Soil moisture redistribution as a mechanism of facilitation in savanna tree–shrub clusters

Plant–soil water relations were examined in the context of a selective removal study conducted in tree–shrub communities occupying different but contiguous soil types (small discrete clusters on shallow, duplex soils versus larger, extensive groves on deep, sandy soils) in a subtropical savanna parkland. We (1) tested for the occurrence of soil moisture redistribution by hydraulic lift (HL), (2) determined the influence of edaphic factors on HL, and (3) evaluated the significance of HL for overstory tree–understory shrub interactions. Diel cycling and nocturnal increases in soil water potential (Ψ_soil), characteristic signatures of HL, occurred intermittently throughout an annual growth cycle in both communities over a range of moisture levels (Ψ_soil=−0.5 to −6.0 MPa) but only when soils were distinctly stratified with depth (dry surface/wet deep soil layers). The magnitude of mean (±SE) diel fluctuations in Ψ_soil (0.19±0.01 MPa) did not differ on the two community types, though HL occurred more frequently in groves (deep soils) than clusters (shallow soils). Selective removal of either Prosopis glandulosa overstory or mixed-species shrub understory reduced the frequency of HL, indicating that Prosopis and at least one other woody species was conducting HL. For Zanthoxylum fagara, a shallow-rooted understory shrub, Prosopis removal from clusters decreased leaf water potential (Ψ_leaf) and net CO₂ exchange (A) during periods of HL. In contrast, overstory removal had neutral to positive effects on more deeply-rooted shrub species (Berberis trifoliolata and Condalia hookeri). Removal of the shrub understory in groves increased A in the overstory Prosopis. Results indicate the following: (a) HL is common but temporally dynamic in these savanna tree–shrub communities; (b) edaphic factors influencing the degree of overstory/understory development, rooting patterns and soil moisture distribution influence HL; (c) net interactions between overstory and understory elements in these woody patches can be positive, negative and neutral over an annual cycle, and (d) Prosopis-mediated HL is an important mechanism of faciliation for some, but not all, understory shrubs.     

Indications of Hydraulic Lift by Pinus halepensis and Its Effects on the Water Relations of Neighbour Shrubs

We measured the stable deuterium isotopic composition of xylem sap, the shoot predawn and midday water potentials, and the leaf δ¹³C of Mediterranean shrubs Pistacia lentiscus, Globularia alypum and Rosmarinus officinalis in a south-oriented transect from a large (12 m tall) Aleppo pine tree, Pinus halepensis. We aimed to study the possibility of hydraulic lift from the deep roots of this pine tree to the shallow soil layers and its influence on these neighbour shrubs. These same traits were also studied in several individuals of the shrub Pistacia lentiscus growing with different types of neighbours: just shrubs, a small (3 – 4 m) pine tree, and the above mentioned large pine tree. The greater the distance from P. halepensis the plants grew, the higher xylem water δD, the lower the soil water content, and, the lower the predawn and midday water potentials were found. These results suggest the existence of an hydraulic lift from deep roots to shallow soil in this big tree. Further indication of this existence is provided by the improved water status of P. lentiscus (higher water potentials and δD, and lower δ¹³C and, therefore, lower water use efficiencies) when growing close to the big pine in comparison with the same shrub species growing close to small pines or just surrounded by other shrubs. Moreover, all these trends occurred in the dry summer season, but disappeared in the wet spring season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Spatial patterns of desert annuals in relation to shrub effects on soil moisture

