Chlorophyta, Chlorophyceae): Life Cycle and Taxonomy
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Fottea 12(1): 75–81, 2012 75 Ettliella tetraspora (Chlorophyta, Chlorophyceae): life cycle and taxonomy František HINDÁK & Alica HINDÁKOVÁ Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 9, SK–84523 Bratislava, Slovakia; e–mail: [email protected]; [email protected] Abstract: The morphology and mode of reproduction in Ettiella tetraspora HINDÁK 1988, a green coenobial alga described from the Czech Republic and Finland, were studied from the plankton of the water reservoir Förmitzspeicher in NE Bavaria, Germany. The morphology of cells and coenobia of the species in this population was in agreement with published data for the species, but its reproduction was not of the Oocystis – type as it has been declared in the relevant literature. The protoplast of mother cells does not divide simultaneously into 4 autospores as in many coccal green algae, but always in two autospores. The first division of the mother cell is perpendicular to the longitudinal axis of the cell and two walled daughter cells arise. Subsequently, the newly formed daughter cells divide again, but this time perpendicular to the new formed cell cross wall. Ultimately, in a new coenobium all four daughter cells are arranged in parallel. Daughter cells remain inside a slightly enlarged mother cell wall (well visible only near the cross wall) and are released by its gelatinisation. Such subsequent production of four autospores from the mother cell represents a special type of multiplication in the green coccal algae known also in some other species [e.g. Tetrastrum komarekii HINDÁK 1977, Willea irregularis (WILLE ) SC H MIDLE 1900, Makinoella tosaensis OKADA 1949]. Key words: autospores subsequent division, coenobial green algae, taxonomy, Oocystaceae, Scenedesmaceae, Germany Introduction autosporulation and therefore he assigned the alga to the family Oocystaceae. When HINDÁK During a workshop devoted to the determination (1988) established this alga as a new genus and of planktic cyanophytes and chlorophytes of species, Ettliella tetraspora, he followed this lakes and reservoirs organised by the Bavarian classification, although the release of autospores Regional Office for the Environment (Bayerisches and remnants of mother cell walls were not Landesamt für Umwelt) in Wielenbach, Germany, observed in detail. It was noticed, however, that in July 2010, samples from several local water the regular Tetradesmus – like clustering of cells bodies were analysed. In the phytoplankton sample of Ettliella is reminiscent of some representatives collected from the water reservoir Förmitzspeicher of the family Coelastraceae or Scenedesmaceae. a rare coccal green alga, Ettliella tetraspora, was In one phytoplankton sample from Bavaria, observed. coenobia of Ettliella tetraspora occurred quite The monotypic genus Ettliella HINDÁK abundantly, thereby offering an opportunity to 1988 including E. tetraspora HINDÁK 1988 was study the process of autosporulation in all its described on the basis of two observations. ETTL stages. These observations led to the conclusion (1965) was the first to find the alga in a fishpond that the mother cell wall does not expand before near Svitavy in Moravia, the Czech Republic, the liberation of autospores as in the Oocystaceae. but designated it as Tetrachlorella cf. alternans Probably ETTL (1965) did not depict expanded mother cell walls around daughter coenobia, but KOR sh IKOV (?nov. var.) (cf. also KOMÁREK & mucilaginous envelopes newly formed around FOTT 1983; HINDÁK 1988). Later, HINDÁK (1988) autocoenobia. observed it in a lake near Jyväskylä, Central Finland. The morphology of cells and colonies and the formation of autospores were similar Material and Methods in material from both localities. Ettl was of the opinion that the mother cell wall expands during Plankton samples were collected from the water 76 HINDÁK & HINDÁKOVÁ : Ettliella tetraspora (Chlorophyta, Chlorophyceae) reservoir Förmitzspeicher (Förmitzsee) near or cells solitary before reproduction, embedded Schwarzenbach a.d. Saale, NE Bavaria, Germany, pH by wide homogeneous hyaline mucilaginous 7.9, July 5, 2010, by Mrs C. Hauenschild, Hof, using a envelopes. Cells in coenobia clustered into tetrads plankton net with 10 µm mesh size (Fig. 1). as in Tetradesmus G.M. SMIT H , i.e. positioned (but Cyanophytes and algae were determined not joined) in parallel next to each other forming according to KOR sh IKOV (1953), HINDÁK (1977, 1980, 1984, 1988, 1990, 2008), KOMÁREK & FOTT a square when viewed from above, with a central (1983), KRAMMER & LANGE –BERTALOT (1986, 1988, opening (Figs 2–4). Cells broadly ovoid or short 1991a, b), KOMÁREK & ANAGNO S TIDI S (1998, 2005), cylindrical, before division