Effects of moderate shade and irrigation with eutrophicated water on the nitrogen economy of Mediterranean oak seedlings

We evaluated the effects of moderate shade (43% vs. 100% of full sunlight) and irrigation with eutrophicated river water (daily vs. alternate-day watering) on growth and nitrogen economy of seedlings of three Mediterranean oak species, two evergreen (Quercus coccifera, Quercus ilex subsp. ballota) and a deciduous (Quercus faginea), grown in pots outdoors. Seedling biomass, N pool, N concentration and N losses by litter fall were measured at the beginning (March 2002) and end (November 2002) of a growing season. All species showed an increase of biomass and N pool under shade and/or high irrigation, while only Q. coccifera – from more arid regions – did the same under full sunlight and low irrigation. At the end of the experiment, biomass of the evergreens was higher in shade than in sun, and in high than in low irrigation, while Q. faginea – from more humid zones – responded to irrigation only. Shade-induced growth was accompanied by a decline in N concentration in the evergreens, but irrigation reduced N concentration only of Q. faginea. Shade, but not irrigation, reduced above-ground N loss. We conclude that both treatments differentially affected the evergreen and the deciduous oaks, probably due to differences in plant hydraulic and stomatal conductance. Although both treatments have similar effects on the growth of evergreens, they produced different effects on seedling N economy, which may have important consequences on future field seedling performance.     

Density Effects of a Dominant Understory Herb,Isoglossa woodii (Acanthaceae), on Tree Seedlings of a Subtropical Coastal Dune Forest

Suppression of tree seedlings by the understory is an important ecological filter with implications for tree diversity and dynamics. In a greenhouse competition experiment, we used seedlings of four canopy species from coastal dune forest (Diospyros natalensis, Euclea racemosa, Sideroxylon inerme and Apodytes dimidiata) to examine the relative competitive effects of the dominant understory herb Isoglossa woodii on seedling performance. We manipulated I. woodii density, light and nutrient levels and measured growth responses. Total seedling biomass decreased with density of I. woodii. The magnitude of biomass suppression with competitor density was similar among tree species. Consequently there was no discernable hierarchy of competitive ranking among tree species. The relative growth rate of seedlings decreased at higher densities of I. woodii and increased at higher nutrient levels but was unaffected by variation in light conditions. Aboveground biomass decreased at higher densities of I. woodii and at higher light levels but increased at higher nutrient levels. Size asymmetric competition for light and nutrients may be the major driver of aboveground interactions between tree seedling and I. woodii. While tree species showed no hierarchy of competitive ability their seedlings exhibited equivalent responses to competition from an understory dominant, permitting species coexistence and the maintenance of species diversity.     

Summer water stress and shade alter bud size and budburst date in three mediterranean Quercus species

Current environmental conditions are known to affect plant growth, morphology, phenology, and therefore, plant performance. However, effects of the previous-year environmental conditions can also affect plant structure by altering bud growth, and proportion and date of budburst. Here, we analysed the effects of previous-year water stress and shade on bud size, percentage, and date of budburst in seedlings of three co-occurring Iberian Quercus species in two independent experiments. Responses of apical, lateral, and basal buds were checked during an annual cycle. In the first experiment, seedlings of two evergreens (Q. coccifera L., Q. ilex subsp. ballota (Desf.) Samp.) and a deciduous-marcescent tree (Q. faginea Lam.) were grown under two levels of summer watering. In the second experiment, seedlings were grown under three light intensities. Soluble sugars and starch in shoots and roots were measured before budburst. Summer drought increased bud size of all species and advanced budburst of Q. ilex and Q. coccifera. Moderate and/or intense shade tended to reduce bud size and delay budburst in all species. These responses seem related to changes in the date of bud formation rather than to the amount of carbon reserves, which were reduced both by drought and shade. Treatments affected percentage of budburst in lateral buds, which was reduced by shade and water stress, probably leading to narrower crowns. These results show that previous-year environmental conditions are relevant for plant phenology and structure. The different responses in budburst date between the deciduous and the evergreens might alter their competition relationships at seedling stage.     