Questions: What are the effects of a shrub (Haloxylon ammodendron) on spatial patterns of soil moisture in different seasons? How does productivity of understorey annuals respond to these effects? Are such effects always positive for annuals under shrubs? Location: South Gurbantunggut Desert, northwest China. Methods: Using geostatistics, we explored seasonal patterns of topsoil moisture in a 12 × 9‐m plot over the growing season. To determine spatial patterns of understorey annuals in response to H. ammodendron presence, biomass of annuals was recorded in four 0.2 × 5.0‐m transects from the centre of a shrub to the space between shrubs (interspace). We also investigated vertical distribution of root biomass for annuals and soil moisture dynamics across soil profiles in shrub‐canopied areas and interspaces. Results: Topsoil moisture changed from autocorrelation in the wet spring to random structure in the dry season, while soil moisture below 20 cm was higher in shrub‐canopied areas. Across all microhabitats, soil moisture in upper soil layers was higher than in deeper soil layers during the spring wet season, but lower during summer drought. Topsoil was close to air‐dry during the dry season and developed a ‘dry sand layer’ that reduced evaporative loss of soil water from deeper layers recharged by snowmelt in spring. Aboveground biomass of understorey annuals was lowest adjacent to shrub stems and peaked at the shrub margin, forming a ‘ring’ of high herbaceous productivity surrounding individual shrubs. To acclimate to drier conditions, annuals in interspaces invested more root biomass in deeper soil with a root/shoot ratio (R/S) twice that in canopied areas. Conclusions: Positive and negative effects of shrubs on understorey plants in arid ecosystems are commonly related to nature of the environmental stress and tested species. Our results suggest there is also microhabitat‐dependence in the Gurbantunggut Desert. Soil water under H. ammodendron is seasonally enriched in topsoil and deeper layers. Understorey annuals respond to the effect of shrubs on soil water availability with lower R/S and less root biomass in deeper soil layers and develop a ‘ring’ of high productivity at the shrub patch margin where positive and negative effects of shrubs are balanced.     

Differential water resource use by herbaceous and woody plant life-forms in a shortgrass steppe community

We conducted a study to test the predictions of Walter's two-layer model in the shortgrass steppe of northeastern Colorado. The model suggests that grasses and woody plants use water resources from different layers of the soil profile. Four plant removal treatments were applied in the spring of 1996 within a plant community codominated by Atriplex canescens (a C₄ shrub) and Bouteloua gracilis (a C₄ grass). During the subsequent growing season, soil water content was monitored to a depth of 180 cm. In addition, stem and leaf tissue of Atriplex, Bouteloua and the streamside tree Populus sargentii were collected monthly during the growing seasons of 1995 and 1996 for analysis of the δ¹⁸O value of plant stem water (for comparison with potential water sources) and the δ¹³C value of leaves (as an indicator of plant water status). Selective removal of shrubs did not significantly increase water storage at any depth in the measured soil profile. Selective removal of the herbaceous understory (mainly grasses) increased water storage in the top 60 cm of the soil. Some of this water gradually percolated to lower layers, where it was utilized by the shrubs. Based on stem water δ¹⁸O values, grasses were exclusively using spring and summer rain extracted from the uppermost soil layers. In contrast, trees were exclusively using groundwater, and the consistent δ¹³C values of tree leaves over the course of the summer indicated no seasonal changes in gas exchange and therefore minimal water stress in this life-form. Based on anecdotal rooting-depth information and initial measurements of stem water δ¹⁸O, shrubs may have also had access to groundwater. However, their overall δ¹⁸O values indicated that they mainly used water from spring and summer precipitation events, extracted from subsurface soil layers. These findings indicate that the diversity of life-forms found in this shortgrass steppe community may be a function of the spatial partitioning of soil water resources, and their differential use by grasses, shrubs, and trees. Consequently, our findings support the two-layer model in a broad sense, but indicate a relatively flexible strategy of water acquisition by shrubs. Received: 23 December 1997 / Accepted: 16 September 1998     

Species-specific patterns of hydraulic lift in co-occurring adult trees and grasses in a sandhill community

Plants can significantly affect ecosystem water balance by hydraulic redistribution (HR) from dry to wet soil layers via roots (also called hydraulic lift, HL, when the redistribution is from deep to shallow soil). However, the information on how co-occurring species in natural habitats differ in HL ability is insufficient. In a field study, we compared HL ability of four tree species (including three congeneric oak species) and two C4 bunch grass species that co-occur in subxeric habitats of fall-line sandhills in southeastern USA. Soil water potentials (_s) were recorded hourly for 3 years both in large chambers that isolated roots for each species and outside the chambers. Outside of root chambers, soil drying occurred periodically in the top 25 cm and corresponded with lack of precipitation during the summer growing season. Soil moisture was continuously available at a 1 m depth. HL activity was observed in three of the tree species, with greater frequency for Pinus palustris than for Quercus laevis and Q. incana. The fourth tree species Q. margaretta did not exhibit HL activity even though it experienced a similar _s gradient. For the C4 bunch grasses, Aristida stricta exhibited a small amount of HL activity, but Schizachyrium scoparium did not. The capacity for HL activity may be linked to the species ecological distribution. The four species that exhibited HL activity in this subxeric habitat are also dominant in adjacent xeric sandhill habitats, whereas the species that did not exhibit HL are scarcely found in the xeric areas. This is consistent with other studies that found greater fine root survival in dry soil for the four xeric species exhibiting HL activity. The differential ability of these species to redistribute water from the deep soil to the rapidly drying shallow soil likely has a strong effect on the water balance of sandhill plant communities, and is likely linked to their differential distribution across edaphic gradients.An erratum to this article can be found at     

