HOME JOURNALS CONTACT

Pakistan Journal of Biological Sciences

Year: 2021 | Volume: 24 | Issue: 4 | Page No.: 537-547
DOI: 10.3923/pjbs.2021.537.547
Vegetation Dynamics and Growth Performance of Nitraria retusa among Different Habitats in Egypt
Ramadan A. Shawky and Yasser A. El-Ameir

Abstract: Background and Objective: Nitraria retusa is a salt-tolerant and drought-resistant shrub located in the Nitrariaceae family. Floristic composition and soil characters in representative habitats of Nitraria retusa were analyzed in terms of habitat variations and vegetation dynamics. Materials and Methods: A total of 12 sites were surveyed and nineteen environmental factors were recognized in three main habitats: sandy dunes, salt marshes and wadi channel. Homogeneity of each stand was secured by visual judgment to comprise uniform habitat dominated by Nitraria retusa. A list of the accompanied species (inside and outside the studied quadrates) was made to give an idea about the plant diversity in the study area. Results: Four main vegetation groups were recorded and their controlling ecological factors were identified. Species diversity gradients in addition to the gradient of human interference were significantly higher in dunes habitat than the other habitats. The growth performance of Nitraria retusa was significantly higher in the sand dunes habitat than in the other habitats. Conclusion: The Egyptian desert's need for judicious utilization and sustainable development. For this, the influence of other environmental factors needs to be analyzed properly understood.

Fulltext PDF Fulltext HTML

How to cite this article
Ramadan A. Shawky and Yasser A. El-Ameir, 2021. Vegetation Dynamics and Growth Performance of Nitraria retusa among Different Habitats in Egypt. Pakistan Journal of Biological Sciences, 24: 537-547.

Keywords: edaphic factors, habitat, performance, vegetation and Nitraria retusa

INTRODUCTION

Nitraria retusa ( Forssk.) Asch. (Arabic name: Ghardag or Ghargad) is a native salt-tolerant and drought-resistant shrub with many erect stems, spreading woody branches, fleshy leaves, white-to-yellowish green flowers and fleshy edible berry-like drupe fruits. Flowering occurs in April and May1,2. It inhibits three types of habitat: the salt marshes where it forms saline mounds that stud the flat ground of the salt marsh, the less saline sand bars (actual chains of sandy hillocks fringing the shoreline) and the channels of some main wadis near the coast3,4.

The positive relationship between habitat heterogeneity and species size structure is amongst the best-documented patterns in ecology5. Greater habitat heterogeneity is associated with greater species diversity for many taxa6,7. Most studies assume habitat specialization through correlative analyses between habitat and species distributions but manipulative experiments or rigorous analytical techniques such as null models can better determine the relative contribution of habitat specialization to species diversity7,8.

Growth performance of the shrubby populations in Egypt has been carried out with previous studies9-12 dealt with the vegetation analysis and species diversity associated with it. No doubt, that variation in the habitats and edaphic factors activities have adversely affect not only species zonation but also species diversity and growth performance of species13.

The aims of this study are analyzing the vegetation dynamics and assessing the soil-vegetation relationships of species associated with Nitraria retusa, also examine the growth performance of Nitraria retusa at habitat heterogeneity.

MATERIALS AND METHODS

Study areas: The study was carried out at Desert Research Center, Egypt from August, 2018 to March, 2019). The main habitat types used for this study were salt marshes (Siwa Oasis) at Lat. 29°13' 41'' N and Long. 25°18' 04'' E, fixed sandy dunes (Northern coast, Marsa Matruh) at Lat. 31°36' 50'' N and Long. 25°50' 10'' E and wadi channel (Suez Gulf) at Lat. 29°16' 20'' N and Long. 32°26' 80'' E (Fig. 1).

Fig. 1:
Map of Egypt showing the studied areas14

Vegetation survey: Vegetation surveys and assessing the ecological situation of plants were carried out in the year 2018 at three sites. Twelve quadrates (each of 20×20 m) were selected to represent the main habitats of Nitraria retusa. Homogeneity of each stand was secured by a visual judgment to comprise uniform habitat dominated by Nitraria retusa. A list of the accompanied species (inside and outside the studied quadrates) was made to give an idea about the plant diversity in the study area. The density and cover of each species have been estimated in each selected stand15. Relative values of density and cover were calculated for each plant species and summed up to give an estimate of its important value (IV) in each stand, which is out of 200. Nomenclature, identification and floristic categories were carried out according to reserachers1,2,16. Life forms were identified according to the scheme17.

Inventory of growth performance: For achieving the growth performance of Nitraria retusa, the height and mean crown diameter of each individual of the studied species in the whole stand was measured (based on 2 diameter measurements/ind.), the size index of each of Nitraria retusa individuals was calculated as the average of its height and diameter (H+D/2). The main, lateral branches and No. of leaves/tree, tree circumference (at DBH), leaf length and leaf area are measured. The number of seedlings was counted9,10.