Competition by herbs as a limiting factor in shrub invasion in grassland: a test with different growth forms

Abstract. We tested the hypothesis that seedling establishment, the critical stage in the invasion of grassland by shrubs, is limited by competition with perennial grasses in seasonally wet/dry savannas. We placed seeds of two invasive exotic shrubs – Cryptostegia grandiflora, a woody vine, and Acacia nilotica, an arborescent legume – into pots with a wide range of existing above- and below-ground herbaceous biomass provided by either a tussock or a stoloniferous perennial grass. We also imposed different levels of watering frequency (5, 10 and 21 d), nutrient addition (+ and -) and grass clipping intensity (no clipping, clipped to 5 cm and clipped to 25 cm). There was no effect of any treatment on shrub seedling emergence or survival and all of the seedlings that emerged survived the 90-d growing period. Herbaceous competition also failed to have an effect on biomass accumulation in shrub seedlings. More frequent watering significantly increased above- and below-ground biomass accumulation for both shrub species and nutrient addition significantly increased Cryptostegia biomass accumulation. Based on these results, we question the proposition that reduction in competition by herbs via livestock grazing has been a significant factor in determining the rate or pattern of exotic shrub increase in the seasonally wet/dry tropics. We also question the suitability of the two-layer soil moisture hypothesis as a basis for management practices to control the ingress of woody species into grasslands and open savannas.     

More than just drought: complexity of recruitment patterns in Mediterranean forests

Understanding community dynamics during early life stages of trees is critical for the prediction of future species composition. In Mediterranean forests drought is a major constraint for regeneration, but likely not the only factor determining the observed spatial patterns. We carried out a sowing experiment aimed at identifying main filters during seed-seedling transition. Specifically, we studied seed fate (predation, fungi infection, emergence) and subsequent seedling performance (mortality during the first summer and overall recruitment after 2 years) of four co-occurring Mediterranean tree species (Quercus ilex, Quercus faginea, Juniperus thurifera, Pinus nigra). We related these processes to the dominant species composition, microhabitat heterogeneity, herb cover and seed mass. The identity of the dominant species in the forest canopy was more important for recruitment than the forest canopy being dominated by conspecific vs. heterospecific species. The patterns we found suggest that biotic interactions such as facilitation (lower mortality under the canopies) and herb competition (during emergence of J. thurifera) are relevant during recruitment. Moreover, our results pointed to ontogenetic conflicts regarding the seed mass of Q. faginea and to density-dependent seed mortality for Q. ilex, rarely described in Mediterranean ecosystems. We propose that our study species experience population growth in forests dominated by heterospecifics where the recruitment success depends on habitat heterogeneity and on moderated biotic and abiotic stresses created by each species. Our results reveal patterns and mechanisms involved in recruitment constraints that add complexity to the well-known drought-related processes in Mediterranean ecosystems.     

Competition between tree seedlings and herbaceous vegetation: support for a theory of resource supply and demand

1 We measured competition intensity (CI) between herbaceous vegetation and tree seedlings ( Quercus macrocarpa and Q. ellipsoidalis ) along an experimental moisture–light gradient. Contrasting theories were tested by comparing variation in competition intensity to changes in neighbour biomass and resource supply and demand.
2 CI based on survival was inversely correlated with net soil water supply (gross supply minus demand by herbaceous vegetation). CI was not positively correlated with either gross resource supply or neighbour biomass, contrary to predictions of Grime's triangular model for plant strategies.
3 Many of the inconsistencies and conflicting results that have characterized the recent literature on plant competition could be eliminated if changes in competition intensity along a resource gradient are compared with changes in net resource supply rather than changes in productivity or neighbour biomass.
4 Tree seedling success in savannas and grasslands may be strongly influenced by the intensity of competition from herbaceous vegetation. Factors that reduce soil water content are likely to increase competition intensity (and reduce seedling success) in these environments, while factors that increase soil water content will favour seedling success through decreased competition for water with herbaceous vegetation.     

