taxonID	type	description	language	source
03F0A80AFFC5A221FF28F9CD6B5E2373.taxon	discussion	Acalles lederi Meyer, 1896 was placed in the newly created monotypic genus Caucasusacalles in 2018 (Stüben 2018). The holotype, a male from Svaneti collected by H. Leder in the collection of E. Reitter, was studied by myself in 2003 in the Hungarian Natural History Museum. The only specimen was prepared by V. Savitsky, the aedeagus was removed and added to the plate (see also in Fig. 3, bottom right). Digital photographs of the habitus and aedeagus were made and have already been published (Stüben et al. 2003: Figs. 333.1 / 333.2). In fact, however, Caucasusacalles lederi is a species complex consisting of at least two further species, as both molecular studies and genital preparations now available show. The inner sac structure (sclerites or socalled ‘ aggonoporium’, Morimoto et al. 2015) in the endophallus, which is unique among the Western Palaearctic Cryptorhynchinae and consists of parallel bars connected on the upper side, is constant within a species, but interspecifically extremely rich in form. In my book on ‘ The Cryptorhynchinae of the Western Palearctic’ (Stüben 2018), I paid great attention to the distinctive and unique internal sac structure of the aedeagus in the description of the new genus Caucasusacalles, and overlooked the fact that I had already illustrated the aedeagus (and habitus) of another species from the C. lederi complex, which was not congruent with the holotype from the Reitter collection mentioned above (Stüben 2018: 24). In reality, this is the adelphotaxon Caucasusacalles subglobosus sp. nov. to be described below. Another species to be separated is Caucasusacalles circularis sp. nov.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC4A226FF28FE206A8827F3.taxon	description	(Fig. 2)	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC4A226FF28FE206A8827F3.taxon	materials_examined	Type material Holotype: 1 ♂, ‘ USSR, Caucasus occ., Kozljacij Brod, Mzymta bank, R. Borovec lgt. / 15 km E Adler pr. Sochi, 29. – 31.5.1985, 500 m’, coll. SDEI, for photo documentation see Fig. 2 // Paratypes: 1 ♂, 1 ♀, same data as for holotype, coll. SDEI, Borovec / 1 ♂, „ USSR [Georgia], Pic [z] unda, 6.69, Kucura ”, coll. Stüben; 2 ♂, 3 ♀, ‘ USSR [Georgia] — Ossetien, Diava [Java] env., V. 87, lgt. Rous’, coll. Stüben & Krátký; 1 ♂, ‘ USSR [Georgia] — Ossetien, Gufta [Gupta], V. 87, lgt. Rous’, coll. Stüben; 1 ♀, ‘ Sotschi, Circassia’, coll. Stüben; 1 ♀, ‘ Russia, Krasnodar, Chvizhepse vill. env., 43 ° 38 ' 32 " N, 40 ° 04 ' 45 " E, 300 m, 10.7.2014, leg. A. V. Kovalev’, coll. ZFMK, DNA: Collecting no. 2754 - PST (in ZFMK), mtCO 1 sequence see Appendix 2. Length. 1.9 – 2.5 mm (without rostrum) Head & Rostrum. Eyes clearly visible from above, round towards forehead when viewed laterally and from the front, but tapering to underside of rostrum; forehead broad, almost as broad as base of rostrum; flattened rostrum of male 3.1 × as long as wide between antennal insertions, densely covered with light-coloured scales in upper part, finely punctured in apical half; rostrum of females 4.0 × as long as wide, scaly only immediately in front of base, otherwise shinier and much more finely punctured towards apex; when viewed laterally rostrum only slightly curved immediately in front of base, otherwise straight to apex in both sexes; antennae short, club-shaped scape not as long as width of dorsum of rostrum between antennal insertions; antennomere 1 of funicle conical, approx. 2 × as long as at widest point, antennomere 2 very narrow, only slightly longer than wide, antennomeres widening from 3 rd to 7 th segment, slightly shorter than wide; club short-oval.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC4A226FF28FE206A8827F3.taxon	description	Pronotum. 