Taiwania 62(3): 321 330, 2017 DOI: 10.6165/tai.2017.62.321 Comparative study of autecological, morphological, anatomical and karyological characteristics of Acanthophyllum ejtehadii Mahmoudi & Vaezi (Caryophyllaceae): a rare endemic in Iran Zahra MALEKI SADABADI, Hamid EJTEHADI*, Parvaneh ABRISHAMCHI, Jamil VAEZI and Mohammad Bagher ERFANIAN TALEII NOGHAN Department of Biology, Faculty of Science, Ferdowsi University of Mashhad, Mashhad, Iran. * Corresponding author's email: hejtehadi@um.ac.ir (Manuscript received 8 December 2016; accepted 28 May 2017; online published 28 July 2017) ABSTRACT: Characterizing traits of a species in its habitat is substantial for planning a practical conservation program. Acanthophyllum ejtehadii Mahmoudi & Vaezi (Caryophyllaceae) is a recently established endemic species for flora of Iran and has a narrow non-conserved natural habitat in Radkan region of Chenaran, Razavi Khorasan Province. The soil and climate of Radkan was studied and the ecological, morphological, anatomical, and karyological aspects of A. ejtehadii were investigated to obtain a comprehensive knowledge about this species and its natural growth conditions. Field observations were performed during the growing seasons in 2014 2015 and 29 vegetation samples were collected as data. Results showed that this plant grows in mound-like sites on clay-loam soils at mean elevation 1279 m.a.s.l. in arid climate. Acanthophyllum ejtehadii is a thorn-cushion form chamaephyte plant. This Plant grow gradually in early-January, the flower unfolds in early-June and the matured seeds are produced in mid-July. A. ejtehadii is a diploid (2n=2x =30) species and has homogenous karyotype. Having unique morphological and anatomical adaptations such as expanded surface roots, reduced leaf area and thickened cuticle, this plant grows successfully in harsh environments. These mechanisms are specific to this specific kind of Acanthophyllum species. The Speciation time of this plant was estimated not more than 200 thousand years ago and if Radkan is preserved from anthropogenic disturbance, this species could expand its distribution area. KEY WORDS: Acanthophyllum ejtehadii, Anatomy, Autecology, Karyology, Morphology. INTRODUCTION Characterizing traits of a species in its habitat is substantial for planning a practical conservation program. These types of studies make a distinct discipline in ecology, called Autecology. Autecology studies mainly concern the relationship of a species to biotic and abiotic conditions in its habitats (Pianka, 2008; Köppler and Hitchmough, 2015). Different properties of a species are called its traits. These traits are representatives of the learnt behaviors in response to different events through the evolutionary history of that species. Autecology gives us the precise information about how a plant species responds to environmental changes (Köppler and Hitchmough, 2015). Iran is one of the large countries in South Western Asia with an area of approximately 1640000 square kilometers. Because of the different climatic conditions, caused by high elevation that surrounds the central parts of Iran, different ecosystems with unique features are created in Iran (Ghahreman and Attar, 1999). These unique natural conditions make Iran a homeland for about 8000 plant species, among them 1727 species are endemic (Jalili and Jamzad, 1999). Although there are a lot of endemic plant species in Iran, unfortunately a lack of knowledge about their characteristics and habitats made them vulnerable to blind human acts. It is necessary to study endemic species traits for planning sustainable usage and managing their natural habitats for conservative goals. Genus Acanthophyllum belonging to Caryophyllaceae has 80-90 species worldwide (Pirani et al., 2014). Acanthophyllum species are dwarf shrubs, perennial, with thorn shaped leaves and white or pink flowers (Schiman-Czeika, 1988). They are mainly distributed in arid lands and foothills of temperate