Direct and indirect interactions among plants explain counterintuitive positive drought effects on an eastern Mediterranean shrub species

Coexistence of woody and herbaceous plants may be governed by a complex set of direct and indirect interactions, whose relative importance have been rarely assessed. We experimentally studied woody species establishment in a mixed plant community by disentangling the potential role of such biotic interactions and the effect of environmental variations on them. Seedling establishment of the common eastern Mediterranean shrub species Sarcopoterium spinosum was investigated under different rainfall and light conditions, combined with the effect of the presence of adult shrubs and annual neighbors. We predicted that seedlings will be directly affected by competition with annuals with increasing water availability, while direct effects of adult shrubs will be positive via amelioration of water stress. Indirect effects were expected beneath shrub canopies due to reduced water stressed and light availability for both annuals and shrub seedlings, which may intensify competition between annuals and shrub seedlings. To test these predictions we performed field and garden experiments in which we combined manipulation of shrub and annual presence with manipulations of water availability and light conditions to simulate the effect of shrub canopy. In contrast to our prediction, shrub seedling establishment was not facilitated but inhibited by adult shrubs because of light limitation. As expected, annuals had direct negative effects on shrub seedlings under wet conditions, which shifted to neutral or positive effects under dry conditions. Thus, interactions among shrubs and annuals, and in particular the release from competition during drought years, leads to a counterintuitive positive effect of drought on shrub seedling establishment. Our findings point to the importance of experimentally studying multidimensional interactions for coexistence of different life forms and to the underestimated role of light for success in water‐limited ecosystems.     

Tree establishment along an ENSO experimental gradient in the Atacama desert

Questions: (1) What are the roles of regional climate and plant growth rate for seedling establishment during ENSO rainy pulses along the western coast of South America? (2) What is the water threshold for tree seedling establishment in these arid ecosystems? Location: Atacama Desert, western South America: Piura (5°10’ S, 80°37’ W), Mejia (17°00’ S, 71°59’ W), Fray Jorge (30°41'S,71°37'W). Methods: We experimentally simulated a gradient of ENSO rainfall in three locations encompassing the total extent of the Atacama Desert to test the relative importance of regional climate for seedling establishment during rainy pulses. We also carried out a common garden experiment to test the role of potential interspecific differences in growth rate among two Prosopis tree species. Results: Water threshold for seedling survival increased towards the south with less than 27 mm required in Piura, 100 mm in Mejia and 450 mm in Fray Jorge. We found that seedling survival and growth rate (shoots and roots) were much higher in Piura than in the other two sites for both Prosopis species. Conclusions: Our results indicate that tree establishment during rainy pulses is more likely to be successful in regions where rain falls during warm months and stimulates fast plant growth, and where loose soil texture facilitates deep root growth and therefore access to more stable water sources.     

Winners and losers: tropical forest tree seedling survival across a West African forest–savanna transition

Forest encroachment into savanna is occurring at an unprecedented rate across tropical Africa, leading to a loss of valuable savanna habitat. One of the first stages of forest encroachment is the establishment of tree seedlings at the forest–savanna transition. This study examines the demographic bottleneck in the seedlings of five species of tropical forest pioneer trees in a forest–savanna transition zone in West Africa. Five species of tropical pioneer forest tree seedlings were planted in savanna, mixed/transition, and forest vegetation types and grown for 12 months, during which time fire occurred in the area. We examined seedling survival rates, height, and stem diameter before and after fire; and seedling biomass and starch allocation patterns after fire. Seedling survival rates were significantly affected by fire, drought, and vegetation type. Seedlings that preferentially allocated more resources to increasing root and leaf starch (starch storage helps recovery from fire) survived better in savanna environments (frequently burnt), while seedlings that allocated more resources to growth and resource‐capture traits (height, the number of leaves, stem diameter, specific leaf area, specific root length, root‐to‐shoot ratio) survived better in mixed/transition and forest environments. Larger (taller with a greater stem diameter) seedlings survived burning better than smaller seedlings. However, larger seedlings survived better than smaller ones even in the absence of fire. Bombax buonopozense was the forest species that survived best in the savanna environment, likely as a result of increased access to light allowing greater investment in belowground starch storage capacity and therefore a greater ability to cope with fire. Synthesis: Forest pioneer tree species survived best through fire and drought in the savanna compared to the other two vegetation types. This was likely a result of the open‐canopied savanna providing greater access to light, thereby releasing seedlings from light limitation and enabling them to make and store more starch. Fire can be used as a management tool for controlling forest encroachment into savanna as it significantly affects seedling survival. However, if rainfall increases as a result of global change factors, encroachment may be more difficult to control as seedling survival ostensibly increases when the pressure of drought is lifted. We propose B. buonopozense as an indicator species for forest encroachment into savanna in West African forest–savanna transitions.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Survival and growth responses of Myricaria laxiflora seedlings to summer flooding

