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Pulmonata Helicidae) Ruthenica, 2014, vol. 24, No. 2: 173-187. © Ruthenica, 2014 Published online November 24, 2014. http: www.ruthenica.com On the morphology of copulative apparatus of some Ariantinae (Pulmonata Helicidae) A.A. SCHILEYKO A.N. Severtzov Institute of Ecology and Evolution of Russian Academy of Sciences, Leninski prosp. 33, Moscow 119071, RUSSIA. E-mail: [email protected] ABSTRACT. The article contains additional data on Abbreviations in figures: At – atrium; DSt – diverticle of the structure of the penial papillae and the atrial stimu- spematheca; F – flagellum; FO – free oviduct; P – penis; Pil – lators of the representatives of 13 species and subspe- pilaster; PP – penial papilla; RP – retractor of penis; PS – cies of 8 (sub)genera of Ariantinae. The existence with- penis sheath; Sph – sphincter; Sti – stimulator; V – “velum”; in the subfamily of five or six variants of structure of the Va – vagina. copulative apparatus has been shown. Papilla of the Abbreviations of Museums: penis or its elements can take on the role of stimulator; HNHM – Hungarian Natural History Museum, Buda- in such cases stimulator mostly is reduced down to full pest, Hungary disappearance. Reproductive tract of some species has NHMW – Naturhistorisches Museum Wien, Austria been re-described. RMNH – National Museum of Natural History, Leiden, The Netherlands Introduction Results When trying to understand the taxonomic struc- Thiessea sphaeriostoma sphaeriostoma ture of any group of organisms inevitably a problem (Bourguignat, 1857) arises of selection of characters on which we can (Fig. 1) rely. Obviously, for analysis only those characters are suitable which provide the basis for a compari- Material: Greece, Thessalia, Magnissia, 4 km WSW of Marathia (32 km SE of Volos); 15 m alt. 28.V.1999. UTM F J son. One can distinguish only distinguishable things: 8232. Leg. et det. E. Gittenberger, D. Uit de Weerd. RMNH if two objects have same color, then the inclusion of Mol. 62488, 43730-Helicidae. 1 specimen. this feature to the analysis of the similarity/differ- ence between objects does not make sense. In the Vas deferens is rather long. Flagellum is about given case (Ariantinae), if to use for identification two times shorter than penis. Epiphallus very short of genera, for example, the structure of a jaw, or a (shorter than flagellum). Penis bulky. Penial retrac- kidney, or an external appearance of the reproduc- tor attached to the apex of a fleshy outgrowth tive tract, we do not get (or almost do not get) any located on the boundary between penis and epiphal- material for analysis of relationship between taxa. lus. Free oviduct somewhat longer than vagina. Earlier the author has published an article where Mucus glands are simple. considered, in particular, the variety of the structure Penis consists of two chambers: proximal one of the penial papillae and stimulators of 15 species contains papilla and separated from distal chamber of Ariantinae [Schileyko, 2013]. Most of them are by a sphincter; distal chamber bears an axial pilas- type species of genera or subgenera. Now I have ter. Penial papilla with almost smooth surface, thick got some material on the representatives of 8 genera walls, and narrow inner slit-like canal that opens on (subgenera) which have been considered in the the tip of the papilla by a small pore. Walls of papilla mentioned paper by only literature data. References filled with loose parenchyma. to the original descriptions and other information Stimulator as such is absent, unless to believe for each taxon are given in my earlier article [Schi- that a strong longitudinal pilaster, located in the leyko, 2013]. Here emphasis is placed on the struc- vagina and bearing the opening of stylophore, is its ture of the penis papilla and stimulator. derivative. The other branch of the pilaster runs into penis. Material and methods Cingulifera cingulata (Studer, 1820) List of the studied material follows the species (Fig. 2) names. Methods have been described in my previ- Material: Strada ponale und Ponale-Fall bei Riva [Italy]. ous article [Schileyko, 2013]. 1892. Leg. et det. R. Sturany. NHMW No. 19159. 1 specimen. 174 A.A. Schileyko Penial papilla is thick-walled, with very strong, vagina. Inner surface of the vagina with strong, superficial semicircular folds of irregular shape. locally swollen, axial pilasters. Inner surface of the penis has a relief correspond- Remark. Taxonomic position of this species is ing to that of papilla. Inner canal of the papilla is slit- not clear. The shape and size of the stimulator of C. like, of irregular form in cross-section, opens on petrovici is very similar to that of C. trizona, but the tip of the organ. Walls of the papilla contain very structure of the penial papilla in these two species is loose parenchyma saturated with numerous lacunae different: in C. petrovici, as stated above, there is an and sinuses. axial pilaster whereas in C. trizona the pilaster is Stimulator is rather small, conical, located in absent; in C. petrovici there is a slit on the surface vagina above atrium. Regular relief on the inner of the papilla; in C. trizona such a slit is absent. surface of the vagina is absent. In the meanwhile I am refraining from solving the problem, since I dissected only one specimen of Wladislawia polinskii (A. Wagner, 1988) each species. (Fig. 3) Causa holosericea (Studer, 1820) Material: Bulgaria, Blagoevgrad prov., Pirin Planina, Mt. (Fig. 5) Vikhren (30 km N of Melnik); alt. 2914 m, 8.IX.1967. Leg. et det. A.W. Janssen. RMNH, Mol. 336248, 43730-Helicidae. 