “H. & A. HOWDEN / COLLECTION / ex. A.Martínez coll.” (typeset with black border). One male paratype at BMNH labeled a) “18234” (handwritten), b) “ La Plata / M Video ” (handwritten), c) “ Fry Coll. / 1905-100.” (typeset). One male paratype at BMNH labeled a) “Monte- / video” (typeset), b) “ 1909–74. ” (typeset), c) “21.” (handwritten). One female paratype at BMNH labeled a) “ Brazil ” (handwritten), b) “2032.” (handwritten), c) “67-45” (handwritten), d) “inflate Reiche / Brasilia” (handwritten), e) “So named / in Reiches / Collection. / C.W.” (typeset). One female paratype at MNHN labeled a) “Plata / Dolle” (handwritten), b) “ MUSÉUM PARIS / 1906 / Coll. Léon FAIRMAIRE” (handwritten). One female paratype at USNM labeled a) “ Montevideo / Cerro / 24. IX. 33 ” (handwritten), b) “ Apterops / translucidus / (Chand. i. litt.)” (handwritten), c) “OLCartwright / Collection / 1959” (typeset). Two female paratypes at USNM labeled a) “ Montevideo / Cerro / 12 IX 34 ” (handwritten). All paratypes listed above also bear a yellow paratype label and a database label with the database number and a barcode. Description of holotype (Figs. 6–10). Male. Length 6.6 mm, width 3.1 mm. Dorsal and ventral color light brown, elytra translucent. Dorsal surface glabrous. Head: surface moderately punctate. Labrum with apex reflexed backward and convexly parabolic medially; underside concave medially. Clypeus with apex straight, abutted to labrum. Frontoclypeal suture visible. Eyes round, slightly protruding laterally. Antennae with 8 antennomeres; antennal club with 3 antennomeres, club distinctly shorter in length to antennomeres 1–5 combined, club surface thickly setose. Maxillae robust, triangular, basal width slightly less than width of mentum; maxillary palpus with 4 palpomeres, length approximately ¾ that of antennae. Mentum prominent, slightly longer than wide, apex straight, surface slightly convex; labial palpus reduced. Pronotum: surface moderately to densely punctate; apical angle acute, basal angle squared. Pronotum widest medially with smooth margins. Elytra: surface moderately to densely punctate, striae punctate but obscured by other surface punctures. Wings: apterous. Venter: metasternum with dense punctures. Pygidium flat; surface moderately to sparsely punctate with microsculpturing. Legs: protibia with 3 apical teeth on outer margin of apical half; 3rd tooth weak. Claws symmetrical with medial tooth. Protibial spur present, metatibial spurs approximately equal in length. Genitalia: parameres long, slender, tapering into needlelike apices (Fig. 10). Variation. Length 5.9–7.7 mm. Dorsal color tan to brown. Females with slightly more inflated, convex abdominal segments. Etymology. This name is in reference to the translucent elytra and an homage to “ Apterops translucidus ”, which was used on older determination labels for some specimens but was never published. It should be treated as an adjective in the nominative singular. Distribution (Fig. 66). URUGUAY (14): Montevideo (8): Cerro, Montevideo; no data (6). ARGENTINA (14): Buenos Aires (14): Azul, La Plata. NO DATA (1): “ Brazil ”. Temporal data. May (13), September (4), October (2). Remarks. Apteroathlia translucida new species is mainly found around the gulf shores of Río de la Plata in Uruguay and Buenos Aires Province, Argentina. One 19th Century specimen is labeled “ Brazil ”, but we consider this record doubtful and in need of further verification. The specimens collected in May were probably collected fresh while those obtained in September and October appear to have been found dead and dried. The conservation status of this species should be investigated since most of the specimens were collected from coastal habitats in highly populated urban areas.

Athlia rotundata Guti��rrez, 1949 Figs. 44���47, 71. Original combination. Athlia rotundata Guti��rrez, 1949: 12. Type locality. ��� Chile: Prov. Valpara��so, Conc��n ��� (Guti��rrez 1949). Type series. Holotype, allotype, and nine paratypes at Museo de Zoolog��a, Universidad de Concepci��n, Concepci��n, Chile (Cekalovic & Artigas 1969). Two paratypes were examined from CMNC and FMNH. Distribution. Fig. 71. Chile: V Regi��n de Valpara��so (based on specimens from CASC, CMNC, FMNH, UMCE)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 293, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Gutierrez, R. (1949) Notas sobre Scarabaeidae neotropicos (Coleoptera Lamellicornia). Anales de la Sociedad Cientifica Argentina, 148, 9 - 35.","Cekalovic, T. & Artigas, J. N. (1969) Catalogo de los tipos de Insecta depositados en la coleccion del Departamento de Zoologia de la Universidad de Concepcion (INCO) (Diciembre, 1968). Boletin de la Sociedad de Biologia de Concepcion, 41, 111 - 133."]}

Key to species of Athlia (based on Saylor 1946b; Gutiérrez 1949; Martínez 1967, 1974) 1. Metathoracic wings reduced and non-functional (slightly lift an elytron to check). Central Chile (VIII Región del Biobío to IX Región de la Araucanía) (Fig. 69)..................................................... Athlia plebeja Burmeister - Metathoracic wings full sized and functional................................................................ 2 2. Pronotum with basal angles evenly rounded, not protruding or angulate (Figs. 30, 34, 46)............................ 3 - Pronotum with basal angles protruding or angulate (Figs. 13, 17, 22, 26, 42, 52).................................... 5 3. Length less than 10 mm. Metatarsomere 1 noticeably shorter than metatarsomere 2 (Fig. 32). Southern Argentina (Neuquén to Chubut, Fig. 69)................................................................... Athlia parvissima Saylor - Length greater than 10 mm. Metatarsomere 1 equal to or slightly shorter than metatarsomere 2 (Fig. 28)................. 4 4. Labrum weakly curved, weakly elevated medially; lateral border of clypeus slightly reflexed; dorsal surface tan to chestnut brown. Central Chile (V Región de Valparaíso, Fig. 79)................................... Athlia rotundata Gutiérrez - Labrum evenly curved and elevated medially; lateral border of clypeus strongly reflexed; dorsal surface light brown to light tan. Southern Argentina (Neuquén to Chubut) and southern Chile (Región de Aisén, Fig. 68).......... Athlia giaii Martínez 5. Labrum not reflexed, on the same plane as clypeal surface. Southeastern Brazil (Santa Catarina, Fig. 67).................................................................................................... Athlia brasilica Saylor - Labrum reflexed above the plane of clypeal surface........................................................... 6 6. Pronotum surface with dense, even punctures (Figs. 22, 42, 52)................................................. 7 - Pronotum surface with sparse, sometimes unevenly distributed punctures (Figs. 13, 26).............................. 9 7. Elytra evenly covered with setae; pronotum with long setae along lateral edges. Central Chile (IV Región de Coquimbo to IX Región de la Araucanía, Fig. 72)........................................................ Athlia rustica Erichson - Elytra glabrous medially and sparsely setose laterally and apically; pronotum with long setae along lateral edge but mainly in apical half........................................................................................... 8 8. Elytra length less than twice the length of head and pronotum combined Northern and central Argentina (Jujuy to Río Negro, Fig. 67).............................................................................. Athlia bruchi Moser - Elytra length greater than twice the length of head and pronotum combined. Central Chile (IV Región de Coquimbo to Región Metropolitana de Santiago, Fig. 70)................................................ Athlia problematica Gutiérrez 9. Length greater than 12 mm; apical and basal pronotal angles well developed (Fig. 26). Southern Brazil (São Paulo, Fig. 67)...................................................................................... Athlia freyi Martínez - Length less than 12 mm; apical and basal pronotal angles weakly developed (Fig. 13). Northeastern Argentina (Buenos Aires, Fig. 67)............................................................................ Athlia bollei Martínez