Myricaria laxiflora, an endangered shrub species distributed along the banks of the Yangtze River in the Three Gorges area, is completely submerged from June to October every year. It is generally assumed that summer flooding has a strong impact on the survival and growth of seedlings. We designed an outdoor randomized block experiment on the responses of seedling survival and growth to different flooding depth and flooding duration treatments during the flood season in the Three Gorges area. Seedling survival rate, aboveground biomass, belowground biomass, total biomass, root depth, length of primary branch and the number of primary and secondary branches were examined.M. laxiflora was found to acclimate to summer flooding by becoming dormant and losing biomass. Seedlings of M. laxiflora ceased growing during the summer flooding season, regardless of the flooding depth and flooding duration they were subjected to. The number of primary and secondary branches, aboveground biomass and total biomass of seedlings was reduced with prolonged flooding. The length of primary branches and aboveground biomass were more sensitive to flooding than other measured parameters and differed significantly between the onset and the end of flooding.In each flooding treatment most seedlings of M. laxiflora survived a flooding period of 2 months and recovered rapidly after the flooding was terminated in September. After 3 months of recovery, aboveground biomass, total biomass and the number of the primary branches increased significantly. Furthermore, seedling survival and growth in the flooding treatments were not significantly different from the controls both during the summer flooding stage and in the recovery stage. All of these results suggest that summer flooding does not affect seedling survival and growth in this species. On the contrary, flooding released seedlings from the stress of drought during summer and facilitated seedling establishment. M. laxiflora appears to cope adaptively with the flooding cycle by going into a state of dormancy during the flood season.     

Does shrub invasion indirectly limit grass establishment via seedling herbivory? A test at grassland‐shrubland ecotones

Question: Does shrub invasion at ecotones indirectly limit grass establishment by increasing mammalian seedling herbivory? Location: Chihuahuan Desert, New Mexico, USA. Methods: We tested the hypothesis that herbivore‐related mortality of seedlings of the dominant perennial grass Bouteloua eriopoda would be highest in shrub‐dominated portions of grassland‐shrubland ecotones. We tested the hypothesis in two Chihuahuan Desert sites featuring similar shrub encroachment patterns but different shrub species, grass cover, and different abundances of small mammals. Within each site we transplanted B. eriopoda seedlings to grass‐dominated, middle, and shrub‐dominated positions of replicate ecotones during the time of year (mid‐summer) when they would naturally appear and monitored seedling fates. We estimated population size/activity of putative small mammal herbivores. Results: Seedlings were killed by mammals in greater numbers in shrubland than in grassland or middle ecotone positions at the site with large herbivore numbers. At the site with low herbivore numbers, most seedlings were killed in middle ecotone positions. The abundance patterns of herbivores did not parallel patterns of seedling herbivory across the ecotones or between sites. Conclusions: Seedling herbivory is an important process and is related to vegetation composition, but the mechanisms underlying the relationship are not clear. We speculate that variation in small mammal foraging behavior may contribute to seedling herbivory patterns. Restoration strategies in the Chihuahuan Desert need to account for the abundance and/or behavior of native herbivores.     

