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«Sitala rumbangensis E. S m i t h, 1895 (Mollusca, Pulmonata) an unexpected inhabitant of the Zoo in Vienna A. A. Schileyko* Abstract A tropical ...»

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©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at

Ann. Naturhist. Mus. W ien, B 111 5-12 Wien, März 2010

"Sitala" rumbangensis E. S m i t h, 1895 (Mollusca, Pulmonata) an unexpected inhabitant of the Zoo in Vienna

A. A. Schileyko*


A tropical species o f land snails has been found in the Zoo o f Vienna. This species has been described from

Kalimantan under the name Sitala rumbangensis E. S m i t h, 1895. The anatomical investigation has shown that actually this species belongs to a new genus, Schwammeria gen.n. (family Helicarionidae). Conchological redescription o f the species, along with a description o f its anatomy and a discussion o f its taxonomic position is presented.

Key words: Gastropoda, Ariophantidae, Schwammeria, new genus, taxonomy, Tiergarten.

Zusammenfassung Eine tropische Landschneckenart wurde im Tiergarten Schönbrunn in W ien gefunden. Sie wurde als die Art Sitala rumbangensis E. S m i t h, 1895 bestimmt, die aus Kalimantan (Borneo) beschrieben wurde. A nato­ mische Untersuchungen zeigten, dass diese Art in eine neue Gattung, Schwammeria gen.n. (Familie Helica­ rionidae) zu stellen ist. Die Art wird anhand ihrer Schale wiederbeschrieben, zusammen mit ihrer Anatomie und ihre systematische Stellung wird erörtert.

Introduction On 18 and 21 April 2008, the author, together with Mag. Anita and Magdalena Eschner, collected in the Tropenhaus of Tiergarten Schönbrunn (Vienna Zoo) 49 specimens of a species which has been determined as Sitala rumbangensis E. S m it h, 1895.

The exact locality from which these snails came is unknown, but I have been told that the plants, soil and debris (pieces of wood, dry leaves) in the Tropenhaus were brought from Borneo (Kalimantan). Evidently, the snails were introduced together with soil and plant material.

Anatomical investigation showed that this species substantially differs from all other Helicarionoidea and must be segregated as a new genus.

Conchological redescription of this species, along with a description o f its anatomy and a discussion o f its taxonomic position is presented.

Material and methods Three specimens were dissected and drawings of the shell and the reproductive tract were made using an Olympus SZ 51 stereo microscope. The material is deposited in the A natoly A. Schileyko, A.N. Severtzov Institute o f Problem s o f Evolution o f Russian A cadem y o f Sciences, Moscow, Russia - asch@ gol.ru ©Naturhistorisches Museum Wien, download des Naturhistorischen

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Naturhistorisches M useum Wien, Austria (NHMW ) and the Zoological Museum of M oscow State University, Russia (ZMMU). Other abbreviations used: NHML, The Natural History Museum, London, UK.

Comparative material examined: Kalielia barrakporensis (L. P f e i f f e r, 1852). "Wewa Inhalagala, Forest Yoda Ela FR [Sri Lanka], alt. 73 m, 2.12.1999", NHML 2008.1052.

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Type species. Sitala rumbangensis E. Sm ith, 1895 Derivatio nominis. The genus is named in honor of Dr. Harald Schwammer (Tiergarten Schönbrunn), who kindly permitted me to collect the material in the Tropenhaus.

Diagnosis. Shell small, trochoid, thin-walled, with filiform keel; upper surface (above keel) finely radially striated, basal surface (below keel) with delicate spiral lines.

Umbilicus closed.

Conchologically, Schwammeria is similar to Kaliella and Sitala\ anatomically it differs from both by its rudimentary penis. It also differs from Kaliella by the presence of a well-developed spermatheca, and from Sitala by the peculiar shape of the spermathecal reservoir.

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Locus typicus. "Rumbang, Sarawak, and Mount Rabong" [Kalimantan = Borneo].

Redescription. Shell (Fig. 1) trochoid, thin, translucent, silky glossy. Outline o f spire very slightly convex (tangent-line not straight), whorls much flattened. Suture shallow, margined. Last whorl not descending, with thread-like peripheral keel that is visible above suture on earlier whorls and becomes weaker toward aperture. Color uniformly corneous. Embryonic whorls with hardly expressed radial wrinklets (nearly smooth), subsequent whorls with very delicate, crowded, not veiy regulär radial wrinklets; basal surface below keel with fine but quite distinct, impressed spiral lines. Aperture semilunate, m oderately oblique, with simple, fragile margins; colum ellar m argin shortly reflexed, scarcely thickened. Um bilical depression shallow, um bilicus completely covered. Whorl number up to 6.5. Height up to 3.7, diameter up to 3.3 mm; dimensions of pictured shell: height 3.7, diameter 3.3 mm.

