{"id":223,"date":"2021-05-22T21:52:37","date_gmt":"2021-05-22T21:52:37","guid":{"rendered":"https:\/\/hnords.de\/wordpress\/?page_id=223"},"modified":"2021-12-10T09:17:01","modified_gmt":"2021-12-10T08:17:01","slug":"alopia","status":"publish","type":"page","link":"https:\/\/hnords.de\/wordpress\/alopia\/","title":{"rendered":"Alopia"},"content":{"rendered":"

Genital studies of enantiomorph taxa pairs of Alopia<\/i><\/b><\/p>\n

Hartmut Nordsieck (III.2021)<\/p>\n

Introduction<\/h4>\n

Alopia<\/i> species from the southern Carpathians, Romania, are left-coiled (L) or right-coiled (R). In some mountains left- and right-coiled taxa, which look like mirror-images = enantiomorphic taxa, occur together. These taxa have been treated until now as species or as subspecies of one species. They are similar in shell and genital characters; in some cases it was shown that they hybridize. If more widely distributed, as in three mountains of the region (Piatra Craiului, Bucegi, Ciuca\u015f), the ranges of the enantiomorphs (and their subspecies) are not separated, but mosaic-like meshed with that of the other enantiomorph, with occasional syntopic occurrences of both enantiomorphs (Nordsieck 2008). Therefore, to avoid imbalances within the genus and obscuration of the diversity, most enantiomorphic taxa are treated here as species (same arguments as for hybridizing species in general, see website article on hybridization).
\nA good example to show the problems of uniting enantiomorphic taxa to species is given by the\u00a0A<\/i>.\u00a0livida<\/i>\u00a0group from the Bucegi Mountains. If one unites the species of that group (A<\/i>.\u00a0straminicollis<\/i>,\u00a0A<\/i>.\u00a0livida<\/i>\u00a0with\u00a0A<\/i>.\u00a0l<\/i>.\u00a0fussi<\/i>,,A<\/i>.\u00a0nixa<\/i>), which hybridize, to one (mega)species (Koch et al. 2020), one must do this also with other enantiomorph taxa pairs of the genus. These megaspecies would then contain subspecies of different rank, the enantiomorphs and the subspecies of the enantiomorphs each. Besides, in the megaspecies\u00a0A<\/i>.\u00a0livida<\/i>\u00a0group, one had to include also\u00a0A<\/i>.\u00a0glorifica<\/i>\u00a0from the Piatra Craiului Mountains (and its enantiomorph\u00a0A<\/i>.\u00a0lischkeana<\/i>, both belonging to another megaspecies), because it hybridizes with\u00a0A<\/i>.\u00a0livida<\/i>\u00a0in that part of the mountains where the latter species occurs (M\u0103gura Mountains) (Nordsieck 2016).<\/p>\n

Genital studies of enantiomorphs<\/h4>\n

In the course of my work on Alopia<\/i>\u00a0the genital organs of 36 species taxa have been investigated. It was of special interest to state if enantiomorphic\u00a0Alopia<\/i>\u00a0taxa exhibit genital differences. The literature dealing with is not sufficient, because descriptions and figures of the genitalia are not exact enough. Therefore, the genitalia of ten enantiomorph taxa pairs have been examined and measured.
\nThe enantiomorphs occur sympatrically, but not syntopically (except taxa pair VI from Bucegi Mountains, Valea Velicanului).
\nThe taxa pairs are ordered from W to E. The names are those of the system presented in Nordsieck (2008).
\nThe preparation numbers are given in brackets. Two specimens each, in some cases three, have been examined.<\/p>\n

I:
\nA<\/i>.\u00a0subcosticollis occulta<\/i>\u00a0(L), Cheile Coste\u015fti near Pietreni (418),
\nA<\/i>.\u00a0hildegardae fortunata<\/i>\u00a0(R), Cheile Bistri\u0163a near Pietreni (417).
\nThe enantiomorphs have a relatively short epiphallus (ratio epiphallus\/penis ~ 1.1) in common.
\nIn\u00a0A<\/i>.\u00a0h<\/i>.\u00a0fortunata<\/i>\u00a0penis and epiphallus are shorter (in comparison ~ 0.8).<\/p>\n

II:
\nA<\/i>.\u00a0glorifica boettgeri<\/i>\u00a0(L), Piatra Craiului Mts.,Valea R\u00e2ului (lower part) (419),
\nA<\/i>.\u00a0lischkeana obesa<\/i>\u00a0(R), Piatra Craiului Mts.,Valea R\u00e2ului (upper part) (420).
\nThe enantiomorphs have a relatively long epiphallus (ratio epiphallus\/penis ~ 1.4) in common.
\nIn\u00a0A<\/i>.\u00a0l. obesa<\/i>\u00a0the proximal parts of bursa copulatrix (diverticulum, bursa + proximal pedunculus) (~ 0.7) and the epiphallus (~ 0.8) are shorter.<\/p>\n

