The Holocene history of low altitude Mediterranean Fagus sylvatica forests in southern France

Questions

As the dominant tree in many European forests, Fagus sylvatica functions as an ecosystem engineer, yet its istory remains little understood. Here we ask: (a) are there indications for its presence in southeast France during the last Glacial period; (b) what was the timing of the expansion and decline of F. sylvatica dominated forests; (c) which factors influenced their dynamics and in particular to what extent did past precipitation changes impact upon them; and (d) at which altitudes did these beech forests occur within the region?

Location

Languedoc, the French Mediterranean area.

Method

This article presents a well dated and high‐resolution pollen sequence covering the last 7,800 years from the Palavas Lagoon in the Languedoc together with a review of Fagus charcoal occurrences in the Languedoc and the lower Rhône Valley, and a review of pollen data from a compilation of 69 sites in southeast France.

Results

The Palavas pollen sequence provides a regional summary of F. sylvatica abundance changes near the Mediterranean coast. Around 6,000 years cal BP , an abrupt transition from small beech populations to well‐developed forests is recorded. The maximum development of beech forests occurred between 4,000 and 3,000 years cal BP , while F. sylvatica started to regress after 3,000 years cal BP .

Conclusion

Scattered F. sylvatica populations probably survived throughout southern France during the last Glacial period. F. sylvatica started to spread around 8,000 years cal BP while beech forests never expanded before 6,000 years cal BP . The complex patterns of F. sylvatica expansion in southern France after 6,000 years cal BP suggests that a combination of global (climate change) and local (human impact) factors were responsible for this major change. Recurrent abrupt climate changes, the aridity trend and human deforestation caused beech forests to decline after 3,000 years cal BP .

     

Interpretation of seasonal changes of xylem embolism and plant hydraulic resistance in Fagus sylvatica

The annual course of xylem embolism in twigs of adult beech trees was monitored, and compared to concurrent changes of tree water status and hydraulic resistances. Xylem embolism was quantified in 1-year-old apical twigs by the hydraulic conductivity as a percentage of the maximum measured after removal of air emboli. Tree and root hydraulic resistances were estimated from water potential differences and sap flux measurements. The considerable degree of twig embolism found in winter (up to 90% loss of hydraulic conductivity) may be attributed to the effect of freeze-thaw cycles in the xylem. A partial recovery from winter embolism occurred in spring, probably because of the production of new functional xylem. Xylem embolism fluctuated around 50% throughout the summer, without significant changes. Almost complete refilling of apical twigs was observed early in autumn. A significant negative correlation was found between xylem embolism and precipitation; thus, an active role of rainfall in embolism reversion is hypothesized. Tree and root hydraulic resistances were found to change throughout the growing period. A marked decrease of hydraulic resistance preceded the refilling of apical twigs in the autumn. Most of the decrease in total tree resistance was estimated to be located in the root compartment.     

One strategy does not fit all: determinants of urban adaptation in mammals

Urbanisation exposes wildlife to new challenging conditions and environmental pressures. Some mammalian species have adapted to these novel environments, but it remains unclear which characteristics allow them to persist. To address this question, we identified 190 mammals regularly recorded in urban settlements worldwide, and used phylogenetic path analysis to test hypotheses regarding which behavioural, ecological and life history traits favour adaptation to urban environments for different mammalian groups. Our results show that all urban mammals produce larger litters; whereas other traits such as body size, behavioural plasticity and diet diversity were important for some but not all taxonomic groups. This variation highlights the idiosyncrasies of the urban adaptation process and likely reflects the diversity of ecological niches and roles mammals can play. Our study contributes towards a better understanding of mammal association to humans, which will ultimately allow the design of wildlife‐friendly urban environments and contribute to mitigate human‐wildlife conflicts.     

