Protistology Re Description of Vannella Mira Schaeffer 1926
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Protistology 2 (3), 178–184 (2002) Protistology Redescription of Vannella mira Schaeffer 1926 (Gymnamoebia, Vannellidae), an often mentioned but poorly known amoebae species Alexey V. Smirnov Marine Biological Laboratory University of Copenhagen, Helsingor, Denmark and Department of Invertebrate Zoology, Faculty of Biology and Soil Sci., St.Petersburg State University Summary Amoebae species Vannella mira Schaeffer 1926 was relatively well described and has often been discovered in various regional faunas. However, there is still no clear understanding as to what we really mean by the name “Vannella mira”. A.A. Schaeffer described this species from marine environment in 1926 (as Flabellula mira), but further researchers applied this name for several brackish and freshwater isolates. F.C. Page redescribed “V. mira” in 1968 from freshwater and in 1976 listed it among freshwater amoebae. Perhaps, it was a misidentification. In 1988 he revised his opinion stating that “long experience has failed to reveal any other gymnamoeba occurring in both fresh and salt water” and applied a specific name Vannella cirifera Frenzel 1892 to the isolate that he previously called V. mira. However, various authors used the name V. mira for different marine and freshwater isolates even after the renaming. The present paper describes a marine amoeba isolated from cyanobacterial mats in France (Camarque) that corresponds in full to Schaeffer’s initial description of “Flabellula mira”. New type material is established for the species V. mira Schaeffer 1926; it remains a type species of the genus Vannella. Key words: amoebae, Vannella, Lobosea, Gymnamoebia, morphology, systematics, Vannellidae Introduction shape always present in endoplasm. No vacuoles. No crystals. Nucleus single. Chromatin mass spherical, An amoebae species “Flabellula mira” was esta about 2 µm in diameter. Nuclear membrane 5 µm in blished by A.A. Schaeffer (1926) with the following diameter. Food vacuoles in small number 2 to 5 µm in diagnosis: “Length in locomotion 25 µm; breadth 15 diameter. No uroid. Food, bacteria. Habitat, seawater, to 25 µm; shape oval to fanshaped in outline; when among bluegreen algae, and elsewhere; common. not in locomotion, shape extremely variable with Tortugas and Key West, Florida.” The above diagnosis numerous pseudopods of a large variety of shapes. Rate is accomplished with a rather detailed description of locomotion rapid, 75 µm per minute. Endoplasm (Schaeffer, 1926, p. 48–49) and eleven line drawings, clear in anterior portion, finely granular in posterior of which six illustrates the floating form, three – part. A small number of refractory grains of irregular transformation from floating to locomotive form, one © 2002 by Russia, Protistology Protistology · 179 shows a sequence of the body shape in locomotion and water (e.g. Page, 1968)”. In his first key to freshwater the last is a summarising coloured drawing of the and soil amoebae (Page, 1976), he illustrated the strain locomotive form (op. cit., fig. 12 and plate 4, fig.1). that he called V. mira with photographs (p. 101) and No type material was established (it was not a rule at give the dimensions according to his data (Page, 1968). that time). Some amoebae in his photographs bore few faecal It seems that this relatively welldescribed (for the pellets. Page (1979) gave a rediagnosis of the genus facilities of that time) species should not produce much Vannella with the same type species – Vannella mira difficulties to systematists, given that up to now species Schaeffer 1926. Page and Blakey (1979) illustrated the distinction within the genus Vannella (to which cell coat of freshwater V. mira. Schaeffer’s species belongs in the modern system of At the same time Bovee and Sawyer (1979) amoebae) is based on the same features that are described V. mira among marine amoebae (distribution: mentioned in Schaeffer’s description. Hopkins (1938) Atlantic coast, US, Florida to Maine, Pacific coast, reported that the marine amoeba which he identified Southern California; coral reefs, Madagascar). The as “Flabellula mira” could be cultured in 5 ppt seawater, distribution mentioned far exceeds that noted by though without forming contractile vacuole. Bovee in Schaeffer (1926) and suggests that Bovee and Sawyer his PhD thesis (1950, cited by Bovee and Sawyer, 1979) isolated this species themselves from various marine mentioned this species from marine habitats, and three locations. According to their description, this species years later (Bovee, 1953a) isolated a strain that he called in locomotion is 15 to 25 µm long by 15 to 25 µm wide, “Flabellula mira” Schaeffer 1926 from a freshwater nucleus 4 to 6 µm in diameter with central endosome 2 pond and noted the presence of contractile vacuole in to 3 µm in diameter. Contractile vacuole absent in the cytoplasm of this strain. He argued that his strain is seawater, 3 to 6 µm in