Life history of Cladophora surera sp. nov. (Cladophorales, Ulvophyceae)

The life history of Cladophora surera sp. Nov., described from the south of the province of Buenos Aires, Argentina, was found to be diplobiontic and isomorphic with haploid gametophytic (n= 24) and sporophytic (2n= 48) plants. Other freshwater species, namely C, suhriana Kutz, and C. callicoma Kütz. have also been reported to be diplobiontic and isomorphic but differing from C. surera by their ploidy level. Plants exhibit great morphological variation, as the number of branches/mm² tends to increase with higher water velocity, thus adopting morphotypes that resemble very different species, that is, C. vagabunda (L.) Hoek, C. glomerata (L.) Kutz., C, laetevirens (Dillw.) Kutz., C. brasiliana Martens, C. dalmatica Kütz., C. vadorum (Aresch.) Kütz. and C. rivularis (L.) Hoek., but clearly differing from them by other morphological parameters; apical cell diameters, The ploidy level 24/48 has not been established for a species of Cladophora. Autogamy is the normal conjugation method and can take place even inside the gametangia before the gametes are released. Isogametes conjugate in a slow behavioral anisogamy, in which the contents of one migrate to the other. Tetra-flagellate zoospores result from meiosis.     

Strategies providing success in a variable habitat: I. Relationships of environmental factors and dominance of Cladophora glomerata

Dominance of macrophytes and their response to environmental factors were studied in the river Ilm, Thuringia/ Germany with special reference to Cladophora glomerata (L.) Kütz. Macroalgae showed a growth peak in spring with C. glomerata, Ulothrix zonata Kütz., Lemanea fluviatilis (L.) C. Agardh and Audouinella sp. being the dominant species. Shortly after this peak, a rapid decline of macrophyte substrate coverage was observed. Only C. glomerata revealed a second growth peak in late summer/early autumn. Frequent disturbances of the macrophyte assemblage by floods resulted repeatedly in an almost complete wash out of benthic organisms. After summer floods C. glomerata was the species that recolonized the substrate. At high‐light sites, faster recovery of C. glomerata was observed as compared to low‐light sites. This is discussed in relation to the life cycle of C. glomerata. Among the physical and chemical parameters that were analysed, irradiance, current velocity, pH, soluble reactive phosphorus and ammonia‐nitrogen accounted for most of the observed patterns of dominance of C. glomerata.     

NOTES ON CLADOPHORA (CHLOROPHYCEAE). II. CLADOPHORA CATENATA (LINNAEUS) KÜTZ., 1843, EMEND. V. D. HOEK, 1963

The species name Cladophora catenata (L.) Kütz. emend, van den Hoek has priority over Cladophora fuliginosa Kütz.     

AN ECOLOGICAL REVIEW OF CLADOPHORA GLOMERATA (CHLOROPHYTA) IN THE LAURENTIAN GREAT LAKES1

Cladophora glomerata (L.) Kütz. is, potentially, the most widely distributed macroalga throughout the world’s freshwater ecosystems. C. glomerata has been described throughout North America, Europe, the Atlantic Islands, the Caribbean Islands, Asia, Africa, Australia and New Zealand, and the Pacific Islands. Cladophora blooms were a common feature of the lower North American Great Lakes (Erie, Michigan, Ontario) from the 1950s through the early 1980s and were largely eradicated through the implementation of a multibillion‐dollar phosphorus (P) abatement program. The return of widespread blooms in these lakes since the mid‐1990s, however, was not associated with increases in P loading. Instead, current evidence indicates that the resurgence in blooms was directly related to ecosystem level changes in substratum availability, water clarity, and P recycling associated with the establishment of dense colonies of invasive dreissenid mussels. These results support the hypothesis that dreissenid mussel invasions may induce dramatic shifts in energy and nutrient flow from pelagic zones to the benthic zone.     

