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Nanotermitodius Howden, 2003 Nanotermitodius Howden, 2003: 393. Type species: Nanotermitodius peckorum Howden, 2003: 396, by original designation. Diagnosis. Nanotermitodius is unique among the rhyparines by having the male sexual dimorphism of a Rhyparus - type mesotibia and Termitodiellus - type metatibia (Skelley 2007). The females have simple, unmodified tibial apices. Other useful characters to diagnose Nanotermitodius include elytral carinae sinuate in dorsal and lateral views, caudal apex of discomedian costa on elytra with an erect tuft of setae, mesoventrite with shallow longitudinal median groove, metaventrite with distinct discolateral fovea on either side of midline, and male protibia distinctly tridentate at apex, female protibia weakly tridentate. The genus also has fused elytra with flight wings reduced to vestigial buds, reduced eyes, and is presently known only from the state of Oaxaca, Mexico. Comments. Howden (2003) provided a generic description, which also acted as the species description for N. peckorum. Thus, a brief diagnosis of the genus listing important delimiting characters was needed. Other traditional characters like body shape, costal development, etc. vary between these two Nanotermitodius enough to overlap with other genera, making these characters less useful for generic delimitation. Skelley (2007) noted that male N. peckorum have Rhyparus - type mesotibia but Termitodiellus - type metatibia. The new species shows this character combination more prominently than N. peckorum. This tibial combination in the males will readily distinguish Nanotermitodius., Published as part of Skelley, Paul E., Smith, Andrew B. T. & Mora-Aguilar, Eder F., 2022, A review of the flightless genus Nanotermitodius Howden, 2003 (Coleoptera Scarabaeidae: Aphodiinae: Rhyparini), pp. 355-364 in Zootaxa 5200 (4) on page 356, DOI: 10.11646/zootaxa.5200.4.4, http://zenodo.org/record/7270716, {"references":["Howden, H. F. (2003) Two new genera of New World Rhyparini (Coleoptera: Scarabaeidae: Aphodiinae). Sociobiology, 42, 391 - 402.","Skelley, P. E. (2007) Generic limits of the Rhyparini with respect to the genus Termitodius Wasmann, 1894 (Coleoptera: Scarabaeidae: Aphodiinae). Insecta Mundi, 9, 1 - 9."]}

Nanotermitodius peckorum Howden, 2003 Figures 4–6, 16 Nanotermitodius peckorum Howden, 2003: 396. Type material examined (2). Holotype male and allotype female (CMNC): “ MEXICO: Oaxaca, 1220m, 26 km E. Valle Nacional, 25.VI.1983, S. & J. Peck, Mont. trop. for. litter Ber. Km 71, 83-50”. Additional material examined (14): MEXICO: Oaxaca: Hwy. 175, 22mi. S. Valle Nacional, 5800’, 3-VI-1983, CW. & L. O’Brien & GB. Marshall, Berlese sifted cloud forest litter (1♂, 1♀ FSCA); 18.7 mi S Valle Nacional, 5200’, 17-VIII-1973, A. Newton, Berlese litter cloud forest (2♂♂, 2♀♀ FSCA; 1♂, 1♀ IEXA; 1♂, 5♀♀ MCZC); 18.7 mi S Valle Nacional, 5200’, 11-18-VIII-1973, A. Newton, leaf litter forest floor (1♀ MCZC). Diagnosis. Nanotermitodius peckorum (Figs. 4–6) differs from N. andersoni in smaller body size (length 2.5–2.6 mm), pronotal lateral margin deeply sinuate, short elytra (1.5 times length of pronotum), which converge posteriorly having more sinuate costae, flight wings vestigial, and other characters including those in key. Distribution. Known only in wet slope of Sierra de Juarez, Oaxaca, Mexico, commonly known as “La Chinantla”, an elevation of 1220–1750 m (Fig. 16). Comments. There appears to be an error in the label data of the type series. The marker “km 71” along the highway 157 Tuxtepec-Oaxaca is 26 km south southwest of Valle Nacional, not east. This area matches the elevation (1220 m) and vegetation (montane tropical forest) as stated on the label. To the east of Valle Nacional the territory is below 800 m and has different lower elevation plant communities. The locality Puerto Antonio is located at km 71–72, we suspect this may be the actual type locality. This error in labeling is supported by data associated with additional specimens that have been recently studied. Discussion. Sierra de Juárez is considered one of the most diverse regions of the state and present one of the two highest number of endemic species of arthropods (González-Pérez et al. 2004). In “La Chinantla”, cloud forest exhibit three variations, in lower elevations is represented by species of Lauraceae, Ilex L. (Aquifoliacee), Podocarpus, Alchornea Sw. (Euphorbiaceae) and with Oreomunnea mexicana (Standley) Leroy (Juglandaceae) (1400–1600 m), O. mexicana is the dominant element between 1600–1800 m of elevation, and cloud forests in higher elevations (1800–2050 m), where O. mexicana and several species of the genus Quercus (Fagaceae) are present as dominant trees (Rzedowski & Palacios-Chávez 1977). Particularly, cloud forests with O. mexicana as the dominant element are considered relict and paleoendemic vegetation established in areas that presumably served as Pleistocene refugia of flora and fauna in Mexico (Rzedowski 1991)., Published as part of Skelley, Paul E., Smith, Andrew B. T. & Mora-Aguilar, Eder F., 2022, A review of the flightless genus Nanotermitodius Howden, 2003 (Coleoptera Scarabaeidae: Aphodiinae: Rhyparini), pp. 355-364 in Zootaxa 5200 (4) on pages 361-362, DOI: 10.11646/zootaxa.5200.4.4, http://zenodo.org/record/7270716, {"references":["Howden, H. F. (2003) Two new genera of New World Rhyparini (Coleoptera: Scarabaeidae: Aphodiinae). Sociobiology, 42, 391 - 402.","Gonzalez-Perez, G., Briones-Salas, M. & Alfaro, A. M. (2004) Integracion del conocimiento faunistico del Estado. In: GarciaMendoza, A. J, Ordonez Diaz, M. J. & Briones-Salas, M. (Eds.), Biodiversidad de Oaxaca. Fondo Oaxaqueno para la Conservacion de la Naturaleza, World Wildlife Fund, Mexico, pp. 449 - 465.","Rzedowski, J. & Palacios-Chavez, R. (1977) El bosque de Engelhardtia (Oreomunnea) mexicana en la region de La Chinantla (Oaxaca, Mexico) - una reliquia del Cenozoico. Boletin de la Sociedad Botanica de Mexico, 36, 93 - 127. https: // doi. org / 10.17129 / botsci. 1161","Rzedowski, J. (1991) El endemismo de la flora fanerogamica mexicana: una apreciacion analitica preliminar. Acta Botanica Mexicana, 15, 47 - 64. https: // doi. org / 10.21829 / abm 15.1991.620"]}

Allsopp, Peter G., Smith, Andrew B.T. (2022): Australian Melolonthini (Coleoptera: Scarabaeidae: Melolonthinae): a third species of Allothnonius Britton, 1978, and notes on the other known species. Zootaxa 5099 (4): 475-484, DOI: https://doi.org/10.11646/zootaxa.5099.4.3

Allothnonius mouldsi Allsopp & Smith, new species (Figs. 1���6, 18) urn:lsid:zoobank.org:act: 769A8492-8798-458D-B914-B49E4FCEF8B1 Type series. Holotype ♂. NORTHERN TERRITORY, Keep River x-ing [Crossing] [15.872��S, 129.182��E], Victoria Hwy [Highway], N.T. 7 Jan. 1986 M.S. & B.J. Moulds | Photographed Specimen [purple label] | QM Reg. No. T 250929 [yellow label]; in QM. Paratype. 1♂, same collection data as holotype [CMNC]. Description. Male holotype (Figs. 1���5). Body 14.5 mm long; head black; pronotum very dark brown to black; scutellum lighter brown; elytra, pygidium, legs and abdominal ventrites brown; antennae pale yellow-brown; palpi reddish brown. Labrum rounded laterally with deep median indentation, not protruding beyond anterior margin of upper clypeus, with a mixture of scattered short and longer setae. Terminal palpomere of maxillary palp slender, about as long as palpomeres 1���2 together, dull area on upper surface. Anterior face of clypeus 5x as wide as long, with line of setiferous punctures across middle and laterally; outline of upper surface broadly rounded with a slight median indentation, without an emargination but reflexed on anterior, surface very coarsely punctured, with flattened, adpressed, white setae, longer, thinner and denser laterally. Frons coarsely and densely punctured, anterior quarter with long, thin, white setae with a few shorter, broader, flattened, white setae in between; median half covered with dense, flattened, white setae. Ocular canthi with longer, pale-yellow setae in addition to flattened, white setae. Antennal club in males 7-lamellate but lamella of antennomere 4 0.8x length of other lamellae; antennomere 3 wedge-shaped. Pronotal width across base 2x mid length, sides obtusely angled behind middle, anterior and posterior angles obtuse, posterior angles slightly rounded, anterior and posterior margins narrowly raised and defined by grooves continuous from side to side; surface coarsely punctured, with short, flattened, white setae varying in size from contained within their punctures to about twice puncture diameter. Scutellum with scattered setae similar to pronotal setae. Elytral sutural and two other striae impressed, surface fairly uniformly punctured, punctures smaller than those on pronotum and each containing a small, flattened, pale seta; lateral margins with fringe of light-brown setae. Propygidium uniformly and densely clothed with recumbent, parallel-sided, flattened, pale-yellow setae. Pygidium with scattered, flattened, white setae, sparser in posterior angle; posterior margin with longer, finer, brown setae. Ventral thorax with dense, long, pale-yellow setae. Legs with scattered pale-yellow setae and a few flattened, white setae. Abdominal ventrites laterally with dense, flattened, white setae, longer and more scattered medially, posterior half of penultimate segment and posterior segment almost bare, bare area on penultimate segment bounded anteriorly by row of long, yellow-brown setae, posterior margin of posterior segment with long, yellow-brown setae. Aedeagus (Figs. 3���5) symmetrical, lateral edges slightly tapering then expanded, rounded and folded dorsally at apices. Variation (paratype). Body 13.8 mm long; head very dark brown to black; pronotum dark brown; scutellum and elytra brown with apices slightly darker; pygidium, ventral thorax, and legs lighter brown; ventrites dark brown except posterior half of penultimate and all of final ventrite lighter brown. Anterior face of clypeus 4.7x as wide as long. Pronotal length 1.8x width. Scutellum laterally with a few scattered setae similar to pronotal setae, centre almost bare. Abdominal ventrites with dense, flattened white setae extending across middle (except on posterior half of penultimate ventrite and posterior ventrite). Female. Unknown. Etymology. Named for Max Moulds, continuing the use in the genus of names that honour one of the collectors. A noun in the genitive case. Distribution. Known only from the type locality (Fig. 18); it has a K��ppen-Geiger climate classification (Beck et al. 2018) of Aw (savannah). Natural history. The type series was collected during early January. Both specimens had moth scales attached, so were presumably taken at light., Published as part of Allsopp, Peter G. & Smith, Andrew B. T., 2022, Australian Melolonthini (Coleoptera: Scarabaeidae: Melolonthinae): a third species of Allothnonius Britton, 1978, and notes on the other known species, pp. 475-484 in Zootaxa 5099 (4) on pages 476-478, DOI: 10.11646/zootaxa.5099.4.3, http://zenodo.org/record/6091678, {"references":["Beck, H. E., Zimmermann, N. E., McVicar, T. R., Vergopolan, N., Berg, A. & Wood, E. F. (2018) Present and future Koppen- Geiger climate classification maps at 1 - km resolution. Scientific Data, 5, 180214. https: // doi. org / 10.1038 / sdata. 2018.214"]}

Allothnonius brooksi Britton, 1978 (Figs. 11���18) Allothnonius brooksi Britton, 1978: 39, Figs. 13, 117���119, 276A; Weir et al. 2019, Fig. 31.156, Plate 60B. Type series. Holotype (by original designation) ♂: AUSTRALIA: QUEENSLAND: Emmett Creek, Cromarty [19.47��S, 147.05��E], 6.xii.[19]65, J.G. Brooks & E.P. Brooks [ANIC, examined]. Paratypes (examined). AUSTRALIA: QUEENSLAND: 20♂, same data as holotype [ANIC]; 4♂, Townsville [19.26��S, 146.82��E], Dodd [SAM]; 4♂ on one card, Emmett Creek, Cromarty, about 20 miles S Townsville, 6.xii.1963, J.G. Brooks & E.P. Brooks [ANIC]; 2♂, Cromarty, Townsville, 6.i.1964, J.G. Brooks [ANIC]; 2♂, Brandon [19.55��S, 147.35��E], 13.viii.1968, I.T. Freshwater [ANIC, QM registration number T250928]. Other specimens examined. AUSTRALIA: QUEENSLAND. 1♂, Emmett Creek, Cromarty, 6.xii.1965 J. A. G. Brooks [QDAF registration number 1-004151]; 1♂, Cromarty, 12 km south, J & E B[rooks] [ANIC]; 1♀, Brandon [19.55��S, 147.35��E], 13.viii.1968, I. T. Freshwater [QM]; 18♂ Airville [19.65��S, 147.35��E], 21.i.[19]70, A. Johnson E. Adams [ANIC]. Redescription Male (Figs 11���14). Body 11���14 mm long; head, pronotum, scutellum, and ventral thorax black; elytra and abdominal ventrites dark red-brown; antennae pale yellow-brown, scape sometimes light brown; palpi and tarsi reddish brown; coxae, femora, and tibiae very dark brown to black. Labrum rounded laterally with slight median indentation, not protruding beyond anterior margin of upper clypeus, with a few scattered, short setae. Terminal palpomere of maxillary palp slender, about as long as palpomeres 1���2 together, with dull area on upper surface. Anterior face of clypeus 4x as wide as long, with setiferous punctures along anterior margin, posterior surface smooth; outline of upper surface broadly rounded, without emargination but reflexed on anterior, surface very coarsely punctured, long, yellow-brown setae intermixed with a few, flattened, adpressed, white setae across the posterior margin and scattered on anterior half. Frons coarsely and densely punctured, covered with dense, long yellow-brown setae intermixed with a few short, adpressed, white setae; ocular canthi with dense fringe of long, stout, yellow-brown setae. Antennal club 6-lamellate, all lamellae about the same length; antennomere 4 slightly wedge-shaped anteriorly. Pronotal width across base 2x mid length, sides obtusely angled in middle, anterior angles widely obtuse, posterior angles obtuse and slightly rounded, anterior and poster margins narrowly raised and defined by grooves continuous from side to side except in middle of posterior margin; surface coarsely punctured, covered with long, erect, yellow-brown setae intermixed with short, flattened, white setae varying in size from contained within their punctures to about 2x puncture diameter. Scutellum with a few scattered setae similar to both pronotal types. Elytra with only sutural stria impressed, surface fairly uniformly punctured, punctures smaller than those on pronotum and each containing a small, pale seta; lateral margins with a fringe of very long, stout, light-brown setae. Propygidium and pygidium uniformly clothed with recumbent, parallel-sided, flattened, white setae. Ventral thorax with dense, long, pale-yellow setae. Legs with scattered, pale-yellow setae and a few flattened, white setae. Abdominal ventrites with long, pale-yellow setae intermixed with denser, flattened, white setae. Aedeagus (Figs. 12���14) with lateral edges almost parallel then rounded at apices. Female (Figs. 15���17). Body 13.8 mm long; head, pronotum, scutellum, and ventral thorax dark brown; elytra and abdominal ventrites red-brown; antennae pale yellow-brown, scape light brown; palpi and legs reddish brown, coxae brown. Labrum rounded anteriorly, not protruding beyond anterior margin of upper clypeus, with a few scattered, short setae. Terminal palpomere of maxillary palp slender, about as long as palpomere 1, with dull area on upper surface. Anterior face of clypeus 4x as wide as long, with setiferous punctures across middle, anterior surface smooth, posterior surface smooth with occasional punctures; outline of upper surface broadly rounded, without emargination but slightly reflexed on anterior, surface very coarsely punctured, with long, yellow-brown setae. Frons coarsely and densely punctured, covered with dense, long yellow-brown setae intermixed with a few short, adpressed, white setae; ocular canthi with dense fringe of long, stout, yellow-brown setae. Antennae of 10 antennomeres, club 6-lamellate but club rounded with lamella of antennomere 5 about 0.5x length of antennomere 9 (the longest antennomere of the club). Pronotal width across base 1.9x mid length, sides rounded, anterior angles widely obtuse, posterior angles slightly obtuse and slightly rounded, anterior and poster margins narrowly raised and defined by grooves continuous from side to side except in middle of posterior margin; surface coarsely punctured, covered with long, erect, yellow-brown setae intermixed with short, flattened, white setae varying in size from contained within their punctures to about 2x puncture diameter. Scutellum with a few scattered setae similar to both pronotal types. Elytra with only sutural stria impressed, surface fairly uniformly punctured, punctures smaller than those on pronotum and each containing a small, pale seta; lateral margins with a fringe of very long, stout, lightbrown setae. Propygidium and pygidium uniformly clothed with recumbent, parallel-sided, flattened, white setae. Ventral thorax with dense, long, pale-yellow setae. Legs with scattered, pale-yellow setae and a few flattened, white setae. Abdominal ventrites with long, pale-yellow setae intermixed with denser, flattened, white setae. Distribution. Known from the coastal area between Townsville and Ayr in northeastern Queensland (Fig. 18). All localities have a K��ppen-Geiger climate classification of Aw (savannah). Natural history. Adults of both sexes have been collected in August and December���January. Female antennae (Fig. 17) are consistent (Allsopp 1990) with other Australian old-Gondwanan Melolonthini (Allsopp 1995) where both genders do not feed as adults, and the females fly little (if at all) and attract males with pheromones (Allsopp 1993). This contrasts with more recent Oriental arrivals, such as Lepidiota spp., where male and female antennal clubs are similar, and mating appears to take place at feeding sites., Published as part of Allsopp, Peter G. & Smith, Andrew B. T., 2022, Australian Melolonthini (Coleoptera: Scarabaeidae: Melolonthinae): a third species of Allothnonius Britton, 1978, and notes on the other known species, pp. 475-484 in Zootaxa 5099 (4) on pages 480-482, DOI: 10.11646/zootaxa.5099.4.3, http://zenodo.org/record/6091678, {"references":["Britton, E. B. (1978) A revision of the Australian chafers (Coleoptera: Scarabaeidae: Melolonthinae) Vol. 2. Tribe Melolonthini. Australian Journal of Zoology, Supplementary Series, 60, 1 - 150. https: // doi. org / 10.1071 / AJZS 060","Weir, T. A., Lawrence, J. F., Lemann, C. & Gunter, N. L. (2019) Scarabaeidae: Melolonthinae Leach, 1819. In: Slipinski, A. & Lawrence, J. F. (Eds.), Australian Beetles. Vol. 2. Archostemata, Myxophaga, Adephaga, Polyphaga (part). CSIRO Publishing, Clayton South, pp. 467 - 507.","Allsopp, P. G. (1990) Sexual dimorphism in the adult antennae of Antitrogus parvulus Britton and Lepidiota negatoria Blackburn (Coleoptera: Scarabaeidae: Melolonthinae). Journal of the Australian Entomological Society, 29, 261 - 266. https: // doi. org / 10.1111 / j. 1440 - 6055.1990. tb 00360. x","Allsopp, P. G. (1995) Biogeography of the Australian Dynastinae, Rutelinae, Scarabaeinae, Melolonthini, Scitalini and Geotrupidae (Coleoptera: Scarabaeoidea). Journal of Biogeography, 22, 31 - 48. https: // doi. org / 10.2307 / 2846071","Allsopp, P. G. (1993) Evidence for sex attraction in three species of Australian canegrub beetles (Coleoptera: Scarabaeidae: Melolonthinae). Coleopterists Bulletin, 47, 51 - 52."]}

