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A Redescription of Remanella Multinucleata (Kahl, 1933) Nov

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A Redescription of Remanella Multinucleata (Kahl, 1933) Nov Europ. .J. Protistol. 32,234-250 (1.996) European Journal of Mav 31, 1996 PROTISTOLOGY A Redescription of Remanella multinucleata (Kahl, 1933) nov. gen., nov. comb. (Ciliophora, Karyorelictea), Emphasizing the lnfraciliature and Extrusomes Wilhelm Foissner Universität Salzburg, lnstitut für Zoologie, Salzburg, Austria SUMMARY The morphology, infraciliature, and extrusomes of Remanella mwbinwcleata (Kahl, 1933) nov. comb. were studied in live cells, in protargol impregnated specimens, and with the scanning electron microscope. The entire somatic and oral infraciliature consists of diki- netids which have both basal bodies ciliated or only the anterior or posterior ones, de- pending on the region of the cell. The right side is densely ciliated. Its most remarkable specialization is a kinety which extends on the dorsolateral margin from mid-body to the tail, where the kinetids become condensed and associated with conspic- uous fibres originating from the ciliated anterior basal bodies. The left side seemingly has two ciliary rows extending along the cell margins. Flowever, detailed analysis showed that these rows arevery likely a single kinety curving around the cell. The oral infraciliature of Remanella isverysimilartothat of Loxodes spp.,i.e.consistsof fourhighlyspecialized and specifically arranged kineties, whose structure is described in detail. Previous inves- tigations could not determine whether or how the nematocyst-like extrusomes of Rema- nella are extruded. The present study shows that they are discharged, thereby assuming a unique, drumstick-like shape because the roundish extrusome capsule remains attached to the despiralized filament. The data emphasize the close relationship between Remanella and Loxodes, earlier proposed by Kahl, and suggest that they emerged from a common ancestor which looked similar to a present day L oxodes. Remanella Kahl, 1933 is a nomen nudum because Kahl, when founding the genus with five new species and one new variety failed to designate any as type. I thus reinstall Remanella nov. gen. for the species assigned to Kahl's invalid taxon and fix R. mubinucleata as type species. Correct names, dates, and authorships are provided for all species described. Introduction ures are highly schematic and hardly show the details required for a proper comparison with Loxodes and Species of the karyorelictid ciliate genus Remanella related genera [1, 3, 10-12]. Using a new fixative, are common ln marlne interstitial and intertidal en- invented by Jean Dragesco, and \Tilbert's protargol vironments 110, 17, 26, 271. Ultrastructural studies technique, I got some excellent preparations showing 137,401on the somatic cortical organization confirmed many previously unrecognized details, which are re- the close relationship withLoxodes, the sole freshwater ported in this study, together with some speculations karyorelictean genus, proposed by Kahl f26l,the foun- on evolution in loxodid ciliates. Furthermore, a de- der of the genus. However, a detailed study of the so- tailed redescription of R. mwltinucleata is provided be- matic and oral ciliary pattern of Remanella was never cause previous studies do not meet the present standard performed, possibly because it was difficult to reveal of ciliate alpha-taxonomy. Last not least, extensive no- with the preparation methods available. Published fig- menclatural changes are required because Kahl, when 09 32 - 47 39 -9 6 -0 03 2-0234$ 3. s 0-0 @ 1.996 by Gustav Fischer Verlag, Stuttgart Infraciliature ol Remanella ' 235 founding the genus with five new species and a new Remanella as new genus to avoid an inflation of names. variety, unfortunately failed to designate any as type. Furthermore, I fix R. multinucleata (Kahl, 1933) nov. Thus, the genus is illegitimate according to the Interna- comb. as type species of the new genus, for which an tional Code of Zoological Nomenclature [25]. improved diagnosis is provided in the discussion. All Remanella species described need to be combined with the new genus. Note that Carey [5], without giv- Material and Methods, Type specimens, ing any reason, quotes Dragesco [10] as combining Terminology author of Kahl's 126l and Dragesco's [8, 9] species although he accepts Kahl [26] as founder of the genus Remanella mwhinucleata occurred in considerable number and Dragesco [10] did not recognize Kahl's [26] mis- in the mesopsammon of the French Atlantic coast at Roscoff. take. Carey's [5] species citation is difficult to under- Samples were collected and treated exactly as described by stand; possibly, he simply did not know that Faur6-Fremiet [14], i.e. the specimens were detached from international rules exist. There are also