III:
\nA.glorifica glorifica<\/i>\u00a0(L), Piatra Craiului Mts., Degetul lui C\u00e2lnic (385),
\nA<\/i>.\u00a0lischkeana lischkeana<\/i>\u00a0(R), Piatra Craiului Mts., Pr\u0103p\u0103stii (386).
\nThe genitalia of the enantiomorphs are different.
\nIn\u00a0A<\/i>.\u00a0l. lischkeana<\/i>\u00a0the vagina (~ 0.7) and the male copulatory organs, especially the distal part of epiphallus (glorifica<\/i>\u00a0ratio proximal\/distal part of epiphallus ~ 1.0,\u00a0lischkeana<\/i>\u00a0~ 1.5) are shorter.<\/p>\n

IV:
\nA.glorifica glorifica<\/i>\u00a0(L), Piatra Craiului Mts., Valea Brusturetului above Cabana Brusturet (Valea Seac\u0103 a pietrelor, 1050 m a. s. l.) (682),
\nA<\/i>.\u00a0lischkeana lischkeana<\/i>\u00a0(deceptans<\/i>) (R), Piatra Craiului Mts., Valea Brusturetului above Cabana Brusturet (Valea Seac\u0103 a pietrelor, 1120 m a. s. l.) (683).
\nThe enantiomorphs have a relatively long epiphallus (ratio epiphallus\/penis ~ 1.8) in common.
\nIn\u00a0A<\/i>.\u00a0l<\/i>.\u00a0lischkeana<\/i>\u00a0the proximal parts of bursa copulatrix (diverticulum, bursa + proximal pedunculus) (~ 0.7 and 0.8, respectively) and the penis and proximal part of epiphallus (both ~ 0.8) are shorter.
\nNote the correspondence to the enantiomorph pair II.
\nThis is the taxa pair, in which according to Feh\u00e9r et al. (2013) the COI sequences are identical.<\/p>\n

V:
\nA<\/i>.\u00a0glorifica galbina<\/i>\u00a0(L), Piatra Craiului Mts., M\u0103gura Mare (southern slope below summit, ~ 1250 m a. s. l.) (700),
\nA<\/i>.\u00a0livida deaniana<\/i>\u00a0(R), Piatra Craiului Mts., M\u0103gura Mare (northern slope between summit and eastern depression, ~ 1250 m a. s. l.) (701).
\nThe genitalia of the enantiomorphs are different.
\nIn\u00a0A<\/i>.\u00a0l<\/i>.\u00a0deaniana<\/i>\u00a0the diverticulum is longer (~ 1.5), the penial caecum shorter (~ 0.65) and the distal part of epiphallus longer (~ 1.4).<\/p>\n

VI:
\nA<\/i>.\u00a0straminicollis straminicollis<\/i>\u00a0(L), Bucegi Mts., Valea Velicanului (upper part, ~ 1700 m a. s. l.) (698),
\nA<\/i>.\u00a0livida hypula<\/i>\u00a0(R), Bucegi Mts., Valea Velicanului (upper part, ~ 1700 m a. s. l.) (699).
\nThe genitalia of the enantiomorphs, which in contrast to the other pairs occur syntopically, are largely corresponding, but especially one character of\u00a0A<\/i>.\u00a0l<\/i>.\u00a0hypula<\/i>, the length of the proximal part of epiphallus, is like that of\u00a0A<\/i>.\u00a0l<\/i>.\u00a0bipalatalis<\/i>\u00a0occurring nearby (668, 679) (see table). This is a further indication of the origin of\u00a0A<\/i>.\u00a0l<\/i>.\u00a0hypula<\/i>\u00a0from\u00a0A<\/i>.\u00a0l. bipalatalis<\/i>\u00a0by hybridization with\u00a0A<\/i>.\u00a0s<\/i>.\u00a0straminicollis<\/i>\u00a0(see Nordsieck 2016).<\/p>\n

In the following table the length ratios of\u00a0A<\/i>.\u00a0straminicollis<\/i>\u00a0and\u00a0A<\/i>.\u00a0livida<\/i> from different localities of Bucegi Mountains are given (two specimens each measured).<\/p>\n

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VII:
\nA<\/i>.\u00a0straminicollis monacha<\/i>\u00a0(L), Bucegi Mts., Cheile Pe\u015fterii (~ 1550 m a. s. l.) (430),
\nA<\/i>.\u00a0livida nubila<\/i>\u00a0(R), Bucegi Mts., Cheile Coteanu (1490 m a. s. l.) (654).
\nThe genitalia of the enantiomorphs are different (see table).
\nIn\u00a0A<\/i>.\u00a0s<\/i>.\u00a0monacha\u00a0<\/i>vagina (~ 1.9), epiphallus (~ 1.6) and its proximal part (~ 1.8) are longer than in\u00a0A.l<\/i>.\u00a0nubila<\/i>.<\/p>\n

