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Phytoplankton is composed of a diverse community responsible for the primary productivity in water supply reservoirs. Due to their role as the basis of the food web as well as because of their short life cycles, the spatial and temporal dynamics of the phytoplankton community can provide important information about other biotic components and on general environmental conditions of an ecosystem (Reynolds, 2006). Additionally, water supply reservoirs are convenient environments to investigate the effects of environmental variables on phytoplankton structure due their highly dynamic conditions (Chellappa et al., 2009; Yang et al., 2017).
Seasonal fluctuations in rainfall alter the water volume stored by reservoirs as well as other environmental variables, such as mixing conditions, turbidity, oxygen and nutrients contents in the water column (Chaves et al., 2013; Mosley et al., 2015; Santos et al., 2017). These changes, in turn, lead to shifts in phytoplankton attributes (Chellappa et al., 2009; Câmara et al., 2015), which reinforce the use of this community to evaluate the environmental status of these aquatic ecosystems. For example, high concentrations of nutrients, high conductivity and increased phytoplankton biomass could be expected in reservoirs and lakes located in semi-arid regions with reduced water volume (Naselli-Flores, 2003; Bouvy et al., 2003; Mosley et al., 2012; Jirsa et al., 2013). On the other hand, since the reduction of water volume occurs along with intensification in sediment resuspension and turbidity, a negative impact on phytoplankton biomass could also be expected (Braga et al., 2015). Additionally, the dominance of few species, with particular concern for the dominance of potentially harmful cyanobacteria, often occurs in semi-arid regions under drought conditions (Bouvy et al., 1999; Brasil et al., 2016).
The phytoplankton classification into functional groups rather than only based on taxonomic affiliations has been often used as tool for environmental characterization of reservoirs (Silva et al., 2015; Rangel et al., 2016; Rodrigues et al., 2017). In the present study we have use Reynolds's functional groups (FGs), when the species with similar sensibilities, tolerances and ecologies affinities are placed together, being each FG selected by a set of environmental variables (habitat template) (Reynolds et al., 2002 reviewed by Padisák et al., 2009). This approach provides some advantages: i) it can be used to a clear characterization of the environment since the habitat template is known to each Reynolds's FG; and ii) it is of easy application because the FGs are already defined for a large amount of species and it does not require biovolume measurements to fitting each species into predetermined FGs, as other functional schemes need, such as that of Kruk et al. (2010). In this way, Reynolds et al. (2002) FG selection was the better adapted to our goal and the data set (species densities and total phytoplankton biomass), available for our studied reservoir. Other particular characteristic of Reynolds's functional scheme is that it requires the identification at the species level and sometimes genus level, an important information for characterizing water supply in reservoirs where potentially harmful cyanobacteria often occurs (Brasil et al., 2016). Additionally, Reynolds's FGs were previously used by Molisani et al. (2010) in the same studied reservoir, allowing a comparative approach.
In this sense, the objective of this work was to evaluate the effect of environmental changes on phytoplankton composition using taxonomic affiliations and Reynolds's functional groups in a large semi-arid reservoir during eighteen months of severe drought, when the water volume continually dropped due to rainfall shortage. Since droughts are increasing in frequency and severity in many regions such as in tropical semi-arid areas (Marengo et al., 2013; Marengo and Bernasconi, 2015), this study may give us clues to understanding the effect of climate change on phytoplankton of artificial reservoirs. These systems play fundamental roles in the Brazilian semi-arid due their multiple uses as drinking water supply and intensive fish-cage aquaculture sites (Cirilo, 2008; Molisani et al., 2015).
Additionally, the results obtained in the present study were compared with those found by Molisani et al. (2010) in the same reservoir, in order to evaluate possible shifts in the phytoplankton composition and biomass from the first years when the Castanhão Reservoir was completely filled (2006/2007) to the period eight years later (2012/2013), when the reservoir suffered nearly 50% of water reduction.
Study area
The study was carried out in the Castanhão reservoir (5.50°S 38.47°W) in the Middle Jaguaribe River watershed, which is located entirely within the semi-arid region in the State of Ceará, NE Brazil (Fig. 1). The total storage capacity of the reservoir is 6.7 billion m3, and the normal operating capacity is 4.45 billion m3. The reservoir reached its full capacity for the first time only in 2004 and covers an area of 325 km2 and is 48 km in length, with a depth exceeding 50 m in some areas (DNOCS, 2014). The Castanhão reservoir accounts for 97% of the total storage capacity from the Middle Jaguaribe River basin and is the largest reservoir in Ceará State. It serves multiple uses, including water supply for drinking water, agriculture and aquaculture (COGERH, 2011). The categories of the World Commission on Dams (2000) classify the Castanhão as a large reservoir.
The semi-arid climate of northeastern Brazil is characterized by annual rainfall means commonly ranging from 400 to 1000 mm, with average of 756.5 mm during the past 80 years (FUNCEME, 2014). Rainfall occurs from January to June and is scarce from July to December.
Although the years of 2012 and 2013 have not been under El Niño influence, a phenomenon associated to lower rainfall in northeastern Brazil, these years were characterized as extreme drought due to the Atlantic dipole (Marengo et al., 2016). Some studies have reported that the heating of the North Atlantic waters, combined with a cooling of the South Atlantic waters, characterizing a positive Dipole, leads to a situation of descending movements of air over the Brazilian Northeast region, inhibiting the formation of clouds (Marengo et al., 2013; Nóbrega and Santiago 2014; Marengo et al., 2016). In fact, it was observed that in the years of 2012 and 2013, rainfall was only 302.3 mm year–1 and 656.5 mm year–1, respectively, significantly below the historical annual average (FUNCEME, 2014).
As a consequence of this period of intense drought during 2012/2013, the Castanhão reservoir showed a very distinctive situation from those observed until 2009, when the reservoir almost reached the limit of its total capacity storage, and even from 2011, when the reservoir volume was about 80% of its capacity (DNOCS, 2014). During the study period, the reservoir volume dropped progressively, reaching 72%, 65%, 55% and 50% of its total capacity in March 2012, August 2012, January 2013 and August 2013, respectively (DNOCS, 2014).
Sampling, environmental variables and chlorophyll a
Sampling campaigns occurred in March and August 2012 and January and August 2013, covering 8 stations located throughout the reservoir (Fig. 1). The stations P1 and P2 corresponded to the inner zone, P3 and P5 to the middle zone, while P6, P7, P8 and P10 to the dam zone. Due to operational issues data from P1, P2 and P3 in March 2012, P1 in January/13 and P3 in August/13 were not collected.
Prior to water collection, major physical and chemical variables were measured in situ in sub-superficial waters (0.5 to 1.0 m depth): dissolved oxygen (YSI 556 probe, YSI Inc., Yellow Springs); water temperature, turbidity, electrical conductivity (Compact-CTD model AST D687; JFE Advantech Co., Ltd., Nishinomiya); pH (Portable 826 pH-meter; Metrohm AG, Herisau); and water transparency with a Secchi disk. Oxygen, water temperature, turbidity, electrical conductivity and pH were measured with an accuracy of 0.1 mg L–1, 0.1°C, 0.1 NTU, 1.0 μS cm–1 and 0.1, respectively.
Water samples collected from the subsurface (1.0 m) layer with a Van Dorn bottles were filtered in 47-mm-diameter AP40 glass fiber filters and analyzed for inorganic nutrients: nitrate-N (Braga et al., 2015), nitrite-N (Bendschneider and Robinson, 1952), ammonium-N (Koroleff, 1970) and phosphate-P (Murphy and Rilley, 1962). Unfiltered samples were used to determine total phosphorus (TP) and total nitrogen (TN) (Valderrama, 1981). All nutrients were quantified in triplicate by visible spectrum spectrophotometry reaching detection limits of 1.0 μg L–1, 10 μg L–1, 0.1 μg L–1, 1.4 μg L–1, 0.1 mg L–1 and 5.0 μg L–1 for phosphate-P, nitrate-N, nitrite-N, ammonium-N, TN and TP, respectively. The chlorophyll a concentrations were used as an estimative of total phytoplankton biomass in each sample and were obtained in a spectrophotometer according to the ISO 10260 (1992) protocol. For chlorophyll a analysis the samples were filtered immediately after sampling in a field lab through 47-mm-diameter AP40 glass fiber filters. More detailed field procedures and analytical methods were described previously by Santos et al. (2017).
