Tornado damage of Quercus stellata and Quercus marilandica in the Cross Timbers,Oklahoma, USA

Questions: The Cross Timbers are a mosaic of savannas, grasslands and upland forests, occupying a significant portion of south‐central North America. Our questions here were (1) how does a severe tornado affect the two most dominant tree species of the area Quercus marilandica and Q. stellata with respect to damage and mortality; (2) how do such patterns vary as a function of tree size? What are the implications of disturbance for codominance in species‐poor systems? Location: The Cross Timbers in Oklahoma, USA. Methods: We established a 14.48‐ha permanent plot following a severe tornado in 2003. We identified, numbered and tagged each tree and recorded its diameter at breast height (DBH), spatial coordinates, status (dead or alive), and damage type. We examined (1) relative abundance before and after the tornado; (2) differences in damage and mortality, and (3) the influence of tree diameter on the probability of damage and mortality for each species. Results: Differences in species identity and tree characteristics were significantly related to tree mortality following the tornado, after accounting for spatial locations. The odds of mortality were 12.0 times greater for Q. marilandica than for Q. stellata. Such greater vulnerability of Q. marilandica versus Q. stellata was also reflected in changes in density and basal area. Tree diameter clearly influenced the damage and mortality pattern in Q. stellata; larger trees sustained more damage and mortality. However, Q. marilandica did not exhibit size‐dependent mortality. Conclusion: The tornado affected the two dominant species differently. The intra‐ and inter‐specific differences in windstorm susceptibility may allow coexistence of the two species and are potentially important in the dynamics of the Cross Timbers. Species more damaged might finally benefit from the wind disturbance due to their resprouting ability.     

ORDINATION AND CLASSIFICATION OF WESTERN OAK FORESTS IN OKLAHOMA

Ordination and classification techniques were used to analyze patterns of forest vegetation, species diversity, and soil type in the Wichita Mountains Wildlife Refuge of southwestern Oklahoma. Cluster analysis based on tree species produced three general community types: 1) Quercus stellata-Q. marilandica forests; 2) Q. stellata forests; and 3) mesophytic forests. A polar ordination produced a gradient of vegetation that corresponded to a moisture gradient. Many high diversity forests were located on loamy drainageway soils or north facing slopes. Tree species diversity (H') was inversely related to the importance of Quercus stellata. Cluster analysis based on species composition of the tree seedlings produced four general community types: 1) Q. marilandica type; 2) Q. marilandica-Q. stellata-Juniperus virginiana type; 3) Ulmus americana-Celtis reticulata-Bumelia lanuginosa type; and 4) Acer saccharum type. The third seedling type occurred almost exclusively on loamy drainageway soils. There was no relationship between stand location on the first axis of the tree ordination and the first axis of the seedling ordination suggesting that trees and seedlings respond differently along the moisture gradient.     

Vegetation ecology of flatwoods on the Illinoian till plain

Abstract. Midwestern flatwoods are open woodlands of level uplands and terraces that occur on impervious subsoil horizons and have seasonally wet and dry soils. Vegetation and soil data from six flatwoods remnants on the Illinoian till plain in Illinois were examined to identify what biotic and edaphic factors control vegetation structure, composition and diversity. Soils at all sites are characterized by the presence of argillic horizons (clay pans). Two soil ordination groups were clearly defined, each characterized by separate parent material origin. Canopy and subcanopy composition are dissimilar at sites with soils developed entirely in loess; more similar at sites on lacustrine soils (including aeolian sand). Quercus stellata was by far the most important tree species followed by Q. marilandica. Q. stellata importance was negatively correlated with Ca, particularly in the A-horizon. Q. marilandica shows a positive correlation to sand and magnesium, particularly in the B-horizon, and was most frequent and abundant on soils where Ca/Mg ratios are lower than 1.0 in the E and B-horizons. Total stem density of woody plants was positively related to total available soil moisture as influenced by depth to clay pan and soil textural differences. Ground-cover species richness and diversity were inversely related to tree and total woody stem density and tree diversity. One site with a recent history of annual prescribed fire had significantly greater ground-cover species diversity compared with the unburned sites. Evidence from stand structure and similarity between canopy and subcanopy strata, if reflecting stationary processes of canopy recruitment, suggest that disturbances historically maintained the importance of oak species in most flatwoods. Prescribed fire appears to maintain or enhance ground-cover species diversity at one flatwoods remnant, suggesting that fire could be used to maintain compositional stability at sites with greater available soil moisture.     

