taxonID	type	format	identifier	references	title	description	created	creator	contributor	publisher	audience	source	license	rightsHolder	datasetID
03AA23621471FF84FF28AC37FEEDABE2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF84FF28AC37FEEDABE2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF84FF28AC37FEEDABE2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF84FF28AC37FEEDABE2.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF81FF28ABCBFF71A9C4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF81FF28ABCBFF71A9C4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF81FF28ABCBFF71A9C4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF81FF28ABCBFF71A9C4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621471FF81FF28ABCBFF71A9C4.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AEB8FE9CAC02.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AEB8FE9CAC02.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AEB8FE9CAC02.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AEB8FE9CAC02.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AEB8FE9CAC02.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213974/files/figure.png	https://doi.org/10.5281/zenodo.15213974	FIG. 3. Ranges of average intragroup (blue) and intergroup (red) K2P Onthophagus species groups genetic distances.Acronyms: ch = O. chevrolati species group, cl = O. clypeatus species group, cr = O. crinitus species group, cu = O. curvicornis species group, di = O. dicranius species group, ga = O. gazellinus species group, he = O. hecate species group, hi = O. hircus species group, ho = O. hoepfneri species group, la = O. landolti species group, me = O. mexicanus species group, na = O. nasutus species group, ve = O. velutinus species group.	FIG. 3. Ranges of average intragroup (blue) and intergroup (red) K2P Onthophagus species groups genetic distances.Acronyms: ch = O. chevrolati species group, cl = O. clypeatus species group, cr = O. crinitus species group, cu = O. curvicornis species group, di = O. dicranius species group, ga = O. gazellinus species group, he = O. hecate species group, hi = O. hircus species group, ho = O. hoepfneri species group, la = O. landolti species group, me = O. mexicanus species group, na = O. nasutus species group, ve = O. velutinus species group.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213976/files/figure.png	https://doi.org/10.5281/zenodo.15213976	FIG. 4. Frequency distribution histograms of intragroup and intergroup overlap of Onthophagus species groups K2P genetic distances for COI-5P mitochondrial barcoding sequences. Intragroup and intergroup genetic distances show no barcoding gap.	FIG. 4. Frequency distribution histograms of intragroup and intergroup overlap of Onthophagus species groups K2P genetic distances for COI-5P mitochondrial barcoding sequences. Intragroup and intergroup genetic distances show no barcoding gap.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213980/files/figure.png	https://doi.org/10.5281/zenodo.15213980	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213982/files/figure.png	https://doi.org/10.5281/zenodo.15213982	FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale.	FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8EFF28AC68FCB7AAFB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8FFF28AAE0FF71AA43.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8FFF28AAE0FF71AA43.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8FFF28AAE0FF71AA43.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8FFF28AAE0FF71AA43.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147BFF8FFF28AAE0FF71AA43.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147AFF8FFF28AAA8FA83A84B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147AFF8CFF28A850FEEBAEEB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147AFF8CFF28A850FEEBAEEB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213980/files/figure.png	https://doi.org/10.5281/zenodo.15213980	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147AFF8CFF28A850FEEBAEEB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213982/files/figure.png	https://doi.org/10.5281/zenodo.15213982	FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale.	FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147AFF8CFF28A850FEEBAEEB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147AFF8CFF28A850FEEBAEEB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147AFF8CFF28A850FEEBAEEB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AEF0FACCAD07.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AEF0FACCAD07.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AEF0FACCAD07.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AEF0FACCAD07.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AEF0FACCAD07.