Flagellate green algae from four water bodies in the state of Rio de Janeiro, Southeast Brazil

Algas verdes flageladas de quarto corpos d'água no Estado do Rio de Janeiro, Sudeste do Brasil

Flagellate green algae from four water bodies in the state of Rio de Janeiro, Southeast Brazil

Algas verdes flageladas de quarto corpos d'água no Estado do Rio de Janeiro, Sudeste do Brasil

Mariângela Menezes^I,*; Carlos Eduardo de Mattos Bicudo^II

^IUniversidade Federal do Rio de Janeiro, Museu Nacional, Laboratório de Ficologia, Quinta da Boa Vista, São Cristóvão, 20940-040 Rio de Janeiro, RJ, Brasil

^IIInstituto de Botânica, Caixa Postal 3005, 01061-970 São Paulo, SP, Brasil

ABSTRACT

Floristic survey of flagellate green algae (Chlorophyceae and Prasinophyceae) from four water bodies in the state of Rio de Janeiro, southeast Brazil. Samples were collected bimonthly from September 1988 to August 1991. Thirty-five taxa of which 31 in Chlorophyceae (24 Chlamydomonadales and seven Volvocales) and four in Prasinophyceae (Polyblepharidales) were identified. Twenty-three taxa are new records for Brazil and 31 are registered for the first time for Rio de Janeiro State. On basis of cell shape, location of the nucleus and degree of longitudinal striations on the chloroplast surface a new combination, Vitreochlamys lefevrei (Bourr.) Menezes & C. Bicudo, is proposed. Integrating studies among morphological analysis on natural/cultured material and characterizing of reproduction/life cycle as well the correlating between the occurrence of the species and environmental conditions are needed in order to better knowledge the degree of phenotypic plasticity within the members of the flagellate green algae in the country.

Key words: Chlorophyceae, continental waters, flora, Prasinophyceae

RESUMO

Inventário florístico das algas verdes flageladas (Chlorophyceae e Prasinophyceae) de quatro corpos d'água do sudeste do estado do Rio de Janeiro, Brasil. Com base em amostras coletadas bimensalmente, entre setembro de 1988 e agosto de 1999, foram identificados 35 táxons distribuídos em 31 Chlorophyceae (24 Chlamydomonadales e sete Volvocales) e quatro Prasinophyceae (Polyblepharidales). Do total de táxons, 33 constituem novos registros para o Brasil e 31 para o estado do Rio de Janeiro. Com base na forma da célula, na localização do núcleo e no grau de estriação longitudinal da superfície do cloroplasto propõe-se Vitreochlamys lefevrei (Bourr.) Menezes & C.Bicudo como nova combinação. Estudos integrados entre análise morfológica de material natural/cultura e caracterizacão da reprodução/ciclo de vida bem como correlação entre a ocorrência ds espécies e condições ambientais são necessários visando melhor conhecimento da plasticidade fenotípica dentro de representantes de algas verdes flageladas no País.

Palavras-chave: águas continentais, Chlorophyceae, flora, Prasinophyceae

Introduction

Few floristic and taxonomic investigations have been carried out on the pigmented flagellate algae in Brazilian freshwater systems. Information concerning these algae is dispersed in general floristic' surveys and plankton studied ecology. Many of these publications, however, include neither illustration nor description of the material studied. To complicate matters, many of the latter publications just refer to the flagellate genus name or, even worse, simply mention them as 'phytoflagellates'.

Strict taxonomic studies of Brazilian phytoflagellates started effectively at the beginning of the last century with the works of Cunha (1913, 1914), who recorded altogether 90 pigmented taxa including descriptions of nine new species of euglenoid flagellates.

Somewhat recently, other phytoflagellate groups have become the object of floristic and taxonomic studies. Knowledge of these organisms in Brazil, however, is still very scarce. Despite of being just a few, such studies have shown there to be a very rich and interesting Brazilian flagellate algal flora, especially in the tropical part of the country. Some of the latter publications include descriptions of new taxa (Castro et al. 1991, Conforti 1993a, b, 1994, Menezes 1996, Menezes & Huszar 1997).

Seventy-six papers have already published on the pigmented flagellate algal flora of Brazil, of which 70% have focused on the Euglenophyceae. Regarding the other flagellate classes, publications can be summarized as follow: nine publications on Chrysophyceae and Synurophyceae, five on the Dinophyceae, three on the Raphidophyceae and Prasinophyceae, and three on the Chlorophyceae and the Cryptophyceae.

The present study is part of the floristic survey of the pigmented flagellate algae of Brazil. It includes the Chlorophyceae and Prasinophyceae from four selected water bodies located in the state of Rio de Janeiro. The paper is aimed at increasing knowledge of the flagellate algal flora from tropical Brazil, and includes critical taxonomic remarks on several taxa, information about species composition of each system, as well as notes on the ecology and geographical distribution of each group.

Material and methods

All water bodies investigated are located in the southern part of the municipality of Rio de Janeiro, state of Rio de Janeiro, southeast Brazil (figure 1). The area includes a coastal plain region and some escarpments from the Precambrian Complex (Serra do Mar). Altitude is around 800 m above the sea level, and relief ranges from plane to slightly waved.

Local climate is tropical with warm and rainy summers and dry winters. The mean annual temperature varies between 18 and 24 ºC and the annual rainfall from 1,200 to 2,000 mm. Dominant soils are hydromorphic latosols, which are associated with litosols and podzols. The major part of the coastal plain consists of lagoons, the Jacarepaguá-Marapendi lagoon complex, that are interconnected by sandy strands. Due to its inclination, the lowland is crossed by diverging rivers which are broken close to the sea by penetration of tides raising pools, swamps, and mangroves. In this wet lowland dominates humid vegetation, dunes, and sandy coastal plains ("restingas"). Escarpment areas are found continuously throughout the coastal strand, where remnants of the Atlantic rainforest can be seen (Golfari & Moosmayer 1980, RADAMBRASIL 1983).

The four water bodies investigated can be described as follows:

1. Camorim Reservoir: located on the Pedra Grande massif, at approximately 430 m altitude (23º03'S and 43º27'W). The reservoir has total surface area of about 22 km², maximum depth 3 m, and alkaline, mesotrophic waters. This reservoir is the only source of water supply for the District of Jacarepaguá.

2. Unnamed oxbow pond: located in the Jacarepaguá lowland in front of the km 42 mark at the roadside of Avenida das Américas (22º50'00"S and 43º38'55"W). It has total surface area of approximately 0.1 km², maximum depth between 1 and 2 m and alkaline, eutropic waters. Extensive growth of macrophytes and mainly of Typha domingensis Person can be observed along its entire shoreline and the water surface is mostly covered by Salvinia sp.

3. Lagoinha: a coastal lagoon located in the Biological Reserve of Jacarepaguá (23º02'23"S and 43º28'97"W). It has total surface area of approximately 0.7 km², and mean depth of about 2.4 m. The bottom is covered with organic sediment mainly of plant origin, and the water is slightly acidic, oligohaline, eutrophic.Along the entire shoreline there is extensive macrophyte vegetation growth represented mostly by Typha domingensis Person, and its water surface is mostly covered with Pistia stratiotes Linnaeus.

4. Lake at the Bosque da Barra: a natural lake located in a preservation "restinga" area at Barra da Tijuca (22º59'02"S and 43º23'30"W). It has total surface area of around 0.2 km², mean depth of about 2 m, and oligotrophic, brown, acidic waters due to the presence of humic substances. Specimens of Nymphaea sp. and Utricularia sp. may sporadically occur along its shore.

Samples were collected from September 1988 to August 1991, every two months, from the shore by filling bottles directly with water and/or sediments using a 25 µm mesh size plankton net as well as by gathering by hand macroscopic filamentous algae and cutting off submerged parts of macrophytes. In every case, material was kept in bottles and immediately examined alive under the light microscope. Three or four days after collection material was invariably fixed and preserved with 3%-5% neutralized formalin solution.

All taxa were described in detail. Taxonomic comments, whenever necessary, are given with illustrations and information on the geographic distribution of each taxon identified in Brazil.

The circumscription of classes is according to Round (1981). The systematic arrangement of orders and families essentially follows that of Ettl (1983). Although the groups names proposed by Ettl (1983) for the genus Chlamydomonas are not to be formed according International Code of Botanical Nomenclature these names have been adopted in this work aiming a more didactic presentation. The concept the species applied was the morphospecies from natural samples. When possible, the delimitation of taxa was complemented by works based on combined analysis of morphology by light microscopy and molecular biology, among them of Nozaki et al. (1998), Nozaki (2003), Nozaki & Kuroiwa ( 1992), Nozaki & Itoh (1994), Pröschold et al. (2001). For more information on the techniques used see Menezes (1994).

Results

The total flagellate green algal flora of the four investigated water bodies was 35 species of which 31 belong to the Chlorophyceae (24 Chlamydomonadales and seven Volvocales) and four to the Prasinophyceae (Polyblepharidales). Twenty-three taxa are new records for Brazil and 31 are registered for the first time for Rio de Janeiro State.

CHLOROPHYCEAE

Chlamydomonadales

Chlamydomonadaceae

Carteria Diesing

Carteria lohammari

Skuja, NovaActa R. Soc. Scient. Upsal. ser. IV, 16(3): 115, pl. 15, fig. 1-4. 1956.

Figures 2-3

 

Cell 21-29 µm diam., spherical, without anterior papilla. Chloroplast cup-shaped, 9-12 radial incisions, unequal rays, pyrenoid 1, spherical or elliptical, slightly displaced towards the posterior cell end. Nucleus central, located above the pyrenoid. Stigma elliptical, at the upper part of cell. Contractile vacuoles 2, apical. Flagella 2-2.5 times longer than the cell diameter.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at the Bosque da Barra, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173642); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n.(R173643); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173644); Camorim Reservoir, 22-I1990, M. Menezes & A.M. Werneck s.n. (R173659).

Carteria lohammari belongs to a species-complex that is characteristic by having cells somewhat circular in outline, with a stelloid, cup-shaped chloroplast with one pyrenoid. The main diagnostic features to sort out species in this complex are the cell free-swimming or epibiont habit and the number of chloroplast incisions. Sometimes, secondary features like cell shape and size, presence of an anterior papilla, pattern of wall decoration, and distance between wall and protoplast can also be used.

According to Ettl (1983), C. lohammari and C. conochlii Skuja belong to the group of epibiontic species. However, C. lohammari has 12-16 chloroplast incisions and a rounded anterior cell pole, whereas C. conochlii has eight chloroplast incisions and a truncate anterior cell pole.

