Maternally and paternally inherited molecular markers elucidate population patterns and inferred dispersal processes on a small scale within a subalpine stand of Norway spruce (Picea abies [L.] Karst.)

The within-population spatial structure of genetic diversity is shaped by demographic processes, including historical accidents such as forest perturbations. Information drawn from the analysis of the spatial distribution of genetic diversity is therefore inherently linked to demographic-historical processes that ultimately determine the fate of populations.All adult trees and saplings in a 1.4-ha plot within a mixed Norway spruce (Picea abies [L.] Karst) stand were characterised by means of chloroplast (paternally inherited) markers, and a large sub-sample of these were genotyped at mitochondrial (maternally inherited) molecular markers. These data were used to analyse the spatial distribution of genetic variation and to compare the patterns corresponding to the two marker types. The plot presented non-homogeneous local stem density in the younger cohorts, and the indirect effect of this source of variation on the spatial genetic structure was investigated. Results suggest that (i) spatially limited seed dispersal induced patchiness in genotype distribution, while pollen flow had a homogenising effect; (ii) deviations from random spatial structure were stronger in the demographically most stable portions of the stand, while they were weaker where sudden bursts of regeneration occurred; (iii) spatially overlapping adult and sapling cohorts displayed the same spatial genetic structure (stronger on stable areas, weaker in portions of the stand undergoing events of intense regeneration), which was substantiated by the influence of local demographic processes. Regeneration dynamics as modulated by demography thus influences the distribution of genetic diversity within the stand both in the younger life stages and in the adult population.     

Fine-scale analysis reveals a potential influence of forest management on the spatial genetic structure of Eremanthus erythropappus

Forest management may have significant effects on forest connectivity and natural population sizes.Harvesting old-growth single trees may also change natural patterns of genetic variation and spatial genetic structure.This study evaluated the impacts of forest management using a silvicultural system of seed trees on the genetic diversity and spatial genetic structure of Eremanthus erythropappus(DC.)MacLeish.A complete survey of 275 trees on four plots was undertaken out to compare the genetic variation of a managed stand with an unmanaged stand.We genotyped all adult and juvenile individuals 60 months after the management and compared the genetic diversity and the spatial genetic structure parameters.Genetic diversity was considered high because of an efficient gene flow between stands.There were no genetic differences between stands and no evidence of inbreeding.Genetic clustering identified a single population(K=1),indicating no genetic differentiation between managed and unmanaged stands.Adult and juvenile individuals of the unmanaged stand were more geographically structured than individuals from the managed one.There was a tendency of coancestry among juveniles at the first class of distance of the managed stand,suggesting a drift of genetic structure possibly caused by management.Understanding early responses to management on genetic diversity and stand structure is a first step to ensuring the effectiveness of conservation practices of tree species.The sustainability of forest management of E.erythropappus on genetic diversity,and more accurately,on spatial genetic structure needs evaluation over time to promote effective conservation of the population size and genetic variability.     

A stochastic spatial regeneration model for Pinus sylvestris

A simulation model of pine regeneration is presented, based on earlier models and research for natural regeneration of Scots pine in Finland. The model is stochastic, reflecting the unpredictable variation in regeneration results observed in nature. Geographic area, site and stand characteristics and soil preparation affect the simulated regeneration result. The model predicts the amount and quality of seed crop, germination and predation of seeds, and survival and growth of seedlings. The effect of parent trees on seed dispersal, and growth and survival of seedlings is accounted for. At the stand level, the model gives results similar to empirical measurements. The simulation example provided illustrates the effect of density and spatial distribution of the parent tree stand on the regeneration result.     

Gene dispersion within a population of Pinus sylvestris

Gene dispersion within a population was studied in a 2‐hectare stand of 74 seed trees and 1100 seedlings. Isozyme markers were used to determine the genetic relationships among parental trees, embryos and seedling progenies. Paternity exclusion analyses were used to study multilocus progeny genotypes from certain parents. Results demonstrate large variation between trees in their capacity to contribute their genes through natural regeneration. The proportion of all genes dispersed into a specific distance interval from the source was estimated. The estimation procedure took into account the background of seedlings where parenthood was not excluded, but where the seedlings turned out to be unrelated. We found little evidence for clusters of progeny seedlings surrounding their parental tree. Within 15 m from a tree, approximately 5 % of the genes originated from that tree and 95 % originated from other trees. We found no evidence for an increase in levels of inbreeding, or genetic substructuring in the seedling progeny stand relative to the parental seed trees. A mathematical model for pollen distribution was developed. According to this model, 90% of pollen was distributed more than 15 m from the source. It is concluded that the marker seed trees had a surprisingly low detectable genetic contribution to the regeneration. It seems likely that a considerable part of the new seedlings have parents among felled trees or from surrounding stands.     