天山林区4种主要灌木夏季水分来源差异

天山林区群落结构相对简单、木本植物种类较少,但天山林区灌木群落中主要木本植物间的水分竞争模式尚不明确,水分利用动态缺乏定量分析。运用稳定同位素技术,对天山林区灌木群落4种主要灌木的茎杆水分及各潜在水源的氢氧稳定同位素组成进行测定,运用IsoSource模型定量分析4种灌木在夏季对各潜在水源的相对利用比例,探讨天山林区灌木群落主要灌木树种水分来源差异及动态变化。结果发现:7月,当浅层土壤含水量充足时,密刺蔷薇、黑果栒子和金丝桃叶绣线菊均大幅度吸收利用浅层土壤水,相对利用比例高于89.3%,异果小檗则相反,即吸收利用各潜在水源(浅层土壤水30.7%、中层土壤水29.4%、深层土壤水25.7%、溪水14.2%,下同);8月,当浅层土壤含水量降低时,密刺蔷薇转移至60-100 cm深层土壤水和溪水,相对利用比例分别为64.8%和27%,黑果栒子和金丝桃叶绣线菊以相似比例吸收利用各潜在水源(33.8%和36.8%、30.9%和29.7%、23.5%和22.3%、11.8%和11.2%),异果小檗则表现出可能吸收利用80-100 cm以下更稳定的深层土壤水;9月,当浅层土壤含水量升高时,4种灌木均大量吸收利用浅层土壤水,相对利用比例高于72.2%。这表明,天山林区灌木群落主要树种可通过可塑性转换水分来源来应对环境水分变异,在时间和空间上有效分割灌丛水源从而减缓对水分资源的竞争压力,从而通过在水分资源利用上的生态位分化促进物种间的共存。     

Deuterium labelling of roots provides evidence of deep water access and hydraulic lift by Pinus nigra in a Mediterranean forest of NE Spain

We studied the tree access to deep water sources and the possibility of hydraulic lift from the deep roots of a Pinus nigra tree to the shallow soil layers in a Mediterranean forest of NE Spain. We also studied the use of hydraulically lifted water by neighboring trees, shrubs, and sprouts. We enriched the roots of a large P. nigra (10 m tall) with deuterium by accessing them from a below ground cave. During the next 3 days we measured stable deuterium isotopic composition of xylem sap, shoot predawn and midday water potentials, and the leaf δ¹³C and δ¹⁵N of the P. nigra tree, neighboring Quercus ilex ballota trees and sprouts, and Juniperus oxycedrus shrubs. The study was conducted both in dry summer and in wet spring. In summer, deuterated water absorbed by deep roots of P. nigra appeared in the stem water of neighboring plants and in surface soil. The most δD-enriched plant xylem sap was found in the enriched P. nigra tree, followed by the Q. ilex sprouts, the small Q. ilex trees and the surface soil (15 cm). All these trends disappeared in the wet spring season, when HDO only slightly appeared in the surface soil. The results show that the studied P. nigra tree accesses deep water source and conducts hydraulic lift in this Mediterranean forest in dry summer but not necessarily in wet spring.     