1.1 × as wide as long, widest at or immediately behind middle, there laterally strongly convex, towards basal and anterior margins — there with a slight depression-narrowed in a straight line; fine punctation covered with large round, light, tightly fitting scales, only occasionally with dark brown, erect bristles condensed into small tufts on disc. Elytra. Inversely ovate, but at most 1.1 x as long as wide; widest at about end of basal two-fifths; striae with fine, elongate punctures clearly narrower than slightly arched intervals, which are several times wider; intervals also predominantly covered with round, light and dark brown scales, which often show a banded pattern, albeit irregularly limited; erect, beige and brown, somewhat flattened scales, which are placed at large intervals in a single row on interstriae are at most 1.5 times as long as wide. Legs. Legs, but especially femora, long; fore femora reaching to base of rostrum, hind femora reaching apex of elytra; covered with spindle-shaped, white and light brown scales, which do not completely cover underlying cuticle; scales protruding slightly on th outer faces of tibiae. Underside. Venter with a deep rostrum channel whose mesosternal receptaculum is semicircular and ends in front of mid-coxae; abdomen with five segments with light-coloured, widely spaced scales, each scale 3 × – 3.5 × as long as wide; 2 nd ventrite strongly punctate, sloping obliquely to narrow 3 rd and 4 th ventrites, only slightly longer than 3 rd and 4 th combined. Aedeagus. Median lobe of aedeagus flatly rounded (if the aedeagus aligned exactly ventrally), but ovally rounded when only the tip rests flat; endophallus consisting of two narrow, closely spaced bars, each with an inwardly inclined long, spine-like tip (Fig 2).	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC2A225FF28FB776CFA22EE.taxon	description	(Fig. 4)	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC2A225FF28FB776CFA22EE.taxon	materials_examined	Type material Holotype: 1 ♂, ‘ Armenia: around Arakelots Monastery ruins, W Kirants, Tavush, 21.5.2022, leaf litter sifting and Winkler extractor, 41,032000 45,066000, deciduous forest, leg. Astrin’, coll. ZFMK, ZFMK-TIS- 8010636, mtCO 1 sequence see Appendix 2; for photo documentation see Fig. 4 // Paratypes: 1 ♂, ‘ Georgia: 9 km E of Tianeti, on Aniskhevi River banks, forest next to river and affluent creek, Mzkheta-Mtianeti, 42,08619 45,04298, 1051 m, leg. Astrin’, coll. Stüben, mtCO 1 sequence see Appendix 2 / 1 ♂, ‘ Georgia: E of Saparlo, Kvemo Kartli, 41,3021 44,3178, Quercus, 16.11.2023, leg. Astrin’, coll. ZFMK, ZFMK-TIS- 8028288, mtCO 1 sequence see Appendix 2 / 1 ♂, 3 ♀, ‘ Georgia, Kharagauli, Merelisi env., 482 m, 41 ° 56 ' 26 " N 43 ° 17 ' 01 " E, 29.8.2019, sifting in mixed forest, lgt. Baňař, Hlaváč, Barjadze & Maghradze’, coll Krátký / 1 ♂, ‘ Georgia: „ Kaukasus, Suram-Pass ”, „ Acalles lederi Meyer <handwriting>, Penecke det. ’, coll. Senckenberg Naturhistorische Sammlungen Dresden / Georgia: 1 ♂, ’ Suram-Pass, 2010, Kulzer ”, “ Acalles choudoiri, Zumpt det. ’, coll. Naturhistorisches Museum in Wien.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC2A225FF28FB776CFA22EE.taxon	description	Description & differential diagnosis Length. 2.1 – 2.5 mm (without rostrum) The new species C. circularis from Armenia (type locality) as well as from central and southern Georgia hardly differs externally from the previously described species C. subglobosus. The few short, erect bristles on the very broad elytral intervals are also conspicuous in this species. Likewise, the adjacent scales on both the pronotum and the elytra are predominantly circular, characteristics that make these two species easily distinguishable from all other Cryptorhynchinae of the Caucasus. While the elytra of C. circularis are somewhat more broadly rounded towards the apex (not slightly obovate as in C. subglobosus) and the rostrum is somewhat more robust, deeper and longitudinally punctate towards the edges (Fig. 4), the only reliable and constant distinguishing feature is the sclerite complex in the endophallus. In C. circularis this consists of widely spaced, parallel, narrow bars connected in the upper part by a pointed, but not so strongly sclerotised bridge as in C. circularis, whereas in C. subglobosus the sclerite consists of closely spaced bars, each with an inwardly inclined long, spine-like tip (compare Fig. 2 vs. Fig. 4). In this character, C. circularis actually resembles C. lederi, the type species of the genus Caucasusacalles (type locality: Georgia, Svaneti Region). However, in C. lederi, the sclerite of the endophallus consists only of two simple, subparallel bars and does not show, as in C. circularis, further sclerotised, angularly separated, characteristic protuberances in the basal area as well as strongly curved ends (compare Fig. 3 vs. Fig. 4). In addition, the median lobe of the aedeagus is considerably shorter in this new species. The elytra of the new species are shorter and more broadly rounded towards the apex, whereas the elytra in most of the C. lederi specimens I have