Twenty individual free-swimming cells were studied. The fact of consistently having spherical cells with smooth wall and never a papilla at the anterior cell pole, led us to the conclusion that all Brazilian populations examined could be C. abiscoensis Skuja sensu Peterfi (1968), young cells of C. disclusa Skuja, or with free-swimming cells of C. conochlii and C. lohammari. Material presently examined showed cell dimensions and number of chloroplast incisions that did not fit exactly any of the above species, but the one most closely matching was C. lohammari. So, despite having never found individual cells attached to any substrate, we believe the material from the state of Rio de Janeiro most closely corresponds to the latter species.

Recorded only for Sweden up to now. New to Brazil. Found in plankton samples from Bosque da Barra Lake and Camorim Reservoir.

Chlamydomonas Ehrenberg

Euchlamydomonas

Chlamydomonasdebaryana

Gorozh. var.

micropapillata

Gerloff, Arch. Protistenk. 94: 455, fig. 17. 1940.

Figures 4-5

 

Cell (17-)18(-20) µm long, (12-)13(-15) µm broad, elliptical-ovoid, with a hemi-spherical, 2-3.2 µm long anterior papilla. Chloroplast cupshaped, smooth, basal thickening filling up nearly 1/2 of cell, pyrenoid 1, spherical, basal. Nucleus anterior, approximately in the middle of cell. Stigma rounded, in the upper part of cell, ca. 2.3 µm diam. Contractile vacuoles 2, apical. Flagella about the length of the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 29-X-1988, M. Menezes s.n. (R172157); the same, 29-X-1988, M. Menezes s.n. (R172158); the same, 10-I-1989, M. Menezes s.n. (R173417); Lagoinha, 10-I-1989, M. Menezes s.n. (R173409); the same, 27-III-1989, M. Menezes & I.C.A. Dias s.n. (R173472).

Reported from Europe. In Brazil this species has previously been recorded for the state of Rio de Janeiro (Menezes 1999). It was found in plankton and metaphyton samples, respectively from Camorim Reservoir and Lagoinha.

Chlamydomonas monoica

Strehlow var.

monoica,

Zeitschr. f. Bot. 21: 627, fig. 2. 1929.

Figures 6-8

Cell (14-)18(-20) µm long, (9-)10(-14) µm broad, elliptical, ovoid or elliptical-ovoid. Cell hyaline, wall with a short anterior papilla, ca. 1.5 µm long. Chloroplast cup-shaped, smooth, basal thickening occupying usually nearly 1/2, sometimes up to 2/3 of the cell, pyrenoid 1, spherical, basal. Nucleus anterior, central. Stigma elliptical, in the upper part of cell, ca. 2.3 µm long. Two apical contractile vacuoles. Flagella about the length of the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 10-I-1989, M. Menezes s.n. (R173417); the same, 19-V-1990, A.M. Werneck s.n. (R173695).

Reported from Europe. New to Brazil. This species has found in both plankton and metaphyton samples from Camorim Reservoir.

Chlamydomonas pertyi

Gorozh., Bull. Soc. Imp. Nat. Moscou, n.s. 5: 1, pl. 1, fig. 13-22. 1891.

Figures 9-13

Cells 16-24 µm long, 12-19 µm broad, broadly ovoid or rounded. Cell hyaline, wall with a hemispherical anterior papilla, sometimes cells enclosed in a mucilaginous sheath. Chloroplast cupshaped, smooth, sometimes with radial, peripheric incisions, basal thickening occupying nearly 1/2 of the cell, pyrenoid 1, spherical or elliptical, basal. Stigma elliptical, in the upper cell part, ca. 2 µm long. Contractile vacuoles 7-10, at the anterior part of cell. Flagella 1.5-2 times the cell length. Gloeocystis-like stage with thick mucilaginous sheath surrounding each cell, all cells inside of a common yellowish mucilaginous envelope, 23-25 µm diameter. Isogametes, zygote globose, with a reddish verrucose wall, 15-19 µm in diameter.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lagoinha, 29-IX-1988, M. Menezes s.n. (R172914); the same, 29-IX-1988, M. Menezes s.n. (R172915); unammed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160); the same, 4-IX-1989, M. Menezes s.n. (R173574); the same, 16X-1989, M. Menezes s.n. (R173610); Lake at Bosque da Barra, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173642); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173643); the same, 27-XII1989, M. Menezes & I.S. Nogueira s.n. (R173644);Camorim Reservoir, 19-V-1990, A.M. Werneck s.n. (R173695); the same, 19-VII-1990, M. Menezes s.n. (R173750).

World-wide distributed, reported from Africa, Asia, Europe, and North America. New to Brazil. Chlamydomonas pertyi was a common species in the area studied, being found in all collecting sites both in plankton and metaphyton samples.

Chlamydomonas tetravacuolata

Ettl, Bot. Tidsskr. 74(4): 212, pl. 15, fig. 1-4. 1980

Figures 14-15

 

Cells 15-18 µm long, 12-14 µm broad, ovoid or elliptical. Cell wall hyaline, with a conical, truncated anterior papilla. Chloroplast cup-shaped, smooth, basal thickening occupying nearly l/4 of cell, pyrenoid 1, elliptical, sometimes rectangular, basal. Nucleus in the middle of the cell, slightly excentric. Stigma elliptical, at the upper part of cell, ca. 2.8 µm long. Contractile vacuoles 4, apical. Flagella ca. 1.5 times the cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unnamed oxbow pond, 4-IX-1989, M. Menezes s.n. (R173574).

This species was only recorded from Denmark. New to Brazil. Present in one plankton sample from an oxbow pond.

Material studied differs from the original description in Ettl (1980) by having a much smaller basal thickening of the chloroplast and sometimes a rectangular pyrenoid. Chlamydomonas tetravacuolashows basal thickening of the chloroplast occupying 1/2 of cell and elliptical pyrenoid.

Group Pleiochloris

Chlamydomonas pseudotarda

Bourr., Hydrobiologia 3: 262, pl. 3, fig. 54. 1951.

Figures 16-21

Cells (25-)28-30 µm long, (11-)13-17 µm broad, oblong, oblong-elliptical or elliptical. Cell wall hyaline, with a hemispherical anterior papilla 1-3 µm long. Chloroplast cup-shaped, smooth, basal thickening occupying 1/3 of cell, pyrenoids 2-13, spherical, irregularly distributed within the edge of chloroplast. Stigma linear, large, in the upper part of cell, ca. 6 µm long. Nucleus in the middle of cell. Contractile vacuoles 2, apical. Flagella ca. 0.5 times of cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 19-V-1990, A.M. Werneck s.n. (R173694); the same, Camorim Reservoir, 19-V-1990, A.M. Werneck s.n. (R173695);the same, Camorim Reservoir, 19-VII-1990, M. Menezes s.n. (R173750).

Chlamydomonas pseudotarda, C. ambiguaGerloff var. minor L.S. Peterfi, and C. saccus Pascher morphologically closely resemble each other. Their circumscriptions are very poorly delimited and based on cell shape and size, stigma morphology, and number of pyrenoids.

According to their original descriptions, C. ambigua var. minor is distinct from C. pseudotarda and C. saccus mainly in having smaller cell dimensions (1216 × 9-11 µm) and lesser number of pyrenoids (1-2). Also, in its sometimes ovoid cells. Chlamydomonas saccus has obovoid-oblong cell and the stigma is greater than in the other two species. Chlamydomonas pseudotarda differs from C. saccus only by having elliptical cell. Regarding cell dimensions and pyrenoid number there are overlap between these two species, i.e. cells 22-30 × 13-20 µm and 4 pyrenoids in C. saccus; cells 26 × 13 µm and 4-6 pyrenoids in C. pseudotarda.

Ettl (1976) distinguished C. pseudotarda from C. saccus and C. ambigua var. minor on the basis of pyrenoid size. Chlamydomonas pseudotarda has pyrenoids with different sizes while C. saccus and C. ambigua var. minor have pyrenoids of the same size.

Populations presently studied showed variable cell shape and pyrenoid number and size, which could relate to all three taxa above. As similar situation was recorded by Jakubiec (1988) in material from Poland, i.e. cells oblong, oblong-elliptical to elliptical, with 2-13 pyrenoids. The specimens with oblongelliptical to elliptical cells had 4-5 differently-sized pyrenoids (figures 18-20), except for those with 2 pyrenoids that had equally-sized pyrenoids (figure 21). Individual oblong-celled organisms had differentlysized pyrenoids that varied in number from 7 to 13 (figure 16).

We decided to attribute the present material from the state of Rio de Janeiro to C. pseudotarda due to the high frequency of cells with more than four pyrenoids, and because we never found specimens typically of C. saccus.

Reported from Europe (Austria, France, Spain). New to Brazil. This species was only found in the Camorim Reservoir plankton.

Group Chlamydella

Chlamydomonas typhlos

Gerloff, Arch. Protistenk. 94(3): 464, fig. 33. 1940.

Figures 22-25

Cells 13-17 µm long, 7-11 µm broad, elliptical. Cell wall hyaline, with an anterior conical papilla 1.5-2 µm long. Chloroplast cup-shaped, smooth, lateral thickening with 1 spherical pyrenoid, sometimes angular in outline, in the middle of the chloroplast. Nucleus in the middle of the cell or slightly displaced to the posterior. Stigma not observed. Contractile vacuoles 2, apical. Flagella about same length as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 19-VII-1990, A.M. Werneck & M.M. Marinho s.n. (R173752).

This species is very similar to C. pseudogloegamaGerloff, from which it differs in the lack of a stigma. Gerloff (1940) had, however, referred to the sporadic occurrence of a pale yellowish stigma in the original description of C. typhlos. All Brazilian specimens presently studied never showed a structure that could be a stigma.

Reported from Germany. New to Brazil. Found in just one plankton sample unit from the Camorim Reservoir.

Group Amphichloris

Chlamydomonas parisii

Bourr., Hydrobiologia 3: 260, pl. 2, fig. 45-46. 1951.

Figures 26-27

 

Cells 20-25 µm long, 11-14 µm broad, elliptical, cylindrical-elliptical or ovoid. Cell wall hyaline, with a conical-truncated, double, anterior papilla. Chloroplast tubular, outer side with vertical ribbons, small rectangular space in the cell middle in which lies the nucleus. Pyrenoids 2, spherical, above and below the space. Stigma elliptical, in the upper part of the cell or behind the nucleus. Contractile vacuoles 2, apical. Flagella 1.3-1.5 times the cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at Bosque da Barra, 27-XII-1990, M. Menezes & I.S. Nogueira s.n. (R173642); the same, 27-XII-1990, M. Menezes & I.S. Nogueira s.n. (R173643); the same, 27-XII-1990, M. Menezes & I.S. Nogueira s.n. (R173644).