Morphological and ecophysiological variation of the hybrid oak Quercus subpyrenaica (Q. faginea × Q. pubescens)

Natural hybridization is common among oaks. We studied the variability of morphological and ecophysiological variables in the hybrid Quercus subpyrenaica and its assumed parental species Q. faginea and Q. pubescens, which co-occur in NE Spain. To assess the fitness of these taxa we studied several ecophysiological variables (hydraulic conductivity, K_h; hydraulic specific conductivity, K_s; leaf specific conductivity, LSC; water potential corresponding to a 50% loss of conductivity, PLC₅₀; efficiency of light absorption, E_a). We performed a correspondence analysis (CA) to ordinate seedlings, grown under homogeneous environmental conditions, according to their plant and leaf morphology. The CA axis 1 synthesized intra-taxon variability, while the CA axis 2 summarized inter-taxa variability. Q. subpyrenaica showed a wide spectrum of forms, but they were overall closer to those of Q. faginea. We defined three phenotypes within the hybrid based on morphology, which were: (i) the robur group (Qs-r; auriculate leaf base, rounded lobe apex); (ii) Q. pubescens (Qs-p; rounded leaf base, acute lobe apex); and (iii) Q. faginea (Qs-f; acute leaf base, acute-spiny lobe apex). The mean values of K_s and PLC₅₀ arranged the hybrid groups in the same order as the ordination based on leaf morphology. The Qs-r group showed the highest values of K_s and PLC₅₀, while the Qs-f group showed the lowest. Both morphologically and ecophysiologically, the hybrids showed a wide range of values, which spanned and even exceeded the variation of parental taxa.     

长白山不同演替阶段温带森林林下草本植物对乔木幼苗的影响

为了解次生针阔混交林和阔叶红松林林下草本植物对幼苗生长和存活的影响,基于长白山次生针阔混交林样地(Ⅰ)和阔叶红松林样地(Ⅱ),以246个1 m×1 m幼苗样方中乔木幼苗为研究对象,通过去除草本植物的对照试验探究草本植物对乔木幼苗高度生长和存活率的影响。结果表明,(1)群落水平上,草本植物去除有助于林下乔木幼苗的高度生长。次生针阔混交林和阔叶红松林中幼苗高度生长量在除草后较对照组均有显著提高,且阔叶红松林中幼苗高度增长在对照组和处理组中均高于次生针阔混交林。(2)去除草本植物对不同年龄级水平乔木幼苗高度生长影响不同。次生针阔混交林中,去除草本显著促进四年生及以上幼苗高度生长,对一至三年生幼苗影响不显著;阔叶红松中去除草本显著促进一至三年生幼苗高度生长,对四年生及以上幼苗影响不显著。(3)除草处理后,水曲柳幼苗高度生长量在两处样地均显著增加,假色槭幼苗高度增长量只在次生针阔混交林中显著增加,而其他幼苗高度增长量只在阔叶红松林中显著增加。(4)次生针阔混交林中,幼苗存活率与草本多度和物种数呈正相关关系,与草本盖度无相关关系;阔叶红松林中幼苗存活率与草本物种数呈正相关关系,与草本多度和盖度无相关关系。结果表明,草本植物会抑制乔木幼苗高度生长;虽可能在一定程度上有助于改善微生境,但未显著提高幼苗存活率。     

Stem xylem features in three Quercus (Fagaceae) species along a climatic gradient in NE Spain