regions (Heywood, 1985) and in the main part of steppes and subalpine vegetation in Central and Southwest Asia (Zohary, 1973). According to Takhtajan (1986), all species of this genus grow in Irano-Tournian floristic region. NE of Iran, Afghanistan and Turkmenistan are speciation and diversification zones of this taxon. Acanthophyllum ejtehadii Mahmoudi & Vaezi is a recently recognized species for flora of Iran whose natural habitat is limited to the northeast of Iran (Mahmoudi-Shamsabad et al., 2012). Molecular phylogenetic analysis supports that this species is a distinct species in Caryophyllaceae (Pirani et al., 2014). Heretofore there is no autecological research on A. ejtehadii. In this study, ecological, morphological, anatomical, and karyological properties along with soil texture, climatic and physiographical properties of the only habitats of A. ejtehadii were recorded, measured and described. This study aimed to present a reliable 321 Taiwania Vol. 62, No. 3 Fig. 1. Geographical location of Radkan region. comprehensive data about this species so as to be of use to its conservation. MATERIALS AND METHODS Study area Acanthophyllum ejtehadii grows in Radkan Region in the northwest of Chenaran city, Razavi Khorasan Province. The distribution of the A. ejtehadii is between 36°50 5.4 and 36°50 12.9 north latitude and between 59°00 56.4 and 59°01 10.1 east longitude (Fig. 1). This area geologically belongs to Kopet-Dagh zone, Shorije formation. Lithological studies have reported shale, gypsum and sandstone with Neocomian age construct as the main bedrock of this area (Moussavi-Harami et al., 2004). Climate and soil properties of Radkan In order to estimate climatic parameters of the study area, POWER’s Climatology Resource for Agroclimatology data for a period of 20 years (1994-2014) were downloaded. De Martonne Aridity Index (Oliver, 2005) and Emberger method (Villeneuve, 1980) were used for climatic classification of Radkan. Using the same data, the Ombrothermic Diagram of this area was also drawn. Physiographical characteristics of the study area including the elevation above sea level, slope aspect and slope degree for A. ejtehadii in the growing region were observed and recorded. Three soil samples near A. ejtehadii individuals from 0-20 cm depth were collected. Soil properties including pH, Electrical Conductivity (EC), texture, Nitrogen, Phosphorus and Potassium content and Calcium Carbonate percentage were investigated in the Pedology Laboratory of Agriculture Faculty of Ferdowsi University of Mashhad, Iran. 322 Autecological study Field sampling for autecological study of A. ejtehadii were performed in the 2014 2015 growing seasons. Considering the sparse distribution of this species, 29 selective plots with 1 square meters area, including one individual in the center of each plot, were used. In each sampling plot, the canopy diameter and cover percentage, phenological periods, revitalization strategies and life form (Raunkiær, 1943) of A. ejtehadii were recorded. The companion plant and insect species were also collected and identified through standard keys. Three random flowering individuals were selected for observing the root elongation pattern, root diameter and root depth. These individuals were also used for further morphological and anatomical studies. Morphological study Morphological characteristics of this species were measured in the Plant Ecology Research Laboratory of Ferdowsi University of Mashhad. Twenty six quantitative and seventeen qualitative characters were selected for morphological description of A. ejtehadii. Quantitative characters were pictured by Dino-Lite Plus AM313T Camera and measured by DinoCapture 2.0 software. In terms of Morphological structure, Mahmoudi et al. (2012) have categorized this species as closely related to A. diezianum. In the present study, the morphological characteristics of A. ejtehadii was compared with A. diezianum, A. pachystegium and A. lilacinum. Anatomical