Proceedings: Shrubland Ecotones; 1998 August 12–14; Ephraim, UT
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Population Dynamics of a Perennial Halophyte Allenrolfea occidentalis Bilquees Gul Darrell J. Weber M. Ajmal Khan Abstract—Demography and population biology of a perennial A. occidentalis plants have to allow salts to enter through halophyte, Allenrolfea occidentalis, was studied during the growing root membranes and the physiology of the plant changes seasons (May to November) of 1996 and 1997. Soil samples were from the wet to dry years due to the increase in soil salinity. collected monthly from the playa throughout the growing seasons Flanagan and Jefferies (1988) reported that as salinity for two years and analysed to determine the conductivity, pH, soil increased, photosynthesis in Plantago maritima declined –2 –1 moistures and ions. During the study period, the population was 17 to 14 μmol m .S while leaf conductance dropped mark- exposed to wide variations in soil salinity from low to high and soil edly. Transpiration and photosynthesis involve gas exchange moisture ranging from very wet to drought levels. Seasonal changes between the plant and atmosphere through the stomata and in dry weight was directly related to soil salinity stress. When are well known to decrease with water stress (Fisher 1976) salinity levels become low, the dry matter production increases. The or salinity (Khan and others 1976). population of A. occidentalis suffered heavy mortality in 1996 due to Halophytic species commonly exhibit quite high concen- the high salinity and temperature stress. Plant growth reached its trations of several salt ions in various plant organs or the maximum in July and succulence decreased with aging of plants. entire plant. Although the salt accumulating nature of Results indicate that community vegetation respond differently to halophytes has been recognized for many years, now it has the environmental changes in 1996 and in 1997. Salinity, tempera- been proven that sodium is essential to the growth of most of ture and precipitation have a major effect on the survival and the chenopodiaceae (Brownwell and Wood 1957; Brownwell growth of A. occidentalis under field conditions. 1965; Moore and Caldwell 1972; Naidoo and Rughunanan 1990; Khan and others 1998). Terrestrial halophytes utilize the controlled accumulation and sequestration of inorganic ions, chiefly Na+ and K+ balanced by Cl–, as the basic The distribution of plant species in saline environments of mechanism by which they adjust the osmotic potential of inland western United States is closely associated with soil their internal tissue to the external salinity (Flowers and water potentials and other factors influencing the level of Yeo 1986; Cheeseman 1988). salinity stress, including microtopography, precipitation, In this study, the environmental conditions as they affect and depth of water table (Young and others 1995). There are growth of a natural population of Allenrolfea occidentalis certain areas in western Utah, where the salt content is high during the growing seasons in an inland salt playa of the enough to form thick salt crusts on the top of the soil making Great Basin desert, are reported. The objective of this study plant growth of any kind impossible, on such areas some of was to study the effect of variation in soil salinity and the more salt tolerant plants survive with little or no compe- temperature on the growth and ecophysiology of A. occiden- tition and Allenrolfea occidentalis (Wats.) Kuntze is one of talis under natural conditions. them (Quigley 1956). Allenrolfea occidentalis is a perennial chenopod, which colonizes extremely saline habitats of the temperate desert basins. Plant communities dominated by Materials and Methods ___________ A. occidentalis offer the extreme in adaptations to survival and growth under moisture stress (Young and others 1995). Study site During the growing season A. occidentalis plants are sub- jected to a great variation in edaphic conditions (Trent and The site chosen for this study is a salt playa east of others 1997). Seasonal variation in soil salinity in saline Goshen, northwestern Utah. It is an area of flat, low-lying habitats is well documented and is directly influenced by the ground in the bottom of a fairly wide valley that spreads out fluctuations in soil moisture levels (Ungar 1973, 1978a; at the southern end into a vast stretch of flat, salt incrusted Waisel 1972). Young and others (1995) showed that the plain. The area contains numerous salt marshes and salt playas with nearly pure stands of Allenrolfea occidentalis. The point centered quarter method (Cottam and Curtis 1956) was used to sample the vegetation over 20 random In: McArthur, E. Durant; Ostler, W. Kent; Wambolt, Carl L., comps. 1999. points and relative frequencies were calculated for each Proceedings: shrubland ecotones; 1998 August 12–14; Ephraim, UT. Proc. species in the community. During the spring of 1996, three RMRS-P-11. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. transects were established in the salt playa. These transects Bilquees Gul is a Plant Ecologist, Department of Botany, University of ran through the community and were approximately paral- Karachi, Karachi-75270, Pakistan. Darrell J. Weber is a Professor of Plant lel to the marsh. Fifteen permanent quadrats (100 100 m2) Pathology, Department of Botany and Range Science, Brigham Young Uni- x versity, Provo, UT 84602. M. Ajmal Khan is a Professor of Plant Ecology, were established on three transects, three replicates on each Department of Botany, University of Karachi, Karachi-75270, Pakistan. 124 USDA Forest Service Proceedings RMRS-P-11. 1999 site. The number of plants surviving in each plot and their cover were counted monthly throughout the growing season from May to November 1996 and 1997. In order to reduce disturbance in quadrats, ten equal sized plants were ran- domly collected from the area in the vicinity of each quadrat every month of the growing season for two years. Plants were separated into leaves, stem and root. Allenrolfea occidenta- lis does not have true leaves but they form jointed, seemingly leafless stems. The green leafy succulent portion is consid- ered here as leaf while woody portion as stem. Fresh and dry weight of the plants was recorded before and after drying the material in an oven at 80°C for 48 h. For ion measurements 0.5 gram of plant material was boiled in 25 ml of water for two hours at 100°C using a dry heat bath. This hot water extract was cooled and filtered using Whatman no. 2 filter paper. One ml of hot water extract was diluted with distilled water for ion analysis. Chloride, nitrate and sulphate ions contents were measured with a DX-100 ion chromatograph. Cation contents Na+, K+, Ca2+ and Mg2+ of the plant organs were analysed using a Perkin Elmer model 360 atomic absorption spectrophotom- eter. The net photosynthesis rate of four replicates per quadrat were taken with an LI-6200 portable photosynthe- sis system (LI-COR, Inc., Lincoln, NE). The level of stress in plants growing in field conditions were measured with CF- 1000 chlorophyll fluorescence measurement system. The water potential was measured using a Plant Moisture Stress Instrument (PMS Instrument Co.). Ten surface soil (to a depth of 15 cm) and 10 subsurface (to a depth of 30 cm) soil samples were collected monthly from the A. occidentalis community during the growing season. Soil moisture was measured by weighing 12 g of samples, oven drying them at 136°C for 24 h and reweighing them to determine the water loss. Percent soil moisture was calcu- lated as percentage weight of water in dry soil. Then for the Figure 1—Seasonal pattern of dry weight (g plant–1) determination of organic contents of the soil these samples of leaf, stem and root of plants collected from an Allenrolfea occidentalis community. Bar represents were dried at 360°C for 24 h. mean ± S. E. Five grams of soil mixed with 25 ml of distilled water were shaken and filtered using Whatman no. 1 filter paper. PH (pH meter) and soil conductivity (model-10 portable conduc- tivity meter) were measured. The results of growth, ion contents, net photosynthesis, water potential and stress A three way ANOVA showed a significant individual effect were analysed using three way ANOVA. A Bonferroni test of plant parts, years and months and their interactions in was carried out to determine if significant (P <0.05) differ- affecting dry weight of A. occidentalis plants. Plants col- ences occurred between individual treatments (SPSS 1996). lected from the field were analysed for leaves, stem and root fresh and dry weights. Allenrolfea occidentalis plants col- lected on August 1996 had greater dry weights than plants Results ________________________ collected in other months (fig. 1). The fresh and dry weights of the plants increased gradually from May to September A three way ANOVA showed a significant individual and then decreased. effect of quadrats F = 28.71, (P <0.0001) and years F = 7.02, A three way ANOVA showed a significant individual effect (P <0.001), while months were not significant in affecting of months (P <0.0001) and years (P <0.0001), while soil basal area. Interactions between quadrats, years and months layers were not significant in affecting pH (table 2). The were also significant (P <0.0001). Phytosociological survey surface soil pH was generally higher and fluctuated more showed that the salt playa community has an almost pure than the corresponding subsurface pH. The subsurface pH population of A. occidentalis a few individuals of Salicornia of the soil changed very little during the growing season rubra, Salicornia utahensis and Distichlis spicata were (table 1 and 2). The decrease in pH of the surface soil present. Quantitative data indicated that A. occidentalis appeared to be correlated to the amount of rainfall. A had consistently high cover in the intermediate plots through- decrease in the pH of the surface soil occurred in May to July out the growing season of 1996 (fig.