•Water stress is likely the main driver of pine mortality in the studied ecotone in Central Spain.•Higher tree sensitivity to spring precipitation and microsites with more water availability increase ...tree survival.•Drought tolerance may determine shifts in species dominance in the studied ecotone.
Drought-induced tree decline and accelerated mortality under global change threaten forests worldwide. Identifying environmental and structural factors driving tree mortality of species with different ecology is essential to understand forest dynamics. We compared three coniferous species and trees with different health status (i.e. healthy, declining and dead) on a xeric Mediterranean ecotone where Pinus pinaster shows widespread decline (defoliation, mistletoe, high mortality) while more drought-tolerant Pinus pinea and Juniperus oxycedrus do not. We analysed individual tree and stand characteristics and regeneration of all species to understand how different abiotic factors at different time scales affected stand species dynamics in relation to tree decline and mortality. Only dead and declining pine trees showed recent multiannual negative radial growth trends. The growth decline started after two extreme droughts, being 10 years longer in the declining species, P. pinaster, than in non-declining P. pinea. We observed no signs of decline or mortality in the most drought-tolerant species, J. oxycedrus. Variables related to moisture availability, like stand dominant height and the tree response to precipitation, increased survival probability in P. pinaster but not in P. pinea. Despite P. pinaster dominating the canopy, its regeneration was very scarce and less abundant than that of co-occurring more drought-tolerant species. Our results suggest that drought-induced mortality was related to species-specific sensitivity to water stress and may be inducing compositional changes towards dominance of more drought-tolerant species. Consequently, at its lower elevation limit P. pinaster might be able to persist only in microsites with higher moisture availability.
The antifeedant effect of six cacalolides and six eremophilanolides was tested against the herbivorous insects Spodoptera littoralis, Leptinotarsa decemlineata, and Myzus persicae. The test compounds ...included several natural products isolated from Senecio madagascariensis (14-isovaleryloxy-1,2-dehydrocacalol methyl ether, 4), S. barba-johannis (13-hydroxy-14-oxocacalohastine, 5; 13-acetyloxy-14-oxocacalohastine, 6) and S. toluccanus 6-hydroxyeuryopsin, 7; 1(10)-epoxy-6-hydroxyeuryopsin, 9; toluccanolide A, 11 and the derivatives cacalol methyl ether (1); cacalol acetate (2); 1-acetyloxy-2-methyloxy-1,2,3,4-tetradehydrocacalol acetate (3); 6-acetyloxyeuryopsin (8); 6-acetyloxy-1(10)-epoxyeuryopsin (10), and toluccanolide A acetate (12). Compound 11 and its derivative 12 exhibited moderate antifeedant activity against S. littoralis; 2, 7-10, and 12 showed strong activity against L. decemlineata, while the aphid M. persicae was moderately deterred in the presence of compounds 1, 4, 8, 10, and 12. The phytotoxic activity of 1-12 on Lactuca sativa was also evaluated. Compounds 2 and 4-12 moderately inhibited seed germination at 24 h, while compounds 1-4, 6, 9, and 10 had a significant inhibition effect on L. sativa radicle length (over 50%).
Most information on the ecology of oak-dominated forests in Europe comes from forests altered for centuries because remnants of old-growth forests are rare. Disturbance and recruitment regimes in ...old-growth forests provide information on forest dynamics and their effects on long-term carbon storage. In an old-growth Quercus petraea forest in northwestern Spain, we inventoried three plots and extracted cores from 166 live and dead trees across canopy classes (DBH ≥ 5 cm). We reconstructed disturbance dynamics for the last 500 years from tree-ring widths. We also reconstructed past dynamics of above ground biomass (AGB) and recent AGB accumulation rates at stand level using allometric equations. From these data, we present a new tree-ring-based approach to estimate the age of carbon stored in AGB. The oldest tree was at least 568 years, making it the oldest known precisely-dated oak to date and one of the oldest broadleaved trees in the Northern Hemisphere. All plots contained trees over 400 years old. The disturbance regime was dominated by small, frequent releases with just a few more intense disturbances that affected ≤20% of trees. Oak recruitment was variable but rather continuous for 500 years. Carbon turnover times ranged between 153 and 229 years and mean carbon ages between 108 and 167 years. Over 50% of AGB (150 Mg·ha−1) persisted ≥100 years and up to 21% of AGB (77 Mg·ha−1) ≥300 years. Low disturbance rates and low productivity maintained current canopy oak dominance. Absence of management or stand-replacing disturbances over the last 500 years resulted in high forest stability, long carbon turnover times and long mean carbon ages. Observed dynamics and the absence of shade-tolerant species suggest that oak dominance could continue in the future. Our estimations of long-term carbon storage at centennial scales in unmanaged old-growth forests highlights the importance of management and natural disturbances for the global carbon cycle.