Soil analysis: From each habitat, a composite soil sample was collected as a profile of 50 cm depth and air-dried. All samples were analyzed for soil texture, pH, Electrical Conductivity (EC), organic carbon, calcium carbonates, Na, K, Ca, chlorides and sulphates18,19.

Data analysis: The floristic data matrix of species was subjected for classification by two-way indicator species analysis (TWINSPAN, version 4.5) and Detrended Correspondence Analysis (DCA) into groups20. The relation between the vegetation and soil gradients was assessed using Canonical Correspondence Analysis (CCA)21. Plant diversity indices included Species Richness (SR), Shannon-Weiner diversity index (H’) and Simpson index. These diversity indices were estimated for each vegetation group22-24. Linear correlations coefficient (r) was calculated for assessing the relationship between the estimated soil variables on one hand and the community variables. The obtained data were statistically evaluated using CANOCO v. 4.5.

RESULTS

Floristic analysis: The total numbers of the recorded plant species surveyed in the present study are 53 species (3 annuals, one biennial and 49 perennial) related to 21 families. The largest families were Asteraceae and Poaceae comprising 7 species each, followed by Fabaceae 6 species, Chenopodiaceae 5 species Brassicaceae, Tamaricaceae and Zygophyllaceae comprising 3 species each, Apiaceae comprising 2 species. Other families were represented in only one species (Appendix 1). Life forms of the species recorded are grouped under eight types (Table 1 and Fig. 2). The majority of the recorded species are chamaephytes (25 species = 47.2%) followed by hemicryptophytes and nanophanerophytes (7 species = 13.2%) then by geophytes and therophytes (4 species = 7.55%) and helophytes (2 species = 3.77%). The lowest value of life forms is recorded as phanerophytes which attained (one species = 1.89%).

Chronological affinities: Chronological analysis of the surveyed flora (Table 2) revealed that 22 species (49% of the total flora) were Monoregional and (28.3% of the total flora) were bi-regional and (13.2% of the total flora) were pluri-regional and (9.43% of the total flora) were worldwide elements Monoregional chorotypes extending their distribution all over the Saharo-Arabian, Sudano-Zambezian and Mediterranean regions amounted to 49% of the recorded flora. On the other hand, Cosmopolitan, Panotropical and Paltropical chorotypes constituted five species. While Saharo-Arabian chorotype, either pure or penetrated other regions, was represented by 34 species of the total recorded flora.

Fig. 2:
Life form spectra of the recorded species in the three habitats dominated by Nitraria retusa
Ph: Phanerophytes, Ch: Chamaephytes, Hm: Hemicryptophytes, Ge: Geophytes, He: Helophytes, Pa: Parasites, Th: Therophytes, Nph: Nanophanerophytes


Table 1: Life form spectra of the recorded species in the three habitats dominated by Nitraria retusa
Life forms
Sand dunes
Salt marshes
Wadi bed
Total number
Ch
15
3
17
25
Hm
6
1
5
7
Nph
5
1
5
7
Ge
4
1
1
4
Th
3
1
1
4
Ph
2
2
2
3
He
2
1
1
2
Pa
1
0
1
1
Total
38
10
33
53
Ph: Phanerophytes, Ch: Chamaephytes, Hm: Hemicryptophytes, Ge: Geophytes, He: Helophytes, Pa: Parasites, Th: Therophytes, Nph: Nanophanerophytes


Table 2: Number of species and percentage of various floristic categories of the habitats dominated by Nitraria retusa
Floristic category
Sand dunes
Salt marshes
Wadi bed
Total number
Type
COSM
3
2
0
3
Worldwide
PAL
1
1
1
1
PAN
1
0
0
1
ME+ES+SA
2
0
1
2
Pluriregional
ME+IT+ES
1
0
1
1
ME+IT+SA
4
1
2
4
IT+SA
1
0
2
3
Biregional
ME+ES
1
0
0
1
ME+IT
1
0
3
3
ME+SA
5
2
7
8
ME
9
0
1
9
Monoregional
SA
7
2
11
13
SA+SZ
2
1
4
4
Total
38
9
33
53
COSM: Cosmopolitan, Pal: Paltropical, Pan: Pantropical, SA: Saharo-Arabian, ME: Mediterranean, SZ: Sudano-Zambezian, IT: Irano-Turanian, ES: Eurosiberian


Fig. 3:
TWINSPAN dendrogram of the 12 sampled stands dominated by Nitraria retusa based on the important values of species
Group I (Nitraria retua group), Group II (Nitraria retusa and Acacia tortilis group), Group III (Nitraria retusa group) and Group IV (Nitraria retusa and Arthrocnemum macrostachyum group)

Classification of vegetation: The application of TWINSPAN classification on 53 plant species recorded in 12 stands representing the study area yielded four vegetation groups (Fig. 3). Two species were recorded with variable presence values in the four groups. It included Nitraria retusa and Zygophyllum.

