taxonID	type	description	language	source
75C9F8D83E375EE5A52275A7A0FB31A7.taxon	description	Figs 6, 54 - 57	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
75C9F8D83E375EE5A52275A7A0FB31A7.taxon	materials_examined	Specimens examined. 1 syntype [Luzon Island] based on images, " Atholus bakeri n. sp. Bickh. / Los Banos, / P. I., Baker. / 1639 " [sex undetermined, measurements not available] (MNHUB).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
75C9F8D83E375EE5A52275A7A0FB31A7.taxon	diagnosis	Diagnosis. This species has lateral pronotal striae interrupted in the anterolateral angle, and strong punctations on its entire pygidium. According to the original description of Bickhardt (1914), Atholus bakeri (Bickhardt, 1914) is most similar to A. torquatus (Marseul, 1854) except that the propygidium and pygidium of A. bakeri (Bickhardt, 1914) are strongly punctate. However, based on the syntype observed, the punctation is not as prominent as described and, in fact quite similar to that of A. torquatus (Marseul, 1854). The punctures are finer towards the apical end of the pygidium. Another distinguishable feature of A. bakeri (Bickhardt, 1914) is the medially straight frontal stria, while in A. torquatus (Marseul, 1854) it is weakly bent inwardly. However, several studied individuals of A. torquatus (Marseul, 1854) likewise seem to have their frontal stria medially straight.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
75C9F8D83E375EE5A52275A7A0FB31A7.taxon	description	Re-description. Body (Fig. 6) oval, moderately convex and black; tibia and antenna rufous. Head: clypeus (Figs 54, 55) slightly crenate on anterolateral margin, apical margin slightly extended; frontal stria medially straight, complete, moderately impressed; eyes large, convex, clearly visible dorsally; mandibles with rounded outer margin curved inwardly; sub-apical tooth on left mandible large; mandibular apex acutely pointed. Pronotum: marginal pronotal stria (Figs 54, 55) laterally complete, continuous onto apical angle and crenate behind head; lateral pronotal stria moderately impressed, slightly crenate; apical end shortened and bent inwardly in a curved hook; lateral portion rather distant from margin; its basal end abbreviated from basal fifth of pronotal length. Elytra: elytral epipleuron (Fig. 57) with few coarse punctures on apical half; marginal epipleural stria present on apical half; marginal elytral stria (Fig. 6) complete, slightly impressed; external and internal subhumeral striae absent; oblique humeral stria lightly impressed on basal third; dorsal elytral striae 1 - 4 complete; dorsal elytral stria 5 and sutural elytral stria present on apical half; disk with fine punctures. Abdomen: propygidium (Fig. 56) moderately covered with coarse, round, and shallow punctures; interspaces with fine punctations; pygidial punctations similar to those of propygidium, becoming sparser apically. Meso- and metaventrite: anterior margin of mesoventrite (Fig. 57) outwardly arcuate; marginal mesoventral stria crenate and complete; meso-metaventral suture clearly impressed, complete, medially angulate; punctations of intercoxal disk of metaventrite similar to those of mesoventrite; longitudinal suture of metaventrite lightly impressed; lateral disk of metaventrite moderately covered with large, round, shallow punctures. Legs: posterior surface of protibia (Fig. 57) flattened and strongly dilated; outer lateral margin with four weak, almost inconspicuous teeth, topped by minute denticles.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
75C9F8D83E375EE5A52275A7A0FB31A7.taxon	distribution	Distribution. Endemic to the Philippines (Mazur 2011).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