ACCUMULATION OF WATER SOLUBLE PHOSPHORUS AND HYDROLYSIS OF POLYPHOSPHATES BY CLADOPHORA GLOMERATA (CHLOROPHYCEAE)1,2

Concentrations of hot-water extractable phosphorus from most samples of Cladophora glomerata (L.) Kütz. were relatively high (0.06–0.68%) and correlated closely with total dissolved P in ambient Lake Michigan water. Cladophora was able to hydrolyze polyphosphates by enzymes found in intracellular, extracellular and cell wall fractions. The intracellular phosphatase activity is pH dependent with the optimal hydrolysis rate at pH 7.8. Secretion of phosphatases is affected by pH, with maximum rate at 7, but affected little by light intensity. Magnesium is the most effective metallic cofactor required for maximal rates of intracellular phosphatase activities.     

COMPARATIVE ULTRASTRUCTURE OF THE VALVES OF SOME CYCLOTELLA SPECIES (BACILLARIOPHYCEAE)1

Cyclotella atomus Hust., C. meneghiniana Kütz., C. comta (Ehr.) Kütz., C. antiqua W. Sm., C. michiganiana Skv., C. ocellata Pant., C. glomerata Bachmann, C. pseudostelligera Hust., and C. stelligera (Cleve et Grun.) V. H., were selected for ultrastructure investigations on the basis of their ready availability. Specimens were examined employing both transmission and scanning electron microscopy. Strutted processes, labiate processes, spines and other special features were surveyed in all species. On the basis of these observations the species of Cyclotella observed seem to be in one of 3 morphological groups, the meneghiniana group, the comta group, and the stelligera group.     

INFLUENCE OF VARIOUS EXPOSURE PERIODS ON THE BIOMASS AND CHLOROPHYLL A OF CLADOPHORA GLOMERATA (CHLOROPHYTA)1

The biomass of Cladophora glomerata (L.) Kütz. was estimated at selected sites in the Colorado River between Glen Canyon Dam, Arizona, and River Kilometer 354. C. glomerata biomass was significantly higher at sites above Lees Ferry (25 km downstream from the Dam) than sites below the Ferry. Biomass and chlorophyll a were significantly reduced when C. glomerata was subjected to one-time exposures to the atmosphere for 12 daylight h in more. Repeated 12/12 h and 24/24 h (exposure/submergence) cycles over a two-week period also showed a significant reduction in biomass. The adaptations of C. glomerata to “stranding” during regulated flows are discussed.     

UPTAKE OF INORGANIC CARBON BY CLADOPHORA GLOMERATA (CHLOROPHYTA) FROM THE BALTIC SEA1

Carbon uptake in the green macroalga Cladophora glomerata (L.) Kütz. from the brackish Baltic Sea was studied by recording changes in pH, alkalinity, and inorganic carbon concentration of the seawater medium during photosynthesis. The use of specific inhibitors identified three uptake mechanisms: 1) dehydration of HCO₃ ^? into CO₂ by periplasmic carbonic anhydrase, followed by diffusion of CO₂ into the cell; 2) direct uptake of HCO₃ ^? via a 4,4′‐diisothiocyanato‐stilbene‐2,2′‐disulfonate‐sensitive mechanism; and 3) uptake of inorganic carbon by the involvement of a vanadate‐sensitive P‐type H ⁺ ‐ATPase (proton pump). A decrease in the alkalinity of the seawater medium during carbon uptake, except when treated with vanadate, indicated a net uptake of the ionic species contributing to alkalinity (i.e. HCO₃ ^? , CO₃^{2 ?} , and OH ^? ) from the medium, where OH ^? influx is equivalent to H ⁺ efflux. This would suggest that the proton pump is involved in HCO₃ ^? transport. We also show that the proton pump can be induced by carbon limitation. The inducibility of carbon uptake in C. glomerata may partly explain why this species is so successful in the upper littoral zone of the Baltic Sea. Usually, carbon limitation is not a problem in the upper littoral of the sea. However, it may occur frequently within dense Cladophora belts with high photosynthetic rates that create high pH and low carbon concentrations in the alga's microenvironment.     