Liogenys schneiderae Cherman, new species Figs. 8, 9. Type material. Holotype male, labeled: [green, handwritten] “4-dens Fbr / 4-dentata / kl. / Pg.[?] / Col. [Colombia]”, [white, typeset and handwritten] “MLU Halle / WB Zoologie / S. –Nr. 8/3/11”, [red typeset and handwritten] “ LIOGENYS / SCHNEIDERAE / HOLOTYPE / Cherman M. A.” (MLUH). Paratypes (81), all bearing the label [yellow, typeset and handwritten] “ LIOGENYS / SCHNEIDERAE / PARATYPE / Cherman M. A.”. One female with the labels: [white handwritten] “ Venezuela ”, [white typeset and handwritten] “ Liogenys quadri- / dens / F det. G. Frey, 1968” (ZMHB). One female with the label: [green typeset] “ Venezuela / F. Kummerow S.” (ZMHB). Two females and one male with the following label: [white handwritten] “ Guyana ” (ZMHB). Two females and two males with the label: [green, outlined, typeset] “Hist. Coll. (Coleoptera) / Nr. 24131 / Liogenys quadridens Burm. / Columb., Moritz. / Zool. Mus. Berlin” (ZMHB). One male with the labels: [white typeset] “24131”, [dark green handwritten] “ Amphicrania / 4 dentata Nr / Columb. Moritz”, [white handwritten] “quadridens / Burm / Melol. 4 dens Fab.” (ZMHB). Two males bearing the labels: [white handwritten] “ Colombia / Mus. Ber. / 4-dentata / Kl.”, [white typeset] “Mus. Western” (ZMUC). One male bearing: [white typeset] “ Coll. Kraatz ”, [white typeset] “ Moser det”, [white typeset] “Dtsch. Ent. Inst. / Eberswalde”, [white outlined green, handwritten] “quadridens. / Guyana /F.”, [white handwritten] “ Liogenys. /Guerin.”, [white handwritten] “ Liogenys / quadridens Fab ”, [white, typeset and handwritten] “ Liogenys quadri- / dens / F det. G. Frey, 1968”, [white handwritten] “ Liogenys /quadridens / F.”, [light green typeset] “DEI Münchenberg / Col – 03566” (SDEI). One male with the following data: [white typeset] “CARIACO / (Edo. SUCRE)”, [white typeset] “ Bordon leg.” / [handwritten] “10-IV-59” [white handwritten] “4”, [white, outlined, typeset] “H. & A. HOWDEN/COLLECTION / ex. A. Martinez coll.” (CMNC). One female with the data: [white typeset] “QUIAMARE[Libertad] / (BARCELONA)”, [white typeset] “5-4-58/BORDON LEG.”, [white, outlined, typeset] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.” (CMNC). Two females with the data: [white, typeset] “ Aragua / Venezuela / V.1978 ” (USNM). One male with the data: [white, typeset] “Chaguarramas / (Edo. Guarico)”, [white typeset] “13-3-60 / BORDON LEG.”, [white, handwritten and typeset] “ Liogenys quadridens ♂ / (Fab.) / A MARTINEZ-DET. 1974”, [white with black border, typeset] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.”, genitalia mounted (CMNC). Two males and four females with the data: [white, typeset] “West Indies: / Tobago: / Scarborough. / 16. vi. 1914. / W.E. Broadway. ” and “Brit. Mus. / 1920–487.” (1 male and 2 females at BMNH; 1 male and 2 females at CMNC). One male with the data: [white, typeset] “West Indies: / Tobago: / Scarborough. / 17. vi. 1914. / W.E. Broadway. ” and “Brit. Mus. / 1920–487.” (BMNH). One female with the data: [white, typeset] “West Indies: / Tobago: / Scarborough. / 10. vi. 1914. / W.E. Broadway. ” and “Brit. Mus. / 1920–487.” (BMNH). One female with the data: [white, typeset] “West Indies: / Tobago: / Scarborough. / 12. vi. 1914. / W.E. Broadway. ” and “Brit. Mus. / 1920–487.” (BMNH). Two females with the data: [white, typeset] “West Indies: / Tobago: / Scarborough. / 15. vi. 1914. / W.E. Broadway. ” and “Brit. Mus. / 1920–487.” (BMNH). Two females with the data: [white, typeset] “West Indies: / Tobago: / 9. vi. 1914. / W.E. Broadway. ” and “Brit. Mus. / 1920–487.” (BMNH). One male with the data: [white, handwritten] “quadridens / Fabr. / 4-dentata, / Klug. / Columbia”, [white, typeset] “So named / in Reiches / Collection. / C.W.”, [green, handwritten] “ ♂ ”, [white, handwritten] “nov. sp.”, [light blue, handwritten] “2258.”, [white, typeset] “67.45” (BMNH). Sixteen males and 15 females with the data: [white, typeset] “ VENEZUELA: Guarico, Hato / Masaguaral, 45kmS Calabozo / 8.57N, 67.58W, GalryForest / #28, 75m, 3-5 June 1988 / uv light, M. Epstein ” (eight males and eight females at USNM, four males and three females at CMNC, two males and two females at DZUP, two males and two females at CEMT). Paratypes of DZUP with the sequence of voucher numbers: [white typeset] “ DZUP / 402753 ” to “ DZUP / 402756 ”. One male and eight females with the data: [white, typeset] “ VENEZUELA: Guarico, Hato / Flores Morades, 45 km S / Calabozo, 8.57N, 67.58W / Galry Forest #2, 75 m / 24-26June 1989, uv light / M. Epstein & M. Rodiguez ” (USNM). One male and two females with the data: [white, typeset and handwritten] “ex? / intercepted / at Miami Fla / 1 May 1964 ” (USNM). One male and two females with the data: [white, typeset and handwritten] “ VENEZUELA: BOLIVAR / Guri, 16.VI.1996 / H.&A. Howden Lt / Wet forest ” (CMNC). One female with the data: [white, typeset and handwritten] “ VENEZUELA: GUARICO / 12km W Valle de la / Pascua, 21-22.VI.96 Lt / H.&A. Howden ” (CMNC). One female with the data: [white, typeset] “VEN: Bolivar, 20km / SW Ciudad Bolivar / 19. VI. 1987, S&JPeck / woodland on sand, UV” (CMNC). One female with the data: [white, typeset] “VENEZ: Anzoategui / 18 km E Pariaguan / 24JUN 1987, 240m / M.A. Ivie, beating” (CMNC). Holotype deposited at MLUH. Eleven paratypes at BMNH. Nineteen paratypes deposited at CMNC. Four paratypes at DZUP, four paratypes at CEMT. One paratype deposited at SDEI. Thirty paratypes deposited at USNM. Ten paratypes deposited at ZMHB. Two paratypes deposited at ZMUC. Additional material. One male (BMNH) missing the head and pronotum with the data: [white, typeset and handwritten] “Scarborough, / Tobago. / 15 May 1914. / W.E. Broadway. / 1914-403”. Diagnosis. Body brown, elongate, sides almost parallel in males (Fig. 8A), slightly wider on posterior third in females; elytra brown; pronotum reddish to purplish brown; clypeus quadridentate, clypeal emargination deep, rounded, and narrow; distance between lateral and anterior clypeal teeth slightly longer than basal width of one anterior tooth; right angle between lateral and anterior clypeal teeth; pronotal corners rounded (Fig. 8B), appearing obsolete especially in females; mesotibia quadrate in cross section (Fig. 8C); pygidium weakly convex; pygidial disc glabrous, with scattered bristles on apex (Fig. 8D); ventrites II, III, and sometimes IV undulate on midline, especially ventrite II (Fig. 8E); in males protarsomere II up to twice the length of protarsomere I in dorsal view; metatibia subbasally produced towards apex; basal region of parameres strongly narrowed, parameres more than five times the length of their apex; apex harpoon-shaped, lateral spine short, projected almost perpendicular to parameres (Fig. 8F). Holotype. Male. Length: 9.4 mm, width: 4.4 mm. Brown. Head: distance between eyes nearly twice the width of one eye; frons longer than clypeus; clypeus rugose, clypeal emargination deep, rounded, and narrow; outer sides of anterior teeth parallel; outer margin of anterior teeth shorter than the eye in dorsal view; clypeus convex laterally and strongly produced forming a conspicuous tooth-like projection; distance between lateral and anterior teeth slightly longer than basal width of one anterior tooth, distance between lateral tooth and anterior margin of eye longer than one eye length, angle between anterior and lateral teeth approximately 90º; canthus not exceeding the outer margin of the eye; distal maxillary palpomere maximum width slightly wider than at apex; fovea shallow, extending past the transverse midline of the palpomere; antenna with 10 antennomeres, club lighter in color and as long as the funicle. Thorax: pronotal anterior margin slightly produced medially, undulate and flanged (Fig. 8B); maximum length of pronotum exceeding the length of tarsomeres I–III combined; disc glabrous, punctures very sparse and coarse, denser towards anterior margin; pronotal posterior corners rounded, appearing obsolete; hypomere with long bristles and thin scales; mesepisternum scaly; anterior portion and sides of metaventrite with bristles; distance between mesocoxae and metacoxae nearly twice the length of the metacoxa; scutellum triangular, few, scattered, fine punctures at the sides. Elytra: shiny, uniform light brown, lighter in color than the pronotum; elytra more than three times longer than the pronotum; elytral suture darker than the elytron and slightly elevated; all four elytral ridges weakly defined. Legs: procoxa scaly, thin scales on outer surface; three protibial teeth, middle and apical teeth equal in size, distance between basal and middle teeth longer than between middle and apical teeth; protibial spur present; mesofemur with a row of long bristles on anterior and posterior margins, mesotibia subquadrate in cross section, surface finely sculptured, mesotibia with two transverse carinae, the apical carina incomplete; metacoxa with bristles throughout, basal articular lobe of metacoxa produced beyond the outer margin of trochanter; inner margin of metatibia carinate and abruptly subbasally produced towards apex, apical inner surface glabrous, metatibial surface coarsely sculptured on outer margin; metatibial transverse carina present posteriorly; protarsomere II very elongate; protarsomeres and mesotarsomeres I–IV enlarged, protarsomeres equal in width to the mesotarsomeres, slightly wider than the metatarsomeres; metatarsomere I shorter than metatarsomere II; claw bifid, outer tooth of a claw longer and narrower than the inner tooth; distance between teeth shorter than the inner tooth. Abdomen: disc of ventrites with sparse bristles; ventrites II–III and sometimes IV with tubercles on disc (Fig. 8E); propygidium with bristles, pygidium slightly convex, subtrapezoidal, wide; pygidial width exceeding distance between spiracles of propygidium; pygidial disc glabrous, few bristles on apex, pygidial disc finely punctate; pygidial apex quadrate. Parameres: basal region dorsally narrowed, narrower than both sections of the parameres at its midline; parameral split at 2/3 the length of the basal region; parameres more than five times the length of their apex; inner margins opened, convergent; apex harpoon-shaped, lateral spine thin, short, projected almost perpendicular to parameres (approximately 80º angle). Parameres concave in lateral view (Fig. 8G). Variation. Female length: 9.2–10.2 mm, width 4.5–5.3 mm. females as the holotype except the clypeal anterior teeth wider; canthus exceeding the outer margin of the eye; pronotal corners more marked; metatibia coarsely sculptured, metatibial apical carina complete; ventrite tubercles more noticeable and pygidium wider, more trapezoidal. Male length: 9.0– 9.5 mm, width 4.4–4.8 mm. Males as the holotype except the head and pronotal punctures coarser; pronotal corners more marked, and pygidium flatter and bent downwards. Etymology. The species is dedicated to Dr. Karla Schneider, biologist and curator of the Zoological Collection of the Martin-Luther-Universität, Halle-Wittenberg, Halle, Germany. Karla made the discovery of this new species possible by loaning the holotype from the MLUH to the ZHMB. This allowed its thorough examination during the visit of M.A.C. to these collections. This name should be treated as a noun in the genitive case. Type locality: Colombia. Geographical distribution. TRINIDAD AND TOBAGO (Scarborough), COLOMBIA, GUYANA, VENEZUELA (Anzoátegui, Bolívar, Guaricó, Sucre). Remarks. Liogenys schneiderae Cherman, new species resembles L. quadridentata mainly in the body shape and size, color of the body and elytra, and in the quadridentate clypeus. However, L. schneiderae differs (L. quadridentata in parenthesis) in the clypeus being more rugose, with the impressions next to the frons more marked (not rugose); antenna with club shorter; anterior margin of pronotum slightly depressed throughout (not as so); elytral suture medially darker than the elytron (medially unicolored with elytron); metatibia flatter with inner margin abruptly subbasally produced towards apex (produced straight towards apex); ventrites I–IV with tubercles (without tubercles); and pygidium never inflated, with apex quadrate (pygidium convex in both sexes, inflated in females, pygidial apex rounded)., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on pages 221-224, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886

Liogenys genieri Smith & Cherman, new species Figs. 3, 9. Type material. Holotype male, labeled: [white, typeset] “ VENEZUELA: Guarico, Hato / Flores Morades, 45 km S / Calabozo, 8.57N, 67.58W / Galry Forest #2, 75 m / 24–26June 1989, uv light / M. Epstein & M. Rodiguez ”, [red, typeset and handwritten] “ LIOGENYS / GENIERI / SMITH & CHERMAN / HOLOTYPE ”, genitalia mounted (USNM). Paratypes (8), all bearing the label [yellow, typeset and handwritten] “ LIOGENYS / GENIERI / SMITH & CHERMAN / PARATYPE ”: One male and six females with the same data as the holotype (one male and one female at CMNC, two females at DZUP, three females at USNM). One male with the data: [white, typeset] “ VENEZUELA: Guarico / Hato Masaguaral / (44km S Calabozo) / May 3–10 1985 / Menke & Carpenter” (USNM). Each female paratype of DZUP with the voucher: [white typeset] “ DZUP / 402748 ” and “ DZUP / 402749 ”. Holotype and four paratypes deposited at USNM. Two paratypes at CMNC. Two paratypes at DZUP. Diagnosis. Body light brown, elongate, sides almost parallel; elytra light brown; pronotum reddish brown (Fig. 3A); clypeus quadridentate; clypeal emargination deep, angulate, and narrow; distance between lateral and anterior clypeal teeth approximately equal to basal width of one anterior tooth; angle between lateral and anterior clypeal teeth approximately 90º; pronotal corners rounded (Fig. 3B); mesotibia subquadrate in cross section; pygidium slightly convex; pygidial disc glabrous, with scattered bristles on apex (Fig. 3D); in males protarsomere II more than twice the length of protarsomere I in dorsal view (Fig. 3C); parameres more than six times the length of their apex; lateral expansion along the parameres only seen subapically; inner margins convergent; apex expanded with backward-facing spine weakly projected, spine length less than one-third length of apex (Fig. 3E). Holotype. Male. Length: 9.6 mm, width: 4.6 mm. Light brown. Head: distance between eyes nearly twice the width of one eye; frons equal in length to clypeus; clypeal emargination deep, angulate, and wide; outer sides of anterior teeth parallel; outer margin of anterior teeth shorter than the eye in dorsal view; clypeus convex laterally and strongly produced forming a conspicuous tooth-like projection; distance between lateral and anterior teeth approximately equal to basal width of one anterior tooth, distance between lateral tooth and anterior margin of eye approximately equal to one eye length, angle between anterior and lateral teeth approximately 90º; canthus not exceeding the outer margin of the eye; distal maxillary palpomere, maximum width twice the width at apex; antenna with 10 antennomeres, club lighter in color and longer than funicle. Thorax: pronotal anterior margin slightly produced medially, undulate, and flanged; disc glabrous, punctures moderately dense and coarse; anterolateral margin reflexed; pronotal posterior corners rounded; hypomere with long bristles; mesepisternum scaly; anterior portion and sides of metaventrite with bristles; distance between mesocoxae and metacoxae nearly twice the length of the metacoxa; scutellum triangular with sparse, coarse punctures. Elytra: shiny, uniform light brown, lighter in color than the pronotum; elytra near three times the length of pronotum; elytral suture darker than the elytron and weakly elevated; all four elytral ridges weakly defined. Legs: procoxa with abundant, thin and long scales, short scales on outer surface; three protibial teeth, middle and apical teeth equal in size, distance between basal and middle teeth longer than between middle and apical teeth; protibial spur present; mesofemur with a row of long bristles on anterior and posterior margins, mesotibia subquadrate in cross section, surface coarsely sculptured on outer margin, mesotibia with two transverse carinae, the apical carina incomplete; metacoxa with bristles throughout, basal articular lobe of metacoxa produced beyond the outer margin of trochanter; inner margin of metatibia straight, carinate, and produced on apex, apical inner surface with bristles, metatibial surface coarsely sculptured; metatibial transverse carina weakly defined, only present posteriorly; protarsomere II elongate; protarsomeres and mesotarsomeres I–IV enlarged, protarsomeres slightly wider than the mesotarsomeres and two times wider than the metatarsomeres; metatarsomere I slightly shorter than metatarsomere II; claw bifid, symmetrical, outer tooth of a claw longer and as wide as the inner tooth; distance between teeth shorter than the inner tooth. Abdomen: disc of ventrites with bristles; propygidium with bristles, pygidium weakly convex, subquadrate, wide; pygidial width approximately equal to distance between spiracles of propygidium; pygidial disc glabrous except for a few bristles on apex, pygidial disc coarsely punctate; pygidial apex oval. Parameres: basal region dorsally narrowed, equal to both sections of the parameres at its midline; parameral split at 2/3 the length of the basal region; inner margins slightly convergent; parameres more than six times the length of their apex; apex harpoon shaped; laterally expanded with backward-facing spine weakly projected straight downward, spine length less than one-third length of apex. Parameres in lateral view straight, not coplanar (Fig. 3F). Variation. Female length: 10.2–11.2 mm, width 5.0– 5.6 mm. Females as the holotype except in the size; pronotum wider, with anterolateral margin distinctly reflexed; maximum width of distal maxillary palpomere slightly wider than at apex. Male length: 9.4–9.6 mm, width 4.0– 4.8 mm. Males as the holotype except in the punctures of pronotum slightly less dense; protibial teeth sharper and longer (worn in holotype). Etymology. This species is named after François Génier, scarab taxonomist from the CMNC, in appreciation for his help in photographing specimens for this paper. Type locality. Hato Masaguaral, Guárico, Venezuela (8.57ºN, 67.58ºW). Geographical distribution. VENEZUELA (Guárico). Remarks. Liogenys genieri Smith & Cherman, new species resembles L. clipeosetosa in the body color and size, shape of the quadridentate clypeus, head and pronotum coarsely punctate, anterior teeth of clypeus long, anterior and lateral teeth forming approximately 90º angle, and posterior corners of pronotum rounded. Males in both species have parameres very long, more than five times the apical length, and apex laterally weakly projected. Liogenys genieri differs (L. clipeosetosa in parenthesis) in the frontoclypeal surface without bristles (with bristles); clypeal emargination angulate, narrower (rounded, wider); pronotum with anterolateral margin reflexed (not as so); apex of pygidium oval (quadrate); males with parameres slightly shorter, lateral expansion along the parameres not visible dorsally and straight in lateral view (lateral expansion entirely visible dorsally)., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on pages 207-209, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886

Compsodactylus mendax Fuhrmann & Smith, new species (Figs. 1���8, 15���16, 19���26) Type material. Male holotype (FSCA, Figs. 1���4), allotype (FSCA, Figs; 6���8), and male paratype (CMNC) all labeled (Fig. 5): ��� PERU: Dept. Lima / Sta. Eulalia 1035m / 30���31 Mar 2011 / J.B. Heppner & C. Carrera ���. These specimens also have our corresponding red holotype and allotype labels and yellow paratype label. Diagnosis. Length greater than 8.0 mm; dorsal surface with metallic green reflections; antennae with 10 antennomers; clypeus elongate with expanded apex in males (Fig. 4); clypeus with short, spiny setae along apex and lateral edges in males; apex of protarsomere I 2 times wider than apex of protarsomere II in males; elytral striae distinctly punctate; male metatibial apex with an inner truncate, twisted process (Fig. 16); genitalia of females with gonostyle, and gonocoxite about 2.5 times longer than wide (Figs. 23���25). Holotype description. Length 9.0 mm, width 4.0 mm at the middle of elytra. Dorsal and ventral surface of head and thorax (except lateral edges of pronotum) black with metallic green reflections (Figs. 1���4); dorsal surface of elytra and lateral edges of pronotum yellowish brown with light metallic green reflections; ventral surface of abdomen shiny reddish brown, ventrite VI with anterior dark stripe; legs bicolored, reddish brown with dark sections and markings. Head with clypeus elongate, strongly expanded at apex (Fig. 4), surface glabrous medially and with short, spiny setae along apex and lateral edges. Frons surface with rugose punctures and densely setose, setae long, thin. Interocular distance 5 times wider than dorsal eye width. Antenna with 10 antennomeres, club longer than antennomeres II��� VI combined. Pronotum surface rugose with long, dense setae, anterior margin with some minute, spiny setae. Elytra surface with moderately dense setae, striae distinctly punctate. Pygidium and ventral surface with dense punctures and hair-like setae. Protibia with 2 outer teeth, protarsomere I greatly enlarged with a padlike granulate ventral surface. Metafemur not strongly enlarged (about 2.1���2.4 times longer than wide, Fig. 15), inner margin without spines or carina and with some spiny setae. Metatibial apex with an inner, twisted process; metatibial spurs absent (Fig. 16). Variation. The allotype is 8.5 mm in length with the following sexually dimorphic characters: clypeus not elongate, without glabrous area, and with apex subquadrate, not expanded; frons with short, spiny setae; protarsomere I cylindrical, not enlarged; metatibial apex without inner process and with 2 spurs. Female terminalia bearing a pair of elongate paraprocts; gonocoxite about 2.5 longer than wide and divided in a proximal and a distal piece; a lateral plate between paraproct and proximal piece of gonocoxite is present, gonostyle present (Figs. 23���25). The elytral suture is yellow in holotype and black with metallic green reflections in the allotype and paratype. Distribution. The holotype, allotype, and a single paratype were collected in Santa Eulalia, Lima Department, Peru (approximately 11.902��S, 76.666��W) at 1035 m. Etymology. From the Latin mendax meaning liar, in reference to the elongate clypeus being reminiscent of Pinocchio���s nose. This name is a noun in apposition. Remarks. Compsodactylus mendax new species is easily distinguished from the other four species in the genus because the clypeal apex is enlarged in males while the other species have a trapezoidal clypeus (Fig. 4); the dorsal surface has green metallic reflections and antennae have 10 antennomeres in females, a character combination not found in other species. Compsodactylus mendax is similar to C. martinezi (Frey, 1972) and C. vallejoi Figueroa & Neita-Moreno, 2019 in that all three species have the apical spine of metatibia thick and projecting perpendicular to length of the tibiae in males (Figs. 13���18) and females have gonostyles. Compsodactylus argentinus and C. parvulus differ from these three species in that the apical spine of metatibia is thin and projecting parallel to the length of the tibia in males (Figs. 9���12) and females do not have gonostyles. Compsodactylus mendax is distinguished from C. martinezi and C. vallejoi by (characters of 1��� C. martinezi and 2��� C. vallejoi given in brackets): dorsal surface with metallic green reflections (1���as C. mendax; 2���without green reflections); antennae with 10 antennomeres (1���nine antennomeres; 2���as C. mendax: Jhon C. Neita Moreno, personal communication); male protarsomere I greatly enlarged (1���slightly enlarged; 2���as C. mendax); male metafemur slightly enlarged and approximately 2.1���2.4 times longer than wide (1 and 2���greatly enlarged and approximately 1.5���1.7 times longer than wide; Figs. 13, 15, 17); male metatarsus somewhat filiform (1���somewhat moniliform; 2���as C. mendax); male pygidium with hair-like setae (1���with scale-like setae; 2���as C. mendax); females with long gonocoxite (Figs. 23���25; 1 ���wide; 2���comma-like)., Published as part of Fuhrmann, Juares & Smith, Andrew B. T., 2021, Compsodactylus Fuhrmann, 2012 (Coleoptera: Scarabaeidae: Melolonthinae) distributional notes and a new species, pp. 387-393 in Zootaxa 4990 (2) on pages 388-391, DOI: 10.11646/zootaxa.4990.2.12, http://zenodo.org/record/5026053, {"references":["Figueroa, L. & Neita-Moreno, J. C. (2019) A new Compsodactylus Fuhrmann (Coleoptera: Scarabaeidae: Melolonthinae) from Peru. Zootaxa, 4560 (3), 587 - 591. https: // doi. org / 10.11646 / zootaxa. 4560.3.12"]}