 Stem xylem features in two evergreen Quercus species (Q. coccifera and Q. ilex) and a deciduous one (Q. faginea) were analysed along an Atlantic-Mediterranean climatic gradient in which rainfall and winter cold experience strong variation. Mean maximum vessel diameter, vessel density, vessel element length, xylem transverse sectional area, Huber value (xylem transverse sectional area per leaf area unit), theoretical leaf specific conductivity (estimated hydraulic conductance per leaf area unit) and total leaf area were determined in 3-year-old branches. Q. faginea presented the widest vessels and the highest theoretical leaf specific conductivity while Q. coccifera showed the lowest total leaf area and the highest Huber value. Studied features did not exhibit significant correlations with mean minimum January temperature in any species but did show significant relationships with rainfall. In Q. coccifera, mean maximum vessel diameter, vessel element length and theoretical leaf specific conductivity increased with higher rainfall while vessel density decreased. Mean maximum vessel diameter and total leaf area in Q. ilex increased with precipitation whereas variables of Q. faginea did not show any significant trend. Results suggest that aridity, rather than minimum winter temperature, controls stem xylem responses in the studied evergreen species. Q. faginea traits did not show any response to precipitation, probably because this species develops deep roots, which in turn makes edaphic and topographic factors more important in the control of soil water availability. In response to aridity Q. coccifera only exhibits adjustment at a xylem level by reducing its water transport capacity through a reduction of vessel diameter without changing the amount of xylem tissue or foliage, whereas Q. ilex adjusts its water transport capacity in parallel to the foliage area. Received: 13 January 1997 / Accepted: 8 April 1997     

Abiotic constraints on the establishment of Quercus seedlings in grassland

High evaporative demand and periodic drought characterize the growing season in midwestern grasslands relative to deciduous forests of the eastern US, and predicted climatic changes suggest that these climatic extremes may be exacerbated. Despite this less than optimal environment for tree seedling establishment, deciduous trees have expanded into adjacent tallgrass prairie within the last century leading to a dramatic land cover change. In order to determine the role of light and temperature on seedling establishment, we assessed carbon and water relations and aboveground growth of first‐year Quercus macrocarpa seedlings exposed to one of three conditions: (1) intact tallgrass prairie communities (control), (2) aboveground herbaceous biomass removed (grass removal), and (3) shade plus biomass removal to reduce light (PFD) to levels typical of the grassland‐forest ecotone (shade). In the 2000 growing season, precipitation was 35% below the long‐term average, which had a significant negative effect on oak seedling carbon gain at midseason (photosynthesis declined to 10% of maximum rates). However, net photosynthesis and stomatal conductance in the shade treatment was ca. 2.5 and 1.5 times greater, respectively, than in control treatment seedlings during this drought. During this period, leaf and air temperatures in control seedlings were similar whereas leaf temperatures in the shade treatment remained below air temperature. A late‐season recovery period, coincident with decreased air temperatures, resulted in increased net photosynthesis for all seedlings. Increased photosynthetic rates and water relations in shaded seedlings compared to seedlings in full sun suggest that, at least in dry years, high light and temperature may negatively impact oak seedling performance. However, high survival rates for all seedlings indicate that Q. macrocarpa seedlings are capable of tolerating both present‐day and future climatic extremes. Unless historic fire regimes are restored, forest expansion and land cover change are likely to continue.     

Survival,growth, and photosynthesis of tree seedlings competing with herbaceous vegetation along a water-light-nitrogen gradient

In herbaceous dominated patches and ecosystems, tree establishment is influenced partly by the ability of woody seedlings to survive and grow in direct competition with herbaceous vegetation. We studied the importance of season long wet and dry spells on the competitive interactions between herbaceous vegetation and oak seedlings along a light and nitrogen gradient in an infertile secondary successional grassland in central North America. We conducted a field experiment in which seedlings of bur oak (Quercus macrocarpa) and northern pin oak (Q. ellipsoidalis) were exposed to two levels of light (full sun and 80% shade), three levels of nitrogen input (0, 5, 15 g m^–1 yr^–1), and three levels of water input (low, medium and high). In addition, seedlings were grown with and without the presence of surrounding herbaceous vegetation under both light and all three water levels. Seedling survival, growth, and rate of photosynthesis were significantly affected by competition with herbaceous vegetation and these effects varied along the multiple resource gradient. Overall, seedling survival of both species was significantly greater in wetter and shaded plots and when surrounding herbaceous vegetation was removed and was lower in nitrogen enriched plots. We found that soil water was significantly affected by varying inputs of water, light, and the presence or absence of herbaceous vegetation, and that seedling survival and rate of photosynthesis were highly correlated with available soil water. Our findings show that the impact of season long wet and dry spells on tree seedling success in grasslands can be affected by light and soil nitrogen availability.     