Large‐scale assessment of regeneration and diversity in Mediterranean planted pine forests along ecological gradients

Aim There is increasing concern regarding sustainable management and restoration of planted forests, particularly in the Mediterranean Basin where pine species have been widely used. The aim of this study was to analyse the environmental and structural characteristics of Mediterranean planted pine forests in relation to natural pine forests. Specifically, we assessed recruitment and woody species richness along climatic, structural and perturbation gradients to aid in developing restoration guidelines. Location Continental Spain. Methods We conducted a multivariate comparison of ecological characteristics in planted and natural stands of main Iberian native pine species (Pinus halepensis, Pinus pinea, Pinus pinaster, Pinus nigra and Pinus sylvestris). We fitted species‐specific statistical models of recruitment and woody species richness and analysed the response of natural and planted stands along ecological gradients. Results Planted pine forests occurred on average on poorer soils and experienced higher anthropic disturbance rates (fire frequency and anthropic mortality) than natural pine forests. Planted pine forests had lower regeneration and diversity levels than natural pine forests, and these differences were more pronounced in mountain pine stands. The largest differences in recruitment – chiefly oak seedling abundance – and species richness between planted and natural stands occurred at low‐medium values of annual precipitation, stand tree density, distance to Quercus forests and fire frequency, whereas differences usually disappeared in the upper part of the gradients. Main conclusions Structural characteristics and patterns of recruitment and species richness differ in pine planted forests compared to natural pine ecosystems in the Mediterranean, especially for mountain pines. However, management options exist that would reduce differences between these forest types, where restoration towards more natural conditions is feasible. To increase recruitment and diversity, vertical and horizontal heterogeneity could be promoted by thinning in high‐density and homogeneous stands, while enrichment planting would be desirable in mesic and medium‐density planted forests.     

Patterns from the past: modeling Public Land Survey witness tree distributions with weights-of-evidence

The Public Land Survey (PLS) witness tree data provide one of the few quantitative data sets of pre-and early-European settlement composition and structure of the forests and woodlands in the western United States. However, quantifying the areal extent of individual woody species from PLS records has proven difficult due to the coarse sampling structure of the data. Several attempts have been made to convert the discrete PLS witness tree data into continuous distributions through the use of various interpolation techniques. While these methods may adequately represent the spatial patterns of individual species over large areas, they fail to consider the numerous environmental covariates that can influence the distribution of individual tree species at finer scales. A more statistically rigorous method calls for combining species–environment relationships to estimate the areal extent of individual species from point data. In this study, we utilize weights-of-evidence (WofE), a discrete multivariate method, to estimate the probable historical distribution of six important woody plant taxa of the cross timbers of south-central Oklahoma. We successfully created posterior probability distribution maps for Quercus stellata, Q. marilandica, Q. velutina, Carya texana, C. illinoinensis, and Juniperus spp. Each posterior probability map was classified into four predictive categories, thereby enabling better estimations of the historical distribution of individual taxon from coarse-resolution PLS data. Model validation indicated that the WofE method effectively estimated the posterior probabilities of all taxa under consideration.     

