Tree dieback, woody plant diversity, and ecosystem driven by topography in semi-arid mountain forests: Implication for ecosystem management.

Mountain landscapes are highly heterogeneous due to topography, notably positions along slope and slope shapes, which control ecosystem mechanisms. We hypothesized that tree dieback is controlled by topography, selecting productive and less diverse communities in lower slopes, and stress-resistant and more diverse communities on upper slopes. Understanding how this heterogeneity drives vegetation patterns should provide benchmarks for ecosystem management of mountain forest dominated by Quercus brantii. Woody communities were sampled along convex vs concave topography (i.e., ridge vs talweg), and with measurements of tree dieback severity, environmental variables (litter depth, soil quality, rock outcrop), stand structure (canopy cover, mistletoe infestation, tree diameter and height, diameter and height differentiations, oaks' number from sprout-clumps or seed-origin), and biodiversity. Slope position was the most significant driver that affected all variables, excepted evenness. Dieback severity was higher on slope shoulders and summits, and lower in lower slopes where trees were the most productive: taller, larger, more homogeneous, and mostly seed-origin. Catena shape affected the diversity and dieback severity, both higher in talwegs, but had no effect on environmental variables and little on stand structure. Outputs indicate that the higher diversity of woody plants is on upper slopes supporting stress-resistant community associated with more severe dieback and mistletoe infection probably because frugivore birds attracted by the shrubs' fruits. Semi-arid forest management must consider the shaped-slope ecosystem heterogeneity by preserving ridges that are more susceptible to tree dieback, and naturally support biodiversity. Restoration measures on lower fertile slopes could be carried out by oak planting or seedlings under the cover of shrubs to counter dieback effects and environmental stresses. In addition, forestry measures can be taken in lower positions for the conversion of coppice to high oak forest to potentially consider a moderate forestry.

Decrease in Soil Functionalities and Herbs' Diversity, but Not That of Arbuscular Mycorrhizal Fungi, Linked to Short Fire Interval in Semi-Arid Oak Forest Ecosystem, West Iran.

The semi-arid forest ecosystems of western Iran dominated by Quercus brantii are often disturbed by wildfires. Here, we assessed the effects of short fire intervals on the soil properties and community diversity of herbaceous plants and arbuscular mycorrhizal fungi (AMF), as well as the interactions between these ecosystem features. Plots burned once or twice within 10 years were compared to unburned plots over a long time period (control sites). Soil physical properties were not affected by the short fire interval, except bulk density, which increased. Soil geochemical and biological properties were affected by the fires. Soil organic matter and nitrogen concentrations were depleted by two fires. Short intervals impaired microbial respiration, microbial biomass carbon, substrate-induced respiration, and urease enzyme activity. The successive fires affected the AMF's Shannon diversity. The diversity of the herb community increased after one fire and dropped after two, indicating that the whole community structure was altered. Two fires had greater direct than indirect effects on plant and fungal diversity, as well as soil properties. Short-interval fires depleted soil functional properties and reduced herb diversity. With short-interval fires probably fostered by anthropogenic climate change, the functionalities of this semi-arid oak forest could collapse, necessitating fire mitigation.

Long-Term Steady-State Dry Boreal Forest in the Face of Disturbance.

We used bioproxies from paleosoils buried within two aeolian dunes to test hypotheses concerning the origin of dry sandy boreal forests in Canada. These forests are dominated today by Pinus banksiana Lamb. One hypothesis is that too frequent Holocene stand-replacing fires would have transformed the original vegetation through extirpation of susceptible species to fire in water stress habitat. Alternatively, the ecosystem would have not changed since the dunes stabilized enough to support forest establishment. The vegetation composition and richness were determined by identification of charcoal and macroremains and radiocarbon dating for the chronology. Both sites revealed a similar history covering 6400 years. Half of the charcoal layers were less than 2500 years old in both sites, indicating that they had been subjected to the same fire history. Data indicated a stable plant composition and richness, although the percentage of Pinus decreased slightly over 4000 years (decreasing rate 1% per century). The fungus Cenococcum geophilum was consistently present, with a stochastic abundance. The vegetation grew under natural fire conditions and soil dryness since 6000 years. The ecosystem was probably not stressed by late-Holocene fires or climate changes, as the multi-millennial steady state reveals a resistant and resilient ecosystem.

Geographic isolation and climatic variability contribute to genetic differentiation in fragmented populations of the long-lived subalpine conifer Pinus cembra L. in the western Alps.

