Responses of Pinus resinosa in Newfoundland to wildfire

Abstract. Natural Pinus resinosa (red pine) stands in Newfoundland are restricted to 22 small, dry, nutrient-poor sites. A short wildfire cycle (15 - 30 yr) of both surface and crown fire regulates stand perimeters and is the main factor in regulating stand development. At the nucleus of current stands < 100 yr old, a few trees > 200 yr occur, usually showing multiple fire scars. Stem char heights confirmed an increased flammability of the stand and tree mortality for fires moving in the upslope position, as well as for mixed Pinus resinosa-Picea mariana stands. All P. resinosa stands are severely nutrient-deficient. Leaf concentrations of N, P and K were below or near the reported critical values. Nutrient concentrations were highest three months after a surface fire, but dropped considerably one year later. A gradual increase to near post-fire levels is achieved four years after fire. Foliar nutrient concentrations were positively correlated with average width of the annual rings. Aspects of the ericaceous understory dynamics and its relation to P. resinosa regeneration are also discussed.     

Explaining the distribution of Pinus spp. in a Canadian boreal insular landscape

Abstract. To evaluate the respective contributions of habitat, fire regime and colonization-extinction processes to the distribution of northern Pinus species, we investigated the distribution of P. banksiana (jack pine), P. resinosa (red pine) and P. strobus (white pine) on 117 islands of Lake Duparquet in northwestern Québec. Stepwise logistic regressions indicated that the extent of xeric areas on the islands was the sole factor predicting jack pine distribution. The distribution of white pine was predicted primarily by the combined effects of distance to the shoreline and elevation, with a smaller effect of area of xeric habitat. The distribution of red pine was predicted by other populations of red pine nearby, with a slightly smaller effect of the combined effects of distance to shoreline and elevation. None of the species completely saturates all available islands nor is any restricted to specific, very exposed aspects. The results suggest that pine is more frequent on islands with characteristics that promote lightning strikes and thus higher fire occurrence. However, absence of pine in several islands may not be explained by abiotic characteristics or recent fire history. The presence of very small populations, together with low invasion potential, suggests that the observed distribution is mainly driven by the process of random extinction. A disequilibrium between present and past fire regimes may explain why northern pines have discontinuous distributions inside their range limits.     

Natural regeneration of Pinus resinosa on burned and unburned sites in Newfoundland

Abstract. Natural regeneration of Pinus resinosa (red pine) seedlings around mature trees was studied in burned and unburned stands. Growth inhibitory effects of the forest organic matter on red pine seedlings was tested by a stair-step experiment using leachate of forest soil monoliths and also by a seed germination bio-assay using forest floor substrates. To test if higher burning temperatures can remove the allelopathic effects of red pine-Kalmia organic matter, a laboratory bio-assay was conducted by germinating red pine seeds on the organic matter burned at 200, 400, 600 and 800°C. Deposition of dry needles and a thick duff layer under red pine stands affected seedling establishment. Red pine seedling establishment increased with the decreasing thickness of duff layer away from the stump of the seed-bearing trees. Wildfire helped in removing the duff layer and increased seedling establishment. A high fuel load within a 0 - 1 m radius around the tree stump caused a deep burn of the organic matter including part of the soil seed reserve. On a burned-over surface, more seedlings established in a band between 1 and 2 m around the stump than inside and outside the band. Primary root growth of red pine was severely inhibited when the seedlings were grown in unburned forest floor organic matter where Kalmia was the principal understory species. Water leachate of a Pinus resinosa-Kalmia soil monolith was inhibitory to red pine seedling growth. In greenhouse conditions, the seedlings grew well in burned-over soil from a Pinus resinosa stand. Burned organic matter from a red pine forest showed an increase in pH with a burning temperature of 600°C. Primary root growth of red pine seedlings was similarly increased with increasing temperature up to 600°C; at higher temperatures the root length of seedlings did not increase any further.     