Dihymenonyx herrerai Guti��rrez, 1949 Original combination. Dihymenonyx herrerai Guti��rrez, 1949: 16. Subsequent combination. Athlia (Dihymenonyx) herrerai (Guti��rrez): Mart��nez 1974: 345. Type locality. ���Chile: Territorio de Magallanes, Punta Arenas��� (Guti��rrez 1949). As mentioned in Mondaca (2007), this locality is erroneous. Type series. Holotype at Museo de Zoolog��a, Universidad de Concepci��n, Concepci��n, Chile (Cekalovic & Artigas 1969) (not examined). Distribution. Fig. 73. Chile: VIII Regi��n del Biob��o (Mondaca 2007)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 296, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Gutierrez, R. (1949) Notas sobre Scarabaeidae neotropicos (Coleoptera Lamellicornia). Anales de la Sociedad Cientifica Argentina, 148, 9 - 35.","Martinez, A. (1974) Nuevas consideraciones sobre el Genero Athlia Erichson, 1835 (Col., Scarabaeidae, Sericinae). Entomologische Arbeiten aus dem Museum G. Frey, 25, 343 - 353.","Mondaca, J. (2007) Revision del genero Dihymenonyx Gutierrez (Scarabaeidae: Melolonthinae: Sericini) con descripcion de dos nuevas especies de Chile. Revista Chilena de Entomologia, 32, 13 - 29. [dated 2006]","Cekalovic, T. & Artigas, J. N. (1969) Catalogo de los tipos de Insecta depositados en la coleccion del Departamento de Zoologia de la Universidad de Concepcion (INCO) (Diciembre, 1968). Boletin de la Sociedad de Biologia de Concepcion, 41, 111 - 133."]}

Athlia bollei Martínez, 1955 Figs. 11–14, 67. Original combination. Athlia bollei Martínez, 1955: 418. Type locality. “ Argentina, provincial de Buenos Aires, Partido de Tandill (Serra-nía)” (Martínez 1955). Type series. Holotype at Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires, Argentina (not examined). Distribution. Fig. 67. Argentina: Buenos Aires (based on specimens from CMNC)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 286, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Martinez, A. (1955) Una nueva especie de Athlia de la Argentina (Col. Scarab. Melolonth. Sericinae). Revista Ecuatoriana de Entomologia y Parasitologia, Guayaquil, 2, 417 - 423."]}

Athlia parvissima Saylor, 1946 Figs. 32���35, 69. Original combination. Athlia parvissima Saylor, 1946: 24. Type locality. ��� Zapala, Neuquen, Argentina ��� (Saylor 1946). Type series. Holotype at CASC labeled a) ���Zapala, / Neuquen ��� (typeset), b) ��� Argentina / Dec 9 1920 ��� (typeset), c) ��� HOLOTYPE / Athlia / PARVISSIMA / L.W.Saylor��� (typeset and handwritten on red paper), d) ���PARVISSIMA / Saylor / DET. / L.W. SAYLOR��� (handwritten and typeset), e) ���L.W. Saylor / Collection��� (typeset), f) ��� California Academy / of Sciences / Type / No. 7923��� (typeset and handwritten), g) ���Southern Neotropical Scarabs / database # AS 2617613 / Athlia parvissima / Saylor, 1946 ♀ / DET: A.B.T. SMITH 2008 ��� (typeset). Distribution. Fig. 69. Argentina: Neuqu��n, R��o Negro, Chubut (based on specimens from CASC, CDFA, CMNC, LEMQ)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 288-289, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Saylor, L. W. (1946) Studies in the melolonthine scarab beetle genera of the American continents. I. Revision of the genus Athlia. Proceedings of the Entomological Society of Washington, 48, 18 - 25.","Smith, A. B. T. (2008) South American Melolonthinae (Coleoptera: Scarabaeidae) classification and nomenclature: some problems and solutions. Insecta Mundi, 60, 1 - 28."]}

Athlia giaii Mart��nez, 1959 Figs. 28���31, 68. Original combination. Athlia giaii Mart��nez, 1959: 23. Type locality. ��� Neuqu��n, Piedra del Aguila ��� (Mart��nez 1959). Type series. Holotype at Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires, Argentina (not examined). Twenty paratypes at CMNC were examined. Distribution. Fig. 68. Argentina: Neuqu��n, R��o Negro, Chubut (based on specimens from CDFA, CMNC, LEMQ). Mondaca (2011) recorded specimens from Chile: XI Regi��n de Ais��n., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 288, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Martinez, A. (1959) Scarabaeoidea neotropica. VI. Algunos Melolonthidae nuevos o poco conocidos (Col.). Neotropica, 5 (16), 23 - 29.","Mondaca, J. (2011) Presencia de Athlia giaii y Myloxena patagonica (Coleoptera: Scarabaeidae: Melolonthinae) en la Patagonia occidental, Chile. Revista Chilena de Entomologia, 36, 43 - 49."]}