Sand dune habitat:

Group I (Nitraria retua group): This habitat was represented by one group, was diversified (34 species) among the recognized groups with 4 stands, with species richness of species/stands of 8.5, Simpson index 0.96 and Shannon-Wiener diversity index of 0.82. The other important species are Ammophila arenaria, Polygonum aviculare, Zygophyllum album and Suaeda vera recorded in this group (Table 3). Stands of this habitat were found on soil rich in its clay, organic matter, pH and CaCO3 and lowest levels of Na and Ca (Appendix 2)

Wadi channel habitat: This type of habitat represented by two groups as follow:

Group II (Nitraria retusa and Acacia tortilis group): It was the smallest among the separated vegetation groups. It was comprised of 8 species recorded from one stands, with species richness of species/stands of 8, Simpson index 0.86 and Shannon-Wiener diversity index of 0.77. The other important species, Lygeum spartum, Ochradenus baccatus, Crucianella maritima, Atriplex halimus, Cistanche phelypaea and Zygophyllum album (Table 3)

Table 3: Plant diversity, dominant and important species in each habitat dominated by Nitraria retusa
Community
Number of stands
Total species
Shannon-evenness
Simpson diversity
Dominant species Other important species
I
4
34
0.82
0.96
Nitraria retusa (47.30±15.94) Ammophila arenaria (13.25±2.36)*
Polygonum aviculare (12.03±11.34)
Zygophyllum album (11.55±8.05)
Suaeda vera (10.89±1.30)
II
1
8
0.77
0.86
Nitraria retusa (50.00) Lygeum spartum (24.40)
Acacia tortilis (46.20) Ochradenus baccatus (23.40)
Crucianella maritima (17.40)
Atriplex halimus (12.40)
Cistanche phelypaea (13.40)
Zygophyllum album (11.00)
III
3
33
0.85
0.89
Nitraria retusa (48.17±11.83) Acacia tortilis (36.43±13.67)
Halocnemum strobilaceum (24.47±3.27)
Zygophyllum album (11.53±6.51)
Reaumuria hirtella (10.47±9.08)
IV
4
16
0.79
0.81
Nitraria retusa (55.50±14.30) Halocnemum strobilaceum (32.20±5.26)
Arthrocnemum macrostachyum (47.73±3.73) Alhagi graecorum (19.33±2.87)
Tamarix aphylla (12.75±6.39)
Phragmites australis (12.33±8.59)
Zygophyllum album (10.60±9.08)
*Values are Means±Standard variation, Group I: Nitraria retua group, Group II: Nitraria retusa and Acacia tortilis group, Group III: Nitraria retusa group and Group IV: Nitraria retusa and Arthrocnemum macrostachyum group


Group III (Nitraria retusa group): The size of this group was represented by the three stands that included 33 species. The average species richness in this group is 11 species/stands, Simpson index 0.89 and Shannon-Wiener diversity index of 0.85. The other important species were Acacia tortilis, Halocnemum strobilaceum, Zygophyllum album and Reaumuria hirtella (Table 3). The stands of this habitat inhabited soil with the highest content of sand, clay and Ca and lowest levels of silt, clay, pH, EC, K and Cl (Appendix 2)

Salt marshes habitat:

Group IV (Nitraria retusa and Arthrocnemum macrostachyum group): This group was the most diversified among the recognized groups. It comprised 16 species recorded from 4 stands, with an average species richness of 4 species/stands, Simpson index 0.81 and Shannon- Wiener diversity index of 0.79. It inhabited soil with the highest water-holding capacity. The other important species were Halocnemum strobilaceum, Alhagi graecorum, Tamarix aphylla, Phragmites australis and Zygophyllum album (Table 3). Stands of this habitat were found on soil the highest content of silt, pH, EC, Na and Cl and the lowest level of clay (Appendix 2)

Ordination of stands: The application of DCA on 12 stands along axes 1 and 2 indicated that the vegetation groups yielded by TWINSPAN classification are distinguishable and have a clear pattern of segregation on the ordination plane except for groups C and D interconnected (Fig. 4).

Fig. 4:
Detrended Correspondence Analysis (DCA) ordination diagram of the 12 sampled stands
Group I (Nitraria retua group), Group II (Nitraria retusa and Acacia tortilis group), Group III (Nitraria retusa group) and Group IV (Nitraria retusa and Arthrocnemum macrostachyum group)

Stands of group I are separated at the upper part of the middle of the DCA diagram. Group II is clear segregated (the right side) along the two axes of DCA. While Group III is separated from the middle part axis 1 of the DCA diagram. On the other hand, group IV is segregated at the Left part along with the axis 2 of the DCA diagram.

Soil-vegetation relationships: The correlation between vegetation and soil characteristics is shown on the ordination diagram produced by Canonical Correspondence Analysis (CCA) of the biplot of species-environment.