Hydraulic lift among native plant species in the Mojave Desert

Hydraulic lift was investigated among native plants in the Mojave Desert using in situ thermocouple psychrometers. Night lighting and day shading experiments were used to verify the phenomenon. Hydraulic lift was detected for all species examined: five shrub species with different rooting depths and leaf phenologies and one perennial grass species. This study was the first to document hydraulic lift for a CAM species, Yucca schidigera. The pattern of diel flux in soil water potential for the CAM species was temporally opposite to that of C₃ species: for the CAM plant, soil water potential increased in shallow soils during the day when the plant was not transpiring and decreased at night when transpiration began. Because CAM plants transport water to shallow soils during the day when surrounding C₃ and C₄ plants transpire, CAM species that hydraulically lift water may influence water relations of surrounding species to a greater extent than hydraulically lifting C₃ or C₄ species. A strong, negative relationship between the percent sand in the study site soils at the 0.35 m soil depth and the frequency that hydraulic lift was observed at that depth suggests that the occurrence of hydraulic lift is negatively influenced by coarse-textured soils, perhaps due to less root–soil contact in sandy soils relative to finer-textured soils. Differences in soil texture among study sites may explain, in part, differences in the frequency that hydraulic lift was detected among these species. Further investigations are needed to elucidate species versus soil texture effects on hydraulic lift. This revised version was published online in June 2006 with corrections to the Cover Date.     

Water use patterns of four co-occurring chaparral shrubs

Summary Mixed stands of chaparral in California usually contain several species of shrubs growing close to each other so that aerial branches and subterranean roots overlap. There is some evidence that roots are stratified relative to depth. It may be that root stratification promotes sharing of soil moisture resources. We examined this possibility by comparing seasonal water use patterns in a mixed stand of chaparral dominated by four species of shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a neutron probe and soil phychrometers to follow seasonal depletion and recharging of soil moisture and compared these patterns to seasonal patterns of predawn water potentials, diurnal leaf conductances, and diurnal leaf water potentials. Our results indicated that 1) Quercus was deeply rooted, having high water potentials and high leaf conductances throughout the summer drought period, 2) Heteromeles/Adenostoma were intermediate in rooting depth, water potentials, and leaf conductances, and 3) Rhamnus was shallow rooted, having the lowest water potentials and leaf conductances. During the peak of the drought, predawn water potentials for Quercus corresponded to soil water potentials at or below a depth of 2 m, predawn water potentials of Heteromeles/ Adenostoma corresponded to a depth of 0.75 m, and predawn water potentials of Rhamnus corresponded to a depth of 0.5 m. This study supports the concept that co-occurring shrubs of chaparral in California utilize a different base of soil moisture resources.     

Spatial distribution of soil nutrients and ephemeral plants underneath and outside the canopy of Porlieria chilensis shrubs (Zygophyllaceae) in arid coastal Chile

Soil nutrients and density and biomass of annual plants underneath and outside the canopy of Porlieria chilensis shrubs were measured at the end of the growing season in a protected arid coastal site in Chile. Levels of soil nitrogen, phosphorus and organic matter were significantly higher underneath than outside the canopies of shrubs. Almost 4 times as many plants occurred outside than underneath shrubs, but no significant differences in total aboveground biomass were found. Several species had higher densities and/or biomass outside rather than underneath shrubs, whereas others showed the oppsite trend. Species richness was lower underneath P. chilensis canopy. The spatial microdistribution of ephemeral species may be explained by differential water and nutrient requirements. Comparison of the patterns observed in our protected site versus surrounding unprotected areas supports the generalization that man, by removing shrubs and trees, has changed a previous heterogeneous spatial distribution of nutrients to a more homogenous one.This is a contribution of the Program of Arid Zone Studies of the Universidad de la Serena     

THE ANNUAL RHYTHM OF CAMBIAL ACTIVITY IN TWO WOODY SPECIES OF THE CHILEAN “MATORRAL”

The annual rhythm of cambial activity is compared in Proustia cuneifolia and Acacia caven, two typical shrubs of the “matorral” in the semiarid region of central Chile. Proustia, a drought deciduous shrub, shows a typical desert cambial rhythm, highly sensitive to rainfall. The growth activity of this species is limited to the periods of adequate moisture. Acacia is an evergreen whose cambial activity is observed almost throughout the year; it is not synchronous with rainfall. Adaptation in this species seems to consist in developing long roots able to tap underground water. These results indicate that both shrubs, although growing together, have different adaptive strategies to the same xerophytic conditions.     