Dihymenonyx Guti��rrez, 1949 Dihymenonyx Guti��rrez, 1949: 16. Type species Dihymenonyx herrerai Guti��rrez, 1949 by original designation. Redescription. Length 9.0���13.0 mm. Elongate-oval, widest about middle of elytra; reddish brown, brown, or dark brown; dorsal and ventral surfaces shiny, glabrous. Head: labrum wide, broadly emarginate and evenly reflexed anteriorly, and completely fused to clypeus with labroclypeal suture more-or-less distinct and terminating in small, marginal notches. Mentum longer than wide, narrowly emarginate in front, with sides broadly arcuate, gradually converging posteriorly. Antennae with 9 antennomeres, club with 3 short, thick lamellae. Pronotum: convex, wider than long, widest medially or slightly behind middle. Elytra: convex and coarsely punctostriate. Metathoracic wings fully developed or reduced in size. Abdomen: without lateral carinae on ventrites. Legs: protibia with three apical teeth on outer margin, basal tooth weak, spur present and shorter than first tarsomere. Claws symmetrical and bifid, each pair flanked on both sides at base by a membranous flap, and with a short, bisetose onychium in between. Metatibial apices with spurs set in notch, longest spur reaching about half the length of first tarsomere. Male genitalia: symmetrical parameres simple in form, not split or with strong sinuations, and somewhat subequal in length of phallobase. Composition. All three species of Dihymenonyx are endemic to Chile (Fig. 73). Remarks. Dihymenonyx was revised by Mondaca (2007)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 295-296, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Gutierrez, R. (1949) Notas sobre Scarabaeidae neotropicos (Coleoptera Lamellicornia). Anales de la Sociedad Cientifica Argentina, 148, 9 - 35.","Mondaca, J. (2007) Revision del genero Dihymenonyx Gutierrez (Scarabaeidae: Melolonthinae: Sericini) con descripcion de dos nuevas especies de Chile. Revista Chilena de Entomologia, 32, 13 - 29. [dated 2006]"]}

Dihymenonyx micropterus Mondaca, 2007 Original combination. Dihymenonyx micropterus Mondaca, 2007: 18. Type locality. ��� Chile, V Regi��n de Valpara��so, camino a Quintay (33��10'32.7''S y 71��32'36.3''W)��� (Mondaca 2007). Type series. Holotype at MNNC (not examined). Six paratypes at CMNC were examined. Distribution. Fig. 73. Chile: V Regi��n de Valpara��so (based on specimens from CASC, CMNC and records from Mondaca 2007)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 296, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Mondaca, J. (2007) Revision del genero Dihymenonyx Gutierrez (Scarabaeidae: Melolonthinae: Sericini) con descripcion de dos nuevas especies de Chile. Revista Chilena de Entomologia, 32, 13 - 29. [dated 2006]"]}

Apteroathlia translucida Smith & Evans, new species Figs. 6���10, 66. http://zoobank.org/urn:lsid:zoobank.org:act:D4F8FA09-1E22-41A0-91F1-CD2945005552 Type locality. Cerro, Montevideo, Uruguay. Type series. One male holotype, 14 male paratypes, and 14 female paratypes. Holotype male (Figs. 6���10) at CMNC labeled a) ���Montevideo / Cerro / 1. X. 33 ��� (handwritten), b) ���H. & A. HOWDEN / COLLECTION / ex. A.Mart��nez coll.��� (typeset with black border), c) ��� Apterops / translucidus / (Cand. in litt.)��� (handwritten). One paratype male at USNM labeled a) ��� Bs As / Azul / V.956. Daguerre ��� (handwritten), b) ��� ARGENTINA / 1968 Colln. / J.Daguerre ��� (typeset over red, white, and green background), c) ��� Apterops / translucidus / (in lit.) Cand. / Vialle det��� (handwritten). One paratype male at CMNC and one parartype female at USNM labeled a) ��� Bs As / Azul / V.956. Daguerre ��� (handwritten), b) ��� ARGENTINA / 1968 Colln. / J.Daguerre ��� (typeset over red, white, and green background). Four male paratypes at CMNC (2) and USMN (2) and six female paratypes at CMNC (3) and USNM (3) labeled a) ��� ARGENTINA / Buenos Aires Prov. / Azul; V.1956 / J. Daguerre��� (typeset). One male paratype at USNM labeled a) ��� Montevideo / Cerro / 12 IX 34 ��� (handwritten), b) ���Apterops / translucidus / (Chand. i. litt)��� (handwritten). One paratype male at CMNC labeled a) ��� Montevideo / Cerro / 1. X. 33 ��� (handwritten), b) ���H. & A. HOWDEN / COLLECTION / ex. A.Mart��nez coll.��� (typeset with black border). Four male and two female paratypes at CMNC labeled a) ���R. O. del / Uruguay ��� (typeset on faded green paper), b) ���H. & A. HOWDEN / COLLECTION / ex. A.Mart��nez coll.��� (typeset with black border). One male paratype at BMNH labeled a) ���18234��� (handwritten), b) ��� La Plata / M Video ��� (handwritten), c) ��� Fry Coll. / 1905-100.��� (typeset). One male paratype at BMNH labeled a) ���Monte- / video��� (typeset), b) ��� 1909���74. ��� (typeset), c) ���21.��� (handwritten). One female paratype at BMNH labeled a) ��� Brazil ��� (handwritten), b) ���2032.��� (handwritten), c) ���67-45��� (handwritten), d) ���inflate Reiche / Brasilia��� (handwritten), e) ���So named / in Reiches / Collection. / C.W.��� (typeset). One female paratype at MNHN labeled a) ���Plata / Dolle��� (handwritten), b) ��� MUS��UM PARIS / 1906 / Coll. L��on FAIRMAIRE��� (handwritten). One female paratype at USNM labeled a) ��� Montevideo / Cerro / 24. IX. 33 ��� (handwritten), b) ��� Apterops / translucidus / (Chand. i. litt.)��� (handwritten), c) ���OLCartwright / Collection / 1959��� (typeset). Two female paratypes at USNM labeled a) ��� Montevideo / Cerro / 12 IX 34 ��� (handwritten). All paratypes listed above also bear a yellow paratype label and a database label with the database number and a barcode. Description of holotype (Figs. 6���10). Male. Length 6.6 mm, width 3.1 mm. Dorsal and ventral color light brown, elytra translucent. Dorsal surface glabrous. Head: surface moderately punctate. Labrum with apex reflexed backward and convexly parabolic medially; underside concave medially. Clypeus with apex straight, abutted to labrum. Frontoclypeal suture visible. Eyes round, slightly protruding laterally. Antennae with 8 antennomeres; antennal club with 3 antennomeres, club distinctly shorter in length to antennomeres 1���5 combined, club surface thickly setose. Maxillae robust, triangular, basal width slightly less than width of mentum; maxillary palpus with 4 palpomeres, length approximately �� that of antennae. Mentum prominent, slightly longer than wide, apex straight, surface slightly convex; labial palpus reduced. Pronotum: surface moderately to densely punctate; apical angle acute, basal angle squared. Pronotum widest medially with smooth margins. Elytra: surface moderately to densely punctate, striae punctate but obscured by other surface punctures. Wings: apterous. Venter: metasternum with dense punctures. Pygidium flat; surface moderately to sparsely punctate with microsculpturing. Legs: protibia with 3 apical teeth on outer margin of apical half; 3rd tooth weak. Claws symmetrical with medial tooth. Protibial spur present, metatibial spurs approximately equal in length. Genitalia: parameres long, slender, tapering into needlelike apices (Fig. 10). Variation. Length 5.9���7.7 mm. Dorsal color tan to brown. Females with slightly more inflated, convex abdominal segments. Etymology. This name is in reference to the translucent elytra and an homage to ��� Apterops translucidus ���, which was used on older determination labels for some specimens but was never published. It should be treated as an adjective in the nominative singular. Distribution (Fig. 66). URUGUAY (14): Montevideo (8): Cerro, Montevideo; no data (6). ARGENTINA (14): Buenos Aires (14): Azul, La Plata. NO DATA (1): ��� Brazil ���. Temporal data. May (13), September (4), October (2). Remarks. Apteroathlia translucida new species is mainly found around the gulf shores of R��o de la Plata in Uruguay and Buenos Aires Province, Argentina. One 19th Century specimen is labeled ��� Brazil ���, but we consider this record doubtful and in need of further verification. The specimens collected in May were probably collected fresh while those obtained in September and October appear to have been found dead and dried. The conservation status of this species should be investigated since most of the specimens were collected from coastal habitats in highly populated urban areas., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 283-285, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674

