{"id":111,"date":"2021-05-21T20:21:24","date_gmt":"2021-05-21T20:21:24","guid":{"rendered":"https:\/\/hnords.de\/wordpress\/?page_id=111"},"modified":"2022-03-23T01:50:36","modified_gmt":"2022-03-23T00:50:36","slug":"charpentieria-itala","status":"publish","type":"page","link":"https:\/\/hnords.de\/wordpress\/charpentieria-itala\/","title":{"rendered":"Charpentieria itala"},"content":{"rendered":"<p style=\"text-align: left;\"><strong><em>Charpentieria itala<\/em>: subspecies, species, superspecies<\/strong><\/p>\n<p style=\"text-align: left;\">Hartmut Nordsieck (V.2021)<\/p>\n<h4 style=\"text-align: left;\"><strong>I. Introduction and history<\/strong><\/h4>\n<p style=\"text-align: left;\">In the revision of the <em>Charpentieria<\/em> species from the southern limestone Alps (Nordsieck 1963: 174-190, as <em>Delima<\/em>) I tried to describe the intraspecific diversity of <em>C. itala<\/em> (G. von Martens 1824) by a subdivision of the species of that region into several subspecies. These are (from W to E) <em>C. i. albopustulata<\/em> (Cristofori &amp; Jan 1832), <em>C. i. latestriata<\/em> (K\u00fcster 1850), <em>C. i. rubiginea<\/em> (Rossm\u00e4ssler 1836), and <em>C. i. serravalensis<\/em> (H. Nordsieck 1963), besides, as transitional forms within <em>C. i. albopustulata<\/em>, but later on treated as subspecies, <em>C. <\/em><em>i. braunii<\/em> (Rossm\u00e4ssler 1836) and <em>C. i. baldensis<\/em> (Charpentier 1852) [already described by Strobel (1851: 25)]. A complex of forms from the same region which in shell morphology and ecological preferences are similar to the related species <em>C. stenzii<\/em> (Rossm\u00e4ssler 1836), but in genital morphology largely correspondent with <em>C. itala<\/em> and in part connected with it by transitional forms, was included in the latter as so-called stenzioid subspecies (Nordsieck 1963: 171-173, 181-187): <em>C. i. clavata<\/em> (Rossm\u00e4ssler 1836), <em>C. i. variscoi<\/em> (Pini 1883), <em>C. i. balsamoi<\/em> (Strobel 1850), <em>C. i. lorinae<\/em> (Gredler 1869). Later on, because of its semispecies character, that complex has been separated as species <em>C. clavata<\/em> (Nordsieck 1979: 259). In the 1963 paper the subspecies of <em>C. itala<\/em> from the remaining distributional range (Monti Berici and Colli Euganei, Provence, Maritime and Ligurian Alps, northern and central Apennines) were not considered. The related species <em>C. dyodon<\/em> (Studer 1820) from Piedmont was not treated either, because at this time its close relationship to <em>C. itala<\/em> was not known. Zilch (1972), in a type catalogue of the Delimini, has also listed the subspecies of <em>C. itala<\/em> outside the southern Alps: <em>C. i. itala<\/em> from Monti Berici and Colli Euganei, and <em>C. i. nigra<\/em> (Issel 1866) and <em>C. i. punctata<\/em> (Michaud 1831) from the remaining range.<br \/>\nSeveral Italian authors called in question the proposed subdivision of <em>C. itala<\/em> into subspecies, with the following arguments:<br \/>\nGiusti &amp; Mazzini (1971: 294): &#8222;Sulla scorta delle sue descrizioni [Nordsieck 1963] abbiamo provato a determinare alcuni materiali dell&#8216; Italia settentrionale, tuttavia, senza riuscirci mai con sicurezza. Tanto variabili sono i caratteri conchiliologici &#8230;&#8220;<br \/>\nBoato et al. (1985: 312): <em>C. itala<\/em> cfr. <em>punctata<\/em>: &#8222;&#8230;, sia questa, che le numerose altre che normalmente si includono nella <em>C. itala<\/em> sono spesso variabili e poco definite&#8220;.<br \/>\nManganelli et al. (1995: 47): &#8222;Molte delle entit\u00e0 sottospecifiche convalidato da Nordsieck (1963 &#8230;) e da Zilch (1972 &#8230;) qui riportate appaiano di validit\u00e0 dubbia&#8220;.<br \/>\nThese statements, however, are only opinions, not substantiated by further investigations.<br \/>\nTherefore, for this chapter the shell characters, on which the subspecies division of <em>C. itala<\/em> was based, have been checked once more, and the revision of 1963 was supplemented by the inclusion of the forms from beyond the southern Alps and of <em>C. dyodon<\/em> from Piedmont. The stenzioid subspecies of <em>C. itala<\/em> = <em>C. clavata<\/em> and the transitional subspecies <em>C. i. allatollae<\/em> (K\u00e4ufel 1928) were not considered in this revision, because new results could not be expected. The examined shell material of <em>C. itala<\/em> has been augmented by a third (from 350 samples for the 1963 paper to 460 for this chapter), and 25 samples of <em>C. dyodon<\/em> have been included. The material is deposited in the collection of the Forschungsinstitut Senckenberg (SMF) and in my own collection (deposited mainly in the Staatliches Museum f\u00fcr Naturkunde Stuttgart).