taxonID	type	description	language	source
E91F87CDFF98587CF68A83712742F980.taxon	description	(Figs 1 A – E)	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF98587CF68A83712742F980.taxon	materials_examined	Type locality. Brasil, Mato Grosso, Rio Bento Gomes, Pantanal; 15 ° 49 ’ S, 56 ° 31 ’ W. Type material. Holotype, female: Brasilien: Mato Grosso | Rio Bento Gomes | (Pantanal) | 1993 – 1994, E. Stuhr leg. || Flussabschnitt bei | Filisterra | 15 ° 49 ’ S | 56 ° 31 ’ W [NHMW].	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF98587CF68A83712742F980.taxon	description	Description of the holotype (female). Dorsum (Fig. 1 A). Body oblong-ovate. Length: 4.85 mm; maximum width: 2.10 mm. Brownish-black, with somewhat lighter head and sides of pronotum; antennae, tarsomeres and mouth parts brown; whole body covered by argillaceous, greyish coating and dorsally with dense long and thick macrosetae. Head (Fig. 1 D) distinctly convex, anteriorly weakly shiny with distinct microreticulation, rest of head covered by argillaceous, greyish coating; transversely trapezoidal; clypeus anterior margin widely, weakly sinuate with sides very widely rounded, not notched before genae, which are obtusely rounded and very distinctly exceeding eyes. Fronto – clypeal suture absent. Clypeus with dense, coarse, shallow punctation bearing long, thick, rounded at apex, brownish macrosetation. Epipharynx (Fig. 1 E) transverse, with sides rounded, anterior margin weakly sinuate, with indistinctly produced corypha bearing four very short and very thin celtes at apex. Acanthopariae, acropariae, prophobae without chaetae. Chaetopariae, adelochaetae, chaetopediae with dense belts of long chaetae; chaetopariae and adelochaetae thick, chaetopediae thin. Epitorma somewhat bell – shaped. Tormae very long. Pronotum entirely covered by argillaceous, greyish coating, distinctly convex, distinctly transverse, very slightly wider than base of elytra, widest in the middle. Anterior angles right-angled; sides very widely rounded, base of pronotum widely rounded before posterior angles. Anterior and posterior margins not bordered, sides not bordered. Sides and posterior margin with dense, long, thick, rounded at apex macrosetation; macrosetation of sides yellowish, lighter than macrosetation of pronotal surface. Punctation of pronotum simple, dense, coarse; punctures bearing long, thick, rounded at apex, brownish macrosetae which are arranged on slight ridges which are similar to some members of the family Trogidae. Scutellum very small, triangular, entirely covered by argillaceous, greyish coating. Base of elytra very weakly and narrowly bordered, with in distinct humeral denticles. Elytra elongate, distinctly convex, widest slightly after the middle. Elytra entirely covered by argillaceous, greyish coating, with ten striae and ten interstriae. Interstriae weakly convex; alternating interstriae with longitudinal, relatively higher tubercles; all interstriae with irregular rows of punctures bearing long, thick, rounded at apex, brownish macrosetation; longitudinal tubercles with tuft of very dense, very long, acute at apex macrosetae. Striae shiny, shallow, with distinct, not coarse, quite dense punctation; punctures of striae not encroaching on interstriae; striae variably connected before apex. Pygidium with similar structure as abdominal ventrites. Legs (Figs 1 A – C) all femora mat, with distinct microreticulation, with dense punctation bearing short, quite thin macrosetae. Profemora distinctly bordered anteriorly and posteriorly, meso- and metafemora bordered only before apex, anteriorly. Protibiae distinctly tridentate laterally, proximally serrulate, with dorsal side smooth; apical spur weakly downwardly bent before apex. Meso- and metatibiae with distinct accessory spine; metatibiae superior apical spur distinctly shorter than basal metatarsomere, latter slightly shorter than following four tarsomeres combined. Claws very short, thin, distinctly arcuate. Macropterous. Venter (Fig. 1 B). Metaventral plate entirely covered by argillaceous, greyish coating, very weakly convex, medially quite distinctly concave, with distinct, narrow, shallow longitudinal line in the middle; with dense punctation bearing short and thin macrosetae. Mesoventral plate small, shiny, strongly microreticulate, not covered by argillaceous, greyish coating; triangular, without any depression. Abdominal ventrites entirely covered by argillaceous, greyish coating; with quite dense, shallow punctation bearing very short, thick macrosetation; anterior margins narrowly, shallowly fluted. Male. Unknown.