75C9F8D83E375EE5A52275A7A0FB31A7.taxon	biology_ecology	Biology. Unknown.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C773C507E3D95545B2E224FEB32C0370.taxon	description	Figs 2, 9 - 14, 38 - 40	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C773C507E3D95545B2E224FEB32C0370.taxon	materials_examined	Specimens examined. 13 ♂♂, 2 ♀♀ and 4 specimens of undetermined sex. Luzon Island, Isabela, Angadanan, Pissay, 1 ♂, 16.44207 ° N, 121.46277 ° E 60 m a. s. l., 2019 - VII- 20 [IS- 19 - IDC- 001], I. N. DELA CRUZ leg.; Pangasinan, Asingan, Bantog, 1 ♂, 15.59384 ° N, 120.41151 ° E 50 m a. s. l., 2019 - VII- 22 [PG- 19 - IDC- 001], I. N. DELA CRUZ leg.; Batangas, Calatagan, Balitoc, 1 ♀, 13.51417 ° N, 120.38138 ° E 10 m a. s. l., 2019 - VI- 26 [BG- 19 - IDC- 001], I. N. DELA CRUZ leg. Mindoro Island, Oriental Mindoro, Mt. Halcon, 1 ex., 2005 - IV. Panay Island, Capiz, Dumarao, Bugsuan, 3 ♂♂, 11.14422 ° N, 122.44405 ° E 76 m a. s. l., 2019 - VIII- 03 [CP- 19 - IDC- 001], I. N. DELA CRUZ leg.; Antique, Patnongon, Igbobon, 1 ♂, 1 ex. [IC- 21 - 20], 10.55434 ° N, 121.59592 ° E - 10 m a. s. l., 2019 - VIII- 02 [AQ- 19 - IDC- 001], I. N. DELA CRUZ leg.; Iloilo, Calinog, Simsiman, 1 ♂, 11.07008 ° N, 122.32289 ° E 70 m a. s. l., 2019 - VIII- 01 [II- 19 - IDC- 001], I. N. DELA CRUZ leg. Guimaras Island, Guimaras, Jordan, Alaguisoc, 1 ♂, 10.37576 ° N, 122.36379 ° E 153 m a. s. l., 2019 - VII- 30 [GU- 19 - IDC- 001], I. N. DELA CRUZ leg. Negros Island, Negros Occidental, La Carlota, La Granja, 1 ♂, 10.23566 ° N, 122.59334 ° E 90 m a. s. l., 2019 - VII- 29 [NC- 19 - IDC- 002], I. N. DELA CRUZ leg.; Negros Occidental, Mt. Canlaon, 1 ex., 1988 - IV- 11 - 30, D. MOHGAN leg.; Negros Oriental, Tanjay, Azagra, 1 ex., 09.29363 ° N, 122.08473 ° E 0 m a. s. l., 2019 - VII- 31 [NR- 19 - IDC- 001], I. N. DELA CRUZ leg. Cebu Island, Cebu, Tuburan, Poblacion, 1 ♂, 10.43204 ° N, 123.49155 ° E 15 m a. s. l., 2019 - VII- 27 [CE- 19 - IDC- 001], I. N. DELA CRUZ leg. Mactan Island, Buyong Maribago, Lapu-lapu City, 1 ex., 1996 - IV- 3, S. SHIMANO leg. Mindanao Island, Agusan del Norte, Butuan, Tiniwisan, 1 ♂, 1 ♀ [IC- 21 - 11], 08.57694 ° N, 125.35521 ° E 20 m a. s. l., 2021 - V- 01 [AN- 21 - IDC- 001], I. N. DELA CRUZ leg.; Taligaman, 2 ♂♂, 08.56894 ° N, 125.38534 ° E 60 m a. s. l., 2021 - VI- 14 [AN- 21 - IDC- 003], I. N. DELA CRUZ leg.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C773C507E3D95545B2E224FEB32C0370.taxon	diagnosis	Diagnosis. Atholus coelestis (Marseul, 1857) is best characterized by its third dorsal elytral stria extending inwardly towards the apical end of the fourth and fifth striae. The slight emargination on the anterior margin of the mesoventrite is also a distinct character of this species. The number of denticles of the protibia (Figs 13, 14), is 11 on the outer margin, one one the inner apical angle, and eight on the outer sublateral margin. The protibial teeth are slightly prominent only on the outer apical angle, topped with three denticles. The number of denticles on the outer margin may range from 11 - 13 denticles. The shape of the gonocoxite of A. coelestis (Marseul, 1857) is slenderer, becoming narrower towards the apex compared to A. philippinensis (Marseul, 1854). Moreover, the presence of a single occipital fovea on the posterior portion of the head of A. coelestis (Marseul, 1857) (Fig. 11) is rather a remarkable character differentiating it from other species that has not been previously described.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C773C507E3D95545B2E224FEB32C0370.taxon	description	Additional description. Female genitalia: anterior portion of valvifers (Figs 38, 39) paddle-shaped; gonocoxite (Fig. 40) elongate, almost 4 x as long as broad, not shovel-like, more narrowed on apical end; inner and outer surfaces differentiated; inner face weakly separated from outer face by elevated lateral ridge; sclerotized setae on apical half of outer face short and sparse; inner face with short setae and moderate setae; apex of gonocoxite with two teeth; gonostyli present, freely articulated; spermathecae multiple, consisting of four sacs; sacs gradually enlarged and elongate, not sclerotized.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C773C507E3D95545B2E224FEB32C0370.taxon	distribution	Distribution. Widely distributed in the Oriental Region including China, Taiwan, Ryukyu Islands (Japan). Also present in the Palearctic Region: Tajikistan and in the Afrotropical Region: Comoros Islands (Mazur 2011).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C773C507E3D95545B2E224FEB32C0370.taxon	biology_ecology	Biology. All individuals of A. coelestis (Marseul, 1857) were collected from dungs of cows and water buffaloes of lowland farms and pastures across all islands of the archipelago. This species may also seem to be moisture-specific, as they were observed to dwell only on more desiccated dungs during field collection.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
