Tree demography suggests multiple directions and drivers for species range shifts in mountains of Northeastern United States

Climate change is expected to lead to upslope shifts in tree species distributions, but the evidence is mixed partly due to land‐use effects and individualistic species responses to climate. We examined how individual tree species demography varies along elevational climatic gradients across four states in the northeastern United States to determine whether species elevational distributions and their potential upslope (or downslope) shifts were controlled by climate, land‐use legacies (past logging), or soils. We characterized tree demography, microclimate, land‐use legacies, and soils at 83 sites stratified by elevation (~500 to ~1200 m above sea level) across 12 mountains containing the transition from northern hardwood to spruce‐fir forests. We modeled elevational distributions of tree species saplings and adults using logistic regression to test whether sapling distributions suggest ongoing species range expansion upslope (or contraction downslope) relative to adults, and we used linear mixed models to determine the extent to which climate, land use, and soil variables explain these distributions. Tree demography varied with elevation by species, suggesting a potential upslope shift only for American beech, downslope shifts for red spruce (more so in cool regions) and sugar maple, and no change with elevation for balsam fir. While soils had relatively minor effects, climate was the dominant predictor for most species and more so for saplings than adults of red spruce, sugar maple, yellow birch, cordate birch, and striped maple. On the other hand, logging legacies were positively associated with American beech, sugar maple, and yellow birch, and negatively with red spruce and balsam fir – generally more so for adults than saplings. All species exhibited individualistic rather than synchronous demographic responses to climate and land use, and the return of red spruce to lower elevations where past logging originally benefited northern hardwood species indicates that land use may mask species range shifts caused by changing climate.     

Regeneration of monsoon rain forest in northern Australia: the sapling bank

Abstract. As part of a wider study examining regeneration pathways in monsoon rain forest vegetation in northern Australia, a one-off, dry season census of saplings was undertaken along transects sampled at each of 33 relatively undisturbed sites broadly representative of the range of regional monsoon rain forest vegetation. Four floristic quadrat groups were derived through TWINSPAN classification. Subsequent analyses involved: (1) comparison of mean dry season stockings of juveniles occurring in different rain forest types, and their structural and environmental correlates; (2) comparison of the contributions of different life forms, and the influence of clonal reproduction in the sapling regeneration banks of different forest types; and (3) exploration of relationships between the distributions of saplings of common tree species with respect to seed bank, floristic, structural, and environmental variables. While data presented here require cautious interpretation given that processes of seedling/sapling recruitment and mortality are highly dynamic, sapling banks were found to be most dense on coarse-textured, moist soils, and least dense on coarse-textured, seasonally dry soils. Canopy cover and fire impact were shown to be highly influential on sapling distribution, especially for saplings of tree species and those growing on seasonally dry sites. Sapling densities were little influenced by proximity to rain forest margins, except for shrubs. The potential for clonal reproduction was significantly greater on dry sites, especially for trees. The majority of saplings sampled were derived from relatively few common, non-clonal, canopy tree species. Sapling distributions of 20 out of 23 common tree species were clumped in the vicinity of conspecific adults; for most species the strength of this relationship was greater than that for any other variable. These data support observations in the literature concerning the distribution of sapling banks in moist and dry tropical forests.     

Grow slowly,persist, dominate—Explaining beech dominance in a primeval forest

Being able to persist in deep shade is an important characteristic of juvenile trees, often leading to a strong dominance of shade‐tolerant species in forests with low canopy turnover and a low disturbance rate. While leaf, growth, and storage traits are known to be key components of shade tolerance, their interplay during regeneration development and their influence on juveniles'' survival time remains unclear. We assessed the ontogenetic effects of these three traits on the survival time of beech (Fagus sylvatica), and Norway and sycamore maples (Acer pseudoplatanus, Acer platanoides) in a primeval beech forest. Biomass allocation, age, and content of nonstructural carbohydrates (NSC) were measured in the stems and roots of 289 seedlings and saplings in high‐ and low‐vitality classes. Saplings experienced a trade‐off between absolute growth rate (AGR) and storage (NSC) as the leaf area ratio (LAR) decreases with biomass development. High LAR but low AGR and low NSC corresponded to beech with a marked ability to persist in deep shade while awaiting canopy release. In turn, a comparably small LAR in combination with a high AGR and higher storage (NSC), as observed in Norway maple and sycamore maple, reduced sapling survival time, thus offering an explanation for beech dominance and maple disappearance in the undergrowth of old‐growth beech forests.     