VIII:
\nA<\/i>.\u00a0nixa\u00a0<\/i>(L), Bucegi Mts., Valea Ob\u00e2r\u015fia Ialomi\u0163ei (~ 2000 m a. s. l.) (425),
\nA.livida fussi<\/i>\u00a0(R), Bucegi Mts., Valea Doamnelor (~ 1850 m a. s. l.) (424).
\nThe enantiomorphs have the genital characters male copulatory organs lengthened, penial caecum reduced in common.
\nIn\u00a0A<\/i>.\u00a0l. fussi\u00a0<\/i>the penis is hardly longer than in\u00a0A<\/i>.\u00a0nixa<\/i>. Its penial caecum is reduced like in that species (in the literature, however, it is given as absent).
\nNote that despite of their shell similarity the male copulatory organs of A<\/i>.\u00a0l<\/i>.\u00a0fussi<\/i>\u00a0are much different from those of other subspecies of\u00a0A<\/i>.\u00a0livida<\/i>. Like in\u00a0A<\/i>.\u00a0nixa<\/i>\u00a0they are similar to those of\u00a0A<\/i>.\u00a0pomatias<\/i>.<\/p>\n

IX:
\nA<\/i>.\u00a0mauritii<\/i>\u00a0(L), Ciuca\u015f Mts., Mun\u0163ele Ro\u015fu (mountain) (426),
\nA<\/i>.\u00a0nefasta<\/i>\u00a0(R), Ciuca\u015f Mts., Mun\u0163ele Ro\u015fu (above Valea Berii) (457).
\nThe enantiomorphs have a relatively long epiphallus (mauritii\u00a0<\/i>ratio epiphallus\/penis ~ 1.8,\u00a0nefasta<\/i>\u00a0~ 1.5) in common.
\nIn\u00a0A<\/i>.\u00a0nefasta\u00a0<\/i>vagina (~ 1.8) and male copulatory organs (penis ~ 1.6, epiphallus ~ 1.4) are longer.<\/p>\n

X:
\nA<\/i>.\u00a0helenae zagani\u00a0<\/i>(L), Ciuca\u015f Mts., Grop\u015foare (SW, ~ 1300 m a. s. l.) (454),
\nA<\/i>.\u00a0helenae helenae<\/i>\u00a0(R), Ciuca\u015f Mts., Grop\u015foare (SW, ~ 1350 m a. s. l.) (453).
\nThe enantiomorphs have a relatively long epiphallus (zagani\u00a0<\/i>ratio epiphallus\/penis ~ 2.0,\u00a0helenae<\/i>\u00a0~ 2.2) in common.
\nIn\u00a0A<\/i>.\u00a0h<\/i>.\u00a0helenae<\/i>\u00a0the proximal parts of bursa copulatrix(diverticulum, bursa + proximal pedunculus) are somewhat longer (~ 1.2).<\/p>\n

It results that the genitalia of most enantiomorphs are largely, but not fully corresponding. The differences are comparable with those of the species taxa of\u00a0Alopia<\/i>\u00a0in general. This concurs with the shell morphology.
\nThe enantiomorphic taxa are found in the same groups of the COI tree of Feh\u00e9r et al. (2013), except of taxa pairs III, V, VI and VII.
\nThe given measurements are those of few specimens. Therefore, measuring must be expanded to check the stated genital differences of the enantiomorphs.<\/p>\n

References<\/h4>\n

Feh\u00e9r, Z., N\u00e9meth, L., Nicoar\u0103, A. & Szekeres, M. (2013): Molecular phylogeny of the land snail genus Alopia<\/i>\u00a0(Gastropoda: Clausiliidae) reveals multiple inversions of chirality. \u2013 Zoological Journal of the Linnean Society,\u00a0167<\/b>: 259-272.<\/p>\n

Koch, E. L., Neiber, M. T., Walther, F. & Hausdorf, B. (2020): Patterns and processes in a non-adaptive radiation:\u00a0Alopia<\/i>\u00a0(Gastropoda, Clausiliidae) in the Bucegi Mountains. \u2013 Zoologica Scripta,\u00a049<\/b>\u00a0(3): 280-294.<\/p>\n

Nordsieck, H. (2008): The system of the genus\u00a0Alopia\u00a0<\/i>H. & A. Adams 1855 (Gastropoda: Stylommatophora: Clausiliidae). \u2013 Mitteilungen der deutschen malakozoologischen Gesellschaft,\u00a079\/80<\/b>: 7-18.<\/p>\n

Nordsieck, H. (2016): Interspecies hybridization in the genus\u00a0Alopia<\/i>\u00a0(Gastropoda, Stylommatophora, Clausiliidae) from southern Carpathians, Romania, demonstrated by shell examination.\u2013 Conchylia,\u00a046<\/b>\u00a0(1-4): 3-15, 3 pls.<\/p>\n

 <\/p>\n

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Genital studies of enantiomorph taxa pairs of Alopia Hartmut Nordsieck (III.2021) Introduction Alopia species from the southern Carpathians, Romania, are left-coiled (L) or right-coiled (R). In some mountains left- and right-coiled taxa, which look like mirror-images = enantiomorphic taxa, occur together. These taxa have been treated until now as species or as subspecies of one species. They are similar in shell and genital characters; in some cases it was shown that they hybridize. If more widely distributed, as in three…<\/p>\n

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