Tree dispersion in oak-dominated forests along an environmental gradient

Summary Spatial pattern was analyzed in seventeen stands of oak-dominated forest to address the hypothesis that species tended to be aggregated under favorable conditions and widely spaced in xeric, nutrient poor conditions. Trees were sampled at 80–100 points in each stand with the distance-to-nearest neighbor method. Soil samples were collected in each stand for analysis of total nitrogen, total phosphorus, total potassium, soil pH, soil texture, and soil organic matter. Growing season precipitation was also recorded from climate stations near each stand. Quercus stellata (Wang.) dominated 10 stands, Q. marilandica (Muenchh.) dominated three stands and these species were codominant in four stands. Principal components analysis identified a soil texture/fertility gradient across the study area. Quercus stellata and all species combined were aggregated in most stands, whereas Q. marilandica was mostly randomly distributed within a stand. Small trees of all species combined tended to be aggregated and large trees were randomly dispersed in all but two stands, suggesting competition. Mean distance between large-large pairs was always greater than mean distance between small-small pairs in all stands, but this difference was only significant in one stand. Correlations between nearest neighbor distance and combined size of nearest neighbors were significant and positive in 12 of 17 stands. In all cases, however, slopes were shallow suggesting that competition is weak in these communities and has a limited effect on spacing of neighboring trees. Contrary to our hypothesis, trees were more aggregated on coarse-textured soils with low organic matter content. For all species combined, degree of aggregation was unrelated to growing season precipitation. Aggregation appears to be common in these forests because environmental stress in many stands reduces growth rates. Trees have not yet reached a size at which competition or other interactions can greatly increase interplant distances and reduce the degree of aggregation. A simple graphical model is developed to describe the relationship between patterns, stress and competition in plant communities.     

Short-term development of ambrosia and bark beetle assemblages following a windstorm in French broadleaved temperate forests

Abstract: In most temperate deciduous forests, windstorm is the main source of dead wood. However, the effects of this natural disturbance on ambrosia and bark beetle communities are poorly known. In managed oak‐hornbeam forests storm‐damaged in France in 1999, we sampled ambrosia (and second bark beetles) by ethanol‐baited window‐flight traps in 2001. By comparing uncleared gaps, undisturbed closed‐canopy controls and seedling‐sapling stands, we investigated the short‐term effects of gap formation, gap size and surrounding landscape to provide a snapshot of scolytid response. Contrary to expectations, neither the abundance nor the richness of ambrosia beetle species was significantly higher in gaps than in undisturbed stands. Few responses in abundance at the species level and only a slight difference in assemblage composition were detected between gaps and closed‐canopy controls. Gaps were more dissimilar from seedling‐sapling stands, than from closed‐canopy controls. More scolytid individuals and species were caught in gaps than in seedling‐sapling stands. Mean local and cumulative richness peaked in mid‐size gaps. Only mid‐size gaps differed from closed‐canopy controls in terms of species composition. We identified generalist gap species (Xyleborus saxesenii, X. cryptographus), but also species significantly more abundant in mid‐size gaps (Platypus cylindrus, Xyloterus signatus). The faunistic peculiarity of mid‐size gaps seemed to be partly related to a bias in oak density among gap size classes. Few landscape effects were observed. Only the scolytids on the whole and X. dispar were slightly favoured by an increasing density in fellings at the 78 ha scale. We did not find any correlation between scolytid abundance and the surrounding closed‐forest percentage area. We confirmed that temperate, deciduous, managed stands did not come under threat by ambrosia and bark beetle pests after the 1999 windstorm. Nonetheless, our data stressed the current expansion in Western Europe of two invasive species, X. peregrinus and especially X. germanus, now the predominant scolytid in the three oak forests studied.     

