The Bolboceratidae, Geotrupidae (Coleoptera, Scarabaeoidea), and Scarabaeinae (Coleoptera, Scarabaeoidea, Scarabaeidae) fauna of Honduras: Taxonomy and distribution Rivera-Duarte, José D. Salomão, Renato P. Solís, Ángel Michel, Mauricio Matamoros, Marcela I. Kohlmann, Bert ZooKeys 2025 2025-10-10 1255 127 206 Kohlmann, Solis & Rivera Kohlmann, Solis & Rivera 2025 Insecta Scarabaeidae Phanaeus Animalia Coleoptera Arthropoda species lempira sp. nov.   Figs 6, 7   Note. (CS, EZ); Honduras, Mexico.   Figure 6. Habitus  Phanaeus lempira sp. nov.Paratype male ( ♂), Parque Nacional de Montaña Celaque (  14,484 624°N, -88,669 43°W), Departamento de Lempira, Honduras. Scale bar: 2 mm.   Type material.     Holotype: male. Honduras.•  Departamento de Lempira. Municipio de Gracias. Parque Nacional de Montaña Celaque. Bosque de Pino-Encino.  14,485 7°N, -88,670 1°W. JD Rivera, RP Salomão, Michel M leg.  17. VI. 23,  2060 m.    Allotype: female. Honduras.•  Departamento de Lempira. Municipio de Gracias. Parque Nacional de Montaña Celaque. Bosque de Pino-Encino.  14,484 1°N, -88,668 8°W. JD Riveraand RP Salomão leg.  17. VI. 23,  2010 m.    Paratypes: Honduras.•  Departamento de Lempira. Municipio de Gracias. Parque Nacional de Montaña Celaque. Bosque de Pino-Encino. JD Riveraand RP Salomão, Michel M leg.  14,565 1°N, -88,632 9°W.  17. VI. 23,  1728 m( 1 male);•   14,492 8°N, -88,656 1°W.  17. VI. 23  1721 m( 1 male);•   14,491 1°N, -88,656 9°W.  17. VI. 23  1696 m( 2 males);•   14,491 8°N, -88,656 6°W.  17. VI. 23  1705 m( 1 male);•   14,492 2°N, -88,656 4°W.  17. VI. 23  1704 m( 1 male);•   14,484 6°N, -88,669 4°W.  17. VI. 23,  2023 m( 2 males);•   14,484 9°N, -88,669 9°W.  17. VI. 23  2037 m( 1 male);•   14,565 1°N, -88,632 3°W.  16. VI. 23  1258 m( 1 male);•   14,484 1°N, -88,668 9°W.  17. VI. 23  2010 m( 2 females).•   Departamento de Francisco Morazán. Municipio de Tatumbla. El Aguacate,  13 ° 59 ' N, 87 ° 06 ' W,  19 / 7 / 1991, leg L. Vásquez. EAPZ 007.359, ( 1 male).•   Departamento de Copán. Municipio de San Pedro. Capucas,  25. 11. 2018. Leg. D. Díaz, J. Orozco, R. Ortega, ( 1 male, 1 female).   Figure 7. Habitus  Phanaeus lempira sp. nov.Paratype female, Parque Nacional de Montaña Celaque ( 14,484 624 ° N, - 88,669 43 ° W), Departamento de Lempira, Honduras. Scale bar: 2 mm.  Holotype, allotype, and two maleand one female paratypesfrom Departamento Francisco Morazánand Copándeposited in: Insect Collection, Escuela Agrícola Panamericana El Zamorano, Honduras. One maleand one female paratypesfrom Departamento de Lempira( 17 - VI- 2023, 2024 and 2010 ma. s. l.) are deposited in the National Museum of Costa Rica, San Jose, Costa Rica. Nine maleand one female paratypesfrom Departamento de Lempirawere deposited in the Insect Collection of the Hydrobiology Laboratory, School of Biology, Universidad Nacional Autónoma de Honduras.   Figure 8. A, B. Last abdominal ventrite of  C. sallei. A. Male; B. Female, which are always black with yellow borders. C, D. Pygidium of  Canthon sallei. C. Male; D. Female.   Diagnosis. Large species, dull with dark blue pronotum. Male pronotum covered with vermiculate sculpturing on disk. Male with pronotal anterior carina an inverted V, very strongly developed and raised in the middle. Male with base of pronotal disk with a raised, smooth transverse carina, slightly triangularly arched at the middle, with a well-developed tooth and a mid-sulcus at its base (Fig. 6). Basal fossae clearly impressed. Elytral striae fine, well-defined; interstriae slightly convex and shiny, with coppery-red reflections. Female with metabasitarsus widened apically; outer ridge of ventral surface distinctly more sinuous than inner ridge (Fig. 12).   Figure 9. The insect collection of the Natural History Museum, London (photos taken by ÁS in 2009). A. The specimens are  Canthon salleifrom Central America, save one misplaced  C. moniliatus, third line, second from left on the right of the specimen with the Turrialba, Costa Rica label. The first two specimens on the first row, left, having a round red label, are the syntypes of  C. sallei. The macular pronotal pattern of the type material exists in Honduras, Nicaragua, and Costa Rica, albeit other macular variations exist for this species as well; B. The four upper left specimens are  Canthon salleifrom Panama, all other seven specimens are  C. triangulatus. The difference in the pronotal maculae are illustrative.   