Diversity of canopy and understorey spiders in north-temperate hardwood forests

1 We characterized and compared diversity patterns of canopy and understorey spiders (Arachnida: Araneae) on sugar maple ( Acer saccharum Marsh.) and American beech ( Fagus grandifolia Ehrh.) in hardwood forests of southern Québec, Canada.
2 We sampled canopies of 45 sugar maple and 45 American beech trees and associated understorey saplings in mature protected forests near Montréal. Samples were obtained by beating the crown foliage at various heights and by beating saplings around each tree.
3 Eighty-two species were identified from 13 669 individuals. Forty-eight species and 3860 individuals and 72 species and 9809 individuals were collected from the canopy and the understorey, respectively.
4 Multivariate analyses (NMDS ordination and NPMANOVA) showed the composition of canopy and understorey assemblages differed significantly, and canopy assemblages differed between tree species. Rank-abundance distribution models fitted to the canopy and understorey data indicated that different mechanisms structure the assemblages in both habitats. Three abundant spider species were significantly more common in the canopy; ten species were collected significantly more often in the understorey.
5 The forest canopy was shown to be an important reservoir for spider diversity in north-temperate forests.     

Canopy gaps and establishment patterns of spruce (Picea abies (L.) Karst.) in two old-growth coniferous forests in central Sweden

The spatial pattern of seedlings, saplings and canopy trees was studied in two spruce (Picea abies (L.) Karst.) forests in central Sweden. Canopy and forest structure were determined in five 0.25 ha plots. Life stage classes were distinguished on the basis of age and size distributions. Ripley's K-function (1977) was used to analyze the spatial patterns within each class. A random distribution of seedlings gave way to a more aggregated pattern on a small scale during the establishment phase. Saplings and sub-canopy trees were strongly aggregated and canopy trees were again randomly distributed within the plots. The proportion of individuals growing in gaps was used as an index of association between the spatial pattern in saplings and sub-canopy trees and the occurrence of small (50–350 m²) canopy gaps. Under the null hypothesis of independence the expected value of this statistic would equal the canopy gap ratio for the stand. Monte Carlo simulation of this statistic, using fixed sapling positions and randomly repositioned canopy gaps, confirmed the importance of canopy gaps for the final success of establishment of spruce. The association of understorey trees with gaps suggest that small gaps are typically closed by recruitment of new saplings from a sapling bank rather than by the release of larger suppressed trees.     

Composition and Dynamics of Functional Groups of Trees During Tropical Forest Succession in Northeastern Costa Rica

We compared the functional type composition of trees ≥10 cm dbh in eight secondary forest monitoring plots with logged and unlogged mature forest plots in lowland wet forests of Northeastern Costa Rica. Five plant functional types were delimited based on diameter growth rates and canopy height of 293 tree species. Mature forests had significantly higher relative abundance of understory trees and slow-growing canopy/emergent trees, but lower relative abundance of fast-growing canopy/emergent trees than secondary forests. Fast-growing subcanopy and canopy trees reached peak densities early in succession. Density of fast-growing canopy/emergent trees increased during the first 20 yr of succession, whereas basal area continued to increase beyond 40 yr. We also assigned canopy tree species to one of three colonization groups, based on the presence of seedlings, saplings, and trees in four secondary forest plots. Among 93 species evaluated, 68 percent were classified as regenerating pioneers (both trees and regeneration present), whereas only 6 percent were classified as nonregenerating pioneers (trees only) and 26 percent as forest colonizers (regeneration only). Slow-growing trees composed 72 percent of the seedling and sapling regeneration for forest colonizers, whereas fast-growing trees composed 63 percent of the seedlings and saplings of regenerating pioneers. Tree stature and growth rates capture much of the functional variation that appears to drive successional dynamics. Results further suggest strong linkages between functional types defined based on adult height and growth rates of large trees and abundance of seedling and sapling regeneration during secondary succession.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp     

