Mediterranean Tree Research Articles

The valid nomen for the tree frog (genus Hyla) of Tunisia and Eastern Algeria.

Mediterranean tree frogs, Hyla gr. meridionalis Boettger, 1874 (Anura: Hylidae) are widespread around the Western Mediterranean Basin, where they naturally occur across the Maghreb (Morocco, Algeria, Tunisia). Individuals of diverse Moroccan origins have been introduced and have expanded throughout the Iberian Peninsula, southern France and northern Italy (Liguria), but also on the Canary and Balearic archipelagos (Recuero et al. 2007; Dufresnes et al. 2019). Early molecular studies uncovered several mitochondrial lineages and suggested a major cryptic diversification within this taxon, with Tunisian and eastern Algerian (Numidia) populations carrying deeply divergent haplotypes compared with the rest of the range (Recuero et al. 2007; Stöck et al. 2008; Stöck et al. 2012). While intron markers showed little differentiation (Stöck et al. 2008; Stöck et al. 2012), genome-wide data obtained from RAD-sequencing have supported the deep split suspected from mtDNA (Dufresnes et al. 2018).

Securing timelines in the ancient Mediterranean using multiproxy annual tree-ring data

Calendar-dated tree-ring sequences offer an unparalleled resource for high-resolution paleoenvironmental reconstruction. Where such records exist for a few limited geographic regions over the last 8,000 to 12,000 years, they have proved invaluable for creating precise and accurate timelines for past human and environmental interactions. To expand such records across new geographic territory or extend data for certain regions further backward in time, new applications must be developed to secure "floating" (not yet absolutely dated) tree-ring sequences, which cannot be assigned single-calendar year dates by standard dendrochronological techniques. This study develops two approaches to this problem for a critical floating tree-ring chronology from the East Mediterranean Bronze-Iron Age. The chronology is more closely fixed in time using annually resolved patterns of 14C, modulated by cosmic radiation, between 1700 and 1480 BC. This placement is then tested using an anticorrelation between calendar-dated tree-ring growth responses to climatically effective volcanism in North American bristlecone pine and the Mediterranean trees. Examination of the newly dated Mediterranean tree-ring sequence between 1630 and 1500 BC using X-ray fluorescence revealed an unusual calcium anomaly around 1560 BC. While requiring further replication and analysis, this anomaly merits exploration as a potential marker for the eruption of Thera.

Variation in morphological and chemical traits of Mediterranean tree roots: linkage with leaf traits and soil conditions

AimsRoot functions are multiple and essential for the growth and survival of terrestrial plants. The aim of this work was to analyse the main trends in the variation of root traits, their coordination with leaf traits and their relationships with soil conditions.MethodsWe measured the variation of 27 fine root traits (five morphological, 20 chemical and two isotopic signatures) in trees of seven species of a mixed plantation in a metal-contaminated and remediated site of Southern Spain.ResultsWe found evidences supporting the existence of a root economics spectrum (RES). However, other dimensions were identified as being independent of the main RES: mainly the variation in the carbon concentration, the accumulation of trace elements associated with tolerance of metal-rich soils, and the fractionation of δ15N as a time-integrated trait of mycorrhizal-mediated nutrition. In general, roots and leaves were functionally coordinated, although most of the trace elements showed strong root-leaf discordance. The soil conditions interacted with the fine root traits in feedback processes. The ability of tree roots to accumulate trace elements and to reduce their translocation to leaves is a desirable trait for the phytoremediation of metal-contaminated soils.ConclusionsRoots are multifunctional. Understanding the variations in the root traits of trees will help us to predict both the responses of forests to global changes, including soil contamination, and the provision of soil-based ecosystem services.

Tree Growth Under Climate Change: Evidence From Xylogenesis Timings and Kinetics.

