Hybridization between Quercus robur and Q. petraea in a mixed oak stand in Denmark

? Hybridization and mating pattern between Quercus robur and Q. petraea was studied in a 5.8 ha mixed forest stand in Jutland, Denmark which comprises in total 135 Quercus robur and 230 Q. petraea trees. Classification of the oak trees into species was performed using canonical discriminant analysis of a range of leaf morphological traits. Adult trees (365) and offspring (582) were genotyped with eight microsatellite markers. Seedlings were sampled in 2003 and acorns were collected in 2004.

? Mating patterns of Q. robur and Q. petraea are expected to be different in the northern range of the distribution area and a larger hybridization rate is expected. It is further expected, that pollination from outside sources will be relatively less in small fragmented forest management systems compared to large scale oak forest. The conclusions should be verified through repeated year to year analysis of the mating pattern.

? Phenological studies revealed that there was no difference in flowering time between species. Data for the adult trees revealed no significant departures from Hardy-Weinberg proportions and there was weak, but significant spatial genetic structure, which supports the idea that the stand is of natural origin. Spatial genetic structure in the first distance class is stronger for Q. petraea. The genetic composition of the offspring was remarkably consistent from year to year. Paternity analysis revealed that, on average, 85% pollination came from fathers within the stand. The direction of the pollen flow varied from year to year. Inter-specific hybridization was high and ranged from 15–17% and from 48–55% for Q. petraea and Q. robur mothers respectively. Paternity analysis revealed that the population was basically outcrossing and only 3.7% of the analysed progeny were the product of selfing. Over the two years of study, approximately 200 trees contributed to the paternity of the next generations.

? The study confirms earlier studies with a greater tendency for Q. robur mothers to produce hybrid seeds than Q. petraea mothers. The rate of hybridization is higher in this Danish stand than in comparable studies elsewhere in Europe. Gene flow from outside sources are relatively low.

     

Variation in wood density and carbon content of tropical plantation tree species from Ghana

Most research on carbon content of trees has focused on temperate species, with less information existing for tropical trees and very little for tropical plantations. This study investigated factors affecting the carbon content of nineteen tropical plantation tree species of ages seven to twelve and compared carbon content of Khaya species from two ecozones in Ghana. For all sample trees, volume of the main stem, wood density, wood carbon (C) concentration and C content were determined. Estimated stem volume for the 12-year-old trees varied widely among species, from 0.01 to 1.04 m³, with main stem C content ranging from 3 to 205 kg. Wood density among species varied from 0.27 to 0.76 g cm^?3, with faster growing species exhibiting lower density. Significant differences in wood density also occurred with position along the main stem. Carbon concentration also differed among tree species, ranging from 458 to 498 g kg^?1. Differences among species in main stem C content largely reflected differences among species in estimated main stem volume, with values modified somewhat by wood density and C concentration. The use of species-specific wood density values was more important for ensuring accurate conversion of estimated stem volumes to C content than was the use of species-specific C concentrations. Significant differences in wood density did exist between Khaya species from the wet and moist semi-deciduous ecozones, suggesting climatic and site factors may also need to be considered. Wood densities for these plantation grown trees were lower than literature values reported for the same species in natural forests, suggesting that the application of data derived from natural forests could result in overestimation of the biomass and C content of trees of the same species grown in plantations.     

The stand dynamics of a mixed coppice forest of shade-tolerant and intermediate species

We investigated stand dynamics of a mixed coppice forest of shade-tolerant and intermediate species, Fagus crenata Bl. and Quercus crispula Bl., in five stands with canopy-stem densities from 420 to 1320 ha⁻¹ (estimated stand age: 37–62 years). The mean basal area (BA), exposed-crown area (ECA) and crown depth (CD) per stem were significantly larger in F. crenata in the three lower density plots. Also Q. crispula had considerably less overtopped stems in the two lower density plots. The mean annual BA increments of the two species were similar in the earlier 20 years; however, in the most recent 5 years, F. crenata had a higher relative growth rate for basal area (RGR_BA) than Q. crispula. The most critical variable of the RGR_BA was the ECA for F. crenata and the relative height (RH) for Q. crispula. We predict that dominance of F. crenata will gradually increase over that of Q. crispula as stands mature. This seems to be caused by lower shade-tolerance despite the similar initial growth rates of Q. crispula compared with F. crenata.     

