Seedling survival of Mediterranean shrubland species in relation to root:shoot ratio, seed size and water and nitrogen use

1. We hypothesized that in Mediterranean plant communities seedling survival of different species during the drought period would be related to their ability to use below-ground resources, particularly water and nitrogen.
2. For 5 years we studied under field conditions the summer seedling survival of 11 dominant species of a Spanish Mediterranean shrubland. We related seedling survival of the different species to their seed size and root allocation estimated as the slope of the function root biomass vs log shoot biomass. We used δ¹³C for the estimation of water-use efficiency and δ¹⁵N and nitrogen concentration to determine the sources of nitrogen utilized. We correlated these variables with root allocation.
3. Seedling survival of the different species was positively correlated with root allocation and seed size. Root allocation was also positively correlated with seed size. δ¹⁵N and nitrogen concentration were also positively correlated with root allocation, but δ¹³C was not.
4. Under the relatively moist conditions occurring during summer 1996, higher root allocation was associated with the use of nitrogen from more nitrogen saturated microsites.     

Tertiary relict trees in a Mediterranean climate: abiotic constraints on the persistence of Prunus lusitanica at the eroding edge of its range

Aim To investigate the ecophysiological traits allowing persistence of a subtropical relict tree (Prunus lusitanica L.) under a dry Mediterranean climate at the eroding edge of its range. Location A glasshouse for the study under controlled conditions and two marginal populations located in riparian forests of central Spain and exposed to summer drought, in contrast to subtropical populations that grow in mountain cloud forests. Methods Two experiments were conducted to assess tolerance to the abiotic conditions found in riparian habitats. In a glasshouse experiment, gas‐exchange and light‐use parameters were measured to evaluate seedling responses to a factorial combination of irradiance (60%, 10%, 2% and 0.5% full sun) and moisture (well watered control and drought stress). In a parallel field experiment, irradiance and soil moisture were measured as predictors of seedling survival at two sites in which half the seedlings were subjected to a threefold increase in summer precipitation by adding water every 10 days. Results  Soil moisture strongly determined survival both in the glasshouse and in the two field sites. In the field, periodic addition of water failed to increase survival. Water‐use efficiency (WUE) increased with drought. Seedlings did not tolerate deep shade (2–0.5%) and their performance and survival were better when exposed to moderate (10%) or high (60%) irradiance. The effect of water stress on seedling performance was stronger at moderate irradiance, moderate at high irradiance and negligible at very low light. Seedling size (height and number of leaves) attained after 1 month of experimental treatments had a positive effect on survival at the end of the summer, hence greater survival was not achieved at the expense of vertical growth. Main conclusions While studies in Macaronesia have shown that P. lusitanica occupies a wide range of moisture conditions along mountain slopes, it behaves as an obligate riparian species in dry peripheral populations. Intolerance to deep shade and tolerance to moderate and high irradiance allow the species to grow in small and moist gaps, or in treeless river sections. Despite the species’ low range filling in marginal, drought‐prone regions, long‐term persistence might have been achieved through shifts to riparian habitats serving as local refuges.     

Ectomycorrhizal colonization and intraspecific variation in growth responses of lodgepole pine

Across different host plant species, the effects of mycorrhizal colonization on host growth parameters can vary, but intraspecific variation in this relationship has rarely been measured. We tested the direction and consistency of the relationship between ectomycorrhizal colonization level and growth responses across seed families of Pinus contorta var. latifolia. Root tips of seedlings from eight full sib seed families varied in levels of ectomycorrhizal fungal colonization from 39% to 100%. We observed positive, negative, or neutral relationships between colonization level and shoot mass, depending on plant family. For the majority of seed families no relationship was observed between colonization level and root mass; however, two seed families showed negative relationships. Shoot height differed only by seed family. Results from our study indicate that the relationship between colonization level and host growth depends on host genotype. We suggest that models of plant intraspecific interactions should consider ectomycorrhizal associations when assessing phenotypic variability.     