Growth and biomass allocation of shrub and grass seedlings in response to predicted changes in precipitation seasonality

Anthropogenic emissions contribute to an annual 0.5% increase in atmospheric CO₂. As global CO₂ levels increase, regional precipitation patterns will likely be altered. Our primary objective was to determine whether a reduction in summer precipitation or an increase in winter/spring precipitation, predicted by global climate change models, will favor the establishment of C₄ grasses or C₃ shrubs in southern savannas. Our secondary objective was to determine how defoliation and microsite light availability interact with altered precipitation regimes to influence grass and shrub seedling growth and biomass allocation patterns. Seedlings of 3 shrub species (Prosopis glandulosa var. glandulosa, Acacia berlandieri, and A. greggii var. wrightii) and 3 grass species (Aristida purpurea var. wrightii, Setaria texana, and Stipa leucotricha) were watered based on probable changes in precipitation in a CO₂ enriched atmosphere (0.6, 0.8, and 1.0 current ambient summer precipitation and 1.0, 1.15, and 1.30 current winter/spring precipitation). Seedlings were defoliated at 3 levels (non-defoliated, single defoliation, and repeated defoliation) within 2 levels of microsite light availability (100 and 50% ambient). Defoliation significantly reduced total shrub and grass seedling biomass. Reducing light availability decreased shrub seedling root:shoot ratio, but total biomass was not significantly affected. Grass seedling biomass and root:shoot ratio decreased when light availability was reduced. Changing the seasonality of precipitation by reducing summer rainfall or increasing winter/spring rainfall did not significantly influence growth or biomass allocation of grass and shrub seedlings in a semiarid savanna. Microsite variations in defoliation intensity and light availability influence seedling growth and biomass allocation more than changing seasonality of precipitation. Shrub and grass seedling establishment and growth on semiarid rangelands are already limited by summer precipitation, so a further reduction as proposed by climate change models will have a limited impact on seedling dynamics.     

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.     

Establishing Native Grasses in a Big Sagebrush–Dominated Site: An Intermediate Restoration Step

Many semiarid rangelands in the Great Basin, U.S.A., are shifting dominance to woody species as a consequence of land degradation including intense livestock grazing and fire suppression. Whereas past rehabilitation efforts in Big sagebrush (Artemisia tridentata) steppes removed the shrub and added introduced forage grasses to successfully shift communities from shrublands to grasslands, current consensus is that native species should be included in restoration projects and that retention of some woody plants is desirable. We examined the potential for interseeding grasses into dense shrub communities as a precursor to thinning shrubs and releasing grasses from shrub interference. We compared seedling establishment of the native grass, Bluebunch wheatgrass (Pseudoroegneria spicata), with that of the Eurasia grass, Crested wheatgrass (Agropyron desertorum), in dense Ar. tridentata stands. Shrubs may play an important role as nurse plants for seedling establishment (reduced solar radiation, “island of fertility” effect) but result in highly contrasting light environments and root interference for seedlings. In experimental plots, we examined effects of Ar. tridentata shade levels (0, 40, 70, and 90% reduction of solar radiation) and initial root exclusion (present/absent) on the establishment and growth of P. spicata and Ag. desertorum seedlings. With this design we evaluated the interference effects of Ar. tridentata on the two grasses and identified the most beneficial microsites for grass restoration in Ar. tridentata–dominated communities. We predicted seedling survival and growth to be greater under moderate shade (40% reduction) and limited root competition than under no or strong shade conditions (0 and 90%) and unrestricted root interactions. Fifty to 85% of the P. spicata and Ag. desertorum seedlings survived the dry summer months of 1995 and 1996 and the intervening winter. Neither shading nor root exclusion from Ar. tridentata affected final seedling survival of either species. Seedling biomass of both grass species was negatively affected by initial root interactions with Ar. tridentata. However, the analysis of seedling biomass variability (coefficient of variation) indicated that in all shade and root‐exclusion treatments, some seedlings of both species developed to large individuals to survive in Ar. tridentata–dominated rangelands. Thus, the use of interseeding techniques shows promise for restoring herbaceous species in dense Ar. tridentata stands and should be given further consideration when shrub retention is an important consideration.     

