Persistence of<Emphasis Type="Italic">Pseudotsuga menziesii</Emphasis> (Douglas-fir) in temperate coniferous forests of the Pacific Northwest Coast,USA

Old-growthPseudotsuga-Tsuga forests of the Pacific Northwest Coast of North America are characterized by the presence of large, old trees ofPseudotsuga menziesii var.menziesii (Douglas-fir). Colonizing soon after a stand-replacing disturbance,P. menziesii persists in these forests, coexisting for centuries with the late-successional species.P. menziesii survives by maintaining emergent status in the uppermost part of the forest canopy, above the crowns of competing late-successional species. After reaching maximum tree height and crown size,P. menziesii maintains shoots and foliage of the established crown by epicormic shoot production. In this review, we propose that attaining emergent status in the upper canopy combined with the process of crown maintenance contributes to the persistence ofP. menziesii into later stages of succession, making this species a long-lived pioneer that between infrequent disturbances can coexist with late-successional species for centuries.     

Age‐related Crown Thinning in Tropical Forest Trees

Gap dynamics theory proposes that treefall gaps provide high light levels needed for regeneration in the understory, and by increasing heterogeneity in the light environment allow light‐demanding tree species to persist in the community. Recent studies have demonstrated age‐related declines in leaf area index of individual temperate trees, highlighting a mechanism for gradual changes in the forest canopy that may also be an important, but less obvious, driver of forest dynamics. We assessed the prevalence of age‐related crown thinning among 12 tropical canopy tree species sampled in lowland forests in Panama and Puerto Rico (total N = 881). Canopy gap fraction of individual canopy tree crowns was positively related to stem diameter at 1.3 m (diameter at breast height) in a pooled analysis, with 10 of 12 species showing a positive trend. Considered individually, a positive correlation between stem diameter and canopy gap fraction was statistically significant in 4 of 12 species, all of which were large‐statured canopy to emergent species: Beilschmiedia pendula, Ceiba pentandra, Jacaranda copaia, and Prioria copaifera. Pooled analyses also showed a negative relationship between liana abundance and canopy gap fraction, suggesting that lianas could be partially obscuring age‐related crown thinning. We conclude that age‐related crown thinning occurs in tropical forests, and could thus influence patterns of tree regeneration and tropical forest community dynamics.     

Gap regeneration of shade-intolerant Larix gmelini in old-growth boreal forests of northeastern China

Abstract. Stand mapping and vertical and oblique tree crown projections were used to study gap characteristics and gap effects on the regeneration and stand development of Larix gmelini. The hypothesis is that waves of advance Larix regeneration are recruited into the canopy layer following the creation of canopy openings. In old-growth Larix forests of the northern Da Xingan Ling Mts., at 52 °N, obliquely projected gaps (OPG) begin at a distance of 60 - 80% of the canopy tree height from the southernmost stems bordering the gap and thus the OPGs may extend beyond the northern boundaries of the vertically projected gaps (VPG). Changes in the environment and resource availability in the OPG result in increased Larix sapling survival. Due to a greater incidence of light, 10 - 30 yr old OPG saplings were more abundant than saplings in either a near-oblique projected gap (NOPG) or in the shadow of obliquely projected crowns (SOPC). The survival of saplings more than 30-yr old was highest when they were found in the OPG of one canopy opening and the VPG of another. This means that, following recruitment into an OPG, saplings then require the space found in a VPG to permit growth into the canopy. Thus, various-sized gaps contribute to the survival of different aged saplings by increasing the complexity of stand structure. Although individuals may regenerate in an OPG, successful recruitment into the canopy requires the available growing space of a VPG. This research suggests that shade intolerant Larix gmelini can maintain its canopy dominance without fire via gap regeneration.     

Loss of oak dominance in dry-mesic deciduous forests predicted by gap capture methods

Gap capture methods predict future forest canopy species composition from the tallest trees growing in canopy gaps rather than from random samples of shaded understory trees. We used gap capture methods and a simulation approach to forecast canopy composition in three old oak forests (Quercus spp.) on dry-mesic sites in southern Wisconsin, USA. In the simulation, a gap sapling is considered successful if it exceeds a threshold height of 13–17 m (height of maximum crown width of canopy trees) before its crown center can be overtopped by lateral crown growth of mature trees. The composition of both the tallest gap trees and simulated gap captures suggests that 68–90% of the next generation of canopy trees in the stands will consist of non-Quercus species, particularly Ulmus rubra, Carya ovata and Prunus serotina. Quercus species will probably remain as a lesser stand component, with Quercus alba and Quercus rubra predicted to comprise about 19% of successful gap trees across the three stands. Several methods of predicting future canopy composition gave similar results, probably because no gap opportunist species were present in these stands and there was an even distribution of species among height strata in gaps. Gap trees of competing species already average 11–13 m tall, and mean expected time for these trees to reach full canopy height is only 19 years. For these reasons, we suggest that dominance will shift from oaks to other species, even though late successional species (e.g., Acer and Tilia) are not presently common in the understories of these stands.     

