Windstorms are natural factors in vegetation dynamics, but their intensity is increasing and undermines the forest resilience. Post-event interventions depend on forest types and management purposes. To promote mixed stands that are less susceptible to windthrow than monocultures, natural tree regeneration is usually recommended. However, wild ungulates attracted by the increased food supply in the new clearings can influence the secondary succession and slow down vegetation recovery. The impact of ungulate pressure on secondary successions in blowdown areas is still poorly known, especially in Mediterranean areas, which is particularly vulnerable to climate stressors. We investigated this topic using a monospecific silver fir (Abies alba) forest of artificial origin in the Apennines as a model system. The forest was left to natural succession after a severe windthrow in 2015, offering the opportunity to establish an ungulate exclosure experiment to analyse vegetation changes over six years. For the whole plant community, cover and height, alpha-diversity (species richness, Shannon and evenness indexes), and species composition were recorded yearly, together with tree seedling density, in open and fenced plots at increasing distances from the forest edge. Ecological and functional traits (proportion of light-demanding and forest specialists, endo-/epi-zoochorous species), and life-forms of the community samples were also analysed. Overall, we found that ungulates significantly influenced the dynamic trajectories of secondary succession. The effects on plant cover and diversity were negative and increased with distance from the forest edge. The presence of ungulates favoured generalist species and endozoochorous taxa. However, distance from the forest edge strongly reduced the latter effect. The average density of tree seedlings was overall high (ca. 10,000/ha), but it was strongly reduced by ungulates for three deciduous species and A. alba, the most browsed species. Natural recolonization after windthrow in Apennine pure silver fir stands may favour the formation of mixed, less susceptible forests, but the current ungulate pressure slows down this process, especially in the areas furthest from the forest edge.
Nursery cultivation practices can be modified to increase resistance to water stress in forest seedlings following field establishment, which may be increasingly important under climate change. We evaluated the morphological (survival, growth) and physiological (chlorophyll fluorescence, leaf water potential) responses to water stress for three ecologically diverse Quercus species (Q. robur, Q. pubescens, and Q. ilex) with varying traits resulting from the combination of growing media (peat, coir) and fertilization (standard, P-enriched, K-enriched). For all species under water stress, seedlings grown in coir had generally higher growth than those grown in peat. Seedlings fertilized with P performed better, particularly for survival; conversely, K fertilization resulted in inconsistent findings. Such results could be explained by a combination of factors. P fertilization resulted in higher P accumulation in seedlings, while no K accumulation was observed in K fertilized seedlings. As expected, the more drought-sensitive species, Q. robur, showed the worst response, while Q. pubescens had a drought resistance equal or better to Q. ilex despite being classified as intermediate in drought resistance in Mediterranean environments.
A user-friendly and affordable broad-band digital Near Infrared (NIR) camera (Canon PowerShot S110 NIR) was compared with a narrow-band reflectance spectrometer (USB2000, Ocean Optics) at leaf scale for monitoring changes in response to drought of three ecologically contrasting Quercus species (Q. robur, Q. pubescens, and Q. ilex). We aimed to (a) compare vegetation indices (VIs; that is: NDVI, Normalized Difference Vegetation Index; GNDVI, Green NDVI and NIRv, near-infrared reflectance of vegetation) retrieved by NIR-camera and spectrometer in order to test the reliability of a simple, low-cost, and rapid setup for widespread field applications; (b) to assess if NIR-camera VIs might be used to quantify water stress in oak seedlings; and (c) to track changes in leaf chlorophyll content. The study was carried out during a water stress test on 1-year-old seedlings in a greenhouse. The camera detected plant status in response to drought with results highly comparable to the visible/NIR (VIS/NIR) spectrometer (by calibration and standard geometry). Consistency between VIs and morpho-physiological traits was higher in Q. robur, the most drought-sensitive among the three species. Chlorophyll content was estimated with a high goodness-of-fit by VIs or reflectance bands in the visible range. Overall, NDVI performed better than GNDVI and NIRv, and VIs performed better than single bands. Looking forward, NIR-camera VIs are adequate for the early monitoring of drought stress in oak seedlings (or small trees) in the post-planting phase or in nursery settings, thus offering a new, reliable alternative for when costs are crucial, such as in the context of restoration programs.