Strategies providing success in a variable habitat: II. Ecophysiology of photosynthesis of Cladophora glomerata

Cladophora glomerata (L.) Kütz. is the dominant filamentous algae of the river Ilm, Thuringia, Germany. For most of the year it can be found at open as well as at shaded sites. Photosynthetic acclimation of C. glomerata to different light intensities was detected by chlorophyll fluorescence measurements and pigment analysis. Cladophora glomerata from highlight sites showed decreased values of efficiency of open photosystem II (F_v/F_m) as compared with C. glomerata from low‐light sites. Winter populations revealed higher F_v/F_m values than summer populations. A light‐induced decrease in efficiency of the closed photosystem II was observed at increasing irradiance intensities. The decrease was higher in C. glomerata from shaded sites compared with plants from open sites. Differences in the photosynthetic electron transport rate of different populations of C. glomerata were shown by photosynthesis–irradiance curves. Summer populations from high‐light sites yielded higher maximum electron transport rates than plants from low‐light sites, whereas winter populations exhibited significantly decreased values compared with the summer populations. Results of the analysis of photosynthetic pigments corresponded with data from chlorophyll fluorescence measurements. In addition to these long‐term acclimation effects, C. glomerata expressed its ability to cope with rapid changes in the light environment by the de‐epoxidation of violaxanthin during exposure to high light intensities.     

COMMUNITY STRUCTURE DYNAMICS OF EPILITHIC AND EPIPHYTIC DIATOMS IN A MISSISSIPPI STREAM1

Epilithic and epiphytic diatoms were collected monthly from 18 June 1979 through 19 May 1980 from both shaded and unshaded sites of Sessums Creek, a shallow, oligotrophic and strongly alkaline stream in northeastern Mississippi. The dominant macroalgae were filamentous green algae belonging to the non-mucilage producing genera Cladophora and Rhizoclonium. Ninety taxa were identified with Navicula minima Grun. and Nitzschia dissipata (Kütz.) Grun. accounting for 50% of all valves counted. Other important taxa were Navicula menisculus var. upsaliensis (Grun.) Grun., N. cryptocephala var. veneta (Kütz.) Rabh., Achnanthes minutissima Kütz. and Cymbella turgida (Greg.) Cl. With few exceptions, the more dominant taxa were equally abundant in the shaded and unshaded sites and also in the epilithon and epiphyton. Species diversity (H') and the number of taxa in a sample in all four habitats showed an identical pattern, being lowest in winter and highest in fall. No one pair of habitats was more structurally similar than any-other pair when compared by a similarity index (SIMI). Apparently, differences in light intensity and substrate type had little effect on diatom distribution in Sessums Creek and it was concluded that the locality sampled supported a single, nearly homogeneous diatom community.     

ABSOLUTE CONFIGURATION ANALYSIS OF THE FLAGELLAR APPARATUS IN CLADOPHORA AND CHAETOMORPHA MOTILE CELLS,WITH AN ASSESSMENT OF THE PHYLOGENETIC POSITION OF THE CLADOPHORACEAE (ULVOPHYCEAE,CHLOROPHYTA)

Absolute configurational analyses of flagellar apparatus components were performed on the motile cells produced by three species of Cladophora, Cl. dalmatica Kütz., Cl. flexuosa (Dillw.) Harv., and Cl. glomerata (L.) Kütz., and by Chaetomorpha aerea (Dillw.) Kütz. There was little variation among the species. All of the flagellar apparatuses demonstrated the ulvophyceous features of 180° rotational symmetry, counterclockwise absolute orientation, and basal body overlap, as well as the alignment of the basal bodies perpendicular to the long axis of the cell. Diagnostic features included the nearly complete absence of C tubules from the basal bodies and the presence of a coarsely striated component dorsal to the two-membered rootlets in all cells, as well as, in quadriflagellate cells, a tetralobate distal fiber, the coaxial arrangement of the lowermost pair of basal bodies, and the presence of a characteristic array of basal-body-associated striated bands. The distal fiber architecture, the presence of a “wing” in the X-membered rootlets, and the “flattening” of the flagellar apparatus components suggests a close relationship of the Cladophoraceae to the Dasycladales, and indicates that these two groups may have shared a common ancestor, possibly ancient in terms of the geological time scale but relatively recent in the context of ulvophyceous evolution. A sizeable phylogenetic gap exists between the Cladophoraceae and uninucleate-celled, presumably primitive members of the Ulvophyceae.     