Liogenys quadridens (Fabricius, 1798) Figs. 6, 9. Melolontha quadridens Fabricius, 1798: 131 (original description). Liogenys quadridens: Burmeister 1855: 14 (redescription); Harold 1869: 1140 (checklist); Bates 1887:155 (checklist); Dalla Torre 1913: 318 (checklist); Blackwelder 1944: 228 (checklist); Frey 1969: 50 (key); Evans 2003: 213 (checklist); Evans & Smith 2009: 181–182 (checklist); Krajčík 2012: 145 (checklist); Cherman et al. 2017: 11, 16; 2019: 62 (taxonomy). Type material examined. Melolontha quadridens female lectotype, present designation (ZMUC): [white handwritten] “4 Dens.”, [white typeset] “zmuc / 00022369”, [white, outlined in red, typeset and handwritten] “ LIOGENYS / QUADRIDENS / (Fabricius, 1798) / LECTOTYPE / Des. Cherman M. A.”. Additional material examined. PANAMA. Coclé: Río Hato, 22.V.1960, light trap, V.J. Tipton, 1 male, 1 female (CNCI). Panamá: Cerro Campana, 820 m, 8º40`N, 79º56`W, 22.V.1976, at blacklight, B.C. Ratcliffe, 2 males, 4 females (UNSM); Cerro Campana, 2600 feet, 10–20.V.1981, J.E. Wappes, 1 male, 1 female (UNSM). Panamá Oeste: Nueva Gorgona, 25.V.1962, R.&F. Zweifel, 1 male, 1 female (AMNH); Sajalices, 17.V.1982, E. Giesbert, 1 male, 3 females (UNSM). Veraguas: Santiago, 18.V.1977, H. Howden, 1 male (CMNC). VENEZUELA. Guárico: San Juan de Los Morros, 28.IV.1938, C.H. Ballou, 1 female (USNM); Monagas: El Tejero, 200 m, 2.V.1964, C. Bordon, 1 male (CMNC). Distrito Capital: Caracas, 1938, G. Vivas-R., 2 males (USNM). Without locality and date, E. Kummerow S., 1 female (ZMHB). Diagnosis. Body light brown, elongate, males with sides parallel (Fig. 6A), females with sides slightly wider on posterior third (Fig. 6B); elytra light brown to yellowish brown, pronotum reddish to purplish brown, head even darker; clypeus quadridentate; clypeal emargination deep, rounded, and wide (Fig. 1A), slightly narrower in females than in males (Fig. 1B); distance between lateral and anterior clypeal teeth longer than basal width of one anterior tooth; angle between lateral and anterior clypeal teeth approximately 90º; mesotibia subquadrate in cross section; pygidium flat or slightly convex, pygidial disc glabrous, with scattered bristles on apex (Fig. 6D); males with protarsomere II less than twice the length of protarsomere I in dorsal view; parameres elongate, up to three times the length of their apex (Fig. 6H), inner margins straight, lateral expansion on outer margins along the parameres partially hidden in dorsal view (Fig. 1E), apex harpoon-shaped, lateral spine projected backwards, slightly convergent to the outer margin of the parameres; spine length slightly more than twice the length of the apex, subapical and proximal part of the apex elevated medially (Fig. 6), abruptly flattened distally at apex and spine (Fig. 6G). Redescription. Length: 9.2–10.8 mm, width: 4.6–5.0 mm. Light brown to brown. Head: dark, distance between eyes nearly twice the width of one eye; frons equal in length to clypeus; clypeal emargination deep, rounded, and narrow; outer sides of anterior teeth parallel; outer margin of anterior teeth shorter than the eye in dorsal view; clypeus convex laterally and strongly produced forming a conspicuous, tooth-like projection; distance between lateral and anterior teeth longer than basal width of one anterior tooth, distance between lateral tooth and anterior margin of eye longer than one eye length, angle between anterior and lateral teeth approximately 90º; canthus exceeding or not the outer margin of the eye; distal maxillary palpomere maximum width twice the width at apex; fovea deep, extending past the transverse midline of the palpomere; antenna with 10 antennomeres, club lighter in color, as long as the funicle in females, longer than the funicle in males. Thorax: pronotal anterior margin slightly produced medially, undulate; disc glabrous, punctures sparse and fine in females, coarser in males; pronotal posterior corners rounded in males, apparently obsolete in females; hypomere with long bristles; mesepisternum scaly; anterior portion and sides of metaventrite with bristles; distance between mesocoxae and metacoxae nearly twice the length of the metacoxa; scutellum triangular, punctures mainly at the sides. Elytra: shiny, uniform yellowish brown to light brown, lighter in color than pronotum; elytra near three times longer than the pronotum; elytral suture slightly darker than elytron and not elevated; all four elytral ridges weakly defined. Legs: procoxa with long bristles on infra-carinal surface and fine scales and short bristles on outer surface; three protibial teeth, middle and apical equal in size, distance between basal and middle teeth longer than between middle and apical; protibial spur present; mesofemur with a row of long bristles on anterior and posterior margins, mesotibia quadrate in cross section, surface coarsely sculptured in females, finely sculptured in males, mesotibia with two transverse carinae, the apical carina complete in females, incomplete in males; metacoxa with bristles throughout and forming a row of scales parallel to the anterior margin (Fig. 6F); basal articular lobe of metacoxa produced beyond the outer margin of trochanter; inner margin of male metatibia carinate and produced on apex, apical inner surface with scattered bristles, metatibial disc coarsely sculptured; metatibial transverse carina and discontinuous longitudinal carina present posteriorly; metatarsomere I and metatarsomere II equal in length; males with protarsomere II as wide as it is long; protarsomeres and mesotarsomeres I–IV equally enlarged, wider than metatarsi; claw bifid, symmetrical, outer tooth of a claw longer and narrower than inner tooth; distance between teeth shorter than the inner tooth. Abdomen: disc of ventrites with sparse bristles; propygidium with bristles, pygidium flat, subtrapezoidal, wide, pygidial width exceeding distance between spiracles of propygidium; pygidial disc glabrous, few bristles on apex, coarsely punctate, roughly in males; slightly depressed medially in males; pygidial apex quadrate. Parameres: basal region slightly wider than both sections of the parameres at its midline; parameral split at 2/3 the length of the basal region; parameres almost three times the length of their apex; inner margins straight; lateral expansion of the outer margins partially hidden, visible from the midline up to the subapical portion; apex harpoon-shaped, lateral spine straight backwards, slightly bent, spine length more than twice the length of the apex, subapical and proximal part of the apex elevated medially, flattened distally at apex and spine (Figs. 1E, 6G). Parameres concave in lateral view (Fig. 6E). Type locality. “ India Orientali” [Burmeister 1855: 14, misinterpreted locality of West Indies]. Species from Panama and Venezuela. Geographic distribution. PANAMA (Coclé, Panamá, Panamá Oeste, Veraguas), VENEZUELA (Distrito Capital, Monagas). Remarks. Liogenys quadridens was redescribed for the first time by Burmeister (1855), who transferred it from Melolontha Fabricius, 1775 to Liogenys. Fabricius (1798) mentioned in the original description that the species is from “ India Orientali” [East Indies], we interpreted that he meant West Indies instead. According to Burmeister (1855) and Harold (1869), L. quadridens occurs in Colombia and Guyana. Bates (1887) stated that this species occurs in “Amazons” but this record should be disregarded as it was surely based on the misidentification of other species of Liogenys. Bates (1887) also cited Panama, “Volcán de Chiriqui 2500 to 4000 feet (Champion)” but these specimens are here identified as L. quadridentata (based on our examination of these specimens in the BMNH). Frey (1969) cited the distributional limits of this species, from Florida (United States of America) (presumably based on port interception specimens) to Pernambuco (Brazil). In his key, Frey (1969) mentioned an intraspecific variation in the pronotal punctures and in the shape of the parameres, which he drew. At NHMB, we found the specimen from Pernambuco and it is here identified as L. piauiensis Cherman, 2017. Therefore, we refute the occurrence of L. quadridens in Brazil. Among the 41 specimens studied from seven collections bearing L. quadridens identification, we found one male from Nueva Gorgona (Panama) matching with the parameres drawn by Frey (1969), and a female of the same collection event which matches with L. quadridens lectotype. Liogenys quadridens has been cited by the United States Department of Agriculture (1969, 1971, 1974) in its list of intercepted plant pests, carried by aircraft, in unknown hosts. The first time cited (1967–1968) it was being carried from Panama, and the following (1971–1972) probably from Venezuela (United States Department of Agriculture 1969, 1974). This species was also cited as a tomato plant host from Maracaibo, Venezuela (Morales- Valles et al. 2003), and there exists reports of this species in imported ripe tomato fruit throughout the European and Mediterranean Plant Protection Organization (EPPO) region. Nonetheless, as it is not likely to be transported in tomatoes it is classified non dangerous (Grousset et al. 2015). In a study on scarab beetles abundance of a tropical dry forest in Colombia, García-Atencia et al. (2015) found that L. quadridens was the most abundant species (1,667 individuals), characterized as an important pest of semestral crops in the Colombian Caribbean. After studying several specimens of the Caribbean region we cannot confirm that all those specimens mentioned on the previous works are L. quadridens. Although we do believe that L. quadridens occurs in Colombia (based on the known distribution in Panama and Venezuela), we did not examine any specimens to confirm this country record., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on pages 214-215, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886, {"references":["Fabricius, J. C. (1798) Supplementum entomologiae systematicae. Proft et Storch, Copenhagen, 572 pp.","Burmeister, H. (1855) Handbuch der Entomologie (Coleoptera Lamellicornia Phyllophaga chaenochela). Vierter Band. Zweite Abtheilung. Theod. Chr. Fr. Enslin, Berlin, 570 pp.","Harold, E. F. von (1869) Scarabaeidae. In: Gemminger, M. & von Harold, E. F. (Eds.), Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus. Vol. 4. E. H. Gummi, Munich, pp. 976 - 1346.","Bates, H. W. (1887) Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. In: Salvin, O. & Godman, F. du C. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 25 - 160.","Dalla Torre, K. W. von (1913) Scarabaeidae: Melolonthinae IV. Pars 50. In: Schenkling, S. (Ed.), Coleopterorum catalogus. Vol. XX. Junk, W., Berlin, pp. 291 - 450.","Blackwelder, R. E. (1944) Checklist of the Coleopterous Insects of Mexico, Central America, The West Indies and South America. Part 2. Smithsonian Institution, Washington, D. C., pp. 189 - 265. https: // doi. org / 10.5479 / si. 03629236.185.2","Frey, G. (1969) Bestimmungstabelle und revision der gattung Liogenys Guerrez (Coleoptera - Melolonthinae - Macrodactylini). Entomologische Arbeiten aus dem Museum G. Frey, 20, 38 - 64.","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211, 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3 2009. Papers in Entomology, 1, 1 - 344.","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031","United States Department of Agriculture [USDA] (1969) List of intercepted plant pests, 1968 (Pest recorded from July 1, 1967, through June 30, 1968). Hayattsville, Maryland. U. S. G. P. O., Washington, D. C., pp. 46 - 58.","United States Department of Agriculture [USDA] (1971) Insects not known to occur in the United States: A Scarab (Liogenys macropelma Bates). Cooperative Economic Insect Report, 21, 335 - 336.","United States Department of Agriculture [USDA] (1974) List of intercepted plant pests, 1972 (Pest recorded from July 1, 1971, through June 30, 1972). Hayattsville, Maryland. U. S. G. P. O., Washington, D. C., pp. 55 - 68.","Grousset, F., Suffert, M. & Petter, F. (2015) EPPO Study on pest risks associated with the import of tomato fruit. EPPO Bulletin, 45, 153 - 156. https: // doi. org / 10.1111 / epp. 12180","Garcia-Atencia, S. P., Martinez-Hernandez, N. & Pardo-Locarno, L. C. (2015) Escarabajos fitofagos (Coleoptera: Scarabaeidae) en un fragmento de bosque seco tropical del departamento del Atlantico, Colombia. Revista Mexicana de Biodiversidad, 86, 754 - 763. https: // doi. org / 10.1016 / j. rmb. 2015.07.009"]}

Liogenys macropelma Bates, 1887 Figs. 5, 9. Liogenys macropelma Bates, 1887: 155 (original description); Nonfried 1892: 268 (checklist); Dalla Torre 1913: 318 (checklist); Blackwelder 1944: 227 (checklist); Frey 1969: 42 (key); United States Department of Agriculture 1971: 335; 1974: 28 (natural history); Evans 2003: 210 (checklist); Evans & Smith 2009: 179 (checklist); Krajčík 2012: 145 (checklist); Cherman et al. 2017: 14, 16, 69, 86 (taxonomy); 2019: 18, 64 (taxonomy), 53–55 (redescription). Liogenys gebieni Moser, 1921: 140 (original description); Blackwelder 1944: 227 (checklist); Frey 1969: 41 (key); Evans 2003: 209 (checklist); Evans & Smith 2009: 178 (checklist); Krajčík 2012: 145 (checklist). New junior subjective synonym. Type material examined. Liogenys macropelma male lectotype (MNHN): [white typeset] “V. de Chiriqui, / 25–4000 ft. / Champion.”, [white handwritten] “ Liogenys / macropelma / Bates ♂ ”, [red typeset] “ SYNTYPE ”, [white handwritten] “micropygus”, [white, outlined in red, typeset] “ LECTOTYPE / Liogenys macropelma / Bates, 1887 / Cherman M. A. 2015”. Genitalia mounted. Six males with the same labels as the lectotype, plus the label: [white, outlined in red, typeset] “ PARALECTOTYPE / Liogenys macropelma / Bates, 1887 / Cherman M. A. 2015”. One female with one different label: [white handwritten] “ Liogenys / macropelma / Bates ♀ ”. Six male and one female paralectotypes (BMNH): [white, typeset] “V. de Chiriqui, / 25–4000 ft. / Champion.”, [white, typeset and handwritten] “B.C.A.Col.II. (2) / Liogenys / macropelma”. One male paralectotype also has the labels: [round, white with red border] “Type”, [white, typeset] “Sp. figured.”. Liogenys gebieni male lectotype, present designation (ZMHB): [white typeset] “Columbien / LaGarita [Los Patios, Norte de Santander] (Gebirge [mountains]) / W. Fritsche / leg. 1912” [genitalia mounted on the label], [white handwritten] “Isonychus [misspelled Liogenys] / gebieni / Typen Mos.”, [red typeset] “TYPUS”, [white typeset] “ Liogenys / gebieni / Mos. ”,. [white, outlined in red, typeset and handwritten] “ LIOGENYS / GEBIENI / Moser, 1921 / LECTOTYPE / Des. Cherman M. A.”. One female paralectotype with the same labels as the lectotype, plus the label: [white, outlined in red, typeset] “ PARALECTOTYPE / Des. Cherman M. A. [handwritten] LIOGENYS / GEBIENI Mos.”. Additional material examined. PANAMA. Colón: Coco Solo, 21.V.1976, B.C. Ratcliffe, 10 males, 20 females (UNSM); Coco Solo Hospital, V.1981, 2 males (UNSM); Fort Gulich, 17.V.1976, at lights, B.C. Ratcliffe, 1 male, 2 females; IV.1981, at lights, 5 males (UNSM); Fort Sherman, 26.IV.1980, D. Engleman, 1 female (UNSM). Panamá: Balboa, 16–17.V.1977, light, H.&A. Howden, 3 males, 1 female (CMNC); 19.VI.1977, H.&A. Howden, 1 male (CMNC); Corazal, 27.IV.1911, A. Busck, 2 males (USNM); Curundú, 8.V.1972, R.&E. Froeschner, 1 male (USNM); Diablo Heights, 3.V.1971, W.E. Bivin, 1 male, 1 female (USNM); Fort Clayton, 15.V.1961, C.E. Yunker, 8 males, 2 females (CNCI); Fort Clayton, V.1961, C.E. Yunker, 1 male (CMNC); La Sabanas, Panama City, IV.1927, at light, J. Zetek, 7 males, 19 females (USNM); Paraiso, 24.IV.1911, A. Busck, 2 males, 1 female (USNM); Punta Paitilla, Panama City, 8º59’N, 79º31`W, 17.V.1970, H. Hespenheide, 1 male (CMNC); Trivoli Hotel, Panama City, 15.IV.1967, H. Hespenheide, 1 male (CMNC). Geographical distribution. PANAMA (Chiriquí, Colón, Panamá), COLOMBIA (Norte de Santander). Remarks. After studying the type series of L. gebieni (ZMHB) (Fig. 5A–C), and comparing its external features and male genitalia (Fig. 5A–F) with that of L. macropelma type series and non-type material (Fig. 5G), we concluded that these specimens belong to the same species. The only difference is the color, which is darker in the L. gebieni types. Nonetheless, the difference in body color is a common intraspecific variation within Liogenys. Herein, we propose L. gebieni Moser as a junior subjective synonym of L. macropelma Bates. The parameres of each L. macropelma and L. gebieni drawn by Frey (1969) seem to be different. However, there were no specimens labeled as L. gebieni among Frey’s material housed at NHMB or USNM, thus, the material used by Frey (1969) to illustrate L. gebieni parameres remains unknown. According to the United States Department of Agriculture (1971, 1974), L. macropelma has been intercepted in Florida, Texas, and Puerto Rico (United States of America) coming by aircraft from El Salvador, Panama, and South America (probably Brazil) several times in a period from 1948–1971, always between April and May (identification verified based on the illustration in the report). Also, it was recorded being a host of okra plant Hibiscus esculentus L. (Malvaceae) coming from Guatemala, Panama, and Venezuela, which were intercepted in Florida, California, and Puerto Rico (United States of America). Several specimens in the USNM collection had been caught in port interceptions from Houston, Texas, originated from Panama, 1966 with no other explanation.Another specimen in the USNM bears the following data: 27 April 1962 port interception from Miami, Florida originating from “South America”. Based on the poor state of taxonomy of the genus Liogenys when these port interceptions took place, all identifications should be considered questionable unless they can be verified using voucher specimens., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on pages 212-214, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886, {"references":["Bates, H. W. (1887) Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. In: Salvin, O. & Godman, F. du C. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 25 - 160.","Nonfried, A. F. (1892) Verzeichniss der seit 1871 neu beschriebenen Glaphyriden, Melolonthiden und Euchiriden. Berliner Entomologische Zeitschrift, 37, 249 - 290. https: // doi. org / 10.1002 / mmnd. 18920370302","Dalla Torre, K. W. von (1913) Scarabaeidae: Melolonthinae IV. Pars 50. In: Schenkling, S. (Ed.), Coleopterorum catalogus. Vol. XX. Junk, W., Berlin, pp. 291 - 450.","Blackwelder, R. E. (1944) Checklist of the Coleopterous Insects of Mexico, Central America, The West Indies and South America. Part 2. Smithsonian Institution, Washington, D. C., pp. 189 - 265. https: // doi. org / 10.5479 / si. 03629236.185.2","Frey, G. (1969) Bestimmungstabelle und revision der gattung Liogenys Guerrez (Coleoptera - Melolonthinae - Macrodactylini). Entomologische Arbeiten aus dem Museum G. Frey, 20, 38 - 64.","United States Department of Agriculture [USDA] (1971) Insects not known to occur in the United States: A Scarab (Liogenys macropelma Bates). Cooperative Economic Insect Report, 21, 335 - 336.","United States Department of Agriculture [USDA] (1974) List of intercepted plant pests, 1972 (Pest recorded from July 1, 1971, through June 30, 1972). Hayattsville, Maryland. U. S. G. P. O., Washington, D. C., pp. 55 - 68.","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211, 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3 2009. Papers in Entomology, 1, 1 - 344.","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031","Moser, J. (1921) Neue Melolonthiden von Mittel- und Sud-Amerika. Stettiner Entomologische Zeitung, 82, 133 - 182. https: // doi. org / 10.1002 / mmnd. 192119210312"]}

Liogenys quadridentata Blanchard, 1851 Figs. 7, 9. Liogenys quadridentatus Blanchard, 1851: 168 (original description); Lacordaire 1855: 269 (checklist); Harold 1869:1140 (checklist); Dalla Torre 1913: 318 (checklist); Krajčík 2012: 145 (checklist). Liogenys quadridentata: Blackwelder 1944: 228 (checklist); Evans 2003: 213 (checklist); Evans & Smith 2009: 182 (checklist). Amphicrania 4-dentata [quadridentata] Klug in litt. Burmeister 1855: 14 (synonym of Liogenys quadridens, nomen nudum). Amphicrania quadridentatus Klug in litt.: Harold 1869: 1140 (synonym of L. quadridens). Type material examined. Liogenys quadridentatus male lectotype, present designation (MNHN): [white handwritten] “292 / 39”, [green handwritten] “ Amphicrania / 4 Dentata.”, [light green typeset] “MUSEUM PARIS”, [green handwritten] “ L. quadridentatus / Cat Mus”, [red typeset] “ HOLOTYPE ”, [white, outlined in red, typeset and handwritten] “ LIOGENYS / QUADRIDENTATA / Blanchard, 1851 / LECTOTYPE / Des. Cherman M. A.”, genitalia mounted. Additional material examined. PANAMA. Chiriquí: Volcán de Chiriquí, 2500–4000 feet, G.C. Champion, 4 males (BMNH). Panamá: La Chorrera, 2 females (BMNH). COLOMBIA. Magdalena: 12 miles W Santa Marta, 28.IV.1973, J.M. Campbell & H.F. Howden, 1 female (CMNC); 12 miles W Santa Marta, 29.IV.1973, J.M. Campbell & H.F. Howden, 1 male, 3 females (CMNC); 12 miles W Santa Marta, 12.V.1973, J.M. Campbell & H.F. Howden, 2 males (CMNC); 18.V.1973, 3 males, 5 females (CMNC); Parque Tayrona, 21 miles E Santa Marta, 15.V.1973, H.F. Howden & J.M. Campbell, 1 female (CMNC); Aracataca, 25.III.28, Darlington, 1 female (AMNH); Santa Marta, 3.V.1928, Darlington, 1 male (AMNH); Bonda, VI, 12 females (CMNH). VENEZUELA. Delta Amacuro: 15 km E Los Casillos, 45 km NE Ciudad Guyana, 26.VI.1985, ultraviolet light, M.A. Ivie, 3 females (CMNC). Guárico: 12 km W Valle de la Pascua, 21–22.VI.1996, light, H.&A. Howden, 1 female (CMNC); Estación Biológica Los Llanos, 12 km S Calabozo, 6–12.II.1969, ultraviolet light, P.&P. Spangler, 16 males, 9 females (USNM). Sucre: Cariaco, 20.V.1959, C. Bordon, 1 female (CMNC). Zulia: Maracaibo, without date, E. v. Jess., 1 male (ZMHB). Without locality and date, Moequerys, 2 females (SDEI); with erroneous locality (Brazil), without date, Mniszech, 1 female (MNHN). Diagnosis. Body light brown, elongate, sides parallel in males (Fig. 7A), slightly wider on posterior third in females; elytra light brown to yellowish brown, pronotum reddish to purplish brown; clypeus quadridentate; clypeal emargination deep, rounded, and narrow (Fig. 1C); anterior margin noticeable bent upwards, especially in females (Fig. 1D); distance between lateral and anterior clypeal teeth as long as basal width of one anterior tooth; acute angle between lateral and anterior clypeal teeth; mesotibia cylindrical in cross section (Fig. 7D); pygidium convex in both sexes, females with pygidium strongly convex medially (Fig. 7E); pygidial disc glabrous, with scattered bristles on apex; males with protarsomere II more than twice the length of protarsomere I in dorsal view; parameres up to three times the length of their apex (Fig. 7F); inner margins opened, slightly convergent; apex harpoon-shaped, lateral spine projected backwards, almost parallel to the outer margin of the parameres; spine length slightly more than twice the length of apex, subapical and part of the apex elevated medially, abruptly flattened at apex and spine. Redescription. Length: 9.4–11.2 mm, width: 4.7–5.2 mm. Brown. Head: distance between eyes nearly twice the width of one eye; frons somewhat swollen; clypeal emargination deep, rounded, and narrow; outer sides of anterior teeth parallel; outer margin of anterior teeth shorter than the eye in dorsal view; clypeus convex laterally and strongly produced forming a conspicuous tooth-like projection; in females anterior margin noticeable bent upwards; distance between lateral and anterior teeth equal to basal width of one anterior tooth, distance between lateral tooth and anterior margin of eye longer than one eye length, acute angle between anterior and lateral teeth; distal maxillary palpomere maximum width wider than at apex; fovea deep, extending up to the transverse midline of the palpomere; antenna with 10 antennomeres, club lighter in color and longer than funicle. Thorax: pronotal anterior margin slightly produced medially; punctures very sparse, denser and coarser than the disc towards anterior margin; pronotal posterior corners rounded; hypomere with long bristles; mesepisternum with sparse, thin scales; anterior portion and sides of metaventrite with bristles; distance between mesocoxae and metacoxae nearly twice as long as metacoxal width; scutellum triangular, sparse punctures at the base and sides. Elytra: shiny, uniform light brown or yellowish brown, lighter in color than the pronotum; elytral suture unicolored with elytron and slightly elevated or flat; all four elytral ridges weakly defined. Legs: procoxa with bristles, fine, thin scales on outer surface; three protibial teeth, middle and apical teeth equal in size, distance between basal and middle teeth longer than between middle and apical teeth; protibial spur present; mesotibia cylindrical in cross section, surface finely sculptured, mesotibia with two transverse carinae, the apical carina incomplete in males, complete in females; metacoxa with bristles throughout; inner margin of male metatibia carinate and produced straight towards apex, apical inner surface glabrous, metatibial disc coarsely sculptured along the outer margin; metatibial transverse carina present posteriorly; protarsomere II elongate; protarsomeres and mesotarsomeres I–IV enlarged, protarsomeres slightly wider than the mesotarsomeres, more than twice the width of metatarsomeres; metatarsomere I slightly shorter and wider than metatarsomere II; claw bifid, outer tooth of a claw equal in length and narrower than the inner tooth; distance between teeth shorter than the inner tooth. Abdomen: disc of ventrites with sparse bristles; propygidium with bristles, pygidium convex, convexity with intraspecific variation, females with pygidium inflated medially; pygidium subtrapezoidal, apex rounded; pygidial maximum width exceeding the distance between spiracles of propygidium; pygidial disc glabrous, few bristles on apex. Parameres: basal region narrowed at the midline; parameral split at 2/3 the length of the basal region; length of parameres nearly three times the length of their apex; inner margins convergent; apex harpoon-shaped, lateral spine projected backwards, slightly divergent from the outer margin of the parameres, spine length slightly more than twice as long as apex, subapical and proximal part of the apex elevated medially, abruptly flattened distally at apex towards the spine. Parameres concave, not coplanar in lateral view (Fig. 7G). Type locality. Not established. Geographic distribution. COLOMBIA (Magdalena), PANAMA (Chiriquí), VENEZUELA (Delta Amacuro, Guárico, Sucre, Zulia). Remarks. The lectotype of Liogenys quadridentata bears a green label “ Amphicrania 4-dentata ”, a name credited to “Klug MSC” by Burmeister (1855) but not formally validated until Blanchard (1851) described the species. Burmeister (1855) listed the name as a junior synonym of L. quadridens. Soon after, Harold (1869) mentioned “ Amphicrania 4-dentata ” as a synonym of L. quadridens, followed by “in litt.”, which means: “new species mentioned in litteris (in correspondence), that will be described later in a separate paper”. Harold (1869) probably did not see the L. quadridentata type series, so he did not associate the name “ Amphicrania 4-dentata ” to the latter. After the mentioned synonymy of “ Amphicrania 4-dentata ” (Burmeister 1855; Harold 1869), L. quadridentata was suggested to be a junior synonym of L. quadridens by later authors (Bates 1887; Dalla Torre 1913). We here confirm that L. quadridentata is a different species from L. quadridens. Liogenys quadridentata differs from L. quadridens (in parenthesis) mainly in the anterior margin of clypeus strongly bent upwards (not strongly bent); anterior and lateral tooth closer and acute angle between them (anterior and lateral tooth less close and right angle between them); mesotibia cylindrical in cross section (mesotibia subquadrate); pygidium noticeably convex, inflated medially in females (pygidium flat in both sexes); males with the protarsomere II longer (as wide as it is long), and the parameres are different in shape. The lectotype of L. quadridentata does not bear any locality data, and Blanchard (1851) did not mention the type locality in the original description. This fact agrees with Lacordaire (1855), who described the locality of L. quadridentata as “Patrie inconnue”. However, Burmeister (1855) and then Harold (1869) mentioned “Columbia” [current Colombia and Panama] as a locality record of “ Amphicrania 4-dentata ”, which is likely the actual distribution of this species. Dalla Torre (1913) cited L. quadridentata from Montevideo (Uruguay). This is here interpreted as an error, which was reproduced by Blackwelder (1944) and subsequent authors. Due to the layout of Blanchard (1851) descriptions when they are contiguous, it is easy to attribute the type locality of one species to the next one. It is most likely that Dalla Torre (1913) attributed the type locality of Liogenys parva Blanchard, 1851 to L. quadridentata., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on pages 217-218, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886, {"references":["Blanchard, E. (1851) Museum d'Histoire Naturelle de Paris. Catalogue de la Collection Entomologique. Classe des Insectes. Ordre des Coleopteres. Tome I. Gide et Baudry, Paris, 240 pp.","Lacordaire, M. T. (1855) Genera des Coleopteres. Famille XXX. Pectinicornes. In: Histoire Naturelle des Insectes. Librairie Encyclopedique de Roret, Paris, pp. 1 - 594.","Harold, E. F. von (1869) Scarabaeidae. In: Gemminger, M. & von Harold, E. F. (Eds.), Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus. Vol. 4. E. H. Gummi, Munich, pp. 976 - 1346.","Dalla Torre, K. W. von (1913) Scarabaeidae: Melolonthinae IV. Pars 50. In: Schenkling, S. (Ed.), Coleopterorum catalogus. Vol. XX. Junk, W., Berlin, pp. 291 - 450.","Blackwelder, R. E. (1944) Checklist of the Coleopterous Insects of Mexico, Central America, The West Indies and South America. Part 2. Smithsonian Institution, Washington, D. C., pp. 189 - 265. https: // doi. org / 10.5479 / si. 03629236.185.2","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211, 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3 2009. Papers in Entomology, 1, 1 - 344.","Burmeister, H. (1855) Handbuch der Entomologie (Coleoptera Lamellicornia Phyllophaga chaenochela). Vierter Band. Zweite Abtheilung. Theod. Chr. Fr. Enslin, Berlin, 570 pp.","Bates, H. W. (1887) Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. In: Salvin, O. & Godman, F. du C. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 25 - 160."]}