Key to species of Athlia (based on Saylor 1946b; Gutiérrez 1949; Martínez 1967, 1974) 1. Metathoracic wings reduced and non-functional (slightly lift an elytron to check). Central Chile (VIII Región del Biobío to IX Región de la Araucanía) (Fig. 69)..................................................... Athlia plebeja Burmeister - Metathoracic wings full sized and functional................................................................ 2 2. Pronotum with basal angles evenly rounded, not protruding or angulate (Figs. 30, 34, 46)............................ 3 - Pronotum with basal angles protruding or angulate (Figs. 13, 17, 22, 26, 42, 52).................................... 5 3. Length less than 10 mm. Metatarsomere 1 noticeably shorter than metatarsomere 2 (Fig. 32). Southern Argentina (Neuquén to Chubut, Fig. 69)................................................................... Athlia parvissima Saylor - Length greater than 10 mm. Metatarsomere 1 equal to or slightly shorter than metatarsomere 2 (Fig. 28)................. 4 4. Labrum weakly curved, weakly elevated medially; lateral border of clypeus slightly reflexed; dorsal surface tan to chestnut brown. Central Chile (V Región de Valparaíso, Fig. 79)................................... Athlia rotundata Gutiérrez - Labrum evenly curved and elevated medially; lateral border of clypeus strongly reflexed; dorsal surface light brown to light tan. Southern Argentina (Neuquén to Chubut) and southern Chile (Región de Aisén, Fig. 68).......... Athlia giaii Martínez 5. Labrum not reflexed, on the same plane as clypeal surface. Southeastern Brazil (Santa Catarina, Fig. 67).................................................................................................... Athlia brasilica Saylor - Labrum reflexed above the plane of clypeal surface........................................................... 6 6. Pronotum surface with dense, even punctures (Figs. 22, 42, 52)................................................. 7 - Pronotum surface with sparse, sometimes unevenly distributed punctures (Figs. 13, 26).............................. 9 7. Elytra evenly covered with setae; pronotum with long setae along lateral edges. Central Chile (IV Región de Coquimbo to IX Región de la Araucanía, Fig. 72)........................................................ Athlia rustica Erichson - Elytra glabrous medially and sparsely setose laterally and apically; pronotum with long setae along lateral edge but mainly in apical half........................................................................................... 8 8. Elytra length less than twice the length of head and pronotum combined Northern and central Argentina (Jujuy to Río Negro, Fig. 67).............................................................................. Athlia bruchi Moser - Elytra length greater than twice the length of head and pronotum combined. Central Chile (IV Región de Coquimbo to Región Metropolitana de Santiago, Fig. 70)................................................ Athlia problematica Gutiérrez 9. Length greater than 12 mm; apical and basal pronotal angles well developed (Fig. 26). Southern Brazil (São Paulo, Fig. 67)...................................................................................... Athlia freyi Martínez - Length less than 12 mm; apical and basal pronotal angles weakly developed (Fig. 13). Northeastern Argentina (Buenos Aires, Fig. 67)............................................................................ Athlia bollei Martínez, Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 285-286, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Saylor, L. W. (1946) Studies in the melolonthine scarab beetle genera of the American continents. I. Revision of the genus Athlia. Proceedings of the Entomological Society of Washington, 48, 18 - 25.","Gutierrez, R. (1949) Notas sobre Scarabaeidae neotropicos (Coleoptera Lamellicornia). Anales de la Sociedad Cientifica Argentina, 148, 9 - 35.","Martinez, A. (1967) El genero Athlia Erichson (Col. Scarabaeidae, Sericinae). Entomologische Arbeiten aus dem Museum G. Frey, 18, 327 - 372.","Martinez, A. (1974) Nuevas consideraciones sobre el Genero Athlia Erichson, 1835 (Col., Scarabaeidae, Sericinae). Entomologische Arbeiten aus dem Museum G. Frey, 25, 343 - 353."]}