Individual Canopy‐tree Species Effects on Their Immediate Understory Microsite and Sapling Community Dynamics

Canopy trees are largely responsible for the environmental heterogeneity in the understory of tropical and subtropical species‐rich forests, which in turn may influence sapling community dynamics. We tested the effect of the specific identity of four cloud forest canopy trees on total solar radiation, canopy openness, soil moisture, litter depth, and soil temperature, as well as on the structure and dynamics of the sapling community growing beneath their canopies. We observed significant effects of the specific identity of canopy trees on most understory microenvironmental variables. Soil moisture was higher and canopy openness lower beneath Cornus disciflora. In turn, canopy openness and total solar radiation were higher beneath Oreopanax xalapensis, while the lowest soil moisture occurred beneath Quercus laurina. Moreover, Chiranthodendron pentadactylon was the only species having a positive effect on litter depth under its canopy. In spite of these between‐species environmental differences, only C. pentadactylon had significant, negative effects on sapling density and species richness, which may be associated to low seed germination and seedling establishment due to an increased litter depth in its vicinity. The relevance of the specific identity of canopy trees for natural regeneration processes and species richness maintenance depends on its potential to differentially affect sapling dynamics through species‐specific modifications of microenvironmental conditions.     

Morphological plasticity of regeneration subject to different levels of canopy cover in mixed-species, multiaged forests of the Romanian Carpathians

Morphological plasticity was studied for advanced regeneration trees in different light environments of the mountainous, mixed-species forests in the Carpathian Mountains of Romania. The primary species in these mixtures were very shade tolerant silver fir (Abies alba Mill.) and European beech (Fagus sylvatica L.), and midtolerant Norway spruce (Picea abies (L.) Karst). Seedlings/saplings of these species were selected for measurements in different stands from two different geographical locations. Various morphological traits (specific leaf area, live crown ratio, crown width to length ratio, terminal to lateral ratio, number of internodal shoots, number of shoots in terminal whorl, stem symmetry, stem orientation, stem forking) for each regenerating tree were measured during summers of 2001 and 2002. Percentage of above canopy light and stand basal area measures were used to assess the available growing space for each seedling/sapling. Regression relationships were developed for the different morphological indicators as a function of these two variables. All species adapted their morphology along the gradient in light and basal area. Spruce seemed to be less adapted to low light conditions than both fir and beech. However, no significant differences in terms of shade tolerance were detected using the above indicators. In really dense stand conditions (less than 20% above canopy light and stand basal area above 36 m² ha⁻¹), probability for stem forking in beech increased. In open, all three species adapted their morphology for vigorous growth. Under such conditions, spruce was better adapted than fir.     

Disparate distribution patterns between canopy and subcanopy life-forms in two temperate North American forests