<br \/>\nIt resulted that the most important characters, those of lunellar and clausilium plate, are not at all too variable and are therefore suited for the definition of subspecies. Obviously, these characters were not considered or not understood by the named authors (because the paper was written in German). In the following they are once more described and figured.<\/p>\n<h4 style=\"text-align: left;\">II. <strong>Results<\/strong><\/h4>\n<p style=\"text-align: left;\">1. Character analysis:<\/p>\n<p style=\"text-align: left;\">The lunellar of <em>C. itala<\/em> (Fig. 1) consists of (posterior) upper palatal plica, lunella and posterior lower palatal plica. An anterior upper palatal plica and an anterior lower palatal plica (= basalis) are absent. The posterior lower palatal plica is named subclaustralis; it is connected with the lunella by an obtuse or nearly right angle. A further posterior lower palatal plica, the lowest one, is named sulcalis. It is in part weakly developed; as a rule, it is not connected with the subclaustralis. The outer edge of the clausilium plate rests against the lunella and the subclaustralis in the closed position.<br \/>\nThere are two modifications of closing apparatus (Nordsieck 1963: 175, Figs. 2-9). In one type (A, Figs. 5-7) the upper palatal plica is connected with the lunella by an angle or a curve; the lunella is less oblique with respect to the shell axis. In the other type (B, Figs. 2-4, 8-9) the upper palatal plica is connected with the lunella by a curve; the lunella is more oblique with respect to the shell axis. There are two possibilities, how the clausilium plate rests against the subclaustralis; this can be observed by an oblique view into the aperture. In type A (Figs. 6-7) the outer edge of the clausilium plate rests against the subclaustralis up to its distal end and does not overlap this plica, in type B (Figs. 8-9) the outer edge of the clausilium plate overlaps that plica (at least in most specimens) at its distal end. <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em>, <em>C.<\/em> <em>i.<\/em> <em>rubiginea<\/em> and outside the southern Alps <em>C. i. punctata<\/em> (see part 3) exhibit the type A, <em>C. i. albopustulata<\/em>, <em>C. i. latestriata<\/em>,<em> C. i. braunii<\/em>, <em>C. i. baldensis<\/em> and outside the southern Alps<em> C. i. itala<\/em> (see part 2) the type B.<br \/>\nAnother shell character used for defining subspecies of <em>C. itala<\/em> is the length of the principal plica. The remaining shell characters, size, shape, sculpture, development of peristome and palatal callus are of minor importance.<\/p>\n<h4 style=\"text-align: left;\"><strong>Fig. 1. L<\/strong><strong>unellar of <em>Charpentieria itala<\/em> (scheme).<\/strong><\/h4>\n<h5 style=\"text-align: left;\"><a href=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/italalunellarschemahomp.jpg\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone wp-image-271 size-medium\" src=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/italalunellarschemahomp-294x300.jpg\" alt=\"\" width=\"294\" height=\"300\" srcset=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/italalunellarschemahomp-294x300.jpg 294w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/italalunellarschemahomp-265x270.jpg 265w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/italalunellarschemahomp.jpg 338w\" sizes=\"auto, (max-width: 294px) 100vw, 294px\" \/><\/a><\/h5>\n<h5 style=\"text-align: left;\">Abbreviations:<br \/>\nlu = lunella; plp = posterior lower palatal plica (subclaustralis); pr = principal plica; pup = (posterior) upper<br \/>\npalatal plica; sc = subcolumellar lamella; sul = lowest palatal plica (sulcalis).<\/h5>\n<p style=\"text-align: left;\"><strong>Figs. 2.-5. <em>C. itala<\/em>.<br \/>\n<\/strong>Actual shell height (mm).<\/p>\n<p style=\"text-align: left;\"><a href=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-2-5-Kopie.jpg\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone wp-image-269 size-large\" src=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-2-5-Kopie-1024x517.jpg\" alt=\"\" width=\"640\" height=\"323\" srcset=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-2-5-Kopie-1024x517.jpg 1024w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-2-5-Kopie-300x151.jpg 300w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-2-5-Kopie-768x388.jpg 768w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-2-5-Kopie-535x270.jpg 535w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-2-5-Kopie.jpg 1417w\" sizes=\"auto, (max-width: 640px) 100vw, 640px\" \/><\/a><\/p>\n<h5 style=\"text-align: left;\">2. <em>C. i. itala<\/em>, Mira (Villa Ducale) (ex SMF 193974), H = 18.9.