Understorey vegetation is a key component of plant biodiversity in forests. Events of extreme drought are a potential major driver of change of this layer. Although these are predicted to increase in frequency and duration due to climate change, their impacts on understorey communities are still unknown. In this paper, we resurveyed a set of 36 plots in mature thermophilous deciduous forests eight months after the end of an extremely dry period in 2016-2017. The plots are part of a European network representing a gradient of overstorey richness from 1- to 5-tree species mixtures, with variable composition. We took advantage of this model system to test the hypothesis that species richness and identity of dominant trees in the overstorey may modulate the impact of drought on understorey cover, diversity and composition. In spring 2018, a ca. 50% reduction of understorey cover occurred in both mono-specific and mixed plots. Species evenness and Shannon alpha-diversity increased in the monospecific plots, due to a reduction of abundance of tree species at juvenile stage and the appearance of non-forest specialists that did not occur in the mixtures. Overstorey species identity effects on understorey diversity were negligible. Overall compositional changes were moderate, though larger in the mixed plots and in those with the less drought-adapted tree species in the overstorey. Ellenberg ecological profile of the community did not change significantly but light values tended to increase in the monospecific plots, contrary to the mixtures. Overall, our findings provide circumstantial support to a higher diversity stability in the understorey of mixed plots shortly after drought, despite compositional variations. More studies and monitoring programs are needed to assess understorey recovery capacity and resilience after such events in the long-term.
Date palms are highly economically important species in hot arid regions, which may suffer ozone (O-3) pollution equivalently to heat and water stress. However, little is known about date palm sensitivity to O-3. Therefore, to identify their resistance mechanisms against elevated O-3, physiological parameters (leaf gas exchange, chlorophyll fluorescence and leaf pigments) and biomass growth responses to realistic O-3 exposure were tested in an isoprene-emitting date palm (Phoenix dactylifera L. cv. Nabut Saif) by a Free-Air Controlled Exposure (FACE) facility with three levels of O-3 (ambient [AA, 45 ppb as 24-h average], 1.5 x AA and 2 x AA). We found a reduction of photosynthesis only at 2 x AA although some foliar traits known as early indicators of O-3 stress responded already at 1.5 x AA, such as increased dark respiration, reduced leaf pigment content, reduced maximum quantum yield of PSII, inactivation of the oxygen evolving complex of PSII and reduced performance index PITOT. As a result, O-3 did not affect most of the growth parameters although significant declines of root biomass occurred only at 2 x AA. The major mechanism in date palm for reducing the severity of O-3 impacts was a restriction of stomatal O-3 uptake due to low stomatal conductance and O-3-induced stomatal closure. In addition, an increased respiration in elevated O-3 may indicate an enhanced capacity of catabolizing metabolites for detoxification and repair. Interestingly, date palm produced low amounts of monoterpenes, whose emission was stimulated in 2 x AA, although isoprene emission declined at both 1.5 and 2 x AA. Our results warrant more research on a biological significance of terpenoids in plant resistance against O-3 stress.
Peat, a non-sustainable resource, is still predominately used in forest nurseries. Coconut coir might provide an alternative, renewable, and reliable growing media but few studies have evaluated this media type in forest nurseries. We assessed the influence of pure coir, in combination with various fertilization regimes, on the growth and physiology of three ecologically diverse Quercus species seedlings (Q. robur, Q. pubescens, and Q. ilex) during nursery cultivation. Seedlings were grown using peat and pure coir in combination with three fertilization treatments (standard, K-enriched, and P-enriched). Data were collected for: (1) growth and physiological traits; (2) detailed above- and below-ground morphological traits by destructive analysis; and (3) NPK content in leaves, shoot and roots, and in the growing media, following cultivation. Peat and coir in combination with the various fertilization treatments affected above- and below-ground morphology and, to a lesser extent, the physiological traits of Quercus seedlings. Large effects of the substrate occurred for most morphological variables, with peat being more effective than coir in all studied species. Fertilization also produced significant differences. The effect of K-enriched fertilization on plant growth was clear across the three species and the two growing media. P-enriched fertilization in peat was the only combination that promoted a higher amount of this element in the tissues at the end of cultivation. Despite their smaller size, seedlings produced in coir were compatible with standard Quercus forest stocktype size, and showed a proportionally higher root system development and fibrosity. Our results suggest that coir can be used as an alternative substrate to grow Quercus species seedlings, and that fertilization can offset coir deficiencies in chemical properties. As several functional traits drive planting performance under varying environmental conditions. according to the Target Plant Concept, coir might thus serve as an acceptable material for seedling cultivation in some cases.