Phytoplankton analysis
Phytoplankton samples were taken from the subsurface layer with a Van Dorn bottle concomitantly with those for nutrients and were immediately fixed with Transeau solution (ratio 1:1). The quantification and identification of phytoplankton organisms were performed according to the Utermöhl (1958) method in an inverted microscope equipped with phase-contrast (Olympus CK2) (Edler and Elbrächter, 2010). The phase-contrast was used to improve the quantification and identification of slender cyanobacteria. The volume of sedimentation chambers was of 2 or 5 mL, depending on the organisms' concentration and sedimentation time ranged from 24 to 48 hours. The counting was standardized to obtain at least 100 organisms (cells, colonies, filaments and trichomes) of the dominant taxa, with the standard error estimated to be 20% and the confidence limit of 95% (Edler and Elbrächter, 2010).
During the counting procedure the organisms were quantified and identified concomitantly at 400x magnification. However, to help in qualitative analysis, at the time of counting, the organisms were photographed and subsequently measured and analyzed according their morphology for identification at the lowest possible taxonomic level using the bibliography cited thereafter. The images were processed using Axio Vision Rel. 4.7 software which allowed the accurate measurement of the organisms, even those slender cyanobacteria with a diameter of about 1.0 μm. When necessary, organisms were also observed under 1000x magnification. In this step, it was possible to identify four pairs of species that were not separated during the counting procedure, due their large morphology similarity. They were i) Planktolyngbya minor and Pl. limnetica; ii) Pseudanabaena catenata and Ps. papillaterminata; iii) Cylindrospermopsis sp. and C. raciborskii (on coiled morphotypes); iv) Synedra acus and Fragilaria delicatissima. Therefore, though in qualitative analysis these species have been discriminated, in statistical analysis these four pairs of species were considered as four taxa because they were previously counted together. It is important to emphasize that, as each pair of species was classified in the same Reynolds's FG, no bias was introduced in data interpretation.
The identification was based on specific bibliography (books and articles) for each phytoplankton group (Komárek and Anagnostidis, 2000; Komárek, 2001; Brassac and Ludwig, 2003; Cronberg and Komárek, 2004; Komárek and Anagnostidis, 2005; Bicudo and Menezes, 2006; Komárek, 2013; McGregor, 2013; and references therein). The identified taxa were assembled into functional groups, using the criteria established in Reynolds et al. (2002) and reviewed by Padisák et al. (2009).
Data analysis
Relative abundance and frequency of occurrence
In order to evaluate the structure of the phytoplankton community, the following indicators were used: density expressed in organisms L–1 (org L–1), relative abundance (RA) and frequency of occurrence (F) expressed as a percentage. The relative abundances (RA) (%) of each taxonomic class, taxa or functional group were calculated based on the density of each one in relation to total density of organisms in the sample. In turn, the frequencies of occurrence of each taxon were calculated based on the numbers of samples where each taxon occurred divided by the total number of total samples. Regarding the frequency of occurrence, each taxon could be classified as: Rare (F ≤ 10%); Common (10% < F ≤ 50%) or Constant (F > 50%).
Statistical analysis
Contents of chlorophyll a and other environmental variables (except pH) were transformed by log (x+1) before all multivariate analysis. For phytoplankton densities (total and of each taxon), data were transformed by fourth root prior to statistical analysis.
The differences among the reservoir zones (inner, middle and dam) relative to environmental variables, total phytoplankton density and chlorophyll a were analyzed using one-way analysis of variance (ANOVA) followed by Tukey's honestly significant different test (T-HSD) at P<0.05 level of probability. In turn, to test the hypothesis of significant differences among the reservoir zones relative to total phytoplankton composition, we used a Permutational Multivariate Analysis of Variance based on Bray-Curtis similarity (PERMANOVA, P<0.05).
Bray-Curtis similarity based on densities of all phytoplankton taxa was also used to access the degree of similarity of phytoplankton community among samples. Cluster analysis based on the Bray-Curtis similarity was performed using UPGMA, and the significance of groups was tested with the Similarity Profile test (SIMPROF). A similarity percentage analysis (SIMPER) was applied to obtain the contribution of each taxa in percent terms to the similarity within each group. The 'characterizing taxa' was defined as those taxa that most contributed to the similarity within each group using a cut-off of 90%. The significance level for SIMPROF and SIMPER was P<0.05.
The SIMPROF results showed that samples could be divided in three significant groups relative to total phytoplankton composition (P<0.05). In order to test the hypothesis of differences among these three sample groups for environmental variables, chlorophyll a and total phytoplankton density, ANOVA was used followed by Tukey's honestly significant different test (P < 0.05).
Redundancy analysis (RDA), a constrained ordination method, was used to examine the relationships between the environmental variables and phytoplankton community and to select the variables that best described the temporal variability of phytoplankton structure. Firstly, two RDA models were performed in order to test the efficiency of the top 'characterizing taxa' to summarize the overall temporal variability of phytoplankton structure and their correlations with environmental variables: one using biotic data of all 102 taxa (response matrix 1) found along the survey; and another using only the top nineteen 'characterizing taxa' (response matrix 2) defined by the SIMPER (cut-off 90%). The length of the first axis (<3.0) of Detrended Correspondence Analysis (DCA) of response matrices data supported the use of RDA.
For both response matrices 1 and 2, the same procedures were adopted to reach the final RDA and to access the relationships between environmental variables (predictor matrix) and phytoplankton structure (response matrix). Firstly, a preliminary RDA was performed with all available environmental variables: Secchi depth (Secchi), turbidity, pH, Temperature, Conductivity, Dissolved Oxygen, Total Phosphorous, Total Nitrogen, phosphate-P, nitrate- N, nitrite-N and ammonium-N. Only the significant variables (P<0.05) were retained in the second RDA. In order to know the collinearity among the explanatory variables, we calculated the variation inflation factor (VIF) of each one. The VIF >10 indicates that a variable is strongly dependent on others and therefore does not have independent information (Oksanen et al., 2015). Only variables with VIF <10 were used in the second RDA model. We tested the marginal significance of the remaining variables by 999 permutations, and only those with significance were used in final RDA. The significance of the first two axis and of the final RDA models were tested by 999 permutations. The adjusted R2 values were computed to obtain the explanatory power of the final RDAs.
As the same results were found for RDAs using the response matrix 1, based on biotic data of all 102 taxa (data not shown), and the response matrix 2, based on the top nineteen 'characterizing taxa', only the last one is shown due its higher power to summarize the variability of phytoplankton composition in the Castanhão reservoir throughout the survey.
The ANOVA and T-HSD tests were performed using the software Statistica® 7.0 (Statsoft Inc., Tulsa, OK, USA). The Bray–Curtis similarity followed by cluster analysis, the SIMPROF test and SIMPER procedure were carried out using the software PRIMER 6.0 (Plymouth Marine Laboratory). PERMANOVA, DCA and RDA analyses were done using the vegan package in R software (Oksanen et al., 2015).
A total of 102 taxa were identified, distributed among the classes Cyanophyceae (cyanobacteria) (34), Chlorophyceae (green algae) (42), Bacillariophyceae (diatoms) (18), Zygnemaphyceae (4) and Euglenophyceae (4), besides one unidentified phytoflagellate taxon. Within this total set of 102 taxa, we discriminated 18 functional groups, being Cyanophyceae in seven codons; MP, S1, L, LM, Lo, SN and H1, Bacillariophyceae in six; P, B, D, MP, C and B, Chlorophyceae in four; P, X1, F and J, Zygnemaphyceae in two; N e NA and Euglenophyceae in one codon; W2. Furthermore, 26 taxa from the total data set were not classified to any functional group described by Reynolds et al. (2002) and Padisák et al. (2009) either because their identification did not occur at the taxonomic level of species or genera or because they have not yet been included within the associations described by those authors. Among the most representative taxa not assigned to any functional group, we highlight Pseudanabaena papillaterminata, Pseudanabaena cf. biceps, Pseudanabaena sp.1 and Myxobaktron sp. This reinforces the fact that the categorization of phytoplankton into Reynolds's functional groups, especially taking into account tropical ecosystems, needs to be revised aiming the inclusion of new species within the previously described groups as well as with the proposition of new functional groups. For this, the ecology of each species still needs to be better understood.
Therefore, taking into consideration the recommendations provided by Padisák et al. (2009) we classified Ps. cf. biceps and Ps. papillaterminata on functional group MP, which included periphytic or epilithic species occasionally found in plankton samples. Komárek and Anagnostidis (2005) described Ps. papillaterminata as a benthonic species. In turn, Ps. cf. biceps is found mainly attached to substrate (Komárek and Anagnostidis, 2005; Douma et al., 2009; Klemenčič et al., 2010).