<br \/>\n3. <em>C. i. itala<\/em>, Vicenza (syntype of. C. i. var. vicentina, ex SMF 196209), H = 21.3.<br \/>\n4. <em>C. i. albopustulata<\/em>, Varone waterfall near Riva (ex SMF 174367), H = 18.3.<br \/>\n5.<em> C. i. serravalensis<\/em>, Vittorio Veneto (ex SMF 265953), H = 18.4.<\/h5>\n<p style=\"text-align: left;\"><strong>Figs. 6-9. <em>C. itala<\/em>.<br \/>\n<\/strong>Actual shell height (mm) = H.<br \/>\na. Body whorl dorsal, view on lunellar.<br \/>\nb. Body whorl frontal, oblique view into aperture. Different position of clausilium plate with respect to the<br \/>\nsubclaustralis marked by an arrow.<\/p>\n<h5 style=\"text-align: left;\"><a href=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-6-9-Kopie.jpg\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone wp-image-270 size-large\" src=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-6-9-Kopie-1024x880.jpg\" alt=\"\" width=\"640\" height=\"550\" srcset=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-6-9-Kopie-1024x880.jpg 1024w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-6-9-Kopie-300x258.jpg 300w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-6-9-Kopie-768x660.jpg 768w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-6-9-Kopie-314x270.jpg 314w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/itala-Lunellar-6-9-Kopie.jpg 1417w\" sizes=\"auto, (max-width: 640px) 100vw, 640px\" \/><\/a><\/h5>\n<h5 style=\"text-align: left;\">6. <em>C. i. rubiginea<\/em>, Bozen (ex coll. Bosch), H = 17.1.<br \/>\n7. <em>C. i. punctata<\/em>, Fivizzano (ex SMF 186744), H = 16.7 (apical part broken).<br \/>\n8. <em>C. i. albopustulata<\/em>, Lugano (ex coll. Bosch), H = 18.4.<br \/>\n9. <em>C. i. braunii<\/em>, Heidelberg (castle) (ex coll. Bosch), H = 15.9.<\/h5>\n<p style=\"text-align: left;\">2.<em> C. i. itala:<\/em><\/p>\n<p style=\"text-align: left;\">G. von Martens (1824: 442, pl. 3, fig. 1) described <em>Clausilia itala<\/em> from Miravecchia (&#8222;&#8230; Garten des aelterlichen Hauses &#8230;&#8220;, according to E. von Martens 1857: 129 between Mira and Dolo). Zilch (1972: 247) regarded the forms from Monti Berici, Mira and Colli Euganei as representing the nominotypical subspecies and listed the specimens collected by me in Mira (SMF 193974, Nordsieck 2007: pl. 13, fig. 6; Fig. 2) as topotypes. The type form from Mira is relatively small and similar to large forms of <em>C. i. albopustulata<\/em> (eastern form, see Nordsieck 1963: 177). The samples from Monti Berici (e. g., Arcugnano) and Vicenza (<em>C. i.<\/em> var. <em>vicentina<\/em> A. Schmidt 1868, syntypes SMF 196209, Fig. 3) have a larger and more ventricose shell. The samples from Colli Euganei (Abano Terme, Teolo) (<em>C. i. <\/em>var<em>. vicentina<\/em> f. <em>tridentina<\/em> O. Boettger 1879, types SMF 167941) have in part a larger shell, but of the same shape as the type form. In contrast to the other samples, in those from Teolo the clausilium plate does not overlap the subclaustralis (type A).<br \/>\nAmong the samples classified by Zilch (: 247) with <em>C. i. itala<\/em>, however, some from Padova are wrongly affiliated to this subspecies. The sample from Padova (botanical garden) (SMF 156968, including the specimen figured by Zilch 1960: fig. 1532) belongs to <em>C. i. braunii<\/em>.<br \/>\n<em>C. i. itala<\/em> does not much differ from <em>C. i. albopustulata<\/em> (eastern form), but is separated as subspecies because of its shell size (on average larger) and its isolated range (see map). The <em>vicentina<\/em> form differs from <em>C. i. <\/em><em>albopustulata<\/em> also by its shell shape (more ventricose); thus the type form of <em>C. i. itala<\/em> is transitional between the <em>vicentina<\/em> form and <em>C. i. albopustulata<\/em>. The form from Teolo, however, resembles <em>C. i. serravalensis.<\/em><\/p>\n<p style=\"text-align: left;\">3. <em>C. i. punctata<\/em>:<\/p>\n<p style=\"text-align: left;\">The various forms of <em>C. itala<\/em> from Provence, Maritime and Ligurian Alps and northern and central Apennines (see map) are characterized by a stronger sculpture and a clausilium plate which does not overlap the subclaustralis (type A, Fig. 7). The form from the Provence (<em>Clausilia\u00a0 punctata<\/em> Michaud 1831) and those from the Apennines (<em>C. alboguttulata<\/em> var. <em>obesa<\/em> Issel 1866 [non L. Pfeiffer] from Lucchese,<em> C. a.