Comparative studies of ectomycorrhiza formation in Alnus glutinosa and Pinus resinosa with Paxillus involutus

 Mycorrhiza ontogeny and details of Hartig net and mantle structure were compared in ectomycorrhizas synthesized in growth pouches between the broad host range fungus Paxillus involutus and the tree species European black alder (Alnus glutinosa) and red pine (Pinus resinosa). In Alnus glutinosa, a paraepidermal Hartig net was restricted to the proximal (basal) portion of first-order laterals; the hypodermal layer appeared to be a barrier to fungal penetration. Phi-thickenings were present in some cortical cells but these were not related to lack of fungal ingress into the cortex. The mantle was often present close to the root apex but in many roots it was loosely organized and patchy. In several instances, the mantle formed around the root apex was only temporary; renewed root growth occurred without the formation of a mantle. In Pinus resinosa, the Hartig net developed between cortical cell layers of monopodial and dichotomously branched first–order laterals. Fungal hyphae in the Hartig net exhibited a complex labyrinthine mode of growth. The mantle had a pseudoparenchymatous structure and covered the root, including apices of dichotomously branched roots. The Paxillus–Pinus resinosa interaction had all the characteristics of a compatible ectomycorrhizal association. The Paxillus–Alnus glutinosa interaction, however, showed only aspects of superficial ectomycorrhizas, including the presence of a minimal (sometimes absent) and mostly proximal Hartig net and variable mantle development. Sclerotia were produced in the extraradical mycelium of Paxillus involutus when associated with either Alnus glutinosa or Pinus resinosa. Accepted: 22 October 1998     

Population age structure of Pinus banksiana at the southern edge of the Canadian boreal forest

Abstract. To assess the effects of site type, forest initiation periods and fire regimes on the dynamics of Pinus banksiana (Jack pine), the age structure of 69 populations of the species was analyzed. Two landscapes with different fire regimes were selected in the southern part of the Canadian boreal forest in Québec: the ‘mainland landscape’ is characterized by a fire regime of large lethal fires, the ‘island landscape’ is affected by a complex fire regime including lethal and non-lethal fires. Age structure was compared between forest initiation periods and site types (mesic mainland, xeric mainland and xeric island) using the Shannon regularity index. An even-aged population structure was found within the first 100 yr following a lethal fire, while after that period the population structure becomes more uneven-aged. Under mesic conditions, populations tend to have an even-aged structure, under xeric conditions an uneven-aged structure. Natural openings present in xeric sites allow for recruitment in the absence of fire. This permits the self-maintenance of Pinus banksiana. Xeric island populations show more uneven-aged structures than xeric mainland populations. The occurrence of non-lethal fires on the islands creates uneven-aged structures. Further, the results suggest that the selection pressure of the island fire regime, favouring non-serotinous and mixed P. banksiana individuals, is one of the factors responsible for a higher recruitment in the absence of fire on islands than on the mainland.     

Vegetation of Sphagnum bogs in highly disturbed landscapes: relative influence of abiotic and anthropogenic factors

Question: Has the vegetation of Sphagnum bogs been affected by more than 200 years of human activities? Location: Bas‐Saint‐Laurent region, southeastern Québec, Canada. Methods: Data (species assemblages, abiotic and spatio‐historical variables) were collected in 16 bogs ranging from 2 to 189 ha, and incorporated in a geographical information system. Major gradients in vegetation composition were identified using DCA. CCA was used to relate vegetation gradients to abiotic and spatio‐historical variables. Results: A clear segregation of species assemblages was observed, from open and undisturbed bogs to forested and highly disturbed sites. Among abiotic factors, tree basal area, water table level and peat thickness had a significant influence on plant species composition. Among spatio‐historical factors, disturbance level, area loss and fire were the most influential factors. Variance partitioning between these groups of factors suggests that spatio‐historical factors had a major influence on peatlands, representing 22% of the variation observed in the plant species assemblages while abiotic factors represent only 17% of the variation. Conclusions: The results highlight the influence of agricultural and other anthropogenic activities on plant assemblages and suggest that even wetlands apparently resistant to disturbances, such as peatlands, can be severely affected by anthropogenic factors. Plant species assemblages of ombrotrophic peatlands of the Bas‐Saint‐Laurent region were, and still are, largely influenced by human activities.     