Early establishment of planted Retama sphaerocarpa seedlings under different levels of light,water and weed competition

Large amounts of former cropland are being abandoned in developedregions. To formulate guidelines for land reclamation programmes, we exploredthe effects of artificial shading, irrigation, and removal of weed competitionon the performance of Retama sphaerocarpa (L.) Boiss.seedlings in a factorial experiment located in an abandoned cropland in CentralSpain. R. sphaerocarpa is of interest for revegetationbecause it is a drought tolerant leguminous shrub that is a major structuralcomponent of the native plant community. Seedling performance was evaluated inthree ways: seedling survivorship, growth, and photochemical efficiency. Wealsomeasured soil moisture and weed biomass production and found that bothincreasedunder artificial shading conditions. Soil moisture increased very slightlywhereweeds were removed. Thus, increased transpiration from weeds outweighed reducedevaporation from soils due to shading by weeds. Artificial shading was the mosteffective treatment for seedling survivorship, followed by removal ofcompetition by weeds. After summer, 34 % of the seedlings survived in the mostfavourable conditions (artificially shaded plots where weeds were removed),compared to ca. 1 % in full-light plots with no removal of weed competition. Apositive effect of irrigation was found for growth of seedling cover and heightin shaded plots. The analysis of photochemical efficiency pointed out therelevance of weed competition removal, and confirmed the usefulness of fastfluorescence transient techniques for the quantification of seedlingperformance. The data suggest that competition between seedlings and weeds wasprimarily for water rather than for light. We conclude that i) artificialshading improved seedling performance, but this is a little practical techniquebecause of its cost; ii) as weeds compete with, rather than facilitate, plantedseedlings, weed clipping around the seedlings is a feasible technique thatwouldimprove seedling survival; and iii) seedling performance could alsoconsiderablyimprove with a higher irrigation than was used in this experiment(75lm^–2 per growth period), provided that weedsare removed.     

The effects of neighbors on the growth and survival of shrub seedlings following fire

Where plant species vie for limited resources, disturbances might preclude competition by releasing a flush of nutrients, or by reducing biomass and thereby diminishing the consumption of resources. However, if new seedlings colonize in clumps, they may still deplete resources within the local aggregations, which may then reduce their growth and survivorship. We investigated competition among seedlings in a burned area by examining the relationship between the performance of newly established shrub seedlings of Ceanothus impressus and (1) the proximity and (2) the identity of their near neighbors. We also investigated the relationship between neighbor proximity and the availability of water. Both survivorship and growth of C. impressus were positively associated with increasing distance to near neighbors, in a manner consistent with resource competition. The availability of water (as determined by pre-dawn xylem pressure potentials) tended to be greater when neighbors were farther away, providing evidence that water was a resource for which plants were competing. This conclusion is reinforced by the finding that the effects of neighbors were stronger in drought years, suggesting that yearly variation in the availability of an important resource (water) can affect the strength of competitive interactions. This suggests that after disturbances, when some resources are apparently abundant on a large scale, competition may be important in determining the small scale patterns of seedling growth and survival.     

What are the effects of substrate and grass removal on recruitment of <Emphasis Type="Italic">Acacia mellifera</Emphasis> seedlings in a semi-arid environment?

Acacia mellifera is one of the most important encroaching woody plants in southern African savannas. Previous studies found that this species encroaches far more readily on rocky areas than on sandy substrates, although it grows larger on sandy substrates. Rocky substrates are known to retain more water than sandy substrates, which may be of vital importance during recruitment in semi-arid and arid environments. A number of studies have also indicated that competition with grasses may reduce the recruitment and biomass of tree seedlings. We created an experiment in a semi-arid environment (mean annual rainfall = 388 mm) that tested for the effects of rockiness on A. mellifera recruitment. We also tested the hypothesis that grasses effectively compete with A. mellifera in this environment by simulating the effect of grazing by clipping grasses from half the plots in both the rocky and sandy treatments. Significantly more A. mellifera seedlings established in plots where grasses were clipped than in control plots. A. mellifera seedlings had greater biomass on sandy substrates than on rocky substrates. No significant interaction effects were found between substrate and grass clipping treatment for either seedling number or biomass. We conclude that A. mellifera seedlings are more likely to encroach in habitats with low grass density, although they may achieve greater biomass on sandy soils. Thus, it may be the lower grass density rather than rockiness, which increases the encroachment observed in naturally rocky habitats. These results are also consistent with our observations that adult A. mellifera trees are larger on sandy soils than on rocky soils.     