Tolerance of broadleaved tree and shrub seedlings to preemergence herbicides

Isoenzymes were used to study the genetic variation of 121 mature trees of Sorbus torminalis in a natural stand and a subset of 21 high yielding seed trees in the same stand used for a commercial harvest. Eight enzyme systems were studied. The genetic variation of the mature stand was higher than that of other stands reported in the literature. However, this level of genetic variation was not represented to the same extent by the seeds of the commercial harvest. Differences between the bud sample taken from the mature stand and the seed sample taken from the commercial subset were particularly obvious with respect to diversity measures and heterozygosities. The hypothetical gametic multilocus diversity (HGMD) of a bud sample of the mature Sorbus torminalis trees was more than 70% higher (94.1) than the value of the seed harvest sample (55.1). Deviations between the genetic structures of both samples were statistically significant. There is no indication that inbreeding is an essential component of the mating system of Sorbus torminalis. The potential causes for the lack of Sorbus torminalis seedlings and saplings originated from seeds and their consequences for regenerating this species are discussed. This research underscores that sampling a forest tree population for commercial harvest can result in a statistically significant reduction in the genetic diversity of the sampled group relative to the overall population.     

Genetic aspects of artificial regeneration of Douglas-fir (Pseudotsuga menziesii) in Bavaria

Douglas-fir has been planted in Bavaria for over 100 years. Its high growth potential and good wood quality combined with its silvicultural characteristics have increased interest in the species which can easily be integrated into existing natural ecosystems. In order to be successful in the establishment of Douglas-fir one must select the best provenance (only the so called “green” or viridis coastal type form is suitable for Bavaria), maintain a broad genetic base and carry out proper silviculture. The goal of the study was to check the origin/provenance (coastal or interior type) of older Bavarian Douglas-fir stands, the respective genetic variation of the stands and also the natural regeneration, the collected seed and the plants raised in the nursery. Possible changes in the genetic composition and diversity due to selective processes in the nursery phase were also investigated. Isoenzyme gene markers were used for the study. The majority of the investigated older stands apparently belong to the coastal type Douglas-fir; however, we also found mixed stands and stands solely of the interior (unsuitable) type. Older stands show large variation in their genetic diversity (v _gam between 19.9 and 90.1). Some of the stands of unsuitable provenance have natural regeneration and others have been selected as seed collecting stands. Through natural regeneration the genetic structure of the older stands is passed on to the following generation. The same is true for reproductive material collected from such stands. In comparison to seed collected from stands in the USA (from stands selected by the EU for seed collection), seed from Bavarian stands is genetically very heterogeneous in respect to the allele variants and the diversity. No significant changes in the genetic population structure were found in the nursery due to sowing or transplanting. The critical stage for obtaining high quality reproductive material of Douglas-fir is the selection of seed collecting stands and the harvesting procedure. The resulting consequences for seed collection and procurement of suitable reproductive material for Douglas-fir are discussed.     

Understanding plantation transformation using a size-structured spatial population model

Plantation transformation is a goal of increasing interest to silviculturalists. The target forest state is characterised by high variance in age and size, and an irregular spatial structure, which leads to inhomogeneous interactions between, and consequent development of, trees in the stand. This presents a difficulty for traditional methods such as yield tables, and demands a more careful consideration of stand dynamics. On the other hand, while forestry has a great heritage of simulation, the level of complexity implemented at an individual level generally precludes direct understanding of stand scale behaviours, and leads to difficulties in verification with appropriate data.A promising approach is the application of relatively simple models developed by ecologists. These can be adapted to yield accurate representations of forest stands, while being highly amenable to analysis. Motivated by data from Scots pine (Pinus sylvestris L.) stands, we here apply a simple spatial birth-death-growth model to the comparison and analysis of transformation strategies for plantation stands. The model captures the effects of neighbours in a way which retains the conceptual simplicity of a generic, analytically solvable model, while allowing insights into the driving factors of population dynamics.Timing and intensity of management interventions, as opposed to their specific criteria, are of primary importance: thinnings of a moderate intensity performed over a long period produced the best results. Variation in the strategy applied leads to more subtle effects which transformation strategies must also take into account, such as the development of variation in size of the remaining trees (increased using spatially correlated thinnings), the survival chances of regeneration and “underplanted” trees, and the overall productivity of the stand (increased using spatially homogeneous crown thinning). Finally, we demonstrate the robustness of model predictions to fundamental choices of model formulation.     