<\/em> var. <em>nigra<\/em> Issel 1866 from Volterra, <em>C. a.<\/em> var. <em>elegans<\/em> Gentiluomo 1868 [non Cantraine] from Vallombrosa) do not much differ and are therefore not separated as subspecies (in contrast to Giusti &amp; Mazzini 1971: 295, Zilch 1972: 251).<br \/>\nThe whole subspecies has to bear the oldest name <em>C. i. punctata<\/em>. This was already proposed by Boato et al. (1985: 312), but as cfr. <em>punctata,<\/em> and Manganelli et al. (1995: 24), but with the statement (: 47): &#8222;L&#8216; attribuzione del materiale della Liguria e dell&#8216; Appennino a <em>C. i. punctata<\/em> permane incerta.&#8220;<br \/>\nThe alleged syntypes of <em>C. punctata<\/em> from the Rossm\u00e4ssler collection (SMF 196374) are much smaller and more ventricose than the type specimens (Michaud 1831: 55, pl. 15, fig. 23) and belong to <em>C. i. braunii<\/em>. This has not been noticed by the previous authors (Rossm\u00e4ssler, O. Boettger) and by Zilch (1972: 251), who listed them as syntypes.<br \/>\nIt is unclear, if <em>Clausilia genei<\/em> Lessona 1880 from Valle Pesio, Piedmont, is a form of <em>C. i. punctata<\/em>, because material is not available.<br \/>\nIt should be checked, if <em>C. i. punctata<\/em> is distributed in Italy beyond the southern border (as indicated in the map), e. g., as far as the Colli Albani, where it is said to occur by E. von Martens (1857: 136).<\/p>\n<p style=\"text-align: left;\">4. Two misidentified forms of <em>C<\/em>. <em>itala<\/em>:<\/p>\n<p>The <em>C<\/em>. <em>itala<\/em> form from Meran and surroundings, South Tyrol, differs much from <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em>, as which it was determined in my 1963 paper (: 177). Unlike <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em>\u00a0it has a long principal plica and a clausilium plate not overlapping the subclaustralis (type A). It thus corresponds with <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em> (the only subspecies in which a long principal plica is present, Fig. 7) and is therefore classified with that subspecies. It may have been introduced by man (with winegrowing?) in the region of Meran, like <em>C.<\/em> <em>i.<\/em> <em>braunii<\/em> from Trento into the region of Brixen (see part 1).<br \/>\nThe <em>C<\/em>. <em>itala<\/em> form from Verona and surroundings, Venetia, differs from <em>C<\/em>. <em>i<\/em>. <em>albopustulata<\/em>, to which it was affiliated in my 1963 paper (: 188), by a more ventricose shell with in part detached aperture. It thus corresponds to <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em>. The forms from the eastern and southern shores of Lake Garda exhibit often a detached aperture, but are more slender than the form from Verona. They are transitional to <em>C<\/em>.<em> i<\/em>. <em>albopustulata<\/em>.<br \/>\nThe form from Udine, Friuli, which has been mentioned as <em>Clausilia alboguttulata<\/em> var. b <em>minor <\/em>by Pirona (1865\u00a0: 692), is strikingly similar to that from Verona, in qualitative shell characters and in the measurements (see Table 1). Nevertheless it has been described by de Mattia et al. (2021) as new subspecies <em>C<\/em>. <em>i<\/em>. <em>zalloti<\/em>. Because of this similarity it is deemed to have been introduced, like other forms of <em>C<\/em>. <em>i<\/em>. <em>braunii.<\/em> Therefore, <em>C<\/em>. <em>i<\/em>. <em>zalloti<\/em> is regarded as a synonym of <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em>.<\/p>\n<p>Table 1. Shell measurements of <em>C<\/em>. <em>itala<\/em> (normal subspecies).<br \/>\nValues of extreme or transitional forms in brackets. As a rule 10 specimens per sample have been measured.<\/p>\n<table>\n<tbody>\n<tr>\n<td width=\"256\">Subspecies (number of samples)<\/td>\n<td width=\"256\">shell height = H (mm)<\/td>\n<td width=\"256\">relative shell width = D\/H<\/td>\n<\/tr>\n<tr>\n<td width=\"256\"><em>C<\/em>. <em>i<\/em>. <em>albopustulata<\/em> (11)<\/td>\n<td width=\"256\">16.8-21.2<\/td>\n<td width=\"256\">0.228-0.243 (- 0.253)<\/td>\n<\/tr>\n<tr>\n<td width=\"256\"><em>C<\/em>. <em>i<\/em>. <em>latestriata<\/em> (3)<\/td>\n<td width=\"256\">16.5-16.9<\/td>\n<td width=\"256\">0.237-0.248<\/td>\n<\/tr>\n<tr>\n<td width=\"256\"><em>C<\/em>. <em>i<\/em>. <em>braunii<\/em> (9)<br \/>\n[Verona, Udine]<\/td>\n<td width=\"256\">16.3-19.4<br \/>\n[16.6, 16.3]<\/td>\n<td width=\"256\">(0.239-) 0.244-0.263<br \/>\n[0.251, 0.255]<\/td>\n<\/tr>\n<tr>\n<td width=\"256\"><em>C<\/em>. <em>i.