Figure 10. Map of the known distribution of  Canthon cyanellus(black rhombus),  C. spinosus(red triangle), and  C. sallei(blue dot).  Canthon cyanellushas no subspecies.   Figure 11. Last abdominal sternite of  Canthon spinosusA. Male; B. Female, which are somber and unicolorous. Pygidium of  C. spinosusshowing setae C. Male; D. Female.   Figure 12. A. Photo of the female metabasitarsus of  Phanaeus melampus. Texas A & M University Insect Collection, ex F. Psota collection, ex Field Museum, ex David Edmonds collection. Photo courtesy of Mario Cupello, no locality label. B. Photo of the female metabasitarsus of  Phanaeus lempira sp. nov.Honduras. Parque Nacional de Montaña Celaque. 14,484 138 ° N, - 88,668 799 ° W.   Description of holotype  (male, Fig. 6).Length 22.1 mm. Width 14.9 mm. Dorsum somber, pronotum and pygidium dark blue, elytra appearing black with interstriae having a red-coppery reflection, underside black. Head horn distinctly swollen basally. Pronotum appearing varnished; covered with black-blue vermiculate sculpturing on the disk; anterior carina an inverted V, very strongly developed and raised in the middle; basal medial portion smooth and raised, forming an extended soft triangular arch, acutely denticulate in its anterior center with a clear medial sulcus at its base, basal fossae present. Elytra with fine striae, with obsolete punctures; interstriae finely convex, appearing almost flat, alutaceous, shiny with coppery-red reflections. Profemora coarsely punctate, mesofemora coarsely punctate at apex, metafemora with a few coarse punctures at apex. Pygidium convex, with many coarse punctures and a faint midline.    Allotype.  Female. Fig. 7. Length 22.1 mm. Width 14.1 mm. Dorsum somber, pronotum with coppery-red reflections on disk and turquoise blue on borders, interstriae with coppery-red reflections and pygidium turquoise blue. Head with very rugose surface; tri-tuberculate cephalic carina with nearly aligned, similar-sized, conical tubercles. Pronotal surface rugose, matte; transverse carina forming an open arch with three conical lobes closely set with a posterior concavity; median lobe higher and more projected than laterals, a small midline carina runs from the median lobe base to the concavity; with two basal impressions. Metabasitarsus widened apically; outer ridge of ventral surface distinctly more sinuous than inner ridge (Fig. 12).   Variation. Males. Length 20.2–23.6 mm. Width 11.9–14.9 mm. Females. Length 20.6–22.1 mm. Width 12.4–14.1 mm.   Etymology. The name is dedicated in honor of the Honduran national hero and symbol of liberty and cultural resistance, who was a captain of the Lenca tribes that fought the Spanish invasion and conquest.   Habitat. This species is distributed between 1528 and 2060 ma. s. l. Pine and pine-oak association are distributed in this park between 1000 and 1500 ma. s. l. on the Atlantic slope, and between 1800 and 2300 ma. s. l. on the Pacific slope ( Mejía Ordóñez and House 2002). Six of the seven pine species of the country occur here,  Pinus oocarpaSchiede ex Schltdl., 1838being the most common one. Mixed forest (pine,  Liquidambar styracifluaL., 1753,  Clethrasp.,  Nectandrasp.) is recorded higher than 1500 ma. s. l. and between 1800 and 2000 ma. s. l. the cloud forest begins on the Atlantic slope of the park, while on the Pacific slope it starts at ca 2300 ma. s. l. The cloud forest is characterized by a significant plant species richness (  Abies guatemalensisRehder, 1939,  Alnussp.,  Pinus pseudostrobusLindl., 1839,  Prunussp.,  Quercus lancifoliaSchltdl. & Cham, 1830,  Taxus globosaSchltdl., 1838), where tree ferns are very common. It is the highest mountain range in Honduras. This park is home to several endemic species of plants, amphibians, shrews, a scarab gold beetle (  Chrysina pastoriCuroe, 1994), and a scarab flower beetle (  Archedinus antoshkai Seidel and Arriaga-Varela, 2018).   