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF98587CF68A83712742F980.taxon	etymology	Etymology. The name of the new species is derived from the genus name Trox Fabricius, 1775 (Scarabaeoidea: Trogidae) (noun in apposition), which has species with similar tuberculate and bristlely elytral intervals.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF98587CF68A83712742F980.taxon	diagnosis	Differential diagnosis. With the whole body covered by argillaceous, greyish coating, Ataenius trox Minkina & Skelley, new species can confidently be included in the imbricatus species group (see Stebnicka 2003, 2007). Within the group, also Ataenius tuberculatus (Schmidt, 1911) has such distinct and large tubercles on the elytral intervals, with long macrosetation of the body. Ataenius trox Minkina & Skelley, new species has completely different body proportions (body much less elongated), much larger tubercles on the elytral intervals that have tufts of macrosetae, while in A. tuberculatus the tubercles are glossy and lack macrosetation. For comparison with A. tuberculatus see Figs 2 A – F.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9A587CF68A80C6278DF8CD.taxon	description	(Fig. 2 A – F)	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9A587CF68A80C6278DF8CD.taxon	materials_examined	Type material examined. None. Additional material examined. 1 male: Venezuela, Lago de Los Patos, Guar., 15 km S Calabozo. 9 – 13. ii. 1969, leg. P. & P. Spangler [ISEA].	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9C5878F68A86DD2375FE22.taxon	description	(Figs 3 A – F)	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9C5878F68A86DD2375FE22.taxon	materials_examined	Type locality. Indonesia: NW Halmahera NW, W slope at Mount Ibu; 1 ° 24 ’ 09 ’’ N, 127 ° 30 ’ 46 ’’ E. Type material. Holotype, male: Indonesia: Halmaheira NW | W slope at Mt. Ibu | 1 ° 24 ’ 09 ’’ N, 127 ° 30 ’ 46 ’’ E | 400 – 900 m, 25. i. 2006 | leg. A. Skale (UWP) [NHMW]. Paratypes (8 unsexed specimens): the same data as holotype [5 specimens in NHMW, 2 specimens in ISEA, 1 specimen in FSCA].	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9C5878F68A86DD2375FE22.taxon	description	Description of the holotype (male). Dorsum (Fig. 3 A). Length: 3.5 mm; maximum width: 1.20 mm. Dark brownish; antennae, tarsomeres and mouth parts brown. Glabrous. Head (Fig. 3 B) distinctly convex, shiny, with trace of microreticulation; transversely trapezoidal; clypeus anteriorly widely, weakly but distinctly sinuate; sides widely rounded, truncate, weakly notched before genae, which are obtusely rounded and very distinctly exceeding eyes. Fronto – clypeal suture absent. Punctures on clypeus regular in size, shape and distribution, moderately coarse, dense. Epipharynx (Fig. 3 E) transverse, with sides rounded, anterior margin truncate, with not so distinctly produced corypha bearing no celtes at apex. Acanthopariae, acropariae, prophobae without chaetae. Chaetopariae, adelochaetae, chaetopediae with dense belts of long, thick chaetae. Epitorma somewhat drop – shaped, shortened basaly. Tormae quite long. Pronotum distinctly convex, distinctly transverse, somewhat wider than base of elytra, widest in the middle, shiny, without microreticulation. Anterior angles rounded, somewhat right-angled; sides nearly parallel, not distinctly visible from above, very weakly sinuate before posterior angles. Anterior margin not bordered, sides distinctly bordered, basal margin very distinctly bordered. Punctation double; smaller punctures eight to ten times smaller than larger, dense, fine, quite regularly distributed, regular in size; larger punctures dense, quite regularly distributed, somewhat irregular in size, posteriorly larger. Scutellum small, triangular, elongate, shiny, without microreticulation, without punctation. Base of elytra not bordered, with small but distinct humeral denticles. Elytra elongate, distinctly convex, paralel sided. Elytra with ten striae and ten interstriae. Interstriae distinctly convex, shiny, without microreticulation, with sparse, irregularly distributed, very fine punctures. Striae shiny, with distinct, very coarse, quite dense punctation; punctures of striae distinctly encroaching on interstriae; striae variably connected before apex. Pygidium with similar structure as abdominal ventrites. Legs (Figs 3 A – C) all femora quite shiny, with weak microreticulation; with simple, very fine and sparse puncatation. Profemora distinctly bordered anteriorly and posteriorly; meso- and metafemora anteriorly and posteriorly not bordered. Protibiae distinctly tridentate laterally, proximally with few serrations; dorsal side smooth; apical spur outwardly curved before apex, somewhat downwardly bent. Metatibiae superior apical spur distinctly longer than basal metatarsomere, latter nearly as long as following two tarsomeres combined. Claws very short, thin, quite