Hotspots of biotic compositional change in lakes along vast latitudinal transects in northern Canada

Ecotones mark zones of rapid change in ecological structure at various spatial scales. They are believed to be particularly susceptible to shifts caused by environmental transformation, making them key regions for studying the effects of global change. Here, we explored the variation in assemblage structure of aquatic primary producer and consumer communities across latitudinal transects in northeastern North America (Québec‐Labrador) to identify spatial patterns in biodiversity that indicated the location of transition zones across the landscape. We analyzed species richness and the cumulative rate of compositional change (expressed as beta‐diversity) of diatoms and chironomids to detect any abrupt shifts in the rate of spatial taxonomic turnover. We used principal coordinates analysis to estimate community turnover with latitude, then applied piecewise linear regression to assess the position of ecotones. Statistically significant changes in assemblage composition occurred at 52 and 55°N, corresponding to the transition between closed‐ and open‐crown forest, and to the southern onset of the forest tundra (i.e., the forest limit), respectively. The spatial distribution of ecotones was most strongly related to air temperature for chironomids and to vegetation‐ and soil‐related chemical attributes of lake water for diatoms, including dissolved organic carbon content and water color. Lakes at mid‐ to high‐latitudes currently face pressures from rapidly rising temperatures, accompanied by large increases in organic carbon inputs from their catchments, often leading to browning and its associated effects. The biota at the base of food webs in lakes located in transition zones are disproportionately affected by the cascading effects of these multi‐factorial changes, concurrent with pronounced terrestrial greening observed in these regions. Similar patterns of biotic shifts have been observed along alpine aquatic transects, indicating the potential for widespread restructuring of cold, high‐altitude and high‐latitude freshwater communities due to global change.     

Discovery of the oldest record of Nitellopsis obtusa (Charophyceae,Charophyta) in North America

Studies of the colonization and spread of invasive species improves our understanding of key concepts in population biology as well as informs control and prevention efforts. The characean green alga Nitellopsis obtusa (starry stonewort) is rare in its native Eurasian range but listed by the United States Geological Survey (USGS) as an aggressive invasive in North America. First documented in North America in 1978 from New York, United States, it has since been reported from numerous inland lakes from Minnesota to Vermont, and from Lake Ontario and inland lakes in southern Ontario, Canada. While the ecological impacts of N. obtusa are not clearly understood in its invasive range, initial results show negative environmental effects. We have discovered a liquid‐preserved herbarium specimen that predates the 1978 records by at least 4 years, and is the first confirmed record of N. obtusa in Québec.     

Environmental factors explaining the vegetation patterns in a temperate peatland

Although ombrotrophic temperate peatlands are important ecosystems for maintaining biodiversity in eastern North America, the environmental factors influencing their flora are only partly understood. The relationships between plant species distribution and environmental factors were thus studied within the oldest temperate peatland of Québec. Plant assemblages were identified by cluster analysis while CCA was used to related vegetation gradients to environmental factors. Five assemblages were identified; three typical of open bog and two characterized by more minerotrophic vegetation. Thicker peat deposit was encounter underlying the bog assemblages while higher water table level and percentage of free surface water distinguished the minerotrophic assemblages. Overall, the floristic patterns observed were spatially structured along the margins and the expanse. The most important environmental factors explaining this spatial gradient were the percentage of free surface water and the highest water-table level. To cite this article: S. Pellerin et al., C. R. Biologies 332 (2009).     

Distributional gradients and variability of macroelement concentrations in the crowns of plantation grownPinus resinosa (AIT.)