湖南栎类天然次生林幼树更新特征及影响因子

以湖南典型栎类天然次生林为研究对象,基于51块样地的调查数据,采用k-means聚类分析划分林分类型,研究湖南不同栎类天然次生林幼树更新特征,分析了湖南不同栎类天然次生林幼树更新指标(幼树密度、幼树平均地径、平均高以及平均冠幅)与环境因子、林分因子的相关性,旨在阐明环境因子、林分因子对幼树更新的影响,以期为湖南不同栎类天然次生林的恢复与经营管理提供理论依据。结果表明:(1)利用聚类分析可将研究区内栎类天然次生林划分为5个类型,包括甜槠锥栗混交林(CC)、亮叶水青冈多脉青冈混交林(FC)、石栎樟树混交林(LC)、枹栎甜槠混交林(QC)、青冈栎混交林(CG)。(2)不同类型栎类天然次生林更新幼树优势种分化明显,物种丰富度差异显著(P0.05)。5种不同栎类次生林幼树密度均未超过500株/hm~2,更新情况较差;幼树数量差异显著(P0.05),为亮叶水青冈多脉青冈混交林石栎樟树混交林青冈栎混交林枹栎甜槠混交林甜槠锥栗混交林;生长情况差异显著(P0.05),为青冈栎混交林亮叶水青冈多脉青冈混交林枹栎甜槠混交林甜槠锥栗混交林石栎樟树混交林。(3)相关分析结果显示,不同类型次生林幼树更新的主要影响因子存在差异。甜槠锥栗混交林中幼树密度与腐殖质厚度呈显著负相关(P0.05);幼树平均高与灌木盖度呈显著正相关(P0.05);幼树平均地径与草本盖度、灌木盖度呈显著正相关(P0.05)。亮叶水青冈多脉青冈混交林中幼树密度与海拔、腐殖质厚度、枯落物厚度呈显著正相关(P0.05),与草本盖度呈极显著正相关(P0.01);幼树平均地径与郁闭度呈显著负相关(P0.05);幼树平均高、幼树平均冠幅与坡位呈显著正相关(P0.05)。石栎樟树混交林中幼树密度与坡向、土壤厚度呈显著正相关(P0.05),其余因子对幼树生长无显著影响。枹栎甜槠混交林中幼树密度与郁闭度、乔木密度呈极显著正相关(P0.01),与坡位呈显著负相关(P0.05);幼树平均冠幅与坡度呈显著负相关(P0.05)。青冈栎混交林中幼树平均地径与土壤厚度呈显著正相关(P0.05),与乔木密度呈极显著正相关(P0.01);幼树平均冠幅与灌木盖度呈显著正相关(P0.05)。     

Differential ecophysiological responses to seasonal drought of three co-existing oak species in northern Greece

Oaks (Quercus spp.) represent the most important broadleaf genus with respect to forest-shaping tree species in the Mediterranean. Considering future climate scenarios (increased drought conditions), the identification of drought tolerant oak species is of great importance for future forest management in this region. The objective of the study was the comparison of physiological status of three economically and ecologically valuable oak species (Quercus ilex, Quercus frainetto and Quercus pubescens) co-existing in natural coppice stands in NE Greece, in response to seasonal drought stress. Measurements were conducted between June and September 2016, every 15–20 days until leaf falling. The parameters studied were predawn leaf water potential and fast chlorophyll fluorescence induction curves (OJIP test), chlorophyll content, and relative water content. Meteorological data from the area were also collected. Photosynthetic parameters such as performance indices (PI_abs and PI_tot) reacted to summer drought conditions, with Q. frainetto showing the lowest values. The discrepancy between species increased with duration of drought period. Q. frainetto revealed the lowest predawn water potential values. The results indicate that Q. frainetto is less suitable for future forestry applications in the studied climate/elevation zone than Q. pubescens and Q. ilex.     

A nuclear DNA barcode for eastern North American oaks and application to a study of hybridization in an Arboretum setting

DNA barcoding has proved difficult in a number of woody plant genera, including the ecologically important oak genus Quercus. In this study, we utilized restrictionsite‐associated DNA sequencing (RAD‐seq) to develop an economical single nucleotide polymorphism (SNP) DNA barcoding system that suffices to distinguish eight common, sympatric eastern North American white oak species. Two de novo clustering pipelines, PyRAD and Stacks, were used in combination with postclustering bioinformatic tools to generate a list of 291 potential SNPs, 80 of which were included in a barcoding toolkit that is easily implemented using MassARRAY mass spectrometry technology. As a proof‐of‐concept, we used the genotyping toolkit to infer potential hybridization between North American white oaks transplanted outside of their native range (Q. michauxii, Q. montana, Q muehlenbergii/Q. prinoides, and Q. stellata) into a horticultural collection surrounded by natural forests of locally native trees (Q. alba and Q. macrocarpa) in the living collection at The Morton Arboretum (Lisle, IL, USA). Phylogenetic and clustering analyses suggested low rates of hybridization between cultivated and native species, with the exception of one Q. michauxii mother tree, the acorns of which exhibited high admixture from either Q. alba or Q. stellata and Q. macrocarpa, and a hybrid between Q. stellata that appears to have backcrossed almost exclusively to Q. alba. Together, RAD‐seq and MassARRAY technologies allow for efficient development and implementation of a multispecies barcode for one of the more challenging forest tree genera.     