模拟干旱胁迫对巴东木莲种子萌发和芽苗生长的影响

巴东木莲原生种群天然更新差,林下鲜见更新幼苗或幼树.为探究其种子萌发和芽苗在生长期对干旱胁迫的耐受性,该文以巴东木莲种子为材料,采用不同质量浓度的聚乙二醇(PEG-6000)模拟干旱胁迫,分析干旱胁迫对其种子萌发、芽苗生长及其相关的生理生化指标的影响.结果表明:(1)不同质量浓度的PEG溶液对巴东木莲种子萌发和芽苗生长...     

Phenotypic plasticity facilitates resistance to climate change in a highly variable environment

Increased summer drought will exacerbate the regeneration of many tree species at their lower latitudinal and altitudinal distribution limits. In vulnerable habitats, introduction of more drought-tolerant provenances or species is currently considered to accelerate tree species migration and facilitate forest persistence. Trade-offs between drought adaptation and growth plasticity might, however, limit the effectiveness of assisted migration, especially if introductions focus on provenances or species from different climatic regions. We tested in a common garden experiment the performance of Pinus sylvestris seedlings from the continental Central Alps under increased temperatures and extended spring and/or summer drought, and compared seedling emergence, survival and biomass allocation to that of P. sylvestris and closely related Pinus nigra from a Mediterranean seed source. Soil heating had only minor effects on seedling performance but high spring precipitation doubled the number of continental P. sylvestris seedlings present after the summer drought. At the same time, twice as many seedlings of the Mediterranean than the continental P. sylvestris provenance were present, which was due to both higher emergence and lower mortality under dry conditions. Both P. sylvestris provenances allocated similar amounts of biomass to roots when grown under low summer precipitation. Mediterranean seedlings, however, revealed lower phenotypic plasticity than continental seedlings under high precipitation, which might limit their competitive ability in continental Alpine forests in non-drought years. By contrast, high variability in the response of individual seedlings to summer drought indicates the potential of continental P. sylvestris provenances to adapt to changing environmental conditions.     

Mechanisms of structural change derived from patterns of seedling emergence and mortality in a semi-natural meadow

Questions: Is seedling emergence limited by the set of viable seeds, by incompatibility between the phenology of seed shedding and timing of mowing, or by dry weather in germination periods? Does seedling mortality fluctuate with season and weather? Location: Negrentino, southern Alps, Switzerland. Methods: Fecundity estimates of the dominant grass Bromus erectus; highly frequent counts of spontaneous seedlings by species and calculation of a community-level average mortality rate across 5 years; species-level records of seed shedding date and measurements of seed mass; measurement of soil moisture. Results: B. erectus produced 143.9 viable seeds/m²/year while the density of its seedlings was a 55 times smaller fraction. Grasses had fewer seedlings than forbs and their phenology of seed shedding was less compatible with mowing date. Soil moisture was a strong determinant of seedling emergence in spring and less so in autumn. Average seedling mortality declined with age of the populations and reached a maximum in an extremely dry summer. In relatively wet summers establishment success was positively related to seed mass. Conclusion: Community structure is susceptible to drought through mechanisms that selectively reduce recruits of coexisting plant functional groups. We propose that (1) more frequent intense droughts tend to reduce species that depend on frequent recruitment from seed, hence favour long-lived clonally spreading species, (2) drought timing selects between species with different germination phenology and drought resistance, and (3) drought impacts can be mitigated by changing management regimes that affect seed shedding.     

