taxonID	type	description	language	source
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	description	(Figs 3 – 6, 11 – 13)	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	materials_examined	Type material. Holotype male, labelled: “ GEORGIA, Racha-Lechkhumi and Kvemo Svaneti region, Racha karst massif, Usholta Cave, dark zone, 21. viii. 2022, E. Maghradze leg. ” (IZISU). Paratypes (21 ♂♂, 19 ♀♀): 1 ♂, 2 ♀♀: same data as holotype (IZISU); 3 ♂♂, 2 ♀♀: same data as holotype, but 25. vii. 2022, S. Barjadze leg. (IZISU); 1 ♂, 1 ♀: same data as holotype, but 04. v. 2023, E. Maghradze leg. (IZISU); 6 ♂♂, 6 ♀♀: same data as holotype, but 23. vi. 2023, E. Maghradze & A. Faille leg. (3 ♂♂, 3 ♀♀ - IZISU; 3 ♂♂, 3 ♀♀ - SMNS); 1 ♂, same data as holotype, with extraction code SMNS-L 2354 (SMNS); 1 ♀: same data as holotype, but 11. ix. 2022, J. Grego lgt., with extraction code RL- 89 (CRL). Spares in ethanol: 1 ♂ - EM 68 (25. vii. 2022, S. Barjadze leg. (IZISU )) and 2 ♂♂, 1 ♀, E. Maghradze & A. Faille leg., 23. vi. 2023 (SMNS).	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	diagnosis	Diagnosis. Medium-sized anophthalmous trechine beetle; body depigmented; coloration homogeneous, yellowish to light brown; body strongly flattened; microsculpture almost indistinct, except on the neck where a few transverse meshes can be distinguished. Head longer than wide; mandibles short and robust; frontal furrows complete; eyes reduced, without trace of ommatidia; submentum with 6 setae. Pronotum strongly cordate, wider than long. Elytra, and appendages elongate, narrow; elytral striae almost effaced; two discal setiferous pores and one preapical pore on each elytron. Abdominal ventrites mostly glabrous with a pair long paramedian and short irregulary sinuated setae around paramedian pair. Habitus as in Figs 3 – 6.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	description	Description. BL: 5.5 – 6.7 mm. Head (Figs 3 – 6): glabrous, longer than wide, longer and slightly narrower than pronotum, HL 1.21 – 2.4 mm, HW 0.96 – 1.24 mm, Index HL / HW 1.2 – 1.5. Temples convex, neck constriction distinct. Two pairs of supraorbital setae present, frontal furrows shallow, divergent anteriad, incomplete, reaching posterior frontal pore. Head with a strong depression behind the second supraorbital setae and reaching the base of the neck (Figs 5 – 6). Eyes reduced to a small whitish area, without trace of ommatidia, two pairs of anterior supraocular setae (Fig. 6). Clypeus quadrisetose; labrum concave, with six setae situated near the anterior margin, extreme ones being the longest and anterior lateral margins with 7 – 10 short setae. Mandibles short and robust, sharp; right mandible tridentate, left bidentate. Mentum bisetose, with a simple, protruding tooth; labial suture complete, not fused; submentum with 6 setae (Fig. 5); maxillary palpi short, glabrous (some specimen with a tiny seta on palpomere II); labial palpi thin and long, glabrous except penultimate palpomere, which is bisetose. Antennae elongate, filiform, pubescent from antennomere II, scape with few setae; AL 4.5 – 5.4 mm, length of antennomeres (from scape to terminal segment, in mm): 1.00 / 0.78 / 1.50 / 1.47 / 1.5 / 1.39 / 1.33 / 1.22 / 1.17 / 1.06 / 1.19. Pronotum: small and strongly cordate, slightly wider than long, PL 0.86 – 1.11 mm., PW 1.05 – 1.27 mm, Index PL / PW 0.71 – 0.9, with narrow protruding base, lateral margins strongly sinuate before hind angles, widest at anterior fourth. Pronotal base deeply foveolate near posterior angles, narrowed, the basal margin much shorter than the anterior; hind angles distinct, prominent, strongly and upwardly protruding; anterior angles widely rounded. Two pairs of long and thick pronotal setae present, antero-lateral pair situated in anterior sixth of pronotal length, basolateral setae near posterior angles (Fig. 3). Elytra (Fig. 3): long and narrow, gradually widened towards apex, EL 3.1 – 3.6 mm, EW 1.85 – 2.24 mm, index EL / EW 1.55 – 1.69. Humeri rounded; elytral striae almost effaced, barely visible. Elytra slightly convex, with flattened disc. Scutellar striola absent, parascutellar seta present. Elytra