A7A3931E063A533E9D6293C74FFDCD51.taxon	description	Figs 5, 50 - 53	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
A7A3931E063A533E9D6293C74FFDCD51.taxon	materials_examined	Specimens examined. Two syntypes of undetermined sex housed in MNHN have been examined by N. Degallier. The following re-description is based on images provided by him.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
A7A3931E063A533E9D6293C74FFDCD51.taxon	diagnosis	Diagnosis. This species is easily distinguished by its almost circular body and absence of sutural elytral striae. Judging by the images of two examined syntypes, this species is clearly distinct in its pattern of dorsal elytral striation, differing from other species by the absence of the fifth or sutural elytral striae. Atholus nitidissimus Desbordes, 1925 (Fig. 5) is similar to A. coelestis (Fig. 2), albeit it is comparatively smaller in size than other species examined.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
A7A3931E063A533E9D6293C74FFDCD51.taxon	description	Re-description. Body (Fig. 5) length: PEL: 2.15 mm; APW: 0.85 mm; PPW: 1.75 mm; EL: 1.15 mm; EW: 1.95 mm. Body almost circular, convex, and black; tibiae and antennae rufous. Head: clypeus (Fig. 50) slightly crenate on anterolateral margin, apical margin projecting; frontal stria round, complete, and moderately impressed; eyes clearly visible dorsally; mandibles with rounded outer margin curved inwardly; mandibular apex acutely pointed. Pronotum: marginal pronotal stria (Fig. 50) laterally complete, continuous onto the apical angle and behind head; lateral pronotal stria moderately impressed; apical end shortened and bent inwardly; lateral portion rather distant from margin; its basal end obsolete on basal sixth of pronotal length. Elytra: external and internal subhumeral striae absent (Fig. 5); oblique humeral elytral stria lightly impressed on basal third; dorsal elytral striae 1 - 3 complete; elytral stria 4 present on apical half or shorter; elytral stria 5 either absent or very short; sutural elytral stria absent. Propygidium and pygidium: propygidium (Fig. 51) moderately covered with coarse, round, and shallow punctures; interspaces with fine punctations; pygidial punctures similar to those of propygidium, albeit slightly denser. Prosternum: anterior margin of prosternal lobe (Fig. 52) round; medio-apical end ascending; marginal prosternal stria impressed, shortly interrupted medially; short striae present on both baso-lateral ends; prosternal lobe with several punctures alongside marginal prosternal stria on both sides; entire disk covered with finer punctures; prosternal suture moderately impressed; prosternal process with few fine punctures; lateral sides descending; lateral prosternal striae deeply impressed; basal half of prosternal process narrow. Meso- and metaventrite: anterior margin of mesoventrite (Fig. 53) truncate; marginal mesoventral stria complete; meso-metaventral suture clearly impressed, complete and carinate; lateral metaventral stria moderately impressed, carinate, extending obliquely and posteriorly, united with oblique humeral stria that inwardly extends from metaventro-metepisternal suture; post-mesocoxal stria extending posteriorly, strongly curved along the posterior mesocoxal margin, almost attaining the metaventro-mesepimeral suture; punctation of intercoxal disk of metaventrite similar to that of mesoventrite; longitudinal suture of metaventrite lightly impressed; lateral disk of metaventrite moderately covered with large, round, shallow punctures. Legs: posterior surface of protibia (Figs 52, 53) flattened and dilated; outer lateral margin with four teeth, topped with minute denticles.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
A7A3931E063A533E9D6293C74FFDCD51.taxon	distribution	Distribution. Endemic to the Philippines (Mazur 2011).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
A7A3931E063A533E9D6293C74FFDCD51.taxon	biology_ecology	Biology. Unknown.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
0EB5F543ECAF5E4D94ECB6C1959E4B83.taxon	description	Figs 1, 7, 8, 35 - 37	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