study In this study, anatomical characteristics of leaf, stem and inflorescence of A. ejtehadii (19 characters) were measured. For softening the woody parts of this plant, selected September 2017 Maleki sadabadi et al.: The characteristics of Acanthophyllum ejtehadii Table 1. Soil parameters of Radkan region Habitat Radkan Soil texture Clay Loam EC (ds/m) 0.48 ± 0.06 pH 8.013 ± 0.05 P (ppm) 6.53 ± 1.68 K (ppm) 403 ± 10 N (ppm) < 0.1 CaCo 3 38 ± 1.73 R Fig. 2. Ombrothermic Diagram of Radkan region, 20 years (1994-2014) data were used for drawing this diagram. Drought season begins in late April and lasts till October. organs were immersed in a mixture of 96% alcohol, water and glycerine in volume proportions 1:1:3 for one month. The softened organs were then fixed in Formalin-Acetic Acid-Alcohol (FAA 1:1:18 v/v). Semi-thin cross sections were manually cut and were then stained by Safranin-Fast Green protocol (Gerach, 1977). For recording leaf epidermis properties of A. ejtehadii, thin-layer crosses of leaves’ inferior surface were used. The prepared crosses were observed using a light microscope (Olympus BH-2) connected to a digital camera (Dino-Eye Am 423) for picturing. Anatomical features were measured by DinoCapture 2.0 software. Karyological study Hand-cut cross sections of radicles (about 1 cm in length) were used for karyological study according to Muller et al. (1991). The fixed slides were observed with Olympus Light Microscope BX-50 and Dino-Eye AM423 was used to picture the slides. Measurements were performed by DinoCapture 2.0. Software. Chromosomal sorting for the karyotype drawing was done in Adobe Photoshop CS5. For karyological analysis and determination of A. ejtehadii karyotype symmetry, five cells in the Metaphase stage were selected and the number of chromosomes and karyotype features including chromosomal formula, long and short chromosomal arm length (L & S), chromosome length (L+S), short/long arm ratio (L/S) and total form percentage (TF %) were calculated through Huziwara (1962) protocol. RESULTS Climatic and soil properties of Radkan area The natural habitat of A. ejtehadii is a mound-like region with an average 15° slope and a mean elevation of 1279 meters above the sea level. Average annual precipitation and temperature of Radkan is 221.23 mm and 13.18 C respectively. This area has arid and cold-arid climate based on De Martonne and Embereger climatic classifications. The Ombrothermic diagram of Radkan is shown in Fig. 2. Radkan’s soil consists of 30% clay, 33% silt and 37% sand grains. Therefore, it can be classified as a clay loom texture based on the United States Department of Agriculture soil classification system (2014). The other assessed soil parameters are noted in Table 1. 323 Taiwania Vol. 62, No. 3 Fig. 3. Acanthophyllum ejtehadii Mahmoudi & Vaezi in Radkan area. A-C Phenological period A. Flowering, B. Developing the mature seeds C. Dormant phase. D-J Morphological characteristics. D. Flowers and inflorescence, E. Cyma, F. Petals, G. Ovary, H. Bracts, I. Bracteole and 7- Calyx teeth. Autecological description of A. ejtehadii The vegetative growth of A. ejtehadii begins in early January and it lasts till May. This plant reaches its flowering peak in early June. Seed development begins 324 at late June and mature seeds are formed in mid-July. At the same time as seed dissemination stage, leaves abscission of individuals begin and plants gradually become dormant (Fig. 3). September 2017 Maleki sadabadi et al.: The characteristics of Acanthophyllum ejtehadii Table 2. 