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•Old-growth forests are essential to understand natural forest dynamics, but are rare in Europe.•Combining inventory and tree-ring analysis informs dynamics and long-term carbon storage.•Small gaps and constant recruitment dominated dynamics in an old-growth oak forest.•Carbon stored in above ground biomass for centuries with trees >400 years old.•Tree longevity and low-intensity disturbance regimes resulted in long turnover times.
We investigated the influence of climate on the ring width and xylem anatomy of two co-occurring pines (
Pinus nigra
Arn. and
P. sylvestris
L.) in the mountains of east-central Spain in order to test ...their utility for dendroclimatic reconstructions. We developed chronologies of ring width, mean lumen diameter and mean cell-wall thickness (in the earlywood, latewood, and the total annual ring) and the number of cells between 1960 and 2006. Drought, expressed as the standardized precipitation-evapotranspiration index (SPEI), was the main climatic driver of tree radial growth, although trees were also sensitive to temperature (negative effect in previous autumn and current summer) and precipitation (with a general positive effect).
P. sylvestris
response was stronger to climate of the current year, whereas the effect of previous-year climate was more important for
P. nigra
. Warm and dry summers reduced ring width, tracheid lumen, and wall thickness in both species, whereas warm winter-spring temperatures had the opposite effect, primarily for
P. sylvestris
. Previous-year or early-season conditions mainly affected earlywood features, whereas latewood was more responsive to summer climate. Overall, climate appeared to be a stronger limiting factor for
P. sylvestris
. During periods of drought, cell-wall thickness was reduced while lumen width increased in the latewood of
P. sylvestris
. This could compromise its hydraulic safety against drought-induced cavitation as our site was close to the southern and dry edge of the species distribution area. Our results suggest that anatomical variables record different and stronger climate information than ring width variables, especially in
P. sylvestris
. Reconstruction models for SPEI at the 3-month scale were developed for July–August and September–October using principal components regression. The best models included anatomical and width variables of both pine species suggesting that tracheid chronologies can be useful for drought reconstructions especially at mesic sites or with species that encode a mixed drought and temperature-precipitation signal.
• We present the longest tree-ring chronology (141 y) of
Quercus ilex
L. (holm oak), and discuss the species climate-growth relationships and the influence of stand density on tree sensitivity to ...climate.
• Similarly to
Quercus suber
L., the most influential climatic variables upon holm oak growth were late spring and early summer precipitation, which enhanced growth, and high temperatures in the previous August and current July, which negatively affected growth.
• High density stands responded to similar climatic factors as low density stands, but their response was generally weaker. Holm oak sensitivity to climate has increased in recent decades, which might be related to increasing temperatures in the region. Sensitivity was higher in low density stands. Additionally, the effect of summer stress on growth seems to have increased during the same period, similarly to other species in the Iberian Peninsula, suggesting that trees are more vulnerable to climatic changes.
• Stand density could buffer the response to climate by smoothing climatic extremes. Nevertheless, the effect of competition might reverse this positive effect at the individual tree level. Precautions should be taken before providing management guidelines regarding the effect of climate change and stand density on holm oak.
Global change potentially increases forest vulnerability. Different abiotic and biotic factors may interact to cause forest decline and accelerated tree mortality. We studied a mixed Mediterranean ...continental forest where Pinus pinaster Ait. (maritime pine) shows widespread decline to analyse the role of different abiotic and biotic factors on health status and growth dynamics both at the individual and plot levels. We also analysed stand composition and regeneration of tree species to check whether there is a change in species dominance. Fungal pathogens were seldom present and we detected no pervasive fungi or insect infestation and no presence of pathogens like Heterobasidion or Phytophthora. Infection of hemiparasite plants like Viscum album L. (mistletoe) can reduce leaf area and its abundance is generally considered an expression of host decline. Yet, the existence among declining trees of high defoliation levels without mistletoe, but not vice versa, suggests that defoliation in response to some abiotic stressor could be a predisposing factor preceding mistletoe infection. Compared to healthy trees, declining and dead trees exhibited higher defoliation rates, smaller needles and lower recent growth with steeper negative trends. Dead and declining trees showed similar negative growth trends since the early 1990s droughts, which we interpreted as early warning signals anticipating mortality of currently declining trees in the near future. Mortality of maritime pine extending across all size classes, the lower presence of this species in the smallest size classes and its lack of regeneration suggest it is potentially losing its current dominance and being replaced by other co-occurring, more drought-tolerant species. Our results unravelled that maritime pine decline seems to be mainly driven by a combination of predisposing and inciting abiotic factors (microenvironment and drought stress) and biotic factors (mistletoe). The absence of widespread fungal pathogens suggests that they may have a minor role on pine decline acting only eventually as contributing factors. Although there could be other interrelations among factors or other biotic agents at play, our results strongly suggest that water stress plays a major role in the decline process of the dominant species on an ecosystem with strong land-use legacies.