Reprint of: Lessons from long-term studies of harvest methods in southwestern ponderosa pine-Gambel oak forests on the Fort Valley Experimental Forest, Arizona, U.S.A.

The Fort Valley Experiment Station (now Fort Valley Experimental Forest) has contributed many long-term studies to forest research. This paper focuses on a “Methods of Cutting” study initiated in 1913 on the Coulter Ranch Unit of Fort Valley and how that long-term study yielded important ecological and management lessons. We quantified the historical and contemporary forest patterns at this ponderosa pine-Gambel oak site, which was harvested using three different harvesting systems in 1913 (seed tree, group selection, and light selection) and was partially excluded from livestock browsing in 1919. Using nine historically stem-mapped permanent plots for the following three stand structural scenarios: 1913 pre-harvest (modeled), post-harvest (actual), and 2003-2006 (actual) conditions, we examined the short- and long-term consequences of harvest and livestock grazing land-use and stand dynamics. We assessed changes in spatial pattern under each harvesting system and in each structural scenario, and lastly, we examined spatial and temporal tree recruitment patterns as observed in the contemporary (2003-2006) conditions. The seed tree harvests effectively converted the spatial patterns from aggregated to random and left few trees, while the group selection and light selection had varying effects, but consistently exaggerated the spatial patchiness of the stand. By 2003-2006, all plots were aggregated at all scales and were one large patch of predominately small trees. Sites that were harvested, but excluded from livestock browsing had 40% more trees in 2003-2006. Contemporary recruitment patterns were significantly aggregated under all harvesting systems, but were most strongly aggregated if the site received a group selection or light selection cut. For group and light selection, pine seedlings initially established in stump patches created by harvesting and then proceeded to fill-in the remaining area, with recruitment rarely found under the residual pine or oak trees. Long-term data sets, such as these established by the Fort Valley Experimental Forest in 1913, are essential for quantifying the impact of historical land-use practices on contemporary forest composition and structure. Ignoring land-use legacies may lead to the misinterpretation of stand dynamics and development, and therefore should be explicitly quantified and incorporated into future management and restoration activities.     

干扰与森林更新

干扰是森林生态系统动态中的主要影响因素。它可破坏或改变原有的环境，对森林群落的发展产生深远的影响。森林对于各种干扰的反应十分复杂，但总的反应可分为两类：一种格局表现为植被结构的重组，另一种格局是通过更新，形成新的植被。因此，森林更新是与各种干扰不能分隔的一个极为重要的生态学过程。本文主要对近年来有关干扰和森林更新的研究进行了综述，分析了干扰的强度、频率和空间范围等方面的变化，以及这些变化对森林产业的作用及其森林的反应，重点讨论了不同的干扰与森林更新机制的关系、干扰对更新的窨格局的影响，干扰与更新周期性的关系、干扰对种子扩散和更新幼苗的影响等方面的问题。     

STUDY ON THE NATURAL REGENERATION EFFECT OF MANCHURIAN ASH BY OPENING UP EFFECT BELT IN DAHURIAN LARCH PLANTATION

NaturalregenerationofmanchurianashusuallytakesplacewheretherearesuitablesoilmoistureandseedbankofsecondarymanchurianashforestintheEasternMountainousAreaofNortheastChina.Sometimes,naturalregenerationofmanchurianashcanbefoundinpureconiferousplantation.Inordertofindoutthepossibilityofintroducingnaturalregenerationofmanchurianashinpureconiferousplantation,trialsandresear-cheshavebeencarriedoutindahurianlarchp1antationbyopeningupeffectbeltinthestands.lntheend,charactersofcommunitystructureandmicr…     

Spatial patterns and genetic structures within beech populations (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) of forked and non-forked individuals

Analyses of distribution patterns and genetic structures of forest stands can address distinct family structures and provide insights into the association of genetic and phenotypic variation patterns. In this study, point pattern analysis and spatial autocorrelation were used to examine the spatial and genetic structures in two naturally generated beech stands, which differ in age, trunk morphology, and stand management. Significant tree clumping was observed at distances up to 20 m in the young forest stand, whereas dispersion at distances under 10 m was observed in the old stand. The spatial analysis based on Ripley’s k function of the two different groups of trees showed that the non-forked trees match in both stands the spatial pattern of all trees while the forked were randomly distributed. Additionally, according to the bivariate analysis, forked trees in both stands were randomly distributed as related to non-forked tree positions. Finally, Moran’s I values were not very high, though significant genetic autocorrelation was identified at distances up to 20 m in the young stand, suggesting the existence of distinct family structures. However, no significant genetic structuring was observed in the old stand. Our findings suggest that spatial genetic patterns are impacted by stand age, environmental factors and human activities. The spatial distribution of forked trees was not clearly associated to family structures. Random effects and also micro-environmental variation could be additional factors explaining forking of beech individuals.     

