Comparing Grazing on Lake Seston by Dreissena and Daphnia: Lessons for Biomanipulation

Biomanipulation measures in lakes, taken to diminish algal blooms, have mainly been restricted to the reduction of zooplanktivorous fish with the aim to stimulate the grazing pressure by native filter feeders such as Daphnia. However, larger filter feeders like the exotic zebra mussel, Dreissena polymorpha, have been suggested as an optional tool because of their high filtering capacity. We compared grazing by two filter feeders, D. polymorpha and Daphnia galeata, offered seston from Lake IJsselmeer, the Netherlands in two consecutive years: 2002 and 2003. The seston in both years was dominated by the colony-forming cyanobacterium Microcystis aeruginosa. The grazing studies were performed under controlled conditions in the laboratory and samples were analyzed on a flow cytometer, making it possible to quantify grazing on different seston components and size fractions, including cyanobacteria, other phytoplankton (green algae, diatoms, etc.), and detritus. No differences in clearance rates, on a per weight basis, were found between the two grazer species. The clearance rate on cyanobacteria (especially <20 μm) was lower in 2003 than in 2002. In 2003, the microcystin concentration of cyanobacteria was higher than in 2002, suggesting that the observed lower clearance rate in 2003 was due to the enhanced toxin content of the cyanobacteria. Zebra mussels, although indiscriminately filtering all seston groups out of the water, positively selected for phytoplankton in their mantle cavity, irrespective of its toxicity, and rejected detritus. Since no differences in clearance rates were found between the two grazer species, we conclude that for biomanipulation purposes of shallow lakes, native species like the daphnids should be preferred over exotic species like zebra mussels. When the seston is dominated by phytoplankton that cannot be filtered out of the water column by Daphnia, however, the use of zebra mussels may be considered. Care should be taken, however, in the choice of the lakes since the mussels may have severe ecological and economic impacts.     

Adaptive strategies of desmids in two tropical monomictic lakes in southeast Brazil: do morphometric differences promote life strategies selection?

Adaptive strategies of desmids in two tropical monomictic lakes were studied in relation to morphometric differences. The study aimed at evaluating desmids' strategies during inter-annual succession patterns in two tropical lakes (Dom Helvécio and Carioca) in southeast Brazil. From monthly samples of freshwater algae collected during five consecutive years (2002–2006) from the two lakes, the desmids were classified according to size, life form, and life strategies (competitive, stress tolerant, and ruderal). Our results indicated that Lake Carioca (19º45′20″S, 42º37′12″W), a small, shallow, turbid, and mesotrophic system showed dominance of invasive/opportunistic taxa (C) and R-strategists (Staurastrum species <20 μm), whereas Lake Dom Helvécio (19º45′–19º48′45″S, 42º33′45″W), a large, deep, and clear system was dominated by ruderal (R) species (Closterium aciculare and Pleurotaenium trabecula >50 μm) and tolerant to stress (S) species (Staurastrum and Staurodesmus species >20 μm). A significant number of desmids from the lakes had low MLD (≤20 μm), low volume (<10³), and high SV^?1 ratio (>1). Consequently, C-strategist species were abundant and formed a group having great adaptive value in the lakes, with higher resilience and capacity to change composition after the turnover period. Stratification stability and atelomixis were the main driving forces to sustain high desmid biomass in these two lakes, despite the greater macrophyte richness and great water column depth at Lake Dom Helvécio and the main factor influencing richness and contribution of complex forms.     

11. The Indo-Malaysian North-Australian phycogeographical region revised

The present knowledge of the freshwater algal flora in the Indo-Malaysian North Australian region is reviewed. More than 4700 taxa have been recorded from this region. Desmids, diatoms and cyanobacteria are among the better studied groups of organisms, while phytoflagellates have received very little attention. Phytoplankton communities in tropical lakes are generally similar to summer communities of temperate lakes. In addition, there is a large number of typical tropical taxa, including pantropical and regional endemic elements. Local endemism occurs in the old Tertiary lakes of the region. The composition of algal communities changes markedly along an altitudinal gradient, and tropical taxa are gradually replaced by taxa characteristic for cool climatic conditions. Biogeographical distribution patterns are exemplified for the desmids and diatoms. Among the more than 2680 desmids recorded from the Indo-Malaysian North Australian region, about 800 have never been found elsewhere. Dispersal by migratory birds and past climatic changes might explain distribution patterns. Because of incomplete taxonomy and a general lack of understanding of the autoecology, distribution and speciation of freshwater algae, however, there remain serious obstacles for detailed biogeographical analyses.     