Quantitative vegetational data of canopy and woody subcanopy species (two life-forms adapted to occupy different strata at maturity) were compared with data collected in two temperate forest ecosystems to determine whether they exhibit a similar pattern of distribution. Tidal freshwater swamps (21 stands) and southern Appalachian forests (19 stands) were examined from data obtained using identical sampling methods. Separate structural analyses of the canopy, sapling, and subcanopy species were compared using the indirect ordination algorithm Detrended Correspondence Analysis. Environmental measurements collected in each stand were assessed for their relationship to the distribution of stands depicted by the ordination diagrams.Canopy trees and saplings showed a similar pattern of distribution, suggesting that the resource requirements of saplings and canopy-statured adults are similar. In contrast, the subcanopy species (species genetically adapted to an understory existence, i.e., shrubs and small understory trees) of neither ecosystem showed any discernable distributional relationship to the canopy or sapling layers (in tidal swamps, there was no clear way to even segregate subcanopy stands into communities). Environmental gradients associated with the subcanopy ordinations differed from those of the canopy and sapling strata in both forest systems, suggesting that subcanopy species partition different resources than do canopy species.If a lack of similarity in distribution patterns between canopy and subcanopy species is universal in temperate forests, then the common practice of combining sapling and subcanopy species in structural analyses may hinder our understanding of subcanopy structural patterns in forests.     

黑龙江省不同地点蒙古栎林生态特点研究

通过对黑龙江省６个地点的天然蒙古栎林的结构和更新特点的分析,蒙古栎林可划分为不同特点的蒙古栎群落,即纯蒙古栎群落、蒙古栎桦树群落、蒙古栎槭树群落、蒙古栎红松群落和蒙古栎松混交林群落,其演替趋势如下：红蒙古栎群落、蒙古栎桦树群落至蒙古栎槭树落、蒙古栎红松群落,再至蒙古栎红松混交林群落,蒙古栎群落类型的多样性主要反应群落不同的演替阶段,造成蒙古栎群落多样的原因是人类活动和自然因素作用的结果,随着群落的演替,蒙古栎的优势地位逐渐被消弱,乔木种类丰富度增多,草本种类丰富度增多;蒙栎的相对密度下降,林内环境由于干燥逐渐变中性至较湿润,蒙古栎幼苗和幼树在总幼苗和幼树中所占的比例下降,耐荫物中色木槭等的幼苗和幼树所占比例上升;在演替过程中,蒙古栎分布格局－聚集分布的聚集程度逐渐降低,并向随机分布的方向发展。     

Differential drought responses between saplings and adult trees in four co-occurring species of New England

Tree-ring characteristics in four species were examined to address whether co-occurring mature trees of different successional status respond differently to drought, and whether saplings of these species have a greater response to drought than mature trees. We examined saplings and mature trees of paper birch, yellow birch, red maple and sugar maple, which varied in successional status (shade-tolerance) and co-occurred at Harvard Forest, Petersham, Mass., USA. Three drought events in 1964–1966, 1981 and 1995 were identified using climate data. For mature trees, there was no significant interspecific difference in relative changes in ring-width index (RWI) during the 1964–1966 and 1995 drought events. However, the interspecific difference was significant in the 1981 drought event. Response function analysis for mature trees showed that the radial growth of sugar maple was mainly controlled by spring and summer precipitation, red maple by spring and summer precipitation and temperature, yellow birch by winter and summer precipitation, and spring and summer temperature, and paper birch by spring and summer precipitation and spring temperature. Saplings of sugar maple and yellow birch, but not red maple and paper birch, showed significant positive correlations between RWI and annual total precipitation. In the 1995 drought event, saplings and mature trees of red maple and paper birch differed significantly in drought responses, but this was not true in sugar maple and yellow birch. Our results do not support a generally greater response in saplings than in mature trees, nor an early- versus late successional difference in drought responses.     

Comparison of the initial demographies of pine and oak populations in rural pine forests in Korea and Japan

In order to understand the human impact upon demographic change in plant population in rural forests, we examined the population trends ofPinus densiflora andQuercus serrata inP. densiflora forests viewing their seedlings and saplings in rural Korea and Japan. The most prominent factor affecting the regeneration of the pine and oak was the intensity of management activity which controlled the vegetation stratification and its light environment. Open spaces, such as graveyard or cleaned area that allow the long-term direct daylight in dry season to accelerate the surface heat of soil, were unfavorable habitats for germination and growth of both species. The negative effects of the presence of litter and evergreen trees were related to the failure of early seedling and sapling growth of pine. Cover of litter is, especially, another factor related to the growth of pine saplings. It is considered that successful germination and sapling growth of pine and oak in early successional stages is determined by vegetation structures and light condition.     