BACKGROUND: Genetic processes shape the modern-day distribution of genetic variation within and between populations and can provide important insights into the underlying mechanisms of evolution. The resulting genetic variation is often unequally partitioned within species' distribution range and especially large differences can manifest at the range limit, where population fragmentation and isolation play a crucial role in species survival. Despite several molecular studies investigating the genetic diversity and differentiation of European Alpine mountain forests, the climatic and demographic constrains which influence the genetic processes are often unknown. Here, we apply non-coding microsatellite markers to evaluate the sporadic peripheral and continuous populations of cembra pine (Pinus cembra L.), a long-lived conifer species that inhabits the subalpine treeline ecotone in the western Alps to investigate how the genetic processes contribute to the modern-day spatial distribution. Moreover, we corroborate our findings with paleoecological records, micro and macro-remains, to infer the species' possible glacial refugia and expansion scenarios. RESULTS: Four genetically distinct groups were identified, with Bayesian and FST based approaches, across the range of the species, situated in the northern, inner and south-western Alps. We found that genetic differentiation is substantially higher in marginal populations than at the center of the range, and marginal stands are characterized by geographic and genetic isolation due to spatial segregation and restricted gene flow. Moreover, multiple matrix regression approaches revealed effects of climatic heterogeneity in species' spatial genetic pattern. Also, population stability tests indicated that all populations had experienced a severe historical bottleneck, no heterozygosity excess was detected, suggesting that more recently population sizes have remained relatively stable. CONCLUSIONS: Our study demonstrated that cembra pine might have survived in multiple glacial refugia and subsequently recolonized the Alps by different routes. Modern-day marginal populations, at the edge of the species' range, could maintain stable sizes over long periods without inbreeding depression and preserve high amounts of genetic variation. Moreover, our analyses indicate that climatic variability has played a major role in shaping differentiation, in addition to past historical events such as migration and demographic changes.

The climate, the fuel and the land use: Long-term regional variability of biomass burning in boreal forests.

The influence of different drivers on changes in North American and European boreal forests biomass burning (BB) during the Holocene was investigated based on the following hypotheses: land use was important only in the southernmost regions, while elsewhere climate was the main driver modulated by changes in fuel type. BB was reconstructed by means of 88 sedimentary charcoal records divided into six different site clusters. A statistical approach was used to explore the relative contribution of (a) pollen-based mean July/summer temperature and mean annual precipitation reconstructions, (b) an independent model-based scenario of past land use (LU), and (c) pollen-based reconstructions of plant functional types (PFTs) on BB. Our hypotheses were tested with: (a) a west-east northern boreal sector with changing climatic conditions and a homogeneous vegetation, and (b) a north-south European boreal sector characterized by gradual variation in both climate and vegetation composition. The processes driving BB in boreal forests varied from one region to another during the Holocene. However, general trends in boreal biomass burning were primarily controlled by changes in climate (mean annual precipitation in Alaska, northern Quebec, and northern Fennoscandia, and mean July/summer temperature in central Canada and central Fennoscandia) and, secondarily, by fuel composition (BB positively correlated with the presence of boreal needleleaf evergreen trees in Alaska and in central and southern Fennoscandia). Land use played only a marginal role. A modification towards less flammable tree species (by promoting deciduous stands over fire-prone conifers) could contribute to reduce circumboreal wildfire risk in future warmer periods.

Keep your feet warm? A cryptic refugium of trees linked to a geothermal spring in an ocean of glaciers.

Up to now, the most widely accepted idea of the periglacial environment is that of treeless ecosystems such as the arctic or the alpine tundra, also called the tabula rasa paradigm. However, several palaeoecological studies have recently challenged this idea, that is, treeless environments in periglacial areas where all organisms would have been exterminated near the glacier formed during the Last Glacial Maximum, notably in the Scandinavian mountains. In the Alps, the issue of glacial refugia of trees remains unanswered. Advances in glacier reconstructions show that ice domes did not cover all upper massifs, but glaciers filled valleys. Here, we used fossils of plant and malacofauna from a travertine formation located in a high mountain region to demonstrate that trees (Pinus, Betula) grew with grasses during the Lateglacial-Holocene transition, while the glacier fronts were 200-300 m lower. The geothermal travertine started to accumulate more than 14,500 years ago, but became progressively more meteogene about 11,500 years ago due to a change in groundwater circulation. With trees, land snails (gastropods) associated to woody or open habitats and aquatic mollusc were also present at the onset of the current interglacial, namely the Holocene. The geothermal spring, due to warm water and soil, probably favoured woody glacial ecosystems. This new finding of early tree growth, combined with other scattered proofs of the tree presence before 11,000 years ago in the western Alps, changes our view of the tree distribution in periglacial environments, supporting the notion of tree refugia on nunataks in an ocean of glaciers. Therefore, the tabula rasa paradigm must be revisited because it has important consequences on the global changes, including postglacial plant migrations and biogeochemical cycles.