LOCAL ADAPTATION OF CERAMIUM TENUICORNE (CERAMIALES,RHODOPHYTA) WITHIN THE BALTIC SEA1

Ceramium tenuicorne (Kützing) Wærn is a red alga that is widely distributed in the brackish Baltic Sea. We studied the growth response of Ceramium to low salinity and nutrient enrichment, using isolades from two regions of the Baltic Sea where the alga approaches its inner distribution limit. Ecotypic differentiation was observed in that differences in growth response among isolates corresponded to salinity conditions in their regions of origin. Isolates from the Gulf of Bothnia (4 psu) had inherently lower growth rates that were not increased when transferred to higher salinity, but were better adapted to very low salinity levels than isolates from the Baltic Proper (7 psu). Further, the results indicate that Ceramium from different regions of the Baltic Sea vary in their responses to nutrient enrichment. The observed differences may be best described as a quantitative difference in the proportion of isolates with hyposaline adaptation. The results indicate that the wide distribution of Ceramium in the Baltic Sea is better explained by the occurrence of locally adapted genotypes than by a generalist life strategy, and provide example of adaptive differentiation in a marine edge environment.     

PHYLOGENETIC RELATIONSHIPS WITHIN CLADOPHORALES (ULVOPHYCEAE,CHLOROPHYTA) INFERRED FROM 18S rRNA GENE SEQUENCES,WITH SPECIAL REFERENCE TO AEGAGROPILA LINNAEI 1

The phylogenetic position of a freshwater green alga, Aegagropila linnaei (Cladophorales, Ulvophyceae), was investigated using nuclear 18S rRNA gene sequences. This alga has usually been called Cladophora aegagropila (L.) Rabenhorst or Cladophora sauteri (Nees ex Kütz.) Kütz. Based on morphology, it was formerly classified into the section Aegagropila or into the subgenus Aegagropila, together with several marine species of the genus Cladophora. This classification is not supported by the present phylogenetic analyses in which two very distinct Cladophorales clades are recognized. Aegagropila linnaei groups together in a well‐supported clade with Cladophora sp., Pithophora sp., Chaetomorpha okamurae, Arnoldiella conchophila, Wittrockiella lyallii, and Cladophora conchopheria. Aegagropila linnaei and its closely related species share some ultrastructural and biochemical characteristics, like pyrenoid structure, carotenoid composition, and cell wall composition. Freshwater species, included in the analysis, were located in two distantly related lineages, indicating that adaptation from a marine to a freshwater habitat has happened at least twice independently in the Cladophorales.     

THE DISTRIBUTION OF EDAPHIC DIATOMS ALONG ENVIRONMENTAL GRADIENTS OF A LOUISIANA SALT MARSH1

Edaphic (sediment-associated) diatoms were collected from five sites along a gradient from slightly brackish to saline marsh on 30 November 1977 and 29 April 1978. Several environmental factors change along this gradient in addition to salinity, including tidal flushing and percent organic content of the soil. A total of 112 taxa were identified, with Amphora exigua Greg., Navicula phyllepta Kütz., N. salinarum Grun., and N. tripunctata (Müll.) Bory being among the five most abundant taxa on both sampling dates. The taxonomic composition of the diatom flora was very similar to those observed for other salt marshes located along the Gulf and Atlantic coasts. Diatom taxa were generally euryhaline and distributed continuously along the gradient from brackish to saline marsh. Species diversity (H¹) and the number of taxa in a sample (S) were also similar to those reported for other North American salt marshes. The number of taxa in a sample was higher in the saline than in the brackish area on both sampling dates while species diversity was greater in the saline area only in April.     

