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Phenology of Sirodotia Suecica (Batrachospermaceae, Rhodophyta) in a High-Altitude Stream in Central Mexico

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Phenology of Sirodotia Suecica (Batrachospermaceae, Rhodophyta) in a High-Altitude Stream in Central Mexico Phycological Research 2009; 57: 118–126 Phenology of Sirodotia suecica (Batrachospermaceae, Rhodophyta) in a high-altitude stream in central Mexicopre_528 118..126 Javier Carmona,* Miriam Bojorge-García, Yenny Beltrán and Rocio Ramírez-Rodríguez Phycology Laboratory, A. P. 70-620, Faculty of Sciences, National Autonomous University of Mexico (UNAM), Ciudad Universitaria, Coyoacán, 04510, México, D.F. shaped filaments that produce new gametophytes SUMMARY through vegetative meiosis (Necchi 1991; Necchi et al. 1993; Kumano 2002). The studies of Pascoaloto and The morphology and phenology of a monoecious popu- Necchi (1990), Necchi and Branco (1999), Necchi and lation of Sirodotia suecica Kylin was evaluated season- Vis (2005) and Carmona et al. (2006) are the most ally in a central Mexican fifth-order high-altitude comprehensive investigations of the reproductive stream. Abundance of gametophytes is positively cor- success of each phase of the life-history of Sirodotia related with concentrations of total dissolved solids, species. Umezaki (1960) and Necchi (1991) reported specific conductivity and total ionic concentration; monoecious and/or dioecious populations without any being present the whole dry season and disappearing in clear relation with the environmental situation. In dio- the rainy season. The gametophytes grew in eutrophic ecious populations of Sirodotia delicatula Skuja (as circumstances and particular microhabitat conditions: Batrachospermum delicatulum) the presence of game- high current velocity (66–122 cm s-1), low irradiance tophyte is associated with high irradiance and low tem- (75–263 mmol photons m-2 s-1) and shallow depth perature, and synchronous development of female and (7–26 cm). Percent cover of gametophytes ranged from male thalli results in a higher proportion of fertilized 5–90% and significant differences in abundance were thalli (Necchi & Branco 1999). In contrast, dioecious not observed when alga was present. Some morphologi- populations of Sirodotia huillensis Welwitsch ex G.S. cal and reproductive characteristics seem to be adap- West et West are restricted to waters of low irradiance tations to high current velocity: abundant secondary and high temperature, and exhibits a low female: male branches, spermatangia and carpogonia. ‘Chantransia’ thallus ratio and a low number of carposporophytic stage, microscopic creeping filaments associated with thalli (Carmona et al. 2006). Furthermore, the the base of the gametophyte, were not observed in ‘Chantransia’ stage in Sirodotia suecica and S. huillen- natural conditions. In terms of reproductive success, sis are usually conspicuous and occur in different envi- the population studied can be regarded as highly effi- ronmental conditions throughout the year (Necchi cient, considering the high fertilized carpogonia rate, 1997; Carmona et al. 2006). similar to monoecious populations in lotic habitats. Two species of Sirodotia have been reported for However S. suecica was not common in the study region North America, S. suecica and S. huillensis (Necchi because it was restricted to particular microhabitat et al. 1993; Vis & Sheath 1999; Carmona & Vilaclara conditions. 2007). Sirodotia suecica is distinguished by the goni- moblast initial arising from the non-protuberant side of Key words: Batrachospermaceae, ecology, monoecious, the carpogonia (Necchi et al. 1993; Carmona & phenology, Rhodophyta, Sirodotia suecica, stream. Vilaclara 2007) and S. huillensis has the gonimoblast arising from the protuberant side of the carpogonial base (Carmona et al. 2004). Additionally, development and life history are described in S. huillensis (Carmona INTRODUCTION et al. 2006), and are poorly known in S. suecica. One monoecious population identified as S. suecica has The genus Sirodotia Kylin has the typical vegetative been reported from Mexico (Carmona & Vilaclara 2007) morphology of members of the family Batrachosper- maceae: a uniseriate main axis with four to six pericen- tral cells producing repeatedly branched fascicles of limited growth. It has a heteromorphic triphasic life *To whom correspondence should be addressed. history consisting of a gametophyte (monoecious and/or Email: [email protected] dioecious), which produces spermatangia in clusters, Communicating editor: G. Zuccarello. carpogonia with elongated trichogynes and the Received 14 December 2007; accepted 10 September 2008. ‘Chantransia’ stage (sporophyte) consisting of tuft- doi: 10.1111/j.1440-1835.2009.00528.x © 2009 Japanese Society of Phycology Phenology of Sirodotia suecica in Mexico 119 and corresponds morphologically and ecologically to the influence