For Myxobaktron sp. we propose its classification as X1 codon, because it reached the better performance along with small green algae of similar morphology belonging to X1 codon (see discussion). However, as Myxobaktron is a genus poorly cited in limnological studies and information about its autoecology is scarce, we prefer to assign it as FG=X1 (?). In the case of Pseudanabaena sp.1, a slender genus of cyanobacteria, which the classification on Reynolds's FG depends on species, it was not proper to assign it to any functional group because it reached maximum density in a distinct moment from the other 'characterizing taxa' of similar morphology. Additionally, due to their close association with Secchi depth, we have avoided its classification in MP or S1, typical codons from turbid environments (Padisák et al., 2009). Therefore, from the list of 102 taxa, where 26 of them were not previously classified to any Reynolds's FGs, 23 taxa remained as unsigned.
Spatial variability
The results from abiotic variables characterizing the inner, middle and dam zones during the sampling period are summarized in Supplementary Tab. 1. Except for turbidity, no significant differences were found between the zones (ANOVA, P>0.05). The turbidity values in the inner zone were significantly higher than those found in middle and dam zones (ANOVA, T-HSD, P<0.05). In the inner zone, the lowest Secchi-disk transparency values were also observed. In all three zones, alkaline pH and highwater temperature (>27.0°C) were observed, as well as, high levels of oxygen (> 6.0 mg L–1) and conductivity (> 300 μS cm–1). On the other hand, the levels of inorganic nutrients were generally low, ranging from non-detectable to maximum concentrations of 11.2 μg L–1, 44.6 μg L–1, 2.5 μg L–1 and 132.4 μg L–1 for phosphate-P, nitrate-N, nitrite- N and ammonium-N, respectively. The TN and TP concentrations ranged from low to intermediated values in each zone, with average values of these nutrients below 30 μg L–1 (TP) and 620 μg L–1 (TN).
Average phytoplankton densities were 4.0 (±3.0) 106 organisms L–1, 6.0 (±4.0)×106 organisms L–1 and 10 (±8.0)×106 organisms L–1 at inner, middle and dam zones, respectively. Although the phytoplankton densities have increased toward the dam zone, no significant differences were found between the three reservoir zones (ANOVA, P>0.05). Similarly, the total phytoplankton biomass (chlorophyll a) increased from the inner to the dam zone, with average values of 2.9±0.5 μg L–1, 4.5±0.7 μg L–1 and 5.0±2.1 μg L–1 at the inner, middle and dam zones, respectively. A significant difference was found between the inner and dam zone (ANOVA, T-HSD, P<0.05), but not between the middle and dam zone (ANOVA, T-HSD, P>0.05).
In all three zones of the reservoir the cyanobacteria were dominant with a relative abundance above 80%, followed by green algae and diatoms with relative abundances ranging from 5 to 10% in each zone. The most abundant functional group in all three zones was S1 (Fig. 2), represented mainly by the cyanobacteria Pseudanabaena limnetica and Planktolyngbya limnetica/Pl. minor. The third more abundant taxon was the cyanobacteria Pseudanabaena sp. 1 (FG=?), not assigned to any functional group (Fig. 2). Thus, the phytoplankton composition throughout the reservoir was very constant and no significant differences were found among the zones by PERMANOVA test (F model=1.4061; r2=0.05325; P=0.234; 999 permutations).
Temporal variability
The cluster analysis based on the densities of the 102 phytoplankton taxa is presented in Fig. 3. In this Figure, one can observe the formation of three significant groups (SIMPROF, P<0.05), showing a clear tendency to temporal clustering of samples in three phases/groups: phase I including all samples from March/12; phase II formed by samples from August/2012 and January/2013, plus one sample from August/2013 (P1); and phase III including all samples from August/2013 (except for P1).
Supplementary Tab. 2 summarizes the results of abiotic variables of the three temporal phases suggested by cluster analysis. No significant differences were found for turbidity, dissolved oxygen, pH, nitrite-N, TP and TN among the phases (ANOVA, P>0.05). In turn, the water temperature was significantly different among the phases, reaching higher values in temporal phases I and III, corresponding to the rainy period (ANOVA, T-HSD, P > 0.05). The higher Secchi-disk depth was found in phase II (ANOVA, T-HSD, P < 0.05), while the higher conductivity and nitrate-N were observed in phase III (ANOVA, T-HSD, P < 0.05). Phase I was distinctive from the others by the lowest phosphate-P concentrations (ANOVA, THSD, P < 0.05). Phase II was also distinctive from phase I by the lowest ammonium-N concentrations (ANOVA, T-HSD, P < 0.05).
A list of the major contributors ('characterizing taxa') to similarity within each group/phase according to the SIMPER procedure (cut-off 90%) and their frequency of occurrence during the survey are presented in Tab. 1. Tab. 2 summarizes the percentage contribution of the major contributors, but with a cut-off of 70%, to the average similarity within each group/phase. In all phases the cyanobacteria were the dominant class, followed by diatoms in phase II and green algae in phase III (Tab. 2).
The mean relative abundance (%) of the main phytoplankton functional groups for the phases/groups during the sampling period are presented in Fig. 4. Phase I is clearly distinct from phase II by the dominance of the functional group S1, represented mainly by cyanobacteria Planktolyngbya limnetica/Pl. minor with relative abundance of 62.3% along with Pseudanabaena limnetica (RA=25%). Other important functional groups in phase I were MP represented mainly by cyanobacteria Ps. catenata/ Ps. papillaterminata (RA=7.5%) and SN represented by Cylindrospermopsis sp./C. raciborskii (RA=1.7%).
In phase II the taxa present in phase I decreased in density and were outcompeted mainly by cyanobacteria Pseudanabaena sp.1 (FG=?) (RA=36.2%). Pl. limnetica/ Pl. minor (S1) continued to be an important cyanobacteria in phase II with relative abundance of 21.3%, followed by Ps. cf. biceps (MP) (RA=8.4%), Cylindrospermopsis sp./C. raciborskii (SN) (RA=3.9%) and Myxobaktron sp. (X1?) (RA=2.6%). Despite in phase II the Cyanophyceae have also been the dominant class, an increase in Bacillariophyceae was observed, mainly due to Nitzschia sp. (D) (RA=5.6%) and Aulacoseira granulata var. granulata (P) (RA=3.5%) (Fig. 4).
Finally, in phase III the functional group S1 increased again in density and relative abundance, mainly due to the cyanobacteria Ps. limnetica (RA=40.3%) and Pl. limnetica/ Pl. minor (RA=14.4%). These taxa were followed by cyanobacteria Ps. catenata/Ps. papillaterminata (MP) (RA=7.4%) and Ps. cf. biceps (MP) (RA=4.6 %). Notwithstanding, different from phase I, other functional groups, such as F (mainly due the green algae Micractinium pusillum, RA=5.9%), D (mainly due the diatom Nitzschia sp. RA=4.9%) and X1 (mainly due the green algae Ankyra judayi, RA=2.5%; and Monoraphidium minutum, RA=2.4%) were also well represented in phase III (Fig. 4).
Phase II was distinct from phases I and III by lower total phytoplankton density (ANOVA, T-HSD, P<0.05), but no significant difference was observed between phases I and III (ANOVA, T-HSD, P>0.05, Tab. 2). In relation to the total phytoplankton biomass (chlorophyll a), no significant differences were found among the phases (ANOVA, P>0.05, Tab. 2), though the average have increased progressively from phase I to III.
Redundancy analysis
The RDA performed with nineteen top 'characterizing taxa' defined by the SIMPER procedure (cut-off 90%), as presented in Supplementary table 3, included Secchi depth (P=0.015), ammonium-N (P=0.010), nitrate-N (P=0.010) and phosphate-P (P=0.005), which showed significant marginal effects and VIF <10 (Fig. 5).
The RDA model selected for these four variables was significant (F=6.6, df=4, P=0.001) with an explanatory power of 46.3 % (R2=0.545; adjusted R2=0.463). The first RDA axis was significant (P=0.001) and mainly positively correlated to ammonium-N (0.5961) and negatively to Secchi depth (-0.7733). Axis 2 was also significant (P=0.001) and negatively correlated with nitrate-N (- 0.4310) and phosphate-P (-0.8657).
The first axis (RDA1) separated in the positive side the sampling units from phases I and III, characterized by the highest concentration of ammonium-N and nitrate-N, respectively (Fig. 5). The sampling units from phase I were also distinctive from the others by the lowest phosphate-P concentrations, which were below the detection limits of the analytical method (see Supplementary Tab. 2). Thus, the lowest phosphate-P in phase I contributed to the best performance of Pl. limnetica/Pl. minor (S1) while the highest ammonium-N favored Ps. catenata/Ps. papillaterminata (MP).
On the other hand, in the negative side of axis 1, the sampling units from phase II were separated and characterized by the highest Secchi depth, lowest inorganic nitrogen and an increase in phosphate-P concentrations. This decline in inorganic nitrogen from phase I to phase II, plus an increase in phosphate-P, could be associated to the best performance of Pseudanabaena sp.1 (FG ?) over other cyanobacteria, though a decrease in the total phytoplankton density have occurred, which also leaded to high Secchi depth.