Fire ecology of a tree glacial refugium on a nunatak with a view on Alpine glaciers.

In paleoecology, the function of biomass as a fire driver has become a focus of attention in cold ecosystems, and concerns have been raised about climate in this context. Little is known about the fire frequency and fire-plant relationships during glaciation when woodlands were limited and the climate was cold. Fire history and tree biomass were reconstructed from sedimentary charcoal and macroremains, respectively, archived in lake sediments from the western Alps. Two nunataks were investigated, both with lacustrine sediments covering the last 21 000 yr at least. During the Last Glacial Maximum (LGM) and the Lateglacial, fires occurred only on the nunatak sheltering woody plants. Cembra pine (Pinus cembra) and larch (Larix decidua) survived above glaciers during the LGM, thus evidencing a biological refugium and supporting the nunatak theory. We highlighted a long-term relationship between fires and dominant trees over the last 21 000 yr, where fire frequencies track the global climate and the local changes in tree biomass. Glacial climate (dry, cold) does not rule out fires. Fuel load and composition were significant fire drivers, with cembra pine dominating during colder periods with rare fires, and larch during the warmer Holocene with frequent fires. These findings increase knowledge of fire ecology in cold environments, and open perspectives in tree population genetics by considering new areas of tree glacial refugia in Europe.

Range-wide genetic structure of maritime pine predates the last glacial maximum: evidence from nuclear DNA.

Using nuclear simple sequence repeats (nuSSRs), we determined the genetic variability in the natural distribution range of maritime pine (Pinus pinaster) in the western Mediterranean region. We analysed the role of global and significant climatic fluctuations in driving the evolutionary diversification of this species. We attempted to determine the impact of the last glacial maximum (LGM) and human activity on genetic variation and to identify the effect of bottlenecks, admixing, migration, time to the most recent common ancestor (TMRCA), and recent splits. A total of 972 individuals were analysed. The sample represented 27 natural populations from the western Mediterranean region, which encompasses most of the natural range of P. pinaster. Using eight nuSSRs, we analysed genetic diversity indices for each population and group of populations. We also examined the interpopulation structure by the frequency and distance method and investigated genetic barriers, signals of historical demographic fluctuations, phylogeographic structure, admixing, rate of mutation, migration, as well as testing the hypothesis of isolation by distance (IBD). Both cluster analyses showed similar population genetic structure with three genetic barriers that divided the samples into four large groups. Intensive migration was only detected during the period of the last glacial maximum (LGM), which permitted the mutation rate of the markers used to be calculated. The majority of the population was found to exhibit signs of a recent bottleneck and its timing showed a clear northeast-southwest geographic distribution. A clearly defined phylogeographic structure (Nst > Gst and Rst > Gst ) under IBD was established, and showed the highest divergence between groups of populations separated by physical barriers, such as the Strait of Gibraltar, the Mediterranean Sea and the Pyrenees. The high level of intergroup genetic differentiation (ΦIS = 20.26) was attributed to a long historical isolation (which occurred before the last 18 000 years) between the principal maritime pine population groups that occurred due to physical barriers that limited pollen and seed transfer, combined with a minimal effective radius of distribution. The low level of genetic diversity among the populations was combined with genetic drift and a recent bottleneck during the period of human activity. Significant migration across barriers was due to spontaneous phenomena during the LGM, which had no significant impact on the genetic structure owing to its relatively short duration and the fragmented species. The phylogeographic structure under the assumption of IBD was well established for P. pinaster in each of the principal population groups.

Bark flammability as a fire-response trait for subalpine trees.

Relationships between the flammability properties of a given plant and its chances of survival after a fire still remain unknown. We hypothesize that the bark flammability of a tree reduces the potential for tree survival following surface fires, and that if tree resistance to fire is provided by a thick insulating bark, the latter must be few flammable. We test, on subalpine tree species, the relationship between the flammability of bark and its insulating ability, identifies the biological traits that determine bark flammability, and assesses their relative susceptibility to surface fires from their bark properties. The experimental set of burning properties was analyzed by Principal Component Analysis to assess the bark flammability. Bark insulating ability was expressed by the critical time to cambium kill computed from bark thickness. Log-linear regressions indicated that bark flammability varies with the bark thickness and the density of wood under bark and that the most flammable barks have poor insulating ability. Susceptibility to surface fires increases from gymnosperm to angiosperm subalpine trees. The co-dominant subalpine species Larix decidua (Mill.) and Pinus cembra (L.) exhibit large differences in both flammability and insulating ability of the bark that should partly explain their contrasted responses to fires in the past.