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•Pinus pinaster shows widespread decline and accelerated mortality.•Decline linked to abiotic stress including climatic legacies but not fungal pathogens.•More drought-tolerant species may substitute currently dominant P. pinaster.
Canopy disturbance events in forests often increase light availability and growth rates for surviving trees. Using ring widths, release-detection methods identify the initiation of rapid growth ...associated with these events enabling reconstructions of forest disturbance history. Conversely, dendroclimate reconstructions minimize these rapid growth responses by detrending entire ring-width time series to resolve underlying climate signals. Incorporating advantages from both of these approaches, we present a canopy disturbance detrending method that quantifies the subsequent, additional growth from release events as a discrete time series. Our method uses radial-growth averaging to detect release events and then power transformation and age-dependent smoothing splines to detrend individual release events, separating canopy disturbance responses from climatic effects. We test our canopy disturbance detrending method on both a coniferous and a broadleaf ring-width chronology from two contrasting temperate forests in eastern North America with known disturbance histories. The resulting disturbance growth time series agrees with the documented forest histories for each forest. Removing the effects of canopy disturbance from the ring-width series in each chronology improves climate correlations with monthly values of temperature and precipitation and may recover more low-frequency variation with past climate compared to other common detrending methods. Our method provides an alternative approach for detrending disturbance events in closed-canopy forests that should be accessible and useful for both ecological and climatological reconstructions.
Mature forests hold maximum live biomass stocks Molina-Valero, Juan Alberto; Camarero, J. Julio; Álvarez-González, Juan Gabriel ...
Forest ecology and management,
01/2021, Volume:
480
Journal Article
Peer reviewed
Open access
•Naturalness is the degree to which forests are driven by natural processes.•Secondary mature and old-growth forests stocked similar live biomass.•Maximum values of live carbon storage may be reached ...earlier than old-growth stage.•Structure and tree-ring data allow to assess long-term changes in forest carbon stocks.
The important role that forests play in the global carbon cycle has led to the implementation of management practices to enhance long-term carbon storage in forests. Obtaining information about how forest biomass varies at different successional stages is therefore essential. In this study, we aimed (i) to determine the relationship between stand live biomass and the degree of naturalness (i.e. the degree to which forests dynamics are driven by natural processes in the absence of anthropogenic influences) in undisturbed, mature forests, and (ii) to establish the upper threshold of live biomass stock that is dependent on site quality, by using the site form (SF) index (dominant height-diameter relationship). We assessed live biomass stocks in a total of 10 undisturbed mixed and pure mature silver fir (Abies alba Mill.) and European beech (Fagus sylvatica L.) stands across the Spanish Pyrenees. We used dendrochronology and chrono-functional indicators to assess the degree of naturalness and to detect growth releases as proxies for past disturbance. We compared data from the mature plots and data from the Spanish National Forest Inventory (NFI), for similar forest types and SF values, to determine whether maximum live biomass values were reached in the plots. We observed that live biomass stock was generally independent of naturalness. The forests characterized by the lowest degrees of naturalness held similar amounts of live tree biomass as those characterized by the highest degrees of naturalness, including old-growth forests. Although some stands had undergone severe disturbance in the past, in all but two plots the live biomass stocks reached maximum values at the respective SF values, unlike in the NFI plots. The study findings contribute to the body of evidence supporting the existence of an upper threshold of live biomass stock represented by undisturbed mature stands. Maximum live carbon stock appears to be reached at earlier stages than the old-growth stage. This finding has consequences for biomass and carbon management and may be useful for developing forest policies involving carbon sequestration aimed at mitigating climate change.
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