Habitat-specific responses of seed germination and seedling establishment to soil water condition in two Rheum species in the high Sino-Himalayas

Knowledge of how germination and seedling establishment respond to soil water condition is crucial for plant conservation under global warming and land-use changes. We tested the flooding and drought tolerance of two plant species with different occurrences along a soil water gradient by assessing seed germination, seedling survival, seedling growth, and root characteristics. In the high Sino-Himalayas, Rheum alexandrae typically occurs in wetlands, R. nobile in scree or open slope with well-drained soils. Seeds and seedlings of the two species were subjected to different soil water conditions in controlled greenhouse experiments. Seed germination in both species was inhibited by high soil water content; however, seeds of R. alexandrae were more tolerant to flooding, especially to submergence. Seedling survival, biomass accumulation, root diameter, and root porosity of R. alexandrae increased significantly with increased soil water content, but submergence was lethal for seedlings. Seedling survival, biomass accumulation, and root length of R. nobile increased significantly in response to reduced soil water content. These results indicate that in the two species, seed germination and seedling establishment in response to different soil water condition are habitat-specific. Because both species are susceptible to moderate changes in soil water condition, their species-specific requirements with respect to this factor should be a consideration when planning their conservation.     

Effects of position, understorey vegetation and coarse woody debris on tree regeneration in two environmentally contrasting forests of north-western Patagonia: a manipulative approach

Aim To investigate the differential effects of position within gaps, coarse woody debris and understorey cover on tree seedling survival in canopy gaps in two old‐growth Nothofagus pumilio (Poepp. & Endl.) Krasser forests and the response of this species to gaps in two forests located at opposite extremes of a steep rainfall gradient. Location Nahuel Huapi National Park, at 41° S in north‐western Patagonia, Argentina. Methods In both study sites, seedlings were transplanted to experimental plots in gaps in three different positions, with two types of substrate (coarse woody debris or forest floor), and with and without removal of understorey vegetation. Survival of seedlings was monitored during two growing seasons. Soil moisture and direct solar radiation were measured once in mid‐summer. Seedling aerial biomass was estimated at the end of the experiment. Results Mid‐summer soil water potential was lowest in the centre of gaps, in plots where the understorey had been removed, and highest at the northern edges of gaps. Direct incoming radiation was highest in gap centres and southern edges, and lowest at northern edges. Seedling mortality was highest in gap centres, in both sites. Coarse woody debris had a positive effect on seedling survival during summer in the mesic forest and during winter in the xeric forest. The removal of understorey cover had negative effects in gap centres during summer. Seedling final aerial biomass was positively affected by understorey removal and by soil substrate in both sites. In the dry forest gaps, seedling growth was highest in northern edges, whereas it was highest in gap centres in the mesic forest. Overall growth was positively related to survival in the xeric forest, and negatively related in the mesic forest. Main conclusions Survival and growth were facilitated by the shade of gap‐surrounding trees only in the xeric forest. Understorey vegetation of both forests facilitated seedling survival in exposed microsites but competed with seedling growth. Nurse logs were an important substrate for seedling establishment in both forests; however, causes of this pattern differed between forests. Water availability positively controls seedling survival and growth in the xeric forest while in the mesic forest, survival and growth are differentially controlled by water and light availability, respectively. These two contrasting old‐growth forests, separated by a relatively short distance along a steep rainfall gradient, had different yet unexpected microenvironmental controls on N. pumilio seedling survival and growth. These results underscore the importance of defining microscale limiting factors of tree recruitment in the context of large‐scale spatial variation in resources.     