Finally, in phase III an increase in inorganic nitrogen, mainly nitrate-N, was observed. This condition can have triggered a recovery of cyanobacteria from functional group S1, but at this time mostly represented by Ps. limnetica (S1), which was also associated to low Secchi depth. In phase III, various green algae from functional groups X1, J and F were also favored by increased nitrate- N, but also phosphate-P.
Other important information shown by the RDA model for some species with little known autoecology were the close association of Pseudanabaena cf. biceps (MP) with the highest concentrations of phosphate-P; and Myxobaktron sp. (X1 ?) with highest nitrate-N and phosphate- P concentrations.
The analysis of the results of spatial and temporal dynamics of the phytoplankton community in the Castanhão reservoir along the years of 2012 and 2013, when the volume of the reservoir progressively decreased from 70% to 50% (Santos et al., 2017), showed a tendency of temporal rather than spatial variation across the zones. In general, the study period could be divided in three phases based on the phytoplankton structure. Phase I included all samples from March/12, corresponding to the rainy season, while phase II included all samples from August/12 and January/ 13, both representing the dry season. In turn, phase III included samples from August/2013, which corresponded to the end of the rainy season, since the rainfall in this year lasted until this month (data not shown). The cluster analysis also revealed a high similarity between the phytoplankton composition in phases I and III, which reinforces the influence of climatic seasonality to drive the temporal dynamic of this community. In fact, the development of phytoplankton community in Northeast of Brazil is strongly structured by the seasonal climate (rainy/dry periods) (Chellappa et al. 2009; Brasil et al., 2016).
The highest values of phytoplankton density were found in phase I, being Pl. limnetica / Pl. minor (S1) the most successful taxa. Other important taxa were Ps. limnetica (S1) and Ps. catenata / Ps. papillaterminata (MP). In this phase, the reservoir was classified as oligotrophic based on the total phosphorous and chlorophyll a concentrations (Santos et al., 2017). It is important to note that the highest phytoplankton density observed in phase I was not associated to a high phytoplankton biomass (chlorophyll a), precisely due to the dominance of these slender filamentous cyanobacteria, especially Pl. limnetica/ Pl. minor. All these three taxa of cyanobacteria belong to the functional groups S1 and MP and are well adapted to high turbidity conditions (Padisák et al., 2009), what was confirmed by their inverse association with Secchi depth in Castanhão reservoir in RDA.
Although Pl. limnetica/Pl. minor (Delazari-Barroso et al., 2007; Silva and Costa 2015) along with Ps. limnetica (Mohebbi et al., 2015) and Ps. catenata / Ps. papillaterminata (Moura et al., 2007; Silva and Costa, 2015) have been commonly found in mesotrophic to hypertrophic reservoirs, these slender cyanobacteria frequently outcompeted other phytoplankton species under conditions of reduced P supply (Villena and Romo, 2003). In this sense, in the present study, their highest densities, especially of Pl. limnetica/Pl. minor, were mainly associated with reduced phosphate-P supply in phase I, which could be due to the strong thermocline in this period (see Santos et al., 2017). Corroborating this condition, Pl. limnetica was previously shown as a strong indicator of a stratified water column with phosphate-P depletion, probably due to its high-affinity phosphate uptake system (Prentice et al., 2015). In the same way, another study performed in a tropical water supply reservoir (Extremoz Lake, NE Brazil) also showed the dominance of Pl. limnetica under low values of phosphate-P (Pinto and Becker, 2014), in accordance with our results.
It is important to highlight that although we have associated the successful of Pl. limnetica/Pl. minor with a condition of stratified water column, the water samples in this study were restrictive to subsurface (1.0 m), and therefore Pl. limnetica/Ps. minor was recorded in the mixed and turbid surface layer of Castanhão reservoir. This is in agreement with the habitat template (turbid mixed layers) described for codon S1, which this species belongs.
In phase II, the cyanobacteria previously dominant in phase I were partly replaced by other cyanobacteria, mainly Pseudanabaena sp.1 (FG=?) and Ps.cf. biceps (MP). The phytoplankton biomass in phase II was similar to the observed in phase I since the dominance was also of slender filamentous cyanobacteria. Nevertheless, in phase II the reservoir was classified as mesotrophic due to the increase in TP concentration probably driven by the thermocline disruption (see Santos et al., 2017).
Pseudanabaena is a genus with tropical planktonic species recorded in lakes and large reservoirs, while Ps. cf. biceps is a species that lives primarily attached to substrate (Komárek and Anagnostidis, 2005; Douma et al., 2009; Klemenčič et al., 2010) and therefore belonging to MP codon. In the Castanhão reservoir, the most remarkable environmental change associated to the emergence of Pseudanabaena sp.1 in phase II (absent in phase I) was the increase in Secchi depth and the decrease in ammonium-N.
On the other hand, Ps. cf. biceps seemed to be mostly favored by high phosphate-P concentrations, since its average density increased progressively from phase I to phases II and III (Supplementary Tab. 3), following the rise of this nutrient's concentrations. Since Ps. cf. biceps belongs to MP, a codon typical of frequently stirred, inorganically turbid shallow lakes (Padisák et al., 2009), it may also have been favored by decreasing in water volume and thermocline disruption.
Other cyanobacteria with close correlation not only with phosphate increase, but also with nitrate-N increase was Myxobaktron sp. (X1 ?) that increased in density from phase I to phases II and III (Supplementary Tab. 3). This genus generally forms a minor component of the planktic cyanobacterial biocenosis, but persisted year-round in the tropical regions of northeastern Australia (McGregor, 2013), such as observed in the Castanhão reservoir.
We raised the hypothesis that Myxobaktron sp. would be classified in codon X1 because it reached the better performance along with small green algae (Monoraphidium circinale, Monoraphidium minutum) with fast growth and similar size (length of 5-10 μm), belonging to X1. Both Myxobaktron sp. and these green algae increased in density in phase III, following the raise in nitrate-N concentrations and the reduction of the reservoir volume. The habitats templates of X1 codon are shallow and hypertrophic/ eutrophic environments (Padisák et al., 2009).
In addition to the dominant cyanobacteria in phase II, we highlight the rise in relative abundance of diatoms represented mainly by Aulacoseira granulata var. granulata (P) and A. granulata var. angustissima (P). The functional group P is associated with higher trophic states and mixing conditions (Padisák et al., 2009), in accordance with phosphate-P increase and thermocline disruption observed during this phase (Santos et al., 2017).
In phase III, the shade-adapted cyanobacteria that dominated in phase I, which were Pl.limnetica / Pl.minor (S1), Ps. catenata/Ps. papillaterminata (MP) and Ps. limnetica (S1), increased again in density, but at this time with dominance of the last one. Ps. limnetica (S1) is a common species in eutrophic reservoirs (e.g. Mohebbi et al., 2015), and in the Castanhão reservoir it seemed to have been more favored mainly by the increase in nitrate- N concentration in phase III. Messyasz et al. (2015) also indicated the dependence of Ps. limnetica to high nitrate- N concentrations.
Another similarity between phases I and III was the absence of Pseudanabaena sp.1, suggesting its sensibility to decreasing water transparency (estimated by Secchi depth). Pseudanabaena sp.1 was also associated with decreasing in ammonium-N concentrations in phase II. It was not possible to assign Pseudanabaena sp.1 to any Reynolds FG, both because the identification did not occur at species level and because it reached the maximum density in a distinct moment (phase II) from the other 'characterizing taxa' of similar morphology (other slender cyanobacteria).
Also in phase III, we highlight among the diatoms Nitzschia sp., belonging to the functional group D, which is indicative of turbid, eutrophic and shallow waters (Reynolds et al., 2002; Padisák et al., 2009; Silva and Costa, 2015). The decrease in water depth, along with the increase in the trophic conditions of the reservoir in phase III, was also suggested by the rise of green algae belonging to the functional group X1, such as A. judayi, M. minutum, M. circinale and S. setigera, and belonging to J functional group, represented by Golenkinia sp. According to Padisák et al. (2009), both functional groups, X1 and J, are indicative of shallow and nutrient-enriched systems.
Another functional group within green algae was group F, represented mainly by M. pusillum. F group is typical of mesotrophic/eutrophic waters with clear epilimnion (Padisák et al., 2009), which contrasted with the turbidity condition indicated by the other species, mainly the filamentous cyanobacteria from functional group S1.