slightly asymmetrical Ko m á r e K & Za p o m n ě l o v á (2007), LENZEN W EGER to kidney–shaped, 8–12(18) × 6–8(10) µm, ends (1999) and RAJANIEMI et al. (2005). Identifications broadly oval. Cell walls thick, smooth, without were performed on living material. Light microscope incrustations or conspicuous polar thickenings. observations were carried out with a Leitz Diaplan Chloroplasts single, bright green, parietal, microscope, and photomicrographs were taken with transversely pot–shaped, with a large central a Wild Photoautomat MPS45. Preserved material in formaldehyde is stored at the Institute of Botany SAS, starch–associated pyrenoid, and in cells in tetrads Bratislava. Attempts to cultivate this organism under oriented toward the external side. Reproduction by laboratory conditions were unsuccessful. autocoenobia with 4 autospores. Coenobia before autosporulation mostly disintegrating into solitary cells, losing their tetrahedrical arrangement of cells Results (Figs 5–8). When autospores are formed, colonies arise temporarily with tetrads of young cells, so Ettliella tetraspora HINDÁK (Figs 2–9) that composite colonies exhibit cell tetrads of a Coenobia free floating, solitary, usually 4–8(16)– similar clustering (Figs 7–9). celled, ovoid to spherical, 40–60 µm in diameter, Although the product of autosporulation is one 4 Fig. 1. Locality where Ettiella tetraspora was collected: the water reservoir Förmitzspeicher in NE Bavaria, Germany (photo C. Hauenschild, summer 2010). Table 1. List of cyanophytes and algae found in the phytoplankton of the water reservoir Förmitzspeicher, Germany. Cyanophyceae Chlorophyceae Aphanocapsa delicatissima W. et G.S. WE S T Carteria globosa KOR sh IKOV ex PA S C H ER Aphanocapsa planctonica (G.M. SMIT H ) KOMÁREK et Chlamydomonas passiva SKUJA ANAGNO S TIDI S Eudorina elegans EH REN B ERG Aphanothece clathrata W. et G.S. WE S T Asterococcus limneticus G.M. SMIT H Aphanothece nidulans RIC H TER in WITTROCK et NORD S TEDT Ankistrodesmus falcatus (CORDA ) RAL fs Chroococcus limneticus LEMMERMANN Ankistrodesmus spiralis (W.B. TURNER ) LEMMERMANN Chroococcus minutus (KÜTZING ) NÄGELI Botryococcus braunii KÜTZING Eucapsis minor (SKUJA ) ELENKIN Closteriopsis acicularis (G.M. SMIT H ) J.H. BELC H ER et Merismopedia glauca (EH REN B ERG ) KÜTZING SW ALE Merismopedia minima BECK Coelastrum pseudomicroporum KOR sh IKOV Microcystis aeruginosa (KÜTZING ) KÜTZING Coenochloris polycoccus (KOR sh IKOV ) HINDÁK Microcystis flos–aquae (WITTROCK ) KIRC H NER Coenocystis planctonica KOR sh IKOV Snowella litoralis (HÄYRÉN ) KOMÁREK et HINDÁK Crucigeniella apiculata (LEMMERMANN ) KOMÁREK Synechococcus endophyticus (W. et G.S. WE S T ) JOO S TEN Desmodesmus abundans (KIRC H NER ) E.H. HEGE W ALD Woronichinia naegeliana (UNGER ) ELENKIN Desmodesmus armatus (CH ODAT ) E.H. HEGE W ALD Oscillatoria limosa AGARD H ex GOMONT Desmodesmus brasiliensis (BO H LIN ) E.H. HEGE W ALD Pseudanabaena catenata LAUTER B ORN Desmodesmus communis (E.H. HEGE W ALD ) E.H. HEGE W ALD Pseudanabaena mucicola (NAUMANN et HU B ER –PE S TALOZZI ) Desmodesmus dispar (BRÉ B I ss ON ) E.H. HEGE W ALD SC hw A B E Desmodesmus maximus (W. et G.S. WE S T ) E.H. HEGE W ALD Anabaena mucosa KOMÁRKOVÁ –LEGNEROVÁ et ELORANTA Desmodesmus serratus (CORDA ) AN, FRIEDL et E.H. Anabaena planctonica BRUNNT H ALER HEGE W ALD Desmodesmus subspicatus (CH ODAT ) E.H. HEGE W ALD et A. Bacillariophyceae SC H MIDT Aulacoseira ambigua (GRUNO W ) SIMON S EN Dictyosphaerium tetrachotomum PRINTZ Cyclotella balatonis PANTOCZEK Ettliella tetraspora HINDÁK Discostella pseudostelligera (HU S TEDT ) HOUK et KLEE Komarekia appendiculata (CH ODAT ) FOTT Melosira varians C. AGARD H Monoraphidium griffithii(B ERKELEY ) KOMÁRKOVÁ – Stephanodiscus medius HÅKAN ss ON LEGNEROVÁ Achnanthes laevis OE S TRU P Oocystella borgei (J. SNO W ) HINDÁK Achnanthes lanceolata ssp. frequentissima LANGE –BERTALOT Oocystella lacustris (CH ODAT ) HINDÁK Achnanthes minutissima KÜTZING Oocystella marssonii (LEMMERMANN ) HINDÁK Achnanthes cf. petersenii HU S TEDT Pediastrum biradiatum MEYEN Achnanthes cf. saccula J. R. CARTER Pediastrum boryanum (TUR P IN ) MENEG H INI Caloneis silicula (EH REN B ERG ) CLEVE Pediastrum duplex MEYEN Cymbella gracilis (EH REN B ERG ) KÜTZING Pediastrum tetras (EH REN B ERG ) RAL fs Cymbella silesiaca BLEI S C H Pseudodictyosphaerium minusculum HINDÁK Eunotia bilunaris (EH REN B ERG ) MILL S Pseudodidymocystis inconspicua (KOR sh IKOV ) HINDÁK Eunotia praerupta var. bidens (EH REN B ERG ) GRUNO W Pseudokirchneriella contorta (SC H MIDLE ) HINDÁK Fragilaria crotonensis KITTON Pseudokirchneriella irregularis (G.M. SMIT H ) HINDÁK Fragilaria tenera (W. SMIT H ) LANGE –BERTALOT Scenedesmus acuminatus (LAGER H EIM ) CH ODAT Fragilaria ulna var. acus (KÜTZING ) LANGE –BERTALOT Scenedesmus arcuatus