49 Morphological characteristics of A. ejtehadii. Quantitative characters are reported in µm (except for plant height which is reported in cm) units. No. 1 2 3 4 5 6 7 8 9 10 11 Mean ± SD Cushion-shape 7.4 ± 2.00 26.69 ± 3.30 Pallid or Gray 4.41 ± 0.48 Opposite Lanceolate 15.13 ± 1.74 1.16 ± 0.11 Dichasial Cymes 23.83 ± 3.70 No. 25 26 27 28 29 30 31 32 33 34 35 Characters Bract length Bract width Bracteole length Bracteole width Petals extra Calyx length Calyx shape Calyx length Calyx width Number of calyx teeth Calyx teeth size Calyx teeth shape Mean ± SD 8.84 ± 1.06 1.22 ± 0.10 7.06 ± 0.87 0.88 ± 0.10 4.35 ± 0.63 Tubular 10 ± 0.89 1.64 ± 0.11 5 Unequal Narrow Triangular 14 ± 2.00 36 Calyx teeth length 1.61 ± 0.19 3 37 Calyx mucronum length 0.63 ± 0.09 17 18 19 20 21 22 Characters Plant growing shape Plant height Flowering branch height Stem color Internodes length Phyllotaxy of leaves Leaves shape Leaves length Leaves width Types of inflorescence Inflorescence diameter Number of cyma partialis in inflorescence Number of flower in cyma partialis Lower floral leaves spreading Superior floral leaves spreading Principal inflorescence peduncle length Flower pedicel length Floral leaves shape Floral leaves length Floral leaves width Bracts spreading Bract’s color at neck 23 12 13 14 Horizontal 38 Number of petal 5 Erect 39 Petal’s neak shape Entire or Sharp 4.51 ± 1.50 40 Petal’s colour Pink 0.77 ± 0.14 Lanceolate Erect 1.62 ± 0.20 Erect Purple 41 42 43 44 45 46 Petal length Petal width Number of Stamens Number of ovules in ovary Filament length Style length Bract’s color at base green 47 Hairs on stem 24 Bract shape Acicular 48 Hairs on leaves 25 Bract length 8.84 ± 1.06 49 Hairs on calyx 26 Bract width 1.22 ± 0.10 14.15 ± 0.72 1.70 ± 0.13 10 4 14.30 ± 0.76 13.98 ± 1.11 Covered with short glandular and unicellular hairs Covered with short glandular hairs Covered with short glandular, unicellular and multicellular hairs 15 16 According to Raunkiær plant life-form (1934), A. ejtehadii is a chamaephyte plant. Average canopy of this taxon is 7.65% in 1 square meter area, with 7 27 cm diameter in flowering individuals. This species distributes by wind dispersion of mature seeds. It must be noted that the dormant individual regenerates in every growing season. Red root of A. ejtehadii extraordinary grows in a shallow system. Tap root grows 5 7 centimeters downward and then divides into 2 3 parts; these parts form the sinker roots, grown at radius of 16 50 cm, near soil surface. 27 plant species belonging to 18 families were observed with A. ejtehadii in the sampled plots. Among these species, Rosa persica Michx. ex Juss., Prunus spinosissima Franch and Artemisia scoparia Waldst were observed in all of the plots. Thrips sp. (Thysanoptera) was abundantly observed in A. ejtehadii flower calyx. Morphological characteristics Acanthophyllum ejtehadii is a cushion form dwarf shrub with 4 11 cm heights in flowering individuals. Its stems are gray or light-gray, covered by globular and unicellular trichomes, with internodes of 4±0.5 cm in length. Leaves are opposite, lanceolate with 1 1.3 mm in length (Fig. 3). Morphological properties are described in Table 2. The bracts in A. ejtehadii and A. diezianum are acicular and erect and the floral leaves are lanceolate (the lower floral leaves are horizontal, while the upper floral leaves are erect). The sprouts in A. ejtehadii branch from the middle of the stem while in the A. diezianum the sprouts branch from the base as well. Moreover, in A. ejtehadii the stem, leaves, peduncle and calyx are covered with short and long glandular hairs and single-celled and multicellular simple hairs, whereas the same organs in A. diezianum are covered with only multicellular simple hairs. In A. lilacinum and A. pachystegium, the floral leaves are subulate (the lower floral leaves and the upper ones are semi-horizontal and curved, respectively) and the bracts are subulate and curved. Anatomical characteristics (Fig. 4) Measured anatomical properties of A. ejtehadii are described in Table 3. Stem and inflorescence cross 325 Taiwania Vol. 62, No. 3 Table 3. Anatomical characteristics of A. ejtehadii. The measured characteristics are reported as Mean ± SD micrometers. Characters Length of cross section Width of cross section Size of Cuticle Size of Epidermal layer Size of Parenchyma layers Size of Sclerenchyma layers Length