<\/em> <em>itala<\/em> (4), <em>vicentina<\/em> form (3)<\/td>\n<td width=\"256\">20.0-22.3, 20.7-21.3<\/td>\n<td width=\"256\">0.231-0.241, 0.254-0.260<\/td>\n<\/tr>\n<tr>\n<td width=\"256\"><em>C<\/em>. <em>i<\/em>. <em>punctata<\/em> (5)<\/td>\n<td width=\"256\">16.4-19.5<\/td>\n<td width=\"256\">0.239-0.243 (-0.260)<\/td>\n<\/tr>\n<tr>\n<td width=\"256\"><em>C<\/em>. <em>i<\/em>. <em>rubiginea<\/em> (6)<\/td>\n<td width=\"256\">16.1-17.3<\/td>\n<td width=\"256\">0.227-0.248<\/td>\n<\/tr>\n<tr>\n<td width=\"256\"><em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em> (6)<\/td>\n<td width=\"256\">18.9-21.1<\/td>\n<td width=\"256\">0.224-0.246<\/td>\n<\/tr>\n<\/tbody>\n<\/table>\n<p>In comparison with the other subspecies <em>C<\/em>. <em>i.<\/em> <em>braunii<\/em> and the <em>vicentina<\/em> form of <em>C<\/em>. <em>i<\/em>. <em>itala<\/em> are characterized by more ventricose shells. The values of the samples from Verona and Udine are nearly identical.<\/p>\n<p style=\"text-align: left;\">5. <em>C. dyodon<\/em>:<\/p>\n<p style=\"text-align: left;\"><em>C. dyodon<\/em>, which was formerly classified with another group (<em>Dilataria<\/em> Vest) in another subfamily (Zilch 1960: 401), has been recognized by an examination of the genitalia (Nordsieck 1972: 38) as closely related to <em>C. itala<\/em>. The shell differs from that of <em>C. itala<\/em> mainly by the reduction of the sutural papillae and a lunellar tending to reduction. Until now, forms transitional to <em>C. itala<\/em> have not been found. Therefore, in spite of its genital similarity to <em>C. itala<\/em>, <em>C. dyodon<\/em> is regarded as an independent species. Its distributional range in Piedmont (see map) fills partly the gap between that of the <em>C. itala<\/em> subspecies from the southern Alps and that of <em>C. i. punctata.<\/em> Therefore I assume that <em>C. dyodon<\/em> has originated from an isolate of the common stem form in that region. <em>C. itala<\/em> (with the stenzioid subspecies = <em>C. clavata<\/em>) and <em>C. dyodon<\/em> may belong to a monophyletic group of closely related species, the superspecies of <em>C. itala.<\/em><br \/>\nFor the diversity within <em>C. dyodon<\/em> see Nordsieck (2002: 36).<\/p>\n<p style=\"text-align: left;\">6. Subdivision of <em>Charpentieria<\/em>:<\/p>\n<p style=\"text-align: left;\">The genus <em>Charpentieria<\/em> includes besides the superspecies of <em>C. itala<\/em> two further species, <em>C. ornata <\/em>(Rossm\u00e4ssler 1836) and <em>C. stenzii<\/em>. <em>C<\/em>. <em>ornata<\/em> differs from <em>C. itala<\/em> by shell characters somewhat more than the subspecies of the latter from each other (Nordsieck 1963: 198). <em>C. stenzii<\/em> differs from <em>C. itala<\/em> in shell morphology, but more in genital morphology (Nordsieck 1963: 191, 201). A subgenus <em>Itala<\/em> O. Boettger, in which <em>C. itala<\/em>, <em>C. ornata<\/em> and <em>C. stenzii<\/em> are united, has therefore been rejected (Nordsieck 2002: 36).<br \/>\nBy DNA analyses (see Appendix) it has been found out that <em>C. dyodon<\/em>, as assumed in the superspecies concept, is in fact more closely related to <em>C. itala<\/em> than <em>C. ornata<\/em>.<\/p>\n<p style=\"text-align: left;\"><strong>Map. Distribution of <em>C. itala<\/em> subspecies and <em>C. dyodon<\/em> (&#8222;stenzioid subspecies&#8220; of C. itala = <em>C. clavata<\/em> not <\/strong><strong>considered).<\/strong><br \/>\nMarked ranges cover the regions in which samples of the respective taxa have either been collected or are mentioned in the literature.<\/p>\n<p style=\"text-align: left;\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone size-medium wp-image-253\" src=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/I_App_1b_clausap.tif\" alt=\"\" width=\"1\" height=\"1\" \/><a href=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/Verbreitung-itala-Orig-Kopie.jpg\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone wp-image-256 size-medium\" src=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/Verbreitung-itala-Orig-Kopie-300x278.jpg\" alt=\"\" width=\"300\" height=\"278\" srcset=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/Verbreitung-itala-Orig-Kopie-300x278.jpg 300w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/Verbreitung-itala-Orig-Kopie-1024x949.jpg 1024w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/Verbreitung-itala-Orig-Kopie-768x712.jpg 768w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/Verbreitung-itala-Orig-Kopie-291x270.jpg 291w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/Verbreitung-itala-Orig-Kopie.jpg 1061w\" sizes=\"auto, (max-width: 300px) 100vw, 300px\" \/><\/a><\/p>\n<h4 style=\"text-align: left;\"><strong>Fig. 10.