THE SEASONAL DISTRIBUTION,ABUNDANCE AND DIVERSITY OF DESMIDS (CHLOROPHYTA) IN A SOFTWATER,NORTH TEMPERATE STREAM1

Suspended and benthic algal communities from a mildly acidic, third-order Rhode Island stream were examined to determine the seasonal distribution, abundance and diversity of the lotic desmids. Within a one-year sampling period, 148 species and 202 subspecific taxa of desmids were identified, representing 23 genera. Species of Cosmarium and Closterium accounted for approximately 70% of the desmids present, and were the most diverse and abundant taxa during all seasons except spring, when Hyalotheca dissiliens was the dominant desmid species. Average abundance and species richness generally were greatest during summer for both suspended and benthic desmids. Most desmids occurred in benthic habitats, and were randomly distributed among substrata. Average seasonal abundance was 7.4 × 10⁴ cells·g^?1 dry wt substratum, among 13 types of substrata. Highest desmid abundance was measured among substrata with intricate morphologies, such as Fontinalis spp., which was associated with 1.2 × 10⁶ desmid cells·g^?1 dry wt substratum, or 1.7 × 10³ cells·cm^?2 substratum. Cell division was observed for 70 desmid taxa, and average seasonal reproduction (based on cell numbers) among all substrata ranged from 4% in winter to 20% during summer. In addition, sexually produced zygospores were found occasionally for H. dissiliens. Desmids were distributed among most substrata examined in this stream, with abundance comparable to reported estimates from softwater lakes and acid bogs. In contrast to established dogma, lotic desmids are not incidental drift organisms, but rather comprise a viable and persistent component of the stream periphyton.     

Phytoplankton assemblages and steady state in deep and shallow eutrophic lakes – an approach to differentiate the habitat properties of Oscillatoriales

Ecological conditions and phytoplankton succession in two shallow hypertrophic lakes (Langer See and Melangsee) and a dimictic, eutrophic lake (Scharmützelsee) in a lake chain in Eastern Germany were analyzed from 1999 to 2001 in order to find situations of phytoplankton steady state assemblages and variables controlling the phytoplankton composition according to Reynolds et al. (2002). Long term background data from 1993 to 2001 suggest steady state conditions in shallow lakes, whereas the deep lake exhibited irregular fluctuations between various phytoplankton stages. Since the phytoplankton composition in the shallow lakes was similar in all the 3 years, it was highly predictable. Steady state conditions dominated by different species of Oscillatoriales were detected during the summer period 1999 and 2000 in Langer See and in Melangsee (see Mischke & Nixdorf, this volume). This dominant assemblage found in both lakes (group S ₁ acc. to Reynolds et al., 2002): Planktothrix agardhii (Gom.) Anagn. et Kom., Limnothrix redekei (Van Goor) Meffert, Pseudanabaena (Lauterb.) is typical in turbid mixed layers with highly light deficient conditions, but it is also regularly dominant in the dimictic lake Scharmützelsee as observed in 1999 and 2001 (Pseudanabaena limnetica (Lemm.) Kom. The Nostocales Cylindrospermopsis raciborskii (Wolz.) Seenayya et Subba Raju and Aphanizomenon gracile (Lemmerm.) Lemmerm. were important in the shallow lakes as well as in lake Scharmützelsee. Nevertheless, the occurrence of filamentous cyanobacteria in the dimictic lake was not regular and an unpredictable change in phytoplankton development was observed in 2000. It is discussed, whether this phenomenon of regular succession in shallow hypertrophic lakes is caused by adaptation to a resilient and an extreme environment or by the pool of species that can live or survive in that environment. This was checked through comparison of the depth of the mixed layer, the mean daily irradiance within this layer and the nutrient resources. Although the nutrient resources in both types of lake are near threshold levels, indicating growth inhibition by dissolved nutrients (DIP, DIN, TIC, DSi), the under water light supply seems to be the key factor favoring the dominance of filamentous cyanobacteria belonging to the functional group S ₁.     