Tree mortality in a natural mixed forest affected by stand fragmentation and by a strong typhoon in northern Japan

Windthrow is recognized as an extremely significant disturbance in many forests. Its effects are increased by stand fragmentation, which exposes the stand to strong winds. In this study, we investigated the change in tree mortality in fragmented stands with the distance from the stand edge, in a conifer–broadleaved mixed forest in northern Japan. We set out experimental plots having three stand sizes (400, 1,600, 6,400 m²) and examined the stand dynamics over 10 years. Tree mortality tended to be higher in smaller stands, although an effect of stand size was found only in the first 5 years of the study. Distance-dependent individual mortality was obvious in Abies sachalinensis, the most vulnerable major tree species, again in the first 5 years, suggesting that wind-risk management should emphasize the area within ca. 20 m from the stand edge. No distant-dependent effect was found, however, in the latter 5 years, in which there was a strong typhoon; tree deaths occurred throughout the stand (irrespective of distance within) as a result of this event. We conclude that the severity of wind can cause the features of wind-induced damage to differ; stand edge effects peculiar to a small forest are unlikely to occur with particularly strong winds, and the effect of fragmentation might therefore be clear only in weaker disturbances.     

Variation in log quality and prediction of sawing yield in oak wood (Quercus robur)

Context

The commercial feasibility of sawmilling depends on the expected volume and value of sawn planks. Models that predict the volume of sawn timber of a particular quality and produced from logs of known characteristics are therefore very useful.

Aims

The objectives were to study variation in sawing yield and to obtain models that predict lumber volume and grade recovery on the basis of easy-to-measure predictor variables of saw logs.

Methods

Forty-six oak trees growing in Galicia (NW Spain) were felled and cut into logs. The logs were visually graded and sawn mainly into quartersawn planks, which were dried, planed and visually graded for structural purposes.

Results

The total volumetric sawing yield was 47.6 %. The sawing yield for planks of structural dimensions (cross-section, 70?×?120 or 70?×?170 mm) was 43.4 %, but decreased to 8.4 % for structural sized and quality grade beams because of wane and biotic damage in many pieces. Log grade did not significantly affect sawing yield in the sample analysed, despite the wide range of diameter over bark at the smallest end in the sampled logs (22–77 cm). The sawing pattern affected total sawing yield (F?=?4.913; p value?=?0.001) and the sawing yield for structural planks (F?=?6.142; p value?=?0.0002); radial sawing with one cut and live sawing of half logs provided the highest yields. Three models were proposed for estimating sawn volume in timber products, with the small-end log diameter over bark as the predictor variable and R _adj ² between 0.31 and 0.78 (p value?<?0.01).

Conclusion

For the purpose of producing oak timber destined for structural use, the presence of bark and sapwood in planks must be reduced in the sawing process; this would decrease the total lumber recovery but increase the timber value yield. Air drying must be accelerated to reduce biotic damage in sawn planks. Geometric mean diameter over bark at the smallest end (d) outperforms other measures as a predictor variable for total or structural sawn timber volume.     

Environmental and plant-based controls of water use in a Mediterranean oak stand

In this work, new information is reported on water relations of the Mediterranean oak species Quercus pyrenaica based on environmental and physiological measurements carried out during the growing seasons of 2006 and 2007. The interest in this species has increased due to its use in reforestation programs and its impacts on the water resources due to the extensive spontaneous afforestation after the abandonment of forest (firewood, charcoal, livestock, etc.) and agricultural activities, in general in some areas in the Mediterranean region and in particular, in the studied area (Sistema Central range, Spain). The objectives were to evaluate the long-term water use of this stand and its limit and to analyse the specific traits to cope with summer drought, especially the use of stem water storage and deep soil water reserves. Tree water stress associated with depletion of soil water reserves was not observed since the oak trees appeared to avoid a marked water stress using water reserves from deeper soil layers as summer drought progresses. The contribution of mean daily stem water storage to transpiration was low (4%), although it could be greater under dryer conditions. Only at the end of summer of 2006, the transpiration and canopy conductance were reduced due to soil drought. Despite the absence of marked water stress an upper limit was found in transpiration (slightly higher than 3 mm day⁻¹). The heavy use of soil water resources by this species (75% of available soil water in this study) should be considered when evaluating the impact of spontaneous afforestation and reforestation programs on water resources.     