Summary The vertical distribution of inorganic nutrient concentrations in red pine were dependent on the foliage age. Older foliage did not show an average vertical gradient while younger foliage did show a significant gradient. Horizontal gradients across foliage age classes in a branch whorl were consistent for all branch whorls, but the relative difference between the concentration of the current foliage and foliage three years-old or older was dependent on crown position. Coefficients of variation (CV) did not show a variability gradient in the crown for nitrogen and phosphorus. Variability of potassium tended to decrease as foliage age increased. Contribution of the University of Florida, Soil Science Department, Gainesville, FL and State University of New York, College of Environmental Science and Forestry, Syracuse, New York. Florida Agricultural Experiment Station Journal Series No. 3017.     

Growth responses of Pinus sylvestris L. to climate from the southeastern limit of its natural distribution area,Turkey

Pinus sylvestris, the most widely distributed pine species, is commonly used in dendrochronological studies. Based on a lack of studies at its southeastern distribution, we analysed the growth responses of P. sylvestris to temperature and precipitation. We selected 13 sites to study the effects of climate on the growth of Scots pine stands throughout a geographic gradient over time. Trees were sampled from pure stands at different elevations and landscape conditions. The linear and non-linear associations between tree-ring widths and climate variables were calculated with locally specific linear correlation analysis and a mixed generalised additive model. Moving window correlation function was also performed to understand the temporal stability of limiting factors on growth from 1930 to 2013. Our findings showed that early spring temperature (March-April) and late spring-early summer precipitation (May-June) are the major drivers of growth at all sites, where high temperature constraints and high precipitation enhances the growth. Moving window correlation analysis highlighted that the response to precipitation was stationary while temperature changed over time. Our non-linear analysis provided a threshold for March-April temperature. The threshold indicates that the relative additional increment sharply increases up to 7 °C and then slightly decreases.     

Riverine wetland vegetation: importance of small-scale and large-scale environmental variation

Abstract. The purpose of this study was to evaluate the relative importance of small-scale variation in abiotic factors and large-scale spatio-temporal variation on the distribution of wetland vegetation of a section of the Upper St. Lawrence River in Québec. Vegetation data have been classified with agglomerative clustering into 11 community types, from Acer rubrum, Acer saccharinum and Fraxinus swamps, to scrubs dominated by Salix petiolaris, Alnus rugosa var. americana or Myrica gale, to Typha, Typha/Lythrum, Carex lacustris and Calamagrostis canadensis marshes. Canonical Correspondence Analysis (CANOCO) suggested that peat thickness and water level are the most important abiotic variables correlated with plant community composition. As a whole, small-scale variation accounts for 25.6 % of the species variation. Another 21.6% is explained by large-scale variation based on location data summarizing large-scale spatial distribution, and historical landscape dynamics differentiated into (a) no net loss of wetlands, (b) net loss of wetland, and (c) changes within wetlands, which are correlated with the actual variation in herbaceous and scrubby vegetation. The interaction between small-scale and large-scale variations explains another 1.7 %. In total, 48.9% of the species variation is explained by the two data sets, leaving 51.1% unexplained. Whereas the omission of some abiotic variables is possible, it is hypothesized that the abiotic conditions measured in this study play an important role, especially in the distribution of forested swamps. In addition, past history, particularly that of human interventions, becomes another important factor leading to the observed importance of large-scale spatio-temporal variables. This is particularly true for Alnus rugosa var. americana shrublands. Time lag between a relative stabilization of species distribution and the reduction of natural disturbances (water level fluctuations and fires) could be a possible cause of the importance of spatio-temporal variables and the undetermined portion of species variation.     