Key to Liogenys species from northern South America and Central America 1. Clypeus bidentate (Fig. 5A–B), only anterior margin of clypeus indented; distal maxillary palpomere evenly convex, without a fovea…................................................................... Liogenys macropelma Bates, 1887 - Clypeus quadridentate (Fig. 1A–D), anterior and lateral margin of clypeus indented; distal maxillary palpomere with an elongate, concave fovea................................................................................ 2 2. Angle between lateral and anterior teeth on clypeus slightly or strongly acute (Fig. 1C); males with lateral expansion along the parameres length entirely visible in dorsal view (Fig. 1F)...................................................... 3 - Angle between lateral and anterior teeth on clypeus right or obtuse (Fig. 1A); males with lateral expansion along the parameres at least partially hidden in dorsal view or absent (Fig. 1E)..................................................... 4 3. Frons and clypeus glabrous (Fig. 1C–D); angle between lateral and anterior teeth on clypeus strongly acute; pronotum moderate coarsely punctate; males with pygidium convex medially (Fig. 7D) and females with pygidium strongly inflated (Fig. 7E)...................................................................... Liogenys quadridentata Blanchard, 1851 - Frons and clypeus with bristles (Fig. 2C); angle between lateral and anterior teeth on clypeus slightly acute; pronotum strongly coarsely punctate; pygidium almost flat, sometimes slightly depressed medially (Fig. 2F)............................................................................................ Liogenys clipeosetosa Cherman, new species 4. Ventrites II, III, and sometimes IV with tubercles medially, tubercles variable in strength (Fig. 8E)...................................................................................... Liogenys schneiderae Cherman, new species - All ventrites smooth, without tubercles.................................................................... 5 5. Clypeal emargination clearly wider than the dorsal diameter of one eye (Fig. 1B)..... Liogenys quadridens (Fabricius, 1798) - Clypeal emargination as wide as or narrower than the dorsal diameter of one eye (Fig. 4B)........................... 6 6. Body and pronotum purplish brown, strongly darker than elytra; clypeal emargination strongly angulate; pronotum with anterolateral margin reflexed (Fig. 3C); pygidium without sexual dimorphism, weakly convex in both sexes.......................................................................... Liogenys genieri Smith & Cherman, new species - Body and pronotum reddish brown, slightly darker than elytra; clypeal emargination rounded; pronotum with anterolateral margin not reflexed, pygidium weakly convex in males and strongly convex to inflated in females....................................................................................... Liogenys granadina Cherman, new species, Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on page 203, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886, {"references":["Bates, H. W. (1887) Insecta. Coleoptera. Vol. II. Part 2. Pectinicornia and Lamellicornia. In: Salvin, O. & Godman, F. du C. (Eds.), Biologia Centrali-Americana. R. H. Porter, London, pp. 25 - 160.","Blanchard, E. (1851) Museum d'Histoire Naturelle de Paris. Catalogue de la Collection Entomologique. Classe des Insectes. Ordre des Coleopteres. Tome I. Gide et Baudry, Paris, 240 pp.","Fabricius, J. C. (1798) Supplementum entomologiae systematicae. Proft et Storch, Copenhagen, 572 pp."]}

Liogenys granadina Cherman, new species Figs. 4, 9. Type material. Holotype male, labeled: [white handwritten] “N. Grenada / Cartagena / coll[?] Chev.”, [red typeset and handwritten] “ LIOGENYS / GRANADINA / HOLOTYPE / Cherman M. A.”, genitalia mounted (ZMHB). Paratypes (45), all bearing the label [yellow, typeset and handwritten] “ LIOGENYS / GRANADINA / PARATYPE / Cherman M. A.”: One male bearing: [white handwritten] “ Venezuela / Orinoco river [?]”, [white typeset] “ Liogenys / [handwritten] quadri- / dens F / [typeset] det. G. Frey, 1968”, genitalia mounted (ZMHB). One female with the labels: [white handwritten] “nov. grenada”, [white typeset] “ Liogenys / [handwritten] quadridens / Mos / typeset] det. G. Frey, 1968” (ZMHB). One male with the data: [white, typeset] “ TRINIDAD: St. George Co. / Monos Island. 10º42’Nx / 61º41’W. 31 MAY 1987 / ROM#872012. M.L. Pickles ” (ROME). One male with the data: [pink, handwritten] “Columbia”, [white, typeset] “Pascoe / Coll. / 93—60”, [white, handwritten] “Telea / 4dentata / [illegible]” (BMNH). One male with the data: [green, handwritten] “Columbia / Lebas.”, [light blue, handwritten] “4-dentatus / Blanch. Cat. / p. 168”, [white, typeset] “So named / in Reiches / Collection. / C.W.”, [light blue, handwritten] “2259”, [green, handwritten] “ ♂ ”, [white, typeset] “67.45”, [white, handwritten] “4-dentatus. / Blanch. Dej. / brevicollis, / Reich. / Carthagena” (BMNH). Seven males and three females with the data [white, typeset] “ VENEZUELA, Guar. / 12kmS Carabozo / 6 II 1969 ”, [white, typeset] “P&PSpangler / Est. Bio. Los / Llanos uv lite” (two males and one female at CMNC, two males and one female at DZUP, three males and one female at USNM). Paratypes of DZUP with the sequence of voucher numbers: [white typeset] “ DZUP / 402750 ” to “ DZUP / 402752 ”. Ten males and seven females with the data [white, typeset] “ VENEZUELA / Guar. 12 Km S. / Carabozo / II-6-12-1969 / P.&P. Spangler ”, [white, typeset] “Est. Biologica / Los Llanos / Black Light” (three males and two females at CMNC, two males and two females at CEMT, five males and three females at USNM). Five males and two females with the data [white, typeset] “ VENEZUELA, Guar. / 12kmS Carabozo”, [white, typeset] “ 6-12 II 1969 / P&PSpangler / BiolSta uv lite” (USNM). One male with the data [white, typeset] “ VENEZUELA, Guar. / 12kmS Carabozo”, [white, typeset] “ 6-12 II 1969 / P&PSpangler” (USNM). One female with the data [white, typeset and handwritten] “Cd Bolivar / 23-6 Venez”, [white, typeset] “EA Klages / Collector” (USNM). One female with the data [white, typeset and handwritten] “Cd Bolivar / 20-6 Venez”, [white, typeset] “EA Klages / Collector”, [white, typeset and handwritten] “ Liogenys / quadri- / dens / det. G.Frey,1968” (USNM). One female with the data [white, typeset and handwritten] “Cd Bolivar / 20-6 Venez”, [white, typeset] “EA Klages / Collector” (USNM). One female with the data [white, typeset] “ VENEZUELA: Guarico / Hato Masaguaral / (44km S Calabozo) / May 3–10 1985 / Menke & Carpenter” (USNM). Two females with the data [white, typeset and handwritten] “ VENEZUELA / Maracay / 450 m. / 15.VI.1949. / H.E. Box ” (USNM). Holotype and two paratypes deposited at ZMHB. Two paratypes at BMNH. Eight paratypes at CMNC. Three paratypes at DZUP. Four paratypes at CEMT. One paratype at ROME. Twenty-five paratypes at USNM. Diagnosis. Body light brown, elongate, sides almost parallel; elytra light brown; pronotum reddish brown (Fig. 4A); clypeus quadridentate; clypeal emargination deep, rounded, and narrow; distance between lateral and anterior clypeal teeth longer than basal width of one anterior tooth; angle between lateral and anterior clypeal teeth 90º or slightly acute; pronotal corners rounded (Fig. 4B); mesotibia quadrate in cross section; pygidium convex; pygidial disc glabrous, with scattered bristles on apex (Fig. 4D); in males protarsomere II more than twice the length of protarsomere I in dorsal view (Fig. 4C); parameres four times the length of their apex; lateral expansion along the parameres only seen subapically; inner margins convergent; apex harpoon shaped, lateral spine projected at approximately a 45º angle, spine length as long as the apex (Fig. 4E). Holotype. Male. Length: 9.2 mm, width: 4.6 mm. Light brown. Head: distance between eyes nearly twice the width of one eye; frons equal in length to clypeus; clypeal emargination deep, rounded, and narrow; outer sides of anterior teeth parallel; outer margin of anterior teeth shorter than the eye in dorsal view; clypeus convex laterally and strongly produced forming a conspicuous tooth-like projection; distance between lateral and anterior teeth longer than basal width of one anterior tooth, distance between lateral tooth and anterior margin of eye longer than one eye length, angle between anterior and lateral teeth approximately 90º; canthus not exceeding the outer margin of the eye; distal maxillary palpomere not seen; antenna with 10 antennomeres, club lighter in color and longer than funicle. Thorax: pronotal anterior margin moderately produced medially, undulate and flanged; maximum length of pronotum exceeding the length of tarsomeres I–III combined; disc glabrous, punctures very sparse and coarse; pronotal posterior corners rounded; hypomere with long bristles; mesepisternum scaly; anterior portion and sides of metaventrite with bristles; distance between mesocoxae and metacoxae nearly twice the length of the metacoxa; scutellum triangular with sparse, coarse punctures, mainly at the sides. Elytra: shiny, uniform light brown, slightly lighter in color than the pronotum; elytra more than three times longer than the pronotum; elytral suture darker than the elytron and weakly elevated; all four elytral ridges weakly defined. Legs: procoxa with abundant, thin and long scales, outer surface with short scales; three protibial teeth, middle and apical teeth equal in size, distance between basal and middle teeth longer than between middle and apical teeth; protibial spur present; mesofemur with a row of long bristles on anterior and posterior margins, mesotibia subquadrate in cross section, surface coarsely sculptured on outer margin, mesotibia with two transverse carinae, the apical carina incomplete; metacoxa with bristles throughout, basal articular lobe of metacoxa produced beyond the outer margin of trochanter; inner margin of metatibia carinate and produced on apex, apical inner surface with sparse bristles, metatibial surface coarsely sculptured; metatibial transverse carina present posteriorly; protarsomere II elongate; protarsomeres and mesotarsomeres I–IV enlarged, protarsomeres wider than mesotarsomeres and two times wider than metatarsomeres; metatarsomere I slightly shorter than metatarsomere II; claw bifid, outer tooth of a claw longer and as wide as the inner tooth; distance between teeth shorter than the inner tooth. Abdomen: disc of ventrites with bristles; propygidium with bristles, pygidium convex, subquadrate, wide; pygidial width exceeding distance between spiracles of propygidium; pygidial disc glabrous and apex with few bristles, pygidial disc coarsely punctate; pygidial apex subquadrate. Parameres: basal region dorsally narrowed, equal to both sections of the parameres at its midline; parameral split at 2/3 the length of the basal region; parameres four times the length of their apex; inner margins convergent; lateral expansion along the parameres only seen subapically; apex harpoon-shaped, lateral spine projected straight, at approximately a 45º angle, spine length as long as the apex. Parameres concave in lateral view (Fig. 4F). Variation. Female length: 9.5–11.4 mm, width 4.8–5.4 mm. Females as the holotype except in the size, head, and pronotum darker than elytra; pronotum wider, with pronotal corners obsolete; maximum width of distal maxillary palpomere slightly wider than at apex; fovea deep, extending up to the transverse midline of the palpomere; scutellum ogival. Male length: 8.4–10.2 mm, width 4.2–5.0 mm. Males as the holotype except in the scutellum ogival; punctures of pronotum and tibiae less marked; elytral suture unicolored with the elytron; ventral expansion of the outer margins of the parameres more apparent. Etymology. This species is named after Nueva Grenada, a defunct nation corresponding to present-day Colombia and parts of many other countries in South America (Brookes 1819). This name should be considered an adjective in the nominative singular. Type locality. Nueva Grenada (Republic of New Grenada, today a region that comprises Panama, many South American countries, and the total territory of Colombia). Geographical distribution. TRINIDAD AND TOBAGO (Saint George), COLOMBIA (Cartagena), VENEZUELA (Guaricó, Maracay). Remarks. Liogenys granadina Cherman, new species resembles L. quadridens in the shape of the quadridentate clypeus and of the pygidium. However, L. granadina is smaller than L. quadridens; the distance between anterior and lateral clypeal teeth is shorter; and in the pronotal corners are defined (obsolete in L. quadridens). Frey (1969) considerated specimens of L. granadina as being L. quadridens, judging by his description in the key, one of the two parameres illustration (the other one corresponds to L. quadridens), and because two paratypes (USNM and ZMHB) bear his determination label. Moreover, Frey (1969) refers to a specimen from Pernambuco, Brazil, which was found at the NHMB. This Brazilian specimen was identified as Liogenys piauiensis Cherman, 2017 by Cherman et al. (2019). There are three specimens in the USNM that were intercepted at the port of Miami, Florida on 1 May 1964 with no other information on the host or origin. This indicates that some previous port interceptions or pest reports for L. quadridens possibly included or even all of them are specimens of L. granadina., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on pages 209-212, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886, {"references":["Brookes, R. (1819) General Gazeteer ... Illustrated with maps ... The fifteenth edition ... with considerable additions and improvements. 15 th Edition. Plummer and Brewis, Printers, Love-Lane, Little-Eastcheap, London, 796 pp.","Frey, G. (1969) Bestimmungstabelle und revision der gattung Liogenys Guerrez (Coleoptera - Melolonthinae - Macrodactylini). Entomologische Arbeiten aus dem Museum G. Frey, 20, 38 - 64.","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031","Cherman, M. A., Basilio, D. S., Mise, K. M. & Almeida, L. M. (2019) Unraveling the puzzle of Liogenys biodiversity: fifteen new species, nomenclatural acts and new geographical records (Coleoptera: Scarabaeidae). Insect Systematics & Evolution, 51, 1 - 69. https: // doi. org / 10.1163 / 1876312 X- 00001040"]}

Compsodactylus argentinus (Moser, 1919) Material examined (all from the CMNC). ARGENTINA: Salta: Departamento Capital, Villa San Lorenzo, ii.1988, 1 male; Coronel Moldes, ii.1945, Francisco Monr��s (collector), 2 males, 1 female; La Vi��a, ii.1992, 2 males; Departamento La Vi��a, La Vi��a, i.1991, M.A. Viana (collector), 1 male, 1 female; Departamento La Vi��a, Ruta 68, 4 km S La Vi��a, i.1988, 10 males, 7 females; Departamento La Vi��a, El Carmen, ii.1993, 25 males, 10 females; Departamento La Vi��a, El Carmen, iii.1993, 7 males. CatamaRCa: Campo del Arenal, ii.1983, L. Pe��a (collector), 8 males, 5 females; Santa Maria Village, 26��41���S, 66��02���W, 14.ii.1983, 3 males; Chafi��an, Weiser (collector), 2 males, 1 female. miSionES: Departamento Capital, Itaemb�� Min��, iv.1993, M.A.F. (collector), 2 males. tuCuman, San Pedro de Colalao, i.1947, P. Arnau (collector), 2 females. CoRDoba, Departamento de Calamuchita, Ca��ada del Sauce (��� El Sauce ���), xii.1938, Manuel J. Viana (collector), 2 males. Distribution (Fig. 26). New provincial records were discovered as follows. Argentina: C��rdoba: Departamento de Calamuchita: ���El Sauce���; Argentina: Misiones: Departamento Capital: Itamb�� Min��. This species was previously known from Catamarca, Salta, and Tucum��n in Argentina. Note. Fuhrmann (2019) noted that the pronotum of Compsodactylus parvulus (Frey, 1970) is dark brown and sometimes the lateral angles and posterior margin have yellow maculae. This same pattern was observed in some specimens of C. argentinus., Published as part of Fuhrmann, Juares & Smith, Andrew B. T., 2021, Compsodactylus Fuhrmann, 2012 (Coleoptera: Scarabaeidae: Melolonthinae) distributional notes and a new species, pp. 387-393 in Zootaxa 4990 (2) on pages 387-388, DOI: 10.11646/zootaxa.4990.2.12, http://zenodo.org/record/5026053, {"references":["Fuhrmann, J. (2019) Taxonomy of Neotropical genera Compsodactylus and Dasyus and notes on claw movement in Macrodactylini (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 4679 (1), 139 - 163. https: // doi. org / 10.11646 / zootaxa. 4679.1.9"]}