weakly arcuate. Macropterous. Venter (Fig. 3 B). Metaventral plate shiny, without microreticulation, very weakly convex, medially quite distinctly concave; with distinct, narrow, quite deep longitudinal line in the middle; with very fine, moderately dense punctation. Abdominal ventrites shiny, with a trace of microreticulation; anterior margins narrowly, but distinctly fluted, except the last one which is not fluted, only deeply bordered; with distinct, quite sparse, very fine punctation. Aedeagus (Fig. 3 F). Parameres the same length as phallobase, weakly downwardly curved before apex, which is regularly rounded in lateral view. Variation. Length from 3.4 to 3.5 mm. Punctation of pronotum slightly variable in density. Female. Metaventral plate of females with less distinct median concavity and apical spur of protibiae less distinctly curved and less downwardly bent.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9C5878F68A86DD2375FE22.taxon	etymology	Etymology. Toponymic; an adjective derived from the Indonesian name of Halmahera Island: Halmaheira, where is the origin of the new species.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9C5878F68A86DD2375FE22.taxon	diagnosis	Differential diagnosis. Known Indonesian species of Saprosites are revised by Stebnicka (2012) with only one species being more recently described (Minkina & Jákl 2024). When identifying S. halmaheiraensis Minkina & Skelley, new species, using the key of Stebnicka (2012), there is a problem at couplet 14, where S. hamaheraensis posesses a mixture of the characters discussed. Due to the sharply protruding genae and sides of pronotum laterally distinctly, deeply bordered, we should proceed to couplet 19, where we can easily continue to couplet 20. At this point, S. halmaheraensis Minkina & Skelley, new species appears externally similar to the widely distributed S. narae Lewis, 1895. Saprosites halmaheraensis Minkina & Skelley, new species can be distinguished from S. narae (as well as from many other Saprosites species from this region) by characters pointed in the key below.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9E5878F68A828B2577F810.taxon	description	(Figs 4 A – F)	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9E5878F68A828B2577F810.taxon	materials_examined	Type material examined. Holotype, female: Indonesien, North Sumatra, Aceh-Prov. Umg. Von Kutacane, Ketambe, 02. – 07. xii. 2013, leg. Horst Rudolph. depository Additional material examined. 1 male: Indonesia: N Siberut, Mentawai Islands, Mount Malancan, 100 m, 18 – 21. i. 2024, leg. Stanislav Jákl [ISEA].	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9E5878F68A828B2577F810.taxon	description	Short complementary description. The same as holotype except: length: 3.4 mm, body blackish, protibiae slightly more inwardly curved, apical spur of protibiae very slightly less outwardly curved and very slightly less downwardly bent, metaventral plate with a more distinct median concavity. Aedeagus (Fig. 4 F). Parameres distinctly shorter than phallobase, regularly rounded at apex, in basal, down part with distinct, characteristic less sclerotized sinuation. Variation. Length from 3.4 to 3.6 mm. Body colour from brownish to blackish. Punctation of pronotum slightly variably distributed. Sexual dimorphism. Female metaventral plate with a less distinct median concavity; protibiae slightly more inwardly curved, apical spur of protibiae very slightly less outwardly curved and very slightly less downwardly bent.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9E5878F68A828B2577F810.taxon	discussion	Remark. Externally, there are no significant differences (except of sexual dimorphism) between the holotype female from Aceh Province in North Sumatra and the male from the Mentawai Islands.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9E5878F68A828B2577F810.taxon	distribution	Distribution. Indonesia: Sumatra and Siberut Islands.	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
E91F87CDFF9E5878F68A87A827B7FBE7.taxon	discussion	Another externally similar to S. halmaheiraensis Minkina & Skelley, new species is also a widely distributed S. japonicus Walterhouse, 1875, which can be easily distinguished by the much less conspicuously protruding genae and characteristic protuberances on the sides of the pronotum, in front of the posterior angles (compare with figs: 1, 2, 8, 11 in Kawai et al. 2005).	en	Minkina, Łukasz, Skelley, Paul (2025): Studies on Eupariini Schmidt, 1810 (Coleoptera: Scarabaeidae: Aphodiinae) with description of the new species of Ataenius Harold, 1867 and Saprosites Redtenbacher, 1857. Zootaxa 5633 (2): 384-393, DOI: 10.11646/zootaxa.5633.2.10, URL: https://doi.org/10.11646/zootaxa.5633.2.10