In this context, the success of M. pusillum (F) in phase III, along with the other green algae belonging to the functional groups X1 and J and the cyanobacteria from S1, could be due to its ability to growth rapidly under favorable conditions, such as increasing in nutrients, in addition to a possible tolerance to high turbidity conditions. This tolerance is indicated by the fact that the functional group F is also found in estuaries, which are naturally turbid environments (Costa et al., 2009). Additionally, M. pusillum could have an adaptive advantage over other species due to its ability to use organic compounds (mixotrophy) (Bouarab et al., 2004).
Similarly, Dias (2009) studying the Arcoverde reservoir (also located in the northeastern Brazil semi-arid zone as well) observed the success of species belonging to functional group F, more specifically M. pusillum, along with others from the functional groups X1 and S1 in the same period (rainy season). In the Arcoverde reservoir, the rainy period in which these functional groups predominated was characterized by high values of turbidity and dissolved oxygen, as well as elevated concentrations of total dissolved phosphorus, nitrite and nitrate (Dias, 2009). Similarly, in the Castanhão reservoir the key-factors to the rise of M. pusillum in phase III were the increase in nitrate-N and phosphate-P concentrations, as shown by the RDA.
Finally, in phase III we call attention to the highest phytoplankton biomass, precisely due to the increase of unicellular and mainly colonial green algae, which were mostly induced by the increase in nutrient concentrations, namely nitrate-N and phosphate-P. In fact, the increase in green algae and diatoms in conditions of high nitrate-N and phosphate-P concentrations could be explained by their reduced uptake rates and thus larger requirements when compared to slender filamentous cyanobacteria. For example, Prentice et al. (2015) showed that the seasonal phytoplankton succession in a large lake was driven by the capacity of phosphate uptake, which resulted into the cyanobacteria dominance under decreased phosphate concentrations during the summer-stratification period and in a diatom/cryptophyte/chlorophyte dominance in surface water with elevated phosphate concentrations. Despite the change in the phytoplankton structure from phase II to III, in this last phase the Castanhão reservoir was also characterized as mesotrophic based on the total phosphorous and chlorophyll a concentrations (Santos et al., 2017).
Overall, our results suggest that in accordance with other studies, shifts in phytoplankton community were associated to changes in nutrient concentrations and water transparency (Medeiros et al., 2015; Costa et al., 2016). Changes in these environmental conditions were closely associated with hydrological changes in the reservoir, namely the reduction in water level due to years of severe drought. However, taking account only the eighteen months of severe drought addressed in this work, the phytoplankton composition seems to have been structured by the climatic rainy / dry regime instead of just by the ongoing decrease in water level.
On the other hand, at least concerning the relative abundance based on density data, this study shows that the dominance of cyanobacteria displayed a decrease from nearly 100% in phase I to 75% in phase III, following the ongoing reduction of water level. This finding agrees with other studies performed in semi-arid reservoirs (Medeiros et al., 2015; Costa et al., 2016), that have shown, in contrast with the expectations, that a reduction in water level would led to an increase in cyanobacterial dominance (Naselli-Flores et al., 2007; Yang et al., 2016), other phytoplankton groups (such as cryptomonads and diatoms) could also improve their performance under scenarios of extreme drought.
In our study, although the cyanobacteria have dominated throughout the eighteen months of drought, the relative abundance of slender shade-adapted cyanobacteria in codon S1 decreased in phase II and III relative to phase I, due the rise of diatoms in codon P and D, beyond greenalgae in codons X1, F and J, most associated to nutrients enrichment environment (Padisák et al., 2009). In this way, the trajectory of phytoplankton succession in water supply reservoirs is not easy to predict, and other particular characteristics such as regional climate (rainy / dry seasons), reservoir type (size and depth), reservoir aging, watershed size and their multiple uses should be considered (Holz, 1997; Medeiros et al., 2015).
In addition to the species already mentioned in the diagnosis of temporal variability of phytoplankton community in Castanhão reservoir, we highlight the common frequency of C. raciborskii (SN) and Cylindrospermopsis sp. (SN) across the study, especially in middle and dam zones during phase I.
Cylindrospermopsis, a taxon belonging to functional group Sn, is associated to mixed environments with high temperatures (Padisák et al., 2009; Bonilla et al., 2012; Soares et al., 2013). Additionally, Cylindrospermopsis is a cyanobacteria with ability to fix nitrogen (Padisák, 1997; Molot, 2017), although this mechanism does not always seem to explain its success in water systems. In fact, C. raciborskii can tolerate variations in nutrient availability, being dominant under a large spectrum of trophic conditions raging from low (Moura et al., 2007) to high nitrogen (Chislock et al., 2014; Silva and Costa, 2015) as well as from low (Bonilla et al., 2012; Prentice et al., 2015) to high phosphorous (Soares et al., 2013). Some studies related this large plasticity of C. raciborskii to the existence of ecotypes with different environmental preferences (Piccini et al., 2011; Bonilla et al., 2012). Although in the Castanhão reservoir, C. raciborskii and Cylindrospermopsis sp. were not dominant species, the highest mean density of these two species together in phase I (Supplementary Tab. 3) was mainly associated to high ammonium-N, low soluble reactive phosphorus and low Secchi depth.
Taking into account the predicted expansion of C. raciborskii in water systems around the world under climate- change scenarios (Bonilla et al., 2012), the knowledge of the temporal dynamic of this species in years of extreme drought conditions brought by the present study is a result of great relevance. Particularly to C. raciborskii, it is very important to know the particular conditions that lead to its high abundance in water supply reservoirs due to its potential for producing cyanotoxins (Sant'Anna et al., 2008). C. raciborskii have been recorded as common and, in fact, dominant in many eutrophic to hypertrophic reservoirs in the Northeast of Brazil (Bouvy et al., 2000; Moura et al., 2007; Silva and Costa, 2015).
Last but not least in importance, we emphasize that, although the shifts in phytoplankton composition seem to be related to extrinsic effects (environmental variables) discussed here, further studies are needed to evaluate to what extent the intrinsic effects (e.g. predation and interspecific competition), could have also influenced the phytoplankton temporal dynamic. Indeed, both intrinsic and extrinsic effects can play a large role in phytoplankton dynamic in reservoirs (Yang et al., 2018).
Comparative analysis: Phytoplankton structure in 2006/2007 versus 2012/2013
In the first years after the complete flooding of the Castanhão reservoir (2006/2007), a previous study was performed by Molisani et al. (2010). Molisani et al. (2010) classified the system as a whole as mesotrophic based on chlorophyll a, Secchi depth and the total phosphorus concentrations, although the inner reservoir zone has been classified as eutrophic. The phytoplankton composition found in this previous period (2006/2007) reflected these mesotrophic/eutrophic conditions.
In turn, in the present survey, according results from Santos et al. (2017), the state index of the reservoir was oligotrophic in phase I and mesotrophic in the others phases. The phytoplankton composition has still indicated mesotrophic/eutrophic conditions, similar to the previous (2006/2007) study, although in phase I the reservoir was dominated by a slender cyanobacteria (Pl. limnetica/Pl. minor) very adapted to face phosphate-P deprivation. Both Molisani et al. (2010) and Santos et al. (2017) adopted the Trophic State Index developed by Lamparelli (2004), thereafter published by Cunha et al. (2013).
In both surveys, the phytoplankton biomass in the inner zone was lower than in the middle and dam zones, which is a common behavior in dam reservoirs (Kimmel et al., 1990). In 2006/2007, chlorophyll a ranged from 3.3±1.4 to 7.5±4.5 μg L–1 (Molisani et al., 2010), while in the present study from 2.9±0.5 to 5.0±2.1 μg L–1. The lower values of phytoplankton biomass observed recently (2012/2013) may be related to the increase in dominance of slender cyanobacteria over other larger groups that were well represented previously (2006/2007), such as diatoms, cryptomonads and green algae. Additionally, these results agree with Kimmel and Groeger (1986) model that predict a tendency of reduction in phytoplankton production in years after the initial phase of high productivity ('trophic upsurge'), which occurs soon after reservoir filling.
Also, similar to Molisani et al. (2010), we used the functional group approach from Reynolds et al. (2002) reviewed by Padisák et al. (2009) to study the phytoplankton dynamic. However, while in Molisani et al. (2010) the main taxa were defined as 'descriptive taxa', which are those with >5% of the total phytoplankton density (organisms L–1) in at least one sample, in this present work the main taxa have been defined as 'characterizing taxa', and they were obtained by the SIMPER procedure. Despite this, adopting the same criterion, when the list of the 'descriptive taxa' of that previous study (Supplementary Tab. 4) is crossed against the list of 'characterizing taxa' used here (Tab. 1), they match well. Therefore, since similar methodologies were adopted to study the phytoplankton community, which were the Reynolds's functional group approach and a close cut-off value to define the main phytoplankton taxa that were used as indicative of environmental conditions in Castanhão reservoir, the data of both studies could be compared.