Soil Base Saturation Combines with Beech Bark Disease to Influence Composition and Structure of Sugar Maple-Beech Forests in an Acid Rain-Impacted Region

Sugar maple, an abundant and highly valued tree species in eastern North America, has experienced decline from soil calcium (Ca) depletion by acidic deposition, while beech, which often coexists with sugar maple, has been afflicted with beech bark disease (BBD) over the same period. To investigate how variations in soil base saturation combine with effects of BBD in influencing stand composition and structure, measurements of soils, canopy, subcanopy, and seedlings were taken in 21 watersheds in the Adirondack region of NY (USA), where sugar maple and beech were the predominant canopy species and base saturation of the upper B horizon ranged from 4.4 to 67%. The base saturation value corresponding to the threshold for Al mobilization (16.8%) helped to define the species composition of canopy trees and seedlings. Canopy vigor and diameter at breast height (DBH) were positively correlated (P < 0.05) with base saturation for sugar maple, but unrelated for beech. However, beech occupied lower canopy positions than sugar maple, and as base saturation increased, the average canopy position of beech decreased relative to sugar maple (P < 0.10). In low-base saturation soils, soil-Ca depletion and BBD may have created opportunities for gap-exploiting species such as red maple and black cherry, whereas in high-base saturation soils, sugar maple dominated the canopy. Where soils were beginning to recover from acidic deposition effects, sugar maple DBH and basal area increased progressively from 2000 to 2015, whereas for beech, average DBH did not change and basal area did not increase after 2010.     

天山云杉天然林不同林层的空间格局和空间关联性

天山云杉(Picea Schrenkiana)林是天山林区的重要针叶林。研究调查了分别代表天山云杉增长型种群和成熟型种群的两块标准地,应用Ripley's K系数法研究了天山云杉更新层、低林层、中林层和高林层的空间格局和不同林层的空间关联性,并应用Moran's I系数法研究了不同林层的林木高度的空间自相关性。结果表明:所有林层的天山云杉活立木都是显著聚集的,只是发生聚集格局的强度和聚集尺度不同。更新层天山云杉具有最大的聚集强度,从更新层到低林层,聚集强度急速降低。两块标准地中天山桦(Betula tianschanica Rupr.)、天山柳(Salix tianschanica Rgl.)和天山云杉活立木的空间关联性的表现不同。研究结果表明:增长型种群中天山桦、天山柳在所有尺度上都抑制天山云杉的生长;而成熟型种群中天山桦、天山柳在1-10 m尺度内对天山云杉种群的生长没有影响,10-40 m表现出正相关,40 m以上没有表现出明显的关联性;同时天山桦、天山柳基本上在所有尺度都抑制天山云杉的更新。在更新层和低林层间表现出正相关;而更新层和枯死木之间的空间关联性为正相关。     

Initial regenerative processes ofDistylium racemosum andPersea thunbergii in an evergreen broad-leaved forest

We studied the initial regenerative processes ofDistylium racemosum andPersea thunbergii, major canopy species, in an old-growth evergreen broad-leaved forest, Kagoshima, southwestern Japan. Although the two species coexisted, the patterns of seed production, seedling emergence, and sapling survival were different between them. During the observation (1989–1991),P. thunbergii produced large numbers of seeds in alternate years, whileD. racemosum had no mast year. The density of established seedlings was high and increased with mass seed production forP. thunbergii, but was fairly low forD. racemosum. The mortality of seedlings and saplings ofP. thunbergii was higher than that ofD. racemosum. In a closed stand, the sapling bank was maintained by dense seedling supply forP. thunbergii, and by extremely low mortality of individuals germinated before the beginning of observation forD. racemosum. Since saplings ofD. racemosum showed continuous height growth beneath the closed canopy, the possibility of recruitment into the upper layer seems to be high. The strategy ofP. thunbergii might be to wait for suitable conditions favoring sapling growth on various site by means of frequent and large seed productivity and wide dispersal of seeds. Thus both species might be able to coexist through environmental heterogeneity.     