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AEF0FACCAD07.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213980/files/figure.png	https://doi.org/10.5281/zenodo.15213980	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AD6CFC62A84B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AD6CFC62A84B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AD6CFC62A84B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8CFF28AD6CFC62A84B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8DFF28A850FAA5AFCB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8DFF28A850FAA5AFCB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8DFF28A850FAA5AFCB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8DFF28A850FAA5AFCB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621479FF8DFF28A850FAA5AFCB.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8DFF28AFD0FCA4AAD7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8DFF28AFD0FCA4AAD7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8DFF28AFD0FCA4AAD7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8DFF28AFD0FCA4AAD7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8DFF28AFD0FCA4AAD7.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8AFF28A91CFA92AC03.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8AFF28A91CFA92AC03.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8AFF28A91CFA92AC03.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8AFF28A91CFA92AC03.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA23621478FF8AFF28A91CFA92AC03.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8AFF28AC68FD7BA93B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8AFF28AC68FD7BA93B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8AFF28AC68FD7BA93B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8AFF28AC68FD7BA93B.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213980/files/figure.png	https://doi.org/10.5281/zenodo.15213980	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8BFF28A920FB90AC97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8BFF28A920FB90AC97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8BFF28A920FB90AC97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8BFF28A920FB90AC97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147FFF8BFF28A920FB90AC97.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213986/files/figure.png	https://doi.org/10.5281/zenodo.15213986	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	FIG. 9. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species, partitioned by codon. An IQ-TREE analysis was performed using default parameters for the Invertebrate Mitochondrial Code. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213969/files/figure.png	https://doi.org/10.5281/zenodo.15213969	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 1. Neighbor-joining tree for the Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any species group. The male protibiae of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213978/files/figure.png	https://doi.org/10.5281/zenodo.15213978	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	FIG. 5. Onthophagus important characters marked with arrows (details discussed in the text). 1, O. atriglabrus; 2, O. orphnoides; 3, O. propraecellens; 4, O. genuinus; 5, O. andersoni; 6, O. crinitus; 7, O. gazellinus; 8, O. onthochromus; 9. O. landolti; 10, O. anthracinus; 11, O. championi; 12, O. cynomysi; 13, O. orpheus orpheus; 14, O. hoepfneri; 15, O. acuminatus; 16, O. incensus; 17, O. cyanellus; 18, O. sanpabloetlorum; 19, O. cartwrighti; 20, O. velutinus. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213980/files/figure.png	https://doi.org/10.5281/zenodo.15213980	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	FIG. 6. Ratio of maximum width to length of metafemur, a larger number indicates a relatively wider femur. 1 O. curvicornis species group; 2 O. hircus species group; 3 O. nasutus species group; 4 O. mexicanus species group; 5 O. eulophus species group; 6 O. velutinus species group; 7 O. landolti species group; 8 O. hecate species group; 9 O. chevrolati species group; 10 O. crinitus species group; 11 O. gazellinus species group; 12 O. hoepfneri species group; 13 O. dicranius species group; O. clypeatus species group. Parts of the box plot: left end of the line, minimum value; right end of the line, maximum value; left side of the box, first quantile; right side of the box, fourth quantile; cross line in the box, median; X-shaped mark, average. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213982/files/figure.png	https://doi.org/10.5281/zenodo.15213982	FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale.	FIG. 7. Male protibiae of the following subspecies: 1, O. orpheus canadensis; 2, O. orpheus pseudorpheus; 3, O. orpheus orpheus. All photos taken in the same scale.