Tissue water relations of three chaparral shrub species after wildfire

Summary We compared the tissue water relations among resprouts and seedlings of three chaparral species during the first summer drought after wildfire. Two of the species, Rhus laurina and Ceanothus spinosus recover after fire by a combination of resprouting and seedling establishment (facultative resprouters), whereas a third species, Ceanothus megacarpus recovers by seedling establishment alone (obligate seeder). Our objectives were to document any differences in tissue water characteristics that might arise between resprouts and seedlings and to test the hypothesis that seedlings of obligate seeders develop more drought tolerant characteristics of their tissues than seedlings of facultative resprouters. We found that resprouts had much higher predawn values of water potential, osmotic potential, and turgor potentials than seedlings. Predawn turgor potentials of resprouts were 1.5 MPa through July and August when turgor potentials for seedlings remained near 0 MPa. During summer months, midday water potentials were 2 to 3 MPa higher for resprouts than seedlings and midday conductances of resprouts were two to five fold greater than those of seedlings. Even though resprouts did not experience severe water stress like seedlings, their tissue water characteristics, as determined by pressure-volume curve analyses, were similar by the peak of the drought in August. Further-more, the tissue water characteristics of seedlings from the obligate seeder, C. megacarpus, were similar to those of facultative resprouters — R. laurina, and C. spinosus. We attribute the observed differences in plant water status between resprouts and seedlings to differences in rooting depths and access to soil moisture reserves during summer drought. We conclude that the higher growth rates, photosynthetic performance, and survivorship of postfire resprouts are primarily a result of higher water availability to resprouting tissues during summer months. It appears that the greater seedling survivorship during summer drought observed for the obligate seeder, C. megacarpus, is not associated with more favorable tissue water characteristics.     

Seedlings and ramets recruitment in two rhizomatous species of Rupestrian grasslands: Leiothrix curvifolia var. lanuginosa and Leiothrix crassifolia (Eriocaulaceae)

Leiothrix curvifolia var. lanuginosa and Leiothrix crassifolia are endemic and sympatric species in the Brazilian rupestrian grasslands, a habitat that has a predominance of sandy and shallow soils with low water retention. Based on the premise that soil moisture is one of the abiotic factors that affects most reproduction in plants, we hypothesized that the flowering phenology events and establishment of sexual and vegetative offspring would occur in the periods of higher soil water availability. We marked 478 ramets distributed among 100 genets of L. curvifolia var. lanuginosa and 693 ramets distributed among 100 genets of L. crassifolia, so that they could be observed monthly along the two rainy seasons from December 2003 to 2004. Both species showed phenological synchrony in the flower heads and seedlings production with soil moisture availability. Seedling mortality was intense in the dry period. Unlike the seedlings, the ramets survived was 100%. The greater capacity of ramets to survive can result from a much greater biomass compared with seedlings, and ramets become adult much faster. We conclude that for a successful seedling establishment, the synchronization with the rainy season was required, and moreover, that repeated seedling recruitment can be important for the maintenance of local populations of these species which suffer from high seedling mortality in the drought period. It is likely that the coincidence of the rainy period with seedling establishment is an important factor that determines the flowering phenological pattern of L. curvifolia var. lanuginosa and L. crassifolia in rupestrian grasslands.     

Constraints to seedling success of savanna and forest trees across the savanna-forest boundary

Tropical savannas and closed forests are characterized by distinct tree communities, with most species occurring almost exclusively in only one of the two environments. The ecology of these two groups of species will largely determine the structure and dynamics of the savanna-forest boundary, but little is known about the ecological and physiological differences that might control their distributions. We performed field and nursery experiments to compare seedling establishment success, predawn leaf water potential, biomass allocation, and root carbohydrate concentration of congeneric species, each composed of one savanna species and one forest species. Seedling establishment of savanna and forest species responded differently to vegetation cover, with forest species having lowest establishment success in the open savanna and savanna species having lowest success in forest. Subsequent survival followed similar patterns, resulting in even greater differences in cumulative success. The low survival of forest species in the savanna appears related to drought stress, as seedlings of forest species had lower predawn leaf water potential than savanna species. Seedlings of savanna species had greater root: shoot ratios and root total nonstructural carbohydrate (TNC) concentration, particularly among evergreen genera. Among evergreen genera, root TNC per shoot mass, which may largely determine resprout capacity, was seven times higher in savanna species than forest species. Although water availability and microclimate may reduce the success of forest species, these factors appear unable to completely exclude forest seedling establishment in savanna. Fire, on the other hand, appears to be a much more absolute constraint to success of forest species in savanna.     