Lessons from long-term studies of harvest methods in southwestern ponderosa pine–Gambel oak forests on the Fort Valley Experimental Forest,Arizona, U.S.A.

The Fort Valley Experiment Station (now Fort Valley Experimental Forest) has contributed many long-term studies to forest research. This paper focuses on a “Methods of Cutting” study initiated in 1913 on the Coulter Ranch Unit of Fort Valley and how that long-term study yielded important ecological and management lessons. We quantified the historical and contemporary forest patterns at this ponderosa pine–Gambel oak site, which was harvested using three different harvesting systems in 1913 (seed tree, group selection, and light selection) and was partially excluded from livestock browsing in 1919. Using nine historically stem-mapped permanent plots for the following three stand structural scenarios: 1913 pre-harvest (modeled), post-harvest (actual), and 2003–2006 (actual) conditions, we examined the short- and long-term consequences of harvest and livestock grazing land-use and stand dynamics. We assessed changes in spatial pattern under each harvesting system and in each structural scenario, and lastly, we examined spatial and temporal tree recruitment patterns as observed in the contemporary (2003–2006) conditions. The seed tree harvests effectively converted the spatial patterns from aggregated to random and left few trees, while the group selection and light selection had varying effects, but consistently exaggerated the spatial patchiness of the stand. By 2003–2006, all plots were aggregated at all scales and were one large patch of predominately small trees. Sites that were harvested, but excluded from livestock browsing had 40% more trees in 2003–2006. Contemporary recruitment patterns were significantly aggregated under all harvesting systems, but were most strongly aggregated if the site received a group selection or light selection cut. For group and light selection, pine seedlings initially established in stump patches created by harvesting and then proceeded to fill-in the remaining area, with recruitment rarely found under the residual pine or oak trees. Long-term data sets, such as these established by the Fort Valley Experimental Forest in 1913, are essential for quantifying the impact of historical land-use practices on contemporary forest composition and structure. Ignoring land-use legacies may lead to the misinterpretation of stand dynamics and development, and therefore should be explicitly quantified and incorporated into future management and restoration activities.     

云冷杉过伐林主要树种结构特征分析

选择长白山针阔比9∶1、8∶2、7∶3的云冷杉过伐混交林,探讨直径结构、空间格局、蓄积生长、林下更新以及植物多样性等林分结构问题,为恢复红松为主的针阔混交林,提出结构调整思路。研究表明:由于过去以不合理的非经营性采伐,红松成主要择伐对象,红松不同年份各径阶株数变化幅度较大,直径18 cm开始出现缺损株数现象,缺乏中、大径组株数,导致母树数量、结实量和种源减少,造成在主要树种中红松更新最差的问题,不利于林分正向演替。不同针阔比混交林空间格局与其主要树种空间格局并非完全一致,在同一林分不同树种间存在差异。针阔比9∶1、8∶2、7∶3的混交林空间格局分别为随机分布、聚集分布和均匀分布。其中,红松分布格局主要呈聚集分布。因此,恢复以红松为主的针阔混交林,以红松和珍贵阔叶树种作为目标树种,采取目标树培育方法,伐除影响其更新和生长的林木,而将云冷杉和一般阔叶树作为伴生树种,逐渐减少云冷杉的优势,最终逐渐恢复成红松阔叶混交林;对针阔比8∶2和7∶3混交林红松分布格局,采取人工补植、间伐等措施,结合天然更新,向随机分布进行调整;采取人工补植、清理母树周围枯枝落叶层等措施人工辅助红松天然更新。     

Narrow genetic base in forest restoration with holm oak (Quercus ilex L.) in Sicily

In order to empirically assess the effect of actual seed sampling strategy on genetic diversity of holm oak (Quercus ilex) forestations in Sicily, we have analysed the genetic composition of two seedling lots (nursery stock and plantation) and their known natural seed origin stand by means of six nuclear microsatellite loci. Significant reduction in genetic diversity and significant difference in genetic composition of the seedling lots compared to the seed origin stand were detected. The female and the total effective number of parents were quantified by means of maternity assignment of seedlings and temporal changes in allele frequencies. Extremely low effective maternity numbers were estimated (Nf _e ≈ 2–4) and estimates accounting for both seed and pollen donors gave also low values (N _e ≈ 35–50). These values can be explained by an inappropriate forestry seed harvest strategy limited to a small number of spatially close trees.     