0EB5F543ECAF5E4D94ECB6C1959E4B83.taxon	materials_examined	Specimens examined. 3 ♂♂, 3 ♀♀. Mindanao Island, Agusan del Norte, Butuan, Taligaman, 3 ♂♂, 3 ♀♀ [IC- 21 - 18], 08.56894 ° N, 125.38534 ° E 60 m a. s. l., 2021 - VI- 02 [AN- 21 - IDC- 002], I. N. DELA CRUZ leg.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
0EB5F543ECAF5E4D94ECB6C1959E4B83.taxon	diagnosis	Diagnosis. Atholus philippinensis (Marseul, 1854) (Fig. 1) is easily distinguished from other Philippine congeners by entire dorsal elytral striae 1 - 3 (fourth stria is incomplete), and dense punctation of propygidium and pygidium. Among Philippine species, it is the largest one in size, with its markedly wider elytra and posterior angles of pronotum. The number of denticles of the outer sublateral margin of protibia is four.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
0EB5F543ECAF5E4D94ECB6C1959E4B83.taxon	description	Additional description. Female genitalia: anterior portion of valvifers (Figs 35, 36) paddle-shaped; gonocoxite (Fig. 37) slightly elongate, almost twice as long as broad, shovel-like; inner and outer surfaces differentiated; inner face moderately separated from outer face by elevated lateral ridge; sclerotized setae on apical half of outer face short and somewhat dense; inner face with short and sparse setae; apex of gonocoxite with two teeth; gonostyli present, freely articulated; spermathecae multiple, consisting of four sacs; sacs gradually enlarged and elongate, not sclerotized.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
0EB5F543ECAF5E4D94ECB6C1959E4B83.taxon	distribution	Distribution. Philippines; Malaysia; Indonesia (Sumatra, Borneo, Java); Myanmar, Vietnam; India (Meghalaya); China (Hainan); Taiwan (Mazur 2011).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
0EB5F543ECAF5E4D94ECB6C1959E4B83.taxon	biology_ecology	Biology. This species occurs in decaying banana stumps and are often found along with some species of Platylister (Platysomatini, Histerinae, Histeridae).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C9A24D84EC4257E5A9AA8C9D1560DAF9.taxon	description	Figs 4, 44 - 49	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C9A24D84EC4257E5A9AA8C9D1560DAF9.taxon	materials_examined	Specimens examined. Seven specimens of undetermined sex. Luzon Island, Laguna, northern Lucena, Kinabuhayan, 7 exs. [IC- 21 - 47], 1994 - V-VI, N. Monreal leg.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C9A24D84EC4257E5A9AA8C9D1560DAF9.taxon	diagnosis	Diagnosis. Atholus pirithous (Marseul, 1873) is generally recognized for its light excavation in the area behind the anterolateral angle of the pronotum.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C9A24D84EC4257E5A9AA8C9D1560DAF9.taxon	distribution	Distribution. Japan, Russia: Far East, China (Guandong, Shanghai), Korea, Taiwan, Vietnam, Nepal, Oman (Mazur 2011); Philippines (new record).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
C9A24D84EC4257E5A9AA8C9D1560DAF9.taxon	biology_ecology	Biology. Unknown.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
18C27BD6FEC55D468701010508D507FF.taxon	description	Figs 3, 15 - 20, 21 - 28, 29 - 34, 41 - 43	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
18C27BD6FEC55D468701010508D507FF.taxon	materials_examined	Specimens examined. 8 ♂♂, 14 ♀♀ and 7 exs. Luzon Island, Bataan, Abucay, Gabon, 8 ♂♂ [IC- 21 - 23], 12 ♀♀ [IC- 21 - 53], 5 exs., 14.42329 ° N, 120.26222 ° E 570 m a. s. l., 2019 - VII- 21 [BA- 19 - IDC- 001], I. N. DELA CRUZ leg.; Laguna, Northern Lucena, Kinabuhayan, 2 ♀♀ [IC- 21 - 49], 1989 - II, N. Monreal leg. Mindoro Island, Oriental Mindoro, Mt. Halcon, 1 ex., 2005 - IV. Palawan Island, Puerto Princesa, Barrio Talabigan, 1 ex., 1979 - III- 24, K. Wada leg.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
18C27BD6FEC55D468701010508D507FF.taxon	diagnosis	Diagnosis. Atholus torquatus (Marseul, 1854) is recognized with a combination of its interrupted lateral pronotal stria in the anterolateral angle, and fine punctations on the apical portion of its pygidium. This species also possesses remarkable teeth of protibia, increasing in size apically. The structure of the female genitalia of this species is described here for the first time, showing its similarity to the shape of the gonocoxite of A. philippinensis (Marseul, 1854), which is broad and shovel-like.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
