Available online freely at www.isisn.org Bioscience Research Print ISSN: 1811-9506 Online ISSN: 2218-3973 Journal by Innovative Scientific Information & Services Network RESEARCH ARTICLE BIOSCIENCE RESEARCH, 2021 18(1): 259-275. OPEN ACCESS

Macrophytic vegetation and its associations in relation to environmental factors inhabiting a large river-channel,

Fawzy Salama1,*, Monier Abd El-Ghani2, Asmaa Mahmoud1 and Ahmed Amro1

1Department of Botany and Microbiology, Faculty of Science, Assiut University, Assiut, Egypt 2Department of Botany and Microbiology, Faculty of Science, Cairo University, , Egypt

*Correspondence: [email protected] Received 16-12-2020, Revised: 21-1-2021, Accepted: 01-02-2021 e- Published: 10-02-2021 The present study was aiming to characterize the floristic composition and structure of vegetation inhabiting different microhabitats along Ibrahimiya Canal, to find the relationship between environmental conditions and associated macrophytic communities, and to assess the role of the prevailing environmental factors that affect the diversity and distribution of vegetation. Between 2017 and 2018, 27 geo-referenced sample plots were selected to conduct this investigation along the main watercourse of Ibrahimiya Canal. The sampling of the vegetation was based on a nested plot design using a stratified sampling method. At each sampling plot, floristic data were collected from 3 different recognized microhabitats: (1) the water body, (2) the embankment slope, and (3) the terrace. Physico-chemical analysis for both soil and water were performed in the sampling plots. For soil, 12 parameters, for water 10 parameters, and 2 diversity indices were estimated. Classification (with TWINSPAN) and ordination (with DCA) were employed to analyze the presence/absence data matrix of 85 species × 27 sampling plots. The relationships between vegetation gradients and the studied environmental variables in both water and soil were examined using CANOCO, and a CCA biplot ordination diagram was elaborated. In TWINSPAN, 5 vegetation groups were identified and separated along the first and second DCA axes. CCA showed that significant variables in soil were EC, K, clay and silt, but in water were K, Na, Cl and Mg. Keywords:, Environmental monitoring, physico-chemical variables, TWINSPAN, CCA, microhabitats, diversity

INTRODUCTION total length of the canals and drains is In Egypt the River is the primary source approximately 47.000 km. of fresh water. It also provides Egypt with a very Rivers are diverse and dynamic systems that fertile and productive land along both the Nile play a crucial role in the complexity of the Valley and Delta regions. Of the total course of landscape (Chovanec et al. 2000). Their the River Nile, only the terminal 1530 km lie within ecological status is influenced by human activities, the borders of the country. It provides a complex affecting the physical properties of the riverbed, system including various forms of water bodies riparian vegetation and land beyond the riparian (lakes, marshes, streams, canals, drains… etc) zone (Petersen, 1998). These changes result in and land forms (plains and valleys). These worsened water quality and altered communities geomorphologic units (territories) encompass a of aquatic organisms, including macrophytes, great variety of climate, vegetation, land use, and which play important roles in energy flow, nutrient also a number of biogeographical regions. The cycling and sedimentation processes (Holmes, et Salama et al. Macrophytic vegetation along Ibrahimiya Canal, Egypt al. 1999; Gaberščik et al.2003). are the most important factor for submerged plant The occurrence patterns of macrophytes may growth and distribution, although nutrient be related to environmental factors that determine enrichment in water could inhibit the growth of the presence of different species. Thus, the some aquatic plants. Johnson and Ostrofsky floristic composition of aquatic plants in a (2004) demonstrated the importance of sediment hydrographic basin can be conditioned by abiotic characteristics in determining macrophyte characteristics and the physiological needs of community structure. Van Donk and Otte (1996) each species (Sass et al., 2010; Ceschin et al., reported that fish grazing on macrophytes affected 2012; Chappuis et al., 2014). The general the internal balance among autotrophic distribution patterns of these plants are mainly components by changing composition and related to variables such as luminosity, nutrient lowering the macrophyte standing crop. availability, temperature, pH, alkalinity, salinity, Middelboe and Markager (1997) and Armengol et speed of currents, water level variation, and al. (2003) reported that water depth is the most landscape elements such as geomorphology and important factor influencing water transparency shading (Janauer et al., 2010; Azzella et al., and hence distribution of submerged plants varies 2014). Biotic and abiotic variables act together on with depth. Water velocity not only affects the a population or an individual, and some abundance of submerged plants but also controls characteristics function as environmental filters for gas exchange processes (Sorrell and Downes, the occurrence of macrophytes (Ferreira et al., 2004). 2015). Aquatic conditions in the canals favour the Aquatic plants tend to respond to local development of abundant and diverse macrophyte environmental conditions (of the river stretch, communities (Caffrey, 1991). These features lake, or basin) faster than other groups of include shallow (<2 m), relatively clear water with organisms such as fish and insects, a response a stagnant or slow flow, deep and soft mud which determines the presence or absence of substrates, little bankside shading and a minimum certain species and life forms (Manolaki and of in-stream disturbance (Caffrey et al., 1998). Papastergiadou, 2013). It has been demonstrated Proliferation of macrophytes in many canal that local factors such as the environment’s sections interferes with amenity exploitation, physical structure and variability in chemical water principally navigation and angling (Caffrey, 1993). variables affect these plants more than other It is, therefore, necessary to implement weed large-scale factors such as climate (Alahuhta et control programmes that will meet the al., 2015). The submerged and floating species requirements of the different user groups. are more susceptible to environmental factors in According to Zahran and Willis (2009), the the aquatic environment because they are directly aquatic weeds of the Nile system of Egypt are in contact with the water, while emergent and some 35 species of 19 genera of 15 families amphibian species still have a great dependence (Täckholm, 1974). The plants are either entirely on the terrestrial ecosystem (Alahuhta et al., submerged, free-floating or their roots may 2013). penetrate the soil at the bottom of the stream. Studies intending to identify main factors Some of these bottom-rooting plants have floating explaining macrophyte occurrence or searching leaves. Zahran and Willis (2003) reported 36 for correlations between species abundance and communities dominated by 8 submerged, 9 environmental parameters in large rivers, like the floating and 19 emerged species that comprised River Nile, face several difficulties. Human the aquatic vegetation of the River Nile system of impacts (such as river regulation, bank Egypt. stabilization, navigation and urbanization, The aims of this paper are: i) to characterize agricultural and industrial activities) are constantly the floristic composition and structure of endanger and alter habitats and make vegetation inhabiting different habitats along ecosystems highly complicated (Birk et al., 2012). Ibrahimiya Canal, ii) to find the relationship Furthermore, vegetation assessment is often between environmental conditions and associated sparse and does not cover the entire length of the macrophytic communities, iii) to assess the role of hundreds of kilometers along river channels while the prevailing environmental factors that affect the the evaluation of macrophyte data can be diversity and distribution of macrophytic problematic in several aspects (Engloner, 2012). vegetation. The distribution and abundance of aquatic plants are influenced by many factors. Nutrients MATERIALS AND METHODS