Differential effect of shade,water and soil type on emergence and early survival of three dominant species of the Atacama Desert

Understanding the regeneration niche of species may allow us to gain insight into how communities are structured. In deserts, the regeneration niche is usually related to spaces beneath shrubs where shade cast by shrubs creates microenvironments that benefit seedlings and where even small amounts of rain may favour germination and establishment. Shade and water may also interact with different types of soils. However, species may have different requirements for germination and seedling survival. We could expect that shrub species with different drought tolerances exhibit different responses to the combination of these factors. We ask if responses of dominant species of the Atacama Desert to abiotic factors (shade, water and soil type) are related to their drought tolerance, a topic not exhaustively explored in shrubs growing in true deserts. We conducted two factorial experiments. The first one was designed to evaluate how shade (microhabitat) in combination with water may affect germination (emergence) and early survival. In the second experiment, we assessed the influence of shade in relation to soil type. Each species responded distinctively to the three variables under study, but in general, their emergence responses were more influenced by water (more water, greater emergence) than by microhabitat or soil type. Survival was influenced both by microhabitat and by water and was higher under shade and abundant water. Soil type affected only one of our species in terms of emergence. Species responses in general depended on their tolerance to stress. In one species, there was indication of a seed–seedling conflict. Our results show similar species responses to environmental constraints but also more or less unique responses that are related to their tolerance to drought and which may ultimately permit species coexistence. We found that shade may not be important for germination but may be crucial for survival in dry years.     

Seed mass and summer drought survival in a Mediterranean-climate ecosystem

We related seed mass to summer drought survival in a Mediterranean-climate ecosystem. Previous experimental evidence linking seed mass and survival under drought is limited and at times contradictory. We tracked summer drought survival among four families/subfamilies at a restoration site in southwestern Australia. We coupled these observations with a glasshouse experiment assessing the growth and root morphology of Acacia and Eucalyptus species, with a range of seed masses, under mild and severe drought compared with a well-watered control. Summer drought survival in the field increased with seed mass across all four families/subfamilies. Seedling root biomass and length increased with seed mass consistently across five harvests over 60 days. Initial survival of seedlings in the glasshouse increased with seed mass and decreased with drought, but there was no interaction between the two. Greater absolute root investment provides a mechanism for both short and longer-term drought survival. Within-species variation in root growth may also affect the relative versus absolute survival advantage of large-seeded species. The benefits of large seed mass for establishment under environmental hazards are often considered to be temporary. Our results show that seed mass was correlated with other traits, including root length, which in turn, increased longer-term drought survival. Traits correlated with seed mass should therefore be considered in explanations of the ecological effect of seed mass variation.     

Differential survival of chaparral seedlings during the first summer drought after wildfire

Summary Big Pod Ceanothus (Ceanothus megacarpus) is an obligate seeder after fire; Laurel Sumac (Rhus laurina) is primarily a resprouter after fire. Both species commonly occur together in mixed stands and are dominant members of the coastal chaparral of southern California. We compared the mean survival of post-fire seedlings of each species during the first summer drought after fire and found C. megacarpus to have a mean survival of 54% while R. laurina had a mean survival of only 0.1%. Rooting dephs were similar between species but predawn water potentials and leaf temperatures were higher for R. laurina seedlings. Leaf temperatures for R. laurina reached a mean value of 46.8° C on hot, summer days, about 5° C higher than seedlings of C. megacarpus. By the end of the first growing season, 92% of all C. megacarpus seedlings had suffered herbivory compared to only 17% of all R. laurina seedlings. Herbivory did not appear to be the immediate cause of seedling mortality. Transect data indicated that full recovery of prefire species composition and density at our study site was likely but the mode of recovery was different for the species examined. R. laurina recovered primarily by sprouting, C. megacarpus totally by seedling establishment and a third species, Adenostoma fasciculatum (chamise), by a combination of sprouting and seedling establishment. We attribute the higher mortality of R. laurina seedlings to the greater sensitivity of its tissue to water stress. It may be that differential survival of shrub seedlings and differential modes of reestablishment after fire play an important role in maintaining species diversity in the chaparral communities of coastal, southern California.     