Taxonomic and ecological characterization of cyanobacteria from some brackish and saline lakes of Southern Transbaikal Region

The species composition of cyanobacteria from 12 lakes of Southern Transbaikal Region was studied. In the studied lakes, a total of 28 species of cyanobacteria were detected, including 10 species previously not retrieved from the lakes of Southern Transbaikal Region. The morphological and ecophysiological characteristics, as well as the phylogenetic position, of pure cyanobacterial cultures were determined. According to the results of 16S rRNA sequencing, the Cya 1, Cya 2, and Cya 10 cultures were identified as representatives of the genus Phormidium; the Cya 5 and Cya 6 cultures were assigned to the genus Nodularia. On the basis of its morphological properties, the Cya 4 culture was identified as Pseudanabaena frigida. The studied microorganisms were moderate alkaliphiles and could grow within a broad salinity range (0–100 g/l NaCl).     

Biological soil crusts determine the germination and growth of two exotic plants

In arid and semiarid ecosystems, the potential threats of exotic invasive species are enhanced due to increasing human activities. Biological soil crusts (BSCs), acting as arid ecosystem engineers, may play an important role in preventing the establishment of exotic invasive plants. Our goal was to examine whether BSCs could inhibit the establishment of probable exotic plant species originating from adjacent grasslands located along the southeast edge of the Tengger Desert. In our study, we investigated the effects of three BSC types (cyanobacteria, lichen, and moss crusts) under two disturbance conditions (intact and disturbed) on the establishment of two exotic plant species (Ceratoides latens and Setaria viridis) using indoor experiments. We found both negative and positive effects of BSCs on the establishment of the two exotic plant species. Compared with the disturbed BSCs, the germination percentages of C. latens and S. viridis were reduced by 54% to 87% and 89% to 93%, respectively, in intact BSCs. In contrast, BSCs significantly promoted the height growth and aboveground biomass of the two exotic plant species (p < .05) by enhancing the soil water and nutrient availability for the exotic plants. Our results confirm that BSCs strongly suppress the rapid expansion of exotic plant populations by inhibiting germination of seed with big size or appendages and have a weak inhibitory effect on exotic plant with small and smooth seeds. This may decrease the threat of propagation of exotic species. In the meantime, BSCs promote the growth of a few successful engraftment seedlings, which increased the beta diversity. Our work suggests that better understanding the two opposing effects of BSCs on the establishment of exotic plant species in different growth stages (germination and growth) is important for maintaining the health and stability of revegetated regions.     

Invasion of exotic piscivores causes losses of functional diversity and functionally unique species in Japanese lakes

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Higher temperatures enhance the effects of invasive sportfish on mountain zooplankton communities

1. Decades of introductions of exotic sportfish to mountain lakes around the world have impoverished them biologically, and this may be exacerbated by global warming. We assessed the current status of invasive salmonids and native zooplankton communities in 34 naturally fishless lakes along an elevational gradient, which served as an environmental proxy for the expected effects of climate change. 2. Our main goal was to explore how climate‐related variables influence the effects of stocked salmonids on the total biomass, species richness and taxonomic composition of zooplankton. We predicted that warmer conditions would dampen the negative predatory effects of exotic brook trout (Salvelinus fontinalis) on zooplankton communities because more temperate lakes contain a greater diversity of potentially tolerant species. 3. Instead, we discovered that the persistence of stocked brook trout in the warmer lakes significantly amplified total zooplankton biomass and species richness. In colder and deeper lakes, zooplankton were relatively unaffected by S. fontinalis, which however persisted better in alpine lakes than at lower elevations after stocking practices were halted over two decades ago. Warmer lake conditions and higher concentrations of dissolved organic carbon (DOC) were significant primary drivers of zooplankton species turnover, both favouring greater species diversity. 4. Our findings of an ecological surprise involving potential synergistic positive effects of climate warming and exotic trout on native zooplankton communities presents a conundrum for managers of certain national mountain parks. Present mandates to eradicate non‐native trout and return the mountain lakes to their naturally fishless state may conflict with efforts to conserve biodiversity under a rapidly changing climate.     

