taxonID	type	description	language	source
803E6543FFDC992EFF4DF8F6FD43FF5D.taxon	description	The subtribe comprises four genera. It is represented in Asia with one species (Aspidoglossa subangulata), known from Japan. The morphology (shape E) of the external female genitalia represents a very different basic pattern. Members of the American genera Ardistomis Putzeys, 1846 and Semiardistomis Kult, 1950 investigated for validation (Tab. 2) show the same shape. Valdés (2012) already recognized the value of the female genitalia and separated the “ ardistomine type ” from the “ clivine type ”. In the absence of data from the American genus Kearophus Dajoz, 2004 the data confirm the distinctness of a separate line.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFDF992EFF4DFEE2FA98F879.taxon	description	The falciform Shape A 1 (Fig. 1) is found in approximately 70 % of the genera and species. In this group, a morphology is found which seems to represent a close shape to the plesiomorphic dimeric form constituting the ‘ nebridian configuration’ (according to Deuve 1993). Depending on the genus, the gonocoxite- 1 and gonocoxite- 2 are more or less fused with a more or less wide membranous area in between. Significant differences between the subgenera Clivina s. str. and Leucocara Bousquet, 2009 are not observed. The most notable variations are as follows: The subgenus Dacca shows more elongated, slender and sharp coxites similar to members of the C. helferi - species group. In the C. mekongensis species-group, gonocoxite- 1 and gonocoxite- 2 are not as intensely fused as in all other species and they are still a bit movable (Fig. 12). That shape is considered as the most plesiomorphic form. This find also justifies highlighting this group by assigning it herewith as C. mekongensis - species group. Rugiluclivina seems to be somewhat more specialized (Fig. 21). The coxites are slightly curved, gonocoxite- 2 is distinctly ensiform widened, and the gonocoxites- 1 are more strongly medially connected to each other (Fig. 22). It somewhat resembles the shape found in Forcipatorina. The following groups have been revised recently, all including figures of the coxites: Ancus (Balkenohl 2016), Clivina castanea - species group (Balkenohl 2021 a), Clypeospinus (Balkenohl 2021 b), Leucocara with the C. semicarinata - species group (Dostal & Bulirsch 2016), Rugiluclivina (Balkenohl 1996, 1999 a, 2015, 2018 a), Dacca (Balkenohl 2020). For Leleuporella, figures of coxites have been provided for two species (Fedorenko 2012, Bulirsch & Magrini 2019). Shape A 1 seems to be distributed over very wide geographic ranges, probably worldwide. The variations with one or two short macrotrichia (shapes A 4 and A 5) are considered to represent derived forms where the large seta is replaced by one or two short macrotrichia. The groups with shape A 4 with slender coxites have been revised recently with figures of the coxites provided: Orictites and its subgenus Semictites (Balkenohl 2017 a, b), and Paracoryza (Balkenohl 2000, 2005). The groups with shape A 5 with monomeric and conspicuously curved and short coxites were revised as well and comprise Syleter (Balkenohl 2021 b), Parasyleter (Balkenohl 2022), Trilophus (Balkenohl 1999 b), and Trilophidius (Balkenohl 2001). Another basic morphology is represented by the small and unusual monomeric foliform gonocoxite and semirectangular glabrous epipleurite without central articulation (shape B, Fig. 2). On a group level, Thliboclivina and Physoclivina have been revised and keyed out recently (Balkenohl 2001, 2018 c, Dostal 2015). Eoclivina is characterised and keyed out by Kult (1959) but without considering characters of the genitalia. The Oriental C. attenuata - species group is supposed to belong to Eoclivina as indicated in Lorenz (2005, 2022). However, this assumption is investigated here the first time. The C. attenuata - species group has not been revised in more than ninety years, with the last available treatment on a species level by Andrewes (1929). It has been found that, apart from the conspicuous differences of characters in the female genitalia, all members of these four species groups share several unusual external characters and combinations of characters not present in the other Clivinini. Based on the striking differences, the sixteen species have been integrated into a new subtribe characterised in this contribution (see below). The third basic pattern shows a monomeric hyaline musaceous gonocoxite and a glabrous rectangular epipleurite without central articulation (shape C, Fig. 47), also representing a different line. The genus Lophocoryza has been revised recently (Balkenohl 2018 b). Halocoryza was treated by Whitehead (1966, 1972) and revisited by Erwin (2011). Coryza is an unrevised genus with partly unclear synonymies, undescribed species, and partly unclear distribution. The fourth basic pattern shows monomeric hyaline elongated foliform gonocoxites and semi-square glabrous epipleurites with complete articulation (shape D, Fig. 55). The shape is found in Afrosyleter, a genus treated by Bulirsch & Magrini (2018) with a figure of the gonocoxites of one species. This shape seems to represent also a different line. However, the data published and the examples at hand are poor and need further insight.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFDE992FFF4DFE72FA03FD98.taxon	description	The subtribe includes currently twenty-four genera with nearly 200 species (Lorenz 2022). Many members of the subtribe show a shape similar to that of Trilophus, Trilophidius and Parasyleter (shape A 5). The subtribe is worth to be investigated separately in detail and has been found to be too huge to be included in this contribution.