GENETIC ADAPTATION AND ACCLIMATION OF PHYTOPLANKTON ALONG A STRESS GRADIENT IN THE EXTREME WATERS OF THE AGRIO RIVER–CAVIAHUE LAKE (ARGENTINA)1

We tested if different adaptation strategies were linked to a stress gradient in phytoplankton cells. For this purpose, we studied the adaptation and acclimation of Dictyosphaerium chlorelloides (Naumann) Komárek et Perman (Chlorophyta) and Microcystis aeruginosa (Kütz.) Kütz. (Cyanobacteria) to different water samples (from extremely acid, metal‐rich water to moderate stressful conditions) of the Agrio River–Caviahue Lake system (Neuquén, Argentina). Both experimental strains were isolated from pristine, slightly alkaline waters. To distinguish between physiological acclimation and genetic adaptation (an adaptive evolution event), a modified Luria‐Delbrück fluctuation analysis was carried out with both species by using as selective agent sample waters from different points along the stress gradient. M. aeruginosa did not acclimate to any of the waters tested from different points along the stress gradient nor did D. chlorelloides to the two most acidic and metal‐rich waters. However, D. chlorelloides proliferated by rapid genetic adaptation, as the consequence of a single mutation (5.4 × 10^?7 resistant mutants per cell per division) at one locus, in less extreme water and also by acclimation in the least extreme water. It is hypothesized that the stress gradient resulted in different strategies of adaptation in phytoplankton cells from nonextreme waters. Thus, very extreme conditions were lethal for both organisms, but as stressful conditions decreased, adaptation of D. chlorelloides cells was possible by the selection of resistant mutants, and in less extreme conditions, by acclimation.     

Invertebrate grazing and riparian shade as controllers of nuisance algae in a eutrophic river

1. Algal growth in lotic systems is controlled either from the bottom‐up (e.g. nutrients and light, which determine growth rates) or from the top‐down (e.g. grazing pressure, which reduces accumulated biomass). Nutrient‐enriched streams that support large and diverse grazing macroinvertebrate populations and those with shaded riparian corridors rarely suffer from excessive algal growth. 2. In this study, the density of benthic algivorous macroinvertebrates was experimentally manipulated in shaded and open nutrient‐enriched stream habitats of the Owennagearagh River, south‐west Ireland. The ability of macroinvertebrate grazers and riparian shade to control benthic algal growth [particularly the nuisance alga, Cladophora glomerata (L. Kütz)] was investigated. Three sites with markedly different concentrations of plant nutrients (one site upstream and two sites downstream of the sewage outfall) were selected. The density of grazing invertebrates colonising ceramic tiles was reduced using high‐voltage localised electric pulses. Replicates of treatment (grazer‐excluded) and control (grazed) tiles were deployed in open and shaded (<25 and >80% canopy cover, respectively) patches of stream bed, in each site. 3. After 2‐week Cladophora cover, periphytic chlorophyll a and biofilm ash‐free dry mass (AFDM) were quantified for all experimental tiles. Values for all three parameters were highest on grazer‐excluded tiles from open patches. Grazed tiles from open patches accrued little Cladophora and had significantly lower levels of chlorophyll a and AFDM. Nutrient inputs were found to have an impact on the density of grazing invertebrates, with higher densities of Baetis nymphs at the most nutrient‐enriched site. 4. Our results demonstrate that in eutrophic, high‐light streams, filamentous algae can quickly accumulate to nuisance levels in the absence of invertebrate grazers. In future, greater attention should be paid to the role of grazing invertebrates in controlling nuisance algae in streams, in addition to algal–nutrient relationships.     

ON THE LIFE CYCLE,CYTOLOGY AND TAXONOMY OF CLADOPHORA CALLICOMA FROM INDIA

The present investigation deals with the cultural observations on the morphology, reproduction, life cycle, cytology and taxonomy of a freshwater Cladophora, C. callicoma (L.) Kütz, from India. There is a regular isomorphic alternation of generations between quadriflagellate zoospore-producing diploid plants and biflagellate isogamete-producing haploid plants, coupled with heterothallism in the latter. Chromosome numbers of n = 12 and 2n = 24 were determined respectively for gametophytes and sporophytes. Twelve bivalents were counted in meiosis where chiasmata varied from one to three. In light of above observations, the affinity of C. callicoma with C. glomerata, to which the former has often been assigned, has been discussed. It is proposed that the C. glomerata complex, which consists of intraspecific polyploid races, might have two distinct lines of evolution with regard to life cycle; one with forms having higher ploidy levels and lacking an alternation of generations, and the other with forms having low ploidy levels and alternation of generations.     