©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at Figs 1-4: Schwammeria rum bangensis (E. Smith, 1895) comb.n.: (1) Shell from Tiergarten Schönbrunn; (2) posterior end o f cephalopodium; (3) jaw; (4) reproductive tract. CH, caudal horn; HD, hermaphroditic duct; P, penis; PR, penial retractor; PVG, perivaginal gland; RS, reservoir of spermatheca; SS, spermathecal stalk; VD, vas deferens.

Cephalopodium (Fig. 2) with blunt caudal projection ("horn") and indistinct caudal foss.

Jaw (Fig. 3) oxygnathous, thin, with wide, blunt projection on cutting edge.

Vas deferens uniformly slender, entering penis subapically at sharp angle. Flagellum or epiphallus absent. Penis very small, simple, rudimentary (Fig. 4 depicts specimen with the largest penis), without sheath; no special inner structure detected. Penial retractor thin, attached to penis apically at entrance of vas deferens. Free oviduct long, its distal half surrounded by distinct perivaginal gland. Vagina as such practically absence because spermathecal stalk is branched off from genital atrium. Basal portion of sper­ mathecal stalk strongly expanded, intemally with circular folds (as seen in transmitted light). This expanded portion passes to a thread-like duct; reservoir of spermatheca elongated, of peculiar sandglass shape (with narrowing in middle). Reservoir lies on male side o f spermoviduct, barely not reaching albumen gland.

©Naturhistorisches Museum Wien, download des Naturhistorischen Annalen unter www.biologiezentrum.at Museums in Wien, B, 111

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On the species name. Dr. Menno Schilthuizen and Dr. Jaap Vermeulen (pers. comm.) suppose, with some reservations, that K. rumbangensis could be a junior synonym of Kaliella barrakporensis (L. P f e i f f e r, 1852). To clarify this problem I am citing the ori­

ginal description of Helix barrakporensis (L. P f e if f e r 1852: 156):

"H. testa subperforata, elevato-trochiformi, tenui, striatula, nitida, pellucida, fusco-cornea; spira conica, acutiuscula; sutura profunda; anfract. 6, convexis, lente accrescentibus, ultimo carinato, non descendente, basi convexiuscula; apertura vix oblique, depressa, subangulato-lunari; perist. simplice, tenui, margine colum ellari brevi, ad perforationem punctiformem reflexiuscula. Diam. 3/4, alt. 3/4 mill. Hab. ad Barrakpore, Indiae (Bacon)."

The shells from the Tropenhaus differ from the above description by at least five characters: 1) spire rather cupola-like than conical; 2) summit obtuse rather than "acutius­ cula"; 3) suture is shallow, by no means profound; 4) whorls flattened, not "convexis";

5) umbilicus definitely closed (shell is not "subperforata"). Above all, Pfeiffer did not mention the presence o f a spiral striation on the basal surface.

"Sitala" rumbangensis E. S m Museum Wien,-download unter www.biologiezentrum.at o f the Zoo in Vienna ©Naturhistorisches i t h, 1895 an unexpected inhabitant 9

S c h il e y k o :

Fig. 6: Comparison o f structure o f reproductive tracts of Conulema, Sitala and Schwammeria gen.n. (schematized). Sarcobelum dotted, caecum shaded.

The original description o f Sitala rumbangensis S m it h, 1895 is as follows: "Testa vix perforata, conica, fusco-cornea, nitida, ad peripheriam filo-carinata, lineis incrementi levibus striata, inferne striis concentricis tenuissimis sculpta; spira obtuse conoidalis;

anfractus 6, convexi, lentea ccrescentes, sutura carina saturatiore marginata sejuncti;

apertura parva, lunata; peristoma tenue, margine columellari fortiter incrassato, leviter reflexo. Diam. maj. 4.5 millim., min. 4, alt. 4.5."

Besides, S m it h (1895) adds an unnamed variety: "Var. Testa typo paulo angustior, anfractibus 6 1/2 striisque basalibus minus conspicuis. Diam. maj. 3.3 millim., min. 3.2, alt. 4. Hab. Mulu Mountain, N. Borneo."

The comparison of the descriptions of the mollusks I investigated with the original one shows no essential differences between them except for the umbilicus: in the original description "teste vix perforate", whereas in my shells the umbilicus closed. It is possible that this difference depends on the quality of the optics: under a simple lens it is difficult to establish whether the umbilicus is closed or barely open.

Shell photographs of the syntypes of Sitala rumbangensis and Helix barrakporensis (deposited in the NHML) which Dr. Naggs kindly sent me, convinced me that my mate­ rial was identified correctly.