In 2006/2007 the main taxa in the inner zone were some diatoms in codons C and MP, cryptomonads in Y, and green algae in J and X1. In turn, in the present work, the same zone was dominated by slender shade-adapted cyanobacteria from the functional group S1, probably related to turbid conditions and the decrease in the availability of inorganic nutrients. Indeed, slender cyanobacteria could be more suitable than green algae, diatoms and cryptomonads to face the reduction of nutrient concentrations, especially phosphate-P (Villena and Romo, 2003; Prentice et al., 2015). This drop in nutrient concentrations in years after the initial phase of reservoir filling agree with the model from Kimmel and Groeger (1986) to describe water quality changes in response to reservoir aging. As consequence of changes in nutrients and light conditions, shifts in phytoplankton composition are expected as well (Holz et al., 1997).
Although water transparency (estimated by Secchi depth) have increased from 2006/2007 to 2012/2013 (in all reservoir zones), the dominance of shade-adapted cyanobacteria (S1) was an indicative that the turbidity could be still considered high for phytoplankton. This increase in water transparency could be, at least in part, also related to a decrease in the relative abundance of large phytoplankton and an increase in dominance of slender cyanobacteria. In addition, S1 is typical of mixing layers and the lower water volume in 2012/2013 may have favored this FG. It is important to note that lower depth of the reservoir could have favored short-term events of higher turbidity due to the effect of strong winds upon a dropping water column depth, which may have not be detected by Secchi data (an instantaneous measurement), but are reflected by phytoplankton community (an integrative proxy). Additionally, the role of other factors, such as changes in zooplankton structure, that direct impact on the phytoplankton size and composition (Yang et al., 2018), cannot be ruled out as drivers of phytoplankton shifts in the Castanhão reservoir and should be investigated as well.
In the middle and dam zones, during 2006 / 2007 the dominance was already of cyanobacteria (Molisani et al., 2010), similar to the present study. The dominance of cyanobacteria belonging to the functional groups S1 and MP, typical of mixing and turbid systems (Padisák et al., 2009), persisted during 2012/2013, while the functional group Lo decreased in abundance. In fact, it is important to note the absence of Synechocystis aquatilis in 2012/2013, since it was an abundant species across the zones from Castanhão reservoir in 2006/2007 (Molisani et al., 2010). Synechocystis aquatilis belongs to functional group Lo, which is adapted to both low and high nutrients (Padisák et al., 2009), but is sensitive to prolonged and deep mixing (Reynolds et al., 2002). In this sense, the absence of the functional group Lo between the 'characterizing taxa' in 2012/2013, along with a reduction in the reservoir water volume, is indicative of best mixing conditions in 2012/2013 than in 2006/2007, associated with reduced volume and shallower waters as suggested by Santos et al. (2017).
From 2006/2007 to 2012/2013 the middle and dam zones also experienced a drop in nutrient concentrations, just as happened in the inner zone. Regardless of this, it is important to take into mind that even with a decrease in nutrient concentrations and phytoplankton biomass, the system remained still mostly dominated by mesotrophic/eutrophic taxa, especially in phase III. In 2012/2013, along the drought months, the system experienced an ongoing increase in trophic state from oligotrophic to mesotrophic, resulting in the rising of the phytoplankton species more associated to a nutrient enrichment environment.
Cylindrospermopsis raciborskii (SN), a potentially toxic cyanobacterium, was found in 2006/2007 and persisted as a frequent species in 2012/2013 in the middle and dam zones. In addition, other species have appeared, such as Cylindrospermopsis sp., which were absent in 2006/2007 (Molisani et al., 2010). Other cyanobacteria observed in the present study and absent in 2006/2007 were Pseudanabaena cf. biceps (MP) and Myxobaktron sp. (X1 ?).
Finally, we highlight the replacement of the dominant diatom Achnanthidium minutissimum (MP) in the middle and dam zones in 2006/2007 (Molisani et al., 2010) by other diatoms represented mainly by Nitzschia sp. (12-25 μm) (D), followed by Aulacoseira granulata (P) in 2012/2013. A. minutissimum is a species of smaller size than A. granulata and Nitzschia sp., which could indicate the more favorable mixing conditions to the diatoms of larger sizes in the middle and dam zones in 2012/2013. Indeed, according to Padisák et al. (2009), A. granulata (P) is associated to mixing layers, while Nitzschia sp. to shallow turbid waters.
Our study demonstrates that the temporal shifts in phytoplankton community in eighteen months of severe drought in a large reservoir from the Brazilian semi-arid region was associated with sub-surface phosphate-P, ammonium- N and nitrate-N concentrations, along with water transparency changes. We demonstrated a shift in phytoplankton composition, from slender cyanobacteria (S1 and MP) favored by turbid and poor phosphate-P conditions in Phase I to other slender cyanobacteria (FG ?) probably favored by increase in Secchi depth and decrease in ammonium-N in phase II. In turn, in phase III the ascension of opportunistic and fast-growing green algae (X1, F and J) were driven by the increase of phosphate-P and nitrate-N in the reservoir.
In general, although some species substitutions have occurred, especially in the inlet zone, when comparing the first years after the completing flooding of the Castanhão reservoir (2006-2007) with eight years later (2012-2013), the phytoplankton composition, with emphasis on the dominance of slender filamentous cyanobacteria, continued to indicate mesotrophic/eutrophic and turbid conditions of the system, though they have been facing the decrease in inorganic nutrients. The large dominance of functional group S1 (typical of turbid and mixing layers) in the last years was an indicative that the turbidity could still be considered high to phytoplankton, favoring the slender shade-adapted cyanobacteria, especially Planktolyngbya limnetica/Pl. minor and Pseudanabaena limnetica.
The dominance of cyanobacteria from the functional group S1 during most of the time in 2012/2013 was also an indicative of increase in mixing conditions in comparison to 2006/2007. On the same way, the best mixing conditions during drought years related to lower reservoir volume, was indicated by the lack of the functional group Lo between the 'characterizing taxa' in 2012/2013, along with the replacement of small diatoms (MP and C) by large diatoms belonging to functional groups D and P.


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Actress Dakota Johnson made sure producers banned everyone except essential crew members while filming racy scenes for Fifty Shades Of Gray after her mother Melanie Griffith told her she had a right to demand a closed set.
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A specially-prepared Volkswagen Beetle has driven at speeds above 200mph for over one mile at the Bonneville Salt Flats, making it the fastest Beetle in the world
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Amid the low thatch of five- and six-story apartment houses on Riverside Drive is a tall stand of buildings between West 155th and 158th Streets, eight- to 13-story apartment houses that stick out like a cottonwood grove on the prairie.
The Riviera, the Vaux Hall, the Sutherland and others were meant to appeal to the professional class that would otherwise take up residence lower down on Riverside or West End Avenue. Today, they remain out of character with their low-rise environment, forming a memorable enclave.
The land here originally plunged down to the Hudson River. When John James Audubon built a frame house in 1840 on land now between 155th and 156th Streets, he was moving to the country; the woodlands were rich with birds, elk and muskrat. Suburban development followed in the 1880s, and the area became known as Audubon Park.
The buildings came about a few years after 1900, when Riverside Drive was extended north of West 155th Street, destroying many houses in its path as it curved inland along its present line, atop an elevated landfill held back by a giant retaining wall. In 1908, Reginald Pelham Bolton, writing in The Record and Guide, complained the new street was "undesirable and even dangerous." But he observed that the newly created land would form a good site for "the highest class of institutional buildings."
Yet it was apartment-house developers who saw possibilities here, and from 1910 to 1915 five large residential buildings went up on this winding three-block stretch: the Vaux Hall (often spelled Vauxhall), 780 Riverside Drive, at 155th Street; the Rhinecliff (occasionally the Rhinecleff), 788 Riverside Drive, at the south corner of 156th; the Riviera, 790 Riverside Drive, between 156th and 157th; the Grinnell, 800 Riverside Drive, between 157th and 158th; and the Sutherland, at the head of the drive at 611 West 158th Street.Continue reading the main story
They were designed by four of the big-gun architects of speculative apartment houses: George and Edward Blum did the Vaux Hall; Schwartz & Gross did the Rhinecliff and the Grinnell; Rouse & Goldstone did the Riviera; and Emery Roth did the Sutherland.
They present an encyclopedia of styles. On the Vaux Hall the Blums used their characteristic brick and inset tile for the Arts and Crafts style for which they are well known. Schwartz & Gross gave the Rhinecliff a standard stocky West End Avenue-style facade, but with two ebullient round-topped shields at the top.