This study investigated visible foliar ozone (O-3) injury in three deciduous tree species with different growth patterns (indeterminate, Alnus glutinosa (L.) Gaertn.; intermediate, Sorbus aucuparia L.; and determinate, Vaccinium myrtillus L.) from May to August 2018. Ozone effects on the timing of injury onset and a plant injury index (PII) were investigated using two O-3 indices, i.e., AOT40 (accumulative O-3 exposure over 40 ppb during daylight hours) and PODY (phytotoxic O-3 dose above a flux threshold of Y nmol m(-2) s(-1)). A new parameterization for PODY estimation was developed for each species. Measurements were carried out in an O-3 free-air controlled exposure (FACE) experiment with three levels of O-3 treatment (ambient, AA; 1.5 x AA; and 2.0 x AA). Injury onset was found in May at 2.0 x AA in all three species and the timing of the onset was determined by the amount of stomatal O-3 uptake. It required 4.0 mmol m(-2) POD() and 5.5 to 9.0 ppm.h AOT40. As a result, A. glutinosa with high stomatal conductance (g(s)) showed the earliest emergence of O-3 visible injury among the three species. After the onset, O-3 visible injury expanded to the plant level as confirmed by increased PII values. In A. glutinosa with indeterminate growth pattern, a new leaf formation alleviated the expansion of O-3 visible injury at the plant level. V myrtillus showed a dramatic increase of PII from June to July due to higher sensitivity to O-3 in its flowering and fruiting stage. Ozone impacts on PII were better explained by the flux-based index, PODY, as compared with the exposure-based index, AOT40. The critical levels (CLs) corresponding to PII = 5 were 8.1 mmol Mm(-2) POD7 in A. glutinosa, 22 mmol m(-2) POD0 in S. aucuparia, and 5.8 mmol m(-2) POD1 in V myrtillus. The results highlight that the CLs for PII are species-specific. Establishing species-specific O-3 flux-effect relationships should be key for a quantitative O-3 risk assessment.
The effect of salinity on seed germination/emergence in narrow-leaved ash (Fraxinus angustifolia) was studied both under field and laboratory conditions, in order to detect critical values to NaCl exposure. Research Highlights: Novel statistical methods in germination ecology has been applied (i) to determine the effects of chilling length and salinity (up to 150 mM NaCl) on Fraxinus angustifolia subsp. oxycarpa seed emergence, and (ii) to estimate threshold limits treating germination response to salinity as a biomarker. Background and Objectives: Salinity cut values at germination stage had relevant interest for conservation and restoration aims of Mediterranean floodplain forests in coastal areas subjected to salt spray exposure and/or saline water introgression. Results: Salinity linearly decreased germination/emergence both in the field and laboratory tests. Absence of germination was observed at 60 mM NaCl in the field (70–84 mM NaCl depending on interpolation model) and at 150 mM NaCl for 4-week (but not for 24-week) chilling. At 50 mM NaCl, germination percentage was 50% (or 80%) of control for 4-week (or 24-week) chilling. Critical values for salinity were estimated between freshwater and 50 (75) mM NaCl for 4-week (24-week) chilling by Bayesian analysis. After 7-week freshwater recovery, critical cut-off values included all tested salinity levels up to 150 mM NaCl, indicating a marked resumption of seedling emergence. Conclusions: Fraxinus angustifolia is able to germinate at low salinity and to tolerate temporarily moderate salinity conditions for about two months. Prolonged chilling widened salinity tolerance.
Maritime pine (Mims pinaster) stands in Italy have recently been invaded by maritime pine bast scale (Matsucoccus feytaudi), resulting in widespread tree mortality. Clearcutting has been applied to progressively remove infected maritime pine, concurrent with planting of holm oak (Quercus ilex) seedlings. Establishment of holm oak regeneration has, however, been severely constrained by fallow deer Mama dame) browsing pressure. We hypothesized that the ability of planted holm oak to grow above the browse line was closely correlated with protective cover provided by natural regeneration. This study was carried out in a coastal protected area in central Italy, where overall performance of planted holm oak was assessed under browsing pressure in relation to the relative distance of seedlings to surrounding vegetation. To interpret the results, four categories of holm oak were identified: (i) dead plants, (ii) sprouts from stump, (iii) repeatedly browsed plants without growth, and (iv) plants that were able to develop at least one tall shoot safe from browsing. The occurrence of the latter was positively related to increasing canopy cover and plant height of surrounding vegetation, especially maritime pine. Density, height, canopy cover, and crown radius of maritime pine, as well as its relative distance from holm oak all promoted the survival and growth of target plants. Complete protection around holm oak was not required for facilitation, but the presence of crown and branches occupying the space above the target plants was crucial. Maritime pine saplings primarily served as a mechanical barrier to browsing and thereby favored the development of holm oak shoots. Site-specific, long-term monitoring studies on competition from and facilitation by surrounding vegetation are required, before implementing under-planting strategies.