Taxonom ic position of the genus Schwammeria. Helix barrakporensis L. P f e i f f e r, 1852 is the type species o f the genus Kaliella B l a n f o r d, 1863 (subsequent designation ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at 10 Anncilen des Naturhistorischen M useums in Wien, B, 111

by G o d w in -A u ste n, 1882). Its anatom y has been studied by G o d w in -A u ste n (1883:

146, pl. 38, fig. 5). B l a n f o r d & G o d w in -A u sten (1908: 259; fig. 81A) have repeated the o riginal anatom ical description and illustration. A ccording to G o d w in -A u ste n (1883), K. barrakporensis has norm ally a developed penis provided w ith calc-sac; no indication o f the presence o f sperm atheca. Besides, G odw in-A usten did not m ention the presence o f a perivaginal gland. Through courtesy o f Dr. Fred N aggs and Dr. Jonathan A b lett I have d issected tw o specim ens o f Kaliella barrakporensis from Sri L anka (unfortunately, both specim ens were retracted). The dissection (Fig. 5) showed that the

description and illustration m ade by G odw in-A usten are correct, except for one detail:

this author did not m ention the perivaginal gland. This organ actually is present, but not very distinct, th at’s why it has been overlooked by G odw in-A usten. The calc-sac in the individual I dissected (Fig. 5) looks like a flagellum, but is filled w ith m ilky-w hite fluid

- a peculiar character o f ariophantoid calc-sacs. Jaw and the stm cture o f the posterior end o f the cephalopodium in K. barrakporensis are very sim ilar to that o f Schwammeria rumbangensis (Fig. 6).

Even if we admit that the snails from the Tiergarten are hemiphallic or aphallic representatives o f Kaliella (the phenomenon of hemiphallia or aphallia is well known among Stylommatophora), the presence of well-developed spermatheca testifies against this species belonging to the same genus as barrakporensis, i.e. to the genus Kaliella.

Many species under the generic name "Kaliella" are known, but unfortunately we know little about their anatomy (except for the type species - barrakporensis). This is why it is impossible to determine how many of them really belong to the genus Schwammeria, which conchologically does not differ reliably from Kaliella. This group needs thorough revision on an anatomical basis.

As the species rumbangensis originally has been placed in the genus Sitala H. A dam s, 1865 (type species - Helix infula B enson, 1848, by original designation), it should be noted that m y m aterial anatom ically differs from this genus by the rudim entary penis (in Sitala the penis is norm ally developed) and the absence o f an epiphallic caecum to w hich the penial caecum is attached (for the anatom y o f Sitala infula see S to lic z k a 1871, pl. 18, figs 5-9).

Among Euconulidae or Helicarionidae, I have not found any genus to which the species with the above-described anatom ical characters could be assigned. I suppose that Schwammeria is a relative of Sitala for three reasons: 1) The shape o f the spermathecal stalk is similar to that of Schwammeria, in particular, the basal section o f the stalk is expanded; 2) The absence o f a vagina because the spermathecal stalk sits on the genital atrium. 3) If we mentally remove the caecum in Sitala infula, we obtain a Situation similar to that observed in Schwammeria.

S to lic z k a (1971: 237, pl. 18, fig. 1^-) describes and illustrates another species - Sitala attegia (B enson, 1859) (from Nyanmar). It differs from S. infula by the presence of a well-developed sarcobelum. Stoliczka's illustrations o f the anatomy o f Sitala infula and S. attegia were recently reproduced by S c h ile y k o (2002: 1290, fig. 1694). Note that S to lic z k a (1871: 237) stated: "The presence or absence o f an amatorial gland (i.e. sar­ cobelum) cannot be accepted as a generic character, which will be evident from what I shall presently say in comparing the generative organs o f C. attegia with those of C.

"Sitala" rumbangensis E. S m iMuseum Wien,-download unter www.biologiezentrum.ato f the Zoo in Vienna ©Naturhistorisches t h, 1895 an unexpected inhabitant 11

S c h il e y k o :

i n f u l a S to lic z k a (1871) assigned these species to the genus Conulema S to lic z k a, 1871: 236 (type species Helix attegia B enson, 1859, by original designation). From a phylogenetic point o f view, I think Stoliczka is correct. B ased on the analysis o f vast m aterial, I have suggested that the disappearance o f accessory organs o f the reproductive tract in Pulm onata, in particular in helicarionoid groups, is a com m on path o f morphological evolution (S c h ile y k o 1991a, b, 2003). If true, the new genus would occupy the last place in the evolutionary row: Sitala attegia (both sarcobelum and caecum are present) - S. infula (sarcobelum is absent, caecum is present) - Schwammeria gen.n.

(both sarcobelum and caecum are absent) (Fig. 6).

With regard to the taxonomical position of the genus Kaliella, it should be noted, that this genus and three other considered genera {Sitala, Conulema and Schwammeria gen.n.), according to the family diagnoses, belong to different families: Kaliella to Euconulidae (Kaliellinae), the three other genera to Helicarionidae.

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