Eutrophication mediates rapid clonal evolution in Daphnia pulicaria

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New insights into the distribution of alien cyanobacterium Chrysosporum bergii (Nostocales,Cyanobacteria)

Invasive and alien cyanobacteria are considered as a serious threat to aquatic ecosystems due to their ability to produce cyanotoxins and outcompeting native species. Among cyanobacteria, Nostocales is a group with strong competitive advantages including the production of resting cells, ability to fix nitrogen or high affinity to phosphorus. A species Chrysosporum bergii has broadened its original distribution from brackish and seawater habitats of Caspian and Aral Seas to northern regions of Europe. The aim of our study was to expand the knowledge on the distribution of C. bergii, its preferred habitats in the invaded areas and examine whether the biomass of C. bergii is related with biomass of other alien or native cyanobacteria. We examined the phytoplankton community composition and the abiotic factors in randomly selected lakes of western Poland. For the first time, we showed that it occurs widely in lakes of this region. However, it occurred in one third of the investigated lakes and its contribution to total phytoplankton biomass was low. It occurred more frequently in shallow lakes with low conductivity and dissolved phosphorus, yet with higher total phosphorus concentration. The most remarkable finding was a negative relationship of C. bergii biomass with the biomass of native A. gracile, but a positive relationship with an invasive cyanobacterium C. raciborskii. This result suggests that these alien species occur under similar environmental conditions in Polish lakes. Moreover, the morphological features of C. bergii in Poland were similar to those of Anabaena bergii var. limnetica Couté et Preisig from Lithuania and Anabaena bergii from Czech Republic.     

Temporal and spatial macrofaunal community changes along a salinity gradient in seagrass meadows of Akkeshi-ko estuary and Akkeshi Bay,northern Japan

Changes in water clarity (secchi disc transparency) in relation to the presence/absence of introduced, exotic fish, including rudd (Scardinius erythrophthalmus), tench (Tinca tinca), perch (Perca fluviatilis), brown bullhead catfish (Ameiurus nebulosus), goldfish (Carassius auratus), and koi carp (Cyprinus carpio) were determined for 49 small, North Island, New Zealand lakes. There was a negative association between water clarity and the presence of exotic fish independent of lake depth. Moreover, a ‘before-and-after’ comparison and examination of case-studies indicated that introductions of exotic fish reduce water clarity. The number of species introduced affected the relationship between lake depth and water clarity but the specific role of each species could not be distinguished because most of the lakes (83%) contained more than one exotic fish species. A model incorporating the known mechanisms by which planktivorous, benthivorous and herbivorous fish can influence water clarity in lakes showed that control over just one species or feeding guild may not result in an improvement in water clarity because of the additive and synergistic effects of different species on lake trophic processes. Handling editor J. Cambray     

Desmids in mountain lakes of the subpolar Urals

The species composition of desmids was studied in eight mountain lakes of the Kozhim and Malyi Patok River basins in the “Yugyd Va” National Park (subpolar Urals, Russia). A total of 98 desmid taxa representing 13 genera belonging to 4 families were identified. Geographical and ecological analyses showed that the desmid flora is typical, with a predominance of cosmopolitan species, planktic-benthic forms, acidophilic and pH indifferent species, and halophobic to salinity indifferent species. The investigated lakes can be classified as pristine and in good ecological condition on the basis of their hydrochemical and algal biodiversity characteristics. Presented at the International Symposium Biology and Taxonomy of Green Algae V, Smolenice, June 26–29, 2007, Slovakia.     

Alkaline soda Lake Velika Rusanda (Serbia): the first insight into diatom diversity of this extreme saline lake

Alkaline soda lakes are unique habitats found in specific geographic regions, usually with dry climate. The Carpathian Basin is one of those regions very important for habitat and biodiversity conservation in Europe, with natural soda lakes found in Austria, Hungary and Serbia. In comparison to other two countries from Central Europe, algal biodiversity studies of saline soda lakes in Serbia are scarce. Lake Velika Rusanda has the highest measured salinity of all saline lakes in the Carpathian Basin and there were no reports of its diatom species richness and diversity till now. We conducted 2-year investigation programme to study biodiversity and seasonal dynamics of diatoms in this lake. A total of 27 diatom taxa were found, almost all of them attached to reed and much less in benthos and plankton. Five new diatom species for Serbia were recorded, Craticula halopannonica, Navicymbula pusilla, Hantzschia weyprechtii, Nitzschia thermaloides and Navicula staffordiae. The last mentioned is new for Europe as well. Lake Velika Rusanda is inhabited mostly by alkaliphilous and halophilic diatoms. Since diatoms are used as bioindicators in soda lakes, our results will improve their further application in ecological status assessment of these fragile habitats in the Carpathian Basin.