Some individuals (5% of total examined) werefound, which were different from the original description of C. parisii by having ovoid-shaped cell and the stigma located behind the nucleus (figure 27).

Recorded from France. New to Brazil. The species was found in the plankton of Bosque da Barra Lake.

Chlamydomonas pseudopertusa

H. Ettl, Arch. Protistenk. 108: 410, fig. 96. 1965.

Figures 28-30

Cells 18-21 µm long, 12-14 µm broad, elliptical or oblong. Cell wall hyaline, with a conical, truncated anterior papilla 1.3-2 µm long. Chloroplast tubular, smooth, rectangular outline to space in middle of cell, in which lies the nucleus. Pyrenoids 2, spherical, located below and above the space. Stigma elliptical, in the upper portion of cell, 3-4 µm long. Contractile vacuoles 2, apical. Flagella 0.5-1.5 times the cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 19-V-1990, A.M. Werneck s.n. (R173695); the same, 20-V-1989, M. Menezes s.n. (R180714).

Regarding cell shape, part of the material studied is similar to C. pertusa Chodat. However, the structure of the papilla was a very stable characteristic that led us to consider the Brazilian specimens identical to C. pseudopertusa.

Reported from Asia, Europe, New Zealand. New to Brazil. This species was collected only from the plankton of Camorim Reservoir.

Chlamydomonas

aff.

glans

Pascher,Arch. Protistenk. 69: 117, fig. 12. 1930.

Figures 31-33

Cells (12-)17(-22) µm long, 7.8-14.4 µm broad, elliptical. Cell wall hyaline, with a conical rounded as well as a truncated anterior papilla, a large pore at the center of a papilla present. Chloroplast tubular, open at both ends, smooth, rarely cup-shaped. Pyrenoid 1, spherical, in the middle of a transverse plate. Nucleus in the posterior space of the chloroplast. Stigma elliptical, lying in the upper cell portion, 1.2-2 µm long. Contractile vacuoles 2, apical. Flagella about the same length as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at Bosque da Barra, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173642); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n.(R173643); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173644).

Material from the state of Rio de Janeiro differs from the original description of the species in its homogeneous and thick cell wall and, sometimes, in the occurrence of a cup-shaped chloroplast. Our material also differs from of C. glans in having a conical rounded sometimes truncated anterior papilla.

Recorded from Europe. New to Brazil. This species was collected from the Bosque da Barra Lake plankton.

Chlamydomonas virgata

Pascher, Arch. Protistenk. 69: 120, fig. 15. 1930.

Figures 34-35

Cells 25-30 µm long, 10-13 µm broad, cylindricalellipsoidal, with a conical anterior cell pole. Cell wall hyaline, with a rounded anterior papilla. Chloroplast tubular, open at both ends, with vertically arranged superficial incisions, 5-7 unequal and parallel lobes, slightly curved towards the cell inside, sometimes interrupted at the transverse plate, with a spherical pyrenoid at the middle of the transverse plate. Nucleus in the posterior space of the chloroplast. Stigma elliptical, lying in the upper cell portion, about 2 µm long. Contractile vacuoles 2, apical. Flagella about the same length as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 10-I-1989, M. Menezes s.n. (R173417).

Part of the present material differs from the original description of C. virgata in that the outer side of the chloroplast is interrupted by incisions that produce lobes of different extent. However, we are of the opinion that localization, extensions, and number of chloroplast incisions and lobes are not good taxonomic characters. Similar conclusions also apply to Carteria lohammarii Skuja that was also identified in the present study.

Reported from Europe. New to Brazil. This species was found in plankton and metaphyton samples from the Camorim Reservoir.

Group Chlorogoniella

Chlamydomonas lunata

Pascher & Jahoda, Arch. Protistenk. 61: 262, fig. 16. 1928.

Figures 36-37

 

Cells 12-15 µm long, 5-7 µm broad, dorsiventral, elliptical in frontal view, ventrally curved, convex on dorsal side, convex in anterior 2/3 or retuse in its mid portion. Cell wall hyaline, with a hemispherical anterior papilla. Chloroplast dorsal, laminar, smooth, with a spherical pyrenoid in the middle of the dorsal side, sometimes below the cell equator. Nucleus central. Stigma elliptical, in the upper part of the cell, ca. 1.8 µm long. Contractile vacuoles 2, apical. Flagella about the same length as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at Bosque da Barra, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173644).

Reported from Austria. New to Brazil. This species was collected just once in the Bosque da Barra Lake plankton.

Chlamydomonas sordida

H. Ettl, Arch. Protistenk. 108: 381, fig. 75. 1965.

Figures 38-40

Cells (7-)10(-16) µm long, (4.6-)5(-8) µm broad, slightly dorsiventral, elliptical-ovoid or ovoid in frontal view, ventrally curved, dorsal margin convex, ventral margin slightly concave to the anterior portion of the cell. Cell wall hyaline, with a hemispherical anterior papilla. Chloroplast laminar, dorsal, smooth with a spherical pyrenoid in the middle of the dorsal part. Nucleus basal, excentric. Stigma elliptical, below the pyrenoid, 1.2-2 µm long. Contractile vacuoles 2, apical. Flagella ca. 1.5 times of cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 19-VII-1990, A.M. Werneck & M.M. Marimho s.n. (R173752).

Present plants from Rio de Janeiro State showed slightly longer cells when compared to the original description.

Reported from Europe. In Brazil this species has previously been recorded for the state of São Paulo (Bicudo 2004). This species was found just once in a plankton sample from Camorim Reservoir.

Chlamydomonas rattuli

Korš., Arch. Protistenk. 78: 590, fig. 48-49. 1932.

Figures 41-46

Cells free-swimming or an epibiont in the mucilage of plankton algae, 7-9 µm long, 2.8-3.2 µm broad, ovoid or club-shaped, sometimes curved, dorsal side convex, ventral side flattened or concave in the anterior third of cell, without papilla or with a small anterior protrusion similar to a membranaceous hemispherical papilla. Chloroplast dorsal, laminar, smooth, with a spherical pyrenoid in the middle of the dorsal part. Stigma punctiform, in the upper part of the cell, dorsal, near to the vacuoles. Contractile vacuoles 2, apical. Flagella 1-1.5 times the cell length. Asexual reproduction while cell is actively motile, by division into 4 daughter cells.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 19-VII-1990, M. Menezes s.n. (R173750).

Chlamydomonas rattuli is closely related to C. gloeophila Bourr. Both species are epibiontic in the mucilage of rotifers and plankton algae when vegetative and differ from each other by the presence of papilla in C. gloeophila. Bourrelly (1951) described and sketched C. rattuli with an anterior papilla, and considered C. gloeophila a synonym of the former. Ten years latter, Huber-Pestalozzi (1961) maintained C. rattuli sensu Korš. and considered C. rattuli described by Bourrelly (1951) to be synonymous with C. gloeophila.

We sometimes found in the same sample on the mucilage of Botryococcus braunii Kützing coexisting with some others that were actively motile. At other times, they were observed free-swimming in the plankton. Epiphytic cells were mostly attached to the substrate by their flagella. The latter were ovoid in shape, with a rounded anterior pole that did not touch the host surface (figures 41-42, 44). Club-shaped cells with their anterior pole attached to the substrate were also observed, which presented a slight short protuberance that was well delimited from the rest of the cell body (figures 43, 45).

Approximately the same variation above was also observed in populations of free-swimming specimens. Very often, epiphytic vegetative cells released from the substrate later on formed zoospores which kept themselves in a motile stage (figure 46).

Reported from Asia, Europe. New to Brazil. This species was found solely in the Camorim Reservoir plankton.

Chloromonas Gobi

Chloromonas

cf.

vesterbottnica

(Skuja) Gerloff & H. Ettl, Beih. Nova Hedwigia 34: 128, pl. 37, fig. 2. 1970 ≡

Chlamydomonas verstebottnica

Skuja,Nova Acta R. Soc. Scient.Upsal. s‚r. 4, 16(3): 140, pl. 20, fig. 9-11. 1956.

Figure 47

 

Cells 20-23 µm long, 11-18 µm broad, ovoid. Cell wall hyaline, with a hemispherical or truncate anterior papilla. Chloroplast cup-shaped, dissected into discoid portions. Nucleus central. Stigma elliptical, at the upper portion of cell, 6-7 µm long. Contractile vacuoles 2, apical. Flagella about the same length as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unnamed oxbow pond, 27-XII-1989, M. Menezes s.n. (R173641).

Our material differs from of C. vesterbottnica in having hemispherical sometimes truncate papilla. Chloromonas vesterbottnica shows a spherical-conical anterior papilla (Ettl 1983)

Reported from Denmark, Rumania, Sweden. New to Brazil. Present species was found in one plankton sample from the oxbow pond.

Gloeomonas Klebs

Gloeomonas tubulosa

(Pascher & Jahoda) H. Ettl, Ber. Nat.-med. Ver. Innsbruck 58: 1970 ≡

Chlamydomonas tubulosa

Pascher & Jahoda,Arch. Protistenk., 61: 274, fig. 27. 1928.

Figures 48-49

Cells 11-14 µm long, 8-10 µm broad, ovoid. Cell wall hyaline, with a short rounded sometimes truncate anterior papilla. Chloroplast cup-shaped, smooth, basal thickening occupying 1/3 of cell, without pyrenoid. Nucleus central. Stigma elliptical, at the upper part of cell, 3-3.5 µm long. Contractile vacuoles 2, apical. Flagella about the same length as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 19-VII-1990, M. Menezes s.n. (R173750); the same, 19-VII-1990, A.M. Werneck & M.M. Marinho s.n. (R173751).

Reported from Austria. New to Brazil. Found in the plankton of Camorim Reservoir.

Vitreochlamys Batko

Vitreochlamys cylindrica

(Skuja) Maidana & Vigna, Physis, sec. B, 46(110): 17. 1988 ≡

Sphaerellopsis cylindrica

Skuja, Symb. Bot. Upsal. 9(3): 93, pl. 9, fig. 20-26. 1948.

Figures 50-51

Cells 28-32 µm long, 14-18 µm broad, oblong, rounded poles, sometimes sides slightly concave. Cell membrane hyaline, wide. Protoplast 20-22 µm long, 7-10 µm broad, ovoid, widely distant from the membrane. Chloroplast urn-shaped, with conspicuous longitudinal striations, posterior lateral thickening, sometimes a spherical pyrenoid in the mid region. Nucleus central. Stigma elliptical, at the upper portion of cell, ca. 2 µm long. Contractile vacuoles 2, apical. Flagella about the same length as the cell, penetrating the cell membrane at both sides of papilla, surrounded by short tubes at the base.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 19-V-1990, A.M. Werneck s.n. (R173694); the same, 19-V-1990, A.M. Werneck s.n. (R173695).