Forest stand dynamics and sudden oak death: Mortality in mixed-evergreen forests dominated by coast live oak

Sudden oak death (SOD), caused by the recently discovered non-native invasive pathogen, Phytophthora ramorum, has already killed tens of thousands of native coast live oak and tanoak trees in California. Little is known of potential short and long term impacts of this novel plant–pathogen interaction on forest structure and composition. Coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica) form mixed-evergreen forests along the northern California coast. This study measured tree mortality over a gradient of disease in three time periods. Direct measurements of current mortality were taken during 2004, representing a point-in-time estimate of present and ongoing mortality. Past stand conditions, c. 1994, were estimated using a stand reconstruction technique. Future stand conditions, c. 2014, were calculated by assuming that, given a lack of host resistance, live trees showing signs of the disease in 2004 would die. Results indicate that coast live oaks died at a rate of 4.4–5.5% year⁻¹ between 1994 and 2004 in highly impacted sites, compared with a background rate of 0.49% year⁻¹, a ten-fold increase in mortality. From 2004 to 2014, mortality rates in the same sites were 0.8–2.6% year⁻¹. Over the entire period, in highly impacted sites, a 59–70% loss of coast live oak basal area was predicted, and coast live oak decreased from 60% to 40% of total stand basal area, while bay laurel increased from 22% to 37%. Future stand structures will likely have greater proportions of bay laurel relative to coast live oak.     

Changes in the spatial structure of oak carbon-based secondary compounds after pine harvesting

In natural plant populations, leaf polyphenols show high intraspecific variation that occurs both temporally and spatially. Leaf phenolics may be induced by diverse ecological factors such as light, nitrogen availability or herbivory attack. Both light and nitrogen availability can show spatial structure in forested stands, meaning that they each have a high degree of autocorrelation, which can determine the appearance of spatial structure in leaf polyphenols. However, the availability of these resources may be drastically changed by forest disturbance, and little is known about the effect of forest disturbance on the spatial pattern and scale of leaf secondary compounds. We hypothesise that the spatial structure of leaf polyphenols in understory vegetation will disappear due to forest harvesting, because these compounds depend on light availability, yet it will remain unaltered for those compounds that either depend on the availability of other resources or are under major genetic control. The study was performed in young pedunculate oak (Quercus robur) populations growing either under a pine canopy (Pinus pinaster) stand or in a pine harvested stand in NW Spain. The spatial structures of green and senescent leaf polyphenols, tannins, non-tannin polyphenols and nitrogen were analysed in both stands using geostatistical analysis. The spatial structures observed for green and senescent leaf polyphenols and tannins in the forested stand disappeared in the harvested stand. However, non-tannin polyphenols, as well as nitrogen, showed spatial structure in both stands. Understanding these changes may be important for the successful recovery of native oak populations growing under pine forests in NW Spain, one of the priorities of the local government. Our results showed that changes in the concentration of leaf secondary compounds after disturbance may be accompanied by differences in their spatial properties, which may have important consequences for ecosystem function.     

Unexpected disproportion observed in species composition between oak mixed stands and their progeny populations

We used morphological analysis to assess species composition of natural regenerations and progeny plantations established from two mixed oak stands in Jamy and Legnica, Poland. Despite equal proportions of pedunculate (Quercus robur) and sessile oak (Quercus petrea) in seed stands, the species composition differed strikingly. In all progeny populations, pedunculate oak dominated, reaching 89.5% and 96.6% in Legnica and Jamy, respectively. However, sessile oak predominated in natural regenerations. Morphological studies indicated a varied number of phenotypically intermediate or mosaic individuals. Among artificial populations, the highest number of putative hybrids was observed in Legnica (average 2.5%) and the lowest in Jamy (average 0.2%). Hybrids in natural regeneration were 1% in Legnica and 8% in Jamy. The disproportionate species composition could result from either unintentional indirect acorn selection during collection or selection in nursery practice. We discuss the role of ecophysiological differences between species in biased species representation in progeny populations.     

Integrating species and successional classes for wood production in a mixed forest restoration system in a neotropical region

As forests in neotropical regions,particularly in developing countries,are devastated,interventions to restore biodiversity and its ecological functions are nee...     

Role of waterlogging-responsive genes in shaping interspecific differentiation between two sympatric oak species

Pedunculate (Quercus robur L.) and sessile oak (Quercus petreae Matt. Liebl.) are closely related species with a widely sympatric distribution in Europe. These two oak species are also known to display different ecological features, particularly related to their adaptation to soil waterlogging. Pedunculate oak grows in humid areas and can withstand high moisture content of the soil, whereas sessile oak requires drier soil with better drainage. The main goal of this study was to explore the role of gene expression contributing to differences in terms of waterlogging tolerance between these two species. We implemented a series of experiments aimed at evaluating whether differentially expressed genes between species are associated with their ecological preferences and underlie adaptive genetic divergence. Rooted cuttings of both species were grown in hydroponic conditions and subjected to gradual root hypoxia. White roots were sampled after 6, 12, 24 and 48 h. Real-time polymerase chain reaction (qPCR) was first used to monitor the expression of 10 known waterlogging-responsive genes, to identify discriminating sampling time points along the kinetics of hypoxia. Secondly, four subtractive suppressive hybridization libraries (sessile vs. pedunculate, pedunculate vs. sessile for early and late responses) were generated to isolate differentially expressed genes between species. A total of 2160 high-quality expressed sequence tags were obtained and annotated, and a subset of 45 genes were selected for qPCR analysis in a second independent factorial experimental design applying two stress durations per two species. Significant differences of gene expression between pedunculate and sessile oaks were detected, suggesting species-specific molecular strategies to respond to hypoxia. This study revealed that the ability of pedunculate oak to maintain glycolysis and fermentation under hypoxic conditions may help explain its tolerance to waterlogging.     