水分条件变化对哀牢山亚热带常绿阔叶林林下幼苗死亡率的影响

为探讨环境水分条件变化对亚热带森林林下乔木幼苗动态的影响,以及木材密度和幼苗在干旱中死亡率的关系,研究了哀牢山常绿阔叶林2005—2011年的总体和主要树种幼苗(黄心树、多果新木姜子、多花山矾、鸭公树和大花八角)的死亡率,并分析了幼苗死亡率和旱季幼苗根系所在土层土壤质量含水量、旱季降水量、旱季降水日数以及木材密度的关系。结果表明:1)幼苗死亡率与旱季降水日数、旱季浅层土壤平均质量含水量有显著的负相关关系,和旱季降水量无显著相关性;2)总幼苗及5种乔木幼苗的死亡率均在2010年(西南干旱)达到有观测以来最高,是2009年的2—10倍,其中多花山矾、黄心树幼苗的死亡率最高,大花八角幼苗的死亡率最低;3)在种间,幼苗2010年干旱中的死亡率和木材密度显著正相关,即木材密度较大的物种幼苗死亡率更高。研究表明林下幼苗由于根系较浅,对降雨变化较为敏感,因而受到了这次干旱的较大影响。由于木材密度较低的树种在干旱中有较低的死亡率,干旱频度和强度的增加可能使低木材密度幼苗的丰富度增加,森林的组成也将受到影响。     

The Effects of Drought and Shade on the Performance,Morphology and Physiology of Ghanaian Tree Species

In tropical forests light and water availability are the most important factors for seedling growth and survival but an increasing frequency of drought may affect tree regeneration. One central question is whether drought and shade have interactive effects on seedling growth and survival. Here, we present results of a greenhouse experiment, in which seedlings of 10 Ghanaian tree species were exposed to combinations of strong seasonal drought (continuous watering versus withholding water for nine weeks) and shade (5% irradiance versus 20% irradiance). We evaluated the effects of drought and shade on seedling survival and growth and plasticity of 11 underlying traits related to biomass allocation, morphology and physiology. Seedling survival under dry conditions was higher in shade than in high light, thus providing support for the “facilitation hypothesis” that shade enhances plant performance through improved microclimatic conditions, and rejecting the trade-off hypothesis that drought should have stronger impact in shade because of reduced root investment. Shaded plants had low biomass fraction in roots, in line with the trade-off hypothesis, but they compensated for this with a higher specific root length (i.e., root length per unit root mass), resulting in a similar root length per plant mass and, hence, similar water uptake capacity as high-light plants. The majority (60%) of traits studied responded independently to drought and shade, indicating that within species shade- and drought tolerances are not in trade-off, but largely uncoupled. When individual species responses were analysed, then for most of the traits only one to three species showed significant interactive effects between drought and shade. The uncoupled response of most species to drought and shade should provide ample opportunity for niche differentiation and species coexistence under a range of water and light conditions. Overall our greenhouse results suggest that, in the absence of root competition shaded tropical forest tree seedlings may be able to survive prolonged drought.     

Lack of recruitment in Lavandula stoechas subsp. pedunculata: a case of safe-site limitation

Lavandula stoechas subsp. pedunculata regeneration depends exclusively on the establishment of new individuals. Seed availability and seedling emergence and survival are therefore critical life stages and processes for species regeneration. In this study, seedling emergence and survival was monitored for two years in the scrub, both in clearings and adjacent to adult plants, and the surrounding perennial grassland, at 1, 3 and 5 m from the scrub. Soil seed bank spatial distribution was also studied for one year in the same two habitats, using the same sampling design. Soil seed availability in the scrub is high regardless of the distance from the adult individuals. On the contrary, the adjacent grassland shows a drastic fall in seed density, and almost no seedlings were observed there. In the scrub, seedling density was negatively related to distance from the three nearest adult plants in the clearings, and positively related to adult plant size beneath the adult Lavandula plants. There was also a negative relationship between seedling density and the percentage of bare soil. Only one seedling survived the first drought period, with no detection of effects of either position with respect to adult individuals or seedling density. We hypothesized that the study populations suffer a lack of appropriate safe sites within the scrubland while in the adjacent perennial grassland, observed low seed availability was added to safe-site limitation. That results in a lack of successful seedling establishments and a poor expansion potential of Lavandula scrublands, whose edges remain static in the short and medium term. As found in other Mediterranean scrubland, recruitment may only occur in years with particularly favourable weather, under disturbance regimes that increase seedling survival probability or when external dispersal agents increased seed availability in adequate places for Lavandula establishment.