Distribution.   Phanaeus lempirais closely related to the Mexican endemic  Ph. melampus Harold, 1863. Edmonds (1994)recorded the latter from three localities in Veracruzand two in Chiapas, and another Chiapaslocality was added by Navarrete-Gutiérrez and Edmonds (2006). The nine Navarrete-Gutiérrez and Edmonds specimens were collected at 145 ma. s. l. during February, August, and October in traps baited with human feces in the Montes Azules Biosphere Reserve in tall and medium evergreen tropical forests. Upon request, Navarrete-Gutiérrez provided us with photos of these specimens. Upon close examination, these individuals showed all the characteristics of the new  Phanaeusspecies described in this paper. Even though we did not see Edmonds’s (1994) Chiapasspecimens, they also come from places covered in wet tropical evergreen forest, El Bosque and Tenejapa, at 1460 ma. s. l., indicating they also most probably belong to this new species. Finally, among the material photographed by Mario Cupello from the Texas A & M University collection, there is one specimenwith the characteristics of  Ph. lempirathat bears a label saying only “ Tabasco, Messico”. This specimen might have been collected around Teapa, where Tabascoreaches the piedmont of the Chiapasmountains. All this suggests that  Ph. lempirais distributed from Chiapasto Honduras (Fig. 13), whereas  Ph. melampusis distributed yonder the Isthmus of Tehuantepec divide – currently known exclusively from Veracruz – and not south of it.   Figure 13. Map of the distribution of  Phanaeus lempira sp. nov.(stars) and  Ph. guatemalensis(dots). Triangles represent specimens that were considered initially to belong to  Ph. melampusby Navarrete-Gutiérrez and Edmonds (2006), but that on photographic inspection show the characteristics of  Ph. lempira.  Lizardo et al. (2022: suppl. fig. 1), on predicted and recorded distributions of  Phanaeuspresents a map of the expected distribution of the  Phanaeus amethystinusspecies group of which this species is a member. These climatically predicted distributions are somewhat at odds with the actual recorded distributions. For example, their model predicts a possible distribution of  Ph. melampus(probable sister species of  Ph. lempira) in the marshy tropical savannah of northeastern Honduras, where no species of  Phanaeushas been registered at present and where the majority of the collected species belong to the roller guild and some  Onthophagustaxa as well. These climatic models usually make distribution predictions that are way off the mark in tropical mountainous areas. As Kohlmann et al. (2007)have already discussed, finer-scale systems, like the Life Zones of Holdridge (1967), are better suited for these tropical areas.   Phenology. This species was collected in June, July, and November using traps baited with human feces.   Taxonomic remarks. This species is closely related to  Ph. melampus, reported from a mountain wet tropical evergreen forest from Veracruzin Mexicocollected at 1500 ma. s. l., and to  Ph. guatemalensisHarold, 1871, reported from forests and adjacent open habitats in the Guatemalan Highlands, distributed from 1500 to 2400 m( Edmonds 1994). The nearest recorded species group specimen to the type locality of this new species is  Ph. guatemalensis, 230 kmaway, suggesting a great geographical separation. The new species  Ph. lempiracan be distinguished from  Ph. melampusby the shape of the smooth, transverse basal area of the male pronotum, which features a sulcus at the mid-base of the pronotum. In  Ph. lempira, this area is slightly triangularly arched (Fig. 6), resembling the shape observed in  Ph. amethystinus. In contrast,  Ph. melampusexhibits a strongly triangular, punctate area that lacks the sulcus at the base of the medial tooth (Fig. 14 A). Additionally, the interstriae of  Ph. melampusare dull and rarely show a bluish cast, whereas  Ph. lempirahas shiny interstriae with coppery-red reflections. Female specimens of  Ph. melampushave a semicircular pronotal transverse carina, while  Ph. lempirafeatures a more open transverse carina (Figs 7, 14 B). The female of  Ph. lempirahas a metabasitarsus widened apically, and the outer ridge of the ventral surface is distinctly more sinuous than the inner ridge, similar to  Ph. amethystinus(Fig. 12 B). Conversely, the female of  Ph. melampushas a narrower metabasitarsus with both ventral ridges running more or less straight (Fig. 12 A).   