seen appear more elongate and are much more ovally rounded towards the apex. In any case, males of the three very similar Caucasusacalles species mentioned here can be easily distinguished by their differently shaped internal structures of the endophallus and specimens can be easily assigned to a species. Molecularly, however, the females of C. subglobosus and C. circularis can be assigned to one of the species, because they are separated by 11 % in the mitochondrial CO 1 subgene (658 pb, Follmer region). This is a very high value (compared to other western Palaearctic species of the Cryptorhynchinae, which differ more clearly in morphological terms) and undoubtedly distinguishes these as different species (for details see Schütte et al. 2023).	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC2A225FF28FB776CFA22EE.taxon	distribution	Distribution. This species is so far known only from specimens from Armenia and central and southern Georgia.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC2A225FF28FB776CFA22EE.taxon	etymology	Derivatio nominis. The species epithet ‘ circularis ’ (Latin) refers to the almost circular elytra.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC0A22BFF28FEDC6D4E214B.taxon	description	urn: lsid: zoobank. org: act: 6 F 7 FE 0 E 9 - 7350 - 4 BC 0 - B 954 - 9625 FCB 3939 D (Fig. 5)	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC0A22BFF28FEDC6D4E214B.taxon	materials_examined	Type material Holotype: 1 ♀, ‘ Russia, Krasnodar: Krasnaya Polyana env., 43 ° 41 ’ 17 ” N, 40 ° 12 ’ 20 ” E, 720 m, 22.07.2014, leg. A. V. Kovalev’, coll. ZFMK, collector no. 2759 - PST, GenBank: MF 426966 (was previously a paratype of A. ossetiensis Stüben, 2018); for photo documentation see Fig. 5 // Paratypes: 1 ♂, ‘ USSR, Caucasus occ., Krasnaya Polyana, 1500 m, leg. Borovec’, ‘ 60 km E Adler pr. Sochi, 30.5.1985 ’, coll. Stüben / 1 ♀, ‘ Russia, Krasnodar: Chvizhepse vill. env., 43 ° 38 ’ 32 ” N 40 ° 04 ’ 45 ” E, 300 m, 11.07.2014, leg. A. V. Kovalev’, coll. Stüben; collector no. 2756 - PST, GenBank: MF 426965 (was previously a paratype of Acalles ossetiensis Stüben, 2018) / 1 ♀, ‘ NW Caucasus, Adygeya rep., 14.7.1998, Kamennomostskij, lgt. Kopecký & Svarc’, coll. Stüben-SDEI (col. 21432). Note. Holotype and one paratype are from the paratype series of Acalles ossentiensis Stüben, 2018 from Krasnodar (Russia). Preliminary remark: Acalles ossetiensis from southern Ossetia, which I described in 2018, also needs to be further split up after molecular analyses, and finally morphological studies made it very clear that it is another independent North Caucasian Acalles species from the Russian Krasnodar Territory and the Republic of Adygea. Both species were very often identified in the past as the very similar Acalles echinatus (Germar, 1824), especially from Eastern Europe, but this is certainly not supported by more recent molecular studies (see Fig. 1 and Appendix 1).	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC0A22BFF28FEDC6D4E214B.taxon	description	Length. 1.5 – 2.3 mm (without rostrum). Head & Rostrum. Eyes large, readily visible in dorsal view, considerably overlapping upper margin of antennal groove; rostrum of females microscopically finely punctate, straight, shiny, long and narrow, length / width ratio of 5 (measured between the insertions of the antennae). Last five funicular antennomeres as long as broad, trapezoidal, not clearly separated from very large, wide club; first two segments elongate. Pronotum. Only slightly longer than wide; widest behind middle, laterally evenly strongly rounded, but narrowing in a straight line directly behind front margin. Pronotal disc (lateral view) evenly rounded over entire length, slightly curved, always clearly separated from elytra. In dorsal view without keels or tubercles, but with strong, deep side by side punctures, without shallow interspaces; each puncture with a short and broad white or brown bristle; broad, short, light scales, which lie close to the underlying cuticle, particularly visible on flanks of pronotum. Elytra. Obovate, narrowly rounded apically; 1.2 × as long as wide; widest at end of first quarter behind base. In lateral view outline of elytra with a flat curve then regularly arcuate toward apex. Base of elytra slightly convex towards pronotum. Integument with white, beige and brown, circular and short oval non-overlapping scales, which do not obscure underlying cuticle; white scaly patch present