diameter in brackish or fresh indistinguishable from Schaeffer’s one and believed that water. These measurements of the contractile vacuole the contractile vacuole is developed under freshwater differ from that given by Page (1968), namely 4.7–7.6 conditions, but is absent in marine isolate. µm. As a source of other data they refer to Schaeffer Actually, this mentioning introduced a misun (1926) and Page (1971). Bovee (1985) gives the same derstanding. Given that “F. mira” may be found in both description and measurements of V. mira, but refers to marine and freshwater, Page (1968), basing on the Schaeffer (1926) only. observations of two freshwater isolates, redefined the Later Page (1988) stated that “long experience has species “F. mira”. He formulated a new, rather detailed failed to reveal any other gymnamoeba occurring in diagnosis of this species (not an emendation of both fresh and salt water” and suggested a name Schaeffer’s one) (Page, 1968, p. 13). According to Vannella cirifera Frenzel 1892 as a substitute for the Page’s definition, “F. mira” has the “greatest dimension strain that he called V. mira in 1968. However, no other of the locomotive form approximately 23–55 µm; motivation except that “Frenzel’s name has priority, greatest dimension usually breadth. Diameter of the and his figures (Frenzel, 1892) greatly resembles those nucleus 4.8–8.3 µm; diameter of nucleolus 2.1–4.1 in this publication” was given (1988, p. 74). One of the µm” (op. cit., p. 14). His strain could grow on the agar photographs of “V. cirifera” given by Page (1988) is the made with up to 10 ppt seawater and one strain, same as for “V. mira” in Page (1976); two other are abnormal in appearance – up to 25 ppt. As a distinctive different. Page (1991, p. 87) gives three new photo feature of his strain he noted the presence of numerous graphs of “V. cirifera”. Page (1988) illustrates the cell faecal pellets, visible on agar surface. No type material coat of “V. cirifera”, stating that, in contrast with V. was established. simplex, this species does now have simple filaments Bovee (1965) redefined Flabellula and established among glycostyles. the genus Vannella for the species that did not fit to the Several authors (Davis et al., 1978; Fernandez et former genus. Former “F. mira” was transferred to the al., 1989; Munson, 1992; FernandezLeborans and newly established taxon as a type species and named Novillo, 1993; Rogerson and Gwaltney, 2000) list V. Vannella mira Schaeffer 1926. Page (1969) recognised mira in the faunas of marine habitats. Michel and Just these changes and wrote that the species “F. mira” (1984) mentioned “V. mira” among the species isolated which he had redescribed in 1968, should therefore from the cooling and rinsing water in dental treatment be identified as Vannella mira (Schaeffer, 1926), thus units – i.e. from freshwater. Ariza et al. (1994) stating identity of his and Schaeffer’s species. Page illustrated two freshwater strains called V. mira. (1971) mentioned this species also as Vannella mira and Growing bulk of knowledge about amoebae noted that “while V. mira (Schaeffer, 1926) … were indicated that species distinction among vannellids is originally described from salt water, amoebae indis very difficult (Page, 1988; Smirnov, 2001). However, tinguishable in form from V. mira was found in fresh there are at least 14 morphologically different species 180 · Alexey V. Smirnov of the genus Vannella (Page, 1991; 1983; Smirnov and (average 26.5 µm). Length/breadth ratio 0.5–1.8 Fenchel, 1996; Smirnov, 2001), corresponding to the (average 1.08). In a few specimens there were small modern (Page, 1979) definition of the genus. This dorsolateral ridges or small lobes at the frontal number does not include V. mira or V. cirifera, because hyaloplasm (Figs 2; 5; 14, D), but most of cell in culture of the difficulties in the species definition mentioned had smooth surface. Some amoebae in culture bore above. The impression is that various authors applied several rounded faecal pellets. Developed floating form the name V. mira to different species. To stop the misuse (Figs 6–9; 14, F–L) always had long, narrowconical of the name of the type species of the genus Vannella, a pseudopodia with rounded tips; their number varied marine amoebae strain that corresponds in full to the from three to eleven. Several very characteristic types original Schaeffer’s description of V. mira is illustrated of the floating form (Fig. 14, F–L) were observed: and described in the present paper; neotype of V. mira 1. Floating form with 3–11 radiating pseudopodia; is established. some pseudopodia may furcate. During the develop ment of the floating form pseudopodia elongate, but Material and Methods sometimes the development may “freeze” and floating amoeba for a long time has short, curved, relatively Vannella mira was isolated from cyanobacterial thick pseudopodia (e.g. Fig. 14, F). mats collected in Camarque, (France, Mediterranean 2. Floating form with all cytoplasm in the conical coast).