Recruitment patterns and northward tree migration through gap dynamics in an old-growth white pine forest in northern Ontario

Decreases in abundances and declines in growth of eastern white pine over the past century due mainly to human activities have resulted in few large intact old-growth white pine forests in Ontario. These stands may be vulnerable to replacement by deciduous species from temperate forests further south, where recruitment in canopy gap disturbances can greatly define the regeneration process. We investigated recruitment dynamics in canopy gaps of an old-growth white pine forest of Temagami, northern Ontario, Canada, the northern limit of the temperate?Cboreal ecotone. White pine, red pine, black spruce and eastern white cedar represented 85?% of the mature canopy abundance, where trees and saplings established equally in gaps and the closed canopy. Balsam fir and paper birch were more abundant in gaps, showing increases of abundance and basal area with increases in gap size representing canopy self-replacement (balsam fir) and autogenic succession (paper birch). Red maple, at its northernmost range limit, was the only species to show linear increases of abundance and basal area with increases in gap size and gap age. This result, along with adult red maples present in gaps but absent from the closed canopy, identifies the establishment of a northward migrating species in gaps as hypothesized for pine forests at the northern limit of this broad ecotone. We discuss how migration pressures, coupled with pine recruitment limitation through reduced fire frequency by regional fire suppression and predicted future increased warming of 2?C4?°C over the next century, threatens replacement of old-growth white pine forests at this latitude with northward migrating tree species found further south.     

Tree distribution pattern and fate of juveniles in a lowland tropical rain forest – implications for regeneration and maintenance of species diversity

Three analyses of species diversity in a lowland dipterocarp forest were conducted to examine whether the nature of forest community dynamics are determined by density-dependent recruitment and mortality of saplings with a data set obtained in a 50 ha plot in Pasoh Forest Reserve. The first analysis examined whether sapling density varied as a function of distance from the nearest conspecific adult. The second analysis assessed the relationship between the spatial distribution patterns of saplings and adult trees. A third analysis examined sapling recruitment and mortality based on data from 2 censuses, taken in 1985 and 1990. Four hundred forty-four species (each with more than 100 individuals) out of the total of 814 species recorded in the plot, were chosen for the analyses. Of these selected species, 56 species showed significant reduction in sapling densities close to the conspecific adults. Within this group, 11 species were in the emergent layer (29.0% of the total species in this layer), 17 were in the canopy layer (10.5%), 18 were in the understory layer (11.3%), and 10 were in treelet and shrub layer (11.8%). In contrast, the sapling densities of 53 species decreased with increasing distance from conspecific adults; 2 of these species were in the emergent layer (5.2% of the total species in this layer), 14 were in the canopy layer (8.6%), 21 were in the understory layer (13.2%), and 16 were in the treelet and shrub layer (18.8%). The saplings of 35 of the 444 total selected species were clumped, while adults were regularly or randomly distributed. Of the remaining species, in 183 species (41.2%), the distributions of both adults and saplings were clumped. Thus, these 2 analyses do not support the prediction that most of the species of lowland tropical forests fail to produce new adults in their vicinity and as a result of this, adult trees are more regularly distributed than their conspecific juveniles (Janzen 1970). In the third analysis, the recruitment of saplings of species in the emergent and canopy layers increased significantly and in proportion with mortality, suggesting that the dominant species suffer higher mortality than do less common species. This trend is not so apparent in the understory, and the treelet and shrub layers. The results imply that a dynamic equilibrium process, which prevents competitive exclusion and maintains space for minor species, may be active among the species in the upper layers (particularly the emergent layer); however, such a dynamic equilibrium condition is not due exclusively to the reduced recruitment of saplings near conspecific adults, and the dynamic equilibrium condition is not prevalent among the lower story species.     