<em> C. i. braunii<\/em> (type form), on a wall near the castle of Weinheim, Germany.<\/strong><\/h4>\n<p style=\"text-align: left;\"><a href=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/braunii-Weinheim-Tiere.jpg\"><img loading=\"lazy\" decoding=\"async\" class=\"alignnone wp-image-257 size-medium\" src=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/braunii-Weinheim-Tiere-300x221.jpg\" alt=\"\" width=\"300\" height=\"221\" srcset=\"https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/braunii-Weinheim-Tiere-300x221.jpg 300w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/braunii-Weinheim-Tiere-367x270.jpg 367w, https:\/\/hnords.de\/wordpress\/wp-content\/uploads\/2021\/05\/braunii-Weinheim-Tiere.jpg 531w\" sizes=\"auto, (max-width: 300px) 100vw, 300px\" \/><\/a><\/p>\n<h4><strong>Appendix: DNA studies on <em>Charpentieria itala<\/em><\/strong><\/h4>\n<p>Subspecies:<\/p>\n<p>The subspecies division of <em>C<\/em>. <em>itala<\/em> (with stenzioid subspecies) from the southern Alps, as proposed in Nordsieck (1963) and once more in this chapter, has been confirmed, at least west of Lake Garda, in the DNA study of Scheel &amp; Hausdorf (2012). The western normal = non-stenzioid subspecies <em>C<\/em>. <em>i<\/em>. <em>albopustulata<\/em> and <em>C<\/em>. <em>i<\/em>. <em>latestriata<\/em> are accepted. In their AFLP network they form four clusters, <em>C<\/em>. <em>i<\/em>. <em>latestriata<\/em> one of them.<br \/>\nThe stenzioids (<em>C<\/em>. <em>i<\/em>. <em>clavata<\/em>, <em>C<\/em>. <em>i<\/em>. <em>variscoi<\/em>, <em>C<\/em>. <em>i<\/em>. <em>balsamoi<\/em>, <em>C<\/em>. <em>i<\/em>. <em>lorinae<\/em>), because of introgression from <em>C<\/em>. <em>itala<\/em>, \u00a0are treated as subspecies of <em>C<\/em>. <em>itala<\/em>. In the AFLP network the stenzioids form five clusters, because C. <em>i.<\/em> <em>trepida<\/em>, which had been included in <em>C<\/em>. <em>i<\/em>. <em>lorinae<\/em> in my 1963 paper, forms a cluster of its own. The transitional forms (<em>C<\/em>. <em>i<\/em>. <em>allatollae<\/em> of the 1963 paper) are assembled in an additional cluster.<br \/>\nThe downgrading of the stenzioids to subspecies of <em>C<\/em>. <em>itala<\/em> is unsatisfactory, because stenzioids behave in part like species, in part like subspecies (see Nordsieck 1963).<br \/>\nAs result of the recently published genome-wide DNA study of Xu &amp; Hausdorf (2020) <em>C<\/em>. <em>itala<\/em> revealed as genetically subdivided into two main clades, a western and an eastern one, separated by Lake Garda.<br \/>\nThe stenzioids are classified within two species. The western stenzioids, because free of genomic shares with <em>C<\/em>. <em>itala<\/em>, are treated as an independent species, while the eastern stenzioids, which exhibit genomic shares with <em>C<\/em>. <em>itala<\/em>, are regarded as subspecies of that species (<em>C<\/em>. <em>i<\/em>. <em>lorinae<\/em>, <em>C<\/em>. <em>i<\/em>. <em>trepida<\/em> included). This separation is also unsatisfactory, because\u00a0 the monophyly of the stenzioids is thus obscured. Besides, also in western stenzioids (<em>C<\/em>. <em>c<\/em>. <em>clavata<\/em>) introgression from <em>C<\/em>. <em>itala<\/em> has been proved (see Scheel &amp; Hausdorf). The transitional form (<em>C<\/em>. <em>i<\/em>. <em>leccoensis<\/em> of the 1963 paper) is neither considered by Scheel &amp; Hausdorf nor by Xu &amp; Hausdorf.<br \/>\nAs concerns the non-stenzioid subspecies in the western clade <em>C<\/em>. <em>i<\/em>. <em>albopustulata<\/em> and <em>C<\/em>. <em>i.<\/em> <em>latestriata<\/em> are accepted as subspecies, the latter, however, only because of its assumed origin by hybridization. The subspecies of the eastern clade are assembled in a subspecies named <em>C<\/em>. <em>i<\/em>. <em>itala<\/em> (because they are not separable in the trees).