Rouse & Goldstone put up the huge Renaissance-style Riviera in three sections, reducing its apparent mass. Schwartz & Gross gave the Grinnell a Mediterranean-style facade with a central drive-in courtyard. And Emery Roth put a fanciful, even funny mansard and copper dome on his Beaux-Arts Sutherland.
Tenants were willing to trade the more central location of the West End Avenue area for the new subway going up Broadway and the spectacular views across the Hudson — the western exposure was completely open until the 1920s. These were big but not fabulous apartments, from five to nine rooms, although according to Matthew Spady, who lives in the Grinnell and has done extensive research on Audubon Park, that building had three duplexes.
The apartments attracted people like "Miss M. E. Berry, Pianist," who made her home in the Riviera. She advertised in The New York Tribune in 1915 that she could be engaged for parties: "My Dance Music Is an Inspiration to the Poetry of Motion." Did she ever play for Harry Sohmer, the president of the piano company that his father founded in 1872? In 1920 he was living at the Vaux Hall.
Servants were rare, although Rafael Brache, a diplomat from the
After World War I a sudden burst of co-oping took place, and in 1920 tenants bought the Vaux Hall for $750,000 from an investor who had paid $700,000 for it the previous year.
The West Side lost caste in comparison with the East Side after that time, and in the 1930s many of the large apartments were subdivided. By the 1960s drugs and crime were a fact of life in the neighborhood, and the once-elegant buildings fell into disrepair.
Things turned around in the 1980s, and a group of friends dedicated themselves to clearing out the vines from the mall that is a distinctive feature of this stretch of Riverside Drive. In 1925 it was named Charles and Murray Gordon Memorial Park, after two men killed in World War I, but in recent years has been known as Riverside Oval.
The vines were winning again by about 2000, and in 2004 Vivian Ducat, who lives in the Riviera — appalled to discover the little oval not only ensnared but buried in vodka bottles, parts of a broken fence and other detritus — called a meeting to revive it. Today Christina Ham Read is the head gardener; she says that in addition to pruning shrubs, this fall volunteers planted grasses and, for a bit of winter greenery, two holly trees.
"To me," Ms. Ducat says, reflecting on the effort, "it's what makes the neighborhood different. I grew up on the West Side, and I never really thought I could make a difference. But here I have."Continue reading the main story


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We've all been there before, the last minute invitation to a entree party after work. This leaves you absolutely no time to go home and refresh your make up. It's completely understandable that in today's world, which is full of the working, career-oriented business women, that our schedules vacate us little to nay beat for tangent up our makeup. Here are some quick and easy tips from makeup professionals to help circle your day time make up look to a fabulous night time look in no time.
The first rank in being able to flawlessly go from day to nighttime is to start with a polished day time look which can treffen added to later on.
It is important to start amidst a fresh base which can be achieved using a good cleanser and moisturizer. To finish preparing the skin, apply a primer. A primer will smooth the base and help the root to stay on for longer. Once you contain prepared the base, follow with a foundation that matches your skin and is evenly blended in. Then finish with a translucent powder.
Next apply make up to your cheeks. Professionals recommend utilizing a cream blush followed near a powder in a pop of color. It is recommended that your cheeks have a just pinched look. You could further top your cheeks off with some translucent powder or bronzer for a illusion of shimmer and staying power to last you all day.
A great day time look for your eyes is a nice neutral tone that you can house on later for your night time look. To achieve this look activate with a mat neutral all over the eye lid. Next apply a shimmery dusting of an eye shadow which will allot you a pretty shine throughout the day and help you to transition this look later. For your day opportunity look, try incorruptible lining your upper lids beside eye liner instead of upper and lower.
Lips also need to start out beside a good foundation. Apply a moisturizing lip balm in the morning and reapply throughout the day. You can add a lip lustrous on culmination of your lip balm for a shimmer and color throughout the day.
Now to turn your beautiful day look into a dramatic night time look; You resolution need to have a few supplies handy. If your face tends to acquire oily throughout the day, have some tissues on hand to blot the oil, then conceal where needed and finish your base touch up with a dusting of powder. You may poverty to add a bit more colour to the apple of your cheeks and bronzing highlighter to chin, cheeks and t-zone. To turn your eyes into that sexy, theatric night time look you will need to have eyeliner, mascara and an eye lash curler. Now is when you can add some eyeliner to your lower lids and reapply to your upper lids if necessary. Next you should crisp your eye lashes and add a coat or two of mascara for that extra ample look. For your lips you can add a pop of color. For example, this season reds and plums are the hottest colors so add a dramatic color to positively spice up your evening time look. With a few bare tips and tricks, you can easily turn your day time review into a night time look.


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Know more about this business than we do? Cool! Please submit any corrections or missing details you may have.Help us make it right
Your one stop solution for Contingency, Small Balance, Telecommunications, Auto Leasing, Retail, Student Loans, Legal, and Commercial Credit Collections
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Nelson Watson & Assoc is located at 80 Merrimack St, Haverhill, MA. This business specializes in General Contractors and has 2 review(s) with a star rating of 1.0.


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SOAX is an open source software tool to extract the centerlines, junctions and filament lengths of biopolymer networks in 2D and 3D images. It facilitates quantitative, reproducible and objective analysis of the image data. The underlying method of SOAX uses multiple Stretching Open Active Contours (SOACs) that are automatically initialized at image intensity ridges and then stretch along the centerlines of filaments in the network. SOACs can merge, stop at junctions, and reconfigure with others to allow smooth crossing at junctions of filaments.
ITK is an open-source, cross-platform system that provides developers with an extensive suite of software tools for image analysis. Developed through extreme programming methodologies, ITK employs leading-edge algorithms for registering and segmenting multidimensional data. It is widely used and contributed in the medical imaging field.
Reproducing an experiment doesn't stop at the bench when images are concerned. Icy is an open source bioimaging software package that aims to provide a framework for authors to share, and others to reproduce, research once the sample hits the microscope. Icy was released in April 2011 and is being developed at the Quantitative Image Analysis Unit at the Pasteur Institute in France by Jean-Christophe Olivo-Marin and his team.
Ported to Drupal by Drupalizing, a Project of More than (just) Themes. Designed by Zsolt Kacso


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A career in private tutoring can be profitable. Judge the various fees of private tutors and the factors that affect the price range.
Archive for the 'Work' Category
Should College Athletes Be Paid?
Paying college athletes is one of the major issues in collegiate sports today. Scrutinize the opinions going around today and decide if paying college athletes is necessary.
Where Should I Send My Demo Cd?
Sending out demo CDs involves more than just mailing the disc to a record company. Uncover the right way of sending out demo CDs and increase your chances of breaking in the music industry.


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Mandore was the capital of the marwar prior the foundation of Incredible Jodhpur which is situated 7-kms north from the Jodhpur junction. The popular local attraction of mandore is its royal based gardens with high rock terraces and most important there are the cenotaphs of the former rulers of Incredible Jodhpur.
The special attraction of these cenotaphs is that instead of usual Chatri-shaped as typical Rajasthan style, there are set in the beautiful landscape mandore garden along with the line of a Hindu temple. Here they are four stories with the fine columns and a unique spire, all in red sandstones. The most appealing dewal is of Maharaja Ajit Singh. Over the hills on a rocky outcrop the cenotaphs of Maharani's are set.
In this victorious Mandore garden you will get to see "the Shrine of the Three Hundred Millon Gods", fully surrounded by colorful bright images of Hindu Gods which gives us a pleasant feeling.