     

An elementary,structural analysis of river phytoplankton

Summary A structural analysis of river phytoplankton has been carried out based upon published studies on 67 rivers. When available on a yearly basis to account for seasonal variability, five structural features have been chosen: species composition, species richness, species dominance, diversity and biomass (total and per taxonomic groups). Despite the high number of reported studies, most of them cover only some of the aforementioned features. As a result of the low amount of studies, tropical rivers are underepresented. No size distribution studies have been carried out on river phytoplankton. The average species richness amounts to 126, being higher in temperate rivers. Roughly one half of each flora is comprised of sporadic species. No statistically significant relationship between species richness and latitude has been found despite the fact that tropical rivers appear to house fewer species than temperate rivers. Also, one half of the support in the floras are either benthic or tychoplanktonic. Diatoms comprise the majority of species numbers in the whole data set but are substituted by desmids in tropical rivers and by green algae when benthic species are not taken into account. There appears to be lower biomass in river phytoplankton than in lakes. Diatoms are also the major taxonomic group comprising total biomass in rivers but they share clearly a lower fraction in tropical rivers. On an average basis, diatoms appear to be more dominant in rivers than in lakes. The time course of diatom dominance occurs close to the summer solstice in tropical rivers whereas is much more lagged in temperate sites. The diversity of river phytoplankton is highly scattered (0.40–4.40 bits ind^–1).     

The microbial plankton of freshwater lakes in the Vestfold Hills,Antarctica

Summary The plankton of twelve freshwater and slightly saline lakes in the Vestfold Hills, Antarctica was sampled in February 1991. All of the lakes are oligotrophic. The chlorophyll a concentrations in the lakes ranged from 0.10–2.69 g · 1^–1. The majority of the phytoplankton were flagellates or picoplanktonic cyanobacteria with the species composition varying between the lakes. Cyanobacteria were found in five of the lakes. Five to 6 species of ciliated protozoa occurred, among them oligotrichs, including the mixotrophic species Strombidium viride. The concentrations of protists and bacteria were an order to several orders of magnitude lower than reported from lower latitude oligotrophic lakes. Low species diversity and low numbers in the plankton characterise these eastern Antarctica lakes which reflects their low nutrient status and isolation.     

Primo contributo alla conoscenza dei laghi appenninici meridionali di alta quota. Il fitoplancton dei laghi del massiccio del Monte Sirino (Appennino Lucano)

Abstract

A preliminary account on some mountain lakes of the Southern Apennines. Phytoplankton investigation on three ponds of the Monte Sirino (Basilicata).—Phytoplankton standing crop and species composition have been studied in three ponds of the Southern Apennines. The observations carried out throughout ten months in 1975, showed a great number of species present in the Remmo pond, mainly diatoms, desmids, green algae, and blue-green algae. In the Sirino pond only diatoms and Chrysophyceae were present.

In comparison, in the Zapano pond, the phytoplankton was scarcely represented but the occurrence of some species of green algae may suggest a eutrophic water condition. On the basis of the peculiarities both of species composition and biomass, the major water bodies may be considered oligotrophic.     

Invasion biology in non‐free‐living species: interactions between abiotic (climatic) and biotic (host availability) factors in geographical space in crayfish commensals (Ostracoda,Entocytheridae)