Reported from Austria, Sweden. New to Brazil. This species was found in the plankton of Camorim Reservoir.

Vitreochlamys fluviatilis

(Stein) Batko, Acta Hydrobiol. 6: 415. 1970 ≡

Chlamydococcus fluviatilis

Stein, Infus. 3 (1): pl. 24, fig. 26-46. 1878.

Figures 52-55

Cells 15-20 µm long, 11-14 µm broad, ellipticalovoid, rounded poles. Cell membrane hyaline, wide. Protoplast 11-13 µm long, 6-8 µm broad, ovoid, elliptical-ovoid or pear-shaped widely separated from the membrane. Chloroplast cup-shaped, smooth, basal thickening occupying 1/3 of protoplast, with a spherical pyrenoid. Nucleus central, slightly displaced to the anterior cell end. Stigma elliptical, at the upper part of cell, ca. 1 µm long. Contractile vacuoles 2, apical. Flagella about the same length as the cell, penetrating the cell membrane at both sides of papilla, surrounded by short tubes at the base.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Bosque at Lake da Barra, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173642); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n.(R173643); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173644).

Some specimens were morphologically close to S. alpina Pascher & Jahoda (=Vitreochlamys alpina?) and V. ordinata (Skuja) Nakazawa, respectively, since having the protoplast at a much shorter distance from the membrane and the elliptical-ovoid protoplasts.

Reported from Hungary, India, Japan, North America, Rumania, Scandinavia, Ukranie. New to Brazil. Material studied was planktonic and collected from the Bosque da Barra Lake.

Vitreochlamys lefevrei

(Bourr.) Menezes & C. Bicudo, comb. nov. ≡

Sphaerellopsis lefevrei

Bourr. Hydrobiologia 3: 266, pl. 5, fig. 99. 1951.

Figures 56-57

Cells 25-28 µm long, 12-15 µm broad, oblong, poles rounded. Cell membrane hyaline, wide. Protoplast 15-18 µm long, 7-9 µm broad, elliptical or elliptical-ovoid, widely distant (about 3,5 µm) from the membrane. Chloroplast cup-shaped, smooth, basal thickening occupying almost 1/2 of protoplast, with a spherical basal pyrenoid. Nucleus anterior, median, sometimes shifted from the longitudinal axis. Stigma elliptical, anterior, 1.5-2 µm long. Contractile vacuoles 2, apical. Flagella about 0.6 times the cell length, penetrating the cell membrane at both sides of papilla, surrounded by short tubes at the base.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 29-IX-1988, A.M. Werneck s.n. (R173160).

The genus has been revised by Nazakawa et al. (2001) based upon strains isolate from different parts of Japan, the Czech Republic, Slovak Republic and Germany using comparative light and electron microscopy and rbcL (large subunit of Rubisco, ribulose-1,5-bisphosphate carboxylase/oxygenase) gene phylogeny. The results of this work showed that the delimitation of the six studied species- V.fluviatilis (Stein) Batko, V. gloeocystiformis (Dill) Nakazawa, V. ordinata (Skuja) Nakazawa, V. nekrassovii (Korš.)Nakazawa, V. aulata (Pascher) Batko, and V. pinguisNakazawa based on the morphological differences used in the traditional taxonomic system were consistent with molecular data: cell and protoplast shape, numbers of contractile vacuoles and pyrenoids, form and position of the stigma, location of the nucleus, degree of longitudinal striations on the chloroplast surface. Besides of these characters Nakazawa et al. (2001) included as criteria for species delimitation the ultrastructure of the pyrenoid and stigma.

Although our material have not been analyzed under electron microscopy the populations previously identified as S. lefevrei belong to the group of species of Sphaerellopsis/Vitreochlamys with chloroplast surface smooth or with inconspicuous striations and nucleus sometimes shifted from the longitudinal axis. However, it can be distinguished from these species by clearly its oblong cell.

Reported from Austria, France and Switzerland. New to Brazil.This species has been found in the plankton of an oxbow pond.

Haematococcaceae

Chlorogonium Ehrenberg

Chlorogonium fusiforme

Matv., Proc. Kharkov A. Gorky State Univ. 14: 62, pl. 4, fig. 43-33. 1938.

Figures 58-59

 

Cells 25-30 µm long, 5-8 µm broad, fusiform, anterior end truncate, posterior end acuminate. Chloroplast laminar, smooth, with a spherical central pyrenoid. Nucleus almost central. Stigma elliptical, anterior, ca. 1.3 µm long. Contractile vacuoles 2, one anterior, the other one posterior to the nucleus. Flagella 0.3-0.5 times the cell length. Asexual reproduction while cell actively motile, by division into 4 daughter cells.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Bosque at Lake da Barra, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173642); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n.(R173643); the same, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173644).

Reported from Austria, Czech Republic, Denmark, Japan, Slovakia, Rumania, Ukraine. In Brazil this species has previously been recorded for the state of São Paulo (Bicudo 2004). The species was collected from the Bosque da Barra Lake plankton.

Haematococcus C. Agardh emend. Flotow.

Haematococcus capensis

Pocock var.

capensis

, Trans. R. Soc. S. Afr. 36(1): 13, fig. 1-2, pl. 1b-c, pl. 2d-g. 1960.

Figures 60-61

Cells (15-)19(-25) µm long, (11-)13(-17) µm broad, elliptical, rounded poles. Cell membrane hyaline, wide, lacking papilla. Protoplast 12-15 µm long, 7-8.5 µm broad, ovoid, distant from the outer membrane (ca. 3-5 µm), with a hyaline anterior end forming a conspicuous mammillae projected, which is distant from the outer membrane of about 2 µm. Coarse radiating protoplasmic processes branched or not, 13-14 hyaline cytoplasmic strands frequently branched and connecting the protoplasmic processes to the outer membrane. Chloroplast cup-shaped, smooth, basal thickening occupying almost 1/2 of protoplast, with a spherical basal pyrenoid. Stigma elliptical, near to the pyrenoid, ca. 1 µm long. Contractile vacuoles numerous, irregularly distributed. Flagella about the same length as the cell, penetrating the cell membrane at both sides of papilla, surrounded by short, divergent tubes at the base.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 22-I-1990, M. Menezes & A.M. Werneck s.n. (R173658).

Present material somewhat differs from the original description of the species in the smaller cell dimensions.

Reported from South Africa. New to Brazil. Found in the plankton of Camorim Reservoir.

Haematococcus droebakensis

Wollenw. var.

droebakensis

, Ber. dt. bot. Ges. 25: 318, fig. 10-15. 1907.

Figures 62-65

Cells (30-)33(-37) µm long, (20-)23(-26) µm broad, elliptical, rounded poles. Cell membrane hyaline, wide, sometimes with a 2-lobed papilla. Protoplast (22-)26(-32) µm long, (20-)23(-26) µm broad, elliptical or ovoid, apart 5-7 µm from the outer cell membrane, anterior end hyaline, projecting into a conspicuous conical-rounded mammilla which is 1.2-3.5 µm from the outer cell membrane. Coarse radiating protoplasmic processes branched or not, mostly with 11 branched hyaline cytoplasmic strands connecting protoplasmic processes with the outer membrane, sometimes cytoplasmic strands absent and the protoplasmic processes touching the cell membrane. Chloroplast tubular, with a central space rectangular in outline, smooth, pyrenoids 2, one anterior, the other one posterior to the space. Hematochrome granules numerous, disperse in the cytoplasm. Nucleus in the space of the chloroplast. Stigma elliptical, in the space of chloroplast, ca. 1 µm long. Contractile vacuoles numerous, irregularly distributed. Flagella about the same length as the cell, penetrating the cell membrane at both sides of papilla, surrounded by short tubes at the base.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 25-III-1990, A.M. Werneck & M.C. Bernardes s.n. (R173686); the same, 20-V-1989, M. Menezes s.n. (R180714).

Some individuals studied showed some features of H. zimbabwiensis Pocock, such as longer flagellar tubes (figure 63) and a small papilla at the anterior protoplast pole (figure 64). Furthermore, about 1% of individual cells examined lacked cytoplasmic strands, the protoplasmic processes being more developed and sometimes touching the cell membrane (figure 65). Latter specimens do not agree with Pocock's (1960) circumscription of the species that states that presence of cytoplasmic strands is a diacritic character for Haematococcus.

Reported from Hungary, South Africa. In Brazil it was previously reported for the Amazonas State (Bittencourt-Oliveira 1993). The species was collected from the Camorim Reservoir plankton.

Haematococcus pluvialis

Flotow

emend

. Wille, Nyt. Mag. Naturvid. 41: 89. 1903.

Figures 66-67

Cells 30-37 µm long, 25-35 µm broad, broadly elliptical or spherical, rounded poles. Cell membrane hyaline, wide. Protoplast 22-25 µm long, 13-16 µm broad, ovoid, 5-7 µm from the outer membrane, anterior pole hyaline, projecting into a conspicuous conical-rounded mammilla, which is 3-7 µm from the outer membrane. Radiating hyaline protoplasmic processes delicate, mostly with 25-52 branched hyaline cytoplasmic strands connecting the protoplasmic processes with the outer membrane, sometimes cytoplasmic strands absent, the protoplasmic processes then touching the cell membrane. Chloroplast cupshaped, smooth, basal thickening occupying 1/3 of protoplast, pyrenoids 3-7, spherical, irregularly distributed. Hematochrome granules numerous, dispersed in the cytoplasm. Nucleus central. Stigma elliptical, at the upper cell part, ca. 1 µm long. Contractile vacuoles numerous, irregularly distributed. Flagella about the same length as the cell, penetrating the cell membrane at both sides of the papilla, surrounded by short divergent tubes at the base.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 20-V-1989, M. Menezes s.n. (R180714).

Registered from Hungary and Slovakia. New to Brazil. The species was found in the plankton of Camorim Reservoir.

Phacotaceae

Pteromonas Seligo

Pteromonas pseudoangulosa

L.S. Peterfi, Strudii si cercet. biol. ser. Bot. 17: 272, pl. 1, fig. 18-19. 1965.