Response of photosynthesis and respiration to temperature under water deficit in two evergreen Nothofagus species

Respiration and photosynthesis were studied in two Nothofagus species with different drought tolerance in order to evaluate the effect of water deficit on foliar carbon balance and the possible role of the alternative pathway on respiratory adjustment. We propose that under severe water deficit the more drought‐tolerant species N. dombeyi is able to decrease its respiration more than the less drought‐tolerant species N. nitida, thus carbon gain could be maintained when photosynthesis is suppressed by drought. Dark respiration (R_d) and carbon assimilation under saturating light (A_sat) were evaluated under seasonal field conditions and during drying and re‐watering cycles under glasshouse. In addition, respiratory pathway changes were evaluated by oxygen isotope fractionation. In the field, N. dombeyi displayed greater light‐saturated photosynthetic capacity than N. nitida, but R_d did not differ between species during summer. In the glasshouse, N. dombeyi displayed an unchanged rate of R_d and increased carbon loss under severe water deficit. Nothofagus nitida displayed a more flexible respiratory response to water deficit, with a lower thermal sensitivity of respiration (decrease in Q₁₀) and a decrease in R_d. This contributed to maintaining leaf carbon balance during the water deficit period. Respiratory electron flow was mainly via the cytochrome pathway for both species and under all treatments, indicating no strong participation of alternative respiration. Our results suggest that under severe water stress, N. dombeyi could be more injured than N. nitida and that the lack of control in the carbon loss under prolonged periods of drought could be limiting for its survival.     

荒漠绿洲过渡带一年生草本植物对干旱胁迫的响应

选取河西走廊荒漠绿洲过渡带典型一年生草本植物雾冰藜（Bassia dasyphylla）、虎尾草（Chloris virgata）和狗尾草（Setaria viridis）为研究对象,设置5个水分梯度（正常水分（CK）,轻度干旱（5d）、中度干旱（10d）、重度干旱（15d）,重度干旱（15d）复水）,分析了3种一年生草本植物生理和形态等性状对干旱胁迫的响应。结果表明：一年生草本植物可以通过生理反应（渗透调节）适应轻度和中度干旱胁迫,而通过个体形态来适应重度干旱胁迫。在轻度和中度干旱处理下,一年生草本植物通过调控叶片渗透调节物质脯氨酸、可溶性蛋白和可溶性糖维持叶片渗透压,提高保水能力,叶绿素含量增加,使丙二醛含量维持在较低的水平,同时,根系活力增强,有效促进了根系水分吸收;而在重度胁迫下,渗透调节物质作用降低,丙二醛含量迅速增加,导致可溶性蛋白含量下降,叶绿素分解加速,植物生长受到抑制,在有限的生物量下,一年生草本植物主要通过根系伸长、根长与茎长的比增加和减小茎长来适应重度干旱胁迫,最终导致了种子百粒重和结种数量下降。     

Why do early-emerging tree seedlings have survival advantages?: a test using acer rubrum (Aceraceae)

In forest understories, tree seedlings that germinate and emerge early in the growing season have significant survival advantages. We hypothesized that (a) early emergence is related to seed size and half-sib family, and (b) increased survival is due to reduced competition for resources. To test these hypotheses, we collected seeds of Acer rubrum from four half-sib families and planted them in the understory of a South Carolina floodplain forest. In one experiment, date of emergence was not related to seed size or half-sib family. However, our method of seed collection may have underestimated variability among half-sib families. In a second experiment, seeds were spread onto 20 understory plots, half of which were trenched to reduce root competition with overstory trees. One-year survival and biomass were greater for early- than for later-emerging cohorts. Trenching changed biomass distribution among aboveground parts and may have increased total biomass, but had essentially no impact on survival or root mass. Variation in plot flooding resulted in large differences in timing of emergence, survival, and biomass. For Acer rubrum, the survival advantage accorded by early emergence may be more related to flooding, light conditions, and genetic heterogeneity than to seed size or avoidance of root competition.     