Liogenys clipeosetosa Cherman, new species Figs. 2, 9. Type material. Holotype male, labeled: [green typeset] “Venezuela / F. Kummerow S.”, [red typeset and handwritten] “ LIOGENYS / CLIPEOSETOSA / HOLOTYPE / Cherman M. A.”, genitalia mounted (ZMHB). Paratypes (10) all bearing the label [yellow, typeset and handwritten] “ LIOGENYS / CLIPEOSETOSA / PARATYPE / Cherman M. A.”. Six female paratypes bearing the same data of the holotype (ZMHB). Three female paratypes with the label: [green, outlined, typeset] “Hist. Coll. (Coleoptera) / Nr. 24131 / Liogenys quadridens Burm. / Columb., Moritz. / Zool. Mus. Berlin” (ZMHB). One female paratype with the data: [white, typeset] “ VENEZUELA: Lara / 20 km. E. Carora / 24-VI-1976 / Holotype and nine paratypes deposited at ZMHB. One paratypes at USNM. Additional material. Three females (FSCA) were port interceptions from Gulfport, Mississippi, United States of America in Musa (Musaceae) (bananas, plantains, etc.) originating from Colombia. We examined photographs of these specimens provided by John Leavengood (United States Department of Agriculture, Tampa, Florida, United States of America). No specific locality was available within Colombia and it is likely that these specimens were attracted to lights during the packing or loading of this commodity. Diagnosis. Body light brown; elongate (Fig. 2A–B), elytra light brown to yellowish brown, pronotum reddish brown, head darker; clypeus quadridentate; clypeal emargination deep, rounded, and wide; anterior teeth sometimes twice the length of the lateral teeth (Fig. 2C); distance between lateral and anterior clypeal teeth longer than basal width of one anterior tooth; angle between lateral and anterior clypeal teeth 90º or acute; clypeus and part of the frons with bristles; pronotum roughly punctate; pronotal corners rounded or subangulate; mesotibia quadrate in cross section (Fig. 2D); pygidium flat or slightly convex, depressed medially (Fig. 2E–F), pygidial disc with short bristles throughout, bristles longer on apex; males with protarsomere II up to twice the length of protarsomere I in dorsal view; parameres elongate, inner margins straight; lateral expansion along the parameres length entirely visible in dorsal view; apex harpoon-shaped, lateral spine projected backwards, slightly convergent compared to the outer margin of the parameres; spine length slightly longer than the apex, subapical and part of the apex not elevated on midline (Figs. 1E, 2G). Holotype. Male. Length: 12.0 mm, width: 5.8 mm. Light brown. Head: darker than pronotum, distance between eyes less than twice the width of one eye; frons equal in length to clypeus; frontoclypeal surface with bristles, clypeal emargination deep, rounded, and wide; outer sides of anterior teeth parallel; outer margin of anterior teeth shorter than the eye in dorsal view; clypeus convex laterally and strongly produced forming a conspicuous, tooth-like projection; distance between lateral and anterior teeth longer than basal width of one anterior tooth, distance between lateral tooth and anterior margin of eye equal to one eye length, angle between anterior and lateral teeth approximately 90º; canthus not exceeding the outer margin of the eye; distal maxillary palpomere, maximum width more than twice the width at apex; fovea shallow, extending past the transverse midline of the palpomere; antenna with 10 antennomeres, club lighter in color and longer than funicle. Thorax: pronotal anterior margin slightly produced medially, undulate and flanged; anteriorly and on posterolateral surface depressed, sulcate medially; pronotal disc glabrous, punctures sparse medially and rugose on anterior margin; pronotal posterior corners rounded; hypomere with few long bristles; mesepisternum scaly; anterior portion and sides of metaventrite with bristles; distance between mesocoxae and metacoxae nearly twice the width of the metacoxa; scutellum ogival, sparse punctures. Elytra: shiny, uniform yellowish brown, lighter in color than pronotum; elytra near three times the length of pronotum; elytral suture slightly darker than elytron and not elevated; all four elytral ridges weakly defined. Legs: procoxa with long bristles on infra-carinal surface, fine scales and short bristles on outer surface; three protibial teeth, middle and apical teeth equal in size, distance between basal and middle teeth longer than between middle and apical teeth; mesofemur with a row of long bristles on anterior and posterior margins, mesotibia quadrate in cross section, surface coarsely sculptured, mesotibia with two transverse carinae, the apical carina incomplete; metacoxa with bristles throughout; basal articular lobe of metacoxa produced beyond the outer margin of trochanter; inner margin of metatibia carinate and produced on apex, apical inner surface with scattered bristles, metatibial surface coarsely sculptured; metatibial transverse carina and discontinuous longitudinal carina present posteriorly; protarsomere II elongate; protarsomeres and mesotarsomeres I–IV enlarged, protarsomeres slightly wider than mesotarsomeres, more than twice the width of metatarsomeres; metatarsomere I and metatarsomere II equal in length; claw bifid, symmetrical, outer tooth of a claw longer and as wide as the inner tooth; distance between teeth shorter than the inner tooth. Abdomen: disc of ventrites with sparse bristles; propygidium with bristles, pygidium flat, subtrapezoidal, wide, pygidial width exceeding distance between spiracles of propygidium; pygidial disc glabrous except for a few bristles on apex, pygidial disc coarsely punctate; depressed medially; pygidial apex quadrate. Parameres: basal region slightly wider than both sections of the parameres at its midline; parameral split at 2/3 the length of the basal region; inner margins straight; lateral expansion along the parameres length entirely visible in dorsal view; apex harpoon-shaped, laterally expanded and blunt projected. Parameres concave in lateral view (Fig. 2H). Variation. Female paratypes. Length: 11.2–12.6 mm; width 5.4–6.3 mm. As the holotype except pygidial apex rounder, wider and somewhat convex, not depressed medially. Some females have the clypeal teeth noticeable longer. One female from Venezuela has a subtrapezoidal pygidium. Etymology. Adjective in the nominative singular. From the Latin clipeus (clypeus) + Latin setosus, from seta (“bristle”). Type locality. Venezuela. Geographical distribution. COLOMBIA, VENEZUELA (Lara). Remarks. Liogenys clipeosetosa Cherman, new species resembles L. quadridens in the body color and size, shape of the quadridentate clypeus, pygidium, and somewhat in the shape of the male parameres. Liogenys clipeosetosa differs (L. quadridens in parenthesis) in the clypeus with bristles (clypeus glabrous), fovea of the distal maxillary palpomere shallow, surface almost flat (fovea deep); pronotal anterior margin flanged (flange less conspicuous or absent); pronotum depressed anteriorly and on posterolateral surface, sulcate medially (not as so); pronotal disc roughly punctate mainly on anterior margin (pronotal disc less coarse and uniformly punctate); males with the protarsomere II longer than wide (as wide as it is long); apex of parameres entirely flat and lateral expansion along the parameres length entirely visible in dorsal view (subapical and proximal part of the apex elevated medially and lateral expansion of parameres partially hidden, visible from the midline up to the subapical portion)., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2021, Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species, pp. 201-226 in Zootaxa 4990 (2) on pages 203-206, DOI: 10.11646/zootaxa.4990.2.1, http://zenodo.org/record/5025886, {"references":["Fabricius, J. C. (1798) Supplementum entomologiae systematicae. Proft et Storch, Copenhagen, 572 pp.","Blanchard, E. (1851) Museum d'Histoire Naturelle de Paris. Catalogue de la Collection Entomologique. Classe des Insectes. Ordre des Coleopteres. Tome I. Gide et Baudry, Paris, 240 pp."]}

Liogenys maxillaricuspis Cherman, new species Figs. 8 A-I; 12 E-F; 13 A, C, G; 15 Type material. Holotype, male, labeled (CMNC): [white typeset] “ ARGENTINA: Rio Negro / Dto Gral Roca, Catriel / 37º52’30’’S 67º5’10’’W / 4 Jan. 2005, 311 m / F.C. Ocampo ”, [red typeset and handwritten] “ LIOGENYS / MAXILLARICUSPIS / HOLOTYPE / Cherman M.A.”, genitalia mounted. Paratypes (84) labeled [yellow typeset and handwritten] “ LIOGENYS /MAXILLARICUSPIS / PARATYPE / Cherman M. A.”: Seven males and six females with the same data of the holotype (DZUP). Three males and two females with the same data of the holotype (CEMT). Thirty one males and twenty eight females with the same data of the holotype (CMNC). The males with the following sequence of vouchers: [typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029648–81 ”. The females with the following sequence of vouchers: [typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029682–713 ”. Eight specimens with the same data as the holotype, and DNA stored in alcohol at -80ºC and the following voucher numbers (CMNC): Four males with the sequence: [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3003308–11 ”. Two females with the sequence: [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3003312–13 ”. One female [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3003306 ”, [typeset] “A.B. T. SMITH DNA / 28S/18S VOUCHER / SPECIMEN SMI-131”. One female: [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3003307 ”, [typeset, outlined black]. “A.B. T. SMITH SCARAB / DNA VOUCHER / SPECIMEN / AS498 / JULY 2005 ”. One male and one female paratype (IADIZA) labeled: [white typeset] “Arg. Rio Negro / Gral. Rocas / Cnia. Catriel 311 m / S. Roig / G. Flores / 04/I/05”, [white typeset] “ 37º52.83’S / 67º56.16’W ”. One male (CMNC): [white typeset] “ ARGENTINA, Pr. San Luis / 18 km S Arizona / 18-23.I.1982 [space] 250 m / H & A Howden”. One male and three females (VMDC): [typeset] “Argentina Chubut / Gaiman, Bryn Gwyn / 4 Noviembre 2008 / Leg. Daniel Rojas L.”. The male with the voucher: [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3002827 ”. The females with the sequence of vouchers: [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3002860–62 ”. Holotype and 68 paratypes at CMNC. Five paratypes deposited at DZUP. Five paratypes deposited at CEMT. Two paratypes deposited at IADIZA. Four paratypes deposited at VMDC. Diagnosis. Body elongate, dull yellow (Fig. 8A); pronotum slightly darker and slightly wider than the humeral distance; margin of clypeus, pronotum and protibia darker; frons swollen, coarsely punctate; antennae with 9 antennomeres; clypeal emargination deep, narrowed, rounded; teeth subangulate (Fig. 12 E–F); clypeal lateral margin convex; pronotal posterior corners rounded (Fig. 8B); elytra with sparse, random bristles; males and females with one metatibial transverse carina complete, inner margin not carinate; pygidium with sparse, long bristles; smooth, as long as wide (Fig. 8E); inner margins of parameres straight, opened, apex spatulate, apically slightly divergent and curved downwards, lateral margins of apex below the plane of the parameres (Fig. 8 F–H). Holotype. Male. Length: 7.8 mm; width: 3.5 mm. Dull yellow. Head: distance between eyes slightly more than twice the width of one eye; frons swollen, equal in length to clypeus; frontoclypeal surface with few bristles; anterior margin of clypeus dark in color (Fig. 12 E–F); clypeal emargination shallow, wide and rounded; outer sides of anterior teeth subparallel; clypeal lateral margin convex, not produced; distal maxillary palpomere, maximum width slightly wider than apex; fovea deep, extending up to the transverse midline of the palpomere; antenna with 9 antennomeres, club lighter in color and longer than the funicle. Thorax: anterior margin of pronotum slightly produced medially, pronotum anteriorly narrowed forming a depressed ring; pronotal disc with punctures coarse and very sparse; posterior corners rounded; hypomere with long bristles; mesepisternum, sides of metaventrite and metacoxae with sparse bristles; distance between mesocoxae and metacoxae less than twice the metacoxa length; scutellum ogival, finely punctate throughout and with bristles. Elytra: disc shiny; bristles on disc disposed randomly; elytral suture and elytron unicolored, not elevated; two inner elytral ridges more defined than the outer ridge, third ridge undistinguishable. Legs: procoxa with bristles and punctate; protibia with three teeth, almost equal in size and equally spaced; mesotibia cylindric, somewhat flattened in cross section; mesotibial surface finely sculptured; mesotibia with two transverse carinae, the apical carina complete; inner margin of metatibia not carinate; surface finely sculptured; metatibia with two transverse carinae, the apical carina complete; metatibial apical spurs with different lengths, the longer spur equal to the diameter of the tibial apex; metatarsomere I slightly shorter than tarsomere II; protarsomeres I–IV slightly enlarged, protarsomere II elongate; claw bifid, symmetrical, superior tooth of claw longer and equal in width to the inferior tooth; distance between teeth longer than the length of inferior tooth. Abdomen: ventrites with uniform and sparse bristles at the disc and sides; propygidium with bristles; pygidium slightly convex; subquadrate; as long as it is wide; pygidial width not exceeding distance between spiracles of propygidium; pygidial disc almost smooth, few punctures at the sides, sparse, long bristles throughout; pygidial apex rounded. Parameres: basal region as wide as both sections of the parameres at its maximum width, parameral split at 2/3 the length of the basal region; inner margins straight, opened, apex spatulate, apically slightly divergent and curved downwards; parameres in lateral view straight, coplanar (Fig. 8I). Variation. Male paratypes. Length: 7.1–9.4 mm; width: 3.4–4.5 mm. As the holotype, except in the clypeal lateral margin produced (the most in male from San Luis) (Fig. 8C), and in the deepness of the pronotal punctures, some of them being shallower. Female paratypes. Length: 8.9–9.0 mm; width: 4.2–4.3 mm. As the holotype except in the size larger, the lateral margin of clypeus sometimes more convex (Fig. 12F) and the pronotal punctures deeper. Etymology. Adjective in the nominative singular. Prefix from the Latin maxillaris (“related to the maxilla: set of paired mouthparts used to manipulate food”) + from Latin cuspis (“sharp point”). Liogenys maxillaricuspis new species is characterized by having a maxilla furnished with only one, strongly sharp, curved teeth, which can be seen with lowest magnification (6x) in ventral view of the head (Fig. 13A, G). This feature is unique among Liogenys species. Type locality. ARGENTINA, Río Negro, Departamento General Roca, Catriel, 37º52’30’’S 67º5’10’’W. Geographical distribution. ARGENTINA (San Luis, Río Negro). Remarks. Liogenys maxillaricuspis new species resembles L. rufoflava in the body size and color, elytra with bristles, and somewhat in the shape of the bristles on the pygidium, although the bristles are sparser and longer. Liogenys maxillaricuspis new species has unique characters for the genus, especially the mouthparts, but also the elytral ridges and the setae on tibial carinae. In addition to the already mentioned single morphology of the maxilla, the ligula of the labium is shorter than other species, with margin almost straight, without lobes (Fig. 13 C–D). Since this new species has most of the synapomorphies of Liogenys (Cherman et al. 2016), we are describing it in this genus. New molecular phylogeny testing Diplotaxini relationships is being carried out and will bring light about the real position of this species among the Liogenys and other Diplotaxini lineages. The male paratype (CMNC) from San Luis province [18 km S Arizona / 18-23.I.1982 [space] 250 m / H & A Howden] shows slight variations in the shape of the clypeus and in the parameres when compared with the Patagonian specimens (Figs. 8C, G). However, we considered it to be L. maxillaricuspis new species because the maxilla has one tooth (Fig. 13 G), which is distinctive of this new species., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 69-70, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172, {"references":["Cherman, M. A., Moron, M. A. & Almeida, L. M. (2016) Phylogenetic relationships within Diplotaxini Kirby (Coleoptera: Melolonthidae: Melolonthinae) with emphasis on Liogenys Guerin-Meneville. Systematic Entomology, 41, 744 - 770. https: // doi. org / 10.1111 / syen. 12188"]}

Liogenys lucialmeidae Cherman, new species Figs. 6, 12 A–B, 15 Type material. Holotype, male, labeled (CMNC): [white typeset] “ ARGENTINA: Mendoza / Reserva La Payunia, Los Relinchos / S36º00’59’’W68º48’39’’ / Jan-6-2003, 1714 m / F.C. Ocampo, A.B. T. Smith”, [red typeset and handwritten] “ LIOGENYS / LUCIALMEIDAE / HOLOTYPE / Cherman M.A.”, genitalia mounted. Paratypes (20) labeled [yellow typeset and handwritten] “ LIOGENYS LUCIALMEIDAE / PARATYPE / Cherman M. A.”: six females with the label (four at CMNC, two at DZUP (402743-44)): [white typeset] “ ARGENTINA: Mendoza / 1 km Reserva Telteca / S32º 18’40’W67º54’08’’ / Jan-04- 2003, 540 m / F.C. Ocampo, A.B. T. Smith ”. One male (IADIZA): [white handwritten] “ Malarg ̹e: A 7 km de / Ranquil Norte, camino / a Barrancas [Neuquen Prov.] / 23- I-79 / Sergio Roig ”, [circle full blue, indicating C. Berg collection]. One male and one female paratypes (IADIZA): [white typeset] “ ARGENTINA: Chubut Peninsula de / Valdés 8km S Punta norte / 42º12’20’’S 63º52’5,9’’W 42 m / 14-I-2010 Coll: G. Flores, G. Cheli, / R. Carrara”. One male paratype (AMNH): [white typeset] “ ARGEN-TINA / El Bolson, Rio / Negro / 19 [space] / A. Kovacs ”. Two males mounted on cards on the same pin (CMNC): [handwritten by A. Martínez] “ ARGENTINA / RIO NEGRO / Cnel. Gómez / J. Grosso-leg. / Coll. Martínez / NOV-945”, [typeset, outlined black] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.”, [typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029587 ”. One male (CMNC): [typeset] “ARG: Nuequin / nr Zapala / 19–21.XII.1965 / A. Martinez ”, [typeset, outlined black] “H. & A. HOWDEN / COLLECTION / Ottawa, Canada”, [typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029588 ”. One male (CMNC): [typeset] “ARG: Nuequin / nr Zapala / 19–21.XII.1965 / A. Martinez ”, [typeset, outlined black] “H. & A. HOWDEN / COLLECTION / Ottawa, Canada ”, [typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029589 ”. One male (CMNC): [typeset] “ 24-XI-1955 / Villa Regina / Pcia. Rio-Negro / Lg. E.Fleiss ”, [typeset, outlined black] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.”, [typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029590 ”. One male (CMNC): [typeset] “ SA. VACA MUERTA / (950 M. S.N.M.) / NEUQUEN-ARG.”, [typeset] “14-XII-74 / LG.M.GENTILI”, [typeset, outlined black] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.”, [typeset with data matrix barcode] “ Cana-dian Museum of / Musée canadien de la / NATURE / CMNEN 00029591 ”. One male and three females (NHMW): [typeset] “ ARGENTINIEN / Neuquen Anelo / 26.10.1981 leg. KOVAC ”. The male with the voucher: [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3002724 ”. The females with the sequence of vouchers: [typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3002787-89 ”. Holotype and 10 paratypes deposited at CMNC. Two paratypes deposited at DZUP. Three paratypes deposited at IADIZA. One paratype deposited as AMNH. Four paratypes at NHMW. Diagnosis. Body elongate; dull yellow, pronotum and elytra unicolored or pronotum slightly darker (Fig. 6A), frons swollen, mainly in females; in males clypeal emargination deep, subangulate to rounded and wide; in females clypeus short and weakly emarginate (Fig. 12 A–B); antennae with 10 antennomeres; pronotum and head equal in length, posterior corners subangulate (Fig. 6B); in males pronotum with maximum length medially, in females pronotum with uniform length throughout (Fig. 12 A–B); elytra semi-opaque, with bristles uniformly, in males sometimes somewhat not uniformly pruinose; pygidium elongate or as wide as it is long, disc smooth, with bristles throughout (Fig. 6C); males with two protibial teeth; metatibia with inner margin weakly carinate towards the apex and one transverse carina incomplete (Fig. 6 D–E); parameres forming a concavity at the midline; slightly narrowed subapically; apex sagittate, curved upwards (Fig. 6 F–H). Holotype. Male. Length: 9.4 mm; width: 4.3 mm. Uniform dull yellow. Head: distance between eyes almost three times the width of one eye; frons equal in length to clypeus, swollen; clypeal emargination deep, subangulate, wide; outer sides of anterior teeth subparallel, apparently following the lateral margin of clypeus, lateral margin straight; distal maxillary palpomere, maximum width two times the apical width; fovea deep, reduced to a point at the widest part of the palpomere; antenna with 10 antennomeres, club lighter in color than funicle. Thorax: anterior margin of pronotum slightly produced medially; pronotal disc with punctures fine and sparse; posterior corners subangulate, angle obtuse; hypomere with short bristles and a few scales at the inner margen; mesepisternum, sides of metaventrite and metacoxae with sparse bristles; distance between mesocoxae and metacoxae twice the metacoxa length; scutellum ogival, with a few punctures. Elytra: semi-pruinose, bristles dispersed randomly; elytral suture and elytron unicolored, weakly elevated; all four elytral ridges weakly defined (Fig. 6A). Legs: procoxa weakly scaly; protibia with two teeth equal in size; mesotibia cylindrical and somewhat flattened in cross section, mesotibial surface finely sculptured; mesotibia with one transverse carina only demarked posteriorly; inner margin of male metatibia weakly carinate towards the apex; surface finely sculptured; metatibia with one transverse carina incomplete; metatibial apical spurs of different lengths and shape, the longer spur equal to the diameter of the tibial apex, the shorter spur wider, truncate, and slightly projected downwards; metatarsomere I slightly shorter and equal in width to tarsomere II; protarsomeres and mesotarsomeres I–IV enlarged, protarsomere II as wide as long; mesotarsomeres less enlarged than the protarsomeres; claw bifid, symmetrical, superior tooth of a claw longer and slightly wider than the inferior tooth; distance between teeth longer than the length of inferior tooth. Abdomen: ventrites with sparse bristles; propygidium with bristles; pygidium slightly convex; subtrapezoidal, as long as wide; pygidial width not exceeding distance between spiracles of propygidium; pygidial disc smooth and with bristles throughout; pygidial apex subquadrate. Parameres: basal region as wide as both sections of the parameres at maximum width, parameral split at 2/3 the length of the basal region; inner margins convex at the dorsal plane, forming a concavity at the ventral plane; slightly narrowed subapically; apex sagittate, curved upwards; parameres convex in lateral view. Variation. Male paratypes. Length: 10.0– 11.4 mm; width: 4.4–5.0 mm. As the holotype except in the lateral margin of clypeus more concave. Female paratypes. Length: 9.6–10.9 mm; width: 4.8–5.5 mm. As the holotype except in the clypeus short and weakly emarginate; pronotum shorter; elytral ridges more marked; and three protibial teeth. Etymology. Noun in the genitive case. The species is dedicated to Dr. Lúcia Massutti de Almeida, a renowned Brazilian Coleopterist and mentor of the first author. Type locality. ARGENTINA, Mendoza, Reserva La Payunia, Los Relinchos, 36º00’59’’S 68º48’39’’S, 1714 m. Geographical distribution. ARGENTINA (Mendoza, Neuquén, Río Negro, Chubut). Remarks. Liogenys lucialmeidae new species resembles L. flavida in the body size and color; shape of the clypeus, especially in females (Fig. 11 C–D); elytra with bristles; males with two protibial teeth and metatibia with one transverse carina incomplete. Liogenys lucialmeidae new species differs in the clypeal emargination deep in males (shallow); antennae with 10 antennomeres (9 antennomeres); pronotum shorter; males with inner margin of metatibia weakly carinate towards the apex (distinctly carinate); metatibia with the transverse carina more strongly defined; and the different shape of the parameres. The new species also resembles L. kuntzeni, especially the females in the shape of the clypeus (Fig. 11 E–F), differing from this species (in parenthesis) in the clypeal emargination rounder in males (angulate); lateral margins of clypeus straight or slightly concave but teeth never parallel (slightly convex, teeth parallel); females with the clypeal teeth even less noticeable and pronotum shorter; pygidium longer; males with one metatibial transverse carina incomplete (complete); and the shape of the parameres are different.

