Male-Produced Sex Pheromone of the Carrion Beetles, Oxelytrum discicolle and its Attraction to Food Sources

Abstract Carrion beetles are part of the great diversity of insects collected on cadavers. In Brazil, beetles of the genus Oxelytrum have great forensic importance in post mortem interval (PMI) estimation. We investigated the system of chemical communication in the attraction of these necrophagous beetles. Gas chromatographic analysis (GC) of female and male aeration extracts revealed the presence of two male-specific compounds, produced in a ratio of 94:6. Bioassays showed that the combination of male produced volatiles and the odor of a food source (carcass volatiles) were attractive to females. Mass and infrared spectral analyses of the male-specific compounds suggested that they were both unsaturated hydrocarbons. Several microderivatizations were carried out with the natural products, and the target structures were identified as (Z)-1,8-heptadecadiene (major) and 1-heptadecene (minor). The structure of the minor component was assigned by co-injection with a commercial standard. A seven-step synthesis was developed to synthesize (Z)-1,8-heptadiene, which co-eluted with the major natural product on three different GC stationary phases. Y-tube olfactometer assays showed that the mixture of synthetic standards in the naturally occurring proportion was slightly attractive to females. The results contribute both to the understanding of the chemical ecology of O. discicolle and to its potential to improve the accuracy of PMI estimation

Diagnosis and key of the main families and species of South American Coleoptera of forensic importance

The objective of this paper is to provide diagnosis and keys of the families and species, with illustrations of the main groups. A table of all related species recorded from South America is presented, including the substrate in which they were collected and their geographical distribution. The list comprises 221 species included in 15 families, of which 70% of the species are from Brazil. Scarabaeidae is the most diverse family with 121 species, followed by Staphylinidae with 68. Also we provide one database of Coleoptera species associated with carcasses in South America

Liogenys Guérin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae) from the southern South American Transition Zone and boundaries: taxonomic overview with four new species

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. Zootaxa 4896 (1): 46-84, DOI: https://doi.org/10.11646/zootaxa.4896.1.

Liogenys Guerin-Méneville, 1831 (Coleoptera: Scarabaeidae: Melolonthinae Diplotaxini) of northern South America and Central America: taxonomic overview with four new species

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. Zootaxa 4990 (2): 201-226, DOI: https://doi.org/10.11646/zootaxa.4990.2.

Liogenys schneiderae Cherman & Basílio & Mise & Frisch & Smith & Almeida 2021, new species

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/502588

Liogenys quadridens

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/502588

Liogenys undetermined

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 speciesPublished 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/502588

Liogenys genieri Smith & Cherman 2021, new species

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/502588