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD19922FF4DF966FB90FBBD.taxon	type_taxon	Type genus Thliboclivina Kult, 1959	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD19922FF4DF966FB90FBBD.taxon	synonymic_list	Taxa included: Thliboclivina Kult, 1959 microphthalma (Burgeon, 1937) diophthalmica (Basilewsky, 1955) amalita Basilewsky, 1964 bartolozzii Balkenohl, 2001 Eoclivina Kult 1959: 176 stat. nov. dumolinii (Putzeys, 1846) (= pumila (Lutshnik, 1926 )) machadoi (Basilewsky, 1955) burgeoni (Kult, 1959) Physoclivina Kult 1959: 214 stat. nov. physopleura (Burgeon, 1935) bulirschi (Dostal, 2015) donabaueriana (Dostal, 2015) dostaliana (Balkenohl, 2018) Sulciclivina gen. nov. attenuata (Herbst, 1806) striata (Putzeys, 1846) sulcigera (Putzeys, 1867) bhamoensis (Bates, 1892) sagittaria (Bates, 1892)	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD19922FF4DF966FB90FBBD.taxon	diagnosis	Subtribe diagnosis. General appearance similar to a small- to medium-sized subcylindrical Clivina. Head with clypeus bisinuate anteriorly in most of the species or more or less straight, in some species fused with clypeal wing; wing small; supraantennal plate conspicuously widened and conspicuously elongated posteriorly, in most of the species as wide as the eye, partially or completely continued over the eye and separating the eye either anteriorly or completely into two parts; frons separated from clypeus by a carina or a sulcus in form of a flat inverted V; frons in some species with second carina in form of an inverted V or with small depression; with two pairs of supraorbital and one pair of clypeal setae; labrum five- or six-setose; eye small, reduced in many species, separated partially or completely into two parts by lateral carina of supraantennal plate; mandible moderately short, flattened, scrobe small, asetose; antennomeres four to ten moniliform or submoniliform. Pronotum with lateral margin running basally in a regular curve up to base, with two lateral setae, base narrow. Proepisternum tumid laterally, visible dorsally. Elytron with striae 1 - 3 free at base, intervals asetose. Mesotibia dorsally with two distinct parallel rows of short protuberances furnished with robust seta each, laterally with row of strong setae, medially with up to five fine setae. All sterna punctured, but to a different degree among the groups. Last visible sternum with the two apical setigerous punctures separated. Female genitalia very small in comparison to body size, with gonocoxite foliform, monomeric, flattened dorsally and ventrally, less sclerotized to hyaline, with few setae; the coxites are medially not attached but separate; epipleurite semi-rectangular and asetose, the median joint is replaced by a broadly angulated and more or less widened flat rod. Differential diagnosis. Distinguished from all other Clivinini by the outstanding external female genitalia, the elongated and widened form of the supraantennal plates separating the eyes anteriorly or completely into two parts, the small pronotal base, and the tuberculate mesotibia.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD19922FF4DF966FB90FBBD.taxon	biology_ecology	Habitat preference. Ecological data are not published. However, among the 640 specimens checked (MRACT, RBINS, CMBB), there were not seldom ecological data noted on the labels. According to these data, Physoclivina, Eoclivina, and African species of Thliboclivina were collected in detritus from vegetation and in humus of swamp forests, whereas the single Oriental Thliboclivina species was found in an open rice field. For the Oriental genus Sulciclivina gen. nov., data indicate the same habitats but also cultivated areas (bamboo forest, swamp forest, garden, plantation, zebu dung, ground vegetation covered by litter). In summary, the subtribe seems to populate humus underground in shaded humid and wet areas.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD19922FF4DF966FB90FBBD.taxon	discussion	Remarks. Clivina arunachalensis Saha & Biswas, 1985 may possibly belong into the subtribe as well. It is tentatively not included because some key characters are not mentioned in the description nor visible on the ink drawing, giving space for interpretation of the position. In the description, a close relationship to members of the genus Pseudoclivina is emphasized. The type material has not been seen or investigated after the description. Inquiries to the place of deposition of the type material remained unanswered.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD19922FF4DF966FB90FBBD.taxon	discussion	Thliboclivina and Physoclivina have been intensively characterised recently on a species and group level (Dostal 2015, Balkenohl 2001, 2018 c). Eoclivina has been characterised by Kult (1959).	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD39924FF4DFBADFEBDFD69.taxon	description	(Figs 2, 6)	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD39924FF4DFBADFEBDFD69.taxon	type_taxon	Type species: Sulciclivina attenuata (Herbst, 1806): 264 (Scarites attenuatus).	