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The study area plant remains and debris, then packed in plastic This study was conducted along Ibrahimiya bags ready for physical and chemical analysis. Canal; an irrigation canal in Egypt that was built in The soil texture was determined by the sieving 1873. Having its head on the left bank of the Nile, method as described by Jackson (1967). The soil in Assiut, it runs northwards for 60 km and then pH and electrical conductivity were measured in a divides in into two main branches; one soil-water extract (1:5 w/v). Soil reaction (pH) was branch is the Bahr Yussef Canal, while the other determined using an electric pH-meter (model pH- is the Ibrahimiyah Canal proper. This 350 km long 206 Lutron), and the electrical conductance (EC) canal, which is undoubtedly one of the largest was measured by means of conductivity meter artificial canals in the world, used to take off from (model 4310 JEN WAY). Sodium and potassium the Nile without any weirs or inlet works on the were determined by the flame emission technique river (the water of the river entered it freely). using a Carl-Zeiss DR LANGE M7D flame According to Morsy et al. (2002), it supplies photometer, calcium and magnesium contents perennial irrigation to 580,000 acres (2,300 km2) were determined volumetrically by the titration and flood irrigation to another 420,000 acres method using 0.01 N EDTA (Upadhyay and (1,700 km2). The discharge of the canal varied Sharma 2005), chloride contents were between 30 and 80 m3 second-1 in summer, and volumetrically determined as AgCl, and soluble between 500 and 900 m3 second-1 in flood time bicarbonate contents were estimated by titration (Barois, 1887). using the method described by Jackson (1967). Available climatic data obtained from the Contents of sulfates were estimated by Assiut University Meteorological Station at Assiut turbidimetry with BaSO4 using barium chloride and over the past nine years (2009 to 2018) indicated acidic sodium chloride solution (Bardsley and that temperature is regular in its seasonality. Lancaster, 1965). Phosphate contents were Winter months are cold and summer months are determined calorimetrically as phospho-molybdate hot. The lowest mean minimum was 11°C (Woods and Mellon, 1941). recorded in January while the highest mean In water, three samples were collected from maximum was 30 °C recorded in June and July. about 10 cm below water surface to avoid floating The relative humidity (RH%) varies markedly in matters. The surface water at sites 1, 3, 4 and 5 the different seasons of the year and in the suffers from uncontrolled pollution by wastes different hours of the day. The highest mean coming from agricultural runoff, sewage effluent, relative humidity was 50% recorded in December fertilizers and oils and detergents factories. In and the lowest mean was 24% recorded in May. these cases, the water, soil and plant samples Rainfall is scanty and irregular in both space and were taken about 50 m away beyond the pollution time. It occurs in the cold season from November sources. Water samples were directly transferred to May but summer is practically rainless. The to the laboratory to be processed for some mean maximum record occurred in December physical and chemical analysis. (0.5 mm), while the lowest was in May (0.01mm). The evaporation rate varied considerably in the Procedures of vegetation sampling and different months of the year. The highest mean analysis was 19.58 mm day-1 recorded in June and the The sampling of the vegetation was based on lowest mean was 6.24 mm day-1 recorded in a nested plot design using a stratified sampling December. method (Müller-Dombois and Ellenberg, 1974), whereby the plots were randomly distributed Collection and analysis procedures of soil and within a relatively homogeneous area in terms of water samples topography, landform and physiognomy of the Between 2017 and 2018, 27 sample plots present vegetation. At each sampling plot, the (geo-referenced using GPS model Trimble recorded species were collected from 3 different SCOUT®) were selected to conduct this habitats that recognized along the Ibrahimiya investigation along the main watercourse of Canal: (1) the water body, (2) the embankment Ibrahimiya Canal (Figure 1). Three soil samples (0 slope, and (3) the terrace (Figure 2). - 50 cm) were collected from each of the 27 sampling plots, mixed to form one composite sample, air-dried, thoroughly mixed and passed through a 2 mm sieve to remove large gravels,

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Figure 1: Location map of Ibrahimiya Canal, showing the studied 27 sampling plots.