Soil water availability and rooting depth as determinants of hydraulic architecture of Patagonian woody species

Adaptations of species to capture limiting resources is central for understanding structure and function of ecosystems. We studied the water economy of nine woody species differing in rooting depth in a Patagonian shrub steppe from southern Argentina to understand how soil water availability and rooting depth determine their hydraulic architecture. Soil water content and potentials, leaf water potentials (Ψ_Leaf), hydraulic conductivity, wood density (ρ_w), rooting depth, and specific leaf area (SLA) were measured during two summers. Water potentials in the upper soil layers during a summer drought ranged from −2.3 to −3.6 MPa, increasing to −0.05 MPa below 150 cm. Predawn Ψ_Leaf was used as a surrogate of weighted mean soil water potential because no statistical differences in Ψ_Leaf were observed between exposed and covered leaves. Species-specific differences in predawn Ψ_Leaf were consistent with rooting depths. Predawn Ψ_Leaf ranged from −4.0 MPa for shallow rooted shrubs to −1.0 MPa for deep-rooted shrubs, suggesting that the roots of the latter have access to abundant moisture, whereas shallow-rooted shrubs are adapted to use water deposited mainly by small rainfall events. Wood density was a good predictor of hydraulic conductivity and SLA. Overall, we found that shallow rooted species had efficient water transport in terms of high specific and leaf specific hydraulic conductivity, low ρ_w, high SLA and a low minimum Ψ_Leaf that exhibited strong seasonal changes, whereas deeply rooted shrubs maintained similar minimum Ψ_Leaf throughout the year, had stems with high ρ_w and low hydraulic conductivity and leaves with low SLA. These two hydraulic syndromes were the extremes of a continuum with several species occupying different portions of a gradient in hydraulic characteristics. It appears that the marginal cost of having an extensive root system (e.g., high ρ_w and root hydraulic resistance) contributes to low growth rates of the deeply rooted species.     

Soil water partitioning contributes to species coexistence in tallgrass prairie

The majority of tallgrass prairie root biomass is located in the upper soil layers (0–25 cm), but species differences exist in reliance on soil water at varying depths. These differences have led to the hypothesis that resource partitioning belowground facilitates species co‐existence in this mesic grassland. To determine if plant water relations can be linked to soil water partitioning as a potential mechanism allowing C₃ species to persist among the more dominant C₄ grasses, we measured differences in the source of water‐use using the isotopic signature of xylem water, volumetric soil water content at 4 depths, and leaf water potentials. Data were collected for seven species representing C₄ grasses, C₃ forbs and C₃ shrubs over three growing seasons at the Konza Prairie (Kansas, USA) to encompass a range of natural climatic conditions. C₄ grasses relied on shallow soil water (5 cm) across the growing season and had midday leaf water potentials that were highly correlated with shallow soil water regardless of soil water availability at other portions of the soil profile (20, 40 and 90 cm). In contrast, C₃ species only used shallow soil water when plentiful at this depth; these species increased their dependence on soil water from greater depths as the upper soil layers dried. Structural equation models describing plant water relations were very similar for the three C₄ species, whereas a unique set of models and drivers were identified for each of the C₃ species. These results support soil water partitioning as a mechanism for species coexistence, as C₄ species in this grassland have relatively consistent dependence on water in shallow soil layers, whereas C₃ species show niche differentiation in water use strategies to avoid competition with C₄ grasses for water in shallow soil layers when this resource is limiting and leaf water stress is high.     

Size-dependent foraging efficiency,cannibalism and zooplankton community structure

Only recently has the importance of positive interactions among plant species in structuring natural communities been supported by experimental evidence. Most studies have focused on interactions between a pair of species at a single life-history stage. In this study positive interactions between a woody nitrogen-fixing shrub (Myrica pensylvanica) and two herbaceous sand dune species (Solidago sempervirens, Ammophila breviligulata) which frequently grow beneath shrub canopies are examined throughout the life cycles of the herbaceous species. Comparisons of S. sempervirens and A. breviligulata growing beneath and outside M. pensylvanica shrubs show that plants growing in association with shrubs are larger, are more likely to flower, produce greater numbers of flowers and seeds, have higher midday xylem water potentials, have higher tissue nitrogen concentrations, and have higher photosynthetic efficiencies. Measurements of environmental conditions show that areas beneath shrubs are more shaded, have lower soil temperatures, and have higher soil nitrogen levels. The results from experimental manipulations designed to test the effects of Myrica shrubs on understory species suggest that the observed differences in plant performance are strongly influenced by canopy shading and soil nutrient enrichment associated with the shrubs. The results demonstrate that M. pensylvanica facilitates growth, reproduction, and recruitment of S. sempervirens and A. breviligulata growing beneath it. This study, one of the few to examine positive interactions at different life-history stages, supports previous predictions that positive interactions may be particularly important in plant communities characterized by physiologically stressful conditions. Received: 21 July 1999 / Accepted: 18 January 2000     