Table 4: Comparison of growth performance (mean±SE) of Nitraria retusa at dunes, Salt marshes and Wadi channel
Habitats
Sand dunes
Salt marshes
Wadi channel
F-value p-value
Characters
Height (cm)
1.26±0.08
0.99±0.05
0.60±0.06
1.16 0.43**
Size index
5.14±0.21
0.35±1.0
0.16±8.38
1.3 0.41**
Trunk circumference
1.51±0.68
0.77±0.03
0.63±0.03
3.2 0.18**
No. main branches
4.08±0.36
5.58±0.58
2.50±0.19
5.45 0.00**
No. lateral branches
135.1±16.5
754.8±67.1
82.7±12.8
4.98 0.00**
Leaf number
6317.3±772.5
3301.3±412.84
1220.9±123.5
1.07 0.00**
Leaf length (cm)
1.73±0.07
1.62±0.06
1.72±0.07
2.13 0.09ns
Leaf area (cm)
1.09±0.03
1.20±0.05
1.11±0.07
1.14 0.41ns
No. seedlings
3.33±1.20
2.33±0.88
3.0±0.58
0.33 0.84ns
Significance levels are shown as **p<0.05


Fig. 5:
Canonical Correspondence Analysis (CCA) ordination diagram of plant species with soil variables. The indicator and preferential species are abbreviated to the first three letters of the genus and species, respectively

As shown in (Fig. 5) it is clear that the percentages of sand, silt, clay, CaCO3, Cl, SO4 and cations (Na, K and Ca) are the most effective soil variables, which showed highly significant correlations with the first and second axes of CCA ordination diagram. The dominant (Nitraria retua) and abundant species (Ammophila arenaria, Polygonum aviculare, Zygophyllum album and Suaeda vera) of group I are separated at the upper right side of the CCA-biplot diagram. These species in group I showed a close relationship with sand, organic matter and CaCO3. The dominant species (Nitraria retusa and Acacia tortilis) and abundant species (Lygeum spartum, Ochradenus baccatus and Crucianella maritima) in groups II and III are separated at the lower left side. These species in groups II and III showed a close relationship with sand and Ca. The dominant species (Nitraria retusa) and abundant species (Halocnemum strobilaceum, Alhagi graecorum, Tamarix aphylla and Phragmites australis) in groups IV are separated at the right side. These species in group IV showed close relationship silt, pH, EC, Na and Cl.

Growth performance of Nitraria retusa: The growth performance of Nitraria retusa was significantly higher in dunes habitats than in the other habitats (Table 4). Nitraria retusa twice as tall had main and lateral branches and attain nearly 2 times than the salt marshes and 6 times than the wadi channel. The size index and trunk circumference were higher in dunes than the other habitats. The total number of leaves compared with those growing in salt marshes and wadi channels. Besides, the reset of growth performance had a high value in the dunes than the other habitats.

DISCUSSION

The study on the relationship between vegetation patterns and their habitat heterogeneity is important to recover and rehabilitate the desert vegetation, stabilize the desert ecosystem and prevent desert expansion, also help in sustainable development. The studied localities represents a natural xeric habitat, which mainly dominated by Nitraria retusa. The natural plant in the present study is composed of 53 species (3 annuals, one biennial and 49 perennials) related to 26 families. The dominance of the perennials may be attributed to the nature of the habitat types, climatic and soil conditions25,26. The major families were Asteraceae, Poaceae, Fabaceae and Chenopodiaceae, which contributed collectively to about 47.2% of the total recorded plant species. This indicated that these four families are leading taxa and constitute the major bulk of the flora of the three habitats. Other researchers27,28 also reported similar results. Asteraceae is the largest and most widespread family of the flowering plants in the world29. In addition, Poaceae recorded the highest number of species because of its ability to grow in various habitats.

The life-form spectra are important physiognomic attributes, which is widely used by ecologists and chronologists in the vegetation and floristic studies30. Life forms of the flora in the present study showed that chamaephytes are the most represented form may be attributed to distinct defense against the physiological stresses and ability of these species to resist sand accumulation and grazing31,32. The life form spectra provide information, which may help in assessing the response of vegetation to variations in environmental factors33. Magurran24 pointed out that taxonomic diversity will be higher in an area in which the species are divided among many genera as opposed to one in which most species belong to the same genus and still higher as these genera are divided among many families as opposed to few.

From the floristic point of view, Egypt is the meeting point of floristic elements belonging to at least four phytogeographical regions: The African Sudano-Zambezian, the Asiatic Irano-Turanian, the Afro-Asiatic Sahro-Arabian and the Euro-Afro-Asiatic Mediterranean34. The floristic analysis of the present study indicated that the Saharo-Arabian taxa are represented by a relatively high percentage of plant species (64.2%) may be attributed to their capability to penetrate this region and to the influence of man and the history of agriculture. These taxa were either Pluriregional, Biregional or Monoregional. This was confirmed by previous stdueis25-27.