Figure 2: Sketch showing the positions of the three recognized habitats at each sampling plot along the Ibrahimiya Canal.

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In each sample plot, presence/absence of direction of maximum variation, with their length each species was recorded, then a presence proportional to the rate of change (Ter Braak, percentage (P%) for each species was calculated. 1986). A Monte Carlo permutation test (499 Voucher specimens of each species were permutations; Ter Braak, 1990) was used to test collected, and identified at the Herbaria of Cairo for significance of the eigenvalues of the first University (CAI) and Assiut University (AST), canonical axis. All data variables were assessed where they were deposited. A complete checklist for normality (SPSS for windows version 16.0) of the recorded species, together with their life prior to the CCA analysis, and appropriate forms and chorological affinities were presented. transformations were performed when necessary Classification and ordination techniques were to improve normality according to Zar (1984). employed to analyze the vegetation. For this After the elimination of bicarbonates and sulfates purpose, presence/absence data matrix was used which have high inflation factors, twelve soil included 85 species and 27 sampling plots. The variables were included in this analysis: coarse floristic data matrix was then subjected to sand (CS), fine sand (FS), silt, clay, soil reaction classification by Two-Way Indicator Species (pH), electrical conductivity (EC), chlorides (Cl), Analysis (TWINSPAN) using the default settings Mg, Ca, Na, K, and PO4. of the computer program CAP (Community The TWINSPAN vegetation groups were Analysis Package, version 1.2) for Windows subjected to ANOVA (One-Way Analysis of (Henderson and Seaby, 1999) using the minimum variance) based on soil variables to find out variance as an algorithm, and a dendrogram was whether there were significant variations among elaborated. groups. Analysis of variance provides an insight The indirect gradient analysis was undertaken into the nature of variation of natural events, using Detrended Correspondence Analysis (DCA; which is possibly of even greater value than the Whittaker, 1967). Preliminary analyses were knowledge of the method as such (Sokal and made by applying the default options of the DCA Rohlfs, 1981). (Hill and Gauch, 1980) in the CANOCO program to check the magnitude of change in species Species diversity composition along the first ordination axis (i.e., Two species diversity measures were length of gradient in standard deviation units). The employed. Species richness (SR) within each default option of the computer program CANOCO separated TWINSPAN vegetation group was software version 4.5 (Ter Braak, 1987, 1990) was calculated as the average number of species per used for all ordinations. The indirect gradient sampling plot. The Shannon-Wiener diversity S analysis was undertaken using Detrended index (H′) was calculated from the formula H′= -Σi Correspondence Analysis (DCA; Whittaker, 1967). Pilog2Pi (Pielou, 1975), where S is the total DCA ordinates both sites and sampling plots in number of species and Pi is the presence terms of species and species in terms of sampling percentage (P%) of the ith species. plots and sites in which they occur from the same two way data matrix and plots the same results on RESULTS the same set of axes. Four principal axes were given of which any two or three can be used to Floristic composition produce the ordination plane. Usually, the first two Altogether, 85 86 plant species (36 37 axes represented the highest proportion of the perennials and 49 annuals), belonging to 70 variability in the data set, expressed in terms of genera from 29 different families were recorded. eigenvalues. Poaceae was the largest family (22 genera and 26 Canonical Correspondence Analysis (CCA) is species), followed by Asteraceae and Fabaceae a multivariate analysis technique developed to (7 genera and 7 species). Eight species in three relate community composition to known variation genera were recorded for Euphorbiaceae, three in the environment. It is a correspondence species and three genera were recorded for analysis that the axes were chosen in the light of Brassicaceae, and five species in two genera the environmental variables (Ter Braak, 1986). In were recorded for Amaranthaceae (Table 1). CANOCO, the relationships between vegetation Asclepiadaceae, Chenopodiaceae, Cyperaceae, gradients and the studied environmental variables Polygonaceae, and Solanaceae exhibited the can be indicated on the ordination diagram same number of the recorded species (2 for produced by CCA biplot. The variables in the CCA each). Potamogetonaceae included two biplot were represented by arrows pointing in the perennials: Potamogeton perfoliatus and P.