Influence of rocky landscape features and fire regime on vegetation dynamics in Appalachian Quercus forests

Question : How do interactions between rocky landscape features and fire regime influence vegetation dynamics? Location : Continental Eastern USA. Methods : We measured vegetation, disturbance and site characteristics in 40 pairs of rocky and non‐rocky plots: 20 in recently burned stands, and 20 in stands with no evidence of recent fire (‘unburned’ stands). Two‐way analysis of variance (ANOVA) was used to assess the main and interaction effects of fire and rock cover on plant community composition. Results : In burned stands, rock cover had a strong influence on vegetation. Non‐rocky ‘matrix’ forests were dominated by Quercus, and had abundant ground cover and advance regeneration of early and mid‐successional tree species. Burned rocky patches supported greater density of fire‐sensitive species such as Acer rubrum, Sassafras albidum and Nyssa sylvatica and had little advance regeneration or ground cover. Quercus had fewer fire scars and catfaces (open, basal wounds) on rocky patches, suggesting that rocky features mitigate fire severity. In unburned stands, differences between rocky and non‐rocky patches were less distinct, with both patch types having sparse ground cover, little tree regeneration, and high understorey densities of relatively shade tolerant A. rubrum, N. sylvatica and Betula lenta. Conclusion : Under a sustained fire regime, heterogeneity in rock cover created a mosaic where fire‐adapted species such as Quercus dominate the landscape, but where fire‐sensitive species persisted in isolated pockets of lower fire severity. Without fire, species and landscape richness may decline as early‐mid successional species are replaced by more shade tolerant competitors.     

An Analysis of Vegetation Restoration on Opencast Oil Shale Mines in Estonia

We compared four types of 30‐year‐old forest stands growing on spoil of opencast oil shale mines in Estonia. The stand types were: (1) natural stands formed by spontaneous succession, and plantations of (2) Pinus sylvestris (Scots pine), (3) Betula pendula (silver birch), and (4) Alnus glutinosa (European black alder). In all stands we measured properties of the tree layer (species richness, stand density, and volume of growing stock), understory (density and species richness of shrubs and tree saplings), and ground vegetation (aboveground biomass, species richness, and species diversity). The tree layer was most diverse though sparse in the natural stands. Understory species richness per 100‐m² plot was highest in the natural stand, but total stand richness was equal in the natural and alder stands, which were higher than the birch and pine stands. The understory sapling density was lower than 50 saplings/100 m² in the plantations, while it varied between 50 and 180 saplings/100 m² in the natural stands. Growing stock volume was the least in natural stands and greatest in birch stands. The aboveground biomass of ground vegetation was highest in alder stands and lowest in the pine stands. We can conclude that spontaneous succession promotes establishment of diverse vegetation. In plantations the establishment of diverse ground vegetation depends on planted tree species.     

Tree growth response to drought and temperature in a mountain landscape in northern Arizona, USA

