identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03F0A80AFFC5A221FF28F9CD6B5E2373.text	03F0A80AFFC5A221FF28F9CD6B5E2373.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Caucasusacalles lederi (Meyer)	<div><p>Caucasusacalles lederi (Meyer) species complex</p><p>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).</p><p>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.</p><p>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.</p></div>	https://treatment.plazi.org/id/03F0A80AFFC5A221FF28F9CD6B5E2373	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Stüben, Peter E.	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.text	03F0A80AFFC4A226FF28FE206A8827F3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Caucasusacalles subglobosus Stuben 2025	<div><p>Caucasusacalles subglobosus Stüben sp. nov.</p><p>urn:lsid:zoobank.org:act: B3E8BDF7-C8FC-4CCD-AB2F-CE893574A381</p><p>(Fig. 2)</p><p>Type material</p><p>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 &amp; 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), mtCO1 sequence see Appendix 2.</p><p>Length. 1.9–2.5 mm (without rostrum)</p><p>Head &amp; 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 3rd to 7th segment, slightly shorter than wide; club short-oval.</p><p>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.</p><p>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.</p><p>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.</p><p>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 3rd and 4th ventrites, only slightly longer than 3rd and 4th combined.</p><p>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).</p></div>	https://treatment.plazi.org/id/03F0A80AFFC4A226FF28FE206A8827F3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Stüben, Peter E.	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.text	03F0A80AFFC2A225FF28FB776CFA22EE.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Caucasusacalles circularis Stuben 2025	<div><p>Caucasusacalles circularis Stüben sp. nov.</p><p>urn:lsid:zoobank.org:act: E2F97031-00B7-4CD8-B25B-FB2852505807</p><p>(Fig. 4)</p><p>Type material</p><p>Holotype: 1♂, ‘ Armenia: <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=45.066&amp;materialsCitation.latitude=41.032" title="Search Plazi for locations around (long 45.066/lat 41.032)">around Arakelots Monastery ruins, W Kirants</a>, Tavush, 21.5.2022, leaf litter sifting and Winkler extractor, 41,032000 45,066000, deciduous forest, leg. Astrin’, coll. ZFMK, ZFMK-TIS-8010636, mtCO1 sequence see Appendix 2; for photo documentation see Fig. 4 // Paratypes: 1♂, ‘ Georgia: 9 km E of Tianeti, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=45.04298&amp;materialsCitation.latitude=42.08619" title="Search Plazi for locations around (long 45.04298/lat 42.08619)">on Aniskhevi River banks</a>, forest next to river and affluent creek, Mzkheta-Mtianeti, 42,08619 45,04298, 1051 m, leg. Astrin’, coll. Stüben, mtCO1 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, mtCO1 sequence see Appendix 2 / 1♂, 3♀, ‘ Georgia, Kharagauli, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=43.28361&amp;materialsCitation.latitude=41.940556" title="Search Plazi for locations around (long 43.28361/lat 41.940556)">Merelisi</a> env., 482 m, 41°56'26"N 43°17'01"E, 29.8.2019, sifting in mixed forest, lgt. Baňař, Hlaváč, Barjadze &amp; Maghradze’, coll Krátký / 1♂, ‘ Georgia: „ Kaukasus, Suram-Pass ”, „ Acalles lederi Meyer &lt;handwriting&gt;, Penecke det.’, coll. Senckenberg Naturhistorische Sammlungen Dresden / Georgia: 1♂, ’ Suram-Pass, 2010, Kulzer ”, “ Acalles choudoiri, Zumpt det.’, coll. Naturhistorisches Museum in Wien.</p><p>Description &amp; differential diagnosis</p><p>Length. 2.1–2.5 mm (without rostrum)</p><p>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 .</p><p>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).</p><p>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.</p><p>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 CO1 subgene (658pb, 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).</p><p>Distribution. This species is so far known only from specimens from Armenia and central and southern Georgia.</p><p>Derivatio nominis. The species epithet ‘ circularis ’ (Latin) refers to the almost circular elytra.</p></div>	https://treatment.plazi.org/id/03F0A80AFFC2A225FF28FB776CFA22EE	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Stüben, Peter E.	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.text	03F0A80AFFC0A22BFF28FEDC6D4E214B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Acalles krasnodarensis Stuben 2025	<div><p>Acalles krasnodarensis Stüben sp. nov.