several other sand grains ml of a 12% MgCl2 solu- the by adding about 5 nomenclatural mistakes. Thus, I decided to revise the tion to about 20 ml sand and sea water. The mixture was then whole genus the benefit sta- gently rotated in a petri dish so that the sand collected in the nomenclature of the for of center and the ciliates could be picked up individually with a bility, the present paper, and future workers. capillary pipette from the clear supernatant. The genus name is derived from Adolf Remane Cells were studied in vivo using a high-power oil immer- (1898-1976), famous German zoologist and one of sion objective and differential interference contrast 119). the discoverers of the marine interstitial fauna. Rema- The infraciliature was revealed by protargol impregnation nella has feminine gender according to article 30b of [19; protocol 2, §Tilbert's method], using a special fixative in- the ICZN [25]. vented by Jean Dragesco (pers. comm.): 5 ml glutaraldehyde L. Remanella brunnea (Kahl, 1933) nov. comb. (pub- (25%),5 mercuric 3 ml aqu- ml saturated, aqueous chloride, lished by Kahl[26,27] without figure; well rede- eous osmium tetroxide (2"/rl, and 1 ml glacial acetic acid are mixed just before use. Specimens are fixed for 15-30 min. scribed in [12]); and washed three times in distilled water. Preparation for 2. Remanella caudata (Dragesco, 1954b) nov. comb. scanning electron microscopy (SEM) was performed as de- (misdated by Carey [5] as 1953 although the jour- scribed by Foissner [19], using the fixative mentioned above. nal clearly states "paru en d6cember 1,954"); Counts and measurements on silvered specimens were per- 3. Remanella dragescoi (Agamaliev, 1966) nov. formed at a magnification of X 1000. In vivo measurements comb.; were conducted at a magnification of X 40- 1000. Although 4. Remanella faurei (Dragesco, 1,954a) nov. comb.l these provide only rough estimates, it is worth giving such 5. Rernanella gigas (Dragesco, 1954a) nov. comb.; in preparations or contract data as specimens usually shrink 6. Remanella granwlosa (Kahl, 1933) nov. comb.; fixation. Standard deviation and coefficient of varia- during (Dragesco, tion were calculated according to statistics textbooks. Draw- 7. Remanella leuii 1960) nov. comb.; ings of live specimens are based on free-hand sketches and 8. Remanella margaritifera (Kahl,1933) nov. comb.; micrographs, those of impregnated cells were made with a 9. Remanella microstoma (Dragesco, 1954b) nov. camera lucida. comb. (misdated by Cprey [5]; see above); No type slides of R. multinucleata mentioned in the '1.0. Remanellaminwta (Dragesco, 1,954a) nov. comb.; literature. Thus, I have deposited two neotype^re slides with spe- 1.L. Remanella mwlticorpusculata (Vacelet, 1961.) nov. cimens prepared as described in the Oberösterreichische comb.; Landesmuseum in Linz (LI), Austria. Relevant specimens 1.2. Remanella mubinucleata (Kahl, 1,933), type of marked a black circle on the cover glass. are by ink genus (see above); Terminology is according to Corliss and Foissner & [6] (Faur6-Fremiet, nov. Rieder with slight modifications, and strictly descriptive 13. Remanella obtwsa 1951) [21], (very belong genus; because ontogenetic data are incomplete and conflicting [28, comb. likely does not to this 30,42). see discussion); All figures are oriented, if not stated otherwise, with the 14. Remanella rugosa (Kahl, 1933) nov. comb.; -1.5. anterior end of the organism directed to top of page. Remanella rugosa var. wnicorpwsculata (Kahl, 1933) nov. comb. (established as "species nova" by Dragesco [12] due to misinterpretation of arti- cle 45g of the ICZN); Results 1.6. Remanella swedmarki (Dragesco, 1954a) nov. comb.; Remanella 1933 is lnualid under Article 13b Kahl, 17. Remanella trichocysla (Dragesco, 1954a) nov. the of ICZN [25] comb. (nom. em.; the original spelling "trichocys- Kahl 126l founded Remanella with five new species tus" cannot be a Greek or latinized noun in appo- and one new variety, none of which, unfortunatelS he sition, as Dragesco possibly had in mind, because designated as type. The genus is thus invalid according the noun is "trichocystis". Thus, the species name to the ICZN. This was overlooked not only by Kahl has to be treated as latinized adjective and coordi- l27l but also by the first reviser of the genus [10] nated with the gender of genus; article 31 of the and later monographers [5, 6]. I thus declare Rema- ICZN); nella Kahl, 1933 to be a nomen nudum, but reinstall 1.8. Remanella unirwgosa (Hartwig,1,973) nov. comb. 236 . W. Foissner .(f ' ,-MA DM MI ,@ ,m '.'a i'.r 10 Figs. 1-10. Remanella mwbinucleata from life (Figs. 1-6, 9, 10) and after protargol (Fig. 7) and wer silver nirrare impreg- nation (Fig. 8). - Fig. l. Rigltlateral view of typical specimen. - Figs. 2, 3. Spiralizöd and undulating specimens. - Figs. a-7. Internal structures. -_Fje.
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