Influence of an overstorey tree (Prosopis glandulosa) on associated shrubs in a savanna parkland: implications for patch dynamics

The arborescent legume, honey mesquite (Prosopis glandulosa), appears to play a central role in patch dynamics of southern Texas savannas by modifying soils and microclimate and by facilitating the ingress, establishment and/or growth of shrubs in its understorey. As an indirect test for the occurrence and persistence of facilitation in mature shrub clusters (patches), we examined the gas exchange, water relations and production of associated shrubs growing in patches where a Prosopis overstorey was present and in patches where Prosopis had succumbed to natural mortality. Surface (0–10 cm) soils associated with shrub patches were enriched in total [N] and [C] compared to soils of neighboring herbaceous zones. However, there were no detectable differences in soil [N] or [C] in patches with and without Prosopis. Foliar [N] and biomass of various shrub species were also statistically comparable for patches with and without Prosopis. These results are in accordance with other studies that indicate the nutrient legacy associated with Prosopis occupation of a patch may persist for decades after its demise. In comparison to plants growing in the absence of Prosopis, leaf water potentials (predawn and midday), and net photosynthesis and water vapor conductance (morning and midday) of outer-canopy sunlit leaves over an annual growth cycle were comparable for two common evergreen shrubs, Zanthoxylum fagara and Berberis trifoliolata, growing in patches with a live Prosopis. These findings indicate that the presence of Prosopis was not enhancing the growth or activity of mature understorey shrubs; facilitation may, therefore, be important only during early stages of cluster development. In addition, we found no indication that the loss of Prosopis has initiated a downward phase in a cyclic succession of patch initiation, growth and death. Rather, the understorey shrubs appear to be able to maintain growth and productivity in the absence of a Prosopis overstorey, and may, therefore, represent persistent components of woody patches on these savanna landscapes.     

白刺沙包浅层土壤水分动态及其对不同降雨量的响应

以乌兰布和沙漠典型白刺沙包为研究对象,使用EC-5土壤水分传感器对其浅层(0—50 cm)土壤水分进行长期连续监测,分析了白刺沙包不同深度土层对不同降雨量的响应及整个生长季的土壤水分动态特征。结果表明:降雨是乌兰布和沙漠白刺沙包土壤水分的最重要补给源,降雨量大小是影响浅层土壤水分补给深度的决定因素。小于10 mm的降雨完全被表层(0—10 cm)土壤吸收,无法补给10 cm以下土壤水分;10—20 mm的降雨对土壤水分的补给深度达到20 cm;20—30 mm的降雨对土壤水分的补给深度达到40 cm,大于30 mm的降雨补给深度可达到50 cm,甚至更深土层。在研究区降雨量以小于20 mm降雨为主的情况下,20 cm以下土层土壤水分逐步恶化,久之将有利于浅根系草本植物的生长,不利于白刺的生长繁殖。因此,这种降雨格局将对浅层土壤水分及植被演替产生重要影响。     

祁连山北坡退化林地植被群落的自然恢复过程及土壤特征变化

采用长期定位观测的方法,研究了祁连山北坡退化林地人工抚育下2001-2008年间植被群落的自然恢复过程和土壤特征变化。结果表明:人为干扰消除后,退化林地群落环境逐渐优化,群落的科、属、种均明显增加,物种成员更替频繁;灌木和乔木物种出现后,群落垂直高度增大,群落结构出现成层现象;群落总体多样性指数呈不断增大的趋势,在空间结构上,Patrick丰富度指数、Shannon-Wiener多样性指数和Simpson优势度指数表现出:草本层>灌木层>乔木层的规律,而Pielou均匀度指数变化相反;土壤含水量、土壤有机碳和全氮含量随植被恢复均不断增加。在实施封育禁牧措施后,退化林地实现了由草本群落-灌木群落-乔木群落方向的快速演替,当恢复到早期的先锋乔灌混交阶段时,群落的物种组成、结构和多样性趋于复杂化,土壤性状也得到一定改善,显示出相对较好的适应性和恢复效果。     