Figures 68-83

 

Cells 12-25 µm long, 10-13 µm broad, in front view bell-shaped, outline irregular, poles truncate, sometimes posterior end rounded. Cell wall composed of 2 overlapping halves, broadly elliptical or slightly hexagonal in lateral view, sides with concave depressions, oblong, elliptical or hexagonal in apical view, rarely quadrangular, sides straight or 1 or both sides with concave depressions, 2 colorless, short, longitudinal wing-like projections on each valve half, sigmoid in apical view. Protoplast 8-12 µm long, 5-10 µm broad, 1-1.5 µm from outer membrane. Chloroplast cup-shaped, smooth, basal thickening taking almost ½ of protoplast, pyrenoid 1, spherical, usually basal or lateral, sometimes 2-5 irregularly distributed. Nucleus central. Stigma elliptical, at the upper portion of cell, ca. 2 µm long. Contractile vacuoles 2, apical. Flagella 1-2 times the cell length. Asexual reproduction while actively motile, by division into 4 daughter cells.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unammed oxbow pond, 30-IV-1990, M. Menezes & L H.S. Silva s.n. (R173160); the same, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173690).

Specimens from the state of Rio de Janeiro showed variation in cell shape that partially agreed with the one registered by Peterfi (1965, 1968) for P. pseudoangulosa L.S. Peterfi. Some specimens resembled those of P. angulosa Lemm. in having one or both sides of the cell with a median concave depression (figures 71, 77-78). Some other ones were similar to P. cordiformis

Lemm. in having a flagellar tube (figures 80-81). In all individuals examined variation was detected in the number of pyrenoids, from one to five, as well as in their location in the cell.

After fixation, some cells showed wall surface somewhat waved (figure 82) that, when the specimens had two or more pyrenoids, were very much similar to those of P. aequiciliata (Gicklhom) Bourr.

Reported from Rumania. New to Brazil. Pteromomonas pseudoangulosa was found in plankton and metaphyton samples from a roadside oxbow pond.

Volvocales

Goniaceae

Gonium O.F. Müller

Gonium formosum

Pascher

in

Pascher, Süsswasserfl. Deutschl. 4: 418. 1927.

Figures 84-85

 

Vegetative colonies, ca. 60 µm long, ca. 60 µm broad, quadrangular, 16-celled. Cells 10-11 µm long, 9-10.5 µm broad, ovoid, connecting processes almost 2/3 of cell length. Chloroplast cup-shaped, smooth, basal thickening taking more than half of cell, pyrenoids 1-2, spherical, basal. Stigma globose. Contractile vacuoles 2, apical. Flagella homodynamic, 2 times as long as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 29-IX-1988, M. Menezes & A.M. Werneck s.n. (R172159); the same, 19-V-1990, A.M. Werneck s.n. (R173695); the same, 27-VIII-1990, M. Menezes s.n. (R173754); Lagoinha, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173657); unammed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160); the same, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173691).

Reported from Hungary and North America. In Brazil it was previously reported for the state of Rio Grande do Sul (Francheschini 1992). This species was found in the plankton from Camorim Reservoir, and in the sediment of the unnamed roadside oxbow pond.

Gonium multicoccum

Pocock, Madräno 13: 1955.

Figures 86-88.

Vegetative colonies, 60-65 µm long, 60-65 µm broad, quadrangular or rhomboidal, 32-celled. Cells 10-14 µm long, (7-)8(-11) µm broad, elliptical or rounded, connecting processes up to 1/2 the cell length. Chloroplast cup-shaped, smooth, basal thickening taking up to half of cell, pyrenoids 3-7, spherical, irregularly distributed in the inner edge of plastid. Stigma globose. Contractile vacuoles 2, apical. Flagella homodynamic, 1-1.5 times the cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unammed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160); the same, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173691).

Reported from Australia, North America. Previously reported for Brazil from Amazonas State (Bittencourt-Oliveira 1993). The species was found in plankton and sediments from a roadside oxbow pond.

Volvocaceae

Eudorina Ehrenberg

Eudorina elegans

Ehrenberg, Phys. Abh. K. Akad. Wiss. Berlin, 1831: 78, pl. 2, fig. 10a-d. 1831.

Figures 89-94

 

Vegetative hollow colonies, mature colonies (50-)100(-150) µm long, 43-110 µm broad, young colonies 60-70 µm diam., daughter colonies (26)30 (-37) µm diam., elliptical or rounded, 32-celled. Daughter colonies surrounded by a mucilaginous sheath. Colonial sheath double, confluent, smooth or waved, posterior pole rounded, truncate or with mammillae. Cells 5-7 µm diam., spherical, surrounded by a mucilaginous sheath. Chloroplast cup-shaped, smooth, basal thickening taking more than half of cell, pyrenoids 1-2 in the young cells, 3-4 in the mature cells, spherical, basal. Stigma globose, decreasing in size towards the posterior cells tiers where it may sometimes be absent. Contractile vacuoles two, apical, and several others randomly distributed. Flagella homodynamic, 1.5-2 times the cell length. Cellular division not synchronous.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at Bosque da Barra, 1-X-1988, M. Menezes s.n. (R172923); the same, 16-X-1989, M. Menezes s.n. (R173614); unammed oxbow pond, 30IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160);the same, 19-VI-1990, M. Menezes & I.S. Nogueira s.n. (R173747); the same, 27-VIII-1990, M. Menezes s.n. (R173758); Camorim Reservoir, 27-VIII-1990, M. Menezes s.n. (R173754).

The presence of an extra cellular matrix of the gelatinous parental envelope, together the maximum numbers of cells in a colony and modes of sexual reproduction have been considered important morphological features for delimiting volvocacean genera based on cultures, ultrastructure and molecular phylogeny studies. Pandorina Bory have up to 16 vegetative cells without the cellular envelopes of the matrix with isogamous reproduction without sperm packets. Within genera with up to 32 cells per colony, like Eudorina Ehrenb. and Yamagishiella Nozaki, each vegetative cell is enclosed by the cellular envelope. Eudorina has anisogamous sexual reproduction with sperm packets while Yamagishiella shows isogamous sexual reproduction without sperm packets (Nozaki & Kuroiwa 1992, Nozaki & Krienitz 2001, Nozaki et al. 1997, 2000, Coleman 2001).

Although we have not observed sexual reproduction in the Brazilian populations, these can be assigned to the genus Eudorina in having more than two contractile vacuoles. These reject the possibility that material belongs to the genus Yamagishiella which has only two contractile vacuoles (H. Nozaki, unpublished data).

Colonial envelope varied both in the young and adult colonies, being smooth, waved, with the posterior pole sometimes truncate. Mammillae at the posterior pole of colonial matrix were only observed in adult colonies, where they became more evident after fixation (figure 94). Frequently, in asexual reproduction the cells showed division not synchronous resulting in an irregular distribution and differentiation of daughter

colonies development in a colony (figure 91).

Cosmopolitan distribution. The species was previously recorded from samples from North, Southeast, and South Brazil (Cunha 1913, Kammerer 1938, Uherkovich & Schmidt 1974, Bittencourt-Oliveira 1993, Huszar 1984, Dewes & Huszar 1986, Huszar et al. 1987, Sant'Anna et al. 1988, Franceschini 1992). Found in plankton and metaphyton samples from Bosque da Barra Lake and a roadside oxbow pond.

Eudorina illinoisensis

(Kofoid) Pascher

in

Pascher, Süsswasserfl. Deutschl. 4: 443, fig. 464-465. 1927 ≡

Pleodorina illinoisensis

Kofoid, Bull. Ill. St. Lab. Nat. Hist., 5: 273, pl. 36-37. 1898.

Figures 95-96

 

Vegetative hollow colonies, (102-)132(-140.6) µm long, (182-)112(-114) µm broad, elliptical, 32-celled. Colonial sheath double, confluent, smooth or with usually posterior, rarely anterior, mammillae. Cells spherical or ovoid, surrounded by a mucilaginous matrix, cells of first anterior tier 9-10 µm long, 5-7.6(-8) µm broad, all remaining cells tiers (19-)20(-23) µm diam. Chloroplast cup-shaped, smooth, basal thickening taking more than 1/2 of cell, pyrenoid 1 in the anterior somatic cells, 3-5, spherical, irregularly distributed in the others cells. Stigma globose, decreasing in size towards the posterior cells tiers. Contractile vacuoles two, apical, and several others randomly distributed. Flagella homodynamic, ca. 1.5 times the cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unnamed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160).

Eudorina illinoisensis (Kofoid) Pascher resembles some species of Pleodorina Shaw, like in P. thompsonii Ott, Nozaki & Coleman and Pleodorina westii Tiffany, in having 16-or 32-celled colonies with small somatic cells. However, thirty-two-celled colonies of E. illinoisensis have four small anterior cells that may or may not divide to form a daughter colony. i.e. facultative somatic cells (Goldstein 1964, Nozaki 1986). Moreover, in 32-celled colonies of P. westii the small somatic cells are nearly equal in number as large reproductive cells and are distributed in both anterior and posterior portions of colony (Tiffany 1935). In 32-celled colonies of P. thompsonii there are 12 small somatic cells that are only at the anterior pole of the colony and do not divide (Nozaki et al. 2006).

Populations studied always showed individual cells having gelatinous sheath that was hexagonal in outline. Sheaths were clearly evident in adult colonies without staining (figure 95). In young colonies, however, sheaths were evident only after staining (figure 96).

World-wide distribution. In Brazil it was previously recorded from North, Southeast, and South areas (Kammerer 1938, Thomasson 1971, Bittencourt-Oliveira 1990). This species was collected from the plankton of an unnamed roadside oxbow pond.

Eudorina unicocca

G.M. Smith, Bull. Torrey bot. Club 57: 363, pl. 17, fig. 3-4. 1930.

Figures 97-99

Vegetative hollow colonies, (95-)123(-133) µm long, (79.8-)95(-114) µm broad, elliptical or elliptical-obovoid, 32-celled. Colonial sheath double, confluent, slightly waved, internal layer formed by individual cells envelopes in the larger colonies, posterior pole with mammillae. Cells 7-11 µm diam., spherical, sometimes surrounded by a mucilaginous hexagonal matrix. Chloroplast cup-shaped, smooth, basal thickening taking more than 1/2 of cell, pyrenoid 1, spherical, basal. Stigma decreasing in size towards the posterior tiers. Contractile vacuoles two, apical, and several others randomly distributed. Flagella homodynamic, about 2 times the cell diameter.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at Bosque da Barra, 1-X-1988, M. Menezes s.n. (R172923); the same, 8-IV-1990, M. Menezes s.n. (R173689); unammed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160).

Based upon the single basal pyrenoid in the cup-shaped chloroplast of the vegetative cells our populations could be assignable to Eudorina unicocca G.M. Smith or Yamagishiella unicocca (Rayburn & Starr) Nozaki (Goldstein 1964, Nozaki et al. 1997). However, the occurrence of more than two contractile vacuoles led us to consider the Brazilian specimens as E. unicocca.