Physiological performance of beech (Fagus sylvatica L.) at its southeastern distribution limit in Europe: seasonal changes in nitrogen, carbon and water balance

To assess the physiological performance of drought-sensitive European beech ( Fagus sylvatica L.) under the dry Mediterranean climate prevailing at its southeastern distribution limit in Europe, we analyzed seasonal changes in carbon, nitrogen and water balance of naturally grown adult trees. We determined the foliar C and N contents, delta13C and delta18O signatures, total soluble non-protein nitrogen compounds (TSNN) in xylem, leaves, and phloem, as well as leaf water potential and photosynthetic quantum yield in northern Greece during 2003. Tissue sampling was performed in May, July, and September, while field measurements were conducted regularly. Climatic conditions for the 2003 growing season fall within the typical range of the studied area. The N- and C-related parameters displayed distinct seasonal courses. TSNN was highest in May in all tissues, and asparagine (Asn) was then the most abundant compound. Thereafter, TSNN decreased significantly in all tissues and both its concentration and composition remained constant in July and September. In both months, glutamate (Glu) prevailed in leaves, gamma-aminobutyric acid (GABA) in phloem exudates from twigs and trunks, and arginine (Arg) in the xylem sap, where loading with amino acids was rather low during that period, amounting to only 0.8 micromol N ml-1 in September. Highest total foliar N and C contents were detected in May, and the elevated abundance of nutrients as well as an increased foliar delta13C signature at the beginning of the growing season is attributed to remobilization processes. The signatures of delta18O, quantum yield and leaf water potentials varied only slightly throughout the growing season. Although summer precipitation at the study site was considerably lower compared to what is usual for typical central European beech forests, no intensive drought responses of the physiological apparatus were detected in the studied beech trees. This suggests efficient internal regulation mechanisms, constantly ensuring a favourable physiological status under the relatively dry Mediterranean climate.     

Twentieth century changes of tree-ring δ13C at the southern range-edge of Fagus sylvatica: increasing water-use efficiency does not avoid the growth decline induced by warming at low altitudes

We aimed to gain knowledge on the changes in intrinsic water use efficiency (iWUE) in response to increasing atmospheric CO₂ concentrations and climate change over the last century. We investigated the variation in the iWUE of mature Fagus sylvatica trees located in the higher, central and lower altitudinal forest limits (HFL, CFA and LFL) of one of the southernmost sites of beech distribution in Europe, the Montseny Mountains in Catalonia (northeast Spain), during the last century by analysing the δ¹³C of their tree rings. Pre‐ and post‐maturation phases of the trees presented different trends in δ¹³C, Δ¹³C, Ci (internal CO₂ concentration), iWUE and basal area increment (BAI). Moreover, these variables showed different trends and absolute values in the LFL than in the other altitudinal sites, CFA and HFL. Our results show the existence of an age effect on δ¹³C in the CFA and HFL (values increased by ca. 1.25‰ coinciding with the BAI suppression and release phases, previous to maturation). These age‐related changes were not found in the LFL, whose beech trees arrived to maturation earlier and experienced drier conditions during the suppression phase. In the last 26 years of comparable mature trees, the increase of iWUE deduced from the Δ¹³C analyses was ca. 10% in LFL, ca. 6% in CFA and not significant in HFL. These results show that climate change towards more arid conditions accounted for these higher Δ¹³C‐values and increases in the LFL more than the continuous increase in atmospheric CO₂ concentrations. This increased iWUE in the LFL did not avoid a decline in growth in these lowest altitudes of this beech southern range‐edge as a result of warming. Furthermore, since there was no apparent change in iWUE and growth in the beech forests growing in the more standard‐adequate environments of higher altitudes in the last 26 years, the rate of sequestration of C into temperate ecosystems may not increase with increasing atmospheric CO₂ concentrations as predicted by most models based on short‐term small scale experiments.     

Water chemistry,landscape, and spatial controls of δ13C and δ15N of zooplankton taxa in boreal lakes: One size does not fit all

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