Aim To understand how tree growth response to regional drought and temperature varies between tree species, elevations and forest types in a mountain landscape. Location Twenty‐one sites on an elevation gradient of 1500 m on the San Francisco Peaks, northern Arizona, USA. Methods Tree‐ring data for the years 1950–2000 for eight tree species (Abies lasiocarpa var. arizonica (Merriam) Lemm., Picea engelmannii Parry ex Engelm., Pinus aristata Engelm., Pinus edulis Engelm., Pinus flexilis James, Pinus ponderosa Dougl. ex Laws., Pseudotsuga menziesii var. glauca (Beissn.) Franco and Quercus gambelii Nutt.) were used to compare sensitivity of radial growth to regional drought and temperature among co‐occurring species at the same site, and between sites that differed in elevation and species composition. Results For Picea engelmannii, Pinus flexilis, Pinus ponderosa and Pseudotsuga menziesii, trees in drier, low‐elevation stands generally had greater sensitivity of radial growth to regional drought than trees of the same species in wetter, high‐elevation stands. Species low in their elevational range had greater drought sensitivity than co‐occurring species high in their elevational range at the pinyon‐juniper/ponderosa pine forest ecotone, ponderosa pine/mixed conifer forest ecotone and high‐elevation invaded meadows, but not at the mixed conifer/subalpine forest ecotone. Sensitivity of radial growth to regional drought was greater at drier, low‐elevation compared with wetter, high‐elevation forests. Yearly growth was positively correlated with measures of regional water availability at all sites, except high‐elevation invaded meadows where growth was weakly correlated with all climatic factors. Yearly growth in high‐elevation forests up to 3300 m a.s.l. was more strongly correlated with water availability than temperature. Main conclusions Severe regional drought reduced growth of all dominant tree species over a gradient of precipitation and temperature represented by a 1500‐m change in elevation, but response to drought varied between species and stands. Growth was reduced the most in drier, low‐elevation forests and in species growing low in their elevational range in ecotones, and the least for trees that had recently invaded high‐elevation meadows. Constraints on tree growth from drought and high temperature are important for high‐elevation subalpine forests located near the southern‐most range of the dominant species.     

Overstory vegetation and successional trends of Land Between The Lakes,USA

133 stands from the uplands of Land Between The Lakes (LBL) were sampled. Stand data were ordinated to identify clinal patterns in the overstory vegetation. A coeno-cline, accounting for 52% of the species variance, was developed from the first ordination axis. Classification of stands identified nine dominance types: Pinus echinata-Quercus spp., Q. prinus, Q. prinus-Q. alba, Q. stellata-Q. alba, Q. velutina-Q. alba, Q. alba, Q. alba-Q. rubra, Acer saccharum-Q. alba-Fagus grandifolia sadF. grandifolia-A. saccharum. Compositionally-stable (climax) and unstable (successional) stands were analyzed separately. Trajectories of unstable stands include the succession of P. echinata to Quercus-dominated communities on xeric sites and of Quercus types to more mesophytic communities (e.g. A. saccharum and F. grandifolia) onmesic sites. These shifts indicate the nature of the future climax overstory composition at LBL. The first axis DCCA ordination for climax stands accounted for 45% of the vegetation variance. Factor Analysis identified two significant suites of variables related to vegetation: soil chemistry/soil moisture and soil texture. Specific variables related to variation in the climax vegetation include soil pH, calcium and magnesium, effective soil depth, available water capacity, percent rock in the soil, slope position, aspect, elevation and distance to the opposing slope. Modal positions of species along the direct gradient ordination were similar in rank order to those derived from the indirect gradient ordination of all stands. The latter ordination represents the present forest vegetation pattern at LBL. Climax stands showed a shift in modes of Quercus spp. toward the xeric end of the gradient and increases in habitat width of mesophytic species. Following a reduction in disturbance, species appear to adjust to changes in competition and available habitat. That is, the dimensions of realized niches (i.e. habitat widths) change with succession. The climax vegetation at LBL appears to show centrifugal organization.     

Contrasting physiological responsiveness of establishing trees and a C4 grass to rainfall events,intensified summer drought,and warming in oak savanna

Climate warming and drought may alter tree establishment in savannas through differential responses of tree seedlings and grass to intermittent rainfall events. We investigated leaf gas exchange responses of dominant post oak savanna tree (Quercus stellata and Juniperus virginiana) and grass (Schizachyrium scoparium, C₄ grass) species to summer rainfall events under an ambient and intensified summer drought scenario in factorial combination with warming (ambient, +1.5 °C) in both monoculture and tree‐grass mixtures. The three species differed in drought resistance and response of leaf gas exchange to rainfall events throughout the summer. S. scoparium experienced the greatest decrease in A_area (?56% and ?66% under normal and intensified drought, respectively) over the summer, followed by Q. stellata (?44%, ?64%), while J. virginiana showed increased A_area under normal drought (+13%) and a small decrease in A_area when exposed to intensified summer drought (?10%). Following individual rainfall events, mean increases in A_area were 90% for S. scoparium, 26% for J. virginiana and 22% for Q. stellata. The responsiveness of A_area of S. scoparium to rainfall events initially increased with the onset of drought, but decreased dramatically as summer drought progressed. For Q. stellata, A_area recovery decreased as drought progressed and with warming. In contrast, J. virginiana showed minimal fluctuations in A_area following rainfall events, in spite of declining water potential, and warming enhanced recovery. J. virginiana will likely gain an advantage over Q. stellata during establishment under future climatic scenarios. Additionally, the competitive advantage of C₄ grasses may be reduced relative to trees, because grasses will likely exist below a critical water stress threshold more often in a warmer, drier climate. Recognition of unique species responses to critical global change drivers in the presence of competition will improve predictions of grass–tree interactions and tree establishment in savannas in response to climate change.     