Oak seedling survival and growth along resource gradients in Mediterranean forests: implications for regeneration in current and future environmental scenarios

Understanding seedling performance across resource gradients is crucial for defining the regeneration niche of plant species under current environmental conditions and for predicting potential changes under a global change scenario. A 2‐year field experiment was conducted to determine how seedling survival and growth of two evergreen and two deciduous Quercus species vary along gradients of light and soil properties in two Mediterranean forests with contrasting soils and climatic conditions. Half the seedlings were subjected to an irrigation treatment during the first year to quantify the effects on performance of an alteration in the summer drought intensity. Linear and non‐linear models were parameterized and compared to identify major resources controlling seedling performance. We found both site‐specific and general patterns of regeneration. Strong site‐specificity was found in the identity of the best predictors of seedling survival: survival decreased linearly with increasing light (i.e. increasing desiccation risk) in the drier site, whereas it decreased logistically with increasing spring soil water content (i.e. increasing waterlogging risk) in the wetter site. We found strong empirical support for multiple resource limitation at the drier site, the response to light being modulated by the availability of soil resources (water and P). Evidence for regeneration niche partitioning among Quercus species was only found at the wetter site. However, at both sites Quercus species shared the same response to summer drought alleviation through water addition: increased first‐year survival but not final survival (i.e. after two years). This suggests that extremely dry summers (i.e. the second summer in the experiment) can cancel out the positive effects of previous wetter summers. Therefore, an increase in the intensity and frequency of summer drought with climate change might cause a double negative impact on Quercus regeneration, due to a general reduction in survival probability and the annulment of the positive effects of (infrequent) ‘wet’ years. Overall, results presented in this study are a major step towards the development of a mechanistic model of Mediterranean forest dynamics that incorporates the idiosyncrasies and generalities of tree regeneration in these systems, and that allow simulation and prediction of the ecological consequences of resource level alterations due to global change.     

Rapid root extension during water pulses enhances establishment of shrub seedlings in the Atacama Desert

Questions: (1) What is the water threshold for shrub seedling establishment in arid scrubland? (2) How do seedling root growth morphological traits affect the water threshold required for seedling establishment? Location: Arid scrubland, Atacama Desert, north‐central Chile. Methods: We conducted a field experiment with nine native shrub species under a gradient of simulated rainfall to test if species with deep root architecture have higher seedling survival rates and establish more successfully during water pulses. Results: Seedling survival rate was very low; only 2% of the 12 150 planted seedlings survived the first summer drought. Seedling survival required at least 206 mm of water, which is twice the average rainfall and roughly equivalent to the precipitation during an ENSO (El Niño Southern Oscillation) event in this region. Seedling survival at the onset of the summer drought was best explained by leaf habit, root length and initial seedling size. However, only Senna cumingii seedlings survived through the first year. S. cumingii seedlings had distinctively longer roots than the other shrub species, enabling them to reach moister soil layers. Conclusions: Water conditions resembling rainy years enhance shrub seedling establishment in the Atacama Desert, but the effects of higher water availability are strongly dependent on the shrub species. Rapid and deep rooting appears to be a very important functional trait for successful first‐year survival in this arid system where water availability is highly episodic.     

Drought resistance and recovery in mature Bituminaria bituminosa var. albomarginata

Few studies have investigated the response of perennial legumes to drought stress (DS) and their ability, following rewatering, to regrow and restore photosynthetic activity. We examined these responses for two genotypes of drought‐tolerant tedera (Bituminaria bituminosa var. albomarginata) and one genotype of lucerne (Medicago sativa). Plants were grown outdoors in 1‐m deep PVC pots with a reconstructed field soil profile, regularly watered for 8 months (winter to mid‐summer), and then moved to a glasshouse where either watering was maintained or drought was imposed for up to 47 days, before rewatering for 28 days. Drought stress greatly decreased shoot dry matter (DM) production in both species. Lucerne plants showed severe leaf desiccation after 21 days of withholding water. Relative leaf water content (RWC = 42%) and midday leaf water potential (LWP = ?6.5 MPa) decreased in tedera in response to DS, whereas leaf angle (85°) and lateral root DM both increased. Proline and pinitol accumulated in tedera leaves during DS, and their concentration declined after rewatering. Nine days after rewatering, previously drought‐stressed tedera had similar RWC and LWP to well‐watered control plants. In tedera and lucerne, 28 days after rewatering, photosynthesis and stomatal conductance were greater than in the well‐watered controls. The lateral root DM for one tedera genotype decreased during the recovery phase but for lucerne, the lateral root DM did not change during either the drought or the recovery phases. Overall, the root systems in tedera showed greater plasticity in response to DS and rewatering than in lucerne. In conclusion, tedera and lucerne showed different physiological and morphological strategies to survive and recover from DS. Proline and soluble sugars may act as a carbon source for regrowth in tedera during recovery. In comparison with lucerne, tedera's more rapid recovery after rewatering should contribute to a greater aboveground DM yield under alternating dry and wet periods. Tedera genotypes are highly heterogeneous and selecting genotypes with enhanced concentrations of pinitol and proline could be a valuable tool to improve plant performance during DS and recovery.     