Assessing paleo-biodiversity using low proxy influx.

We developed an algorithm to improve richness assessment based on paleoecological series, considering sample features such as their temporal resolutions or their volumes. Our new method can be applied to both high- and low-count size proxies, i.e. pollen and plant macroremain records, respectively. While pollen generally abounds in sediments, plant macroremains are generally rare, thus leading to difficulties to compute paleo-biodiversity indices. Our approach uses resampled macroremain influxes that enable the computation of the rarefaction index for the low influx records. The raw counts are resampled to a constant resolution and sample volume by interpolating initial sample ages at a constant time interval using the age∼depth model. Then, the contribution of initial counts and volume to each interpolated sample is determined by calculating a proportion matrix that is in turn used to obtain regularly spaced time series of pollen and macroremain influx. We applied this algorithm to sedimentary data from a subalpine lake situated in the European Alps. The reconstructed total floristic richness at the study site increased gradually when both pollen and macroremain records indicated a decrease in relative abundances of shrubs and an increase in trees from 11,000 to 7,000 cal BP. This points to an ecosystem change that favored trees against shrubs, whereas herb abundance remained stable. Since 6,000 cal BP, local richness decreased based on plant macroremains, while pollen-based richness was stable. The reconstructed richness and evenness are interrelated confirming the difficulty to distinguish these two aspects for the studies in paleo-biodiversity. The present study shows that low-influx bio-proxy records (here macroremains) can be used to reconstruct stand diversity and address ecological issues. These developments on macroremain and pollen records may contribute to bridge the gap between paleoecology and biodiversity studies.

Vegetation limits the impact of a warm climate on boreal wildfires.

Strategic introduction of less flammable broadleaf vegetation into landscapes was suggested as a management strategy for decreasing the risk of boreal wildfires projected under climatic change. However, the realization and strength of this offsetting effect in an actual environment remain to be demonstrated. Here we combined paleoecological data, global climate models and wildfire modelling to assess regional fire frequency (RegFF, i.e. the number of fires through time) in boreal forests as it relates to tree species composition and climate over millennial time-scales. Lacustrine charcoals from northern landscapes of eastern boreal Canada indicate that RegFF during the mid-Holocene (6000-3000 yr ago) was significantly higher than pre-industrial RegFF (AD c. 1750). In southern landscapes, RegFF was not significantly higher than the pre-industrial RegFF in spite of the declining drought severity. The modelling experiment indicates that the high fire risk brought about by a warmer and drier climate in the south during the mid-Holocene was offset by a higher broadleaf component. Our data highlight an important function for broadleaf vegetation in determining boreal RegFF in a warmer climate. We estimate that its feedback may be large enough to offset the projected climate change impacts on drought conditions.

Fire, fuel composition and resilience threshold in subalpine ecosystem.

BACKGROUND: Forecasting the effects of global changes on high altitude ecosystems requires an understanding of the long-term relationships between biota and forcing factors to identify resilience thresholds. Fire is a crucial forcing factor: both fuel build-up from land-abandonment in European mountains, and more droughts linked to global warming are likely to increase fire risks. METHODS: To assess the vegetation response to fire on a millennium time-scale, we analyzed evidence of stand-to-local vegetation dynamics derived from sedimentary plant macroremains from two subalpine lakes. Paleobotanical reconstructions at high temporal resolution, together with a fire frequency reconstruction inferred from sedimentary charcoal, were analyzed by Superposed Epoch Analysis to model plant behavior before, during and after fire events. PRINCIPAL FINDINGS: We show that fuel build-up from arolla pine (Pinus cembra) always precedes fires, which is immediately followed by a rapid increase of birch (Betula sp.), then by ericaceous species after 25-75 years, and by herbs after 50-100 years. European larch (Larix decidua), which is the natural co-dominant species of subalpine forests with Pinus cembra, is not sensitive to fire, while the abundance of Pinus cembra is altered within a 150-year period after fires. A long-term trend in vegetation dynamics is apparent, wherein species that abound later in succession are the functional drivers, loading the environment with fuel for fires. This system can only be functional if fires are mainly driven by external factors (e.g. climate), with the mean interval between fires being longer than the minimum time required to reach the late successional stage, here 150 years. CONCLUSION: Current global warming conditions which increase drought occurrences, combined with the abandonment of land in European mountain areas, creates ideal ecological conditions for the ignition and the spread of fire. A fire return interval of less than 150 years would threaten the dominant species and might override the resilience of subalpine forests.