Effect of gap size on mid-rotation stand structure and species composition in a naturally regenerated mixed broadleaf forest

This paper is an assessment of the effect of gap size on stand structure and species composition 48 years following treatment in a mixed broadleaf upland forest. Established in 1960, the study tests three circular openings, 15.2 m (0.02 ha), 45.7 m (0.16 ha), and 76.2 m (0.46 ha). Forty-eight years following treatment (2008) basal area, top height, and quadratic mean diameter were significantly lower in 15.2 m openings. Maple (Acer spp.) species had the highest mean importance value across treatments (0.40). Trends suggest that species composition of dominant and codominant trees among opening sizes may have been influenced by shade tolerance adaptations of the species groups present. Whereas 15.2 m openings were dominated by shade tolerant maple species, 45.7 and 76.2 m openings produced a mixture of commercial species including shade intolerant species such as yellow-poplar (Liriodendron tulipifera L.), trees of intermediate shade tolerance like oak (Quercus spp.), and shade tolerant maple. Data further suggest the density of overstory oak was highest in the intermediate opening size (45.7 m), while yellow-poplar increased in the larger opening sizes. Evaluation of species shifts between 1981 and 2008 showed that relative basal area of maple increased across all treatments. Relative basal areas from 45.7 to 76.2 m openings suggest declines in yellow-poplar and other non-commercial species were balanced by increases in oak and maple.     

Crack propagation in mixed mode in wood: a new specimen

Summary For studies on wood structures it is important to have a reliable specimen for mixed mode loading (I+II) of a crack. A new type of specimen derived from the compact tension shear (CTS) specimen proposed by Richard for isotropic materials has been studied. A finite element modelization for pine (Pinus pinaster) gives calibrations in the RL, TL and TR directions. Fracture experiments show a fair agreement between stress intensity factors obtained by calibrations and by the compliance method.     

A stand density index for complex mixed species forests in the northeastern United States

Quantifying stand density is important for accurate prediction of net biomass and carbon accumulation, for estimating growth and mortality risks of trees, stands, and regions, and for the management of forests for multiple goods and services. Building on previous work relating maximum stand density to wood specific gravity, we develop a stand density equation for the mixed species forests of the northeastern United States using data from the US Forest Service, Forest Inventory and Analysis program. We used quantile regression, in concert with a quantile selection and evaluation procedure, to ensure conformity between our density measure and previously developed guidance for well-studied stand types. The resulting strictly additive relative density measure appears to provide reasonable prediction of maximum density even for plantations of exotic conifers in the region. The results suggest that maximum stand densities after accounting for wood specific gravity may be lower in northeastern North America than in the south or west.     

Variation in density and moisture content of wood and bark among twenty Eucalyptus grandis progenies

Summary Variation in density and moisture content of Eucalyptus grandis wood and bark is described utilizing the trees established for a half-sib progeny test in southern Florida. Differences in density and moisture content among progenies were significant. Variations in wood properties were largely due to the genetic differences. No relationship was found between wood density and wood moisture content. Implications for genetic manipulation of wood quality for this species were discussed.     

Corresponding patterns of site quality,decline and tree growth in a sessile oak stand

This study focuses on two rarely studied aspects of oak decline: relations with site characteristics and effects on tree growth. The study was carried out in a 5.5 ha stand in Hungary which is strongly affected by oak decline. The nearly pure sessile oak (Quercus petraea) stand of mostly coppice origin was 90 years old at the beginning of the study. Within-stand site heterogeneity was described by the herbaceous vegetation. Four ecological site types were distinguished by the species composition of herbs, and characterized by the ecological indicator values of the species. Tree growth between 1987 and 1993 was measured, and tree vigour was estimated from visual characteristics five times between 1987 and 1993. Potential volume increment of declining trees was estimated with the growth rates of healthy trees of the same size. Volume increment loss caused by oak decline was also assessed. Significant positive relationships were found between tree vigour and tree size and between tree vigour and tree growth. The growth of seriously declining trees dropped to almost one-half of that of healthy ones. Growth reduction of living trees at the stand level amounted to 5.4%, whereas growth reduction of all trees, including those that died during the observation period, amounted to 19.9% of the potential growth. Tree size and growth were greater on better sites. A strong relationship was also found between tree vigour and site type, but sessile oak was more susceptible to decline at better sites.     