Cells in colonies may or may not have individual gelatinous sheaths. Individual sheaths clearly observed without staining were detected in adult colonies, which also had more developed posterior mammillae.

Reported from Central and North America, Switzerland. In Brazil, it was previously recorded from North, Southeast, and South areas (Uherkovich & Rai 1979, Stankiewiez 1980, Huszar 1984, 1985, Dewes & Huszar 1986, Bittencourt-Oliveira 1993). This species was found in plankton, neuston and metaphyton samples from Bosque da Barra Lake and from an unnamed roadside oxbow pond.

Volvulina Playfair

Volvulina steinii

Playfair, Proc. Linn. Soc. N. S. W. 40(2): 337, pl. 43, fig. 3-4, 8. 1915.

Figures 100-101

 

Vegetative colonies compact, (42-)45(-68) µm diam., globose to almost perfectly spherical, 16-celled. Colonial sheath sometimes formed by confluence of cellular envelopes. Cells (6-)9(-10) µm long, (11)12-13(-15) µm broad, hemispherical or lenticular in front view, spherical in polar view, peripherically distributed in the colonial envelope in alternating tiers of 4 cells each, surrounded by a mucilaginous, hexagonal envelope, which may sometimes be confluent. Chloroplast cup-shaped, smooth, basal thickening taking almost the whole cell, pyrenoid absent. Stigma elliptical, at the anterior 1/3 of the cell, largest in the 1^st tier, reduced in the 2^nd and 3^rd tiers, lacking in the posterior one. Contractile vacuoles 4-6, apical. Flagella about 2 times as long as the cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unammed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160); the same, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173691).

According to Starr (1962), V. steinii Playfair differs from V. pringsheimii Starr mainly in the presence of pyrenoids in the vegetative cells of the latter species. However, light and electron microscopy studies carried out by Nozaki (1982) and Nozaki et al. (1987) have shown that V. pringsheimii has pyrenoids always at the bottom of the chloroplast, and pyrenoids do not vary with culture age. Vegetative cells of V. steinii, however, may also have pyrenoids located at the chloroplast edge but only in some culture conditions. Nozaki and collaborators also observed that during the daughter colonies formation of V. pringsheimii the parental pyrenoid divided and was distributed to each one of the cells of the newly formed colonies. In V. steinii, pyrenoids do not divide but remain in one of the daughter cells of newly formed colonies. Moreover, V. pringsheimii has two contractile vacuoles while V. steinii shows two to eight (Ettl 1983).

In the Rio de Janeiro State material, some colonies were similar to those of V. pringsheimii in having the colonial sheath distinct from the cellular one. However, in all colonies studied cells with 4-6 contractile vacuoles without pyrenoids were observed, what led us to identify the Brazilian material with V. steinii.

World-wide distribution. In Brazil, it was previously reported from Amazonas State (Gessner 1931, Bittencourt-Oliveira 1993). This species was found in the plankton and sediments from an unnamed roadside oxbow pond.

Yamagishiella Nozaki

Yamagishiella sp.

Figures 102-105

 

Vegetative colonies compact, mature colonies 40-45 µm long, 35-42 µm broad, young colonies 20-25 µm long, 15-20 µm broad, daughter colonies 8-15 µm long, 5-10 µm broad, elliptical, rounded or oblong, 16-celled. Daughter colonies surrounded by a mucilaginous sheath. Mature cells 10-15 µm long, 8-15 µm broad, young cells 5-7 µm long, 4-5 µm,daughter cells 3-5 µm long, 2.5-3.5 µm broad, pearshaped. Chloroplast cup-shaped, smooth or dissected into vertical lobes, basal thickening taking more than 1/2 the cell, pyrenoid 1, spherical, basal. Stigma globose, decreasing in size in posterior cells tiers. Contractile vacuoles 2, apical. Flagella homodynamic, 1-1,5 times the cell length.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unammed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173160); the same, 19VI-1990, M. Menezes & I.S. Nogueira s.n. (R173747); the same, 27-VIII-1990, M. Menezes s.n. (R173758);Camorim Reservoir, 27-VIII-1990, M. Menezes s.n. (R173754).

Our populations were identified initially as Pandorina morum (O.F. Müller) Bory in having compact colonies with up to 16 cells. However, the presence of the transparent vesicle inside the parental colonial envelope (figures 103a) and in the daughter colonies (figure 103b, ^c) is not characteristic of Pandorina where the cell division takes place in a 'keystone shaped space' in the parental gelatinous matrix only (Angeler 1998, Nozaki & Kuroiwa 1992). The presence of this gelatinous (extracellular) matrix of vegetative colonies and the two contractile vacuoles indicated that Rio de Janeiro material can be assigned to Yamagishiella unicocca or even represent a new species of this genus. Since we have not worked with cultures and not observed sexual reproduction it was impossible to define a better circumscription of Brazilian populations (see comments of E. elegans).

 

Reported from Europa, Asia, South America, North America China, Finland, Chile, Austria, Germany, Ukraine, Japan, and North America. The material was collected from the plankton of Camorim Reservoir and metaphyton of a roadside oxbow pond.

PRASINOPHYCEAE

Polyblepharidales

Pedinomonadaceae

Monomastix Scherffel

Monomastix ophisthostigma

Scherfell, Arch. Protistenk. 27: 107, fig. 1-46. 1912.

^{Figures 106-112}

Cells 14-21 µm long, 6-10 µm broad, oblong, ovoid or elliptical-oblong, slightly dorsiventrally compressed. Chloroplasts usually 2, sometimes 1, parietal, cup-shaped, a deep vertical incision dividing the chloroplast in to 2 lateral lobes, pyrenoid 1, spherical, in the middle of each lobe. Posterior part of cell with (1-)3-4 cylindrical trichocystis, 3-5 µm long, parallel to each other or almost so, sometimes absent. Stigma elliptical, 1.5 µm long, in the posterior part of cell. Contractile vacuole 1, apical.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Camorim Reservoir, 29-IX-1988, M. Menezes & A.M. Werneck s.n. (R172159); the same, 19-V-1990, A.M. Werneck s.n. R173695); the same, 27-VIII-1990, M. Menezes s.n. (R173754); the same, 20-V-1989, M. Menezes s.n. (R180714); unammed oxbow pond, 30-IV-1990, M. Menezes & L.H.S. Silva s.n. (R173691).

Fifty cells were observed and most (ca. 80%) had exactly the morphology and dimensions described in Scherffel (1912). Some individuals (about 20% of total specimens seen), however, resembled young cells of M. ophistostigma described by Belcher (1965) in having relatively shorter cells (ca. 11 µm long), a single chloroplast divided in two portions and in the number (2-4) of trichocysts (^{figures 109-110}). In May 1990, oblong ellipsoidal individual cells with rounded poles, trychocysts number varying from 0-3 were found (^{figures 111-112}), might be identified as M. minuta.

As far as we now, M. ophistostigma is only reported from Europe; new to Brazil. This species was collected from the plankton of Camorim Reservoir.

Pedinomonas Koršikov

Pedinomonas minutissima

Skuja, Nova Acta R. Soc. Scient. Upsal. Ser. IV, 16(3): 99, pl. 9, fig. 25. 1956.

^{Figures 113-114}

Cells 3-4 µm long, 2-3.5 µm broad, flattened laterally, elliptical to discoid in frontal view. Chloropast cup-shaped taking nearly the entire cell, pyrenoid 1, spherical, basal. Stigma punctiform, approximately at the cell equator. Flagellum about as long as the cell diameter.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, unnamed oxbow pond, 27-XII-1989, M. Menezes s.n. (R173641).

Cosmopolitan in distribution. In Brazil the species was previously recorded from a brackish coastal lagoon in the state of Rio de Janeiro (Menezes & Domingos 1994, Domingos & Menezes 1998). Pedinomonas minutissima was found only once in a plankton sample from the roadside oxbow pond.

Polyblephariadaceae

Nephroselmis Stein

Nephroselmis discoidea

Skuja, Sym. Bot. Upsal. 9(3): 65, pl. 5, fig. 12-18. 1948.

^{Figures 115-116}

Cells 5.5-9 µm long, 6-12 µm broad, flattened laterally, outline globose in front view, sometimes elliptical, anterior end round, with a apical swelling. Chloroplast cup-shaped, taking half of the cell, pyrenoid 1, spherical, basal. Contractile vacuole 1, apical. Stigma small, near the insertion of flagella. Flagella heterodynamyc, unequal in size, the shorter one about the cell length, the longer one ca. 1.5 times as long as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at Bosque da Barra, 16-X-1989, M. Menezes s.n. (R173616); Lagoinha, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173648).

Reported from Europe; new to Brazil. The species was found in two plankton samples from the Bosque da Barra Lake and in one of metaphyton from Lagoinha.

Tetraselmidales

Tetraselmidaceae

Tetraselmis Sein

Tetraselmis bichlora

(H. Ettl & O. Ettl) Norris

et al

., Bot. Mag. Tokyo 93: 336. 1980 ≡

Platymonasbichlora

H. Ettl & O. Ettl, Arch. Prostistenk. 105: 281, fig. 1. 1961.

^{Figures 117-118}

Cells 20-24 µm long, 12-16 µm broad, dorsiventrally flattened, broadly elliptical in frontal view, kidney-shaped in lateral view, anterior end rounded, with a deep apical depression. Chloroplasts 2, lateral, laminar, filling the entire cell, without pyrenoid. Two irregular rows of rounded trichocysts extending towards the posterior part of cell, others dispersed in the cytoplasm periphery. Stigma elliptical or rounded, short distance below the cell equator. Contractile vacuoles 2, apical. Flagella about as long as the cell.

Material studied: BRASIL. Rio de Janeiro: Rio de Janeiro, Lake at Bosque da Barra, 27-XII-1989, M. Menezes & I.S. Nogueira s.n. (R173644).

Reported from Europe; new to Brazil. This species was gathered just once, from the plankton of Bosque da Barra Lake.

Discussion

The circumscription of unicellular and colonial volvocacean species is problematic because most of their original descriptions were based on light microscope of a few specimens from natural samples without information of the intraspecific variation of morphological characters or of the life history (Pröschold et al. 2001, Nozaki & Krienitz 2001, Nozaki et al. 2006). Several species have been recently analyzed using combined morphology, reproduction, and molecular data, however, a reasonable part of these results are divergent with respect to their taxonomy and many questions still remain concerning their revisions that must be solved before accepting their new generic names for clades (Colemann 2002, Hoham et al. 2002, Pollio et al. 2005, Nozaki et al. 2006).