of vascular bundle Width of vascular bundle Size of Phloem Size of xylem Length of Pith Width of Pith 959.256 ± 17.76 805.21 ± 4.57 3.93 ± 0.28 12.27 ± 3.42 17.10 ± 4.96 193.74 ± 22.82 466 ± 7.67 346.70 ± 9.2 29.47 ± 3.38 64.34 ± 8.79 380.60 ± 3.93 161.73 ± 11.41 827.71 ± 10.22 589.70 ± 10.05 3.73 ± 1.31 7.59 ± 1.23 21.39 ± 8.49 143.97 ± 13.98 440.51 ± 22.73 283.94 ± 17.20 22.46 ± 4.70 75.86 ± 7.35 278.48 ± 27.21 100.153 ± 7.179 Fig. 4. Anatomical characteristics in Cross-section of A. ejtehadii. A-C. Leaf. D-F. Stem. 326 903.84 ± 9.38 449.23 ± 16.5 3.67 ± 0.86 13.21 ± 0.91 68.17 ± 14.67 333.36 ± 17.54 108.56 ± 14.63 42.40 ± 6.58 - September 2017 Maleki sadabadi et al.: The characteristics of Acanthophyllum ejtehadii sections of this species are ellipsoid, both including (from outer to inner) epidermis with thick cuticle, with 2 4 layers of parenchymal cells, sclerenchyma, vascular bundles, and pith with parenchymal cells having calcium oxalate crystals (Fig. 4). Palisade parenchyma cells with calcium oxalate crystals lie down beneath the epidermis layer in a leaf’s cross sections. Sclerenchyma cells with vascular bundles fill the mid area (Fig. 4). Acanthophyllum ejtehadii’s leaf epidermis have 88.48 ± 10 µm length and 49.94 ± 11 mm width, its cell walls are anticlinal curling. Anomocytic stomata of this plant have 30.81 (Fig. 5). Fig. 5. Lower epidermis of A. ejtehadii. This species has an anomocytic stomata. Karyological characteristics Acanthophyllum ejtehadii is a diploid species with 2n=2x=30 chromosomes and x=15 basic chromosome number. Its longest and shortest chromosomes have 2.96 and 2.03 µm length, respectively. The karyotypic formula of this species is 12m + 3sm and TF=43.175 (Fig. 6). Full list of A. ejtehadii karyological features are noted in Table 4. Discussion In this study, ecological, morphological, anatomical, and karyological characteristics of A. ejtehadii, a recently known species to the world’s flora along with the features of its only habitat in the northeast of Iran, were studied. Radkan drought season begins at late-April and continues for near 6 months till late-October. The mean evaporation rate in drought seasons is high, where the water stress can limit the plant growth. Water stress adversely changes many normal functions of plant species. For successful survival and reproduction, plants must respond to these changes. With two Table 4. Karyological properties of A. ejtehadii. L is the longest arm length, S is the shortest arm length. m is the metacentric and sm is the submetacentric chromosome Pairs 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 L 1.50 1.51 1.56 1.48 1.68 1.37 1.31 1.29 1.29 1.47 1.47 1.24 1.26 1.16 1.24 S 1.40 1.30 1.12 1.15 0.92 1.20 1.23 1.09 1.03 0.85 0.79 0.99 0.94 0.97 0.84 L+S 2.90 2.81 2.68 2.63 2.60 2.57 2.54 2.38 2.32 2.32 2.26 2.23 2.21 2.13 2.08 L/S 1.07 1.17 1.39 1.29 1.82 1.14 1.06 1.19 1.26 1.72 1.87 1.25 1.34 1.19 1.47 Type m m m m sm m m m m sm sm m m m m different strategies of drought escape and drought tolerance, plants are adapted to water deficiency (Taiz and Zeiger, 2010; Kooyers, 2015). Rapid development for efficient grow in wet season is a cost-effective mechanism for escaping harsh conditions of water shortage (Mitra, 2001). Drought escape is the response of A. ejtehadii to dry season, with rapid growing and early senescence. It seems that the drought tolerance mechanism is specific to this species as well as the other Acanthophyllum spp.. For instance, flowering stages of A. yasamin-nassehiae Joharchi & Pirani and A. pachystegium (grown in Radkan region) occur in late summer (Pirani et al., 2013) and early July (field observations), respectively. Many desert plants use this mechanism to avoid harsh environmental conditions caused by dry seasons (De Micco and Aronne, 2012). Shortening the life cycle is not the only adaptation of A. ejtehadii in arid environments. Reducing leaf surface, thickening cuticle and developing multilayer sclerenchyma