with two pairs of long and thick discal and one similar preapical setiferous pore, preapical pair located near elytral base in stria 3, discal pores situated in the anterior half of elytral length of stria 2 and 3. Humeral group of umbilicate series aggregated, pores 1 – 4 slightly separated from marginal gutter; pore 2 equidistant from pores 1 and 3, distance between pores 2 and 3 longer than between pores 3 and 4. Legs: Long and slender; protibiae without sulcus on dorsal side, antennal cleaning organ well developed; first two tarsomeres of male protarsi dilated at internal margins; tarsi pubescent, claws long and acute. Abdominal ventrites (Fig. 4): mostly glabrous, central part with a few (6 – 19) irregularly situated short, and a pair of long paramedian setae; last ventrite for both sexes with two pairs of setae, one pair longer and one relatively shorter, the difference between female and male is the arrangement of setae, for female they make transverse row (anterior is longer), male setae are longitudinal (posterior is longer). Male genitalia: median lobe of aedeagus relatively narrow in lateral view, wide at base, broadly curved; apical part gradually curved upward, elongate, with an upturned apex (Figs 11 – 12). Internal sac (endophallus) armature poorly sclerotized, paired and symmetrical, covered with a broad field of uniformly sized microtrichia, very similar to that of allied taxa. Parameres with four long, straight apical setae each. Female genitalia: gonocoxite II triangular, distinctly sclerotized, slightly longer than wide, with 4 ensiform spines on the dorsal and lateral surface and small apical groove that harbours two short nematiform setae; gonocoxite I simple, wide, slightly sclerotized, with 4 to 5 long setae on the inner margin and 2 to 4 relatively short randomly distributed setae, on ventral projection; laterotergite less sclerotized with 13 – to 15 setae on the inner margin and 10 – to 16 on the basal half (Fig. 14).	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	diagnosis	Differential diagnosis. Inotrechus usholtaensis sp. nov. differs from the other two Inotrechus species (see Identification Key) by the following combination of characters: a) submentum with 6 setae in the new species, whereas 8 in I. injaevae and I. kurnakovi; b) head large, rounded in the new species, whereas narrow and relatively long in I. injaevae and I. kurnakovi; c) mandibles short and robust in the new species, while I. injaevae and I. kurnakovi have long and slender mandibles; d) reduced eyes in the new species, with a small whitish area on top and no trace of ommatidia, similar to I. kurnakovi, whereas I. injaevae has reduced eyes with a short black line and no ommatidia; e) pronotum distinctly wider than long in the new species and I. injaevae, while, in I. kurnakovi, it is almost as long as wide; f) elytral striae barely visible, indistinct in the new species, while clearly visible in I. injaevae and I. kurnakovi.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	etymology	Etymology. Named after the type locality, Usholta Cave.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	distribution	Distribution. So far, the species is only known from its type locality, Usholta Cave in the Racha karst massif, Racha-Lechkhumi and Kvemo Svaneti regions, Western Georgia (Map 1). Caves in this area are poorly sampled, and I. usholtaensis sp. nov. may be found in additional localities in the future.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C513FFDD5D9EF099B8F1FC6B.taxon	biology_ecology	Ecology. The species is troglobitic and found throughout the cave, typically near the subterranean river on gravel banks, wood debris brought from outside (incl. rests of old wooden ladder), or under stones. The fauna of Usholta Cave is poorly studied, with only seven species recorded to date: Triphosa dubitata (Linnaeus, 1758) (moth), Acanthophorella valerii Antić, 2023 (millipede), Deuteraphorura sp. and Plutomurus weinerae Barjadze, Kováč & Parimuchová, 2024 (springtail), Rilaena zakatalica Snegovaya & Chemeris, 2005 (harvestman), Neobisium cf. verae (Lapschoff, 1940) (pseudoscorpion), and