18C27BD6FEC55D468701010508D507FF.taxon	description	Re-description. Male and female. Body length: PEL: 3.13 - 4.32 mm; APW: 1.11 - 1.47 mm; PPW: 2.35 - 2.90 mm; EL: 1.89 - 2.74 mm; EW: 2.66 - 3.56 mm. Body (Figs 3, 15 - 17) oval, moderately convex and black; tibiae, antennae, mouthparts and apical elytral margin rufous. Head: apical margin of clypeus (Fig. 18) short, entire and slightly forward, but anterolateral margin widely crenate; frontal stria rounded, complete and deeply impressed; disk sparsely clothed with fine punctures, separated by 2 - 3 x their diameter; interspaces with alutaceous microsculpture; occipital fovea absent; labrum dorsally finely punctate, raised and transversely long; short labral fringe (Lackner, 2010) present antero-laterally; mandibles covered with fine and even punctures, outer margin rounded, curved inwardly; sub-apical tooth on left mandible large; mandibular apex acutely pointed; eyes large and convex, clearly visible dorsally. Pronotum: marginal pronotal stria laterally complete, continuous onto apical angle and behind head; lateral pronotal stria (Fig. 19) deeply impressed, slightly crenate and complete; lateral stria rather distant from margin, its basal end abbreviated to basal fourth of pronotal length; apical end bent inwardly behind apical angle; anterior pronotal stria absent; disk with sparse microscopic punctures, wholly covered with alutaceous microsculpture; area behind apical angles bare; posterior margin without row of coarse punctures; ante-scutellar region with a single short longitudinal puncture. Elytra: basal margin with a row of short, longitudinal striae; elytral epipleuron sparsely clothed with fine punctures, with few, coarse punctures on apical half; marginal epipleural stria present on apical half; marginal elytral stria complete, moderately impressed; external subhumeral stria (Fig. 22) generally absent, occasionally noticeable on basal half, abbreviated on basal eighth; internal subhumeral stria absent; oblique humeral elytral stria slightly impressed on basal third; dorsal elytral striae 1 - 4 (Fig. 21) complete; elytral stria 5 present on apical half; sutural elytral stria abbreviated on basal third; elytral disk covered with sparse, fine punctures, separated by 3 - 4 x their diameter; medio-basal area with alutaceous ground sculpture. Propygidium and pygidium: propygidium (Fig. 26) densely covered with coarse, round and shallow punctures, about 25 µm in diameter, separated by 1 - 4 x their diameter; interspaces with irregular, sparse and fine punctations, separated by 2 - 3 x their diameter; surface with alutaceous sculpture; pygidial punctation (Fig. 25) similar to that of propygidium, coarse punctures of pygidium becoming sparser and finer apically; interspaces with fine punctations. Prosternum: prosternal lobe with anterior margin (Fig. 23) round; medio-apical end of prosternal lobe ascending; marginal prosternal stria deeply impressed, carinate and shortly interrupted medially; short striae present on both baso-lateral corners; lobe with few setiferous coarse punctures inside and outside of marginal stria on both sides, separated by their 1 - 2 x their diameter; disk covered with sparse, finer punctures on apical half; prosternal suture lightly impressed; prosternal process covered with few, setiferous fine punctures; lateral sides descending; lateral prosternal striae deeply impressed and complete; lateral disk with several coarse setiferous punctures; basal half narrow; posterior margin of basal lobe strongly emarginated. Meso- and metaventrite: anterior margin of mesoventrite outwardly arcuate (Fig. 24); marginal mesoventral stria complete, carinate, sparsely crenate; stria behind anterolateral angle present; mesoventral disk sparsely clothed with fine punctures separated by 4 - 5 x their diameter; meso-metaventral suture clearly impressed, complete and medially angulate; lateral metaventral stria deeply impressed, carinate, extending obliquely and posteriorly, united with oblique stria which inwardly extends from basal third of metaventro-metepisternal suture; post-mesocoxal stria extending posteriorly and strongly curved along posterior mesocoxal margin, almost