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'Chantransia' Stages of Sirodotia Huillensis

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'Chantransia' Stages of Sirodotia Huillensis Blackwell Publishing AsiaMelbourne, AustraliaPREPhycological Research1322-08292006 Japanese Society of PhycologyJune 2006542108115Original ArticleEcology and morphology of Sirodotia huillensisJ. Carmona et al. Phycological Research 2006; 54: 108–115 Ecology and morphological characterization of gametophyte and ‘Chantransia’ stages of Sirodotia huillensis (Batrachospermales, Rhodophyta) from a stream in central Mexico Javier Carmona,1* Gustavo Montejano1 and Orlando Necchi Júnior2 1Phycology Laboratory, A. P. 70–620, Faculty of Sciences, National Autonomous University of Mexico (UNAM), Ciudad Universitaria, Coyoacán 04510, México DF, Mexico; and 2São Paulo State University, Zoology and Botany Department, 2265-15054-000 São José do Rio Preto, São Paulo, Brazil Key words: Batrachospermales, ‘Chantransia’ stages, SUMMARY ecology, gametophyte, morphology, Rhodophyta, Siro- dotia huillensis, stream. The morphology and phenology of Sirodotia huillensis was evaluated seasonally in a central Mexican first- order calcareous stream. Water temperature was constant (24–25°C) and pH circumneutral to alkaline (6.7–7.9), and calcium and sulfates were the dominant INTRODUCTION ions. The gametophyte stages were characterized by the The genus Sirodotia was established by Kylin (1912) presence of a distinctive mucilaginous layer, a marked based on specimens of Sirodotia suecica from Osby, difference in phycocyanin to phycoerythrin ratio Skane in Sweden. Batrachospermum huillense Wel- between female and male plants, and the presence of witsch ex W. et G. S. West (1897) from Lopollo, Huilla, a carpogonia with a large trichogyne (>60 µm). Occa- Angola in Africa was assigned to the genus Sirodotia as sionally three capogonia were observed on a single Sirodotia huillensis by Skuja (1931). It has the typical basal cell. The ‘Chantransia’ stages were morphologi- vegetative morphology of members of the family Batra- cally similar to those described for the other members chospermaceae (Necchi et al. 1993; Kumano 2002), of Batrachospermales. A remarkable observation was but it is distinguished by its asymmetrical carpogonia the formation of dome-shaped structures, consisting of with elongate conical or club-shaped tricogynes and prostrate filaments that are related with the develop- indeterminate carposporophyte filaments extending ment of new gametophytes. Chromosome numbers were over the axial cortical filaments. Species delimitation n = 4 for fascicle cells, cortical filament cells and is based on the size, arrangement and shape of sper- dome-shaped cells, and 2n = 8 for gonimoblast fila- matangia, carpogonia and gonimoblast filaments (Nec- ment cells and ‘Chantransia’ stage filaments. Gameto- chi et al. 1993; Kumano 2002). Molecular tools have phytes and ‘Chantransia’ stages occurred in fast current been used to determine phylogenetic relationships velocities (60–170 cm/s) and shaded (33.1–121 µmol using sequence analyses for ruBisCo large and small 2 photons/m /s) stream segments. The population fluctu- subunits (rbcL, rbcS) genes (Vis & Sheath 1999). Eight ated throughout the study period in terms of percentage species have been recognized within the genus world- cover and frequency: the ‘Chantransia’ stages were wide (Kumano 1982; Necchi et al. 1993; Vis & Sheath most abundant in the rainy season, whereas gameto- 1999): Sirodotia yutakae Kumano, Sirodotia segawae phytic plants had the highest frequency values during Kumano, Sirodotia sinica Jao, Sirodotia geobelii the dry season. These results were most likely a result Entwisle et Foard, Sirodotia delicatula Skuja, Sirodotia of fluctuations in rainfall and related changes in current suecica Kylin, S. huillensis Welwitsch ex West et velocity. Some characteristics of this population can be viewed as probable adaptations to high current veloci- ties: the mucilaginous layer around plants that reduces drag; potential increase in fertilization by the elongate and plentiful trichogynes and abundant dome-shaped structures producing several gametophytes. *To whom correspondence should be addressed. Email: [email protected] Communicating editor: K. Okuda. Received 28 January 2005; accepted 13 November 2005. Ecology and morphology of Sirodotia huillensis 109 G. S. West and Sirodotia gardneri Skuja ex Flint. Only and ASTM (1989). The variation coefficient (VC) was two species have been reported for North America: used to determine whether environmental parameters S. huillensis and S. suecica. Vis and Sheath (1999) were consistent (those that do not change along time conclude that S. suecica and Sirodotia tenuissima are and space in a water body, usually VC, VC < 5–10%) or conspecific and morphological characters used to dis- variable (those that do change along time and space, tinguish between these species are not phylogenetically usually because they are affected by dilution/evapora- informative, because the sequence data for ruBisCo tion process and biological activity, sensu Margalef (rbcL, rbcL-S spacer, rbcS) genes revealed little varia- 1983). A descriptor was calculated by the equation: tion among specimens of the two species. One popula- VC = s/a·100, where s = standard deviation and tion identified as S. huillensis has been reported from a = average. Mexico (Carmona et al. 2004) and corresponds mor- Observations were made on the natural substrata phologically and ecologically with the description of the (boulders) directly on the river bed. Temporal variation species. was monitored using the quadrat technique (Necchi Reproduction, development and life history are et al. 1995; Necchi 1997; Necchi & Branco 1999), poorly known in Sirodotia and few studies have been which evaluates the influence of variables at micro- published. Vegetative and reproductive characteristics habitat level (current velocity, depth and underwater of Sirodotia species are reported by Necchi (1991), irradiance) on the vegetative and reproductive charac- Necchi et al. (1993) and Kumano (2002). Balakrish- teristics of the populations (proportion of both female nan and Chaugule (1975, 1980) found elimination and male plants and of the ‘Chantransia’ stages). Ten cells in the life history of Batrachospermaceae, whereas sampling units were distributed in ten equally spaced Necchi and Carmona (2002) describe somatic meiosis intervals along the stream segment (58 m2). Microhab- and development of juvenile gametophytes. Necchi itat variables were measured in situ at the center of and Sheath (1992) report chromosome numbers for each sampling unit: current velocity and irradiance S. delicatula (as Batrachospermum delicatulum) as were measured as close to the alga as possible using, n = 3, 2n = 6. Ecological studies of the Batrachosper- respectively, a Swoffer 2100 current velocity meter and males focus primarily on Batrachospermum species a Li-Cor LI-1000 quantum meter with a flat subaquatic (Sheath & Hambrook 1990; Hambrook & Sheath 1991; sensor of photosynthetically active radiation. The num- Necchi 1997; Necchi & Branco 1999). S. huillensis ber of plants was recorded within each sampling unit appears to be more abundant in desert and tropical by visual estimate (Necchi 1997), with a 175 cm2 regions, in warm, neutral to alkaline and high ion con- viewfinder. Frequency of occurrence was calculated tent waters (Necchi et al. 1993; Carmona et al. 2004). from the number of sampling units with algal thalli However, nutrient and microhabitat requirements for compared with the total units sampled (10). Twenty this species are virtually unknown. thalli were randomly selected (2–3 in each sampling The present investigation was conducted to describe unit). morphological and cytological details, as well as the Plants were preserved in 3% formaldehyde and Car- environmental conditions favoring the occurrence of noy’s solution for subsequent analysis in the laboratory. gametophytes and the ‘Chantransia’ stages based on For chromosome observations, plants were stained by laboratory culture and field monitoring for one popula- Wittmann’s hematoxylin (Necchi & Sheath 1992; tion of S. huillensis in central Mexico. Sheath & Cole 1993). Chromosome counts were made on the basis of a minimum of 10 observations for each MATERIALS AND METHODS cell type. In addition, extra cellular polysaccharides were stained with 0.3% Alcian-Blue, in 3% acetic acid The material used in the present study was collected at pH 2.5 (Sheath & Cole 1990). For microscopic anal- from the Santa Anita River, a fast-running stream yses and photographic documentation of cytological through a calcareous region at an altitude of 160 m characters, we used an Olympus BX51 microscope, (Montejano et al. 2000) in a tropical region from cen- with an SC35 microphotography system. Morphometric tral Mexico (21°58′N, 99°11′W). Fieldwork was con- characters were measured in 20 replicates; the number ducted from November 2003 to May 2004, including of replicates was determined using the equation: n = the most important parts of the seasonal cycle, the (s/Eξ)2, where s = standard deviation, E = standard rainy and the dry season. One sampling was carried out error (0.05 and ξ= average (Southwood 1978). at the end of the rainy season (November) and two Cultures were initially obtained from germination of during the dry season (January and May). The following carposporangia. Isolation into culture followed the pro- environmental variables were measured: temperature, cedures described previously (Necchi & Zucchi 1997). pH and specific conductance with a Conductronic PC- The isolates were kept in 20:1 water–soil
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