Identification key to Liogenys from the southern South American Transition Zone and boundaries * *Note on dimorphism: Males have protarsi and mesotarsi flat (wide or not) and females have all tarsi cylindrical. 1. Length of antennal club two times longer than the funicle or almost so (Fig. 10D); head, pronotum, and scutellum dark purplish brown, elytra yellow (Fig. 10A); elytra glabrous; metatibia with inner margin not carinate in both sexes........................................................................................... Liogenys xanthocera Harold, 1869 - Length of antennal club as long as or slightly longer than funicle; head, pronotum, scutellum, and elytra unicolored, yellow or light brown, elytra sometimes slightly lighter in color than pronotum; elytra glabrous or with bristles; male with metatibia weakly or strongly carinate toward apex....................................................................2 2. Head more coarsely punctate than pronotum (Fig. 8B); pronotum wider than elytra at its humeral callus; galea with a single, long, hook-like tooth (Fig. 13G); antenna with 9 antennomeres; elytra with bristles............................................................................................... Liogenys maxillaricuspis Cherman, new species - Head as punctate as or less coarsely punctate than pronotum; pronotum as wide as or slightly narrower than elytra at its humeral callus; galea with four or five teeth (Fig. 13E), the superior tooth slightly larger than the other teeth; antenna with 9–10 antennomeres; elytra glabrous or with bristles................................................................... 3 3. Pygidium longer than wide (Fig. 1E) and always with bristles.................................................. 4 - Pygidium as wide as or wider than long, glabrous or with bristles................................................6 4. Antenna with 10 antennomeres; body length up to 8.5 mm; clypeus emargination in general deep, eventually shallow, narrower than the distance between eyes in both sexes; protibia with three teeth in both sexes; males with inner margin of metatibia carinate towards apex or not................................................................................ 5 - Antenna with 9 antennomeres; body length more than 8.5 mm; clypeus emargination noticeably shallow, females with emargination narrower than the distance between eyes, males with emargination as wide as the distance between eyes (Fig. 4F); protibia in males with two teeth and in females with three teeth; males with inner margin of metatibia carinate......................................................................................... Liogenys flavida Moser, 1918 5. Head as wide as or narrower than posterior margin of pronotum (Fig. 9C); clypeus as long as frons; vertex finely to coarsely punctate; males with inner margin of metatibia carinate straight towards apex; protarsi noticeably enlarged, protarsomere II wider than long.............................................................. Liogenys rufoflava Moser, 1918 - Head distinctly wider than posterior margin of pronotum (Fig. 1B); clypeus noticeably shorter than frons; vertex smooth (Fig. 1B); males with inner margin of metatibia not carinate; protarsi slightly enlarged, protarsomere II longer than wide..................................................................... Liogenys brachyclypeata Cherman, new species 6. Elytra and pygidium glabrous; males with inner margin of metatibia carinate towards apex; protarsomere II noticeably wider than long; apex of parameres lanceolate in a single plane (Figs. 3F, 9F)................... Liogenys flaveola Moser, 1924 - Elytra and pygidium with bristles; males with inner margin of metatibia weakly or not carinate, if carinate, carina never reaching the apex; protarsomere II as long as or longer than wide; apex of parameres lanceolate or not, if lanceolate, never in a single plane (Figs. 2F, 5F, 7F)................................................................................. 7 7. Elytra dull yellow, never pruinose or if so, not uniformly, pronotum unicolored with elytra; males with metatibial transverse carinae incomplete (Fig. 6E).............................................................................8 - Elytra light brown, uniformly pruinose, pronotum dark brown, darker than elytra; metatibia with one or two transverse carinae complete in both sexes (Fig. 5D.......................................................................... 9 8. Clypeal lateral margin with a blunt projection (Fig. 2D), convex; clypeal teeth well developed in both sexes, teeth acute; protibia with three teeth in both sexes; males with protarsomere and mesotarsomere II equally wide; spurs of metatibia uneven in shape (Fig. 2E)................................................................ Liogenys calcarata Frey, 1970 - Clypeal lateral margin not produced, straight or concave, clypeal teeth well developed only in males, teeth broad; protibia in males with two teeth and in females with three teeth; males with protarsomere II noticeably wider than mesotarsomere II; spurs of metatibia even in shape (Fig. 6C).................................. Liogenys lucialmeidae Cherman, new species 9. Elytra shiny or semiopaque, never pruinose; ventrites and pygidium with few bristles; males with two protibial teeth and females with three teeth.......................................................... Liogenys kuntzeni Moser, 1921 - Elytra opaque, noticeably pruinose (Fig. 7E); ventrites and pygidium with thick, scale-like bristles; both sexes with three protibial teeth.......................................................... Liogenys martinezi Cherman, new species, Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 48-49, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172, {"references":["Moser, J. (1918) Neue Amerikanische Melolonthiden. Stettiner Entomologische Zeitung, 79, 95 - 167.","Moser, J. (1924) Beitrag zur Kenntnis der Melolonthiden (Col.). Stettiner Entomologische Zeitung, 84, 114 - 136.","Frey, G. (1970) Beitrag zur kenntnis der Sudamerikanischen Melolonthiden (Col.). Entomologische Arbeiten aus dem Museum G. Frey, 21, 281 - 287.","Moser, J. (1921) Neue Melolonthiden von Mittel- und S ʾ d-Amerika. Stettiner Entomologische Zeitung, 82, 133 - 182. https: // doi. org / 10.1002 / mmnd. 192119210312"]}

Liogenys brachyclypeata Cherman, new species Figs. 1; 13B, D; 15 Type material. Holotype, male, labeled (CMNC): [white typeset] “ ARGENTINA, Mendoza / 40km. N. San Rafael, 1100m., 6.XII.1983 / L. Peña ”, [red typeset and handwritten] “ LIOGENYS / BRACHYCLYPEATA / HOLO-TYPE / Cherman M.A.”, genitalia mounted. Paratypes (42) labeled [yellow typeset and handwritten] “ LIOGENYS / BRACHYCLYPEATA / PARATYPE / Cherman M. A.”: Two males and seven females with the same data as the holotype (one male at DZUP (402736), four females at LEMQ, one male and three females at CMNC). Four female paratypes (CEMT): [white typeset] “ ARGENTINA: Rio Negro / Dto. Gral Roca, Catriel / 37º52,83’S 67º50,16’W / 4 Jan. 2005. 311 m / F. C. Ocampo ”. Seven males and 15 females (four males and 10 females CMNC, two males and four females DZUP (402737-42), one male and one female CEMT): [white typeset] “ ARGENTINA: Mendoza / Reserva Ecológica de Nacuñan / S34°02′42ʺ W67°54′34ʺ / Jan- 17-2003, 824 m / F.C.Ocampo, A.B. T.Smith”. One female (CMNC): [white typeset] “ ARGENTINA: Rio Negro / Dto Gral Roca, Catriel / 37°52′30ʺ S 67°50′10ʺ W / 4 Jan. 2005. 311 m / F.C. Ocampo ”, [white typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029592 ”. One female (CMNC): [white typeset] “ ARGENTINA: Rio Negro / Dto Gral Roca, Catriel / 37°52′30ʺ S 67°50′10ʺ W / 4 Jan. 2005. 311 m / F.C. Ocampo ”, [white typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NATURE / CMNEN 00029593 ”. One female (CMNC): [white typeset] “ ARGENTINA: Rio Negro / Dto Gral Roca, Catriel / 37°52’30’’S 67°50’10’’W / 4 Jan. 2005. 311 m / F.C. Ocampo ”, [white typeset with data matrix barcode] “Canadian Museum of / Musée canadien de la / NA-TURE / CMNEN 00029594 ”. One female (CMNC): [white typeset] “ ARGENTINA: Rio Negro / Colonia Catriel; Ruta Nac. 151 / 37º52’33’’S 67º49’50’’W / 315 m; 23 Jan. 2006; at light / F.C.Ocampo, E.Ruiz, G.Zalazar ”, [white typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3003257 ”, [white typeset] “A.B. T. SMITH DNA / BARCODE VOUCHER / SPECIMEN ABTS693 ” (DNA specimen stored in alcohol at -80 ºC). One female paratype (IADIZA) with the labels: [white typeset] “Arg. Rio Negro / Gral. Rocas / Cnia. Catriel 311 m / S. Roig / G. Flores / 04/I/05”, [white typeset] “ 37º52.83’ S / 67º56.16’W ”. One female (IADIZA): [white typeset] “ ARGENTINA: Chubut Biedma / Península de Valdes Ea San Pablo / 30 km SW Pto. Pirámides 42º 41’ / 43.2’’S 64º 10’ 42,7’’W 66 m / 06-I-2010 Coll: G. Flores, G. Cheli, / R. Carrara”. One female (IADIZA): [white typeset] “ ARGENTINA: Chubut / Península Valdez, Golfo Nuevo / Ea. San Pablo, 50 m / 27 Nov. 2005 ”, [white typeset with data matrix barcode] “SCARABS OF / SOUTHERN / SOUTH AMERICA / SSSA3002680 ”. Holotype and 22 paratypes deposited at CMNC. Seven paratypes deposited at DZUP. Four paratypes deposited at LEMQ. Six paratypes deposited at CEMT. Three paratypes deposited at IADIZA. Diagnosis. Body elongate, yellow; pronotum slightly darker; pronotum and elytra with bristles throughout, elytra pruinose on its margins (Fig. 1A); antenna with 10 antennomeres; frons very swollen and longer than clypeus; clypeal emargination deep, narrow, and rounded; teeth short, subparallel; clypeal lateral margin barely produced in males; almost straight in females; pronotum as long as one half its width, posterior margin narrower than anterior margin (Fig. 1B); elytra with only the sutural and the outer ridge distinguishable; protibia with three teeth (Fig. 1C); mesotibia and metatibia with one transverse carina complete (Fig. 1D); pygidium elongate, with long, decumbent bristles (Fig. 1E); males with protarsomeres slightly enlarged (Fig. 1D); paramere length at least three times the length of the apex; apex lanceolate in a single plane (Fig. 1F). Holotype. Male. Length 8.7 mm; width: 3.9 mm. Yellow. Head: distance between eyes three times the width of one eye; frons longer than clypeus, swollen; frontoclypeal surface with long bristles; clypeal emargination deep, narrow, and rounded; outer sides of anterior teeth subparallel; clypeal lateral margin weakly produced; distance between clypeal lateral projection and anterior tooth longer than the basal width of anterior tooth, distance between clypeal lateral projection and anterior margin of eye half the length of one eye, margin continuous, not forming a sharp angle between outer side of anterior teeth and clypeal lateral projection; distal maxillary palpomere, maximum width more than two times the apical width; fovea deep, very small, not reaching the transverse midline of the palpomere; antenna with 10 antennomeres, club lighter in color and equal in length to the funicle. Thorax: anterior margin of pronotum slightly produced medially, pronotum anteriorly slightly narrowed forming a depressed ring strongly impressed; disc with bristles throughout, mainly at the sides; pronotal disc with punctures coarse and sparse; posterior corners obtusely angulate; pronotal posterior margin narrower than anterior margin; hypomere with long bristles; mesepisternum, sides of metaventrite, and metacoxae with sparse bristles; distance between mesocoxae and metacoxae less than twice the metacoxa length; scutellum ogival, finely punctate throughout and with bristles. Elytra: disc shiny, pruinose along the margins, mainly on posterior and lateral surfaces; bristles on disc disposed randomly; elytral suture and elytron unicolored, elevated; elytral ridges undistinguishable, exept for the outer ridge. Legs: procoxa with bristles and punctate; protibia with three teeth, middle and apical teeth robust and equal in size, distance between basal and middle teeth slightly less than middle and apical teeth; mesotibia cylindric in cross section; mesotibial surface finely sculptured; one transverse carina almost complete; inner margin of metatibia not carinate towards apex; surface finely sculptured; metatibia with two transverse carinae, the apical carina almost complete; metatibial apical spurs subequal in size, the longer spur slightly longer than the diameter of the tibial apex, the smaller spur narrower and truncate; metatarsomere I equal in size to tarsomere II; protarsomeres I–IV slightly enlarged, protarsomere II elongate; claw bifid, symmetrical, superior tooth of claw longer and equal in width to the inferior tooth; distance between teeth equal to the length of inferior tooth. Abdomen: ventrites with uniform bristles on the disc and sides; propygidium with bristles; pygidium flat; subquadrate; elongate; pygidial width not exceeding distance between spiracles of propygidium; pygidial disc finely punctate and with long decumbent bristles throughout; pygidial apex subquadrate. Parameres: basal region wider than both sections of the parameres at maximum width, parameral split at 2/3 the length of the basal region; inner margins convex; paramere length at least three times the length of the apex; apex lanceolate-like shaped in one plane; parameres in lateral view straight, coplanar (Fig. 1G). Variation. Male paratypes. Length: 8.6–9.0 mm; width: 3.9–4.0 mm. As the holotype except in the clypeal lateral margin not produced. Female paratypes. Length: 8.0– 9.2 mm; width: 3.8–4.2 mm. As the holotype except in the clypeal lateral margin straight; pronotum shorter and coarsely punctate, and elytra less pruinose. Etymology. Adjective in the nominative singular. From Ancient Greek βραχύς (brakhús, “short”) + Latin clypeata (“relative to clypeus”). The name is referring to the clypeus shorter than the frons, a rare feature in Liogenys. Type locality. ARGENTINA, Mendoza, 40 km. N. San Rafael [34°10’40.9’’S 68°20’41.3’’W], 1100 m. Geographical distribution. ARGENTINA (Mendoza, Río Negro, Chubut). Remarks. Liogenys brachyclypeata new species resembles L. rufoflava in the body size and color, antenna with 10 antennomeres, elytra with bristles, protibia with three teeth in both sexes, and pygidium elongate and with bristles. Liogenys brachyclypeata new species differs from L. rufoflava (in parenthesis) mainly in the frons noticeable swollen and smooth, longer than clypeus (less swollen and punctate, as long as clypeus); clypeal emargination deep (shallow); pronotum shorter; lateral margin of pronotum with concavity more acute (unique among the Liogenys, which have broader concavity); pronotal posterior margin slightly narrower than the anterior margin and the head, with sharp corners (wider than head, subangulate corners); mesotibiae and metatibiae with one transverse carina complete (as in L. kuntzeni; in L. rufoflava males with transverse carina incomplete or only posteriorly); bristles of pygidium longer, and in males protarsomeres slightly enlarged (distinctly enlarged). The shape of the parameres closely resembles those of L. rufoflava, but in L. brachyclypeata new species they are longer, while the apex is thinner and narrower., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 49-51, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172

Liogenys flaveola Moser, 1924 Figs. 3, 11 A–B, 14 Liogenys flaveolus Moser, 1924: 122; Dallas 1926: 41 (teratologic case); Frey 1969: 47 (key). Liogenys flaveola: Blackwelder 1944: 227 (checklist); Evans 2003: 209 (checklist); Evans & Smith 2009: 177 (checklist); Krajčík 2012: 144 (checklist); Cherman et al. 2017: 4 (generic history). Liogenys kadleci Frey, 1970: 281; Evans 2003: 210 (checklist); Evans & Smith 2009: 178 (checklist); Krajčík 2012: 145 (checklist); Cherman et al. 2016: 759 (systematics); Cherman et al. 2017: 5, 6 (generic history, taxonomy); new synonym. Type material. Male holotype of Liogenys flaveola (ZMHB): [white, typeset, outlined] “RCA. ARGENTINA / Gob. Chubut / 190 [space] / C. Bruch ”, [white handwritten] “Camarones / J. Wiederrecht ”, [white handwritten] “ Liogenys / flaveolus / Typen Mos”, [red typeset] “Typus”, [white typeset] “ Liogenys / flaveolus / Mos.”. Male holotype of Liogenys kadleci (NMHB): [white, typeset] “ ARGENTINA / Villa Regina / Rio Negro / 20.XI.63 leg. Pena”, [red typeset] “ TYPE ”, [white, typeset and handwritten] “Type / Liogenys / kadleci / n. sp. / det. G. Frey, 1970 ”, genitalia mounted. Male paratype (ZMHB): [white, typeset] “ ARGENTINA / Villa Regina / Rio Negro / 20.XI.63 leg. Pena”, [red typeset] “ PARATYPE ”, [white, typeset and handwritten] “Type / Liogenys / kadleci / n. sp. / det. G. Frey, 1970 ”. Male paratype (CMNC): [white, typeset] “ ARGENTINA / Villa Regina / Rio Negro / 20.XI.63 leg. Pena”, [red typeset] “ PARATYPE ”, [white, typeset and handwritten] “ Liogenys / kadleci / det. G. Frey, 1970 / n sp”, [white, typeset] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.”, [white, typeset] “Southern Neotropical Scarabs / database # AS 2614382 / Liogenys kadleci / Frey, 1970 ³ / DET: A.B. T. SMITH 2008” Male paratype (CMNC): [white, typeset] “ ARGENTINA / Villa Regina / Rio Negro / 20.XI.63 leg. Pena”, [red typeset] “ TYPE ”, [white, typeset and handwritten] “Type / Liogenys / kadleci / det. G. Frey, 1970 / n sp”, [white, typeset] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.”, [white, typeset] “Southern Neotropical Scarabs / database # AS 2614381 / Liogenys kadleci / Frey, 1970 ³ / DET: A.B. T. SMITH 2008”. Non-type material (67). ARGENTINA. Buenos Aires: Without locality and collector, 1 male (MLPA); Monte Hermoso, XII.1955, A. Martínez, 1 female (DZUP); Puán, Felipe Solá, XII.1953, A. Martínez, 1 male (CMNC); Mendoza: without locality, 9.XI.1987, L. Peña, 1 male (CEMT); Pata Mora (36 km W), 36º58’20’’S 69º6’19’’W, 7.I.2003, 1145 m, F.C. Ocampo & A.B. T. Smith, 1 male (CMNC); San Rafael, 6.XII.1988, L. Peña, 1 female (INPA); Río Negro: without locality, date and collector, 1 male (MLPA); General Roca, 24.XI.1920, A. Wetmore, 1 male (CASC); Villa Regina, XII.1961, A. Maller, 3 females (1 CMNC, 2 DZUP); 20.XI.1963, L. Peña, 1 male (NHMB); San Antonio Oeste, 15.XII.1966, E.I. Schlinger & M.E. Irwin, at lights, 7 males and 10 females (CASC); San Antonio Oeste, XII.1992, A. Martínez, at lights, 5 males (CMNC); San Antonio Oeste, Los Grutas, XII.1992, A. Martínez, 2 males and 4 females (CMNC); Valcheta, XII.1992, A. Martínez, 1 male (CMNC); Chubut: Península de Valdés, Golfo Nuevo, Estancia San Pablo, 42°41’42’’S, 64°10’46’’W, 27.XI.2005, 50 m, 15 males and 3 females (CMNC); Península de Valdés, Golfo Nuevo, Estancia San Pablo, 42°41’42’’S, 64°10’46’’W, 15.XII.2005, 50 m, G. Cheli, 1 male, 1 female (CMNC); Puerto Madryn (13 KM N), 14.XII.1966, M.E. Irwin & E.I. Schlinger, 1 female (CASC); Puerto Pirámides, XII.1992, 44 males and 39 females (CMNC). Diagnosis. Body elongate, sides parallel in males, wider on posterior third in females; dull yellow, pronotum shiny, unicolored with elytra; elytra shiny or semiopaque, glabrous (Fig. 3A); clypeal emargination shallow, wide and rounded; clypeal lateral margin concave; antenna with 10 antennomeres; pronotal posterior corners subangulate to rounded (Fig. 3B); protibia with three teeth equally spaced; mesotibia quadrate in cross section (Fig. 3D); pygidium flat; subtrapezoidal; pygidial disc smooth and glabrous (Fig. 3C); in males inner margin of metatibia carinate towards apex and curved frontwards; the smaller metatibial spur wider and ½ the length of the larger, apex slightly curved frontwards (Fig. 3E), protarsomere II very wide; inner margins of parameres convex; four times the length of the apex; apex lanceolate in a single plane (Fig. 3F). Redescription. Length: 9.0– 10.7 mm; width: 4.7–4.9 mm. Dull yellow. Head. distance between eyes slightly more than twice the width of one eye; frons swollen, as long as the clypeus; clypeal emargination shallow, wide and rounded; outer sides of anterior teeth subparallel or follow the lateral margin of clypeus; clypeal lateral margin concave, clypeus in males (Fig. 11A) with teeth longer and broader than in females (Fig. 11B); distal maxillary palpomere, maximum width slightly wider than the apex; fovea deep, very small, only at the base; antenna with 10 antennomeres, club lighter in color than funicle. Thorax: anterior margin of pronotum straight; pronotal disc with punctures coarse and sparse; posterior corners subangulate to rounded; hypomere with long bristles; mesepisternum, sides of metaventrite and metacoxae with bristles, disc of metaventrite with spine-like bristles; distance between mesocoxae and metacoxae twice the metacoxa length; scutellum ogival, finely punctate at the sides. Elytra: shiny, glabrous; elytral suture and elytron unicolored, slightly elevated; inner two pairs of elytral ridges more defined than the outer ridges. Legs: procoxa with bristles and punctate; protibia with three teeth, middle and apical teeth equal in size, basal teeth slightly shorter; three teeth equally spaced; mesofemur with thick, erect bristles along the inferior margin; mesotibia quadrate in cross section, mesotibial surface almost smooth; mesotibia with two transverse carinae present posteriorly, in females the apical one complete; metafemur with thick, erect bristles along the inferior margin; inner margin of metatibia carinate towards apex and curved frontwards; surface finely sculptured, almost smooth; metatibia with two transverse carinae present posteriorly; metatibial apical spurs of different lengths, the longer spur equal to the diameter of the tibial apex, males with the smaller spur half the length of the longer spur, apex truncate, slightly curved frontwards, females with longer, wider spurs and apex rounded; metatarsomere I slightly shorter and wider apically than metatarsomere II; protarsomeres and mesotarsomeres I–IV enlarged, protarsomere II very wide; protarsomeres almost two times wider than the mesotarsomeres; claw bifid, symmetrical, superior tooth of claw longer and as wide as the inferior; distance between teeth equal to the length of inferior tooth. Abdomen: ventrites uniformly with bristles; propygidium with a few bristles; pygidium flat; subtrapezoidal; pygidial width not exceeding distance between spiracles of propygidium; pygidial disc smooth and glabrous; pygidial apex subquadrate. Parameres: basal region slightly wider than both sections of the parameres at its maximum width, parameral split at 2/3 the length of the basal region; inner margins convex; apex lanceolate in a single plane; parameres in lateral view straight, not coplanar (Fig. 3G). Type locality. Liogenys flaveola: ARGENTINA. Chubut, Camarones [44°47’56.8’’S, 65°42’36.2’’W]. Liogenys kadleci: ARGENTINA. Río Negro, Villa Regina [39°06’01.2’’S, 67°05’19.6’’W]. Geographical distribution. ARGENTINA (Buenos Aires, Mendoza, Neuquén, Río Negro, Chubut). Remarks. Liogenys flaveola is most similar to L. calcarata (see remarks of L. calcarata). We compared the holotypes of L. flaveola and L. kadleci, the former only differs in the outer sides of anterior clypeal teeth following the lateral margin, and the pronotal posterior corners are rounded. Unfortunately, the parameres of the holotype of L. flaveola are lost. Since the non-type material of L. kadleci showed intraspecific variations that match with the holotype of L. flaveola, we concluded that L. kadleci is a junior synonym of L. flaveola. Dallas (1926) described the teratology found in a female specimen of L. flaveola from Río Negro (Argentina), collected in 1922 by Carlos Bruch. It consisted in the right protarsus bifid, ramified from the second tarsomere, with a normal pair of claws on the inner tarsus and a double pair of claws on the outer one. This kind of deformation is called melomelia, the presence of more than the usual number of limbs, or their ramification (Dallas 1926). We were unable to find the Dallas collection., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 54-56, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172, {"references":["Moser, J. (1924) Beitrag zur Kenntnis der Melolonthiden (Col.). Stettiner Entomologische Zeitung, 84, 114 - 136.","Dallas, E. D. (1926) Un Liogenys flaveolus Moser, monstruoso. Revista de la Sociedad Entomologica Argentina, 1, 41 - 42.","Frey, G. (1969) Bestimmungstabelle und revision der gattung Liogenys Guerrez (Coleoptera-Melolonthinae-Macrodactylini). Entomologische Arbeiten aus dem Museum G. Frey, 20, 38 - 64.","Blackwelder, R. E. (1944) Checklist of the Coleopterous Insects of Mexico, Central America, The West Indies and South America. Part 2. Smithsonian Institution, Washington, D. C., pp. 189 - 265. https: // doi. org / 10.5479 / si. 03629236.185.2","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211 (1), 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3. Papers in Entomology, 1, 1 - 344.","Krajcik, M. (2012) Checklist of the World Scarabaeoidea. 5 th Edition. Animma. x, Plzen, 278 pp.","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031","Frey, G. (1970) Beitrag zur kenntnis der Sudamerikanischen Melolonthiden (Col.). Entomologische Arbeiten aus dem Museum G. Frey, 21, 281 - 287.","Cherman, M. A., Moron, M. A. & Almeida, L. M. (2016) Phylogenetic relationships within Diplotaxini Kirby (Coleoptera: Melolonthidae: Melolonthinae) with emphasis on Liogenys Guerin-Meneville. Systematic Entomology, 41, 744 - 770. https: // doi. org / 10.1111 / syen. 12188"]}