of variables at the microhabitat level the description of this species. Sirodotia suecica (current velocity, depth and underwater irradiance) on appears to have a global distribution, being more abun- the vegetative and reproductive characteristics of the dant in temperate climates, neutral to alkaline pH, low population. The sampling site consisted of a stream specific conductance waters and in clean to polluted segment of 58 m2. Each sampling unit was a circle of conditions (Necchi et al. 1993; Carmona & Vilaclara 10 cm radius (area = 314 cm2). Type and size of sam- 2007). Sirodotia suecica has been collected in streams pling units were defined from preliminary tests and from Europe (Kylin 1912; Israelson 1942; Kwandrans previous research (Necchi et al. 1995; Carmona et al. et al. 2002), North America (Flint 1948; Necchi et al. 2006). Sampling consisted of 10 quadrats, each sepa- 1993; Carmona et al. 2004; Carmona & Vilaclara rated by 1 m and its location was determined by a table 2007), Asia (Umezaki 1960; Kumano 1982) and of random numbers. On all sampling dates the micro- Oceania (Entwisle & Foard 1999). However, the sea- habitat characteristics were recorded. Microhabitat sonal dynamics and microhabitat requirements for this variables were measured in situ at the center of each species are virtually unknown. The present investigation sampling unit: current velocity and irradiance were was conducted to describe the phenology and environ- measured as close as possible to the algae, using a mental conditions which favored the occurrence of Swoffer 3100 current velocity meter and a Li-Cor gametophytes based on field monitoring of a population LI-1000 quantum meter with a flat subaquatic sensor of S. suecica in a high-altitude stream from central of photosynthetically active radiation (PAR), respec- Mexico. tively. The number of thalli (gametophytes) within each sampling unit was recorded by visual estimation 2 MATERIALS AND METHODS (Necchi 1997), with a 175 cm viewfinder. ‘Chantran- sia’ stage was not observed in natural conditions, it was The material used in the present study was collected found during microscopic observations of the gameto- from the Amanalco river; a fifth-order stream segment phyte. A description of the ‘Chantransia’ stage was in a mountainous (elevation 1890 m) region of central realized. Thirty thalli were randomly selected (three in Mexico (19°13′N, 100°07′W). Field work was con- each sampling unit) and preserved in 3% formaldehyde ducted from May 2006 to June 2007, including the for subsequent analysis in the laboratory. The following most contrasting parts of the seasonal cycle; that is, the morphological and phenological characteristics were rainy and dry seasons. Three samplings were carried out defined from preliminary tests and previous research: early, in the middle and at end of the dry season: thalli height; number of secondary branches; whorl December, February and May, respectively (December diameter; axial cell length; number of carpogonia, sper- being the coldest month) and two more during the matangia and carposporangia; the ratio of carpogonia: initial and middle parts of the rainy season (June and fertilized carpogonia (Necchi et al. 1993; Necchi September). Temperature and specific conductance 1997; Entwisle & Foard 1999; Necchi & Branco 1999; were measured with a Conductronic (Puebla, Mexico) Carmona & Vilaclara 2007). PC-18 conductivity meter. Dissolved nutrients (30 mL Five replicates were measured in 1 cm of thallus for was filtered in situ with 0.45 and 0.22 mm pore diam- the calculation of secondary branches and 20 for whorl eter membranes and preserved with chloroform and diameter and axial cell length. To avoid inclusion of frozen) were measured in the laboratory with a multi- incompletely developed reproductive structures, the channel analyzer, following standard titration. Water apical region (top 1 cm) was removed (Necchi & Branco samples for dissolved inorganic nitrogen (DIN) and 1999). Segments were squashed prior to examination, soluble reactive phosphorous (SRP) were kept in cold and spermatangia and carposporangia were counted in conditions until analyses were completed (APHA- replicates of five in an area of 50 mm using a microme- AWWA-WPCF 1980). Water samples for determination ter. The number of carpogonia and fertilized carpogonia of anions and pH were preserved frozen in the dark, were counted around the filament. In addition, mac- whereas samples for cations were preserved with 40% roalgae, diatoms and aquatic invertebrates were iden- nitric acid (pH 2–3). Determination of carbonates was tified from each thallus to examine species associations carried out with the titration method, chlorides by the (Merrit & Cummins 1996; Wiggins 1996). An Olympus selective electrode method, hardness with the titration BX51 microscope with a DP12 microphotography method of ethylenediaminetetraacetic acid
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