Athlia Erichson, 1835 Athlia Erichson, 1835: 266. Type species Athlia rustica Erichson, 1835 by monotypy. Synonym. Rivera Germain, 1903: 392. Type species Athlia plebeja Burmeister, 1855 by monotypy. Redescription. Length 6.5–17.0 mm. Elongate and robust, widest at about elytral declivity; uniformly pale to medium yellowish brown, or brown or reddish brown; dorsal surface dull, sometimes with a faint, glaucous coating on pronotal and elytral surfaces, or weakly shiny, sometimes faintly iridescent, and with or without short, scattered, and appressed setae. Head: labrum wide, broadly emarginate and evenly reflexed anteriorly, and completely fused to clypeus with labroclypeal suture more-or-less distinct and terminating in small, marginal notches. Mentum longer than wide, narrowly emarginate in front, with sides gradually converging posteriorly. Antennae with 9 antennomeres, club with 3 short, thick lamellae. Pronotum: convex, wider than long, widest medially or slightly behind middle. Elytra: disc of each with four weak and narrow costae, interstriae broad and confusedly punctate. Metathoracic wings fully developed or reduced in size (brachypterous). Abdomen: with lateral carinae on ventrites. Legs: protibia with three apical teeth on outer margin, spur present and shorter than protarsomere 1. Claws symmetrical and incised, with short, bisetose onychium in between. Metatibial apices with spurs set in notch, longest spur reaching half to three-quarter point of metatarsomere 1. Male genitalia: symmetrical parameres simple in form, not split or with strong sinuations, and somewhat shorter than phallobase. Composition. All seven known species of Athlia are endemic to the southern half of South America. Remarks. Saylor (1946b) and Martínez (1967, 1974) previously reviewed this genus but a detailed taxonomic inventory of all specimens in collections is needed. We predict that new species of Athlia will be found in southern Brazil and northern Argentina, where endemism is high and specimens are poorly represented in collections., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 285, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Erichson, W. F. (1835) Neue Sudamerikanische Kafergattungen aus der familie der Blatterhorner. Archiv fur Naturgeschichte, 1, 256 - 270.","Germain, P. (1903) Apuntes entomologicos. Anales de la Universidad. Republica de Chile, 112 - 113, 391 - 445.","Burmeister, H. C. C. (1855) Handbuch der Entomologie. Fierter Band. Besondere Entomologie. Fortsetzung. Zweite Abtheilung. Coleoptera Lamellicornia Phyllophaga chaenochela. Theod. Chr. Friedr. Enslin, Berlin, x + 569 pp.","Saylor, L. W. (1946) Studies in the melolonthine scarab beetle genera of the American continents. I. Revision of the genus Athlia. Proceedings of the Entomological Society of Washington, 48, 18 - 25.","Martinez, A. (1967) El genero Athlia Erichson (Col. Scarabaeidae, Sericinae). Entomologische Arbeiten aus dem Museum G. Frey, 18, 327 - 372.","Martinez, A. (1974) Nuevas consideraciones sobre el Genero Athlia Erichson, 1835 (Col., Scarabaeidae, Sericinae). Entomologische Arbeiten aus dem Museum G. Frey, 25, 343 - 353."]}

Dihymenonyx suboblongus Mondaca, 2007 Original combination. Dihymenonyx suboblongus Mondaca, 2007: 20. Type locality. “ Chile, VIII Región del Biobío, Provincia de Ñuble, Cerro Cayumanqui (36°42'08.5''S y 72°30'36.2''W)” (Mondaca 2007). Type series. Holotype at MNNC (not examined). Distribution. Fig. 73. Chile: VIII Región del Biobío (Mondaca 2007)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 296, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Mondaca, J. (2007) Revision del genero Dihymenonyx Gutierrez (Scarabaeidae: Melolonthinae: Sericini) con descripcion de dos nuevas especies de Chile. Revista Chilena de Entomologia, 32, 13 - 29. [dated 2006]"]}

Apteroathlia Smith & Evans, new genus http://zoobank.org/urn:lsid:zoobank.org:act:778D2B0C-E445-46BC-84CB-338DA4F2F7F6 Type species. Apteroathlia translucida Smith & Evans, new species, here designated. Description. Length 5.9���9.4 mm. Ovate, widest behind middle of elytra; testaceous to black, or black with testaceous appendages and partly or entirely testaceous pronotum; surfaces moderately to strongly shiny, and mostly glabrous. Head: labrum wide, broadly emarginate, evenly and shallowly reflexed anteriorly, completely fused to and slightly below plane of clypeus exposing anterior clypeal margin; labroclypeal suture distinct and terminating in small, marginal notches. Mentum longer than wide, narrowly emarginate in front, with sides gradually converging posteriorly. Antennae with 8 antennomeres, club with 3 short, thick lamellae. Pronotum: convex, wider than long, widest medially. Elytra: convex and without costae or distinct interval, punctostriate. Metathoracic wings absent. Abdomen: with lateral carinae on ventrites. Legs: protibia weakly tridentate on outer margin, spur present and reaching to basal third or half of protarsomere 1. Claws symmetrical and toothed with a short, bisetose onychium in between. Metatibial apices with spurs set in notch, each set on either side of tarsal articulation, longest spur reaching about two-thirds to three-quarters the length of first tarsomere. Male genitalia: symmetrical parameres simple in form, not split or with strong sinuations, and subequal in length to phallobase. Etymology. Apteroathlia means ���wingless Athlia ��� to indicate the similarities with the genus Athlia. The name is feminine in gender to match Athlia. Composition. Apteroathlia nox new species and Apteroathlia translucida new species. Remarks. The two known species in this genus are both endemic to the region around R��o de la Plata in Uruguay and Argentina., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 281-282, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674

Athlia bruchi Moser, 1924 Figs. 19–23, 67. Original combination. Athlia bruchi Moser, 1924: 121. Type locality. “ Argentina: Cordoba ” (Moser 1924). Type series. Lectotype (Figs. 19–23) at ZMHB labeled a) “Rep ARGENTINA / Prov. Cordoba / 1911 / C. Bruch” (typeset), b) “ Athlia / Bruchi / Type Mos.” (handwritten), c) “Bruchi Mos.” (handwritten), d) “SYNTYPUS / Athlia / bruchi Moser, 1924 / labelled by MNHUB 2009” (typeset, orange label), e) “ ATHLIA / BRUCHI / MOSER, 1924 / LECTOTYPE / A.B.T. SMITH” (handwritten and typeset, red label). Lectotype here designated. A lectotype is designated to preserve the stability of the nomenclature by selecting a single specimen as the namebearing type for this species. Since Moser (1924) did not explicitly state how many specimens he examined, it is impossible to know the exact composition of the type series. Distribution. Fig. 67. Argentina: Buenos Aires, Córdoba, Jujuy, La Pampa, Neuquén, Río Negro, San Luis (based on specimens from CASC, CMNC, ZMHB)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 286-287, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Moser, J. (1924) Beitrag zur Kenntnis der Melolonthiden (Col.). (XIII). Stettiner Entomologische Zeitung, 84, 114 - 136."]}