Tree growth is one of the most studied aspects of tree biology, particularly secondary growth. In the Mediterranean region, cambial activity is mostly determined by water availability. Climatic projections for the Mediterranean region predict more frequent and intense droughts, and longer periods without precipitation. To investigate tree growth under the predicted scenarios of climate change, a water manipulation experiment was conducted in a maritime pine stand (Pinus pinaster Aiton). In 2017, fifteen trees were divided into three groups: control, rain exclusion, and irrigation. Drought conditions were simulated by installing a continuous plastic sheet on the forest floor from March to September. Trees under irrigation treatment were watered twice a week in September. Cambial activity and xylem formation was monitored every 10 days from February 2017 until March 2018. Cell production was maximal around the spring equinox in all treatments. Trees under rain exclusion decreased cell production rates, xylogenesis duration, and latewood cell wall thickness. The extra irrigation in September did not produce noticeable differences in xylogenesis compared to trees in the control treatment. The synchronization of maximum cambial division rates around the vernal equinox (spring) could allow Mediterranean trees to mitigate the impact of summer drought. With the predicted increase in drought intensity and frequency, lower tree productivity, carbon sequestration, and wood biomass are expected.

Changes in soil organic matter and microbial communities after fine and coarse residues inputs from Mediterranean tree species

Trees residues decomposition is a key step in forested ecosystems. In the context of sclerophyllous Mediterranean forests, the effects of residue quality on soil organic matter and microbial communities is still poorly documented. Using a mesocosm experiment, we explored microbial responses (enzyme activities, basal respiration, molecular fingerprints and catabolic profiles) and variations in soil chemical properties (pH, CaCO3, organic C and N contents, quality of soil organic matter via 13C CP/MAS NMR) induced by coarse woody debris (W) or fine residues (i.e. twigs -leaves/needles, hereafter TL) from mono-specific or mixed stands of Pinus halepensis and Quercus ilex, P. sylvestris and Q. pubescens. After a nineteen-month incubation period, we found that the origin of residues (Pinus spp. stand, Quercus spp. stand or mixed stand) shaped the genetic structure of both fungal and bacterial communities. TL inputs led to higher soil N content and increased enzyme activities and bacterial biomass. Aromatic compounds and C to N ratio increased with W inputs, strongly shaped fungal communities and increased tyrosinase activity. Thus, the type of residue and the tree species identity (Quercus spp. vs Pinus spp.) appeared to modulate the balance between fungal and bacterial biomass and soil microbial diversity. Our results revealed that logging residues management, by modulating the type of residues remaining on forest floor, could have significant impacts on soil functioning by modifying both chemical properties (e.g. soil C/N ratio) and microbial structure and functions. These findings need to be supported by in natura studies since deciphering such mechanisms controlling C fluxes in forest soils has important consequences under the context of climate change.

Comparison of phenological traits, growth patterns, and seasonal dynamics of non-structural carbohydrate in Mediterranean tree crop species

Despite non-structural carbohydrate (NSC) importance for tree productivity and resilience, little is known about their seasonal regulations and trade-off with growth and reproduction. We characterize the seasonal dynamics of NSC in relation to the aboveground phenology and temporal growth patterns of three deciduous Mediterranean species: almond (Prunus dulcis (Mill.) D. A. Webb), walnut (Juglans regia L.) and pistachio (Pistacia vera L.). Seasonal dynamics of NSC were synchronous between wood tissues from trunk, branches and twigs. Almond had almost identical levels and patterns of NSC variation in twigs, branches and trunks whereas pistachio and walnut exhibited clear concentration differences among plant parts whereby twigs had the highest and most variable NSC concentration, followed by branches and then trunk. While phenology had a significant influence on NSC seasonal trends, there was no clear trade-off between NSC storage and growth suggesting that both were similarly strong sinks for NSC. A temporal trade-off observed at the seasonal scale was influenced by the phenology of the species. We propose that late senescing species experience C allocation trade-off at the end of the growing season because of C-limiting thermal conditions and priority allocation to storage in order to survive winter.

Water use strategy affects avoidance of ozone stress by stomatal closure in Mediterranean trees-A modelling analysis.