辽东栎林内不同小生境下幼树植冠构型分析

以黄土高原黄龙山林区辽东栎林内3个小生境（林下、林隙、林缘）下辽东栎天然更新幼树为研究对象,采用典型抽样法对辽东栎幼树侧枝、叶片和树冠的空间分布状况以及生物量分配状况进行调查分析,探讨微生境与幼树植冠构型特征的关系,明确辽东栎幼树对不同小生境的适应策略,为栎林经营和林分结构优化提供理论依据。结果显示：（1）3种生境下辽东栎幼树构型发生了可塑性变化,林下幼树树冠层次比较单一,林隙与林缘的幼树树冠层次更加丰富。（2）由林下至林缘,幼树的树高、枝下高呈逐渐减小的趋势,而地径变化趋势与之相反;幼树的冠幅、树冠面积、树冠率呈先增加后减小的趋势,并且林下与林隙、林缘的差异显著;幼树的总体分枝率、逐步分枝率、枝径比呈先增加后减小的趋势。（3）3种生境下,幼树的一级枝的枝长、直径与倾角随着树高的增加而呈减小的趋势,但3种生境的差异不显著;林下一级枝主要分布在冠层中上部,而林隙与林缘一级枝主要分布在冠层中下、中上部。（4）由林下至林缘幼树叶长、叶宽、单叶面积和比叶面积逐渐降低,而单株叶数、叶总面积、叶面积指数呈先增大后减小趋势;与其他2种生境相比林下叶片分布趋于冠层上部。（5）幼树地上部分生物量中林下主干生物量占83%,枝和叶生物量只占17%;而林隙与林缘虽然各部位生物量有所差异但比例基本一致,其中主干占66%左右,枝和叶生物量占34%左右。研究表明,林隙生境下幼树的构型优于林缘和林下生境,在今后栎林的经营中,可以通过适当间伐来增加林隙数量,为森林更新和结构的优化提供有利条件。     

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)。     

Woodland canopy structure and the light response of juvenile Quercus lobata (Fagaceae)

I investigated competition for light between canopy plants and juvenile valley oaks (Quercus lobata Nee) in a mixed-broadleaf woodland of California's northern Coast Ranges. Canopy effects on understory light supply were separated among the overlying adult valley oak, the adult's woody understory, and neighboring trees and shrubs through a series of light sampling surveys and measurements of the number, size, and spatial distribution of neighboring plants. Light supply in the understory was primarily influenced by neighboring plants, with no detectable effect of the overlying adult valley oak. Light supply in the understory averaged 25% full sun due to a high frequency of canopy gaps and a typically open understory. Seedling response to understory light supply was investigated in an experimental sunfleck gradient (10%, 19%, and 100% full sun). Between 10% and 100% full sun, seedling growth increased by 90% and the shoot:∗∗∗root ratio changed from 1.561 to 0.607. Shade seedlings were also taller and produced fewer, larger, and thinner leaves than seedlings grown in full sun. A field survey of the spatial distribution and crown morphology of saplings and young adults found 1) the distance between young valley oaks and neighboring overstory trees to increase with neighbor size, and 2) crowns of the young oaks to be skewed away from neighbors. Although shading by the canopy was only moderate, canopy effects on understory light supply may restrict juvenile recruitment of valley oak in this woodland.     

Canopy disturbance patterns and regeneration of Quercus species in two Ohio old-growth forests

This study was conducted to determine the abundance of Quercus species, the spatial pattern of Quercus regeneration, the current canopy disturbance pattern, and their interrelationship in two old-growth deciduous forests in Ohio (Goll Woods and Sears-Carmean Woods). Acer saccharum and Fagus grandifolia had the greatest density and basal area in both forests, yet the largest trees (by basal area) present at each site were Quercus spp. Quercus spp. appeared to be decreasing in abundance in both sites. Though Quercus seedlings were common, few Quercus saplings or subcanopy trees were present. The current disturbance regimes were dominated by small canopy gaps created by death of 1–2 trees; canopy gaps 100 m² in size were rare and only 2.5–2.8% of the forest area was covered by recognizable canopy gaps. No significant differences in the density of Quercus seedlings or saplings were found between gaps and non-gap areas at either site. Though no significant barrier to seedling establishment appeared to exist, the present disturbance regimes are not well suited for the growth of Quercus into the subcanopy size class or the recruitment of Quercus into the canopy. The most frequent gapmakers in Goll Woods were Tilia americana and Acer saccharum, and those in Sears-Carmean Woods were A. saccharum and F. grandifolia. The species most frequent as gap fillers were A. saccharum (in both sites) and F. grandifolia (in Goll Woods). These results suggest that A. saccharum will continue to increase in abundance, and Quercus decrease in abundance, in these two old-growth stands.     