DNA SEQUENCE VARIATION IN THE RIBOSOMAL INTERNAL TRANSCRIBED SPACER REGION OF FRESHWATER CLADOPHORA SPECIES (CHLOROPHYTA)1

Freshwater species of Cladophora (Chlorophyta) are globally distributed and occupy an unusually wide range of ecological habitats. Delineating species is difficult because most easily observed morphological traits are highly variable and because sexual reproduction has not been clearly documented. Synthesizing ecological data on freshwater Cladophora species is problematic because it is unclear whether freshwater Cladophora species comprise many genetically distinct species or a few ecologically and morphologically variable and/or plastic species. We determined nucleotide sequences of the internal transcribed spacer (ITS) region of the nuclear ribosomal cistron of freshwater Cladophora species from a wide range of habitats and geographic locations. We compared these sequences to those derived from culture collections of C. fracta and C. glomerata, the two most commonly reported freshwater Cladophora species. Cladophora fracta and C. glomerata had very similar ITS sequences (95.3%). All other sequences were identical to those from the C. fracta or C. glomerata culture collections with the exception of one California sample that was similar to both C. fracta (95.6%) and C. glomerata (92.4%). ITS genotypes did not correlate with morphology or geography. This analysis shows that common freshwater Cladophora species comprise very few (possibly one) ecologically and morphologically variable species.     

Elongation and mat formation of <Emphasis Type="Italic">Chara aspera</Emphasis> under different light and salinity conditions

Former laboratory results indicate that shoot elongation at low light intensities of Chara aspera is absent already at 10 psu which is within the physiologically optimal salinity range for brackish water populations. To investigate if similar restrictions occur in the field, density and morphology of C. aspera were compared between three freshwater and three brackish water sites along its depth range. The lower depth limit of C. aspera varied considerably among sites (30–600 cm) related to turbidity. Light availability at the lower depth limit corresponded to about 15% of surface irradiance in freshwater and brackish water with lower salinity (3.4 psu). Total length increased and fresh weight:length ratio decreased with depth at these sites indicating shoot elongation related to lower light availability. Due to shoot elongation, light availability was far higher at the upper parts of the shoot than at the bottom in the turbid sites. Light availability at the lower depth limit was higher (about 40%) at two sites with higher salinity (7–8 psu), where no shoot elongation was observed at the lower depth limit. Instead, the plants were stunted and often covered with filamentous algae or shaded by other rooted submerged macrophytes indicating competitive disadvantages of C. aspera at higher salinities. As growth in high densities (mat formation) exposes the plants to severe self-shading, it is suggested that shoot elongation is a prerequisite to mat formation. Dense vegetation of C. aspera was found only in freshwater and brackish water with lower salinity. Single, richly branched plants occurred in clearwater sites with higher salinity. C. aspera was not found in “double stress” environments with both high turbidity and high salinity: We asume that the species is a poor competitor under these conditions. Our results indicate that morphological differences between freshwater and brackish water populations of C. aspera are at least partly explained by salinity rather than genetic differences.     

Tolerance to salinity and thermal stress by larvae and adults of the serpulid annelid Ficopomatus enigmaticus

The serpulid annelid Ficopomatus enigmaticus is a widely distributed invader of shallow‐water, brackish habitats in subtropical and temperate regions, where it has numerous damaging ecological and economic effects. Its distributional pattern suggests that temperature and salinity play important roles in limiting its distribution, but because other factors often covary with these, drawing strong conclusions from these patterns is difficult. In an effort to more clearly identify the effects of these factors, we examined tolerance to acute thermal (16–28°C) and salinity (0–35 psu) stress by larvae (5‐day exposure, unfed) and adults (14‐day exposure, unfed) of F. enigmaticus in the laboratory experiments. Larvae showed higher mortality at the highest temperature tested 28°C; adult survival was unaffected by temperature. Neither larvae nor adults survived exposure to pure freshwater (0 psu), but survived well at salinities ranging 3.5–35 psu. In addition, high salinity did not slow tube growth in adults. These results suggest that salinity stress, in particular, does not directly limit the distribution of F. enigmaticus to low‐salinity habitats. Experimental work on the distribution of F. enigmaticus is uncommon in the literature, but is likely needed to identify the abiotic or biotic factors that limit the distribution of this frequently invasive species.