The results of the vegetation analysis had then been related to environmental data. Alternatively, vegetation habitat relationships have been derived from a single analysis of combined floristic and environmental variables21. The phytosociological investigation revealed that the vegetation structure of the three habitats was classified by TWINSPAN into four (I-IV) groups distributed in the three habitats. Each group comprises many sampling stands, which are similar in terms of vegetation and characterized by dominant and/or codominant species as well as, by many indicator and/or preferential species. Group I represented the fixed sand dunes habitat was dominated by Nitraria retusa, group II and III represented the wadi channel habitat, was co-dominated by Nitraria retusa and Acacia tortilis, group IV represented the salt marshes habitat, was dominated by Nitraria retusa. The identified vegetation groups, in the present study, were more or less agree with the previously mentioned studies10,11,14,34.

The most important soil gradients correlated with the distribution of vegetation as recognized by previous studies26,28,35 are soil salinity (EC), moisture gradient, soil fertility, organic carbon and phosphorus content, soil texture (sand, silt and clay) and pH value. In the present study, the application of Canonical Correspondence Analysis (CCA biplot) indicated that, the most important soil variables correlated with the distribution of vegetation types in the study area include soil texture (sand and silt), calcium carbonate, organic carbon, Na and Cl.

In this study, Leaf number, No. main and lateral branches of Nitraria retusa have been found to differ significantly among different habitats, however, plant height, size index and a number of seedlings can be directly compared among habitat types. The results of the study is similar to the findings of previous studies9,10,12,36 that the differences in growth performance can be the result of varying environmental factors (e.g., salinity, nutrient limitation and soil characteristics) among habitat types. Understanding the relationship between the edaphic factors and the distribution of plant species helps us to apply this finding in management, reclamations and development of semi-arid land grass ecosystems.

CONCLUSION

The present study provides an analysis of the floristic composition and vegetation structure of Nitraria retusa community growing in different habitats of Egypt to help in the management and conservation of these natural resources. It shows a wide soil range and occupies diverse habitats. Therefore, the conservation of natural habitats of this desert is of vital importance. The recorded 53 plant species in the present study can play a vital role in the economic and medicinal purposes. Hence, the Egyptian desert needs for judicious utilization and sustainable development. The distribution and growth performance of plant species is subjected to varying conditions of salinity concentrations, nutrient levels and substrate structure. However, the influence of other environmental factors needs to be analyzed properly understood.

SIGNIFICANCE STATEMENT

This study discovers the effect of habitat heterogeneity on the growth and vegetation associated Nitraria retusa that can be beneficial for the economic and medicinal purposes of studying species. This study will help the researcher to uncover the critical areas of plant growth and distribution that many researchers were not able to explore. Thus, a new analysis on growth performance of Nitraria retusa may be arrived at.