高寒沙地典型固沙植物在沙丘不同地貌部位的水分利用特征

乌柳是高寒半干旱沙地植被恢复中常用的灌木树种,对乌柳水分利用来源的研究有助于理解沙地人工生态系统的稳定性维持及可持续发展。在青海湖湖东沙地,以沙丘不同地貌部位（迎风坡、丘顶和背风坡）生长的乌柳为研究对象,利用IsoSource模型分析植物生长季内的水分利用来源。研究结果表明：在生长季内,乌柳主要利用3个层次的土壤水,但不同地貌部位的乌柳在不同生长季利用不同层次的土壤水。生长季初期,迎风坡的乌柳以深层（60-150cm）土壤水为主要水源（50.28±18.11）%,而丘顶和背风坡的乌柳主要利用表层20cm的土壤水,利用比例分别为（79.96±7.59）%和（53.47±6.47）%。生长季中期,迎风坡的乌柳更多地利用中层20-60cm土壤水（45.52±26.91）%,丘顶的乌柳主要利用表层土壤水（45.17±20.14）%,而背风坡的乌柳以深层土壤水为主要水源（39.26±23.28）%。生长季末期,迎风坡的乌柳仍以（46.07±27.17）%的比例利用中层土壤水,丘顶的乌柳转而利用深层土壤水（44.25±26.23）%,而背风坡的乌柳却以表层土壤水作为主要水源（42.57±18.78）%。不同沙丘地貌部位乌柳对土壤水分的利用模式与其根系分布特征及其下方的土壤含水量具有相关性。     

沙丘多枝柽柳灌丛根层土壤含水量变化特征与根系水力提升证据

分析干旱区深根型荒漠植物的根层土壤水分是揭示荒漠植物与土壤水分关系机理的重要方面。在黑河中游一片风沙侵蚀区域的多枝柽柳(Tamarix ramosissima)人工林地中, 对表层0.3 m到3 m深的土壤不同深度的含水量进行了连续的动态观测。结果显示, 多枝柽柳根系层土壤含水量可以分为明显不同的3层: 浅层(0.2-1.7 m深)相对湿润层、中间(1.7-2.7 m深)相对干层和深层(2.7 m以下)有效含水层。在多枝柽柳生长盛期, 浅层相对湿润层土壤含水量呈现明显的昼夜变化特征, 同时, 在晚上植物根系与浅层土壤之间存在正水势梯度, 这说明存在根系水力提升现象。水力提升是干旱气候下根层浅层土壤含水量保持相对湿润的主要原因, 并因此维系浅层根系的发育, 也为多枝柽柳具备的防风固沙功能提供了可能的解释。据初步估算, 多枝柽柳根系水力提升占每天耗水量的5%-8%, 耗水的主要水分来源仍然是充足的土壤深层有效含水层。     

Reproductive traits of two closely related species-pairs on adjacent,different soil types in South African Fynbos

High levels of edaphic endemism and soil-related beta-diversity in Agulhas Plain fynbos communities suggest that reproductive traits of plants growing on different fynbos soils would be related to differences in soil regime. We investigated reproductive traits in two closely related Proteaceae species-pairs growing on adjacent soil types: Protea obtusifolia and Leucadendron meridianum occurring in shallow pockets of limestone-derived soils, and P. susannae and L. coniferum on the adjacent, uniformly deep colluvial sands.We found that species growing on the limestone soil comprised smaller plants, with fewer cones and seeds per plant, than species on the colluvial sands. These differences suggest that the small soil pockets of limestone soil limit plant size, in turn limiting the number of reproductive structures. Annual variation in cones and seed was the same in all species. There were no consistent trends in degree of serotiny, or sex allocation across soil types. The higher cone and seed predation levels of both limestone species than the colluvial sands species were ascribed to the higher plant densities of the former leading to lower insect search times. It was concluded that there were no overall patterns in reproductive traits that could be ascribed to differences in soil regime, other than through size-related effects. Fire regime is likely to have played a more important role in determining reproductive traits.Department of Statistical Sciences, University of Cape Towm