Coexistence and interference between a native perennial grass and non-native annual grasses in California

Little is known about the potential for coexistence between native and non-native plants after large-scale biological invasions. Using the example of native perennial bunchgrasses and non-native annual grasses in California grasslands, we sought to determine the effects of interference from non-native grasses on the different life stages of the native perennial bunchgrass Nassella pulchra. Further, we asked whether N. pulchra interferes with non-native annual grasses, and whether competition for water is an important component of these interspecific interactions in this water-limited system. In a series of field and greenhouse experiments employing neighbor removals and additions of water, we found that seedling recruitment of N. pulchra was strongly seed-limited. In both field and greenhouse, natural recruitment of N. pulchra seedlings from grassland soil was extremely low. In field plots where we added seeds, addition of water to field plots increased density of N. pulchra seedlings by 88% and increased total aboveground N. pulchra seedling biomass by almost 90%, suggesting that water was the primary limiting resource. In the greenhouse, simulated drought early in the growing season had a greater negative effect on the biomass of annual seedlings than on the seedlings of N. pulchra. In the field, presence of annuals reduced growth and seed production of all sizes of N. pulchra, and these effects did not decrease as N. pulchra individuals increased in size. These negative effects appeared to be due to competition for water, because N. pulchra plants showed less negative pre-dawn leaf water potentials when annual neighbors were removed. Also, simply adding water caused the same increases in aboveground biomass and seed production of N. pulchra plants as removing all annual neighbors. We found no evidence that established N. pulchra plants were able to suppress non-native annual grasses. Removing large N. pulchra individuals did not affect peak biomass per unit area of annuals. We conclude that effects of interference from non native annuals are important through all life stages of the native perennial N. pulchra. Our results suggest that persistence of native bunchgrasses may be enhanced by greater mortality of annual than perennial seedlings during drought, and possibly by reduced competition for water in wet years because of increased resource availability. Received: 12 November 1998 / Accepted: 4 August 1999     

Effects of herbivory and herb interference on oak establishment in a semi-arid temperate savanna

Abstract. Early establishment of Quercus emoryi seedlings in the grassland phase of a semi-arid temperate savanna was evaluated in southern Arizona, USA. Controlled-environment experiments indicated that Quercus seedlings are tolerant of frequent clipping, but undipped seedlings produce more biomass than clipped seedlings. Seedlings were much less tolerant of clipping or defoliation in the field than in the growth chamber, and most seedlings died after being clipped or defoliated by herbivores. Invertebrate herbivory, which is rarely mentioned as an important factor limiting seedling establishment, was the leading cause of seedling mortality in the field study. Below-ground interference from herbs had a greater impact on seedling survival and growth than above-ground interference.     

Competitive effects of the invasive shrub,Lonicera maackii (Rupr.) Herder (Caprifoliaceae), on the growth and survival of native tree seedlings