Appendix 1: Floristic composition of the three habitats dominated by Nitraria retusa in f Egypt, and mean of the importance values (out of 200) of the recorded species in different vegetation groups resulting from TWINSPAN classification
Habitat type
Vegetation groups
Family Species Life span Life form Chorotype Sand dunes Salt marshes Wadi bed I II III IV
Amaryllidaceae Pancratium maritimum (L.) Per G ME + 4.88±6.03
Apiaceae Deverra tortuosa (Desf.) DC. Per Ch SA + + 6.80±5.83 1.20±2.08
Asclepiadaceae Asclepias sinaica (Boiss.) Muschl. Per Ch ME+SA + 3.43±5.95
Asteraceae Achillea fragrantissima (Forssk.) Sch. Bip. Per Ch SA+IT + 2.64±4.58
Artemisia judaica L. Per Ch SA + 3.61±4.19 2.88±0.68
Centaurea aegyptiaca L. Bi Th SA + 9.15±8.13 7.54±5.31
Echinops spinosissimus (Turra) Per H ME+SA + + 0.23±0.47 5.20±7.26
Hyoseris radiata (L.) Per H ME + 1.91±2.80
Inula crithmoides (L.) Per Ch ME+ES+SA + + 3.90±3.49 2.50±4.33 2.87±1.84
Otanthus maritimus (L.) Hoffmanns. and Link Per Ch ME + 3.57±4.13
Boraginaceae Echium angustifolium (Mill.) Per H ME + 2.50±2.56
Brassicaceae Cakile maritima (Scop.) Ann. Th ME+ES + 1.01±1.117 1.55±1.04
Farsetia aegyptia Turra Per Ch SZ+SA + 0.22±0.39
Moricandia nitens (Viv.) Per Ch IT+SA + 1.40±2.43
Caryophyllaceae Gymnocarpus decandrum Forssk Per Ch ME+SA
Chenopodiaceae Atriplex halimus (L.) Per Nph ME+SA + + 5.91±1.47 12.4 2.76±4.77
Arthrocnemum macrostachyum (Moric.) Per Ch ME+SA + + + 2.31±4.00 47.73±3.73
Halocnemum strobilaceum (Pall.) M.Bieb. Per Ch ME+IT+SA + + + 24.47±3.27 32.20±5.26
Salsola kali L. Ann. Th COSM + + 2.62±3.19 8.53±10.58
Salsola tetrandra Forssk. Per Ch SA + + 7.08±8.78 1.56±1.50
Suaeda vera (Forssk. ex J.F. Gmel.) Per Ch ME+SA+ES + 10.89±1.30 8.76±0.74
Cistaceae Helianthemum lippii (L.) Pers. Per Ch ME+IT+SA + + 1.99±1.63 0.21±0.36
Convolvulaceae Cressa cretica L. Per H ME+IT + 0.39±0.68
Ephedraceae Ephedra alata Decne. Per Ch ME+SA + 2.18±2.49
Euphorbiaceae Euphorbia paralias (L.) Per Ch ME + 4.24±2.87
Fabaceae Acacia tortilis (Forssk.) Hayne Per Ph SA+SZ + 46.2 36.43±13.67
Alhagi graecorum Boise. Per H PAL + + + 0.58±1.15 1.59±2.76 19.33±2.87
Ononis vaginalis (Vahl.) Per Ch IT+SA + 7.43±9.48
Prosopis juliflora (Swartz) DC Per Ph SA + + 5.05±5.84
Retama raetam (Forssk.) Webb and Berthel Per Nph ME+IT+SA + 4.38±6.08
Lotus polyphyllus (E.D.) Clarke. Per Ch SA + 7.84±6.97
Nitrariaceae Nitraria retusa (Forssk.) Asch. Per Ph SA + + + 47.30±15.94 50 48.17±11.83 55.50±14.30
Peganum harmala L. Per H ME+IT+ES + + 1.15±1.34 1.92±2.25
Orobanchaceae Cistanche phelypaea (L.) Per P ME+SA + + 4.78±5.61 13.4 0.06±0.10
Plumbaginaceae Limonium pruinosum (L.) Chaz. Per H SA + + 1.78±3.55 0.42±0.73
Poaceae Aeluropus lagopoides (L.) Per Ch ME+IT+SA + 8.93±2.90
Ammophila arenaria (L.) Per Ch ME + 13.25±2.36
Elymus farctus (Viv.) Per G ME + 2.45±2.89 1.88±2.67
Lygeum spartum (L.) Per G ME+IT + + 0.15±0.30 24.4
Pennisetum divisum (Forssk. ex J.F.Gmel.) Henrard. Per He SA + + 1.25±1.45 0.05±0.09
Phragmites australis (Cav.) Trin. Per He COSM + + 12.33±8.59
Sporobolus pungens (Schreb.) Per G PAN + 0.37±0.74
Polygonaceae Polygonum aviculare (L.) Ann Th COSM + 12.03±11.34
Resedaceae Ochradenus baccatus Delile. Per Nph SA + 23.4 3.63±6.29
Rubiaceae Crucianella maritima (L.) Per Ch ME + + 1.38±2.77 17.4 6.83±11.84
Solanaceae Lycium shawii (Roem. and Schult.) Per Nph SA+SZ + + 6.93±8.12 3.19±4.63
Tamaricaceae Reaumuria hirtella Jaub. and Spach Per Ch ME+IT + 10.47±9.08
Tamarix aphylla (L.) Karst. Per Nph SA+SZ + + + 3.13±6.25 0.97±1.69 12.75±6.39
Tamarix nilotica (Ehrenb.) Bunge Per Nph SA + 6.44±5.69 3.44±4.31
Thymelaceae Thymelaea hirsuta (L.) Per NPh ME + 6.63±8.00
Zygophyllaceae Fagonia arabica L. Per Ch SA + 0.87±1.51
Fagonia mollis Delile Per Ch SA + 4.17±3.76
Zygophyllum album (L.) Per Ch ME+SA + + + 11.55±8.05 11 11.53±6.51 10.60±9.08
Per.: Perennial, Ann.: Annuals, Bi: Biennial, Th.: Therophytes, Ch. = Chamaephytes, H: Hemicryptophytes, He: Helophytes, G: Geophytes, Nph: Nanophanerophytes, COSM: Cosmopolitan, PAL: Paltropical, PAN: Pantropical, ME: Mediterranean, SA: Saharo-Sindian, IT: Irano-Turanian, ES: Euro-Siberian, SZ: Sudano-Zambezian