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Salama et al. Macrophytic vegetation along Ibrahimiya Canal, Egypt trichoides. The remaining familes were Glinus lotoides, Persicaria salicifolia and monospecific with one species in each family. The Polypogon viridis). Equal contributions (3 species) largest genera were Euphorbia (6 species), from each of Saharo-Arabian and Sudano- Amaranthus (4 species), and Setaria (3 species). Zambezian, Saharo-Arabian and Irano-Turanian, Seven genera represented by two species and Mediterranian and Euro-Siberian origins were (Chenopodium, Cyperus, Echinochloa, Setaria, indicated. The pluriregional taxa (11 species, Polypogon, Sorghum and Potamogeton. 12.9%) constituted species with wide Figure (3A) shows the life forms of the geographical range of distribution. Combinations recorded plant species according to Raunkiaer's of Mediterranean, Irano-Turanian and Euro- system (Raunkiaer, 1937). The total number of Siberian, Mediterranean, Saharo-Arabian and species recorded in the study area was 85 which Irano-Turanian, and Mediterranean, Saharo- can be categorized into 6 different life forms. Arabian and Sudano-Zambezian elements were Therophytes (57.6%) constituted the largest represented (Table 1). Cosmopolitan (19) and number of species (49 species), they included tropical species (12 palaeotropical and 11 among others, the most common weeds in Egypt pantropical) constituted 42 42 species or 49.4% as Amaranthus graecizans, A. hybridus, A. lividus, of the total recorded flora. Dactyloctenium aegyptium, Digitaria sanguinalis, Dinebra retroflexa, Echinochloa colona, Classification of the vegetation Polypogon monspeliensis, Portulaca oleracea, TWINSPAN analysis classified the floristic Sonchus oleraceus, Stellaria media, Trianthema composition of this study area to 5 vegetation portulacastrum, and Urtica urens. The groups (VGs). At the first TWINSPAN level hemicryptophytes (10.6%) included 9 species, (Figure 4), the 27 sampling plots were classified e,g., Alhagi graecorum, Convolvulus arvensis, into two major groups characterized by Imperata cylindrica, Mentha longifolia, Dactyloctenium aegyptium, Ziziphus spina-christi Paspalidium geminatum, and Persicaria salicifolia. and Senecio aegyptius. The second level was Geophytic species (7.1%) and represented by dominated by Dalbergia sissoo, Sorghum 6 species such as Cynodon dactylon, Cyperus halepense and Malva parviflora. The former two rotundus, Desmostachya bipinnata, and species separated groups A and B from the other Saccharum spontaneum. Phanerophytes (10 VGs. Also Echinochloa colona separated group B species; 11.8%) represented by arboreal species from A. On the other hand, Senecio aegyptius, such as Acacia nilotica, Dalbergia sissoo, Salix Phragmites australis and Pesicaria salicifolium mucronata, Sesbania sesban, Tamarix nilotica separated group E from C and D. Myriophyllum and Zizphus spina-christi. Hydrophytes (8.2%) spicatum, Phragmites australis, Phoenix represented by 7 species such as Myriophyllum dactylifera, Cynodon dactylon, Echinochloa spicatum, Potamogeton perfoliatus and stagnina, Pluchea dioscoridis and Malva parviflora P.otamogeton trichoides. Finally, cChamaephytes were represented in all groups. (4.7%) were represented by Chrozophora plicata, Group A (24 species, 2 sampling plots) was Cynanchum acutum, Sorghum halepense and dominated by Myriophyllum spicatum, Pluchea Panicum repens. dioscoridis, Phoenix dactylifera and Sonchus oleraceus (P = 100%). Species richness was Chorological affinities 15.0±8.485 species plots-1, and Shannon-Wiener The recorded species were classified diversity index was 2.62±0.60 (Table 2). Sampling according to their chorological affinities as shown plots of this group inhabited soil rich in coarse in Figure (3B). The monoregional species were sand, fine sand, silt and electric conductivity. The represented by the Saharo-Arabian species (3 water sampling plots showed high values in pH, species, 3.5%), and the Sudano-Zambezian Na and K. Panicum repens and Setaria viridis species (3 species, 3.5%). The five species with were consistent to this group. Eighteen sporadic American origin (5.9%) included Conyza species (P = 33%) were recorded in one sampling bonariensis and Trianthema portulacastrum. The plot. Irano-Turanian species was Pennisetum glaucum. Group B (60 species, 6 sampling plots) was The biregional species included (19 species, dominated by Phragmites australis, Cynodon 22.4%) belonged to two geographical origins dactylon, Pluchea dioscoridis, Malva parviflora (Table 1), 9 of them were of Mediterranean and and Sonchus oleraceus. The co-dominant species Irano-Turanian regions (e.g., Alternanthera (P = 66.7%) were Myriophyllum spicatum, sessilis, Amaranthus lividus, Cichorium endivia,