<br \/>\nThis lumping is not in accordance with the shell morphology, especially the development of lunellar and clausilium plate (types A and B), by which the eastern subspecies are well differentiated (see part 1). Besides, <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em> exhibits a long principal plica, by which it differs from the other subspecies. According to the authors the subspecies differ morphologically hardly from one another, which means that these morphological differences have not been considered. In the tree of all <em>C<\/em>. <em>i<\/em>. populations examined (fig. 5) the eastern subspecies are separable, if the samples are correctly determined. <em>C<\/em>. <em>itala<\/em> from the region of Verona had been determined by me (Nordsieck 1963) as <em>C<\/em>. <em>i<\/em>. <em>albopustulata<\/em>, now (see part 4) classified as <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em>, not as <em>C<\/em>. <em>i<\/em>. <em>rubiginea<\/em><em>, <\/em><em>C<\/em>. <em>itala<\/em> from Schio as <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em>, not as <em>C<\/em>. <em>i<\/em>. <em>rubiginea<\/em>. <em>C<\/em>. <em>itala<\/em> from Eisack valley (Atzwang) could be a form of <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em> like the neighbouring one of Klausen-Waidbruck. A good example that the subspecies can be distinguished is the correction of the determination of <em>C<\/em>. <em>itala<\/em> from Meran (see part 4), which has been confirmed by the study of Xu &amp; Hausdorf.<br \/>\nIt is interesting that in the tree fig. 5 along the branch of <em>itala<\/em>-E\u00a0 (concluded from figs. 2A and B) admixture from <em>C<\/em>. <em>i<\/em>. <em>albopustulata<\/em> (of <em>itala<\/em>-W) decreases across <em>C<\/em>. <em>i<\/em>. <em>baldensis<\/em> to <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em> from Verona, <em>C<\/em>. <em>i<\/em>. <em>itala<\/em> and <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em> (Schio). Admixture from typical <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em> decreases across <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em> (Meran) to <em>C<\/em>. <em>i<\/em>. <em>serravalensis<\/em> (Schio). The similarity of <em>C<\/em>. <em>i.<\/em> <em>baldensis<\/em>, <em>C<\/em>. <em>i<\/em>. <em>braunii<\/em> from Verona and <em>C<\/em>. <em>i<\/em>. <em>itala<\/em> with <em>C<\/em>. <em>i<\/em>. <em>albopustulata<\/em> concerning the closing apparatus (type B) is parallel to the admixture from W to E.<br \/>\nBecause of the morphological and genetical differences I think that it is not appropriate to lump these subspecies together.<\/p>\n<p>Species:<\/p>\n<p>Scheel &amp; Hausdorf (2012) have stated that <em>C<\/em>. <em>ornata<\/em> is the sister group of <em>C<\/em>. <em>itala<\/em> (+ stenzioid subspecies) and thus more closely related to <em>C<\/em>. <em>itala<\/em> than <em>C<\/em>. <em>stenzii<\/em>. <em>C<\/em>. <em>dyodon<\/em>, however, was not included in this study.<br \/>\nIn the DNA study of Xu &amp; Hausdorf (2020)\u00a0 it is shown that <em>C<\/em>. <em>dyodon<\/em>, as assumed in the superspecies concept proposed in part 5, is in fact more closely related to <em>C<\/em>. <em>itala<\/em> than <em>C<\/em>. <em>ornata.<\/em><\/p>\n<h4 style=\"text-align: left;\"><strong>References<\/strong><\/h4>\n<p style=\"text-align: left;\">Boato, A., Bodon, M. &amp; Giusti, F. (1985): Molluschi terrestri e d&#8217;acqua dolce delle Alpi Liguri. \u2013 Lavori della Societ\u00e0 Italiana di Biogeografia, (NS) 9 [1982]: 237-371.<\/p>\n<p>De Mattia, W., Reier, S. &amp; Haring, E. (2021): Morphological investigation of genital organs and first insights into the phylogeny of the genus <em>Siciliaria<\/em> Vest, 1867 as a basis for a taxonomic revision (Mollusca, Gastropoda, Clausiliidae). \u2013 ZooKeys, 1077: 1-175. https:\/\/doi.org\/10.3897\/zookeys.1077.67071.<\/p>\n<p style=\"text-align: left;\">Giusti, F. &amp; Mazzini, M. (1971): Notulae Malacologicae XIV. I molluschi delle Alpi Apuane. Elenco delle specie viventi con descrizione di una nuova specie: <em>Vitrinobrachium baccettii<\/em> n. sp. \u2013 Lavori della Societ\u00e0 Italiana di Biogeografia, (NS) 1 [1970]: 202-335, 9 pls.<\/p>\n<p style=\"text-align: left;\">Manganelli, G., Bodon, M., Favilli, L. &amp; Giusti, F. (1995): Gastropoda Pulmonata. In: Minelli, A., Ruffo, S. &amp; La Posta, S. (ed.): Checklist delle specie della fauna italiana, 16: 1-60. Bologna.<\/p>\n<p style=\"text-align: left;\">Martens, E. von (1857): Reisebemerkungen \u00fcber einige Binnenschnecken Italiens. \u2013 Malakozoologische Bl\u00e4tter, 4: 120-155.<\/p>\n<p style=\"text-align: left;\">Martens, G. von (1824): Reise nach Venedig, 2: VI + 664 pp., 9 pls. Ulm.<\/p>\n<p style=\"text-align: left;\">Michaud, A.-L.-G. (1831): Compl\u00e9ment de l&#8217;Histoire des mollusques terrestres et fluviatiles de la France, de J. P. R. Draparnaud. \u2013 VI+116+12 pp., pls. 14-16. Verdun.