Ecological investigations on the northern distribution range of Hippocrepis comosa L. in Germany

The objective of this study was to identify the causes of the limits of the geographical range of Hippocrepis comosa. Along a gradient from the northwestern distribution boundary towards the distribution centre in Germany, 46 field plots were established where growth and microclimate were monitored simultaneously. In total, 11 vegetative and generative traits and 7 microclimatic parameters were recorded over 7 time intervals during the vegetation period, together with 14 general site parameters. Regression analyses were performed between all traits and environmental parameters in a certain interval. At the beginning of the growing season the best positive correlation coefficients for vegetative growth were observed with soil temperature. From the end of May to the middle of June, vegetative growth rates decreased and showed the best positive correlation with soil water content. Despite credible relationships between vegetative growth and microclimate, their contribution toward explaining the northern distribution boundary was found to be limited, because no correlation with the distance from the distribution boundary was observed. The only growth parameter that showed both a positive correlation with distance from the distribution boundary and a significant correlation with microclimate was the percentage of seed setting, which increased towards the distribution centre and was correlated with air temperature. Further field observations on plots outside the actual range of Hippocrepis comosa revealed no microclimatic reasons as to why the species was absent from these sites. This shows that the environmental parameters are in no way deterministic for the range limit. The frost hardiness of Hippocrepis comosa was studied in additional laboratory experiments in which significant damage was not found above −18 °C for adult plants and above −14 °C for seedlings, which is remarkably low and too low to be relevant for the northwestern distribution boundary. Another field experiment revealed that seedling establishment exhibited a positive relationship to soil water content, which became more favourable towards the range boundary. It is concluded that temperature, particularly air temperature, makes the largest contribution to explaining the northern distribution boundary of Hippocrepis comosa in Germany and that mainly generative reproduction is affected. This revised version was published online in August 2006 with corrections to the Cover Date.     

Biodiversity and resilience of arthropod communities after fire disturbance in temperate forests

Changes in ecosystem functions following disturbances are of central concern in ecology and a challenge for ecologists is to understand the factors that affect the resilience of community structures and ecosystem functions. In many forest ecosystems, one such important natural disturbance is fire. The aim of this study was to understand the variation of resilience in six functional groups of invertebrates in response to different fire frequencies in southern Switzerland. We measured resilience by analysing arthropod species composition, abundance and diversity in plots where the elapsed time after single or repeated fires, as determined by dendrochronology, varied. We compared data from these plots with data from plots that had not burned recently and defined high resilience as the rapid recovery of the species composition to that prior to fire. Pooling all functional groups showed that they were more resilient to single fires than to repeated events, recovering 6–14 years after a single fire, but only 17–24 years after the last of several fires. Flying zoophagous and phytophagous arthropods were the most resilient groups. Pollinophagous and epigaeic zoophagous species showed intermediate resilience, while ground-litter saprophagous and saproxylophagous arthropods clearly displayed the lowest resilience to fire. Their species composition 17–24 years post-burn still differed markedly from that of the unburned control plots. Depending on the fire history of a forest plot, we found significant differences in the dominance hierarchy among invertebrate species. Any attempt to imitate natural disturbances, such as fire, through forest management must take into account the recovery times of biodiversity, including functional group composition, to ensure the conservation of multiple taxa and ecosystem functions in a sustainable manner.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

How needle phenology indicates the changes of xylem cell formation during drought stress in Pinus sylvestris L.

Temperature, water availability and photoperiod are the primary drivers of tree phenological processes. However, there is lack of information how the spring temperature and summer drought affect phenology of both cambium and needles. We evaluated the interplay between xylem and needle development of mature Scots pine trees in three consecutive years differing in the spring temperatures and onset and duration of summer drought. Cambial activity began on the day of a year (DOY) 83–87, while the bud break occurred on DOY 113–119, when also the secondary cell wall of tracheids started its formation. While the timing of bud break correlated with the sum of effective temperatures, the beginning of cambial zone activity did not. The needles were fully unfolded around DOY 170, which corresponded to the transition between earlywood and latewood tracheids. Summer drought did not affect needle development, but it changed the rate of production and morphology of latewood tracheids. Latewood tracheids from the year 2015 when the drought was longest (i.e. until the second half of August in 2015 compared to the duration from June until the middle of July in the two other years) were 32% narrower and they had 34% thinner cell walls than in 2014 and 2016. The improvement of tree water status in July resulted in the formation of intra-annual density fluctuations (IADF). The interplay between needle and cambium phenology provided an insight into how the tree allocates the resources with varying temperature and soil water availability.     