are some anatomical features considered as a response to dry environments. Reduction in leaf number and leaf surface are considered as an effective strategy to reduce evaporation from surface of the leaves. (Lobato et al., 2008; Osuagwu et al., 2010). Thickening of Cuticles and mesophyll in response to water stress was also reported in Stipa lagascae (Boughalleb et al., 2015). Acanthophyllum ejtehadii’s root growth system is one of the outstanding specific adaptation mechanisms of this plant in response to its habitats. The primary roots are relatively short and the first order lateral fibrous roots are fairly long and grow near the soil surface. These types of root, which are considered as type II, are also seen in Ferocactus wislizeni (Cannon, 1949). These types of roots allow plants to collect surface waters during seasonal precipitation i.e., before evaporation. Since the soil texture of Radkan is 327 Taiwania Vol. 62, No. 3 Fig. 6. The karyological characteristics in A. ejtehadii. A. Metphasic cell. B. Karyotype. clay-loam, it is susceptible to waterlogging, due to the weak infiltration, causing poor root aeration (Gill et al., 2004). Roots of A. ejtehadii can escape the waterlogging conditions through growing near the soil surface, a mechanism that was also seen in marsh plants kilogram nitrogen and phosphorus, is considered as a poor soil in terms of the nutrient content (Esu, 1998;Kparmwang et al., 2001). This soil is rich in potassium and calcium carbonate with 403 mg/kg and 38 percent, respectively (Rosen et al., 1998). The vegetation of Radkan region is relatively poor and it seems that the poor soil nutrient content in Radkan doesn’t allow plants to thrive in this region. It was also reported that A. microphyllum can grow in the same soil conditions (Shokri et al., 2003). Acanthophyllum ejtehadii is a chamaephyte cushion plant. Different life forms of plant can be attributed to the adaptation to the climatic conditions of its habitats (Asaadi, 2009). This vegetative form of plants is a possible indicator of unfavorable climatic and poor soil conditions (Barik and Misra, 1998; Malik et al., 2007). Acanthophyllum ejtehadii grows in all slightly steep slope fields of Radkan region. It is also reported that A. microphyllum grows in clay loam soils with poor nutrient content and in all inclined fields (Shokri et al., 2003). Field observation revealed that A. ejtehadii expands its distribution range only by sexual reproduction. Maleki et al. (2015a) reported that the germination percentage of A. ejtehadii’s small seeds were 100 by scarification treatment in Vitro. Wind-induced-dispersion of A. ejtehadii’s seeds may result in their scratching by surrounding bed rocks. This natural scarification facilitates the germination of seeds. This strategy helps A. ejtehadii to avoid intraspecific competition by inhibition of seed germination under maternal canopy (Maleki et al., 2015a). Regarding the shape and orientation of the bract and floral leaves, Mahmoudi et al. (2012) have considered 328 (Atwell et al., 1999). This mechanism isn’t routine for all the Acanthophyllum species. For instance, A. pachystegium has a taproot with a little peripheral growth. Radkan’s soil, with its 0.5 and 6.53 grams per the A. ejtehadii similar to A. diezianum, and in terms of hairs similar to A. pachystegium. Based on the results of the current study branching pattern of buds and characteristics of hairs would be suitable distinguishing characters to identify A. ejtehadii from A. diezianum. The shape and orientation of the bracts and floral leaves are also proper differential properties to identify A. ejtehadii from both A. lilacinum and A. pachystegium. There are extensive data on Caryophyllaceae anatomical characteristics. hair type (Davis, 1967), Calcium oxalate crystals in parenchyma (Schweingruber, 2007), and oval seeds with curved embryo (Schiman-Czeika, 1988; Ghahraman, 2004) are the notable features of Acanthophyllum sp. These