Both ozone (O3 ) and drought can limit carbon fixation by forest trees. To cope with drought stress, plants have isohydric or anisohydric water use strategies. Ozone enters plant tissues through stomata. Therefore, stomatal closure can be interpreted as avoidance to O3 stress. Here, we applied an optimization model of stomata involving water, CO2 , and O3 flux to test whether isohydric and anisohydric strategies may affect avoidance of O3 stress by stomatal closure in four Mediterranean tree species during drought. The data suggest that stomatal closure represents a response to avoid damage to the photosynthetic mechanisms under elevated O3 depending on plant water use strategy. Under high-O3 and well-watered conditions, isohydric species limited O3 fluxes by stomatal closure, whereas anisohydric species activated a tolerance response and did not actively close stomata. Under both O3 and drought stress, however, anisohydric species enhanced the capacity of avoidance by closing stomata to cope with the severe oxidative stress. In the late growing season, regardless of the water use strategy, the efficiency of O3 stress avoidance decreased with leaf ageing. As a result, carbon assimilation rate was decreased by O3 while stomata did not close enough to limit transpirational water losses.

Are Mediterranean trees well known? “Juniperus turbinata” (Cupressaceae), a common but misunderstood taxon / Les arbres de Méditerranée sont-ils bien connus ? « Juniperus turbinata » (Cupressaceae), un taxon commun mais incompris

Among Mediterranean trees, “ Juniperus turbinata” remains a little-known taxon. Described in 1844 (J. turbinata Guss.) it was ignored in many floras and checklists until very recently despite the fact that since the 1980s morphology and phytochemistry studies have confirmed its taxonomic individuality. The conclusions of recent molecular studies also point in this direction. The present work specifies its chorology by country or geographical sectors, allowing an updated world distribution map. Another map illustrates the distribution of the other two taxa in this group (“ J. phoenicea” and “ J. canariensis”). Aspects concerning karyology, reproductive biology and biological interactions are discussed, as well as its ecology : we thus present a summary table of the main phytosociological groups up to the level of alliance in which this taxon is characteristic or in which it just fits. Another focus concerns threats and conservation. Finally, a discussion on the complex taxonomy of red junipers (J. phoenicea sensu lato) allows us to justify our taxonomic treatment for the taxa of this group. Three subspecies (subsp.) are recognized here, a new nomenclatural combination is proposed and an identification key of infraspecific taxa is provided. Thus, Juniperus phoenicea L. subsp. turbinata (Guss.) Nyman should be considered as a remarkable circum-Mediterranean taxon often endangered or declining in its range which stretches along the Mediterranean coast from southern Portugal and Morocco to the eastern Mediterranean (large islands included), including the high continental mountains of North Africa, then ending to Middle-East in Saudi Arabia. Various pictures illustrate this juniper in different landscapes, ecological and geographical situations.

Spring-Summer Drought Induces Extremely Low Radial Growth Reactions in North-Tyrrhenian Pinus pinea L.

Abstract Climate projections predict shifts in environmental conditions, with cascade effects on forest growth dynamics. As such, Pinus pinea L., an ecologically important low-elevation Mediterranean tree, can be threatened by drought events. The occurrence of negative stem growth anomalies (“negative pointer years”, or NPY) and its relation to climatic conditions were analyzed, as well as the influence of extreme dry spells upon the species growth. NPY were temporally independent among the analyzed forest stands, likely due to local factors. We observed that NPY depended on dry and hot conditions during the spring-summer period at both sites, while differences in the NPY-climate reflected the species medium term dendroclimatological signal. Extremely dry years directly reduced stem growth rates. Water stress differentially affected growth at each site, likely reflecting local adaptation to droughts. Because of the increasing drought trend expected for the Mediterranean basin, our findings must be considered regarding the conservation and management of these forests.

Conventional method and microwave drying kinetics of Laurus nobilis leaves: effects on phenolic compounds and antioxidant activity

Abstract Laurus nobilis constitutes one of the usually studied Mediterranean trees due to its therapeutic effects, nutritional properties, and ornamental use. In the current investigation, a comparative study was conducted between two types of drying methods, a microwave-assisted drying (MWD) (from 180 to 900 W) and a conventional method (open-air and oven-drying using temperatures from 40 °C to 120 °C). Drying kinetics were carried out as a function of moisture content according to time. Total Phenolic Content (TPC) as well as reducing power of obtained products were determined. Kinetic results showed that the time required for dried bay leaves became shorter as temperature and microwave power increased; drying with the microwave was much more efficient than conventional methods (open-air and oven-drying). Antioxidants of the Laurus nobilis leaves were better conserved using open-air and microwave drying. Our results underlined a significant correlation between TPC and reducing power. Microwave drying using 300 W during 130 s was the most efficient combination that provided dried bay leaves with the highest content of phenolic compounds and antioxidant activity.