Ground seed density patterns under conditions of strongly overlapping seed shadows in <Emphasis Type="Italic">Abies alba</Emphasis> Mill. stands

In three subsequent years, seeds were collected from 66 to 75 seed traps in three Abies alba stands that differed in vertical structure and the number of potentially reproducing trees. The objective was to compare empirical patterns of seed density with a random (Poisson) model, which assumes that seed density would be the same in every location, and a binomial negative model that is appropriate for over-dispersed data. The seed density patterns were tested for spatio-temporal independence. The effect of some dispersal variables (number of mother trees and seed fall seasons, dispersal distance, fecundity rate) on seed density pattern was studied in a simple simulation experiment. Several local stand density measures (stem density, basal area, distance-weighted indices, canopy openness) were also tested for their ability to predict local seed density. The Poisson model was rejected in all the stands studied, and a much better fit achieved using the binomial negative model. The simulation study revealed that random seed distribution may occur only at considerable dispersal distances, concave seed shadows, and at very low fecundity rates. The seed density patterns identified exhibited both spatial and temporal correlation, which can probably be linked to the arrangement of mother trees and variation in their seed output. To a minor degree, this seed density pattern also correlated with local stand density.     

Spatial and temporal patterns in structure,regeneration, and mortality of an old-growth ponderosa pine forest in the Colorado Front Range

Effective management and restoration of ponderosa pine forests requires an understanding of the heterogeneity of contemporary and historical stand structures. We assessed spatial and temporal patterns of tree establishment, mortality and size structure over a 30-year period in an old-growth ponderosa pine stand in the mid-montane zone of the Colorado Front Range. We analyzed spatial patterns and spatial associations using Ripley's K(t) and K₁₂(t) and then modeled the patterns using point process models. Forest age structure was estimated by aging a sub-sample of trees in the stand. Climate appeared to play a significant role in the coarse-scale temporal pattern of regeneration events. Stand structure (distribution of patches, light availability, and seed trees) influenced the spatial and temporal pattern of more recent regeneration events. Patchy regeneration resulted in spatial independence and some segregation of size classes. Older trees in the stand (40–55 cm dbh) exhibited some regularity in their spatial distribution at short distances indicating that patterns of mortality had been historically patchy. Contemporary patterns of mortality were mostly patchy, and mountain pine beetles caused a significant amount of mortality in the 1970s and 1980s. Both establishment and mortality retained spatial patterns that were somewhat consistent with pre-settlement forests, despite changes in driving processes.     

三峡库区马尾松天然林林分结构特征分析

以三峡库区无干扰马尾松天然林为研究对象,调查了群落物种组成和天然更新状况,并选择角尺度、混交度和大小比数3个空间结构参数及直径分布等非空间结构参数,对马尾松天然林林分结构特征进行分析,为马尾松次生林的科学经营提供理论依据。研究结果表明:研究地区马尾松群落物种数量丰富,共出现维管束植物117种,林下天然更新状况良好,共有树种24种,数量达5 221株·hm^-2,其中高度小于100 cm的幼苗占48.9%。林分随着直径增大,林木株数逐渐减少,呈典型异龄林直径分布特征。林分平均角尺度为0.508,林木整体分布格局属随机分布;林分平均混交度为0.60,整体处于中度以上混交;林分平均大小比数为0.50,大部分林木个体分化程度较高,处于不同竞争地位的林木个体数量基本相等。优势树种马尾松属轻度聚集分布,接近中度混交,且60.2%的个体处于竞争优势或亚优势地位。     

Stand structure of an uneven-aged fir–beech forest with an irregular diameter structure: modeling the development of the Belevine forest, Croatia

This study focuses on the problem of irregular diameter structure in a silver fir–beech selection (plenter) forest with a “surplus” of large diameter trees and a lack of natural regeneration and small diameter trees. We sampled 274 plots (900 m² each) in the Belevine research site (266.24 ha) in the mountain region Gorski Kotar (Croatia), where diameter (dbh) distribution, diameter increment, and natural regeneration were analyzed in detail. A low density of natural regeneration, weak annual recruitment of small (10 cm dbh) diameter trees (only five trees per hectare), delayed diameter growth of trees, and a low annual rate of trees reaching the next dbh class were attributed to the current irregular dbh structure. The stand development prediction for the next 50 years is based on a simulation model, which considers the current diameter structure, increment, recruitment, and future cutting regime. Intensive cutting in the first of five 10-year cutting cycles (intensity higher of 25%) is needed to initiate natural regeneration and to accelerate growth of young silver fir trees. In the next 50 years, the irregular diameter structure will be gradually improved.     

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.