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Phoenix dactylifera, Dactyloctenium aegyptium dioscoridis, Ziziphus spina-christi and Setaria and Echinochloa colona. It was represented by viridis (Table 3). The average species richness the highest species richness of 20.83±8.73 was 16.00±4.58 species plots-1, and Shannon- species plots-1, and the highest Shannon-Wiener Wiener diversity index was 2.75±0.28. It inhabited diversity index of 2.94±0.52 (Table 2). The soil soil with the highest content of clay, salinity (EC) contents of clay, pH, K, Mg and HCO3 were the and the lowest sand content. The contents of highest amongst the others, while the lowest water showed highest values of Ca and Cl, the mean was recorded in fine sand. Water contents lowest Na, K, HCO3 and SO4. Four sporadic of Ca were the highest amongst the others, while species were consistent to this group (Avena the lowest mean was recorded in electric fatua, Potamogeton trichoides, Pennisetum conductivity. 32 sporadic species, included glaucum and Stellaria media). amongst others, Convolvulus arvensis, Rumex dentatus, Sisymbrium irio, Desmostachya Ordination of sampling plots bipinnata and Cyperus alopecuroides. Thirteen Application of Detrended Correspondence species showed consistency to this group (e.g., Analysis (DCA) to the vegetation data of Dactyloctenium aegyptium, Typha domingensis. Ibrahimiya canal (Figure 5) revealed the Tamarix nilotica and Urtica urens (Table 3). segregation of the 5 vegetation groups along DCA Group C (41 species, 10 sampling plots) axis 1 (Eigenvalue = 0.308) and DCA axis 2 characterized by Myriophyllum spicatum and (Eigenvalue = 0.267). The cumulative percentage Phragmites australis (P = 100%), and co- variance of species data of the first two DCA axes dominated by Phoenix dactylifera and was 22.4%. Sampling plots of groups A separated Potamogeton perfoliatus. Three consistent toward the negative side of DCA axis 1, while species to this group were Chenopodium murale, those of groups B separated along its positive Chrozophora plicata and Glinus lotoides. It has end. Meanwhile, the groups C, D and E were the lowest species richness of 11.30±4.99 species transitional in their ordination between the other plots-1, and the lowest Shannon-Wiener diversity VGs. index of 2.31±0.56. Soil gradients of this group were intermediate among other VGs, while they Correlation between soil variables and recorded the highest pH and the lowest silt vegetation content. Water contents of SO4 in the sampling Significant differences in the examined soil plots were the highest amongst the others. variables within the separated vegetation groups Group D (51 species, 6 sampling plots) was were demonstrated in Table (2). Soil contents of dominated by Myriophyllum spicatum, and co- silt and clay showed significant differences dominated by Sonchus oleraceus, Pesicaria between groups at p≤0.01. Water gradients within salicifolium and Senecio aegyptius (P = 83.3%). the 5 groups did not show any significant Its species richness was 18.50±6.89 species differences except for Cl, and Shannon-Wiener sampling plots-1, and Shannon-Wiener diversity index (p ≤ 0.01). Figure (6A) showed the CCA index was 2.85±0.41. The soils were ordination biplot with vegetation groups (A-E), and characterized by rich contents of coarse sand, Na, the examined soil variables. It can be noted that, K, Ca, Mg and Cl, whereas the water sampling sampling plots of group (A) showed correlation plots were characterized by rich contents of Mg with silt fraction, while those of groups (B) and (E) and PO4. On the other hand 25 sporadic species showed a correlation with silt, EC, Na and PO4. were recorded (e.g. Rumex dentatus, Anagallis Meanwhile sampling plots of groups (C) and (D) arvensis, Sisymbrium irio and Desmostachya showed high correlation with clay and almost all bipinnata). The four consistent species were estimated cations and anions. Mentha longifolia, Datura stramonium, Euphorbia The inter–set correlations resulted from prostrata and Melilotus indicus. Canonical Correspondence Analysis (CCA) of the Group E (31 species, 3 sampling plots) was examined soil variables were shown in Table (4). characterized by Myriophyllum spicatum, Malva CCA axis 1 was highly positively correlated with parviflora and Senecio aegyptius. Eight co- EC and highly negatively correlated with K. This dominant species included, amongst others, axis can be interpreted as EC-K gradient. CCA Phragmites australis, Echinochloa stagnina, axis 2 was highly positively correlated with clay Chenopodium album, Desmostachya bipinnata and highly negatively with silt. Thus, this axis can and Imperata cylindrica. Sporadic species be interpreted as clay–silt gradient. comprised 18 species such as Pluchea

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A B M+ES IT, 1.2 M+SA+SZ, 1.2 Cult., 1.2 SZ 3.5 3.5 M+SA+IT M+SA, 1.2 SA+SZ, 3.5 2.4

SA+IT, 3.5 COSM, 22.4 SA, 3.5

AM, 5.9 PAL, 14.1

PAN, 12.9 M+IT+ES, 9.4 M+IT, 10.6

Figure 3: Proportionate of plant life forms (A) and chorological analysis (B) of recorded species along Ibrahimiya Canal

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E col

Figure 4: TWINSPAN dendrogram showing classification of the studied 27 sampling plots of Ibrahimiya Canal, with the five vegetation groups (A-E). D aeg = Dactyloctenium aegyptium, Z spi = Ziziphus spina-christi, S aeg = Senecio aegyptius, D sis = Dalbergia sissoo, S hal = Sorghum halepense, M par = Malva parviflora, E col = Echinochloa colona, P aus = Phragmites australis, P sal = Persicaria salicifolium.

Figure 5: DCA diagram showing the distribution of the 27 sampling plots along the Ibrahimiya Canal within their five vegetation groups (A-E).

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A

0.8

Clay 2 axis CCA

Mg pH Ca PO4 CCA axis 1 EC Cl CS K FS Na GP A GP B

Silt GP C GP D H' GP E SR

-0.8 -1.0 1.0

B

GP A

1.0 GP B GP C

Cl 2 axis CCA GP D GP E

PO4

Na EC Ca CCA axis 1 K SO4

HCO3

pH

Mg

-1.0 -1.0 1.0

Figure 6: Canonical correspondence analysis (CCA) biplot of axes 1 and 2 showing the distribution of the 27 sampling plots of Ibrahimiya Canal, together with their vegetation groups (A-E) and soil variables (A), and water variables (B).

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Table 2: Mean values, standard deviations (±) and ANOVA values of the soil and water variables in the vegetation groups (A-E) of Ibrahimiya canal. EC=Electric conductivity (mS.cm-1), soil fractions (%), CS=Coarse sand, FS=Fine sand, SR=Species richness and H’=Shannon-Wiener index, *= p < 0.05, **=p < 0.01. Soil ions expressed by mg. g-1 soil DW, water ions expressed by mg l-1.