Ulata argentina Saylor, 1945 Figs. 61���65, 74. Original combination. Ulata argentina Saylor, 1945: 442. Type locality. Pucar��, Lago Lacar, Neuqu��n, Argentina (approximately 40.16��S, 71.63��W). The original type locality was ��� Bariloche, Rio Negro, Argentina ��� (Saylor 1945). Type series. Neotype at CMNC labeled a) ���ARGENTINA / NEUQUEN / Lago Lacar / Pucar�� / Shajovskogleg. / Coll. Mart��nez.- / NOV.-964��� (handwritten), b) ���H. & A. HOWDEN / COLLECTION / ex. A.Martinez coll.��� (typeset with black border), c) ��� Ulata ♂ / argentina / Saylor / A. MARTINEZ-DET. 1965��� (handwritten and typeset), d) ���Southern Neotropical Scarabs / database # AS 2615414 / Ulata argentina / Saylor, 1945 ♂ / DET: A.B.T. SMITH 2 008��� (typeset with black border), e) ��� ULATA / ARGENTINA SAYLOR, 1945 ♂ / NEOTYPE ��� (handwritten and typeset with black border on red paper). Neotype here designated. Saylor (1945) stated that this species was described using only the holotype, which was presumably deposited in his personal collection (although no collection deposition was explicitly stated it was Saylor���s common practice to keep holotypes in his personal collection). Saylor���s personal collection was donated to the CASC in 1963 (Ratcliffe 2016), but this institution has no record of the Ulata argentina holotype (the rest of Saylor���s primary types are catalogued and kept in the type collection). The first author carefully searched the entire CASC scarab collection for the holotype, but found nothing. Therefore, we conclude that the holotype has been lost. Due to the historic instability of the classification of this taxon, the neotype is designated with the express purpose of clarifying the taxonomic status of this species and genus. The neotype is consistent with the original description and originated from relatively close to the original type locality (less than 80 km distance). Distribution. Fig. 74. Argentina: Neuqu��n, R��o Negro; Chile: V Regi��n de Valpara��so, Regi��n Metropolitana de Santiago, VIII Regi��n del Biob��o, IX Regi��n de la Araucan��a (based on the original type locality and specimens from AMNH, BMNH, CMNC, FMNH, TMSA, UMCE, VMDM)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 297-299, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Saylor, L. W. (1945) Studies in the melolonthine scarab beetle genera of the American continents. No. IV. A new genus from Argentina. Revista de Entomologia, 16, 441 - 444.","Ratcliffe, B. C. (2016) Lawrence W. Saylor, coleopterist extraordinaire. Coleopterists Bulletin, 70, 279 - 287. https: // doi. org / 10.1649 / 0010 - 065 X- 70.2.279"]}

Athlia brasilica Saylor, 1946 Figs. 15���18, 67. Original combination. Athlia brasilica Saylor, 1946: 23. Type locality. ���Nova Galicia, S. Catarina, Brazylja ��� (Saylor 1946). Type series. Holotype at CASC labeled a) ���Nowa Galicja / Brazylja,S. Catharina ��� (typeset), b) ��� HOLOTYPE / Athlia / BRASILICA / L.W.Saylor��� (typeset and handwritten on red paper), c) ���BRASILICA / Saylor / DET. / L.W. SAYLOR��� (handwritten and typeset), d) ���L.W. Saylor / Collection��� (typeset), e) ��� California Academy / of Sciences / Type / No. 7922��� (typeset and handwritten). One paratype at CASC (examined). Distribution. Fig. 67. Brazil: Santa Catarina (based on specimens from CASC, CMNC)., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 286, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Saylor, L. W. (1946) Studies in the melolonthine scarab beetle genera of the American continents. I. Revision of the genus Athlia. Proceedings of the Entomological Society of Washington, 48, 18 - 25."]}

Athliini Smith & Evans, new tribe http://zoobank.org/urn:lsid:zoobank.org:act:1EF3248F-813E-4307-AD07-43E4F3F75677 Type genus. Athlia Erichson. Diagnosis. Labrum transverse, originating below and fused to anterior clypeal margin so that they are coplanar; labroclypeal suture evident. Clypeus and frons are coplanar. Ocular canthus present, margin continuous with lateral clypeal margin. Antennae with 8 or 9 antennomeres, club with three short, thick, subequal lamellae. Mentum with insertions of labial palpi not exposed. Pronotum with anterior margin membranous. Abdomen with 6 ventrites, 2–5 equal in length and not strongly narrowed medially, 5 and 6 not separated by a membrane, with (Apteroathlia, Athlia) or without carinae on sides, and a distinct propygidial-sternal suture. Propygidium plain, fused to tergite with suture more-or-less evident, spiracle not within or immediately adjacent to suture. Procoxae conical and protuberant, anterior height greater than basal width. Protibial spur present. Metepimeron triangular with a distinct ridge. Outer surfaces of mesotibiae and metibiae with indication of partial or complete (Athlia) transverse carinae. Metatibial apex transverse, inner margin distinctly notched with two spurs, each set above and below tarsal articulation so that the first tarsomere passes between them. Opposing claws symmetrical, incised, toothed, or simple, with a short and bisetose onychium; claws of Dihymenonyx with a fleshy lobe underneath each side of claw. Composition. Apteroathlia Smith & Evans, new genus, Athlia, Dihymenonyx, and Ulata. Distribution. Southern half of South America. Notes. Athliini new tribe is distinguished from all Neotropical Sericoidini and Australian Liparetrini (and similar Australian tribes) by the distinct labrum that is fused to and coplanar with the anterior clypeal margin, a morphological feature that superficially resembles that found in the Sericini. Sexual dimorphism in this tribe is very subtle and difficult to detect. The antennal club is reduced in both sexes (generally less than half the length of the funicle) and there are no sexually dimorphic characters in the head or legs. The convexity of the abdomen is generally sexually dimorphic but the differences are subtle since the abdomen in both sexes appears convex or inflated. Three of the genera in this tribe (Apteroathlia, Athlia, Dihymenonyx) include species with reduced or absent metathoracic wings, thus indicating a general evolutionary trend of flightlessness in this group for species living in arid habitats. Scholtz (2000) hypothesized that some scarab taxa have an increased evolutionary propensity towards flightlessness when they evolve in certain environments such as deserts, mountains, and islands. The immature stages have not been described for any species in this tribe. Studying the larval natural history and morphology of this group may provide insights into the placement of this taxon among other Melolonthinae tribes., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 280-281, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Scholtz, C. H. (2000) Evolution of flightlessness in Scarabaeoidea (Insecta, Coleoptera). Mitteilungen aus dem Museum fur Naturkunde in Berlin. Deutsche Entomologische Zeitschrift, 47, 5 - 28."]}