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213984/files/figure.png	https://doi.org/10.5281/zenodo.15213984	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	FIG. 8. Maximum-likelihood gene tree for the Western Hemisphere Onthophagus species. Bootstrap values were calculated using the rapid bootstrap algorithm of RaxML under the GTRCAT model with 10,000 replicates and run on the CIPRES portal. See Table 2 for an explanation of the specimen codes.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213972/files/figure.png	https://doi.org/10.5281/zenodo.15213972	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	FIG. 2. A summarized neighbor-joining tree for Western Hemisphere Onthophagus species, based on K2P genetic distances comparisons between COI-5P mitochondrial barcoding sequences. Figs. 23–25 belong to the O. eulophus species group. Figs. 44 and 63 are not assigned to any group. The male metafemora of the following species are depicted: 1, O. acuminatus; 2, O. batesi; 3, O. incensus; 4, O. nitidor; 5, O. stockwelli; 6, O. striatulus striatulus; 7, O. tristis; 8, O. basicarinatus; 9, O. hircus; 10, O. marginicollis; 11, O. nyctopus; 12, O. onorei; 13, O. osculati; 14, O. ptox; 15, O. ranunculus; 16, O. atriglabrus; 17, O. genuinus; 18, O. rostratus; 19, O. sharpi; 20, O. tapirus; 21, O. villanuevai; 22, O. violetae; 23, O. anewtoni; 24, O. arnetti; 25, O. browni; 26, O. championi; 27, O. guatemalensis; 28, O. mexicanus; 29, O. cartwrighti; 30, O. velutinus; 31, O. alluvius; 32, O. altivagans; 33, O. anthracinus; 34, O. brachypterus; 35, O. chryses; 36, O. landolti; 37, O. lecontei; 38, O. schaefferi; 39, O. concinnus; 40, O. cynomysi; 41, O. hecate hecate; 42, O. medorensis; 43, O. orpheus orpheus; 44, O. polyphemi sparsisetosus; 45, O. chevrolati; 46, O. cyanellus, 47, O. hippopotamus; 48, O. oaxacanus; 49, O. reyesi; 50, O. salvadorensis; 51, O. sanpabloetlorum; 52, O. skelleyi; 53, O. subtropicus; 54, O. totonicapanus; 55, O. vespertilio; 56, O. crinitus; 57, O. gazellinus; 58, O. onthochromus; 59, O. aciculatulus; 60, O. coscineus; 61, O. hoepfneri; 62, O. igualensis; 63, O. lebasi; 64, O. pennsylvanicus; 65, O. subanaeus; 66, tuberculifrons; 67, O. barreti; 68, O.breviconus; 69, O. contrapositus; 70, O. dicranius; 71, O. dorsipilulus; 72, O. humboldti; 73, O. kohlmanni; 74, O. micropterus; 75, O. neomirabilis; 76, O. orphnoides; 77, O. petenensis; 78, O. subcancer; 79, O. andersoni; 80, O. belorhinus; 81, O. grataehelenae; 82, O. limonensis; 83, O. maya; 84, O. praecellens; 85, O. propraecellens; 86, O. rhinolophus; 87, O. rhinophyllus; 88, undescribed species of Onthophagus; 89, O. viridivinosus; 90, O. xanthomerus. See Table 2 for an explanation of the specimen codes. See Table 7 for the author names of the species listed.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213974/files/figure.png	https://doi.org/10.5281/zenodo.15213974	FIG. 3. Ranges of average intragroup (blue) and intergroup (red) K2P Onthophagus species groups genetic distances.Acronyms: ch = O. chevrolati species group, cl = O. clypeatus species group, cr = O. crinitus species group, cu = O. curvicornis species group, di = O. dicranius species group, ga = O. gazellinus species group, he = O. hecate species group, hi = O. hircus species group, ho = O. hoepfneri species group, la = O. landolti species group, me = O. mexicanus species group, na = O. nasutus species group, ve = O. velutinus species group.	FIG. 3. Ranges of average intragroup (blue) and intergroup (red) K2P Onthophagus species groups genetic distances.Acronyms: ch = O. chevrolati species group, cl = O. clypeatus species group, cr = O. crinitus species group, cu = O. curvicornis species group, di = O. dicranius species group, ga = O. gazellinus species group, he = O. hecate species group, hi = O. hircus species group, ho = O. hoepfneri species group, la = O. landolti species group, me = O. mexicanus species group, na = O. nasutus species group, ve = O. velutinus species group.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
03AA2362147EFF88FF78ADA5FA0DADDC.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/15213976/files/figure.png	https://doi.org/10.5281/zenodo.15213976	FIG. 4. Frequency distribution histograms of intragroup and intergroup overlap of Onthophagus species groups K2P genetic distances for COI-5P mitochondrial barcoding sequences. Intragroup and intergroup genetic distances show no barcoding gap.	FIG. 4. Frequency distribution histograms of intragroup and intergroup overlap of Onthophagus species groups K2P genetic distances for COI-5P mitochondrial barcoding sequences. Intragroup and intergroup genetic distances show no barcoding gap.	2025-03-17	Kohlmann, Bert;Solís, Ángel		Zenodo	biologists	Kohlmann, Bert;Solís, Ángel			