Drought survival,summer dormancy and dehydrin accumulation in contrasting cultivars of Dactylis glomerata

To study survival under prolonged and severe drought in the perennial grass Dactylis glomerata we compared dormant, resistant and sensitive cultivars (cvs.) in both field and glasshouse experiments. Water status, membrane stability and expression of dehydrins were assessed in the immature leaf bases, which are the last surviving organs. Analysis of leaf elongation and senescence of aerial tissues showed that dormancy was exhibited by the potentially dormant cultivar (cv.) only in the field. This cultivar exhibited a high survival rate, similar levels of dehydration and expression of a low-molecular weight (22–24 kDa) dehydrin in both drought and irrigated plants, whether fully dormant or not. At the same level of soil water deficit, there were no differences between the non-dormant drought resistant and drought sensitive cultivars in plant water status and membrane stability. However, the accumulation of dehydrins as drought progressed was markedly different between these cultivars and was associated with their contrasting survival. The possible role of the major low-molecular dehydrins in maintenance of cell integrity under dehydration is discussed with reference to both summer dormancy and survival under severe drought.     

The influence of soybean planting density on dinitrogen fixation and yield

We investigated the effect of planting density on soybean (Glycine max (L.) Merr.) yield in glasshouse and field experiments. Because net canopy photosynthesis increases with increasing plant density, we hypothesized that increasing planting density would result in increasing rates of dinitrogen fixation in soybeans and higher yields per unit land area.In glasshouse studies, Wayne variety soybeans were planted in 10-cm diameter pots, 1 plant pot^-1 in matrices of 10-, 15-, 20-, 25-, or 30-cm equidistant intervals. Bradyrhizobium japonicum inoculum was added to half of the plants in each treatment. Replicate measurements of total stem height, internode lengths, leaf mass, stem mass, root mass, nodule number, nodule mass, and nitrogenase activity were obtained at 3, 6, and 9 weeks post-emergence. Fruits were harvested and counted at week 14. As planting density increased there were (1) altered morphology and growth rates, (2) increased apparent specific nodule activity (SNA), (3) decreased nodule number and mass, and (4) nearly constant fruit and seed production/plant. Expressed on a unit area basis, nitrogen influx and yield increased geometrically as planting density increased, with maximum values observed for 10-cm plantings.Field studies of Wayne, Stein, Williams, and Gold Harvest soybean varieties were made in 1985. Plots were established containing 100 plants spaced at 10-, 20-, and 30-cm distances. Measurements made during the growing season and at harvest established the same relative trends identified from the glasshouse studies. Increasing plant densities resulted in higher yields per unit land. Varietal differences were almost significant.     