Seventeen new microsatellites for Tamarix gallica and cross-amplification in Tamarix species.

PremiseMicrosatellite markers were developed for the western Mediterranean tree Tamarix gallica (Tamaricaceae) as part of a study of its genetic diversity and structure.Methods and ResultsSeventeen microsatellite markers were developed for T. gallica, 14 of which were polymorphic. These microsatellites have di‐, tri‐, and tetranucleotide repeats with 1–13 alleles per locus and population. Levels of observed and expected heterozygosity ranged from 0.000 to 0.900 and from 0.000 to 0.863, respectively. Six microsatellites showed significant deviations from Hardy–Weinberg equilibrium in at least one population. Cross‐amplification in 19 Tamarix species showed a wide transferability to other species of the genus.ConclusionsThe 14 new polymorphic microsatellite markers will be used to assess the genetic diversity and population genetic structure of T. gallica. Additionally, the successful cross‐species amplification suggests their potential usefulness for investigating species delimitation and population genetics in the genus Tamarix.

A comprehensive, genus-level time-calibrated phylogeny of the tree flora of Mediterranean Europe and an assessment of its vulnerability

ABSTRACT We produced the first time-calibrated phylogeny of all 64 native tree genera occurring on the European side of the Mediterranean Basin. This phylogeny is based on 3 plastid DNA sequences (rbcL matK and trnH-psbA), 4 recognized fossil dates and 10 secondary calibrations. Based on the inferred topology, we then tested whether the investigated tree flora exhibits phylogenetic clustering in both life-history traits known to influence reproduction and species’ vulnerability to extinction. Our topology and the estimated dates mostly conform to published partial phylogenies and are highly congruent with the Angiosperm Phylogeny Group classification except for some minor incongruences including the still debated phylogenetic position of Magnoliids. The four strictly endemic genera of the Mediterranean Basin (Chamaerops, Phillyrea, Spartium and Tetraclinis) all showed emergence dates (11–72 Ma) long before the onset of the Mediterranean climate. We did not find any imprints of phylogenetic sorting processes on the life-history traits we studied, except for the mode of seed dispersal, which showed a clustered distribution across our topology. The presence of species at risk of potential extinction within a given genus was randomly distributed along the phylogenetic tree. Species with deficient data were significantly nested within a few of the most recently evolved angiosperm genera. Our analysis closes knowledge gaps and provides a valuable basis for studying the biogeographical and ecological processes that have generated the Mediterranean tree flora. It can also inform conservation planning strategies that aim at broadening traditional taxonomy-focused perspectives with components of evolutionary history and phylogenetic singularity.

Carbon Limitation and Drought Sensitivity at Contrasting Elevation and Competition of Abies pinsapo Forests. Does Experimental Thinning Enhance Water Supply and Carbohydrates?

Stand-level competition and local climate influence tree responses to increased drought at the regional scale. To evaluate stand density and elevation effects on tree carbon and water balances, we monitored seasonal changes in sap-flow density (SFD), gas exchange, xylem water potential, secondary growth, and non-structural carbohydrates (NSCs) in Abies pinsapo. Trees were subjected to experimental thinning within a low-elevation stand (1200 m), and carbon and water balances were compared to control plots at low and high elevation (1700 m). The hydraulic conductivity and the resistance to cavitation were also characterized, showing relatively high values and no significant differences among treatments. Trees growing at higher elevations presented the highest SFD, photosynthetic rates, and secondary growth, mainly because their growing season was extended until summer. Trees growing at low elevation reduced SFD during late spring and summer while SFD and secondary growth were significantly higher in the thinned stands. Declining NSC concentrations in needles, branches, and sapwood suggest drought-induced control of the carbon supply status. Our results might indicate potential altitudinal shifts, as better performance occurs at higher elevations, while thinning may be suitable as adaptive management to mitigate drought effects in endangered Mediterranean trees.