Invasive plants are often associated with reduced cover of native plants, but rarely has competition between invasives and natives been assessed experimentally. The shrub Lonicera maackii, native to northeastern Asia, has invaded forests and old fields in numerous parts of eastern North America, and is associated with reduced tree seedling density in Ohio forests. A field experiment was conducted to test the effects of established L. maackii on the survival and growth of transplanted native tree species. The experiment examined above-ground competition (by removing L. maackii shoots) and below-ground competition (by trenching around transplanted seedlings). The effects of above-ground competition with L. maackii were generally more important than below-ground competition, though both were detected. Shoot treatment was the key determinant for the survival of all species except P. serotina, whereas trenching only enhanced survival for A. saccharum caged and P. serotina, and only in the shoot removal treatment. For the surviving seedlings, L. maackii shoot removal increased growth of A. saccharum seedlings protected with cages, but actually reduced the growth of unprotected Q. rubra and A. saccharum seedlings, indicating that L. maackii shoots confer some protection from deer browsing. Significant interactions between root and shoot treatment on Q. rubra growth parameters, specifically greatest growth in the shoot present & trenched treatment, is attributed to protection from deer browsing combined with release from below-ground competition. Despite this protective function of L. maackii shoots, the overall effect of this invasive shrub is increased mortality of native tree seedlings, suggesting it impacts the natural regeneration of secondary forests.     

Summer-drought constrains the phenology and growth of two coexisting Mediterranean oaks with contrasting leaf habit: implications for their persistence and reproduction

This study analyses how coexisting evergreen and deciduous oaks adjust their phenology to cope with the stressful Mediterranean summer conditions. We test the hypothesis that the vegetative and reproductive growth of the winter deciduous (Quercus faginea Lam.) is more affected by summer drought than that of the evergreen [Quercus ilex L. subsp. ballota (Desf.) Samp.]. First, we assessed the complete aboveground phenology of both species during two consecutive years. Shoot and litter production and bud, acorn and secondary growth were monitored monthly. Second, we identified several parameters affected by summer conditions: apical bud size, individual leaf area (LA), leaf mass per area (LMA) and acorn yield in both species, and leaf-fall in Q. faginea; and analysed their variation over 10 years. Q. ilex performed up to 25% of shoot growth and most leaf development during summer, whereas Q. faginea completed most of both phenophases during spring. Secondary growth was arrested in summer under drought conditions. Approximately, 30–40% of bud and 40–50% of acorn growth was undertaken during summer in both species. Summer drought related to differences in LA, LMA and leaf senescence, but not to acorn yield. Both species had similar year-to-year patterns of acorn production, though yields were always lower in Q. faginea. Bud size decreased severely in both species during extremely dry years. In Q. ilex, bud size tended to alternate between years of large and small buds, and these patterns were followed by opposite trends in stem length. In Q. faginea, bud size was more stable through time. Q. ilex was more phenologically active during summer than Q. faginea, indicating a higher tolerance to drought. Furthermore, bud and fruit growth (the only two phenophases that both species performed during summer) were more severely affected by summer drought in Q. faginea than in the evergreen. The differential effects of summer drought on key phenophases for the persistence (bud growth) and colonization ability (fruit production) of both species may have consequences for their coexistence.     

Seedling competition between native cottonwood and exotic saltcedar: implications for restoration

Altered hydrology of southwestern United States rivers has led to a decline in native cottonwood (Populus deltoides). Areas historically dominated by cottonwood have been replaced by invasive saltcedar (Tamarix chinensis). Restoration of historic hydrology through periodic flooding of riparian areas has been a means of restoring native species. However, due to similarity in germination requirements of cottonwoods and saltcedars, flooding may create an unwanted increase in the number of saltcedar seedlings. Therefore, we evaluated competitive aspects of these co-occurring species in an extant riparian habitat in the arid southwestern US. We measured effects of competition between cottonwood and saltcedar seedlings and among cottonwood seedlings during the first growing season following seedling establishment in 360, 0.5 × 0.5-m plots at the Bosque del Apache National Wildlife Refuge, New Mexico. We used five interspecific density treatments and five intraspecific density treatments. Cottonwood seedling biomass and height were twice that of saltcedar seedlings across all density treatments. As density of cottonwood increased, intraspecific competition increased in severity and biomass of cottonwood seedlings decreased. At 4 plants/0.25 m², cottonwood seedlings had the greatest biomass; whereas, survival was highest at 10 plants/0.25 m². Our results support greenhouse studies and suggest that if favorable germination conditions are established for cottonwood in floodplains, saltcedar seedlings that cogerminate could be outcompeted by native cottonwood seedlings.