In invasion processes, both abiotic and biotic factors are considered essential, but the latter are usually disregarded when modeling the potential spread of exotic species. In the framework of set theory, interactions between biotic (B), abiotic (A), and movement‐related (M) factors in the geographical space can be hypothesized with BAM diagrams and tested using ecological niche models (ENMs) to estimate A and B areas. The main aim of our survey was to evaluate the interactions between abiotic (climatic) and biotic (host availability) factors in geographical space for exotic symbionts (i.e., non‐free‐living species), using ENM techniques combined with a BAM framework and using exotic Entocytheridae (Ostracoda) found in Europe as model organisms. We carried out an extensive survey to evaluate the distribution of entocytherids hosted by crayfish in Europe by checking 94 European localities and 12 crayfish species. Both exotic entocytherid species found, Ankylocythere sinuosa and Uncinocythere occidentalis, were widely distributed in W Europe living on the exotic crayfish species Procambarus clarkii and Pacifastacus leniusculus, respectively. No entocytherids were observed in the remaining crayfish species. The suitable area for A. sinuosa was mainly restricted by its own limitations to minimum temperatures in W and N Europe and precipitation seasonality in circum‐Mediterranean areas. Uncinocythere occidentalis was mostly restricted by host availability in circum‐Mediterranean regions due to limitations of P. leniusculus to higher precipitation seasonality and maximum temperatures. The combination of ENMs with set theory allows studying the invasive biology of symbionts and provides clues about biogeographic barriers due to abiotic or biotic factors limiting the expansion of the symbiont in different regions of the invasive range. The relative importance of abiotic and biotic factors on geographical space can then be assessed and applied in conservation plans. This approach can also be implemented in other systems where the target species is closely interacting with other taxa.     

Comparative analysis of rotifer community structure in five subtropical shallow lakes in East China: role of physical and chemical conditions

Worldwide, there have been few comparative studies on rotifer communities in subtropical lakes. We studied changes in rotifer community structure over 1 year and its relationship to several physicochemical variables in five subtropical shallow lakes in East China, covering a nutrient gradient from mesotrophy to moderate eutrophy. In these lakes, the genera Brachionus, Lecane, and Trichocerca dominated the rotifer species composition, and Polyarthra dolichoptera, Keratella cochlearis, Filinia longiseta, T. pusilla, and Anuraeopsis fissa were the dominant species. With increased nutrient loading, total rotifer abundance and species dominance increased, indicating that rotifer abundance might be a more sensitive indicator of trophic state than species composition. Comparative analyses of the six rotifer community indices calculated in this study and redundancy analysis (RDA) revealed that the two slightly eutrophic lakes and the other two moderately eutrophic lakes exhibited a high degree similarity in community structure. This suggests that the trophic state of a lake determines the rotifer community structure. In contrast, in the two moderately eutrophic lakes, the mass ratios of TN:TP and the contents of TP suggested N-limitation and cyanobacteria dominance in phytoplankton communities might be possible. In these lakes TN played a more important role in shaping the rotifer community according to stepwise multiple regression and RDA. RDA analysis also suggested that rotifer species distribution was strongly associated with trophic state and water temperature, with water temperature being the most important factor in determining seasonality.     

Steady state of phytoplankton assemblage in the tropical Lake Catemaco (Mexico)

Phytoplankton of the tropical lake Catemaco (Veracruz, Mexico) showed similar species composition during samplings from 1993 to 1995. There were two small dominant cyanobacterial species Cylindrospermopsis catemaco Kom.-Legn. et Tavera and Cylindrospermopsis philippinensis (Taylor) Kom., and a group of larger algae and cyanobacteria that were always present, however in smaller numbers: Aulacoseira granulata (Ehr.) Simons morphotype curvata, A. cf. italica (Ehr.) Simons mf. curvata, Fragilaria construens (Her.) Grun., Achnanthes minutissima Kütz., Planktolyngbya circumcreta (G.S.West) Anagn. et Kom., Chroococcus microscopicus Kom.-Legn. et Cronberg. Moreover we found several other scarcely present species. The percentage of total biomass of the two dominant species of Cylindrospermopsis varied between 34 and 81%, but they accounted for 80 to 95% of abundance. Apart from geomorphological features and climate conditions, biological variables played an important role. Fish-stock was formed by filter-feeding native herbivorous species of fish Dorosoma petenense (Günther), Bramocharax caballeroi (Contreras et Rivera), Astyanax mexicanus (Filippi), and an introduced, also herbivore Oreochromis niloticus (L.). Feeding activity of fish removed large species of algae and cyanobacteria as well as detrital remnants and zooplankton from the water. Smaller, inedible cyanobacteria remained in the water and formed the stable portion of the phytoplankton, dominant both in biomass and abundance. CANOCO analysis of samples and species variability demonstrated results of competition between two species of Cylindrospermopsis: steady state during the dominance of C. catemaco lasting probably for the whole year 1993 (one dry and one wet season) and steady state during the dominance of C. philippinensis in 1994 and 1995. According to the functional classification of phytoplankton suggested by Reynolds et al. (2002), Catemaco dominant assemblage would belong to the functional group S _N.