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD39924FF4DFBADFEBDFD69.taxon	materials_examined	Type locality: Assam, India. Genus diagnosis. Size 5.4 – 9.5 mm. Colour piceous. Head with clypeus distinctly widened, bisinuate anteriorly, lateral angles slightly projecting, fused with clypeal wing, supraantennal plate conspicuously widened, elongated posteriorly, separated from clypeus by distinct notch, clypeal suture developed as sulcus in form of a flat inverted V, clypeus and frons separated from supraantennal plate by deep furrow; frons with central depression developed in some species horseshoe-like; with pair of clypeal seta and two pairs of supraorbital setae; eye moderately protruding laterally, supraantennal plate overtopping eye in anterior third; gena flattened, size about one sixth of eye; mandible flattened, wide; labrum six setose, antenna with segments five to ten slightly longer than wide. Pronotum as long as wide, with reflexed lateral margin running from anterior angle up to base, posterior angle rounded off without tooth, marked by the posterior marginal seta, lateral channel of equal width, median line narrow, joining base; proepisternum visible basally in dorsal view; base conspicuously narrow, flange at base indistinct. Elytron long oval to subcylindrical, with striae one to three free at the base, striole short, indistinct, lateral channel with row of umbilical setigerous punctures complete; third interval without setigerous punctures. Hind wing fully developed or reduced. Mesotibia dorso-laterally with row of ten short protuberances and in parallel dorso-medially with row of four protuberances, the apical ones slightly larger, all furnished with robust seta, laterally with row of twelve strong setae, medially with up to ten fine setae. Male genitalia robust, with aedeagus distinctly curved, apical part with or without widened lamella. Female genitalia (Fig. 2) small in comparison to body size, with gonocoxite foliform, monomeric, flattened, usually with a long apical nematiform seta and depending on the species with three to six additional nematiform setae positioned at the median side, epipleurite without setae and without central joint.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD39924FF4DFBADFEBDFD69.taxon	diagnosis	Differential diagnosis. See key to the genera of the subtribe. Moreover, the antennomeres five to ten are slightly longer than wide (around L / W 1.07) (Eoclivina and Physoclivina L / W around 1.0; Thliboclivina L / W around 0.8), the pronotum with the anterior angles are slightly more acute (more rounded off in Eoclivina), the lateral marginal channel of the pronotum which runs in parallel to the margin is not widened between the two lateral setigerous punctures and the base is much narrower compared to the other three genera (see ratio WP / WB provided in the key). Some species of the Neotropic Clivina subgenus Paraclivina Kult, 1947 show slight similarities of the widened and elongated supraantennal plates and also an inverted V is found on the frons of the head (e. g., Clivina bipustulata (Fabricius, 1798), C. fassatii Kult, 1947). However, these species belong to a different group due to their distinctly falcate coxites, the triangle like and plurisetose epipleurite with complete joints, the striae one to five free at the base of the elytron, the presence of setae on interval three of the elytron, and the non-tuberculate mesotibia.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD39924FF4DFBADFEBDFD69.taxon	etymology	Etymology. The name refers to the sulcus on the frons of the head and is composed of the Latin noun “ sulcus ” (= furrow) and Clivina, the lead genus of the tribe.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD39924FF4DFBADFEBDFD69.taxon	distribution	Distribution. Centre of the Oriental region. Due to the partly incorrect synonymies in Andrewes (1929), the species of this genus are in need of reassessment.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD59924FF4DFC98FC95FB7B.taxon	distribution	The genus Ancus Putzeys is known to occur in the Neotropical region with three species and in the Oriental region with four species. Oriental members have been revised recently including a full genus description (Balkenohl 2016).	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD59924FF4DFC98FC95FB7B.taxon	description	When checking the external female genitalia of African members for this contribution, the species Clivina svenssoni Basilewsky, 1946 from Africa came to my attention once again. For some time, it had been assumed that the species may belong in the genus Ancus. This assumption has been challenged now by comparing the gonocoxites and epipleurites with those of the Oriental members, and they are indeed extraordinary similar. The most similar species regarding the coxites is A. hiekei Balkenohl, 2016, with only minute differences in the positon of the setae. Consequently, the species is assigned to the genus Ancus.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD59924FF4DFB2CFDBFFA7E.taxon	diagnosis	Differential diagnosis. The species differs from all other species mainly by the surface of the pronotum. Instead of the two sulci, the African species shows a wide paramedian and paralateral longitudinal cloud of distinct punctures and the Neotropical species displays smaller longitudinal bands of punctures. In addition, the abdomen does not show punctures as in the other members but a distinct isodiametric reticulation.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
803E6543FFD59924FF4DFB2CFDBFFA7E.taxon	distribution	Distribution. Mali and Senegal.	en	Balkenohl, Michael (2022): On the female gonocoxites of Asian, Arabian, and African Clivinini with realignment of the Thliboclivina-, Physoclivina-, and Eoclivina-species groups, and notes on the higher systematics of the tribe Clivinini Rafinesque, 1815 (Coleoptera, Carabidae). Zootaxa 5190 (1): 99-126, DOI: https://doi.org/10.11646/zootaxa.5190.1.4