2 Material: Austria, Tirol, Stubaital, Neustift, Milders; alt. specimens. ca. 1000 m; on walls along road, 9/28.VIII.1957. Leg. et det. C.O. Regteren Altena van. RMNH, Mol. 6738 43730-Heli- Vas deferens moderately long. Flagellum very cidae. 2 specimens. long, about two times longer than penis+epiphallus. Epiphallus rather thin. Penis consists of 2 parts: Vas deferens is comparatively short, entering narrowed, twisted distal part, the curves of this part epiphallus subapically. Epiphallus long, thickened in are fixed by numerous thin muscle fibers (as in its proximal section. Flagellum is short, rudimenta- Causa – see below); proximal part is bulky, thin- ry. Penis consists of two distinct parts: narrowed, walled, surrounded by semitransparent sheath. Pe- twisted distal part, the curves of this part are fixed nial retractor attached to epiphallus well above pe- by numerous thin muscle fibers; proximal part is nis. Free oviduct and vagina are not long, of about swollen, spherical, very thin-walled, surrounded by equal length. Mucus glands forked. very thin, nearly transparent sheath. Penial retractor Penial papilla is of irregular shape, thick-walled, attached to middle section of epiphallus. Free ovi- with few superficial axial grooves and minute apical duct long, curved; vagina somewhat shorter, its pore. Inner surface of penis with longitudinal, swol- inner surface lacks regular relief (almost smooth). len, locally thickened pilasters; rounded fossae on Mucus glands are simple. the papilla surface correspond to these thickenings. Penial papilla short to very short, thick-walled, Walls of the papilla contain loose parenchyma and its walls filled with loose parenchyma. Papilla sur- small lacunae. Internal canal of the papilla narrow, face bears deep circular grooves; relief of inner in form of 3- or 4-rayed star. surface of the penis corresponds to these grooves. Stimulator not large, ovate, occupies transverse Inner canal of the papilla is slit-like, roughly semilu- position in the atrium. Inner surface of vagina sup- nar in cross-section, open terminally. plied with one or two smoothed, broad longitudinal Stimulator is moderately large, lamellar, located folds. obliquely-transversely in the atrium, directed to- ward the penis. Cattania petrovici (A.J. Wagner, 1914) Kosicia intermedia (C. Pfeiffer, 1828) (Fig. 4) (Fig. 6) Material: Albania, 2010/55, Shkodër district, Prokletije Material: Am Füâ des kleinen Tal am Plöken-Pass, Kar- Mts., Breg-Lumi, Shalë River in the village, 360 m, N nische Alpen [Austria], 16.7.1907. Leg. et det. R. Sturany. 42º18.258’ E 19º47.814’, 23.05.2010. Leg. et det. Z. Fehér, NHMW No 44.658. 1 specimen. D. Murányi, Zs. Újvári. HNHM. 1 specimen. Austria, Karnische Alpen, Plöckenpass, 07.08.2008. Penial papilla is large, with circular grooves in its NHMW No SpID#181, inID#2108. From project “Alpine basal part. On the proximal part of the papilla there landsnails”. 1 specimen. is a vast, wide longitudinal slit; on the bottom of the Vas deferens moderately long, entering epiphal- slit lies a wide axial pilaster (similar to that in lus apically. Flagellum markedly longer then Dinarica pouzolzi or Arianta arbustorum – see penis+epiphallus. Epiphallus short, thick, narrowed Schileyko, 2013, Figs. 12 and 15). Orifice of epiphal- in proximal part. Penis is bulky, not long. Penial lus opens at the base of this pilaster. Walls of the retractor attached to the middle of epiphallus. In papilla contain a loose parenchyma and large lacu- one of dissected specimens there was an additional nae. branch of the retractor that connects the main Stimulator large, fleshy, conic, directed to the branch with the atrium, in the other specimen the Copulative apparatus in Ariaphantinae 175 FIG. 1. Thiessea sphaeriostoma. A – reproductive tract. B – inner structure of atrium and vagina. C – inner structure of penis. Asterisk – stylophore opening. РИС. 1. Thiessea sphaeriostoma. A – репродуктивный тракт. B – внутреннее строение атриума и вагины. C – внутреннее строение пениса. Звёздочка – отверстие стилофора. FIG. 2. Cingulifera cingulata. A – inner structure of vagina. B – inner structure of penis. РИС. 2. Cingulifera cingulata. А – внутреннее строение вагины. В – внутреннее строение пениса. 176 A.A. Schileyko FIG. 3. Wladislawia polinskii. A – reproductive tract. B – distal part of penis. C – inner structure of vagina. D – inner structure of penis and papilla from other side.
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