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|Ban Ki-moon arrives in Rwanda to discuss upcoming report on DR Congo rights violations
UN News - September 7, 2010
|Secretary-General Ban Ki-moon arrived today in Kigali, where he plans to meet with Rwandan leaders to discuss the soon-to-be-released United Nations report on serious human rights violations committed in the Democratic Republic of the Congo between 1993 and 2003.
|UN says peacekeepers 'failed' DR Congo rape victims
BBC News - September 7, 2010
|UN peacekeepers have "failed" the victims of mass rape in eastern DR Congo, a senior UN official has said. Atul Khare told the Security Council that the scale of systematic rape by armed rebels was far worse than feared. Atul Khare told the Security Council that the scale of systematic rape by armed rebels was far worse than feared.
|Government Launches Investigation into Latest Boat Accidents
VOA News - September 6, 2010
|Government spokesman Lambert Mende said the government is investigating the causes of two separate boat accidents over the weekend that killed at least 24 people with up to 60 others missing. Mende said the government is questioning those rescued from the waters after the accident to find out how they were able to get onto a cargo boat.
|Senegal ease pass DR Congo in Lubumbashi
Cafonline.com - September 5, 2010
|The Teranga Lions of Senegal got off to a winning start in their 2012 African Cup of Nations qualifiers with an away victory over the DR Congo in Lubumbashi. The Senegalese won by 4 goals. The boys of Amara Traoré opened scores at the 5th minute through Moussa Sow before captain Mamadou Niang added the second eleven minutes after.
|UN human rights chief announces release date for DR Congo Mapping Report
Office of the High Commissioner for Human Rights - September 2, 2010
|The UN High Commissioner for Human Rights Navi Pillay announced Thursday that the report of the Mapping Exercise documenting the most serious human rights violations in the Democratic Republic of the Congo between 1993 and 2003 will be made public on 1 October 2010.
|UN mission speaks out after human rights defender is kidnapped
UN News - August 31, 2010
|The United Nations peacekeeping force in the Democratic Republic of the Congo has expressed deep concern after the apparent abduction of a human rights defender who had spoken out against alleged rights violations by members of the military.
|Esperance beat Mazembe to inch closer
Cafonline.com - August 29, 2010
|Tunisian giants, Esperance inched closer to the semi finals of the Orange CAF Champions League with a 3-0 win over holders TP Mazembe in their Group A clash on Saturday in Rades.
|Key excerpts from UN report on Rwandan army genocide in DR Congo
BBC News - August 27, 2010
|The systematic attacks, in particular killings and massacres perpetrated against members of the Hutu ethnic group, are described extensively in section I of the report. These attacks resulted in a very large number of victims, probably tens of thousands of members of the Hutu ethnic group, all nationalities combined.
|UN report says Rwandan army committed genocide in DR Congo
BBC News - August 26, 2010
|The BBC has seen a draft UN report that says crimes by the Rwandan army and allied rebels in Democratic Republic of Congo could be classified as genocide. The report details how they targeted Rwandan Hutu refugees and Congolese Hutus in DR Congo, from 1993-2003. It lists human rights violations committed by security forces from all countries involved in what has been called an "African world war".
|UN Security Council calls for urgent action over mass DR Congo rape
BBC News - August 26, 2010
|The UN has said everything possible must be done to prevent atrocities like the recent rape of more than 150 women and children in the DR Congo. In an emergency session of the Security Council, Secretary General Ban Ki-moon said Congolese authorities had to fully investigate what had happened. The Council also said peacekeepers in the area should have done more to protect local people from rebels.
All news pages


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Mia Farrow launches fund for Darfur survivorsDecember 3rd, 2007 - 5:23 pm ICT by admin
London, Dec 03 (ANI): Concerned about the plight of survivors from the Darfur region of Sudan, actress and campaigner Mia Furrow has launched a new fund to support them.
The Fund4Darfur will use the money that is donated to help survivors in Darfur and those who have fled to the UK.
Miss Farrow said, at the launch at a hotel in north London, she was both baffled and frustrated by the indifference of British people to the plight of those in Darfur.
She was affected by the arrest of Gillian Gibbons a British schoolteacher teaching at Unity High School in Sudan. She termed the imprisonment as an "insanity and cruelty" of the Sudanese regime.
"We have to look at Darfur through the prism of Rwanda not the prism of Iraq, then it is very clear what we must do," Daily Snack quoted her as saying.
"The situation with Gillian Gibbons demonstrates the palpable insanity and cruelty of that regime, and this is only one woman. Imagine what would happen to her out of the public eye," she added.
The Aegis Trust, a leading anti-genocide charity, is running the new fund.
The Trust said that more than half of the villages in Darfur have been destroyed, two thirds of the population were reliant on some form of humanitarian aid, and more than 2.5 million people had been displaced by the conflict. (ANI)
Tags: aegis, british people, british schoolteacher, campaigner, darfur region of sudan, furrow, gibbons, gillian, humanitarian aid, insanity, launch, mia farrow, north london, plight, prism, public eye, sudanese regime, survivors, unity high school


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I just made a raised bed garden for my balcony. I posted a picture on Instagram and Facebook and got a bunch of questions asking what we used, how we put it together, etc.
So I made a short video explaining exactly what we did. I will make more videos as we go, explaining what to fill your raised bed garden with, what to plant in your raised bed garden, and the progress of our garden.
If you want to know when I am posting these videos, just sign up to be on the list and start receiving emails right away.


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I listen closely to my Buyers and Sellers and translate their needs and desires into an effective, practical system designed to achieve these unique goals. My sharp analytical ability is balanced with a warm sense of humour and a down-to-earth approach. I invite clients to ask any and all questions that they may have and I always provide the honest, straight-forward answer. I take great pride in the relationships that I form with my clients and value the trust that they have instilled upon me. My clients get the best of both worlds - I am pleasant and accommodating, while remaining results-oriented and thorough.
Having lived my entire life within the Greater Vancouver area, I have become exceptionally well-versed in Vancouver and neighborhood knowledge. Let me help you discover your next home in areas like Vancouver, Burnaby, New Westminster and beyond. Or, if you're seeking a more rural lifestyle, let me introduce you to Whistler, Squamish, and more.
My business is primarily derived by way of referals, but I'm always open to meeting with new clients and forming new long-term relations. If you or any of your friends or family are thinking of buying or selling a home, please give me a call with their name and direct contact number and I will follow up with them. My combined system of online and traditional marketing gives me the edge needed to make this possible!
Call me today and put my skills to work for you!


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New exhibition dates
The Lensflair Group December–January Sheffield Showcase display has come to an end, but we'll be exhibiting a new selection of framed prints in a Sheffield Showcase shop window from 6 to 27 April. The display will be at the junction of Pinstone Street and Cambridge Street in the city centre (purple pin on the map), and prints will be again for sale from the Sheffield Showcase office.
Looking a lot further ahead, we're very pleased to have been offered the opportunity to exhibit Peak District images at Castleton Visitor Information Centre in April 2016. More about that nearer the time.
Keywords: Lensflair Group exhibition
No comments posted.
Recent PostsLensflair Group offers: Christmas event 9 December News from Lensflair Group Major update for our website Lensflair's Castleton exhibition extended Lensflair Castleton exhibition opens Lensflair: The Peak Collection – Castleton exhibition New Lensflair greetings cards launched Christmas display – unique gifts for a friend or yourself? Lensflair Group prints on sale 17 October Lensflair Group displays coming up this autumn
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|Message view
|« Date » · « Thread »
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|Martin Sebor <se...@roguewave.com>
|Re: library and build sizes on Windows
|Mon, 19 Nov 2007 05:46:35 GMT
I've committed a script to build and run our examples, locales, and tests: http://svn.apache.org/viewvc?view=rev&revision=596034 You can see the expected format by running on UNIX. We're not using it for our nightly builds yet but I plan/hope to transition to it this week. Once that's done we need to do the same for Windows. Martin Martin Sebor wrote: > > Farid Zaripov wrote: >>> -----Original Message----- >>> From: Martin Sebor [mailto:firstname.lastname@example.com] On Behalf Of Martin Sebor >>> Sent: Monday, November 12, 2007 7:18 AM >>> To: firstname.lastname@example.com >>> Subject: library and build sizes on Windows >>> >>> I enhanced the cross-build scripts to extract the date of the >>> build and the sizes of the library and of the bildspace >>> (before and after running the clean target). This works on >>> UNIX but not on Windows where the date has a different format >> >> On unix: >> ----------------------- >> ### date: >> Wed Oct 31 09:38:50 UTC 2007 >> ----------------------- >> >> On windows: >> ----------------------- >> Wed Nov 07 22:23:49 2007: Loading solution... >> ----------------------- >> >> The difference in date format is the only UTC before year on unix :) > > Right. The problem is the line above it. The script looks for the RE > "^### date:" and takes the whole line below it as the date. We could > have it look for the Windows-specific string on Windows (and strip > what follows the timestamp) and the UNIX one on UNIX but I'd rather > we standardize on the same data format than complicate the script > logic. > >> >>> and the sizes aren't being reported. Would it be possible to >>> change the Windows infrastructure to add this info to the logs? >> >> The cleaning targets performing by the BATMAN, so to add the sizes >> to the logs the BATMAN scripts should be changed. > > Okay. We'll move this part (the script that runs the whole build) > to stdcxx very soon so we'll have direct control over its format. > > Martin > > -- View this message in context: http://www.nabble.com/library-and-build-sizes-on-Windows-tf4788960.html#a13828231 Sent from the stdcxx-dev mailing list archive at Nabble.com.
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Brand: E's Closet
Product Code: E429 coral
Availability: In Stock
Product Code: E429 coral
Availability: In Stock
$52.00 $29.99
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Exclusively E's! If you couldn't tell, we're really into backless maxis. Soft, stretchy material lays beautifully with a cinched waist separating a coral bottom and a white backless top.
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