Apteroathlia nox Smith & Evans, new species Figs. 1–5, 66. http://zoobank.org/urn:lsid:zoobank.org:act:70 ABBE 95-B0D0-4C43-A2DF-4038525D507C Type locality. Playas de Juancho, Madariaga, Buenos Aires, Argentina. Type series. One male holotype, 4 male paratypes, and 2 female paratypes. Holotype male (Figs. 1–5) at CMNC labeled a) “ ARGENTINA / BUENOS AIRES / Do MADARIAGA / Playas de Juancho / Coll. Martínez / Nov. -941” (handwritten), b) “H. & A. HOWDEN / COLLECTION / ex. A.Martínez coll.” (typeset with black border). Two male paratypes at CMNC labeled a) “ ARGENTINA / BUENOS AIRES / Pdo. MADARIAGA / Playas de Juancho / Coll. Martínez / Nov. -941” (handwritten), b) “H. & A. HOWDEN / COLLECTION / ex. A.Martínez coll.” (typeset with black border). Two male paratypes and one female paratype at CMNC labeled a) “ ARGENTINA / BUENOS AIRES / S. Ventana / Gallardo-leg. / Coll. Martínez / 11-VIII-61 ” (handwritten), b) “H. & A. HOWDEN / COLLECTION / ex. A.Martínez coll.” (typeset with black border). Ono female paratype at CMNC labeled a) “ ARGENTINA / BUENOS AIRES / Villa Gesell / Horacio-leg. / Coll. Martínez / Ene.-976” (handwritten), b) “H. & A. HOWDEN / COLLECTION / ex. A.Martínez coll.” (typeset with black border). All paratypes listed above also bear a yellow paratype label and a database label with the database number and a barcode. Description of holotype (Figs. 1–5). Male. Length 7.0 mm, width 3.1 mm. Head (at base), elytra, ventral surface black; head (apex), pronotum, legs dark tan. Dorsal surface glabrous (except sparse row of setae adjacent to apex of clypeus). Head: clypeus densely punctate, frons moderately punctate. Labrum with apex reflexed and concave medially; underside concave medially. Clypeus with apex straight, abutted to labrum. Frontoclypeal suture visible. Eyes round, weakly protruding laterally. Antennae with 8 antennomeres; antennal club with 3 antennomeres, club approximately half the length of antennomeres 1–5 combined, club surface with some setae. Maxillae robust, triangular, basal width slightly less than width of mentum; maxillary palpus with 4 palpomeres, length approximately approximately half that of antennae. Mentum prominent, slightly longer than wide, apex straight, surface slightly convex; labial palpus reduced. Pronotum: surface moderately to sparsely punctate; apical angle slightly acute, basal angle rounded. Pronotum widest medially with smooth margins. Elytra: surface sparsely punctate, striae punctate but somewhat obscured by other surface punctures. Wings: apterous. Venter: metasternum with dense punctures. Pygidium flat; surface sparsely punctate with microsculpturing. Legs: protibia with 3 apical teeth on outer margin of apical half. Claws symmetrical, each with a minute medial tooth (metatarsal claws appear simple). Protibial spur present, metatibial spurs approximately equal in length. Genitalia: parameres long, curving and tapering into needle-like apices (Fig. 5). Variation. Length 7.0– 9.4 mm. Pronotum black, dark tan, or black lateral and dark tan/brown medially. Claws all appear simple on the paratype specimens. Females with slightly more inflated, convex abdominal segments. Etymology. The Latin word nox means night and is used here in reference to the night-like darkness of the dorsal and ventral surfaces. This name is an adjective in the nominative singular. Distribution (Fig. 66). ARGENTINA (7): Buenos Aires (7): Playas de Juancho, Sierra de la Ventana, Villa Gesell. Temporal data. January (1), August (3), November (3). Remarks. Apteroathlia nox new species is endemic to the southern part of Buenos Aires Province, Argentina. The holotype specimen has claws with a minute, medial tooth but the tooth was not seen in other specimens. This perhaps indicates that the tooth is easily worn off., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on page 282, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674

Athlia problematica Guti��rrez, 1950 Figs. 40���43, 70. Original combination. Athlia problematica Guti��rrez, 1950: 270. Type locality. ��� Chile: Prov. Valpara��so, Conc��n ��� (Guti��rrez 1950). Type series. Holotype, allotype, and eight paratypes at Museo de Zoolog��a, Universidad de Concepci��n, Concepci��n, Chile (Cekalovic & Artigas 1969). Two paratypes at BMNH, one paratype at FMNH, and two paratypes at CMNC were examined. Distribution. Fig. 70. Chile: IV Regi��n de Coquimbo, V Regi��n de Valpara��so, Regi��n Metropolitana de Santiago (based on specimens from AMNH, BMNH, CASC, CMNC, FMNH, UMCE). The IV Regi��n de Coquimbo record is based on a single AMNH specimen collected at Combarbal�� on 11 November 1992 by Rosen, Sharkov, and Snyder. Additional specimens are needed to corroborate the Regi��n de Coquimbo distributional record., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 292-293, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Gutierrez, R. (1950) Notas sobre Scarabaeidae Chilenos (Coleoptera Lamellicornia). Arthropoda, 1 (2 / 4), 267 - 278.","Cekalovic, T. & Artigas, J. N. (1969) Catalogo de los tipos de Insecta depositados en la coleccion del Departamento de Zoologia de la Universidad de Concepcion (INCO) (Diciembre, 1968). Boletin de la Sociedad de Biologia de Concepcion, 41, 111 - 133."]}