Liogenys xanthocera Harold, 1869 Figs. 10, 13F, 14 Liogenys pallidicornis Blanchard, 1851: 168; Lacordaire 1855: 269 (generic redescription); Frey 1969: 42 (key). Liogenys xanthocerus Harold 1869a: 123 (new replacement name for L. pallidicornis); Harold 1869b: 1140 (checklist, synonymy); Kolbe 1907: 5 (checklist); Bruch 1911: 200 (checklist); Dalla Torre 1913: 318 (checklist); Frey 1969: 42 (key; junior synonym of L. pallidicornis [sic]); Krajčík 2012: 145 (checklist); Cherman et al. 2017: 4 (generic history). Liogenys xanthoceris [sic]: Frey 1969: 57 (synonym of L. pallidicornis [sic]). Liogenys xanthocera: Blackwelder 1944: 227 (checklist); Evans 2003: 215 (checklist); Evans & Smith 2009: 183 (checklist). Type material. Male lectotype of Liogenys pallidicornis here designated (MNHN): [white handwritten] “6051 / 34”, [light green, typeset and handwritten] “Patagonie / d’Orbigny / MUSÉUM PARIS ”, [green handwritten] “ L. pallidicornis. / Cat. Mus. / Baie de San Blas (Patagonie) / M. D’Orbigny”, [white typeset] “Southern Neotropical Scarabs / database # AS 2615155 / Liogenys xanthocera / Harold 1869 ³ / DET:A.B. T. SMITH 2008”, [red, outlined, typeset and handwritten] “ LIOGENYS / PALLIDICORNIS / BLANCHARD, 1851 / LECTOTYPE / Des. Cherman M. A.”. Male paralectotype (MNHN) bearing the labels: [white handwritten] “6051 / 34”, [white handwritten] “1052”, [light green typeset and handwritten] “Patagonie / d’Orbigny / MUSÉUM PARIS ”, [white typeset] “Southern Neotropical Scarabs / database # AS 2615156 / Liogenys xanthocera / Harold 1869 ³ / DET: A.B. T. SMITH 2008”, [red, outlined, typeset and handwritten] “ LIOGENYS / PALLIDICORNIS / ³ / BLANCHARD, 1851 / PARALECTOTYPE / Des. Cherman M. A.” Non-type material (14). ARGENTINA. Patagonia: without locality, date, and collector, 1 female (ZMHB); Buenos Aires: Puán, Felipe Solá, XII.1944, A. Martínez, 2 males (CMNC); Puán, Felipe Solá, II.1946, A. Martínez, 4 males (CMNC); Puán, Felipe Solá, A. Martínez, XII.1951, 3 males (MZSP); Puán, Felipe Solá, XI.1959, A. Martínez, 1 male (MZSP); Puán, Felipe Solá, XII.1960, A. Martínez, 2 males (MZSP); Estancia Barraú, 30 km SW Villa Iris, XI.1946, A. Martínez, 1 male (CMNC). Diagnosis. Body elongate; body, head, and pronotum in general dark purple; elytra glabrous, shiny yellow, sides subparallel (Fig. 10A); clypeal emargination deep, narrow and rounded; clypeal lateral margin convex and produced, obtuse angle between outer side of anterior teeth and clypeal lateral projection (Fig. 10C); antenna with 10 antennomeres, yellow, club nearly twice as long as the funicle; posterior corners subangulate, obtuse; mesotibia cylindrical in cross section (Fig. 10D); pygidium slightly convex; wide, subtrapezoidal; pygidial disc glabrous or with unconspicous bristles throughout (Fig. 10E); males with inner margin of metatibia not carinate; transverse carina weakly defined; protarsomere II elongate; parameres narrowed at the midline; splitted after the midline; inner margins straight; apex spatulate (Fig. 10F). Redescription. Length: 7.3–7.6 mm; width: 3.8–3.9 mm. Dark purple in body, head, and pronotum, elytra dull yellow. Head. distance between eyes more than three times the width of one eye; clypeal emargination deep, narrow, and rounded; outer sides of anterior teeth parallel; clypeal lateral margin convex and produced, forming a rounded projection; distance between clypeal lateral projection and anterior tooth longer than the basal width of anterior tooth, distance between clypeal lateral projection and anterior margin of eye longer than one eye; obtuse angle between outer side of anterior teeth and clypeal lateral projection; antenna with 10 antennomeres, club nearly twice as long as and lighter in color than funicle. Thorax: anterior margin of pronotum straight; pronotal disc with punctures coarse, shallow and sparse; posterior corners subangulate, obtuse; hypomere, mesepisternum, sides of metaventrite, and metacoxae with sparse, long bristles; distance between mesocoxae and metacoxae less than twice the metacoxa width; scutellum ogival to rounded, punctate throughout. Elytra: shiny, glabrous; elytral suture and elytron unicolored, elevated; two inner pair of elytral ridges more defined than the other two ridges. Legs: procoxa with bristles; protibia with three teeth, middle and apical teeth equal in size, all three teeth equally spaced; mesotibia cylindrical in cross section, mesotibia dorsally smooth with two transverse carinae, the apical carina complete; inner margin of male metatibia not carinate; surface finely sculptured; metatibia with one transverse carina present posteriorly and weakly defined dorsally; metatibial apical spurs of equal lengths, equal to the diameter of the tibial apex; metatarsomere I shorter than metatarsomere II; in males protarsomeres and mesotarsomeres I–IV enlarged, protarsomere II long; protarsomeres and mesotarsomeres equally enlarged; claw bifid, symmetrical, superior tooth of claw almost three times longer and narrower than the inferior tooth; distance between teeth longer than the length of inferior tooth. Abdomen: ventrites with sparse bristles; propygidium glabrous; pygidium slightly convex; subtrapezoidal; pygidial width not exceeding distance between spiracles of propygidium; pygidial disc with inconspicuous bristles, finely and very sparsely punctate; pygidial apex subquadrate. Parameres: basal region wider than both sections of the parameres at its maximum width, parameres narrowed at the midline; parameral split after the midline; inner margins straight; apex spatulate, angulate; parameres straight or slightly convex in lateral view (Fig. 10G). Type locality. ARGENTINA. Baie de San-Blas (Patagonie) [Buenos Aires, Bahia de San Blas, 40°33’8’’S 62°14’4’’W]. Geographical distribution. ARGENTINA (Buenos Aires). Remarks. Liogenys xanthocera is easily recognized among the other Liogenys by its characteristic dark body, pronotum, and scutellum contrasting with the elytra dull yellow; and by the club nearly twice the length of the funicle. The teeth of the galea are reduced (Fig. 13F), as in L. sinuaticeps Moser (Cherman et al. 2016, 2017). The specimen from 30 km SW Villa Iris, Puán, Buenos Aires [38°21’15.8’’S 63°20’37.4’’W] shows variations in the body color, entirely dull yellow, the lateral margin of clypeus more produced, and the parameres strongly convex. As we have only this specimen with these single features, we presume that those differences are only intraspecific variations, needing further material to determine if it is a different species. The lectotype (Fig. 10B) was collected Bahía San Blas, Buenos Aires, Argentina during the period from 14 January–18 February 1829 by A.D. d’Orbigny. The region around the nearby village of “Patagones” (now known as Viedma, Río Negro) was collectively called Patagonia (Patagonie) at the time (see Smith 2017). The “1052” code number on the paralectotype corresponds with an entry in the original field notes of d’Orbigny (available from the MNHN library). D’Orbigny made these observations (text transcribed and translated): “we found this species in great numbers in January and February. They emerged from the earth every day, in numbers. We never saw them during the day, when they were hiding underground”. Blanchard (1851) described two species in Liogenys with the epithet “ pallidicornis ”, the first one in page 167 and the other in the next page. Harold (1869) corrected this homonym by replacing the name of the second one described, as L. xanthocera. The first one described, Liogenys pallidicornis Blanchard, 1851 is a valid name (Cherman et al. 2017)., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 74-82, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172, {"references":["Blanchard, E. (1851) Museum d'Histoire Naturelle de Paris. Catalogue de la Collection Entomologique. Classe des Insectes. Ordre des Coleopteres. Tome I. Gide et Baudry, Paris, 240 pp.","Frey, G. (1969) Bestimmungstabelle und revision der gattung Liogenys Guerrez (Coleoptera-Melolonthinae-Macrodactylini). Entomologische Arbeiten aus dem Museum G. Frey, 20, 38 - 64.","Harold, E. F. von (1869 a) Abanderungen vergebener Namen. Coleopterologische Hefte, 1, 122 - 125.","Harold, E. F. von (1869 b) Scarabaeidae. In: Gemminger, M. & von Harold, E. F. (Eds.), Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus, Vol. 4. E. H. Gummi, Munich, pp. 976 - 1346.","Kolbe, H. (1907) Coleopteren. In: Ergebnisse der Hamburger Magalhaensische Sammelreise 1892 / 93. II Band. Arthropoden. L. Friederichsen & Co., Hamburg, pp. 1 - 125.","Bruch, C. (1911) Catalogo sistematico de los Coleopteros de la Republica Argentina. Revista del Museo de la Plata, 17, 181 - 225. https: // doi. org / 10.5962 / bhl. title. 8799","Dalla Torre, K. W. von (1913) Scarabaeidae: Melolonthinae IV. Pars 50. In: S. Schenkling (Ed.), Coleopterorum catalogus. Vol. XX. Junk, W., Berlin, pp. 291 - 450.","Krajcik, M. (2012) Checklist of the World Scarabaeoidea. 5 th Edition. Animma. x, Plzen, 278 pp.","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031","Blackwelder, R. E. (1944) Checklist of the Coleopterous Insects of Mexico, Central America, The West Indies and South America. Part 2. Smithsonian Institution, Washington, D. C., pp. 189 - 265. https: // doi. org / 10.5479 / si. 03629236.185.2","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211 (1), 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3. Papers in Entomology, 1, 1 - 344.","Moser, J. (1924) Beitrag zur Kenntnis der Melolonthiden (Col.). Stettiner Entomologische Zeitung, 84, 114 - 136.","Moser, J. (1918) Neue Amerikanische Melolonthiden. Stettiner Entomologische Zeitung, 79, 95 - 167.","Moser, J. (1921) Neue Melolonthiden von Mittel- und S ʾ d-Amerika. Stettiner Entomologische Zeitung, 82, 133 - 182. https: // doi. org / 10.1002 / mmnd. 192119210312","Frey, G. (1970) Beitrag zur kenntnis der Sudamerikanischen Melolonthiden (Col.). Entomologische Arbeiten aus dem Museum G. Frey, 21, 281 - 287.","Cherman, M. A., Moron, M. A. & Almeida, L. M. (2016) Phylogenetic relationships within Diplotaxini Kirby (Coleoptera: Melolonthidae: Melolonthinae) with emphasis on Liogenys Guerin-Meneville. Systematic Entomology, 41, 744 - 770. https: // doi. org / 10.1111 / syen. 12188","Smith, A. B. T. (2017) Southern South American Trogidae (Coleoptera): verification of type specimens and notes on geographic distribution. Revista Chilena de Entomologia, 43, 81 - 103."]}

Liogenys kuntzeni Moser, 1921 Figs. 5, 11 E–F, 14 Liogenys kuntzeni Moser, 1921: 139; Blackwelder 1944: 227 (checklist); Gutiérrez 1949: 20 (geographic records); Gutiérrez 1951: 132, 134 (key, redescription); Frey 1969: 48 (key); Evans 2003: 210 (checklist); Evans & Smith 2009: 178 (checklist); Krajčík 2012: 145 (checklist); Cherman et al. 2016: 759 (systematics); Cherman et al. 2017: 4, 16 (generic history, taxonomy). Type material. Male holotype of Liogenys kuntzeni (ZMHB): [white typeset] “ Chili ”, [white handwritten] “ Liogenys / kuntzeni / Typen / Mos”, [red typeset] “Typus”, [white typeset] “ Liogenys / kuntzeni / Mos.”, genitalia mounted. Non-type material (53). CHILE. Valparaíso: Algarrobo, I.1987,L. Peña, 2males (INPA); 21.VII.1951, [33°22’S, 71°40’W], Kuschel & L.E. Peña, 1 male (FMNH); NW Algarrobo. Mirasol, 22.I.1978, L. Peña, 1 male (INPA); NW Algarrobo. Mirasol, 12.I.1975, [33°19’26.93’’S, 71°38’48.88’’W], NW Algarrobo. Mirasol, 12.I.1975, P. Vidal, 6 males and 1 female; NW Algarrobo. Mirasol, 16.I.1975, 4 males and 2 females (FMNH); Horcones, [32°43’12’’S, 71°29’24’’W], 19. II.1963, L.E. Peña, 2 males (FMNH), Viña del Mar, [33°1’30.78’’S, 71°33’3.73’’W], 1–31.X.1975, Herrera, 3 males (UMCE); Zapallar, Cachagua, II.1977, Alcaino, 1 male (MNNC); Llay-Llay, [33°51’S, 70°58’W], 1–30.XI.1934, without collector; 1 male (MNHN); San Antonio, [32°53’S, 71°15’W], without date and collector, 1 male (NHMUK); Santiago: San Cristobal, without date, Zapata, 3 males (UMCE); La Dehesa, I.1975; without collector, 1 specimen; El Principal, IX.1953, L.E. Peña, 1 female (FMNH); [Refugio] Lo Valdés. Cajón del Maipo, 16.I.2005, 2200 m, Ramirez, 1 male (MNNC); Maule: Curicó, Cajón de Teno [near Quilpoco], 3.I.1980, 2000 m, J. R. Barriga, 2 males (USNM); Talca, Rio Maule, 2.X–2.XI.1956, 1400 m, L.E. Peña, 1 male (FMNH); Los Cipreses [Romeral], 14.I.1968, 1000 m, L.E. Peña, 2 males, 1 female (CMNC); Ñuble: 50 km E San Carlos [Punilla Provence, near San Fabián], 26.XII.1950, Ross & Michelbacher, 1 male (NHMB); Biobío: Laguna del Laja [Los Ángeles], 16.I.1987, L. Peña, 3 males and 1 female (INPA); 2.I.1987, Perez de Arce, 1 female (MNNC); Abanico, I. 1948, 800 m, without collector, 3 males (CMNC, MNNC, NHMB); XI.1948, without collector, 1 male (MNNC); 10.I.1948, G. Kuschel, 1 male (MZUC); Los Barros, I.1948, 1800 m, G. Kuschel, 1 male. (MNNC); Concepción, 15.XII.1962, Fetes, 1 male (CNCI); Concepción. Hualqui, 23.XII.1944, C. Junge, 3 males (USNM). Diagnosis. Body light brown, elongate, sides parallel in males, slightly wider on posterior third in females; pronotum slightly darker and shinier than elytra (Fig. 5A); elytra bearing inconspicuous bristles, more visible on the outer ridge and lateral margins; clypeal emargination deep, subangulate, and very wide; clypeal lateral margin straight in males, slightly convex in females; antenna with 10 antennomeres; posterior corners subangulate (Fig. 5B); males with two protibial teeth, sometimes with a vestigial basal tooth such as in females; mesotibia cylindrical in cross section and somewhat flattened, males with one transverse carina complete (Fig. 5D), females with two transverse carinae, the apical carina complete; pygidium slightly convex; subtrapezoidal, pygidial disc smooth and with sparse, random bristles (Fig. 5C); males with inner margin of metatibia weakly carinate present only on apical portion (Fig. 5E); parameres with apex lanceolate, forming and ear-like wing laterally (Fig. 5F). Redescription. Length: 8.8–9.9 mm; width: 4.5–5.0 mm. Light brown. Head. distance between eyes slightly more than twice the width of one eye; frons swollen, equal in length to clypeus; clypeal emargination deep, wide and subangulate, in males almost as wide as the distance between eyes; outer sides of anterior teeth subparallel or following the lateral margin of clypeus, lateral margin straight in males, convex in females; in males teeth of clypeus longer and more defined (Fig. 11E) than in females (Fig. 11F); distal maxillary palpomere, maximum width less than two times the apical width; fovea deep, not reaching the transverse midline of the palpomere; antenna with 10 antennomeres, club lighter in color than funicle. Thorax: anterior margin of pronotum slightly produced medially; pronotum anteriorly slightly narrowed forming a depressed ring; pronotal disc with punctures coarse and sparse; posterior corners subangulate; hypomere with long and short bristles; mesepisternum, sides of metaventrite and metacoxae with bristles; distance between mesocoxae and metacoxae twice that of metacoxa width; scutellum rounded, smooth or with a few punctures. Elytra: shiny, with inconspicuous bristles disposed randomly; elytral suture and elytron unicolored, elevated; all four elytral ridges barely noticeable. Legs: procoxa barely scaly and punctate; males with two protibial teeth, sometimes with a weakly defined third basal tooth such as in females, middle and apical teeth equal in size, all three teeth equally spaced; mesotibia cylindrical and somewhat flattened in cross section, mesotibial surface finely sculptured; males with one mesotibial transverse carina only posteriorly, females with two transverse carinae, the apical carina complete; inner margin of metatibia not carinate in males or females; surface finely sculptured; metatibia with two transverse carinae, the apical carina complete, more defined in females; metatibial apical spurs of different lengths, the longer spur equal to the diameter of the tibial apex; metatarsomere I in males slightly shorter and equal in width to metatarsomere II, in females metatarsomere I and metatarsomere II approximately equal in length; in males protarsomeres and mesotarsomeres I–IV enlarged, protarsomere II elongate; mesotarsomeres less enlarged than the protarsomeres; claw bifid, symmetrical, superior tooth of claw longer and slightly wider than the inferior tooth; distance between teeth slightly longer than the length of inferior tooth. Abdomen: ventrites with a few bristles on the disc; propygidium with bristles; pygidium slightly con-vex; subtrapezoidal, as long as it is wide; pygidial width not exceeding distance between spiracles of propygidium; pygidial disc smooth and with a few scattered bristles; pygidial apex subquadrate. Parameres: basal region as wide as both sections of the parameres at its maximum width, parameral split at 2/3 the length of the basal region; inner margins convex; apex lanceolate-like, base of apex with outer margin widely expanded, forming an ear-like flap; parameres in lateral view straight, not coplanar (Fig. 5G). Type locality. CHILE. Geographical distribution. CHILE (Valparaiso, Santiago, Maule, Ñuble, Biobío). Remarks. Liogenys kuntzeni differs from the other Liogenys in the following combination of characters: clypeal emargination angulate (sometimes worn out and if so, subangulate); metatibia with one transverse carina complete in males and females (different from L. flaveola and L. calcarata, males with carina absent or incomplete, respectively); males with two protibial teeth instead of three (as in L. flavida), but in some specimens a vestigial third basal tooth was found; and inner margin of metatibia very weakly carinate, present on apical portion. Liogenys kuntzeni could be also misidentified with L. lucialmeidae new species and L. martinezi new species (see remarks of both new species). Frey (1969) mentioned Río Negro (Argentina) as a locality record, but this record was not verified. Evans (2003) and Evans & Smith (2009) excluded the Argentina record in their catalog, because it is not reliable and needs to be verified with voucher specimens. An Argentinian male specimen from Neuquén, Pilmatué [Laguna Pimatué], Co. [Cerro] Huayilon [-38.25, -70.3833333] 1500 m Gentili, I.1968) was found at the CMNC. Many features match with L. kuntzeni, including the protibia with two teeth, metatibia with one transverse carina complete; and the apex of the parameres with ear-shaped expansions. However, in this specimen these expansions are shorter than expected in L. kuntzeni, which is more similar to L. martinezi new species, but without the pruinose elytra. Due to the uncertainty we decided not to consider this specimen as a new record of L. kuntzeni. The eastern strip of the Andean Maule province is located along western Neuquén (Argentina), so it should not be surprising to find L. kuntzeni in this region. More collecting events are required to confirm if L. kuntzeni occurs there., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 59-62, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172, {"references":["Moser, J. (1921) Neue Melolonthiden von Mittel- und S ʾ d-Amerika. Stettiner Entomologische Zeitung, 82, 133 - 182. https: // doi. org / 10.1002 / mmnd. 192119210312","Blackwelder, R. E. (1944) Checklist of the Coleopterous Insects of Mexico, Central America, The West Indies and South America. Part 2. Smithsonian Institution, Washington, D. C., pp. 189 - 265. https: // doi. org / 10.5479 / si. 03629236.185.2","Gutierrez, R. (1949) Notas sobre Scarabaeidae neotropicos (Coleoptera Lamellicornia). Anales de la Sociedad Cientifica Argentina, 148, 9 - 35.","Gutierrez, R. (1951) El genero Liogenys en Chile. Revista Chilena de Entomologia, 1, 129 - 145.","Frey, G. (1969) Bestimmungstabelle und revision der gattung Liogenys Guerrez (Coleoptera-Melolonthinae-Macrodactylini). Entomologische Arbeiten aus dem Museum G. Frey, 20, 38 - 64.","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211 (1), 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3. Papers in Entomology, 1, 1 - 344.","Krajcik, M. (2012) Checklist of the World Scarabaeoidea. 5 th Edition. Animma. x, Plzen, 278 pp.","Cherman, M. A., Moron, M. A. & Almeida, L. M. (2016) Phylogenetic relationships within Diplotaxini Kirby (Coleoptera: Melolonthidae: Melolonthinae) with emphasis on Liogenys Guerin-Meneville. Systematic Entomology, 41, 744 - 770. https: // doi. org / 10.1111 / syen. 12188","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031"]}