Effect of canopy closure on pollen dispersal in a wind-pollinated species (Fagus sylvatica L.)

The effect of non-reproductive trees and saplings as a physical barrier to pollen dispersal in wind-pollinated species?? forests has not received enough attention in the literature so far. The neighborhood seedling model was used to fit pollen dispersal models for beech at different stages of gap recolonization and to elucidate the effect of saplings as a physical barrier on pollen dispersal at local scale. Phenological overlap of leaf emergence, and pollen release as well as wind directionality patterns were also examined. As a case study, we used a mixed beech-oak forest that was managed as open woodland until 1974. The ban on entry of cattle has led to the recolonization of empty spaces by seedlings and saplings of beech (Fagus sylvatica L.) and two oak species (Quercus petraea (Matts.) Liebl. and Q. pyrenaica Willd.) and, at last, to canopy closure. The average pollen dispersal distance for the first plants that regenerated in the gaps was almost twice those found for recently installed seedlings and seeds collected in traps, supporting the hypothesis that the understory may act as a physical barrier to pollen dispersal. Although a substantial part of effective pollination directionality is at random, horizontal winds and vertical anabatic winds may explain some of this directionality. At the time of beech pollen release, leaves of beech and sessile oak are fully developed, enhancing pollen interception by the saplings. Explicit models of pollen dispersal for wind-pollinated trees should incorporate the effect of canopy closure caused by growth of saplings and account for leaf phenology of co-occurring species in the forest.     

Norway Maple (Acer platanoides) Invasion of a Natural Forest Stand: Understory Consequence and Regeneration Pattern

Norway maple (Acer platanoidesis) is invasive in a natural stand in suburban Ithaca, NY. To determine the understory pattern and consequences of a Norway maple invasion, I compared density and species richness under Norway maples and sugar maples (Acer saccharum). Mean sapling density was significantly lower (P<0.0027) under Norway maples (3.64/100 m²±1.6 SE) than under sugar maples (19.4/100 m²±4.4 SE). Mean sapling species richness was significantly lower (P<0.0018) under Norway maples (0.7/32 m²±0.18 SE) than under sugar maples (2.6/32 m²±0.48 SE). Likewise, Norway maple regeneration is more frequent under sugar maples than sugar maple regeneration: 57% of sugar maple plots had Norway maple saplings while 0% of Norway maple plots had sugar maple saplings. Two significant plot effects were found for presence–absence: Norway maple saplings grow under Norway maples with a significantly lower frequency (P<0.03) than under sugar maples; sugar maple saplings grow under Norway maples with a significantly lower frequency (P<0.000) than under sugar maples. Across the site, Norway maple saplings were the most abundant (29 saplings for 480 m²). The success of Norway maple regeneration and the reductions in total stem density beneath Norway maples is most likely the result of its strong competitive abilities, notably its high shade tolerance and abundant seed crops.     

Estimation of tree replacement patterns in an Appalachian Picea-Abies forest

Abstract. Patterns of tree species replacement in a Picea-Abies forest, determined by several different methods, are compared and the methods are assessed. Methods are grouped as either understory-based or gap-based estimates of replacement. The understory-based methods characterize canopy-understory interactions with spatial statistics, sapling density measurement, sapling frequency measurement, and successor sapling identification beneath live canopy trees. The gap-based methods include sapling density measurement, sapling frequency measurement, and successor sapling identification in tree-fall gaps. Methods except those based on frequency indicate a strong trend of replacement of all canopy species by Abies. Understory-based methods may underestimate canopy recruitment of intolerant trees, while gap-based methods relying on sapling density or frequency may overestimate recruitment of intolerant trees. Estimates based on the selection of successor saplings in the understory or in gaps are reliable. Gap successor estimates consider the process of gap capture and are useful in analyses of forest dynamics.