Relationships between the density of two potential restoration tree species and plant species abundance and richness in a degraded Afromontane forest of Kenya

In recent years, there have been considerable efforts to restore degraded tropical montane forests through active restoration using indigenous tree species. However, little is known about how these species used for restoration influence other species. In this study, two potential restoration species, Albizia gummifera and Neoboutonia macrocalyx, are investigated with regard to the relationship between their density and the abundance and richness of other plant species. The study was conducted in a degraded forest consisting of disturbed transition zones and secondary forest. Our results show positive relationships between the density of A. gummifera and the abundance of tree seedling and sapling richness in the transition zones and in the secondary forest. Shrub richness was negatively related to the density of A. gummifera. Abundance and richness of tree saplings and shrubs were positively related to N. macrocalyx density both in the transition zones and in the secondary forest. Herb species richness declined with N. macrocalyx density in the transition zones but increased with N. macrocalyx density in the secondary forest. The positive relationships between the density of the two tree species and species richness of other woody species suggest that both A. gummifera and N. macrocalyx can be suitable for active restoration of degraded mountain forests within their natural range.     

Factors affecting species richness of tree regeneration in mixed‐wood stands of central Maine

Question: Two questions about within‐stand spatial variability are addressed in this paper. How does species richness of tree regeneration respond to small‐scale ecological gradients, and what effect does natural Abies balsamea abundance have on the species richness of other tree regeneration? Location: A long‐term, gap‐silviculture experiment, Acadian mixed‐wood forest, Maine, USA. Methods: Eight stands treated with and without gap harvesting were sampled to capture sub‐stand heterogeneity of understorey tree regeneration concurrently with patterning of local stand conditions. Spatial and non‐spatial models were developed to test the relationships between two response variables [species richness of small (height ≥0.1 m, but <0.75 m) and large (height ≥0.75 m, but <1.4 m) regeneration] and five explanatory variables (depth to water table, percentage canopy transmittance, A. balsamea regeneration density, and overstorey basal area and species richness). Results: Despite high unexplained variance for all models, consistent associations among variables were found. Negative associations were found between: (1) the species richness of small regeneration and A. balsamea regeneration density and (2) the species richness of large regeneration and overstorey basal area. Positive associations were found between: (1) the species richness of small regeneration and both overstorey basal area and species richness and (2) the species richness of small and large regeneration and canopy transmittance. Conclusions: Promoting tree species diversity in Acadian mixed‐wood stands may not be achievable through the use of gap‐harvesting alone if the density of understorey Abies balsamea is not reduced either naturally or through silvicultural intervention.     

Oak trees and soil interactions in Mediterranean forests: a positive feedback model