A new, isolated and endangered relict population of dwarf pine (Pinus mugo Turra) in the northwestern Alps.

Dwarf pines were discovered in 2004 during a paleoecological survey in the Mont Cenis massif (Savoy, France). These dwarf pines are the sole natural and spontaneous population in the NW French Alps of Pinus mugo Turra, ssp. mughus (Scop.) O. Schwarz. The population, fragile in light of the individual numbers, is currently isolated, but likely results from populations that would have covered larger areas during the Lateglacial or the early-Holocene, from the SW Alps (France) toward the Mont Cenis, throughout the Susa valley (Italy). With a fragmented distribution area of dwarf pine, the future of the Mont Cenis population seems altered due to important necroses observed on the topmost part of pine crowns, except for individuals that do not grow in the local ski station area. We stress the need for a conservation program to preserve this natural population, the sole known in the NW French Alps.

Long-term fire frequency not linked to prehistoric occupations in northern Swedish boreal forest.

Knowledge of past fire regimes is crucial for understanding the changes in fire frequency that are likely to occur during the coming decades as a result of global warming and land-use change. This is a key issue for the sustainable management of forest biodiversity because fire regimes may be controlled by vegetation, human activities, and/or climate. The present paper aims to reconstruct the pattern of fire frequency over the Holocene at three sites located in the same region in the northern Swedish boreal forest. The fire regime is reconstructed from sedimentary charcoal analysis of small lakes or ponds. This method allows fire events to be characterized, after detrending the charcoal influx series, and allows estimation of the time elapsed between fires. The long-term fire regime, in terms of fire-free intervals, can thus be elucidated. At the three sites, the mean fire-free intervals through the Holocene were long and of similar magnitude (approximately 320 years). This similarity suggests that the ecological processes controlling fire ignition and spread were the same. At the three sites, the intervals were shorter before 8600 cal yr BP (calibrated years before present), between 7500 and 4500 cal yr BP, and after 2500 cal yr BP. Geomorphological and vegetation factors cannot explain the observed change, because the three sites are located in the same large ecological region characterized by Pinus sylvestris-Ericaceae mesic forests, established on morainic deposits at the same elevation. Archaeological chronologies also do not match the fire chronologies. A climatic interpretation is therefore the most likely explanation of the long-term regional pattern of fire. Although recent human activities between the 18th and the 20th centuries have clearly affected the fire regime, the dominant factor controlling it for 10000 years in northern Sweden has probably been climatic.

[Fire and Pinus mugo Turra communities in the western Alps (Susa Valley, Italy) during the Lateglacial-Holocene transition: an evidence of refugia area]. / Incendies et peuplements à Pinus mugo Turra dans les Alpes occidentales (Val de Suse, Italie) durant la transition Tardiglaciaire-Holocène: une zone refuge évidente.

In western Italian Alps, small distinct populations of Pinus mugo Turra raise some questions concerning its ecological status and dynamics in the occidental Alps. This note present new palaeobotanical data based on cone imprints of Pinus mugo, identified in travertine systems located in the Val di Susa and dated back to the Late Dryas (11506+/-66 BP) and the Early Holocene (10145+/-225 et 9475+/-670 BP). Heliophilous species and charcoal fragments were also identified, testifying to the oldness of wildfires in this region. The data support the hypothesis that this zone was a refuge area of this pine during the last glaciation. Here we discuss about the postglacial dynamics of the Pinus mugo in the occidental Alps.

[Proof of the indigenous nature of Populus alba L. in the western Mediterranean Basin]. / Preuve de l'indigénat de Populus alba L. dans le Bassin méditerranéen occidental.

Around the western Mediterranean Basin, the ecological status of Populus alba, whether indigenous or introduced, is controversial. This note presents new palaeobotanical data based on analyses of leaf imprints from a travertine formation located in southern France. This travertine presents two levels with Populus alba imprints. The oldest level is dated back by 14C to the Early Holocene, i.e., 8890 +/- 70 BP. This demonstrates that Populus alba is an autochthonous species of the southern-France vegetation, removing speculations reporting that its distribution area was greatly influenced by Roman civilization. Finally, we discuss this new data in regard to other Pleistocene and Holocene deposits circum the Mediterranean Basin and in Europe, where this species was identified.