Appendix 2: Soil variables of the three studied habitats dominated by Nitraria retusa
Soil texture Chemical characteristics
Habitat Sampling sites Sand Silt Clay OM CaCO3 pH EC Cl SO42 Na+ Ca2+ K+
Sand dunes 1 74.7 20.5 4.8 1.34 43 7.76 1452 2.4 0.79 2.08 1.1 0.73
2 75.1 19.7 5.2 1.31 49 7.67 1389 2.33 0.81 1.99 1.2 0.75
3 75.5 18.9 5.6 1.28 55 7.58 1326 2.26 0.83 1.9 1.3 0.77
4 75.9 18.1 6.0 1.25 61 7.49 1263 2.19 0.85 1.81 1.4 0.79
Mean 75.30b 19.30b 5.40a 1.30a 52.00a 7.63b 1357.50b 2.30b 0.82a 1.95c 1.25c 0.76b
±SE 0.13 0.26 0.13 0.01 1.94 0.03 20.33 0.02 0.01 0.03 0.03 0.01
Wadi cannel 5 88.9 5.75 5.35 0.55 16.7 7.21 354 0.29 0.07 2.79 7.82 0.33
6 87.3 8.6 4.1 0.51 14.1 7.25 285 0.32 0.08 2.76 7.44 0.37
7 85.7 11.45 2.85 0.47 11.5 7.15 316 0.35 0.09 2.73 7.06 0.41
8 84.1 14.3 1.6 0.43 8.9 7.05 253 0.38 0.11 2.7 6.68 0.45
Mean 86.50a 10.03c 3.48a 0.49b 12.80b 7.17c 302.00c 0.34c 0.09b 2.75b 7.25a 0.39c
±SE 0.52 0.92 0.4 0.01 0.84 0.02 10.79 0.01 0 0.01 0.12 0.01
Salt marshes 9 78.3 20.5 1.2 0.21 10.6 7.85 2134 4.74 0.05 4.03 4.1 1.35
10 76.1 22.6 1.3 0.34 14.3 7.78 2100 4.55 0.06 3.89 3.6 1.27
11 73.9 24.7 1.4 0.47 18 7.71 2066 4.36 0.07 3.75 3.1 1.19
12 71.7 26.8 1.5 0.6 21.7 7.64 2032 4.17 0.08 3.61 2.6 1.11
Mean 75.00b 23.65a 1.35c 0.41b 16.15b 7.75a 2083.00a 4.46a 0.07b 3.82a 3.35b 1.23a
±SE 0.71 0.68 0.03 0.04 1.19 0.02 10.97 0.06 0 0.05 0.16 0.03
LSD0.05   3.28*** 4.33*** 1.57*** 0.17*** 8.96*** 0.16*** 94.20*** 0.24*** 0.03*** 0.20*** 0.76*** 0.11***
OM: Organic matter and EC: Electrical conductivity

REFERENCES

  • Tackholm, V., 1974. Students Flora of Egypt. 2nd Edn., Cairo University Cooperative Printing Co., Beirut
    Direct Link    


  • Boulos, L., 2000. Flora of Egypt, Volume 2: Geraniaceae-Boraginaceae. Al-Hadara Publishing, Cairo, Egypt, ISBN-13: 978-9775429223, Pages: 352
    Direct Link    


  • Kassas, M. and W.A. Girgis, 1965. Habitat and plant communities in the Egyptian desert: VI. The units of a desert ecosystem. J. Ecol., 53: 715-728.
    CrossRef    Direct Link    


  • Kassas, M. and M.A. Zahran, 1967. On the ecology of the red sea littoral salt marsh. Ecolo. Mono., 37: 297-315.
    CrossRef    Direct Link    


  • Stein, A., K. Gerstner and H. Kreft, 2014. Environmental heterogeneity as a universal driver of species richness across taxa, biomes and spatial scales. Ecol. Lett., 17: 866-880.
    CrossRef    Direct Link    


  • Clark, D.B., D.A. Clark and J.M. Read, 1998. Edaphic variation and the mesoscale distribution of tree species in a neotropical rain forest. J. Ecolo., 86: 101-112.
    CrossRef    Direct Link    


  • Harms, K.E., R. Condit, S.P. Hubbell and R.B. Foster, 2001. Habitat associations of trees and shrubs in a 50-ha neotropical forest plot. J. Ecol., 89: 947-959.
    CrossRef    Direct Link    


  • Messmer, V., G.P. Jones, P.L. Munday, S.J. Holbrook, R.J. Schmitt and A.J. Brooks, 2011. Habitat biodiversity as a determinant of fish community structure on coral reefs. Ecology, 92: 2285-2298.
    CrossRef    Direct Link    


  • Shaltout, K.H. and M.A. Ayyad, 1988. Structure and standing crop of Egyptian Thymelaea hirsuta populations. Vegetatio, 74: 137-142.
    CrossRef    Direct Link    


  • Shaltout, K.H., M.G. Shededw, H.F. El-Kady and Y.M. Al-Sodanyz, 2003. Phytosociology and size structure of Nitraria retusa along the Egyptian Red Sea coast. J. Arid Environ., 3: 331-345.
    CrossRef    Direct Link    


  • Al-Sodany, Y.M., 2003. Size structure and dynamics of the common shrubs in Omayed Biosphere Reserve in the western Mediterranean coast of Egypt. Ecolo. Mediterr., 29: 39-48.
    CrossRef    Direct Link    


  • Galal, T.M., 2011. Size structure and dynamics of some woody perennials along elevation gradient in wadi gimal, red sea coast of Egypt. Ecologia, 1: 56-67.
    CrossRef    Direct Link    


  • Shawky, R.A., 2018. Effect of edaphic factors on the vegetation zonation in some littoral and Inland salt marshes of Egypt. Bot. Lith., 24: 202-210.
    CrossRef    Direct Link    


  • Issawi, B. and E.S. Sallam, 2018. Stratigraphy and facies development of the pre-Cenozoic sediments in southern Egypt: A geodynamic approach. Arab. J. Geosci., Vol. 11.
    CrossRef    