Belowground competition in a broad-leaved temperate mixed forest: pattern analysis and experiments in a four-species stand

We investigated fine root biomass and distribution patterns in a species-rich temperate Carpinus–Quercus–Fagus–Tilia forest and searched for experimental evidence of symmetry or asymmetry in belowground competition. We conducted extensive root coring and applied the recently introduced in situ-root growth chamber technique for quantifying fine root growth under experimentally altered intra- and interspecific root neighbourhoods in the intact stand. In 75% of all soil cores, fine roots of more than two tree species were present indicating a broad overlap of the root systems of neighbouring trees. Quercus trees had more than ten times less fine root biomass in relation to aboveground biomass or productivity (stem growth) and a much higher leaf area index/root area index ratio than Carpinus, Fagus and Tilia trees. The root growth chamber experiments indicated a high belowground competitive ability of Fagus in interspecific interactions, but a low one of Quercus. We conclude that (1) interspecific root competition is ubiquitous in this mixed stand, (2) root competition between trees can be clearly asymmetric, and (3) tree species may be ranked according to their belowground competitive ability. Fagus was found to be the most successful species in belowground competition which matches with its superiority in aboveground competition in this forest community.     

Armillaria species distribution on symptomatic hosts in northern hardwood and mixed oak forests in western Massachusetts

Armillaria spp. are some of the most important forest pathogens in mixed hardwood forests of southern New England, yet their role as prominent disturbance agents is still not fully appreciated. We investigated the distribution of Armillaria species across eight separate stands of northern hardwood and mixed oak forests in western Massachusetts. We were specifically interested in the Armillaria species distribution from live, symptomatic hosts and not in determining overall incidence in the forest. From 32 plots (16 within each forest type), 320 isolates were collected. Armillaria was routinely encountered causing disease of live trees. In total, 89% (286/320) of all isolations came from live hosts exhibiting symptoms of root and butt rot. Overall, A. gallica was the dominant species in each forest type, making up 88/160 (55%) isolates from northern hardwood and 153/160 (96%) of all isolations from mixed oak stands. However, northern hardwood forests showed much greater species diversity, as A. calvescens, A. gemina, A. ostoyae, and A. sinapina were all found. At one site, a northern hardwood forest surrounding a high elevation spruce-fir forest, A. ostoyae was the most abundant species encountered. All five Armillaria species were found causing disease of live hosts, including A. gemina, a species considered by some as weakly virulent. Armillaria gallica was found on 22/23 tree species’ sampled, and was found most often causing butt rot.     

Analysis and modeling of spatial stand structures. Methodological considerations based on mixed beech-larch stands in Lower Saxony

The first part of this paper highlights spatial stand structure as the central stand characteristic and introduces methods of pattern identification. This involves two nearest-neighbour methods for the identification of stand structures, i.e., the aggregation index R by Clark and Evans [Clark, Ph.J. and Evans, F.C., 1954. Distance to nearest neighbor as a measure of spatial relationships in populations. Ecology 35 (4) 445–453.] for univariate patterns and the segregation index S by Pielou [Pielou, E.C., 1977. Mathematical Ecology. Wiley.] for bivariate patterns. Both were used to describe the structure of 53 experimental plots of mixed beech-larch stands in Lower Saxony which provided the data base for this investigation. The second part of the study deals with the development of the STRUGEN stand structure generator, designed for the modeling and reproduction of spatial stand structures. To generate stand structures, a two-dimensional homogeneous Poisson process is used as well as a set of two-dimensional distribution functions which determine mixture type and intermingling intensity of main and associated tree species. Moreover, a distance function secures minimum distances between competing neighbouring trees. Consequently, the produced pattern is the result of a combination of an inhomogeneous Poisson process (for generating mixture units) and a hard-core process (for securing minimum distances between neighbours). The STRUGEN generator was designed and successfully used for the investigation of 53 mixed beech-larch stands. It provides initial values and stand structures for distance-dependent single-tree models from estimated qualitative stand characteristics. STRUGEN is a useful tool and allows initial, pragmatic steps towards fully utilising available qualitative and quantitative information to diagnose the state of a forest and to predict its growth.