Influence of Nitrogen Supply and Water Stress on Growth and Nitrogen, Phosphorus, Potassium and Calcium Contents in Pearl Millet

Influence of supra-optimal concentrations of N on growth and accumulation of N, K, P and Ca in the shoots and roots in Pennisetum glaucum (L.) R.Br. under water stress was assessed in a pot experiment under glasshouse conditions. Thirty four-day-old plants of two lines, ICMV94133 and WCA-78, were subjected to 224, 336, or 448 mg(N) kg^–1(soil) and soil moisture 100 or 30 % of field capacity for 30 d. Increasing soil N supply decreased growth of both lines under water deficit. Nitrogen content in the shoots of both lines was not affected by supra-optimal levels of N or different watering regimes, but in contrast, the root N content was increased consistently in WCA-78 with increase in soil N content. Shoot P content increased considerably in WCA-78 at the two higher N contents, but it was significantly lower at drought stress than at well-watered treatment. In contrast, shoot or root P content in ICMV94133 did not differ under both watering regimes. Potassium content in the shoots of WCA-78 was considerably increased at the two higher N contents under drought conditions. Root K content was increased in WCA-78 at the highest N content under well-watered conditions, whereas the reverse was true in ICMV94133. Calcium content in the shoots of ICMV94133 was higher under drought stress compared with that at well-watered conditions, but such pattern was not observed in WCA-78. However, root Ca content increased in both lines with increase in N supply.     

Growth features of Acacia tortilis and Acacia xanthophloea seedlings and their response to cyclic soil drought stress

Seedlings of Acacia tortilis (Forsk) Hyne and Acacia xanthophloea Benth. were raised under controlled glasshouse conditions. Control plants were watered daily while other treatments involved withholding water for 2, 4 and 6 days with 1‐day rehydration to container capacity. Compared to A. tortilis, A. xanthophloea seedlings showed higher leaf area, relative growth rates and total dry weight production under adequate water supply conditions. However, with increased water stress, A. xanthophloea seedlings could not alter their pattern of carbon allocation, retaining their root : shoot (r : s) ratio of about 0.5. By comparison, A. tortilis seedlings shifted carbon allocation to the roots, leading to a r : s ratio of 1.5 in water‐stressed seedlings, compared to 0.5 in the control plants. The ability of A. tortilis to reallocate carbon to the roots away from the shoots and to actually increase root growth compared to A. xanthophloea was a dehydration postponement strategy that may be important in species survival during drought.     

Alleviation of Summer Drought Boosts Establishment Success of <Emphasis Type="Italic">Pinus sylvestris</Emphasis> in a Mediterranean Mountain: An Experimental Approach

We performed an irrigation experiment to study the impact of summer drought on Pinus sylvestris establishment at its southernmost distribution limit. Watering was done during the first growing season simulating mesic summer conditions, and we monitored the consequences for survival and growth during the first growing season and the delayed consequences on the second growing season. In addition, we considered the heterogeneity created by the microhabitats, where seeds are found after dispersal (bare soil, under shrubs, and under adult pines). Summer drought was the main mortality factor in all the microhabitats. Watering increased emergence and doubled seedling survival compared to non-watered control sites. Differences were even higher when the cumulative effect on emergence and survival was considered, with an overall recruitment of 22.4% in watered plots vs. 7.9% in control. Irrigation increased growth in bare soil and under shrubs, but had scant effect on growth under pines, suggesting that radiation was the limiting factor in this microhabitat. The positive effect of irrigation on growth parameters persisted during the second growing season despite water was not added the second year, showing delayed consequences of drought on seedling performance. Summer drought thus limits Pinus sylvestris establishment in these southernmost forests by reducing both recruitment and growth. This might lead to the development of a remnant dynamic in these relict populations under the current regional increase in dryness and rainfall variability associated with global warming.     