Liogenys calcarata Frey, 1970 Figs. 2, 13E, 14 Liogenys calcaratus Frey, 1970: 283; Krajčík 2012: 144 (checklist). Liogenys calcarata: Evans 2003: 207 (checklist); Evans & Smith 2009: 176 (checklist); Cherman et al. 2016: 759 (systematics); Cherman et al. 2017: 16 (taxonomy). Type material. Male holotype of Liogenys calcarata (NMHB): [white, typeset] “ ARGENTINA / 20 km N. La Rioja / 24.9.68 leg. Pena”, [red typeset] “ TYPE ”, [white typeset and handwritten] “Type / Liogenys / calcaratus / n. sp. / det. G. Frey, 1970 ”, genitalia mounted. One male paratype (NHMB), five males and one female paratypes (AMNH) with the same data of the holotype. Male paratype (ZMHB): [white, typeset] “ ARGENTINA / 20 km N. La Rioja / 24.9.68 leg. Pena”, [red typeset] “ PARATYPE ”, [white, typeset and handwritten] “ Liogenys / calcaratus / n. sp. / det. G. Frey, 1970 ”. Four male and two female paratypes (CMNC): [white, typeset] “ ARGENTINA / 20 km N. La Rioja / 24.9.68 leg. Pena”, [yellow handwritten and typeset] “ LIOGENYS / CALCARATUS / FREY / PARATYPE ”, [white, typeset and handwritten] “ Liogenys / calcaratus / det. G. Frey, 1970 / n sp”, [white, typeset] “H. & A. HOWDEN / COLLECTION / ex. A. Martinez coll.”. Non-type material (12). ARGENTINA. San Juan: El Baldecito, Valle Fertil, XII.1964, A. Martínez, 1 male (CMNC); Buenos Aires: Estancia Barraú (30 km SW Villa Iris), XII.1946, A. Martínez, 1 male (CMNC); Puán, Felipe Solá, XII.1953, A. Martínez, 1 female (CMNC); San Luis: Arizona (18 km S), 250 m, 18–23.I.1982, H. & A. Howden, 1 female (CMNC); Mendoza: San Rafael, 6.XII.1988, L. Peña, 4 females (INPA); Río Negro: Coronel Gómez, I.1958, A. Martínez, 2 females (NHMUK); Chubut: Dolavon, Angostura, 1. V.2011, 43°18’36’’S, 65°42’36’’W, D. Rojas Lanus, 1 female (VMDC). Diagnosis. Body elongate; dull yellow (Fig. 2A), pronotum shiny, unicolored with elytra; elytra shiny or semiopaque, with bristles, inconspicuous bristles throughout the elytron and also on frontoclypeal surface; clypeal emargination deep, wide and rounded; clypeal lateral margin convex, blunt projected; antenna with 10 antennomeres; pronotal posterior corners rounded (Fig. 2B); protibia with three teeth, distance between basal and middle teeth longer than the distance between middle and apical teeth; mesotibia quadrate in cross section (Fig. 2D); pygidium flat; subtrapezoidal; pygidial disc smooth, inconspicuous bristles throughout, bristles longer towards the apex (Fig. 2C); in males inner margin of metatibia weakly carinate towards apex; metatibial apical spurs of different lengths and shape, the longer spur equal to the diameter of the tibial apex, the shorter spur wider and projected laterally (Fig. 2E); protarsomere II very wide; inner margins of parameres concave; parameres three times the length of the apex; apex lanceolate, outer margins expanded in a plane below at the base (Fig. 2F). Redescription. Length: 8.2–9.5 mm; width: 3.4–4.8 mm. Dull yellow. Head. Distance between eyes twice the width of one eye; frons swollen, longer than the clypeus, frontoclypeal surface with inconspicuous bristles; clypeal emargination deep, wide and rounded; outer sides of anterior teeth subparallel; clypeal lateral margin convex, blunt projected; canthus exceeding the outer margin of the eye; distal maxillary palpomere, maximum width slightly wider than the apex; fovea deep, very small, only at the base; antenna with 10 antennomeres, club lighter in color than funicle. Thorax: anterior margin of pronotum slightly produced medially; pronotal disc with punctures fine and sparse; posterior corners rounded; hypomere with short bristles; mesepisternum, sides of metaventrite and metacoxae with bristles; disc of metaventrite with abundant spine-like bristles; distance between mesocoxae and metacoxae twice the metacoxa length; scutellum ogival, smooth. Elytra: shiny or semiopaque, inconspicuous bristles throughout the elytron, a row of short bristles on the fourth ridge; elytral suture and elytron unicolored, slightly elevated; all four elytral ridges barely noticeable. Legs: procoxa with bristles, weakly scaly and punctate; protibia with three teeth equal in size; distance between basal and middle teeth longer than the distance between middle and apical teeth; mesofemur with thick, erect bristles along the inferior margin; mesotibia quadrate in cross section, mesotibial surface smooth; mesotibia with two transverse carinae present posteriorly; metafemur with thick, erect bristles along the inferior margin; males with inner margin of metatibia weakly carinate towards apex; surface finely sculptured, almost smooth; metatibia with two transverse carinae present posteriorly; males with metatibial apical spurs of different lengths and shape, the longer spur equal to the diameter of the tibial apex, the shorter spur wider and projected laterally; metatarsomere I shorter and equal in width to tarsomere II; protarsomeres and mesotarsomeres I–IV enlarged, protarsomere II very wide; protarsomeres less than twice wider than the mesotarsomeres; claw bifid, symmetrical, superior tooth of claw longer and wider than the inferior tooth; distance between teeth longer to the length of inferior tooth. Abdomen: ventrites with erect bristles at the disc, more abundant on ventrites II and III; propygidium with few bristles; pygidium flat; subtrapezoidal; pygidial width not exceeding distance between spiracles of propygidium; pygidial disc smooth, inconspicuous bristles throughout, a few longer bristles towards the apex; pygidial apex subquadrate. Parameres: basal region slightly wider than both sections of the parameres at its maximum width, parameral split at 2/3 the length of the basal region; inner margins concave; three times the length of the apex; apex lanceolate, outer margins weakly expanded in a plane below at the base; parameres in lateral view convex, coplanar (Fig. 2G). Type locality. ARGENTINA. 20 km N La Rioja [near El Cantadero, 29°14’28.9’’S 66°51’20.3’’W]. Geographical distribution. ARGENTINA (La Rioja, San Juan, Buenos Aires, Mendoza, San Luis, Río Negro, Chubut). Remarks. Liogenys calcarata resembles L. flaveola in the body color, size, and in the shape of tibiae and pygidium. Liogenys calcarata differs from L. flaveola (in parenthesis) in the clypeal emargination deep (shallow); pronotal punctures finer; pygidial disc with bristles (glabrous); males with metatibial spurs uneven, the shorter spur wider and asymmetrically projected (truncate but not projected). The parameres of both species have apex lanceolate, in L. calcarata the outer margins are expanded in a plane below (in one plane). This species also resembles L. kuntzeni in the shape of the apex lanceolate, and at its base the outer margin is also expanded in a plane below. In the case of L. calcarata, the expanded portion at the base of the apex is very narrow., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 52-54, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172, {"references":["Frey, G. (1970) Beitrag zur kenntnis der Sudamerikanischen Melolonthiden (Col.). Entomologische Arbeiten aus dem Museum G. Frey, 21, 281 - 287.","Krajcik, M. (2012) Checklist of the World Scarabaeoidea. 5 th Edition. Animma. x, Plzen, 278 pp.","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211 (1), 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3. Papers in Entomology, 1, 1 - 344.","Cherman, M. A., Moron, M. A. & Almeida, L. M. (2016) Phylogenetic relationships within Diplotaxini Kirby (Coleoptera: Melolonthidae: Melolonthinae) with emphasis on Liogenys Guerin-Meneville. Systematic Entomology, 41, 744 - 770. https: // doi. org / 10.1111 / syen. 12188","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031"]}

Liogenys rufoflava Moser, 1918 Figs. 9; 11G, H; 14 Liogenys rufoflavus Moser, 1918: 105; Frey 1969: 48 (key); Krajčík 2012: 145 (checklist); Cherman et al. 2017: 4 (generic history). Liogenys rufoflava: Blackwelder 1944: 228 (checklist); Evans 2003: 214 (checklist); Evans & Smith 2009: 182 (checklist); Cherman et al. 2017: 20 (natural history). Type material. Male lectotype of Liogenys rufoflava here designated (ZMHB): [white typeset] “ Argentinien ”, [white handwritten] “ Liogenys / rufoflavus / Typen ³ / Moser”, [red typeset] “Typus”, [white typeset] “ Liogenys / rufoflavus / Mos.”, [red, outlined, typeset and handwritten] “ LIOGENYS / RUFOFLAVUS Mos / LECTOTYPE / Des. Cherman M. A.”, genitalia mounted. Female paralectotype bearing the labels: [white typeset] “ Argentinien ”, [white handwritten] “ Liogenys / rufoflavus / Typen ♀ / Moser”, [red typeset] “Typus”, [white typeset] “ Liogenys / rufoflavus / Mos.”, [red, outlined, typeset and handwritten] “ LIOGENYS / RUFOFLAVUS Moser, 1918 / PARA-LECTOTYPE / Des. Cherman M. A.” Non-type material (88). PARAGUAY. Boquerón: without locality, 12.XII.1947, Hunger, 1 male and 2 females (CMNC); Parque Nacional Teniente Inciso, administración, 21º12’S, 61º39’W, 18.IX. 2003, 253 m, B. Garcete, 1 male and 1 female (DZUP). ARGENTINA. Salta: La Viña, 24.IX.1988, Peña, 1 female (INPA); Santiago del Estero: Tintina, XI.1946, without collector, 2 females (MZUC); Río Hondo, Termas, IX.1990, A. Martínez, 2 females (CMNC); Río Hondo, VIII.1956, 1 male (CMNC); Bords du Rio Salado, Icaño, Mistol Paso, XII.1908 – III.1909, E. R. Wagner, 1 female (MNHN); Chuna Pampa [10 km from Lugones railway Rosario-Tucumán], XII.1908 – III.1909, E. R. Wagner, 4 females (MNHN); La Rioja: 20 km N La Rioja, 25.IX.1968, 1 male (AMNH); Guayapa, Patquia [30º01’S 66º57’W], XII.1964, A. Martínez, 2 males (CMNC); Guayapa, Patquia, III.1955, without collector, 1 female (MZUC); Patquia, IX.1945, Breyer, 1 male and 1 female (CMNC); Patquia, III.1948, Breyer, 1 female (CMNC); Catinzaco, 18.XII.1970, Roig & Maury, 1 male and 1 female (IADIZA); Chepes (6 km E), 10.XII.1978, at light, Woodruff, Runnacles, Cordo, 1 male (CMNC); San Juan: El Baldecito, Valle Fertil, XII.1964, Martínez, 6 males and 1 female (CMNC); Marayes, 13.XII.1964, A. Martinez, 1 female (DZUP); 2 males and 1 female (NHMB); Mendoza: without locality, 9.XI.1987, L. Peña, 1 female (CEMT); without locality, 21.X.1982, L. Peña, 4 males and 1 female; without locality, 9. XI.1987, L. Peña, 1 male (INPA); Ñacuñan, 34º02’S, 67º55’W, 22.XI.1997, Lagos, 1 male and 1 female (IADIZA); Reserva Telteca, IX–XI.1996, Flores, Roig, 2 females (IADIZA); San Rafael, 6.XII.1988, L. Peña, 1 male and 6 females (INPA); Malvinas, 10.IX.1987, L. Peña, 3 females (INPA); Estación Pedregal, without date; Jensen-Haarup, 3 males and 2 females (ZMUC); La Pampa: Caleu Caleu (Río Colorado), II.1966, Martínez, 1 female (CMNC); Neuquén: Covunco, 19.XII.1963, 1 specimen (CMNC); Zapala (near), 19–21.XII.1965, A. Martínez, 1 male (CMNC); Pucará, 7.XII.1954, Pastrana, 1 female (CMNC); Parque Nacional Lanin, Cerro Malo, 19.XII.1954, Pastrana, 2 males and 1 female (CMNC); Río Negro: Catriel, 311 m, 4.I.2005, F.C. Ocampo, 5 females (CMNC); San Antonio Oeste, Los Grutas, XII.1973, C. Bordón, 1 male (CMNC); Villa Regina, 20.XI.1963, L. Peña, 1 male (CMNC); Chubut: Península de Valdés, Golfo Nuevo, Estancia San Pablo, 42°41’42’’S, 64°10’46’’W, 27.XI.2005, 50 m, 1 female (CMNC). CHILE: Biobío: Laguna del Laja, 16.I.1987, L. Peña, 3 males and 7 females (INPA). Diagnosis. Body light brown, elongate, sides subparallel, shiny, head and pronotum slightly darker than elytra; frontoclypeal surface and pronotum with inconspicuous bristles, elytra uniformely with bristles (Fig. 9A); clypeal emargination shallow, wide, and rounded; clypeal lateral margin convex and barely produced in males, not produced in females (Fig. 11 G–H); antennae with 10 antennomeres; pronotal posterior corners subangulate (Fig. 9C); protibial with three equally-spaced teeth; mesotibia quadrate or subquadrate in cross section (Fig. 9D), males with one mesotibial transverse carina only posteriorly, females with two transverse carinae, the apical carina complete; males with metatibial transverse carina present posteriorly, females with two transverse carinae, the apical carina complete; inner margin of male metatibia carinate straight towards apex; pygidium slightly convex; elongate, apex rounded; pygidial disc finely punctate and with bristles throughout (Fig. 9E); parameres very short, apex lanceolate in a single plane (Fig. 9F). Redescription. Length: 7.4–8.9 mm; width: 3.7–3.9 mm. Light brown. Head. distance between eyes three times the width of one eye, wider in females; frons swollen, equal in length or slightly longer than clypeus; frontoclypeal surface sometimes with short bristles; clypeal emargination shallow, wide, and rounded, in males as wide as the distance between eyes; outer sides of anterior teeth parallel in males; subparallel in females (Fig. 11 G–H); clypeal lateral margin convex and barely produced in males, distance between clypeal lateral projection and anterior tooth as long as the basal width of anterior tooth, distance between clypeal lateral projection and anterior margin of eye shorter than one eye width, margin continuous, not forming a sharp angle between outer side of anterior teeth and clypeal lateral projection; distal maxillary palpomere, maximum width more than two times the apical width; fovea deep, very small, not reaching the transverse midline of the palpomere; antenna with 10 antennomeres, club lighter in color than funicle. Thorax: anterior margin of pronotum slightly produced medially; inconspicuous bristles throughout the pronotal surface, more visible at the sides; pronotal disc with punctures coarse and sparse; posterior corners subangulate, angle obtuse; hypomere with short and a few long bristles; mesepisternum, sides of metaventrite and metacoxae with bristles; distance between mesocoxae and metacoxae less than twice the metacoxa width; scutellum ogival, finely punctate throughout. Elytra: shiny on disc, weakly pruinose at the outer margins; bristles dispersed uniformly; elytral suture and elytron unicolored, weakly elevated; all four elytral ridges weakly defined. Legs: procoxa scaly and punctate; three protibial teeth, middle and apical equal in size, all three teeth equally spaced; mesotibia quadrate in cross section in males, subquadrate in females; mesotibial surface finely sculptured; mesotibia with two transverse carinae, the apical carina only demarked posteriorly in males, complete in females; inner margin of male metatibia carinate straight towards apex; surface coarsely sculptured; metatibia with two transverse carinae, in females the apical carina complete; metatibial apical spurs of different lengths and shape, the larger spur slightly longer than the diameter of the tibial apex, in males the smaller spur wider and truncate; metatarsomere I equal in size to metatarsomere II; males with protarsomeres and mesotarsomeres I–IV enlarged, protarsomere II wide; mesotarsomeres almost one-half width of protarsomeres; claw bifid, symmetrical, superior tooth of claw longer and narrower than the inferior tooth; distance between teeth equal to the length of inferior tooth. Abdomen: ventrites uniformly with bristles at the disc and sides; propygidium with bristles; pygidium flat or slightly convex; subquadrate; elongate; pygidial width almost equal to the distance between spiracles of propygidium, slightly narrower; pygidial disc finely punctate and with bristles throughout, bristles short and decumbent; pygidial apex subquadrate to rounded. Parameres: basal region wider than both sections of the parameres at maximum width, parameral split at 2/3 the length of the basal region; inner margins convex; parameres very short, length up to twice the length of the apex; apex lanceolate in a single plane; parameres in lateral view straight, not coplanar (Fig. 9G). Type locality. ARGENTINA. Mendoza Geographical distribution. PARAGUAY (Boquerón); ARGENTINA (Salta, Santiago del Estero, La Rioja, San Juan, Mendoza, La Pampa, Neuquén, Río Negro, Chubut); CHILE (Biobío). Remarks. Liogenys rufoflava resembles L. flavida in the body size, elytra with bristles, shape of pygidium elongate; and the frons swollen, comparing mainly the females. This species differs in the pronotum somewhat darker in color than the elytra; head, pronotum, and elytra with more abundant bristles; antenna with 10 antennomeres instead of 9; elytra approximately 2.5 times longer than the pronotum (in L. flavida the elytra are more than three times longer than the pronotum); in males the protibia has three teeth instead of two, protarsomere II is wider; and the parameres are different in shape. This species also resembles L. brachyclypeata new species, especially when comparing the females, although L. rufoflava can be distinguished mainly by the width of the head, narrower than the posterior margin of the pronotum. Liogenys rufoflava also have some features in common with L. maxillaricuspis new species (for more details see remarks of these new species). The first record of L. rufoflava in Chile is noteworthy, as it is the only Liogenys occurring in Chile that is not endemic to this country. Further phylogeographical studies will bring light in order to discover if L. rufoflava is older than the Andes formation, as well as its ancestral relationship with the other species from the same region., Published as part of Cherman, Mariana A., Basílio, Daniel S., Mise, Kleber M., Frisch, Johannes, Smith, Andrew B. T. & Almeida, Lúcia M., 2020, Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species, pp. 46-84 in Zootaxa 4896 (1) on pages 73-74, DOI: 10.11646/zootaxa.4896.1.2, http://zenodo.org/record/4359172, {"references":["Moser, J. (1918) Neue Amerikanische Melolonthiden. Stettiner Entomologische Zeitung, 79, 95 - 167.","Frey, G. (1969) Bestimmungstabelle und revision der gattung Liogenys Guerrez (Coleoptera-Melolonthinae-Macrodactylini). Entomologische Arbeiten aus dem Museum G. Frey, 20, 38 - 64.","Krajcik, M. (2012) Checklist of the World Scarabaeoidea. 5 th Edition. Animma. x, Plzen, 278 pp.","Cherman, M. A., Mise, K. M., Moron, M. A., Vaz-de-Mello, F. Z. & Almeida, L. M. (2017) A taxonomic revision of Liogenys occurring in Brazil with an interactive key and remarks on New World Diplotaxini (Coleoptera, Melolonthidae). ZooKeys, 699, 1 - 120. https: // doi. org / 10.3897 / zookeys. 699.12031","Blackwelder, R. E. (1944) Checklist of the Coleopterous Insects of Mexico, Central America, The West Indies and South America. Part 2. Smithsonian Institution, Washington, D. C., pp. 189 - 265. https: // doi. org / 10.5479 / si. 03629236.185.2","Evans, A. V. (2003) A checklist of the NewWorld chafers (Coleoptera: Scarabaeidae: Melolonthinae). Zootaxa, 211 (1), 1 - 458. https: // doi. org / 10.11646 / zootaxa. 211.1.1","Evans, A. V. & Smith, A. B. T. (2009) An electronic checklist of the New World chafers (Coleoptera: Scarabaeidae: Melolonthinae) version 3. Papers in Entomology, 1, 1 - 344."]}

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."]}

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 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

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 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."]}

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."]}

Madecorphnus grandis Frolov, Smith, and Akhmetova, new species (Figures 1���5.) Type material. Holotype: male at FMNH labeled ��� MADAGASCAR: Befin- / gotra (11.0 km WSW), / R��s Anjanaharibe-Sud, / 14��45���S, 49��27���E, / 1565m, montane rain- / forest, 16.XI.1994 // FMHD #94-58, Winkler / extraction of sifted litter / (leaf mold, rotten wood) / B. L. Fisher #1232 (1-50) / FIELD MUS. NAT.���. Description. Holotype, male (Figs. 1���5). Body length 7.5 mm (without mandibles). Color uniformly dark brown, almost black, legs and underside of body somewhat paler. Right mandible 2 times longer than left, without tooth behind apex (Fig. 3). Labrum subtrapezoidal, with slightly rounded sides, length about 1/8 width (in dorsal view). Clypeus slightly asymmetrical, apically obtuse, with 2 long and 3 shorter setae on the apical margin. Canthus and frontal suture indistinct. Clypeus slightly depressed apicomedially. Head without traces of frontoclypeal suture, finely punctate with minute punctures separated by greater than 4 times their diameter. Pronotum approximately 1.5 times wider than long, widest medially. Disc of pronotum convex, without any depressions, tubercles, or ridges. Punctation on pronotum similar to that on head. Margins with relatively wide border, lateral margins with 4 long setae: 1 seta on basal angle, 1 seta approximately in the middle of lateral margin, and 2 setae on the apical angle. Scutellum triangular, angulate apically, about 1/12 length of elytra. Elytra convex, with distinct humeral and apical umbones, widest at basal third. First stria distinct and reaching the apex of elytron, other striae indistinct. Elytra with double punctation: entire surface with minute punctures similar to those on head and pronotum; disc with larger, sparse, elongate, setigerous punctures. Epipleura with long, sparse, brown setae. Base of elytron with border from scutellum to humeral callus. Wings fully developed. Protibiae with 3 outer teeth, lateral margin basad of outer teeth not crenulate. Apex with robust, spur-like seta and a few smaller setae basally. Mesothoracic and metathoracic legs similar in shape to each other. Longer tibial spur shorter than mesotarsomeres 1���2 in mesothoracic legs and as long as metatarsomeres 1���2 in metathoracic legs. Apices of parameres rounded in lateral view (Fig. 4). Lateral teeth of parameres poorly developed, situated approximately in middle of lateral paramere margin. Endophallus with 2 large, robust spinules with a sclerotization near base of one of spinules, and 2 smaller, comma-shaped sclerites (Fig. 5). Female unknown. Diagnosis. This species is similar to M. peyrierasi Frolov, 2010 and, especially, M. perinetensis Frolov, 2010 in the shape of the endophallus armature but differs in having comma-shaped sclerites longer than in other species, wider and rounded in lateral view apices of parameres, and in less developed lateral teeth of parameres. The new species is also 1.5���2.0 mm longer than M. peyrierasi and M. perinetensis specimens and somewhat longer than the holotype of M. falcatus Paulian, 1992, the largest Madecorphnus species known so far. Distribution. The new species is known from a tropical rainforest in the Anjanaharibe-Sud Reserve, in mid-altitude escarpments of northern Madagascar (Fig. 6). The nearest Madecorphnus locality is that of M. barclayi Frolov, 2012 in the northern part of the Masoala Peninsula (Frolov 2012), some 100 km southeast of Anjanaharibe-Sud. The similar species M. peyrierasi and M. perinetensis occur some 500 km to the south (Fig. 6). Etymology. Adjective in the nominative singular. From the Latin grandis for large, denoting the relatively large size of the species., Published as part of Frolov, Andrey V., Smith, Andrew B. T. & Akhmetova, Lilia A., 2017, A new species of the genus Madecorphnus (Coleoptera: Scarabaeidae: Orphninae) from Madagascar in Zootaxa 4341 (2), DOI: 10.11646/zootaxa.4341.2.7, http://zenodo.org/record/1039495, {"references":["Paulian, R. (1992) Un nouveau genre et une nouvelle espece d'Orphnidae de Madagascar (Coleoptera, Scarabaeoidea). Revue Francaise d'Entomologie, New Series, 14, 169 - 171.","Frolov, A. V. (2012) Three new species of scarab beetle genus Madecorphnus Paulian, 1992 (Coleoptera: Scarabaeidae: Orphninae) from Madagascar. Proceedings of the Zoological Institute RAS, 316, 159 - 165."]}