<\/p>\n<p style=\"text-align: left;\">Nordsieck, H. (1963): Zur Anatomie und Systematik der Clausilien, II. Die Formenbildung des Genus <em>Delima <\/em>in den S\u00fcdalpen. \u2013 Archiv f\u00fcr Molluskenkunde, 92 (5\/6): 169-203.<\/p>\n<p style=\"text-align: left;\">Nordsieck, H. (1972): Zur Anatomie und Systematik der Clausilien, XI. Neue Formen und taxonomische Revision einiger Gruppen der Alopiinae. \u2013 Archiv f\u00fcr Molluskenkunde, 102 (1\/3): 1-51, pls. 1-5.<\/p>\n<p style=\"text-align: left;\">Nordsieck, H. (1979): Zur Anatomie und Systematik der Clausilien, XXI. Das System der Clausilien, II: Die rezenten europ\u00e4ischen Clausilien. \u2013 Archiv f\u00fcr Molluskenkunde, 109 (4\/6): 249-275.<\/p>\n<p style=\"text-align: left;\">Nordsieck, H. (2002): Contributions to the knowledge of the Delimini (Gastropoda: Stylommatophora: Clausiliidae). \u2013 Mitteilungen der deutschen malakozoologischen Gesellschaft, 67: 27-39.<\/p>\n<p style=\"text-align: left;\">Nordsieck, H. (2007): Worldwide Door Snails (Clausiliidae), recent and fossil. \u2013 214 pp., 20 pls. Hackenheim (ConchBooks).<\/p>\n<p>Pirona, J. (1865): Prospetto dei molluschi terrestri e fluviatile finora raccolti nel Friuli. \u2013 Atti del Reale Istituto veneto di scienze, lettere ed arti, (3) 10: 675-708.<\/p>\n<p style=\"text-align: left;\">Scheel, B. &amp; Hausdorf, B. (2012): Survival and differentiation of subspecies of the land snail <em>Charpentieria itala<\/em> in mountain refuges in the Southern Alps. \u2013 Molecular Ecology, 21: 3794-3808.<\/p>\n<p>Schmidt, A. (1868): System der europ\u00e4ischen Clausilien und ihrer n\u00e4chsten Verwandten. \u2013 175 pp., 1 tab. Kassel (Fischer).<\/p>\n<p style=\"text-align: left;\">Strobel, P. (1851): Notizie malacostatiche sul Trentino. \u2013 114 pp. Pavia.<\/p>\n<p style=\"text-align: left;\">Xu, J. &amp; Hausdorf, B. (2020): Repeated hybridization increased diversity in the door snail complex <em>Charpentieria<\/em> <em>itala<\/em> in the Southern Alps. \u2013 Molecular Phylogenetics and Evolution. <a href=\"https:\/\/doi.org\/10.1016\/j.ympev.2020.106982\">https:\/\/doi.org\/10.1016\/j.ympev.2020.106982<\/a>.<\/p>\n<p style=\"text-align: left;\">Zilch, A. (1959-1960): Gastropoda. Teil 2. Euthyneura. \u2013 In: Schindewolf, O. H. (ed.), Handbuch der Pal\u00e4ozoologie, 6 (2): I-XII, 1-834. Berlin-Nikolassee.<\/p>\n<p style=\"text-align: left;\">Zilch, A. (1972): Die Typen und Typoide des Natur-Museums Senckenberg, 50: Mollusca: Clausiliidae (2): Alopiinae: Delimini (1) . \u2013 Archiv f\u00fcr Molluskenkunde, 102 (4\/6): 247-259, pl. 11.<\/p>\n<p style=\"text-align: left;\">\n","protected":false},"excerpt":{"rendered":"<p>Charpentieria itala: subspecies, species, superspecies Hartmut Nordsieck (V.2021) I. Introduction and history In the revision of the Charpentieria species from the southern limestone Alps (Nordsieck 1963: 174-190, as Delima) I tried to describe the intraspecific diversity of C. itala (G. von Martens 1824) by a subdivision of the species of that region into several subspecies. These are (from W to E) C. i. albopustulata (Cristofori &amp; Jan 1832), C. i. latestriata (K\u00fcster 1850), C. i. rubiginea (Rossm\u00e4ssler 1836), and C&#8230;.<\/p>\n<p class=\"read-more\"><a class=\"btn btn-default\" href=\"https:\/\/hnords.de\/wordpress\/charpentieria-itala\/\">Weiterlesen<span class=\"screen-reader-text\"> Weiterlesen<\/span><\/a><\/p>\n","protected":false},"author":1,"featured_media":0,"parent":0,"menu_order":0,"comment_status":"closed","ping_status":"closed","template":"","meta":{"footnotes":""},"class_list":["post-111","page","type-page","status-publish","hentry"],"_links":{"self":[{"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/pages\/111","targetHints":{"allow":["GET"]}}],"collection":[{"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/pages"}],"about":[{"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/types\/page"}],"author":[{"embeddable":true,"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/users\/1"}],"replies":[{"embeddable":true,"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/comments?post=111"}],"version-history":[{"count":38,"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/pages\/111\/revisions"}],"predecessor-version":[{"id":322,"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/pages\/111\/revisions\/322"}],"wp:attachment":[{"href":"https:\/\/hnords.de\/wordpress\/wp-json\/wp\/v2\/media?parent=111"}],"curies":[{"name":"wp","href":"https:\/\/api.w.org\/{rel}","templated":true}]}}