In this work morphological characteristics used to distinguish species of flagellate green algae were, in general, often adequate for that purpose and the majority of examined populations were morphologically well defined and easy to distinguish under light microscope. However, the variability detected for some morphological features only from field observations and the absence of reproductive aspects becomes confusing and a clear delineation of some taxa was not possible like Chlamydomonasaff. glans, Chloromonas cf. vesterbottnica and Yamaguishiella sp. Possibly many of these organisms here identified can represent new taxa for the science or only morphotypes but would be necessary to analyze better the phenotypic living population characteristics from field observations and cultured material to determine the range of their sexual reproduction, size and shape.

Observation of cell habit did not help in the identification of Carteria lohammari and Chlamydomonas rattuli, both species being described in the literature as epibiontic on zooplankton or phytoplankton organisms. Skuja (1948) had already reported free-swimming vegetative cells of C. lohammari forming zoospores in the absence of substrate and for other epibiontic species of Chlamydomonas, especially during the warm season of the year. According to Burkholder (1996), such a kind of interaction apparently is not obligatory since the alga may also be epizoic. This could be well interpreted as a dispersal mechanism or adaptation strategy to increase the organism's access to nutrients, to escape predators, or to get an advantageous location for growing. Then, it is possible to understand the epibiontic habit as a facultative phase that would constitute an adaptation process under stressed environmental conditions.

Variation in the number and extend of the chloroplasts in Carteria lohammari and Chlamydomonas virgata, as well as in the pyrenoid number and size in Chlamydomonas pseudotarda needs to be re-evaluated. Apparently, these characteristics could be related to the growth and/or physiology of the organism. However, in the present study increase of pyrenoid number was closely related to the increase of cell dimensions, especially with the cell breadth. Concerning size, different pyrenoid development stages in the same specimen might correspond to the formation of secondary pyrenoids in adult cells, as it was already observed in some Hematoccocaceae (Nozaki et al. 1998).

In relation to occurrence of identified taxa in different communities and environments our results agreed with literature data based on morphospecies criteria. Among the present unicellular species of Chlorophyta (24 of Chlorophyceae and four of Prasinophyceae), all 27 were always found in the plankton, mainly in Camorim Reservoir. Colonial volvocacean, in particular species of Gonium and Volvulina, were more frequent in the plankton, metaphyton, and sediments of the oxbow pond. Flagellated green algae are found in a variety of differing aquatic habitats. In such environments their life cycle provides for rapid colonization and further growth (Desnitski 2000, 2002, 2003). Particularly, Chlamydomonadales and colonial representatives of the Volvocales are frequently associated with small, mild alkaline, shallow water bodies. The latter group (Volvocales) is also often associated with high solar radiation (Menezes 1999, Pocock 1933a, b, Reynolds 1997) hot dry weather, warm water temperatures (Znachor & Jezberova 2005).

From the geographical distribution view based on phenotypic approach available in literature the flagellate chlorophyte flora (taxa size < 0,15 mm) was represented mainly by taxa that are common in temperate climates and that constituted about 57% of the total number of identified taxa. 17% of identified taxa were common to temperate and tropical climates, and 12% were typical of warmer climate. Cosmopolitan and worldwide distributed taxa accounted for 14% of total taxa identified.

Even retaining the concept of morphospecies our findings did not support the main idea about the distribution of microrganisms, "cosmopolitan hypothesis", which predicts that free-living microbes will be present in all kinds of environments where they can live due to unrestricted dispersal capabilities (Finley 2004, Fenchel 2005). This means that freeliving microbes have a lack of biogeography pattern and a low global diversity (Logares 2006). This capacity of ubiquitous dispersal is driven by huge population sizes leading to low probability of local extinction and, then the most microorganisms with size smaller than 1 mm have worldwide distribution while those larger than 10 mm are much less abundant and rarely cosmopolitan (Fenchel & Finlay 2003, Finlay & Fenchel 2004). Although some molecular studies support the "cosmopolitan hypothesis" (Daugjberg et al. 2000) for microalgae this idea is controversy because more recently available data suggest that micro-organisms (including protists and algae) have distribution patterns similar to those known from higher plants and animals, and that these patterns reflect historical (Gondwanan/Laurasian), ecological (tropical/temperate), and continental/local conditions (Coleman 2002, Foissner 2006, Luo et al. 2006).

In summary, our results may reflect, just how little is known about the chlorophytes flora from tropical and subtropical regions of the World. It is important to highlight the limited sampling of tropical species in the studies encompassing more complete description of intraspecific variation on natural and cultured material and/or more advanced tools like molecular phylogenetic.

Then, further investigations of Brazilian green flagellates should proceed, at least, to integrate morphological analysis on natural and cultured material in order to analyze the degree of phenotypic plasticity, to observe the characteristics of reproduction/life cycle, and correlate them to environmental conditions.

Acknowledgements

Authors wishes express their thanks to the CNPq, Conselho Nacional de Desenvolvimento Científico e Tecnológico, and CAPES, Coordenadoria de Aperfeiçoamento de Pessoal do Ensino Superior, for partial financial support.