</p><p>urn:lsid:zoobank.org:act: 6F7FE0E9-7350-4BC0-B954-9625FCB3939D</p><p>(Fig. 5)</p><p>Type material</p><p>Holotype: 1♀, ‘ Russia, Krasnodar: <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=40.205555&amp;materialsCitation.latitude=43.688057" title="Search Plazi for locations around (long 40.205555/lat 43.688057)">Krasnaya Polyana</a> 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: MF426966 (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: <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=40.079166&amp;materialsCitation.latitude=43.642223" title="Search Plazi for locations around (long 40.079166/lat 43.642223)">Chvizhepse</a> 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: MF426965 (was previously a paratype of Acalles ossetiensis Stüben, 2018) / 1 ♀, ‘NW Caucasus, Adygeya rep., 14.7.1998, Kamennomostskij, lgt. Kopecký &amp; Svarc’, coll. Stüben-SDEI (col. 21432) .</p><p>Note. Holotype and one paratype are from the paratype series of Acalles ossentiensis Stüben, 2018 from Krasnodar (Russia).</p><p>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).</p><p>Length. 1.5–2.3 mm (without rostrum).</p><p>Head &amp; 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.</p><p>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.</p><p>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.</p><p>Legs. Long, front femora reaching foremargin of eyes, hind femora extended beyond elytral apex, covered with long, narrow, white, only slightly raised bristles.</p><p>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.</p><p>Female genitalia. See Fig 5 and differential diagnosis.</p><p>Differential diagnosis</p><p>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).</p><p>Molecularly, these two species are relatively far apart in the mtCO1 gene (659bp). 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).</p><p>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).</p><p>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.</p><p>Derivatio nominis. This species is named after the Russian province of Krasnodar (Krasnodarskij kraj).</p></div>	https://treatment.plazi.org/id/03F0A80AFFC0A22BFF28FEDC6D4E214B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Stüben, Peter E.	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.text	03F0A80AFFCEA233FF28FC396D8420E7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Cryptorhynchinae	<div><p>Image key to the wingless Cryptorhynchinae of the Caucasus</p><p>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.</p><p>1 Base of each elytron strongly S -shaped, protruding towards the pronotum with corners that extend far forward. Length: 3.0– 4.9 mm .................................................................... Acallocrates colonnellii Bahr, 2003</p><p>1* Base of the elytron straight or only slightly convex over the entire length protruding towards the pronotum.............. 2</p><p>2 Male with a distinct (strongly sclerotised) sclerite (= inner sac structure) in the endophallus.......................... 3</p><p>2* Males without a recognisable, strongly sclerotised (more or less complex) sclerite in the endophallus (instead sometimes with a blurred, ‘cloudy’ appearing endophallus consisting of many tiny hooks).......... Onyxacalles &amp; Echinodera ........ 11</p><p>3 Males with a strongly sclerotised, elongate, parallel-sided or X-shaped sclerite in the endophallus, basally (sometimes also apically) with a closed or semicircularly open structure (see couplet 2)......................... Kyklioacalles ………. 4</p><p>3* Males with sclerite in the endophallus differently shaped (without a circular or only semicircularly opened structure)...... 6</p><p>--- Kyklioacalles ---</p><p>4 Elytra with patches of pale scales (‘eyespots’) absent, instead with light-coloured, irregularly delimited, cloudy markings on the elytra and a prominent X-shaped inner sac structure of the aedeagus; tiny species: &lt;2.3 mm ............................................................................................... Kyklioacalles reitteri (Meyer, 1896)</p><p>4* Elytra with two patches of pale scales (‘eyespots’) at the centre of the elytra and a strongly sclerotised, elongate, parallel-sided sclerite in the endophallus (basal or apical with a circular structure)............................................. 5</p><p>5 Larger species,&gt; 3.5 mm; with a long, bottle-necked tip of the aedeagus (couplet 4*, left). Length: 3.5–5.2 mm ........................................................................... Kyklioacalles ocellus Stüben &amp; Astrin, 2025</p><p>5* Smaller species, &lt;3.0 mm; with a rounded tip of the aedeagus (couplet 4*, right). Length: 2.0–3.0 mm.................................................................................... Kyklioacalles chaudoirii (Hochhuth, 1847)</p><p>6 Pronotal disc and elytra with conspicuously circular, large, partly overlapping scales, which largely obscure the underlying cuticle. Elytral intervals with short, widely protruding, vertically erect bristles................ Caucasusacalles ………. 7</p><p>6* Pronotal disc and elytra predominantly with narrower, drop-shaped, slender or short-oval adjacent scales, which usually do not obscure the dark brown, shiny underlying cuticle. Elytral intervals either with dense comb-like brush or tufts or only a few strongly inclined, club-shaped bristles........................................................ Acalles ………. 9</p><p>--- Caucasusacalles ---</p><p>7 Sclerite of the endophallus with a parallel-sided basal structure consisting of two narrow, closely spaced bars, each apical with an inwardly inclined long, spine-like tip. Length: 1.9–2.5 mm ............. Caucasusacalles subglobosus Stüben sp. nov.</p><p>7* Sclerite of the endophallus with a parallel-sided basic structure consisting of widely spaced bars, which are connected apically by a pointed, weakly sclerotised bridge.................................................................... 8</p><p>8 Sclerite of the endophallus with only two simple, subparallel bars (couplet 7*, left); median lobe of the aedeagus much longer, but at least 2× longer than wide. Length: 1.8–2.2 mm ........................... Caucasusacalles lederi (Meyer, 1896)</p><p>8* Sclerite of the endophallus with further angularly separated, characteristic protuberances in the basal area as well as strongly curved ends (couplet 7*, right); the median lobe of the aedeagus considerably shorter, nearly as wide as long. Length: 2.1–2.5 mm .............................................................. Caucasusacalles circularis Stüben sp. nov.</p><p>--- Acalles ---</p><p>9 Elytral intervals with numerous, towering condensations or tufts of bristles on the elytral intervals; inner sac of the aedeagus in the shape of a Greek ‚Omega’. Length: 3.0–4.0 mm................................ Acalles (s.l.) milleri Reitter, 1883</p><p>9* Elytral intervals without such tufts of bristles or conspicuous narrowly arranged accumulations of bristles on the elytral intervals; inner sac of the aedeagus consisting of two simple parallel stripes.............................................. 10</p><p>10 Rostrum of females much shorter, at most 4.1 times as long as wide between antennal insertions; elytra shorter, more rounded towards the apex (couplet 9*, left); punctation of the pronotum finer, less deep and more separated on the disc; scales narrower. Length: 1.8–2.3 mm ......................................................... Acalles ossetiensis Stüben, 2018</p><p>10* Rostrum of females at least 5 times as long as wide between antennal insertions; elytra more elongate, pointed oval or inverted ovate (couplet 9*, right); pronotum with dense, broad, deep punctures, scales much wider, drop-shaped. Length: 1.5–2.3 mm .................................................................... Acalles krasnodarensis Stüben sp. nov.</p><p>11 Eyes large, clearly protruding (up to half the diameter of the eye) over the upper edge of scrobe; median lobe of aedeagus (ventral) short, only slightly longer than wide............................................ Onyxacalles ………. 12</p><p>11* Eyes small, not exceeding the upper edge of the scrobe; median lobe of the aedeagus long (‘rocket-shaped’), many times longer than wide......................................................................... Echinodera ………. 14</p><p>--- Onyxacalles ---</p><p>12 Elytra with strong tubercles on the outermost 3 intervals; elytral bristles inconspicuous, short, very fine, strongly inclined and curved; median lobe of aedeagus tapering to a pointed triangle when viewed ventrally. Length: 2.8–4.0 mm............................................................................. Onyxacalles gibbosus Stüben &amp; Astrin, 2025</p><p>12* Elytra without strong tubercles (outer intervals at most jagged); elytral bristles broader, club-shaped, longer and forming a ‘crown-shape’; median lobe of aedeagus more rounded or with a bottleneck-like tip............................... 13</p><p>13 Elytral intervals with punctures deep and broad, especially on the outermost striae, extended to the intervals, giving the elytra a jagged outline; median lobe of the aedeagus simply pointed (couplet 12*, left). Length: 2.2–3.8 mm ....................................................................................... Onyxacalles caucasicus (Reitter, 1891)</p><p>13* Elytral intervals elevated, but without deeply and heavily punctate surface; median lobe of the aedeagus with a bottleneck-like tip (couplet 12*, right). Length: 3.3.– 3.5 mm .......................... Onyxacalles georgius Stüben &amp; Krátký, 1918</p><p>--- Echinodera ---</p><p>14 Eyes smaller and oval; clearly placed on the sides of the head; aedeagus with an ‘S-shaped’ long, tapering, not sclerotized (“translucent”) apex; integument of the smaller species (&lt;3.4 mm) (light) brown... Subgenus: Echinodera s.str. ………. 15</p><p>14* Eyes larger and mostly round; visible as a narrow strip when viewed from above; aedeagus either simply rounded or protruding bottle-shaped at the apex (but never with an S -shaped, transparent tip); clearly larger, dark brown to black species (&gt; 4 mm)............................................................................. Subgenus: Ruteria ………. 18</p><p>--- Echinodera s. str. subg.---</p><p>15 Spine at the end of the hind tibia of males curved inwards at almost a right angle. Length: 2.2–3.0 mm........................................................................................ Echinodera davidiani Savitsky, 1999</p><p>15* Spine at the end of the hind tibia of males straight or weakly curved............................................ 16</p><p>16 Elytra longer, obovate, apex rounded narrowly oval; elytral bristles very short (at most 1.5× longer than wide)). Length: 2.6–3.4 mm ............................................................. Echinodera orientalis (A. &amp; F. Solari, 1907)</p><p>16* Elytra wider, short ovally rounded with apex more broadly rounded; elytral bristles longer (2×–4× longer than wide)..... 17</p><p>17 Elytral bristles very long, 3.5×–4× longer than wide (couplet 16*, left); striae and the punctures between the outermost elytra intervals narrow; median lobe of aedeagus strongly curved towards the tip when viewed ventrally. Length: 2.2 mm ..................................................................... Echinodera borjomiensis Stüben &amp; Astrin, 2025</p><p>17* Elytral bristles short, only 2×–2.5 x longer than wide (couplet 16*, right); elytral striae strongly punctate between the narrower outermost elytra intervals; median lobe of aedeagus nearly flat when viewed laterally. Length: 1.9–3.1 mm .................................................................................... Echinodera horridula (Reitter, 1888)</p><p>--- Ruteria subg. ---</p><p>18 Sides of the pronotum with light brown and white scales; aedeagus with a narrow, elongate apex. Length: (2.7) 4.0– 5.6 mm. Note. According to Savitsky (1997), the central, southern and southeastern European species is also said to occur in Georgia (the species was described from Austria, Styria). Savitsky mentions only 2 females and 1 male (it remains unclear whether the aedeagus shown is from this male.). In order to take this unusual disjunct distribution into account, the species is also mentioned here for the Caucasus, despite considerable doubt (to facilitate subsequent identification)........................................................................................ Echinodera (Ruteria) hypocrita (Boheman, 1837)</p><p>18* Sides of the pronotum with dark brown or black scales; aedeagus not protruding like a bottle neck at tip. Note. So far, molecular data from this group are only available for E. teplovi . Therefore, I follow here the exclusively morphological differential diagnosis of Savitsky, 1997, as far as it seemed comprehensible to me after examination of the type material (see illustrations of the habitus of holotypes). I supplement with illustrations of the aedeagus; however, further studies are urgently needed here on the generally poorly researched species of the subgenus Ruteria from the Caucasus ............................................................................... Echinodera anatolica group ………. 19</p><p>19 Elytra with 7th stria at end of basal 1/4 abruptly ending; mesosternal part of receptaculum with distinct tubercle; aedeagus slender, tapered. Length: 4.0– 5.5 mm .............................. Echinodera (Ruteria) kakhetica (Savitsky, 1997) (probably a synonym of E. hyrcania, see comment Stüben, 2018: 367)</p><p>19* Elytra with 7th stria not abruptly ending at the end of basal ¼; mesosternal part of receptaculum usually without tubercle.... .................................................................................................. 20</p><p>20 Middle part of metathorax and ventrite 1 covered with dissected scales; uncus of fore tibia in male beveled triangular, split (see couplet 20); apical part of ventrite 1 strongly sloping, at nearly right angle to plane of ventrite 2; aedeagus broader, obtuseangled (dorsal view). Length: 4.1–4.9 mm ......................... Echinodera (Ruteria) meskhetica (Savitsky, 1997)</p><p>20* Middle part of metathorax and ventrite 1 covered with elongate undissected scales; uncus of fore tibia in male of different shape; apical part of ventrite 1 slightly or moderately sloping to plane of ventrite 2................................ 21</p><p>21 Body of male 1.78–1.85, of female 1.77–1.91, times as long as wide; anterior part of receptaculum covered with hairs and admixture of scales divided into several filiform lobes; uncus of fore tibia in male wide; aedeagus wider towards the tip. Length: 4.0– 5.5 mm .................................................. Echinodera (Ruteria) hyrcanica (Savitsky, 1997)</p><p>21* Body of male 1.58–1.77, of female 1.59–1.80, times as long as wide; anterior part of receptaculum covered with hairs and sparse narrow scales divided into 2 filiform lobes; uncus of fore tibia in male weakly dilated; aedeagus with clearly bent tip (lateral view). Length: 3.9–5.0 mm.................................. Echinodera (Ruteria) teplovi (Savitsky, 1997)</p></div>	https://treatment.plazi.org/id/03F0A80AFFCEA233FF28FC396D8420E7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		MagnoliaPress via Plazi	Stüben, Peter E.	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