Laemostenus (Antisphodroides) ljovushkini Vereschagina, 1985 (carabid beetle) (Barjadze et al. 2015, 2024; Antić et al. 2023). Based on above mentioned differences, all three representatives of the genus Inotrechus can be easily identified using the following key:	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	description	(Figs 17 – 21)	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	materials_examined	Type material. Holotype male, labelled: “ GEORGIA, Racha-Lechkhumi and Kvemo Svaneti region, Racha karst massif, Kvedi II Cave, dark zone, 24. v. 2023, E. Maghradze leg. ” (IZISU). Paratypes: (4 ♂♂, 4 ♀♀): 2 ♂♂, 2 ♀♀: same data as holotype, but 07. xii. 2023 (IZISU); 1 ♂, 1 ♀: same data as holotype, but 07. xii. 2023, (SMNS); 1 ♀: same data as holotype, but 12. ix. 2022, J. Grego leg., with extraction code RL- 88 (CRL). Stored in ethanol: 1 ♂ - EM 119 (07. xii. 2023, E. Maghradze leg. (IZISU )).	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	diagnosis	Diagnosis. Medium sized Trechine beetle, with the character states of the genus Duvalius subgen. Duvalius Delarouzée, 1859 (sensu Jeannel, 1928). Body elongated, depigmented, reddish-brown, glabrous, with strongly reduced eyes Microsculpture on head, submentum with 6 or 7 setae, pronotum and elytra well distinct, composed of transverse meshes. Trapezoid pronotum, almost as wide as long. Elongate, ovoid elytra with well developed striae; indistinct, widely rounded humeri. Male protarsi with the first two tarsomeres dilated and toothed at their internal magrins. Habitus as in Figs 17 – 19.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	description	Description. BL: 4.97 – 5.8 mm. Head (Figs 17 – 19): glabrous, longer than wide, distinctly longer and narrower than pronotum, HL 1.05 – 1.2 mm, HW 0.85 – 0.99 mm, Index HL / HW 1.15 – 1.32. Temples convex, neck constriction distinct. Frontal furrows deep, complete, reaching neck constriction. Head with two pairs of anterior supraocular setae and two pairs of long supraorbital setae, anterior pair behind middle of head length, posterior pair at hind part of head near the neck. Mandibles relatively long and slender, acutely pointed, the right one with tridentate basal teeth. Eyes flattened, strongly reduced, with only few ommatidia (Fig. 19). Clypeus quadrisetose; labrum concave, with six setae near the anterior margin, extreme ones being the longest and anterior lateral margins with 4 to 6 short setae. Mentum bisetose, with a bifid, protruding tooth; labial suture complete, mentum and submentum not fused; the latter with 6 or 7 setae; maxillary palpi short, glabrous; labial palpi glabrous except penultimate palpomere, which is quadrisetose. Antennae elongate, filiform, stretching backward reaching middle of elytra; pubescent from antennomere ii, scape bearing a few setae; AL 3.26 – 3.76 m.; length of antennomeres (from scape to terminal segment, in mm): 1.00 / 0.89 / 1.36 / 1.29 / 1.29 / 1.25 / 1.10 / 1.03 / 1.00 / 0.96 / 1.21. Pronotum: Trapezoid, almost as wide as long, PL 0.87 – 1.05 mm, PW 1.08 – 1.21 mm. Index PL / PW 0.81 – 0.96; with maximum width in anterior fifth. Lateral sides slightly rounded and strongly sinuate before posterior angles, which are distinct and protruding; anterior angles rounded, obtuse. Lateral furrows well developed, deep, with two pairs of setae; anterolateral setae situated in the anterior fifth, basolateral pair before hind angles. Basal impressions deep, not punctuated; median furrow weakly marked, visible in middle of pronotum (Figs 17 – 18). Elytra: ovoid, elongate, EL 2.84 – 3.19 mm, EW 2.11 – 1.83 mm, Index EL / EW 1.55 – 1.58; with maximum width before the middle of elytral length. Humeri indistinct, widely rounded, scutellum small, flat; single pair of parascutellar seta present; scutellar striola absent. Elytral disc convex, elytral striae well-developed, distinctly punctate. Elytra with two pairs of long and thick discal and one similar preapical setiferous pore, preapical pair located near elytral base in stria 3 and 4, discal pores situated in the anterior half of elytral length of stria 3 and second pairs of stria 2 and 4 (Figs 17 – 18). Legs: relatively short; protibiae with sulcus on dorsal side, antennal cleaning organ well developed. First two tarsomeres of male protarsi distinctly dilated and toothed at their internal margins. Tarsal claws long and slender, pointed at apex. Abdominal ventrites: Glabrous, with a pair of short paramedian setae; last ventrite with one seta each side in males, two seta each side in females. Male genitalia: Median lobe of aedeagus very narrow, elongated, and gently curved in lateral view; slightly wider at basal bulb, subparallel, almost straight in dorsal view. Apex elongate, shortly curved upward. Internal sac of aedeagus (endophallus) poorly sclerotized. Parameres slender, each with four long, straight apical setae (Figs 20 – 21). Female genitalia: gonocoxite II, distinctly sclerotized, unguiform, slightly longer than wide, with 3 – 4 ensiform spine on the dorsal and lateral surfaceand apical groove that harbours two small nematiform setae; Gonocoxites I simple, triangular, weakly sclerotized, with 4 – 5 long setae on the inner margin, near the corner and 3 – 5 setae on the distal margin; laterotergite less sclerotized with 12 – 15 setae near the inner margin and 6 – 7 in the basal half on the disc (Fig. 13).	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	diagnosis	Differential diagnosis: Duvalius gregoi sp. nov. is the fourteenth species of the genus described from the Caucasian region. All Caucasian Duvalius species belong to the subgenus Duvalius s. str. and exhibit significant variation in morphological traits, reflecting various stages of adaptation to the subterranean environment. In general, the body is more or less flattened and varies in the following morphological characteristics: • colour ranges from piceous / reddish-brown to pale testaceous; • eyes from strongly reduced, flat with almost no ommatidia to slightly reduced, prominent eyes; • legs and antennae from thin and long to thicker and shorter. To date, five Duvalius species have been recorded from Georgia: D. abyssimus Reboleira & Ortuño, 2014, D. iljukhini (Dolzhanski & Ljovuschkin, 1985), D. kurnakovi Jeannel 1960, D. megrel Belousov, 1991 and D. sokolovi Ljovuschkin, 1963 (Belousov, 1991; Belousov & Zamatajlov 1995; Barjadze et al. 2019; Reboleira & Ortuño, 2014). Duvalius gregoi sp. nov. appears to be most closely related to D. sokolovi Ljovuschkin, 1963 from the Abkhazeti region, Trechus’ Cave. Both species share similar reddish-brown body coloration but can be easily distinguished by the following combination of characters: a) Body size: the new species is smaller, with a body length of 4.97 – 5.8 mm, while D. sokolovi measures 6.0 – 6.2 mm in length. b) Pronotum: trapezoid and slightly wider than long in the new species, whereas in D. sokolovi, the pronotum is longer than wide.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	etymology	Etymology. Patronymic, dedicated to our dear friend Jozef Grego (Horná Mičiná, Slovakia) enthusiastic speleologist and biospeleologist, specialist on subterranean snails and discoverer of this new species.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	distribution	Distribution: The species is known only from the type locality, Kvedi II Cave, located in the Racha karst massif, Racha-Lechkhumi and Kvemo Svaneti regions, Western Georgia (Map 1). The caves in this area are biologically underexplored, and D. gregoi sp. n. may be found in adjacent localities.	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
03CB2A29C517FFD05D9EF52DBDA2FC8B.taxon	biology_ecology	Ecology: The species is troglomorphic and was found in a small crack at the end of the cave. The fauna of Kvedi II Cave is poorly studied; so far, only one species of snail, Mucronaria kartvelica Mumladze, Grego & Szekeres, 2023, has been recorded from this cave (Mumladze et al. 2023). All Duvalius species known so far from the Caucasus can be identified using the following key:	en	Maghradze, Eter, Barjadze, Shalva, Lohaj, Roman, Faille, Arnaud (2025): Two new Trechini (Coleoptera: Carabidae) from Georgian caves, Caucasus. Zootaxa 5609 (1): 83-96, DOI: 10.11646/zootaxa.5609.1.5, URL: https://doi.org/10.11646/zootaxa.5609.1.5