Athlia plebeja Burmeister, 1855 Figs. 36–39, 69. Original combination. Athlia plebeja Burmeister, 1855: 125. Subsequent combination. Rivera plebeja (Burmeister): Germain 1903: 392. Type locality. “ Chili ” (Burmeister 1855). Type series. Syntypes presumably at Martin-Luther-Universität, Halle, Germany (not examined). Synonym. Athlia rivera Saylor, 1946: 23. Type locality: “ Chile, Concepcion” (Saylor 1946). Type series: holotype male at CASC labeled a) “ CHILE / Concepcion / P.HerbstS.” (typeset) on upperside and “ 30.x.04.” (handwritten) on underside, b) “Moser determ. / Athlia n.sp. ” (typeset and handwritten), c) “ HOLOTYPE / Athlia / RIVERA / L.W.Saylor” (typeset and handwritten on red paper), d) “ RIVERA / Saylor / DET. / L.W. SAYLOR” (handwritten and typeset), e) “L.W. Saylor / Collection” (typeset), f) “ California Academy / of Sciences / Type / No. 7924” (typeset and handwritten), g) “Southern Neotropical Scarabs / database # AS 2617639 / Athlia plebeja / Burmeister, 1855 ♂ / DET: A.B.T. SMITH 2008 ” (typeset). One paratype at CASC erroneously labeled “ Peru ” (examined). Distribution. Fig. 69. Chile: VII Región del Libertador General Bernardo O’Higgins, VIII Región del Biobío, IX Región de la Araucanía (based on specimens from BMNH, CASC, CMNC, FMNH, MNHN, USNM). Two century-old specimens were also seen labeled “Limache” (CASC) and “ Valparaíso ” (CMNC) but we consider these records as erroneous. The Valparaíso region is far outside the distribution of this species., Published as part of Smith, Andrew B. T. & Evans, Arthur V., 2018, Taxonomic review of Athliini (Coleoptera: Scarabaeidae: Melolonthinae), a new tribe of scarab beetles endemic to South America, pp. 279-308 in Zootaxa 4471 (2) on pages 289-292, DOI: 10.11646/zootaxa.4471.2.3, http://zenodo.org/record/1439674, {"references":["Burmeister, H. C. C. (1855) Handbuch der Entomologie. Fierter Band. Besondere Entomologie. Fortsetzung. Zweite Abtheilung. Coleoptera Lamellicornia Phyllophaga chaenochela. Theod. Chr. Friedr. Enslin, Berlin, x + 569 pp.","Germain, P. (1903) Apuntes entomologicos. Anales de la Universidad. Republica de Chile, 112 - 113, 391 - 445.","Saylor, L. W. (1946) Studies in the melolonthine scarab beetle genera of the American continents. I. Revision of the genus Athlia. Proceedings of the Entomological Society of Washington, 48, 18 - 25.","Smith, A. B. T. (2008) South American Melolonthinae (Coleoptera: Scarabaeidae) classification and nomenclature: some problems and solutions. Insecta Mundi, 60, 1 - 28."]}

Evans, Arthur V. (2017): Book Reviews. The Coleopterists Bulletin 71 (2): 298-300, DOI: 10.1649/0010-065X-71.2.298, URL: http://dx.doi.org/10.1649/0010-065x-71.2.298

Evans, Arthur V., Schnepp, Kyle E. (2012): Notes on the distribution and habitat of Omethes marginatus LeConte (Coleoptera: Omethidae). Insecta Mundi 2012 (244): 1-6, DOI: http://doi.org/10.5281/zenodo.5174541, {"references":["Branham, M. A., and J. W. Wenzel. 2001. The evolution of bioluminescence in cantharoids (Coleoptera: Elateroidea). Florida Entomologist 84(4): 565-586.","Branham, M.A. and J. W. Wenzel. 2003. The origin of photic behavior and the evolution of sexual communication in fireflies (Coleoptera: Elateroidea). Cladistics 19(1): 1-22.","Drury, C. 1902. A revised list of the Coleoptera observed near Cincinnati, Ohio, with notes and references on localities, bibliographical references, and a description of new species. Journal of the Cincinnati Society of Natural History 20: 107-189.","Evans, A. V. 2009. Phyllophaga spreta (Horn), a rare species of June beetle new to the fauna of Virginia, North Carolina, and Pennsylvania (Coleoptera: Scarabaeidae). Banisteria 33: 37-42.","Fleming, G. P. 2002. Ecological communities of the Bull Run Mountains, Virginia: baseline vegetation and floristic data for conservation planning and natural area stewardship. Natural Heritage Technical Report 02-12. Virginia Department of Conservation and Recreation, Division of Natural Heritage; Richmond, VA. 274 p. (Unpublished report submitted to the Virginia Outdoors Heritage Foundation).","Holland, J. D. 2006. Cerambycidae larval host condition predicts trap efficiency. Environmental En- tomology 35: 1647-1653.","Holland, J. D. 2010. Isolating spatial effects on beta diversity to inform forest landscape planning. Landscape Ecology 25: 1349-1362","Malkin, B. 1945. A supplement to the New York state list of Coleoptera, No. 6. Additions and corrections. Journal of the New York Entomological Society 53: 92-116.","McCormick, J. 1962. Vascular flora of Shades State Park and Pine Hills Natural Area, Indiana. Bul- letin of the American Museum of Natural History 123 Article 7: 353-422.","Ramsdale, A. S. 2002. 63. Omethidae LeConte 1861. p. 197-201. In: R. H. Arnett, Jr., M. C. Thomas, P. E. Skelley, J. H. Frank (eds.). Volume 2. American Beetles. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press; Boca Raton, FL. 861 p.","Ramsdale, A. S. 2010. 4.16 Omethidae, LeConte. p. 149-152. In: R. A. B. Leschen, R. G. Beutel, and J. F. Lawrence (eds.). Coleoptera, Beetles. Volume 2: Morphology and systematics (Elateroidea, Bostrichiformia, Cucujiformia partim). Handbook of Zoology. Arthropoda: Insecta. De Gruyter; Berlin. 786 p.","Scott, R. K. 2009. The vascular flora of Turkey Run State Park, Parke County, Indiana. Proceedings of the Indiana Academy of Science 118(1): 55-75","Ulke, H. 1902. A list of the beetles of the District of Columbia. Proceedings of the United States National Museum 25: 1-57."]}

Evans, Arthur V. (2008): Book Review. Illustrated Key to the Longhorned Woodboring Beetles of the Eastern United States. The Coleopterists Bulletin 62 (3): 425-426, DOI: 10.1649/1113.1