Differential effect of drought regimes on the seedling performance of six floodplain grassland species

• The performance of seedlings is crucial for the survival and persistence of plant populations. Although drought frequently occurs in floodplains and can cause seedling mortality, studies on the effects of drought on seedlings of floodplain grasslands are scarce. We tested the hypotheses that drought reduces aboveground biomass, total biomass, plant height, number of leaves, leaf area and specific leaf area (SLA), and increases root biomass and root‐mass fraction (RMF) and that seedlings from species of wet floodplain grasslands are more affected by drought than species of dry grasslands.
• In a greenhouse study, we exposed seedlings of three confamilial pairs of species (Pimpinella saxifraga, Selinum carvifolia, Veronica teucrium, Veronica maritima, Sanguisorba minor, Sanguisorba officinalis) to increasing drought treatments. Within each plant family, one species is characteristic of wet and one of dry floodplain grasslands, confamilial in order to avoid phylogenetic bias of the results.
• In accordance with our hypotheses, drought conditions reduced aboveground biomass, total biomass, plant height, number of leaves and leaf area. Contrary to our hypotheses, drought conditions increased SLA and decreased root biomass and RMF of seedlings. Beyond the effects of the families, the results were species‐specific (V. maritima being the most sensitive species) and habitat‐specific. Species indicative of wet floodplain grasslands appear to be more sensitive to drought than species indicative of dry grasslands.
• Because of species‐ and habitat‐specific responses to reduced water availability, future drought periods due to climate change may severely affect some species from dry and wet habitats, while others may be unaffected.

     

Species-mediated soil moisture availability and patchy establishment of Pseudotsuga menziesii in chaparral

The occurrence of mature individuals of Pseudotsuga menziesii in stands of Arctostaphylos species mark the initial stages of mixed evergreen forest invasion into chaparral in central coastal California. We planted two cohorts of P. menziesii seeds at three sites under stands of two Arctostaphylos species and Adenostoma fasciculatum in order to determine whether first-year seedling emergence and survival, particularly during the regular summer drought, underlie the spatial distribution of mature trees observed in chaparral. Regardless of the chaparral species they were planted under, P. menziesii seeds that were not protected from vertebrate predation displayed very little emergence and no survival. In contrast, emergence of P. menziesii that were protected from vertebrate predators was much higher but still did not significantly differ among the three chaparral species. However, survival of protected seedlings under Arctostaphylos glandulosa was much greater than under A. fasciculatum, with intermediate survival under Arctostaphylos montana. While mortality of protected seedlings due to insect herbivory, fungal infection, and disturbance displayed no consistent patterns, summer drought mortality appeared to drive the patterns of survival of P. menziesii under the different chaparral species. These emergence, mortality, and survival data suggest that spatial patterns of P. menziesii recruitment in chaparral are driven by first-year summer drought seedling mortality, but only in years when seeds and seedlings are released from vertebrate predation pressure. Because the first-year drought mortality and survival patterns of P. menziesii seedlings differed strongly depending on the chaparral species, we examined the additional hypothesis that these patterns are associated with differences in the availability of soil moisture under different chaparral species. Both higher survival and lower drought mortality of P. menziesii seedlings were associated with higher soil water potential under Arctostaphylos stands during the summer drought, especially in the subsurface soil. The data suggest that Arctostaphylos stands, particularly stands of A. glandulosa, ameliorate xeric summer conditions to a degree that facilitates first-year establishment of P. menziesii and strongly influences spatial distribution of mature trees. Received: 18 September 1998 / Accepted: 23 December 1998     

Seed size: phylogeny and adaptation in two closely related Proteaceae species-pairs

Summary We studied seed size, seed nutrient status and seedling growth of two closely related fynbos Proteaceae species-pairs growing on juxtaposed soils of different nutrient and moisture status. Seeds had a greater mass and higher phosphorus and nitrogen contents for species occurring on limestone (higher nutrient and moisture contents) than those on the colluvial sands (lower nutreints and moisture). This trend was found within, but not across genera, stressing the importance of phylogeny in interpreting adaptations. It would be difficult to test for the effects of either nutrients or moisture separately, since the same advantage of enhanced seedling size, and hence survival in a stressed environment, applies to both factors. The increased root: shoot ratios (using lengths) of the Leucadendron species relative to the Protea species are interpreted as an attempt to overcome a phylogenetic constraint that results in smaller seed size in the former genus.