on scutellum and just behind base on first interval, a few more irregularly limited light patches scattered on elytra; white crossband visible on first intervals of elytral declivity behind middle. Striae deeply punctate, intervals increasingly convex towards lateral margins; bearing a row of a few short, strongly inclined, white and brown bristles that are twice as long as wide; dark brown shiny cuticle not obscured by scales. Legs. Long, front femora reaching foremargin of eyes, hind femora extended beyond elytral apex, covered with long, narrow, white, only slightly raised bristles. Underside. Rostrum channel deep, very narrow between procoxae, about half as wide as channel at anterior margin of prosternum; channel ending in semicircular mesosternal receptaculum between the mesocoxae. All abdominal ventrites covered with light, long, widely spaced bristles, which lie close to cuticle. Female genitalia. See Fig 5 and differential diagnosis.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC0A22BFF28FEDC6D4E214B.taxon	diagnosis	Differential diagnosis Acalles krasnodarensis is closely related to Acalles ossetiensis Stüben 2018 from South Ossetia, but differs from it in a number of characteristics. Only females are used here for comparison: Rostrum of females at least 5 times longer than wide between the antenna insertions (in O. ossetiensis much shorter, at most 4.1 times longer than wide); dense, broad and deep pronotal punctation (which in A. ossetiensis is finer, less deep and characterised in places on the disc by broad shiny spaces); elytra more elongate, pointed oval, inverted ovate (versus shorter, more rounded towards the apex in A. ossetiensis); outer elytral intervals in particular strongly convex (versus only slightly convex); pronotum scales, especially well recognisable on the flanks, broader, short oval (versus clearly slimmer, long oval or spindle-shaped in A. ossetiensis); styli of the gonocoxite narrower and almost twice as long in the new species; and cornu of the spermatheca more curved (flatter in C. ossetiensis). Molecularly, these two species are relatively far apart in the mtCO 1 gene (659 bp). The p-distance here is between 7.9 % and 9.1 %, a comparatively very high value for the species of the subtribe Tylodina (see Schütte et al. 2023 for more details). To the next related taxon Acalles echinatus (Germar, 1824), a population near Moscow, with which both species from the Caucasus have obviosly always been equated due to the similar shape of the aedeagus, this value is 18.5 % (Fig. 1, Appendix 1). Immediately before this article went to press, R. Borovec (Czech Republic) provided me with a very small male (1.5 mm) from the type locality of Krasnaya Polyana area of the Krasnodar Territory (Russia), which L. Dieckmann had already (erroneously) identified as Acalles echinatus Germar in 1986. In contrast to this species, which is mainly distributed in Central and Eastern Europe, and Acalles ossetiensis, the median lobe of the aedeagus of A. krasnodarensis is clearly broader ventrally and more curved over its entire length when viewed laterally (see Figs. 5 and 6, top right).	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC0A22BFF28FEDC6D4E214B.taxon	distribution	Distribution. This species is so far known only from the eastern Black Sea coast in the vicinity of the town of Chvizhepse in the Russian province of Krasnodar, but it is likely to have a much wider distribution.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFC0A22BFF28FEDC6D4E214B.taxon	etymology	Derivatio nominis. This species is named after the Russian province of Krasnodar (Krasnodarskij kraj).	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
03F0A80AFFCEA233FF28FC396D8420E7.taxon	discussion	Note. A reliable and quick identification can be made via the aedeagus of the males (see also Fig. 1). Only then should the females be assigned on the basis of the elytra scales and bristles from the same locality. Sifting of wingless Cyptorhynchinae, the main method by which they are collected, always contains a large number of specimens, almost always including males.	en	Stüben, Peter E. (2025): New species and an image key to the wingless Cryptorhynchinae of the Caucasus (Coleoptera: Curculionidae: Tylodina) - a hypothesis on the spread of forest and open land species in the Western Palaearctic. Zootaxa 5647 (3): 235-259, DOI: 10.11646/zootaxa.5647.3.2, URL: https://doi.org/10.11646/zootaxa.5647.3.2