Questions: What is the spectrum of variability of chemical elements in a Mediterranean forest ecosystem across the different compartments? Do co‐existing tree species with different leaf chemical composition and nutrient cycling distinctly modify soil conditions? Could these species‐specific, tree‐generated soil changes create a potential positive feedback by affecting long‐term species distribution? Location: Mixed oak forests of southern Spain, Los Alcornocales Natural Park. Methods: We sampled and chemically analysed five different ecosystem components: leaves, leaf fall, litter and superficial (0–25 cm) and sub‐superficial (25–50 cm) soil beneath the canopies of evergreen Quercus suber and deciduous Q. canariensis trees. We used multiple co‐inertia analysis (MCoA) to conjointly analyse the patterns of variability and covariation of eight macro‐ and micronutrients determined in each of the sampled ecological materials. We implemented a path analysis to investigate alternative causal models of relationships among the chemical properties of the different ecosystem components. Results: Variability in the concentration of chemical elements was related to the nature of their biogeochemical cycles. However, the rank of element concentration was consistent across ecosystem components. Analysis of co‐inertia (MCoA) revealed that there was a common underlying multivariate pattern of nutrient enrichment in the ecosystem, which supported the hypothesis of a separation in biogeochemical niches between the two co‐existing oak species, with Q. canariensis having richer plant tissues and more fertile soil directly under each tree than Q. suber. The feasibility of a potential tree–soil positive feedback model was the only statistically validated among several alternative (non‐feedback) models tested. Conclusions: In the studied Mediterranean forests, oak species distinctly modify soil fertility conditions through different nutrient return pathways. Further investigation is needed to address whether these tree‐generated soil changes could affect seedling establishment and ultimately influence species distribution.     

Radial growth and climate responses of white oak (Quercus alba) and northern red oak (Quercus rubra) at the northern distribution limit of white oak in Quebec, Canada

Aim The objectives of this study were: (1) to compare radial growth patterns between white oak (Quercus alba L.) and northern red oak (Quercus rubra L.) growing at the northern distribution limit of white oak; and (2) to assess if the radial growth of white oak at its northern distribution limit is controlled by cold temperature. Location The study was conducted in three regions of the Ottawa valley in southern Québec. All stands selected were located at the northern limit of distribution of Q. alba. Methods Twelve mixed red and white oak stands were sampled and increment cores were extracted for radial growth analyses. For each oak species, 12 chronologies were derived from tree‐ring measurement (residual chronologies). Principal components analysis and redundancy analysis were used to highlight the difference between radial growth in both species and to determine their radial growth–climate association. Results There was little difference between the radial growth of each species; Q. alba, however, exhibits more year‐to‐year variation in growth than Q. rubra. More than 65% of the variance in radial growth was shared among sites and species. Both species showed a similar response to climate, which suggested that the limit of distribution of Q. alba might not be determined by effects on growth. Both species had a classic response to climate and drought in the early growing season. Main conclusions The northern distribution limit of Q. alba does not appear to be directly controlled by effects on growth processes as indicated by the similarities in radial growth and response to climate between the two species. The location of the stands on southern aspects suggested that cold temperature could have been a major factor controlling the distribution limit of Q. alba. However, it is speculated that stands growing on southern aspects may be more prone to forest fires or to drought, which would favour the maintenance and establishment of oaks, and of Q. alba in particular. Models relating the northern distribution limits of species to broad climate parameters like annual mean temperature will need to be reviewed to incorporate more biologically relevant information. Such assessments will in turn provide better estimates of the effect of climate changes on species distribution.     

Impacts of drought on tree mortality and growth in a mixed hardwood forest

Abstract. The tree and shrub species on a 16-ha watershed in the Coweeta Basin were sampled in 1984 and again in 1991 to determine the effects of drought on tree species composition and basal area growth. Mortality and radial growth were determined for tree species within three community types that represent a moisture gradient from moist to dry: cove-hardwoods > mixed-oak > oak-pine. Tree mortality from 1984 to 1991 was 20% and 23% in the cove-hardwoods and mixed-oak communities, respectively, compared to only 12% in the oak-pine type. With the exception of Oxydendrum arboreum and Robinia pseudoacacia, the oaks had higher percentage mortality than any other genus; Quercus velutina had 29%, 37%, and 20% mortality in the cove-hardwoods, mixed-oak, and oak-pine types, respectively; Quercus prinus had 23% mortality in the mixed-oak type; Quercus coccinea had 36% mortality in the mixed-oak type; and Quercus marilandica had 27% mortality in the oak-pine type. Mortality occurred mostly in the small-size class individuals (< 10 cm in diameter) for all species, suggesting that thinning was still an important process contributing to mortality 29 yr after clearcutting. Although growth of Liriodendron tulipifera was much higher than growth of either Quercus prinus or Quercus coccinea, growth in Liriodendron was significantly reduced by the 1985–88 drought and no growth reduction was observed for these two dominant Quercus species during the same time period.