Vegetation groups Soil parameters F-value A B C D E CS 4.88±2.85 5.78±4.91 2.94±3.85 8.38±9.47 0.31±0.02 1.37 FS 6.78±2.95 6.55±6.18 3.45±3.41 6.08±6.22 0.49±0.07 1.21 Silt 49.67±6.97 47.81±4.38 28.17±9.85 30.96±12.23 31.06±19.56 4.40* Clay 38.66±1.17 39.86±10.88 65.45±14.40 54.58±26.17 68.14±19.49 2.90* pH 8.31±0.21 8.37±0.18 8.58±0.27 8.45±0.21 8.37±0.24 1.14 EC 3.41±2.58 2.20±1.90 2.67±2.86 2.84±3.62 4.70±3.08 0.41 Na 0.50±0.59 0.42±0.46 0.74±0.93 0.82±0.82 0.54±0.67 0.26 K 0.091±0.09 0.12±0.14 0.12±0.12 0.18±0.23 0.07±0.04 0.34 Ca 0.61±0.20 0.94±0.84 1.15±0.99 1.32±1.02 1.23±0.73 0.29 Mg 0.28±0.093 0.31±0.16 0.33±0.19 0.42±0.31 0.28±0.10 0.34 Cl 0.55±0.14 0.65±0.34 0.80±0.60 0.81±0.44 0.60±0.16 0.27 HCO3 11.18±1.44 12.20±2.32 12.00±2.24 11.35±1.75 11.86±0.59 0.2 Water parameters

pH 8.38±0.07 8.37±0.13 8.24±0.13 8.35±0.06 8.24±0.133 1.86 EC 0.23±0.01 0.22±0.005 0.23±0.01 0.23±0.003 0.23±0.01 0.064 Na 58.33±21.21 57.23±36.06 48.00±16.12 43.33±6.99 41.11±22.20 0.507 K 6.30±2.73 6.27±3.33 5.55± 1.70 4.99±0.69 4.72±1.96 0.456 Ca 45.00±2.36 52.78±5.74 50.67±6.05 47.22±4.91 52.22±5.09 1.323 Mg 25.42±1.44 21.69±4.57 23.38±5.85 26.77±2.99 22.37±5.38 0.986 Cl 68.04±4.19 74.45±3.93 76.62±7.04 82.34±9.62 224.83±131.67 9.06** HCO3 105.00±21.21 121.67±13.29 118.00±16.19 121.67±16.02 103.33±11.55 1.174 SO4 0.07±0.02 0.07±0.01 0.09±0.08 0.06±0.01 0.05±0.02 0.7 PO4 0.05±0.00 0.05±0.00 0.10±0.15 0.85±1.26 0.26±0.36 1.76 Diversity indices SR 15±8.48 20.83±8.73 11.30±4.99 18.50±6.89 16.00±4.58 2.3 H' 2.62±0.60 2.94±0.52 2.31±0.56 2.85±0.41 2.75±0.28 1.94

Table 4: Inter–set correlation of CCA analysis for the soil and water variables, together with eigenvalues and species–environment correlations in Ibrahimiya Canal,. fFor abbreviations, see Table (2). NI= Not included due to its high inflation factor.

Soil variables Water variables

Axes Axis 1 Axis 2 Axis 1 Axis 2 Eigenvalues 0.28 0.255 0.257 0.24 Species–environment 0.989 0.968 0.966 0.966 correlations Coarse sand 0.008 -0.035 - - Fine sand 0.156 -0.115 - - Silt 0.368 -0.432 - - Clay -0.298 0.341 - - pH -0.266 0.068 -0.178 -0.517 EC 0.38 -0.0003 -0.071 0.09 Na -0.343 -0.144 -0.463 0.127 K -0.66 -0.116 0.52 -0.028 Ca -0.495 0.026 0.265 0.066 Mg -0.427 0.078 -0.122 -0.68 Cl -0.55 -0.067 -0.073 0.61 HCO3 NI NI 0.069 -0.352 PO4 0.099 0.008 0.103 0.286 SO4 NI NI -0.0028 -0.056 Species Richness (SR) -0.207 -0.695 - - Shannon index (H') -0.17 -0.643 - - Monte Carlo permutation test (499 permutation) A test for significance with an unrestricted for the eigenvalue of axis 1 found to be significant