High-frequency seasonal variation of leaf fluorescence and reflectance in Mediterranean forest species under natural environmental conditions

The strategies of two evergreen Mediterranean tree species ( Arbutus unedo L. and Quercus ilex L.) to face annual temperature variability and extreme event responses has been monitored in continuum by leaf fluorescence and reflectance. The effect of chilling spells and the accumulation of cold days with chilling and freezing temperatures on winter photo-inhibition was discussed in terms of modulation capacity for fluorescence parameters to daily temperature and irradiance conditions. Throughout the winter Q. ilex resulted less photoinhibited than A. unedo , showing a more dynamic response in all fluorescence parameters and a higher non-photochemical quenching capacity. These characteristics effectively act to maintain a higher electron transport capacity in Q. ilex than in A. unedo .

The Carob tree: biogeography, ecology and cultural history of a neglected Mediterranean tree

The Carob tree: biogeography, ecology and cultural history of a neglected Mediterranean tree

Responses of photosynthesis and component processes to drought and temperature stress: are Mediterranean trees fit for climate change?

Global warming is raising concerns about the acclimatory capacity of trees and forests, especially in Mediterranean-type ecosystems. The sensitivity of photosynthesis to temperature is a key uncertainty for projecting the magnitude of terrestrial feedbacks on future climate change. While boreal, temperate and tropical species have been comparatively well investigated, our study provides the first comprehensive overview of the seasonal acclimatory responses of photosynthesis and its component processes to temperature in four Mediterranean climax species under natural conditions. We quantified seasonal changes in the responses of net photosynthesis (Anet), stomatal conductance (gs), mesophyllic conductance (gm) and electron-transport rate (Jcf), and investigated their sensitivity to drought and temperature stress in sunlit and shaded leaves of four Mediterranean tree species (Quercus ilex L., Pinus halepensis Mill., Arbutus unedo L. and Quercus pubescens Willd.). Sunlit leaves, but not shaded leaves, showed a pronounced seasonality in the temperature responses of Anet, gs, gm and Jcf. All four species and variables showed a remarkably dynamic and consistent acclimation of the thermal optimum (Topt), reaching peaks in summer ~29-32 °C. Changes in the shape of the response curves were, however, highly species-specific. Under severe drought, Topt of all variables were on average 22-29% lower. This was accompanied by narrower response curves above all in P. halepensis, reducing the optimal range for photosynthesis to the cooler morning or evening periods. Wider temperature-response curves and less strict stomatal control under severe drought were accompanied by wilting and drought-induced leaf shedding in Q. ilex and Q. pubescens and by additional branch dieback in A. unedo. Mild winter conditions led to a high Topt (~19.1-22.2 °C), benefitting the evergreen species, especially P. halepensis. Seasonal acclimation of Anet was explained better by gs and gm being less pronounced in Jcf. Drought was thus a key factor, in addition to growth temperature, to explain seasonal acclimation of photosynthesis. Severe drought periods may exceed more frequently the high acclimatory capacity of Mediterranean trees to high ambient temperatures, which could lead to reduced growth, increased leaf shedding and, for some species such as A. unedo, increased mortality risk.

Tree-rings in Mediterranean pines – can we ascribe them to calendar years?

A tree ring is defined as a band of xylem-cell layers produced in one year. In Mediterranean trees cambium often produces several bands of alternating early- and latewood during one calendar year, resulting in Intra-Annual Density Fluctuations (IADFs). We present some characteristics of growth layers in Pinus pinea (stone pine) growing on Vesuvius Mountain (near Naples, Italy). The trees showed uninterrupted wood production from January 2015 till the end of January 2016. The ring formed in this period contained five different layers of wood types. Therefore, the wood formed between January and March 2015 would be falsely ascribed to latewood of the tree ring 2014, and the wood formed between March 2015 and the end of January 2016 would be classified as tree ring 2015. We also showed differences in cambium activity in late autumn between years: in November 2015 and 2018 the cambium was still active and not active anymore, respectively. Furthermore, we present the structure of the wood formed in the year 2017 affected by a wildfire in July. Such cases where the growth layer cannot be exactly ascribed to a calendar year should be considered in dendrochronological studies of Pinus pinea and other Mediterranean species.