Figure 14. Habitus  Phanaeus melampusA. Male; B. Female. Veracruz, Orizaba, Mexico. Institut royale des Sciences naturelles de Belgique, through Roger Damoiseau. Photos courtesy of W. D. Edmonds.   Phanaeus lempiracan be separated from  Ph. guatemalensisby the interstriae, flat or weakly convex in  Ph. lempira, raised mid-longitudinally, appearing costate in  Ph. guatemalensis. The pronotoum of  Ph. lempirafemales have asperate punctures posteromedially, whereas in  Ph. guatemalensisthe area is coarsely punctate to rugo-punctate, appearing roughened. With this new species, the number of taxa in the  amethystinusspecies group (  amethystinus,  blackalleri,  guatemalensis,  melampus) increases to five. In the work of Lizardo et al. (2022)  Phanaeus genieri, a member of the  amethystinusspecies group, is considered in their distribution prediction analysis, although this species has not yet been reinstated as a valid species. Gillett and Toussaint (2020), based on molecular genetic analysis, considered that this species group appeared during the Miocene around 17 Ma. However, modern species formation seems to have been relatively recent during the Pleistocene, ~ 1 Ma. 5839161301 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 2060 14.4857 Bosque de Pino-Encino 41044 -88.6701 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira holotype 5839161308 2023-06-17 JD Rivera & RP Salomao Honduras Municipio de Gracias 2010 14.4841 Bosque de Pino-Encino 41044 -88.6688 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira allotype 5839161310 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 1728 14.5651 Bosque de Pino-Encino 41020 -88.6329 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira paratype 5839161309 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 1721 14.4928 Bosque de Pino-Encino 41042 -88.6561 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira paratype 5839161312 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 1696 14.4911 Bosque de Pino-Encino 41043 -88.6569 Parque Nacional de Montana Celaque 2 2 Departamento de Lempira paratype 5839161302 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 1705 14.4918 Bosque de Pino-Encino 41042 -88.6566 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira paratype 5839161303 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 1704 14.4922 Bosque de Pino-Encino 41042 -88.6564 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira paratype 5839161313 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 2023 14.4846 Bosque de Pino-Encino 41044 -88.6694 Parque Nacional de Montana Celaque 2 2 Departamento de Lempira paratype 5839161311 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 2037 14.4849 Bosque de Pino-Encino 41044 -88.6699 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira paratype 5839161306 2023-06-16 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 1258 14.5651 Bosque de Pino-Encino 41020 -88.6323 Parque Nacional de Montana Celaque 1 1 Departamento de Lempira paratype 5839161304 2023-06-17 JD Rivera & RP Salomao & Michel M Honduras Municipio de Gracias 2010 14.4841 Bosque de Pino-Encino 41044 -88.6689 Parque Nacional de Montana Celaque 2 2 Departamento de Lempira paratype 5839161307 1991-07-19 EAPZ L. Vasquez Honduras 13.983334 El Aguacate 1289 -87.1 Municipio de Tatumbla EAPZ 007.359 1 1 Departamento de Francisco Morazan paratype 5839161305 2018-11-25 D. Diaz & J. Orozco & R. Ortega Honduras Capucas Municipio de San Pedro 2 1 1 Departamento de Copan paratype