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(P = 0.049). act as sand controllers and bank retainers (e.g. Dalbergia sissoo, Ziziphus spina-christi, Phoenix Correlation between water contents and dactylifera and Pluchea dioscoridis). Grasses with vegetation wide range of distribution included Sorghum Figure (6B) showed the CCA ordination biplot halepense, Dactyloctenium aegyptium, with vegetation groups (A-E), and the examined Phragmites australis, Cynodon dactylon and water variables. It can be noted that, sampling Echinochloa stagnina were prevail. plots of group A and B showed correlated with pH The invasive hydrophyte Myriophyllum and Na. Whereas sampling plots of group C and spicatum was dominated along the Ibrahimiya D were highly correlated with almost all the Canal (Madsen, 1998). In Egypt, it vigorously estimated water ions (except Na). Members of invaded the southern provinces of the country group E were affected by Cl and PO4 only. The (Fayed, 1985) till it reached Lake Nasser at inter–set correlations resulted from Canonical Aswan (Ali, 2000). According to El-Kholy (1989), Correspondence Analysis (CCA) of the examined M. spicatum is invading the River Nile from Aswan water variables were shown in Table (4). CCA to Giza, and its presence in the Nile Delta has not axis 1 was highly positively correlated with K and been observed. Zahran and Willis (2003) reported highly negatively correlated with Na. So, this axis it as invasive weed in Lake Nasser (High Dam can be interpreted as K-Na gradient. CCA axis 2 Lake). M. spicatum might release allelopathic was highly positively correlated with Cl and highly substances (Nakai et al. 2000) which may reduce negatively with Mg. Thus, this axis can be growth in some other macrophytes. The alkalinity interpreted as Cl–Mg gradient. of water and soil of Ibrahymiya Canal may be the main factor that caused flourish of M. spicatum DISCUSSION (Toetz, 1997). It can also be noted the In Egypt, where a warm climate prevails most disappearance of some aquatic species such as of the year, the hydrophytes of the River Nile and Elodea canadensis, Wolffia hyalina, Zannichellia its irrigation and drainage systems are greatly palaustris, Nymphaea coerulea, Azolla filiculoides, developed (Zahran and Willis, 2009). The plants Lemna gibba, Eichhornia crassipes, of the Nile system of Egypt (about 553 species) Ceratophyllum demersum, Potamogeton crispus represent about 29.9% of the total flora of Egypt. and Ludwigia stolonifera which might be attributed Some 149 species are present in the Nile valley, to the severe human activities, and uncontrolled 291 occur in the Nile Delta and 64 characterize discharge of industrial wastes from factories along the Nile Fayium (Hassib, 1951). the Ibrahimiya Canal (Abd El-Ghani et al., 2010). In this study;, the floristic composition showed The life form spectrum is thought to be either that 85 86 species (37 perennials and 49 annuals) hereditary adjustment to environment (Zahran and were recorded along Ibrahimiya Canal belonging Willis, 2009). In this study, the dominance of to 70 genera from 29 different families. Poaceae, therophytes over other life forms seems to be a Asteraceae, Fabaceae and Euphorbiaceae response to the hot–dry climate, topographic constituted the main bulk (about 50%) of the variation and biotic influence (Heneidy and Bidak, studied flora. This result is consistent, more or 2001). The predominance of therophytes and less, with studies of Quézel (1978) in North Africa, relatively low representation of hydrophytes in and Mashaly (1987) in the north–east Nile Delta. such habitat may be due to the human impact Also, these families were reported to be the most which leads to destruction of this vegetation. Also, frequent in the Egyptian flora and were reported the high disturbance of these habitats may due to earlier by Springuel (1981), Springuel and Murphy the repeated removal of the silt and weeds from (1990), Mashaly et al. (2009), Abd El-Ghani et al. the water courses during routine maintenance (2010 and 2011), Ali (2014) and Amer et al. (Shaltout et al. 1995; Faried and Amro, 2016). (2015) as the most frequent families in the studied The occurrence of some rhizomatous geophytes Nile islands. This observation was not restricted to (e.g. Cynodon dactylon and Phragmites australis) the Nile islands but extended to agro-ecosystem might be coupled with their resistant to in Egypt as reported by Shaheen (2002), Abd El- decomposition under water submergence. Similar Ghani and Fawzy (2006) and Faried and Amro conclusion has been reached by El-Demerdash (2016). (1984), Mashaly (1987) and Shaltout et al. (1994). The floristic composition of the microhabitats The mixture of different floristic elements distinguished in this study was characterized by represented by variable numbers of species can several arboreal species (trees and shrubs) which be attributed to human impact, history of

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Salama et al. Macrophytic vegetation along Ibrahimiya Canal, Egypt agriculture and capability of certain floristic topography and vegetation over the time have a elements to penetrate the study area from several profound effect on the plant communities and adjacent phytogeographical regions (Mashaly, other biological systems that they support. 1987; Shalaby, 1995). It might be expected that Classification (TWINSPAN) and ordination the Mediterranean species would occur mainly analyses (DCA and CCA) resulted in the close to the Mediterranean coast including that of segregation of 5 vegetation groups (communities). the Nile Delta. However they frequently occur in Myriophyllum spicatum dominated four groups (A, the middle and south Nile Delta, and C, D and E). It was associated with Phoenix depression (Kosinová, 1975). According to dactylifera, Pluchea dioscoridis, Sonchus Kassas (1955), the Eu-Mediterranean species oleraceus and Ziziphus spina-christi in group A. range corresponds to two climatic belts, viz. the Meanwhile Phragmites australis, Sonchus Mediterranean coastal belt and . The oleraceus and Malva parviflora shared the other Mediterranean element and its combinations of three communities with M. spicatum in this study area (28.2%) considered to be lower communities (C), (D) and E, respectively. On the than those recorded in the Nile Delta (42.5%; other hand, sample plots of group B (located Mashaly et al. 2009). This relatively high between Dairout and Manfalout, Figure 1) percentage indicated the penetration of the dominated by grasses (Phragmites australis, Mediterranean phytogeographical belt in Egypt Cynodon dactylon and Echinochloa stagnina), southwards, and may represent a transitional weeds of arable lands (Malva parviflora and sector between the Eu-Mediterranean and the Sonchus oleraceus), the shrub Pluchea Saharo-Arabian regions. dioscoridis and the tree Ziziphus spina-christi. In the current study, the recognized Canal banks communities along Assiut vegetation groups showed some similarities in Governorate (hyper-arid region) were, somewhat, their floristic composition to those described by similar to those recorded in Nile Delta (Abd El- Serag and Khedr (1996) and Mashaly et al. (2001, Ghani et al. 2010 and 2011). However in the this 2003, 2009) with respect to the macrophytic Nnorthern part of the Nile, Abd El-Ghani et al. communities in the water bodies and along canal (2010) classified the canal banks’ vegetation of and drain banks in Nile Delta region. The species Nile Delta to 4 groups (1) Spirodella polyrrhiza- composition in this study was classified according Lemna minor (2) Potamogeton pectinatus- to their affinity to the three identified microhabitats Myriophyllum spicatum (3) Phragmites australis- (water, slopes and embankments). The most Lemna gibba with Echinochloa stagnina diversified habitat was the slopes (57 species, (emerged), Eichhornia crassipes (floating), and Table 1), while the least was the water bodies with Potamogeton pectinatus and Ceratophyllum three hydrophytic species (Potamogeton demersum (submerged) were common associates perfoliatus, P. trichoides and Myriophyllum (4) Phragmites australis-Eichhornia crassipes with spicatum). The low species composition of the two floating macrophytes (Pistia stratiotes and water bodies may be related to the fluctuations of Nymphaea lotus) as characteristic associates. water level, cleaning practices and human The vegetation group (B) was the most activities such as boating and fishing, while the diversified one and its associates inhabited soil high species composition of the slopes can be with rich contents in clay, pH, K, Mg, HCO3. The related to habitat heterogeneity as they cover presence of these minerals in high concentrations most of the environmental gradients from the may be the reason of this high diversity. On the open water bodies to terraces. This supports the other hand, the low diversity of group (C) species view that increasing habitat heterogeneity may due to the low content of silt. CCA revealed increases species diversity (Nilsson et al. 1991). that the right-half of the biplot represented group In this context, our results are in line with that (B) plots with soils rich in contents of silt, EC, Na obtained by Abd El-Ghani (2001) from the islands and PO4. Generally, EC, K, clay and silt may be of the middle part of the Nile Valley. Meanwhile, the most important environmental gradients which Shaltout et al. (1994) reached to the same those control the distribution and diversity of this conclusion in the canals and drains of middle Nile flora. Abd El-Ghani et al. (2010 and 2011) also Delta region. observed the high correlations between canal- Species distribution is generally determined bank communities of Nile Delta with EC, NO3, by climate and influenced by biotic factors and soil HCO3 and organic matter. characteristics (Parker, 1991). The differences in soil conditions produced by interaction of climate,