attaining metaventro-mesepimeral suture; punctures of metaventral disk similar to those of mesoventrite; a row of coarse punctures present along inside lateral metaventral stria; longitudinal suture of metaventrite lightly impressed; lateral disk of metaventrite moderately covered with setiferous large round and shallow punctures; interspaces with sparse, coarse to fine punctations; mesepimeron, metepimeron and lateral disk of first abdominal ventrite with dense setiferous, large punctures; interspaces with few coarse to fine punctations; metepisternum with sparse punctures on apical half; punctation of intercoxal disk of first abdominal ventrite similar to that of metaventrite; lateral stria deeply impressed, slightly carinate and complete. Legs: anterior face of protibia (Fig. 27) flattened, dilated and clothed with few, fine ocelloid punctures; basal to median area with weak strigate sculpture; outer lateral margin with four teeth, becoming stronger apically; topped by minute denticles; protarsal groove shallow, with few coarse punctures; anterior protibial stria lightly impressed; inner marginal stria present on basal half, along stria a slightly depressed with row of coarse punctures present; near tarsal insertion with two spine-like tarsal denticles; another one, more distant and longer, located at inner anterior angle; protibial spur moderately long, wider on basal margin, approximately half the length of protarsus; posterior face of protibia (Fig. 28) with sparse, fine punctures and strigate ground sculpture from basal to median surface; number of denticles on outer margin eight, one on inner apical angle, outer sublateral margin three or four; median posterior stria moderately impressed and abbreviated on apical end; inner posterior stria moderately impressed with row of sclerotized setae, terminating in three inner posterior denticles; inner margin of setae present on apical half, with a row of short setae on basal half; inner margin with strigate ground sculpture; profemur sparsely clothed with fine, ocelloid punctations; surface with lightly strigate ground sculpture; marginal stria complete; anterior stria present on apical half; femoral stria almost complete, shortened on basal end; posterior margin with large punctations; a row of setae present on both basal and apical ends. Genitalia: aedeagus (Figs 29, 30) moderately slender, apically slightly curved ventrad; parameres relatively longer, about as almost as thrice the length of phallobase, slightly fused on basal half; median lobe sclerotized; eighth tergite (Figs 33, 34) entire, with longitudinal fold on both lateral sides; ninth tergite (Figs 31, 32) with lateral folds; tenth tergite dorsally longitudinally divided; spiculum gastrale almost as same length as ninth tergite. Anterior portion of valvifers (Figs 41, 42) paddle-shaped; gonocoxite (Fig. 43) slightly elongate, almost as twice as long as broad, shovel-like; inner and outer surfaces differentiated; inner face moderately separated from outer face by elevated lateral ridge; sclerotized setae on apical half of outer face short and slightly dense; inner face with short and sparse setae; apex of gonocoxite with two teeth; gonostyli present, freely articulated; spermathecae multiple, consisting of four sacs; sacs gradually enlarged and elongate, not sclerotized.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
18C27BD6FEC55D468701010508D507FF.taxon	distribution	Distribution. Widespread in the Oriental Region including Indonesia, Myanmar, Laos, Thailand, Vietnam, India, Nepal, and China (Sichuan) (Mazur 2011); Philippines (new record).	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
18C27BD6FEC55D468701010508D507FF.taxon	biology_ecology	Biology. Atholus torquatus (Marseul, 1854) were collected within the dung of cows located in a higher elevation and semi-forested area. The substrate also differs from A. coelestis (Marseul, 1857), as A. torquatus (Marseul, 1854) was typically observed in soggy, moist dung.	en	dela Cruz, Ian Niel, Ohara, Masahiro (2023): Revision of the genus Atholus Thomson, 1859 (Coleoptera, Histeridae, Histerinae) from the Philippines with additional records. ZooKeys 1158: 1-26, DOI: http://dx.doi.org/10.3897/zookeys.1158.100518, URL: http://dx.doi.org/10.3897/zookeys.1158.100518