Ecological effects of changes in fire regimes in Pinus ponderosa ecosystems in the Colorado Front Range

Question: What is the relative importance of low‐ and high‐severity fires in shaping forest structure across the range of Pinus ponderosa in northern Colorado? Location: Colorado Front Range, USA. Methods: To assess severities of historic fires, 24 sites were sampled across an elevation range of 1800 to 2800 m for fire scars, tree establishment dates, tree mortality, and changes in tree‐ring growth. Results: Below 1950 m, the high number of fire scars, scarcity of large post‐fire cohorts, and lack of synchronous tree mortality or growth releases, indicate that historic fires were of low severity. In contrast, above 2200 m, fire severity was greater but frequency of widespread fires was substantially less. At 18 sites above 1950 m, 34 to 80% of the live trees date from establishment associated with the last moderate‐ to high‐severity fire. In these 18 sites, only 2 to 52% of the living trees pre‐date these fires suggesting that fire severities prior to any effects of fire suppression were sufficient to kill many trees. Conclusions: These findings for the P. ponderosa zone above ca. 2200 m (i.e. most of the zone) contradict the widespread perception that fire exclusion, at least at the stand scale of tens to hundreds of hectares, has resulted in unnaturally high stand densities or in an atypical abundance of shade‐tolerant species. At relatively mesic sites (e.g. higher elevation, north‐facing), the historic fire regime consisted of a variable‐severity regime, but forest structure was shaped primarily by severe fires rather than by surface fires.     

Conifer seedling recruitment in a southeastern Canadian boreal forest: the importance of substrate

Abstract. In order to explain conifer species recruitment in Canada's southeastern boreal forest, we characterized conifer regeneration microsites and determined how these microsites vary in abundance during succession. Microsite abundance was evaluated in deciduous, mixed and coniferous stands along a 234-yr postfire chronosequence. Conifers were most often found in relatively well-illuminated microsites, devoid of litter, especially broad-leaf litter, and with a reduced cover of lower vegetation (< 50 cm tall). Although associated with moss-rich forest floor substrates, Abies balsamea was the most ubiquitously distributed species. Picea glauca and especially Thuja occidentalis seedlings were frequently found on rotten logs. Light measurements did not show differences among seedling species nor between stand types. The percentage cover of broad-leaf litter decreased significantly during succession. Also, rotten logs covered with moss occupied a significantly larger area in the mid-successionnal stands than in early successional deciduous or late successional coniferous stands. The results suggest that the presence of specific forest floor substrate types is a factor explaining low conifer recruitment under deciduous stands, conifer codominance in the mid-successional stage, and delayed Thuja recolonization after fire. Results also suggest that some facilitation mechanism is responsible for the observed directional succession.     

Strategies in Mediterranean grassland annuals in relation to stress and disturbance

Abstract. Morphological (size and shape) and functional (growth, reproduction and phenology) attributes are used to characterize 42 annual species of Mediterranean grasslands according to their strategy. Principal component analysis of the matrix of 42 species x 9 attributes shows that the main trend of variation is related to plant size. Larger species have larger seeds, lower relative growth rates and lower reproductive output. The second and third trends of variation are related to plant shape. Ordination of species shows differences in shape between taxa and growth forms (grasses, legumes, forbs). The relative abundance of species with different attributes vary with the level of stress (water and nutrient availability) and disturbance (grazing and ploughing). Size is related to stress, with larger plants dominating in productive habitats and smaller ones in the most unproductive. Disturbance is related to shape and phenology, since grazing favours species with low canopies and ploughing favours species with shorter life cycles, that are usually small in size. Relations between plant attributes and habitat characteristics are examined within the more general framework of plant strategy theory.