Literature cited

Received: 17.10.2007; accepted: 18.09.2008

• Angeler, D. 1998. Life cycle studies and culture experiments for identification of doubtful volvocacean organisms - observations on Pandorina and Eudorina Hydrobiologia 369/370: 269-275.
• Belcher, J.H. 1965. An investigation of three clones of Monomastix Scherffel by light microscopy. Nova Hedwigia 9: 72-82.
• Bicudo, C.E.M. 2004. Criptógamos do Parque Estadual das Fontes do Ipiranga, São Paulo, SP. Algas, 18: Chlorophyceae (Volvocales). Revista Brasileira de Botânica 27: 85-102.
• Bittencourt-Oliveira, M.C. 1993. Ficoflórula do Reservatório de Balbina, Estado do Amazonas, II: Ulothricales e Volvocales (Chlorophyceae) e classe Oedogoniophyceae. Revista Brasileira de Biologia 53: 469-475.
• Bourrelly, P. 1951. Volvocales rares ou nouvelles. Hydrobiologia 3: 251-281.
• Burkholder, J.A. 1996. Interactions of benthic algae with their substrata. In: R.J. Stevenson, M.L. Bothwell & R.L. Iowe (eds.). Algal ecology: freshwater benthic ecossystems. Aquatic Ecology Series. Academic Press, California, pp. 235-297.
• Castro, A.A.J., Bicudo, C.E.M. & Bicudo, D.C. 1991. Criptógamos do Parque Estadual das Fontes do Ipiranga, São Paulo, SP. Algas 2: Cryptophyceae. Hoehnea 18: 87-106.
• Coleman, A.W. 2001. Biogeography and speciation in the Pandorina/Volvulina (Chlorophyta) super clade. Journal of Phycology 37: 836-851.
• Coleman, A.W. 2002. Comparison of Eudorina/Pleodorina ITS sequences of isolates from nature with those from experimental hybrids. American Journal of Botany 89: 1523-1530.
• Conforti, V. 1993a. Study of the Euglenophyta from Camaleão lake (Manaus-Brazil) 1. Trachelomonas Ehr. Révue d'Hydrobiogie Tropicale 26: 3-18.
• Conforti, V. 1993b. Study of the Euglenophyta from Camaleão lake (Manaus-Brazil) 2. Strombomonas Defl. Révue d'Hydrobiogie Tropicale 26: 187-197.
• Conforti, V. 1994. Study of the Euglenophyta from Camaleão lake (Manaus-Brazil) 3. Euglena Ehr., Lepocinclis Perty, Phacus Duj. Révue d'Hydrobiogie Tropicale 27: 3-21.
• Cunha, A.M. 1913. Contribuição ao conhecimento da fauna de protozoários do Brasil, Brasil. Memórias do Instituto Oswaldo Cruz 5: 101-122.
• Cunha, A.M. 1914. Contribuição para o conhecimento da fauna de protozoários do Brasil. Memórias do Instituto Oswaldo Cruz 6: 169-179.
• Daugbjerg, N., Hansen, G., Larsen, J. & Moestrup, Ø. 2000. Phylogeny of some of the major genera of dinoflagellates based on ultrastructure and partial LSU rDNA sequence data, including the erection of three new genera of unarmored dinoflagellates. Phycology 39: 302-317.
• Desnitski, A. 2000. Development and reproduction of two species of the genus Volvox in a shalow temporary pool. Protistology 1: 195-208.
• Desnitski, A. 2002. Dormant stages of the green flagellate Volvox in a natural habitat. Russian Journal Developmental Biology 33: 107-109.
• Desnitski, A. 2003. A review on green microalgae of the genus Pleodorina (in Russian with English summary). Vestnik Sankt-Peterburgskogo Universiteta 3: 98-102.
• Dewes, J. & Huszar, V.L.M. 1986. Influência dos despejos lançados na Sanga da Barbuda sobre as águas do Lago da Barragem Santa Barbara, Pelotas, RS. In: C.E.M. Bicudo, A.C. Teixeira & J.G. Tundisi (eds.): Algas:energia do amanhã. Anais do Simpósio Internacional. Instituto Oceanográfico, São Paulo, p.141-161.
• Domingos, P. & Menezes, M. 1998. Taxonomic remarks on planktonic phytoflagellates in a hypertrophic tropical lagoon (Brazil). Hydrobiologia 369/370: 297-313.
• Ettl, H. 1976. Die Gattung Chlamydomonas Nova Hedwigia, Suppl. 49: 1-1122.
• Ettl, H. 1980. Beitrag zur Kenntnis der Susswasseralgen Danemarks. Svensk Botanisk Tidskrift 74: 179-222.
• Ettl, H. 1983. Chlorophyta, I. Phytomonadina. In: H. Ettl, J. Gerloff, H. Heynig & D. Mollenhauer (eds.). Süsswasserflora von Mitteleuropa, Band 9. Gustav Fischer, Stuttgart.
• Finley, B.J. 2004. Protist taxonomy: an ecology perspective. Philosophical Transactions Royal Society of London, B, 359: 599-610.
• Finlay, B.J. & Fenchel, T. 2002. Microbial eukaryote species. Science 297: 337.
• Finlay, B.J. & Fenchel, T. 2004. Cosmopolitan metapopulations of free-living microbial eukaryotes. Protist 155: 237-244.
• Fenchel, T. 2005. Cosmopolitan microbes and their "cryptic" species. Aquatic of Microbiology Ecology 41: 49-54.
• Fenchel, T. & Finlay, B.J. 2003. Is microbial diversity fundamentally different from biodiversity of larger animals and plants? European Journal of Protistology 39: 486-490.
• Foissner, W. 2006. Biogeography and dispersal of micro-organisms: a review emphasizing protists. Acta Protozoologica 45: 111-136.
• Franceschini, I.M. 1992. Algues d'eau douce de Porto Alegre, Brésil (les Diatomophycées exlues). BibliothecaPhycologica 92: 1-81.
• Gerloff, J. 1940. Beitrage zur Kenntnis der variabilität und Systematik der Gattung Chlamydomonas Archiv für Protistenkunde 94: 311-500.
• Gessner, F. 1931. Volvulina Playfair aus dem Amazonas. Archiv für Protistenkunde 74: 259-261.
• Goldstein, M. 1964. Speciation and mating behavior in Eudorina Journal of Protozoology 11: 317-344.
• Golfari, L. & Moosmayer, H. 1980. Manual de reflorestamento do Estado do Rio de Janeiro. Banco de Desenvolvimento do Estado do Rio de Janeiro. Secretaria Planejamento e Coordenação Geral, Governo do Estado do Rio de Janeiro, Rio de Janeiro.
• Hoham, R.W., Bonome,T.A., Martin, C.W. & Leebens-Mack, J.H. 2002. A combined 18S rDNA and rbcL phylogenetic analysis of Chloromonas and Chlamydomonas (Chlorophyceae, Volcocales) emphasizing snow and other cold-temperature habitats. Journal of Phycology 38: 1051-1064.
• Huber-Pestalozzi, G. 1961. Das Phytoplankton des Süsswassers: Systematik und Biologie. Chlorophyceae. Teil 5. Volvocales. In: A. Thienemann (ed.). Die Binnengewässer, Band 16. E. Schweizerbart'sche Verlagsbuchandlung, Stuttgart.
• Huszar, V.L.M. 1984. Contribuição ao conhecimento das algas planctônicas do Lago da Barragem Santa Bárbara, Pelotas, Rio Grande do Sul, Brasil. Phycologia latinoamericana 2: 169-201.
• Huszar, V.L.M. 1985. Algas planctônicas da Lagoa de Juturnaíba, Araruama, RJ, Brasil. Revista Brasileira de Botânica 8: 1-19.
• Huszar, V.L.M., Silva, L.H.S. & Nogueira, I.S. 1987. Fitoplâncton de rede da Lagoa de Cima, Campos (Rio de Janeiro, Brasil): uma contribuição a seu conhecimento. Neritica, Suppl. 2: 75-104.
• Kammerer, G. 1938. Volvocalean und Protococcalen aus dem unteren Amazonasgebiet. Sitzungsberichte der Akademie der Wissenschaften Wien, sér. math. 147: 183-228.
• Logares, R.E. 2006. Does the global microbiota consist a few cosmopolitan species? Ecología Austral 16: 85-90.
• Luo, W., Pflugmacher, S., Pröschold, T., Walz, N. & Krienitz, L. 2006. Genotype versus phenotype variability in Chlorella and Micractinium(Chlorophyta, Trebouxiophyceae). Protist 157: 315-333.
• Menezes, M. 1994. Fitoflagelados pigmentados de quatro corpos d'água da região sul do município do Rio de Janeiro, Estado do Rio de Janeiro, Brasil. Tese de Doutorado, Universidade de São Paulo, São Paulo.
• Menezes, M. 1996. New species of pigmented flagellates from southeastern Brazil. Archiv für Protistenkunde 147: 101-105.
• Menezes, M. 1999. Flora ficológica da Quinta da Boa Vista, Rio de Janeiro, Brasil: taxonomia e estratégias de Chlorophyceae flageladas em um lago artificial com déficit hídrico. Hoehnea 26: 107-120.
• Menezes, M. & Domingos, P. 1994. La flore planctonique d'une lagune tropicale (Brésil). Revue D'hydrobiologie Tropicale 27: 273-297.
• Menezes, M. & Huszar, V.L. 1997. Bitrichia amazonica,a new species of Chrysophyceae (Stylococcaceae) from Amazonian region, northern Brazil. Algological Studies 85: 13-22.
• Nakazawa, A., Krienitz, L. & Nozaki, I. 2001. Taxonomy of the unicellular green algal genus Vitreochlamys (Volvocales), based on comparative morphology of cultured material. European Journal of Phycology 36: 113-128.
• Nozaki, H. 1982. Morphology and reproduction of Japanese Volvulina steinii (Chlorophyta, (Volvocales). Journal of Japanese Botany 57: 105-113.
• Nozaki, H. 1986. Notes on microalgae in Japan (10). Eudorina illinoisensis (Chlorophyta, Volvocales). Japanese Journal of Phycology 34: 114.
• Nozaki, H. 2003. Origin and evolution of the genera Pleodorina and Volvox (Volvocales). Biologia 58: 425-431.
• Nozaki, H., Hara, Y. & Kasaki, H. 1987. Light and electron microscopy of pyrenoids and species delimitation in Volvulina (Chlorophyta, Volvocaceae). Journal of Pycology 23: 359-364.
• Nozaki, H. & Itoh, M. 1994. Phylogenetic relationships within the colonial Volvocales (Chlorophyta) inferred from cladistic analysis based on morphological data. Journal of Phycology 30: 353-365.
• Nozaki, H, Otha, N., Morita, E. & Watanabe, M.M. 1998. Toward a natural system of species in Chlorogonium(Volvocales, Chlorophyta): a combined analysis of morphological and rbcL gene sequence data. Journal of Phycology 34: 1024-1037.
• Nozaki, H & Krienitz, L. 2001. Morphology and phylogeny of Eudorina minodii (Chodat) Nozaki et Krienitz, comb. nov. (Volvocales, Chlorophyta) from Germany. European Journal of Phycology 36: 23-28.
• Nozaki, H. & Kuroiwa, T. 1991. Morphology and sexual reproduction of Gonium multicoccum (Volvocales, Chlorophyta) from Nepal. Phycologia 30: 381-393.
• Nozaki, H. & Kuroiwa, T. 1992. Ultrastructure of the cellular matrix and taxonomy of Eudorina, Pleodorina and Yamagishiella gen. nov. Phycologia 31: 529-541.
• Nozaki, H., Ito, M., Sano, R., Uchida, H., Watanabe, M.M., Takahashi, H. & Kuroiwa, T. 1997.Phylogenetic analysis of Yamagishiella and Platydorina (Volvocaceae, Chlorophyta) based on rbcL gene sequences. Journal of Phycology 33: 272-278.
• Nozaki, H., Misawa, K., Kajita, T., Kato, M., Nohara, S. & Watanabe, M.M. 2000. Origin and evolution of the colonial Volvocales (Chlorophyceae) as inferred from multiple, chloroplast gene sequences. Molecular Phylogenetics and Evolution 17: 256-268.
• Nozaki, H., Ohta, N., Morita, E. & Watanabe, M.M. 1998. Toward a natural system of species in Chlorogonium (Volvocales, Chlorophyta): a combined analysis of morphological and rbcl gene sequence data. Journal of Phycology 34: 1024-1037.
• Nozaki, H., Ott, D.F. & Coleman, A. 2006. Morphology, molecular, phylogeny and taxonomy of two new species of Pleodorina (Volvocaceae, Chlorophyceae). Journal of Phycology 42: 1072-1080.
• Peterfi, L.S. 1965. Date noi la cunoasterea algelor din R.P. Romana. Strudii si Cercetari in Biologice, Seria Botanica 17: 269-280.
• Peterfi, L.S. 1968. Studies on the taxonomy and ecology of the Romanian Volvocales 2. Nova Hedwigia 16: 216-249.
• Pocock, M.A. 1933a. Volvox and associated algal from Kimberley. Annals of the South African Museum 16: 473-521.
• Pocock, M.A. 1933b. Volvox in South Africa. Annals of the South African Museum 16: 523-646.
• Pocock, M.A. 1960. Haematococcus in Southern Africa. Transactions of the Royal Society of South Africa 36: 1-55.
• Pollio, A., Cennamo, P., Ciniglia, C., De Stefano, M., Pinto, G. & Huss, V.A.R. 2005. Chlamydomonas pitschmannii Ettl, a little known species from thermoacidic environments. Protist 156: 287-302.
• Pröschold, T., Marina, B. Schlösser, U.G. & Melkonian, M. 2001. Molecular phylogeny and taxonomic revision of Chlamydomonas (Chlorophyta). I. Emendation of Chlamydomonas Ehrenberg and Chloromonas Gobi, and description of Oogamochlamys gen. nov. and Lobochlamys gen. nov. Protist 152: 265-300.
• RADAMBRASIL. 1983. Levantamento de recursos naturais. Ministério das Minas e Energia, Rio de Janeiro.
• Reynolds, C.S. 1997. Vegetation processes in the pelagic: a model for ecosystem theory. Excellence in Ecology 9. Ecology Institute, Oldenford.
• Round, F.E. 1981. The ecology of algae. Cambridge University Press, Cambridge.
• Sant'Anna, C.L., xavier, M.B. & Sormus, L. 1988. Estudo qualitativo do fitoplâncton da Represa da Serraria, Estado de São Paulo, Brasil. Revista Brasileira de Biologia 48: 83-102.
• Scherffel, A. 1912. Zwei neue trichocystenartige Bildungen führende Flagellaten. Archiv für Protistenkunde 27: 94-128.
• Skuja, H. 1948. Taxonomie des Phytoplanktons einiger Seen in Uppland, Schweden. Symbolae Botanicae Upsaliensis 9: 1-399.
• Stankiewicz, E.H. 1980. Flórula no conteúdo estomacal do Pseudocurimata gilberti (Quoy & Gaimard, 1824). Dissertação de Mestrado, Universidade Federal do Paraná, Curitiba.
• Starr, R.C. 1962. A new species of Volvulina Playfair. Archiv für Microbiologie 42: 130-137.
• Tiffany, L.H. 1935. Homothallism and other variations in Pleodorina californica Shaw.Archiv für Protistenkunde 85: 140-144.
• Uherkovich, G. & Rai, H. 1979. Algen aus den Rio Negro und seinen Nebenflüssen. Amazoniana 6: 611-638.
• Uherkovich, G. & Schmidt, G.W.F. 1974. Phytoplankton taxa in dem Zentralamazonischen Schwemm landsee Lago do Castanho. Amazoniana 5: 243-283.
• Znachor, P. & Jezberova, J. 2005. The occurrence of a bloom-forming green alga Pleodorina indica (Volvocales) in the downstream reach of the River Malse (Czech Republic). Hydrobiologia 541: 221-228.

Publication Dates

• Publication in this collection
  13 May 2013
• Date of issue
  2008

History

• Accepted
  18 Sept 2008
• Received
  17 Oct 2007