  • Canfield, R., 1941. Application of the line interception method in sampling range vegetation. J. Forest., 39: 288-394.
    CrossRef    Direct Link    


  • Boulos, L., 1995. Flora of Egypt, Checklist. 1st Edn., Al-Hadara Publishing, Cairo, Egypt, Pages: 283
    Direct Link    


  • Raunkiaer, C., 1937. Plant Life Forms. 1st Edn., The Clarendon Press, Oxford
    Direct Link    


  • Allen, S., H.M. Grimshaw, J.A. Parkinson and C. Quarmby, 1974. Chemical Analysis of Ecological Materials Steward E. Allen, Editor (By Stewart E. Allen, H. Max Grimshaw, John A. Parkinson, and Christopher Quarmby with special contributions on Organic Pesticides by John A. Parkinson; X-ray Fluorescence Spectrometry by Colin C. Evans; Data Processing and Statistical Analysis by H. Max Grimshaw and David K. Lindley) John Wiley & Sons, A Halsted Press Book, New York, 1974, pages x + 565, $39.95 ISBN 0-470-02318-X Oxford University Press (OUP), New York, ISBN: 0-470-02318-X Pages: 565
    CrossRef    Direct Link    


  • Ryan, J., G. Estefan and Abdul Rashid, 2001. Soil and Plant Analysis Laboratory Manual. 2nd Edn., ICARDA, Syria, Pages: 172
    Direct Link    


  • Hill, M.O. and P. Šmilauer, 2005. TWINSPAN for Windows version 2.3. Centre for ecology and hydrology; University of South Bohemia, Huntingdon and České Budějovice. https://www.scirp.org/(S(i43dyn45teexjx455qlt3d2q))/reference/ReferencesPapers.aspx?ReferenceID=1567381


  • Ter Braak, C.J.F., 1987. The analysis of vegetation-environment relationships by canonical correspondence analysis. J. Plant Ecol., 69: 69-77.
    CrossRef    


  • Whittaker, R.H., 1972. Evolution and measurement of species diversity. Taxon, 21: 213-251.
    CrossRef    Direct Link    


  • Pielou, E.C., 1975. Ecological Diversity. John Wiley & Sons, New York, USA., ISBN:9780471689256, Pages: 174
    CrossRef    Direct Link    


  • Magurran, A.E., 1988. Ecological Diversity and its Measurements. 1st Edn., Princeton University Press, Princeton, New Jersey, ISBN: 978-94-015-7358-0, Pages: 179
    CrossRef    Direct Link    


  • Shaltout, K., L. Hassan and E. Farahat, 2005. Vegetation-environment relationships in south Nile Delta. Taeckholmia, 25: 15-46.
    CrossRef    Direct Link    


  • El-Amier, Y.A. and O.M. Abdulkader, 2015. Vegetation and species diversity in the northern sector of Eastern Desert, Egypt. West Afr. J. of App. Ecol., 23: 75-95.
    Direct Link    


  • Abd-El-Ghani, M.M., 2000. Floristics and environmental relations in two extreme desert zones of Western Egypt. J. Global Ecol. Biogeogr., 6: 499-516.
    CrossRef    Direct Link    


  • Shawky, R. and S. Keilani, 2020. Vegetation and seed bank in relation to microhabitat of wadi naghamish, north western coast of Egypt. Egypt. J. Desert Res., 70: 11-24.
    CrossRef    Direct Link    


  • Walter, D.R. and D.J. Keil, 1996. Vascular Plant Taxonomy. Kendall Hunt Publishing, Dubuque, USA
    Direct Link    


  • Cain, S.A. and G.M. de Oliveira Castro, 1959. Manual of Vegetation Analysis. Harper and Brothers, New York, USA., Pages: 325
    Direct Link    


  • Danin, A. and G. Orshan, 1990. The distribution of Raunkiaer life forms in Israel in relation to the environment. J. Veg. Sci., 1: 41-48.
    CrossRef    Direct Link    


  • Danin, A., 2012. Plants of desert dunes. Springer Science & Business Media, Springer, Berlin, Germany, ISBN: 978-3-642-60975-6, Pages: 177
    Direct Link    


  • Ayyad, M.A. and R.E.M. El-Ghareeh, 1982. Salt marsh vegetation of the Western Mediterranean Desert of Egypt. Vegetatio, 49: 3-19.
    CrossRef    Direct Link    


  • Zahran, M.A. and A.J. Willis, 2009. The Vegetation of Egypt. Springer, New York, USA
    Direct Link    


  • Salama, F., M.A., El-Ghani and N., El-Tayeh, 2013. Vegetation and soil relationships in the inland wadi ecosystem of central Eastern Desert, Egypt. Turk. J. Bot., 37: 489-498.
    CrossRef    Direct Link    


  • Quezel, P., 1978. Analysis of the flora of Mediterranean and Saharan Africa. Ann. Missouri Bot. Garden, 65: 479-534.
    CrossRef    Direct Link    

  • © Science Alert. All Rights Reserved