specific characteristics are also observed in A. ejtehadii. Maleki et al. (2015 b) reported that this species has curved embryo, oval seeds, and pantopolyporate spherical pollen grains, which is a general characteristic in Acanthophyllum taxa (Mahmoudi-Shamsabad et al., 2013). Acanthophyllum ejtehadii is a diploid species and has homogenous karyotype with 12m+3sm formula. Ghaffari (2004) studied karyological features of 17 Acanthophyllum species, and reported that this taxon has x=14-15 basic chromosome numbers. Most species of Oligosperma and Macrostegia sections are diploid with 2n=2x=30. However, Acanthophyllum and Plesiosperma sections are tetraploid (2n=4x=60) and hexaploid (2n=6x=90), respectively. It was also reported that polyploidy played an important role in speciation and evolution of this genus. September 2017 Maleki sadabadi et al.: The characteristics of Acanthophyllum ejtehadii ACKNOWLEDGEMENTS The authors express their appreciation to the Research Council of Ferdowsi University of Mashhad for the financial support of the research project (Project No. 3/30192). Moreover, the authors gratefully acknowledge Mr. M. Hassanzadeh for his constructive comments during the language edition of the paper. The authors would like to thank Dr. A. Pirani for her comments during the revision of this manuscript. LITERATURE CITED Asaadi, A. M. 2009. Floristic study of Firuzeh watershed (North Khorasan province). Res. J. Bio. Sci. 4: 1092-1103. Atwell, B. J., P.E. Kriedemann and C.G.N. Turnbull. 1999. Plants in Action: Adaptation Nature, Performance in Cultivation. MacMillan Education Australia, Melbourne. Barik, K.L. and B.N. Misra. 1998. Biological spectrum of a grassland ecosystem of South Orissa. Ecoprint 5: 73-7. Boughalleb, F., R. Abdellaoui, Z. Haddad and M. Neffati. 2015. Anatomical adaptations of the desert species Stipa lagascae against drought stress. Biologia (Bratislava) 70(8): 1042-1052. Cannon, W. A. 1949. A tentative classification of root systems. Ecology 30(4): 542-548. Davis, P.H. 1967. Flora of Turkey. Edinburg University Press, Edinburgh. De Micco, V., and G. Aronne. 2012. Morpho-Anatomical Traits for Plant Adaptation to Drought. In: R. Aroca (Ed.), Plant Responses to Drought Stress. Springer-Verlag Berlin, Heidelberg. pp. 37-61. Oliver, J.E. 2005. The Encyclopedia of World Climatology. Springer Dordrecht, Berlin, Heidelberg, New York. Esu, I.E. 1991. Detailed soil survey of NIHORT farm at Bunkure, Kano State, Nigeria. Institute of Agricultural Research, Zaria. Gerlach, D. 1977. Botanische Mikrotechnik. Thieme Verlag, Stuttgart. Ghaffari, S.M. 2004. Cytotaxonomy of some species of Acanthophyllum (Caryophyllaceae) from Iran. Biologia (Bratislava) 59: 53-60. Ghahreman, A. 2005. Plant Systematics Cormophytes of Iran: volume 1, University Publication Center, Tehran. Ghahreman, A. and F. Attar. 1999. Biodiversity of Plant Species in Iran: volume 1. Central Herbarium of Tehran University, Faculty of Science, Tehran. Gill, J. S., J. Tisdall, I.G.M. Kusnarta and B.M. McKenzie. 2004. Physical properties of a clay loam soil mixed with sand. Third Australian New Zealand Soils Conference. Heywood, V.H. 1985. Flowering plants of the World. Croop Helm. London. Huziwara, Y. 1962. Karyotype analysis in some genera of Compositae. VIII. Further studies on the shromosome of Aster. Am. J. Bot. 49(2): 116-119. Jalili, A. and Z. Jamzad. 1999. Red Data Book of Iran. Research Institute of Forests and Rangelands, Tehran. Kooyers, N.J. 2015. The evolution of drought escape and avoidance in natural herbaceous populations. Plant Sci. 234:155-162. Köppler, M.R. and J.D. Hitchmough. 2015. Ecology good, autecology better; Improving the sustainability of designed plantings. J. Landsc. Archit. 10: 82-91. Kparmwang, T., B.A. Raji, A.C. Odunze and V.O. Chude. 2001. Properties, classification and agricultural potential of Ustults, and Tropepts on a sedimentary toposequence in the Benue. Nigerian J. Soil Res. 2: 58-65. Lobato, A.K.S., C.F. Oliveira Neto, B.G. Santos Filho, R.C.L. Costa, F.J.R. Cruz, H.K.B. Neves and M.J.S. Lopes. 