N : P stoichiometry and habitat effects on Mediterranean savanna seasonal root dynamics

Abstract. Mediterranean grasslands are highly seasonal and co-limited by water and nutrients. In such systems, little is known about root dynamics which may depend on individual plant properties and environment as well as seasonal water shortages and site fertility. Patterns of root biomass and activity are affected by the presence of scattered trees, grazing, site management, and chronic nitrogen deposition, all of which can affect nutrient ratios and potentially cause development of nitrogen : phosphorus (N : P) imbalances in ecosystem stoichiometry. In this study we combined observations from minirhizotrons with root measurements from direct soil cores and ingrowth cores, along with measures of above-ground biomass to investigate seasonal root dynamics and root : shoot ratios in a Mediterranean tree–grass “savanna”. We investigated responses to soil fertility, using nutrient manipulation (N∕NP addition) and spatial microhabitat treatments between open-pasture and microhabitats under the tree canopy. Root dynamics over time were also compared with indices of above-ground growth drawn from proximal remote sensing. Results show distinct differences in root dynamics and biomass between treatments and microhabitats. Root biomass was higher with N additions, but did not differ from the control with NP additions in early spring. By the end of the growing season root biomass had increased with NP in open pastures but not higher than N added alone. In contrast, root length density (RLD) in pastures responded stronger to the NP than N-only addition, while beneath trees root biomass tended to be higher with only N. Even though root biomass increased, the root : shoot ratio decreased under nutrient treatments. Timing of root and shoot growth was reasonably well paired, although in autumn root growth appeared to be substantially slower than “regreening” of the system. We interpret these differences as a shift in community structure and/or root traits under changing stoichiometry induced by the fertilization. We also consider seasonal (phenology) differences in the strength and direction of effects observed.

Root Biomechanical Traits in a Montane Mediterranean Forest Watershed: Variations with Species Diversity and Soil Depth

Plant roots play a key role in stabilizing slopes, particularly in the Mediterranean region, characterized by rough and unstable terrain. However, forest species differ in their stabilizing capacities. The purpose of this study is to fill the gap of knowledge on root biomechanical properties of relevant Mediterranean trees and shrubs in relation to slope stability. Root specimens of typical montane Mediterranean tree and shrub species were sampled in Southern Italy. Root characteristics, such as tensile strength (Tr) and root area ratio (RAR), were assessed from live roots sampled in trenches, while root cohesion was calculated. Power law functions yielded the best fit for the relationship of Tr versus root diameter; however, no significant relationship was found between root strength and root moisture content. RAR varied amongst different tree and shrub species. Roots of Quercus cerris L. were the most resistant to breaking under tension, while roots of Ilex aquifolium L. had the highest tensile strength among all shrub species. Results provide quantitative information on the role of root systems of montane Mediterranean forest species in stabilizing soils and will improve modeling of landslide susceptibility to the prevention and mitigation of natural hazards in mountain environments.

Effect of monospecific and mixed Mediterranean tree plantations on soil microbial community and biochemical functioning

Soil microbial communities (SMCs) are important drivers of forest ecosystem functions and services. To optimize afforestation practices, it is important to understand how pure and mixed plantations can affect SMCs and their functioning. Therefore, the objective of this work was to investigate the SMC structure and function of six Mediterranean woody species grown as monocultures, and of six selected 1:1 combinations. SMC structure was assessed through the abundance estimation of different culturable heterotrophic microbial populations (i.e. bacteria, fungi, actinomycetes, Pseudomoas spp. and aerobic spore-forming bacteria) and the Biolog community level physiological profile, while soil biochemical functioning was estimated by quantifying soil dehydrogenase (DHG) activity.Species richness (SR) did not increase the abundance of culturable heterotrophic microbial populations, nor the number of carbon sources metabolized by the microbial communities (Biolog richness) or DHG activity. However, SR significantly decreased the potential metabolic activity (Biolog AWCD) of soil microbial populations. Leaf type significantly affected soil microbial and biochemical features, particularly for monocultures where conifer plots were characterized by a significantly higher fungal/bacterial ratio, reduced AWCD and richness as well as reduced DHG values. Significant interactions between tree species in mixed plots were noticed as well as significant correlations between plant community weighted mean (CWM) traits and soil microbial and biochemical features. Both SMC structure and function were clearly influenced by tree plantations and, importantly, microbial populations of monocultures and mixed plantations, and their functioning, were governed by different CWM traits.