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CONCLUSION diversity around springs and wells in five The relationship between the environment oases of the Western Desert, Egypt. Int J (soil and water) and the macrophytic vegetation Agric Biol 8(2): 249-255. that inhabited different microhabitats along Abd El-Ghani MM, Abou-El-Enain M, Aboel Atta Ibrahimiya Canal, Assiut Governorate of Egypt A and Asaad E. (2011). Habitat was studied. Three different habitats were heterogeneity and soil-vegetation relations in recognized, varied in their floristic composition south of the Nile Delta, Egypt. Ecol Medit and community structure: (1) the water body, (2) 37(1): 53-68. the embankment slope, and (3) the terrace. The Abd El-Ghani MM, El-Fiky AM, Soliman A and most diversified habitat was the slopes, while the Khattab A. (2010). Environmental least was the water bodies. The invasive relationships of aquatic vegetation in the hydrophyte Myriophyllum spicatum was fresh water ecosystem of the Nile Delta, dominated along the Ibrahimiya Canal. The Egypt. Afr J Ecol 49: 103-118. application of CCA indicated that the distribution Alahuhta J. (2015). Geographic patterns of lake of species in these habitats was controlled by macrophyte communities and species edaphic variables such as fine sediments (silt and richness at regional scale. J Veg Sci 26(3): clay), electric conductivity and K. Water chemical 564-575. analysis showed the significance of K, Na, Cl and Alahuhta J, Kanninen A, Hellsten S, Vuori KM, Mg. Human disturbances along the Ibrahimiya Kuoppala M and Hämäläinen H. (2013). Canal played a paramount role in disappearance Environmental and spatial correlates of of several hydrophytes that were recorded in community composition, richness and status different parts along the River Nile. of boreal lake macrophytes. Ecol Indic 32:

172-181. CONFLICT OF INTEREST The authors declared that present study was Ali AH. (2014). Ecology and flora of plants of the performed in absence of any conflict of interest. Nile Islands in the area between Aswan and Esna, MS Thesis, Aswan University, Aswan, AUTHOR CONTRIBUTIONS Egypt. MAEG and FS designed and jointed the field Ali MM. (2000). Aquatic macrophytes in Egyptian trips with AA in data collection and management. Nubia pre- and post-formation of Lake MAEG and FS also wrote the manuscript. All Nasser. In: JF Craig, eds, Proceedings of statistical and multivariate analyses of the data ICLARM Conference on Sustainable Fish were carried out by MAEG. FS, MAEG and AA Production Lake Nasser: Ecological Basis reviewed the manuscript. All authors read and and Management Policy, Aswan, Egypt, pp approved the final version. 61–65. Amer W, Soliman A. and Hassan W. (2015). Floristic composition of Nile islands in Middle Copyrights: © 2021@ author (s). Egypt, with special reference to the species This is an open access article distributed under the migration route. J Amer Sci 11(6): 14-23. terms of the Creative Commons Attribution License Armengol J, Caputo L, Comerma M, Feijoó C, (CC BY 4.0), which permits unrestricted use, Garcīa JC, Marcé R, Navarro E and Ordońez distribution, and reproduction in any medium, J. (2003). Sau reservoir’s light climate: provided the original author(s) and source are relationships between Secchi depth and light credited and that the original publication in this extinction coefficient. Limnetica 22: 195–210. journal is cited, in accordance with accepted Azzella MM, Rosati L, Iberite M, Bolpagni R and academic practice. No use, distribution or Blasi C. (2014). Changes in aquatic plants in reproduction is permitted which does not comply the Italian volcanic-lake system detected with these terms. using current data and historical records. Aquat Bot 112: 41-47. REFERENCES Bardsley, CE and & Lancaster JD. (1965). Abd El-Ghani MM and Fahmy AG. (2001). Sulfur. Methods of Soil Analysis: Part 2 Analysis of aquatic vegetation in islands of Chemical and Microbiological Properties, 9, the Nile valley (Egypt). Int J Ecol Environ Sci 1102-1116. 27: 1-11 Barois EJ. (1887). Irrigation in Egypt, translated Abd El-Ghani MM and Fawzy AM. (2006). Plant by AM Miller. Government Printing Office,

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