2008. Physiological and biochemical behavior in soybean (Glycine max cv. Sambaiba) plants under water deficit. Aust. J. Crop. Sci. 2: 25-32. Mahmoudi-Shamsabad, M., J. Vaezi, F. Memariani and M.R. Joharchi. 2012. A new species and a new record of Acanthophyllum C.A.Mey. (Caryophyllaceae) from northeast of Iran. Iranian J. Bot. 18: 59-63. Maleki Sadabadi, Z., H. Ejtehadi, P. Abrishamchi and J. Vaezi. 2015a. Compare the features of pollen species Acanthophyllum ejtehadii Mahmoudi and Vaezi with its relative species. Second National Conference on Environmental, Natural Resources Engineering and Sustainable agricultural management. Maleki Sadabadi, Z., H. Ejtehadi, P. Abrishamchi and J. Vaezi. 2015b. Evaluation of germination and micro-morphological characters of Acanthophyllum ejtehadii Mohmoudi and Vaezi seed. Second National Conference on Environmental, Natural Resources Engineering and Sustainable agricultural management. Malik, Z.H., F. Hussain, and N.Z. Malik. 2007. Life form and Leaf Size Spectra of Plant Communities Harbouring Ganga Chotti and Bedori Hills During 1999-2000. Int J Agric Biol. 6: 833-838. Mitra, J. 2001. Genetics and genetic improvement of drought resistance in crop plants. Cur. Sci. 80:758 – 763. Moussavi-Harami, R., A. Mahbobi, M. Khanehbad, M. Akhlaghi, M. Khajeh-Yazdi, and Gh. Rostamizadeh. 2004. The relationship between sedimentology and texture parameters of the downstream side of Radkan River Basin, Northwest Chenaran. Proceeding of Seventh conference of Geological Society of Iran (article in Persian). Müller, M., D. Grill, B. Druskovic and J. Pradiz. 1991. A cytogenetic method for examining the vitality of spruces. Phyton 31: 143-155. NASA POWER. 2012. NASA Climatology Resource for Agroclimatology, [homepage on the Internet], NASA POWER Agroclimatology, (cited 10 August 2015), http://power.larc.nasa.gov/cgi-bin/cgiwrap/solar/agro.cgi. Osuagwu, G.G.E., H.O. Edeoga and A.N. Osuagwu. 2010. The influence of water stress (drought) on the mineral and vitamin potential of the leaves Ocimum gratissimum L. Recent Res. in Sci. Technol. 2(2): 27-33. Pianka, E.R. 2008. Autecology. In: (Jørgensen, S. E. and Fath B. D. Eds.). Encyclopedia of Ecology volume 1 Elsevier, Oxford. pp. 285-287. Pirani, A., M.R. Joharchi and F. Memariani. 2013. A new species of Acanthophyllum (Caryophyllaceae) from Iran. Phytotaxa 92(1): 20-24. Pirani, A., Sh. Zarre, B.E. Pfeil, Y.J.K. Bertrand, M. Assadi and B. Oxelman. 2014. Molecular phylogeny of Acanthophyllum (Caryophyllaceae: Caryophylleae), with 329 Taiwania emphasis on infrageneric classification. Taxon 63(3): 592-607. Raunkiaer, C. 1934. The Life Forms of Plant and Statistical Plant Geography. Claredon Press. Oxford. Rosen, C. J., P.M. Bierman, and D.R. Eliason. 2008. Soil Test Interpretation and Fertilizer Management for Lawns, Tturf, Gardens, and Landscape Plant [homepage on the Internet], University of Minnesota, Extension Service [cited 20 March 2015]. http://www.extension.umn.edu/garden/yard-garden/soils/so il-test-interpretations-and-fertilizer-management/. Schiman-Czeika, H. 1988. Flora Iranica, volume 163. Akademische Druck, Graz, Austria. Schweingruber, F.H. 2007. Stem anatomy of Caryophyllaceae, Flora 202(4): 281-292. 330 Vol. 62, No. 3 Shokri, M., M.A. Bahmanyar and M.R. Tatian. 2003. An Ecological Investigation of Vegetation Cover in Estival Rangeland of Hezarjarib (Behshahr). Iranian J. Nat.l Resour. 56: 131-142. Soil Survey Staff. 2014. Keys to Soil Taxonomy 12th edition. United States Department of Agriculture. Taiz, L. and E. Zeiger. 2010. Plant Physiology. Sinauer Associates, Sunderland. Takhtajan, A.L. 1986. Floristic Regions of the World. University of California Press, Berkley. Villeneuve, G.O. 1980. Glossaire de météorologie et de climatologie. Les Presses de I'Université Laval, Quebec. Zohary, M. 1973. Geobotanical foundations of the Middle East, CRC Press, Stuttgart.