Small gap dynamics in the southern boreal forest of eastern Canada: Do canopy gaps influence stand development?

Question: To what extent do small‐scale disturbances in the forest canopy, created by natural disturbance agents, affect stand development? Doubts exist as to whether small canopy openings have any real effect on the understory tree recruitment, especially in boreal forests. Location: Conifer and mixed stands in the Gaspesian region in eastern Québec. The main natural disturbance agents are recurring outbreaks of Choristoneura fumiferana (eastern spruce budworm) and winds. Methods: Linear transects in 27 sites were used to describe the gap (< 0.1 ha) regime parameters, including gap fraction, gap size and change in disturbance severity through time. Three stand types were distinguished, based on a gradient of abundance of tree host species for the eastern spruce budworm. The impact of gaps was evaluated on the basis of changes in the number, the period of recruitment, and the composition of tree saplings present within gap areas. Changes were measured along the gap size gradient, and according to the pattern of recent budworm epidemics. Results: The gap fraction is highly variable (18%‐64%) and is on average relatively high (42%). Gap sizes have a positively skewed distribution. In most cases the growth rate among gap filling saplings increased sufficiently to date disturbance events. The composition and the structure of understory trees were affected by gap formation. The number of shade‐intolerant tree species did increase during or following periods of particularly severe canopy disturbances. However, the establishment or survival of shade intolerant species was not restricted to larger gaps or more intensely disturbed periods. Conclusions: In sub‐boreal forests of Eastern Canada, small scale disturbances in the tree canopy influence stand regeneration dynamics, but not to the extent that parameters such as sapling composition and recruitment patterns depend on gap regime characteristics.     

Shoot growth and tree architecture of saplings of the major canopy dominants in a warm-temperate rainforest

Sapling density, shoot growth, and sapling architecture were studied in five major canopy dominants both under closed canopy and gaps in a warm-temperate rainforest. The five species showed wide variations in distribution, shoot growth, and sapling architecture. Distylium racemosum and Quercus acuta had significantly higher sapling densities under closed canopy than in gaps. Castanopsis sieboldii and Machilus thunbergii had significantly higher sapling densities in gaps than under closed canopy. Quercus salicina showed no significant difference in sapling density between the two habitats. Under closed canopy, C. sieboldii and M. thunbergii had wider crowns than the other species. Distylium racemosum had the greatest number of terminal shoots among the species. Quercus acuta had a branchless small crown. Quercus salicina showed intermediate values in crown width, depth and the number of terminal shoots among the species. Distylium racemosum showed the greatest height-growth rate among the species under the closed canopy, but was the slowest in gaps. Castanopsis sieboldii and M. thunbergii showed the greatest height-growth rates among the species in gaps. Quercus salicina showed the slowest height-growth rates both under closed canopy and gaps. All of the five species showed low mortality under closed canopy. For the major canopy dominants: (i) sapling architecture may not be an important factor in determining mortality but it may be important for height-growth rate; and (ii) sprouting helps saplings to survive until gap formation and facilitates rapid growth in the gaps.     

Gap-scale disturbance processes in secondary hardwood stands on the Cumberland Plateau,Tennessee, USA

Disturbance regimes in many temperate, old growth forests are characterized by gap-scale events. However, prior to a complex stage of development, canopy gaps may still serve as mechanisms for canopy tree replacement and stand structural changes associated with older forests. We investigated 40 canopy gaps in secondary hardwood stands on the Cumberland Plateau in Tennessee to analyze gap-scale disturbance processes in developing forests. Gap origin, age, land fraction, size, shape, orientation, and gap maker characteristics were documented to investigate gap formation mechanisms and physical gap attributes. We also quantified density and diversity within gaps, gap closure, and gap-phase replacement to examine the influence of localized disturbances on forest development. The majority of canopy gaps were single-treefall events caused by uprooted or snapped stems. The fraction of the forest in canopy gaps was within the range reported from old growth remnants throughout the region. However, gap size was smaller in the developing stands, indicating that secondary forests contain a higher density of smaller gaps. The majority of canopy gaps were projected to close by lateral crown expansion rather than height growth of subcanopy individuals. However, canopy gaps still provided a means for understory trees to recruit to larger size classes. This process may allow overtopped trees to reach intermediate positions, and eventually the canopy, after future disturbance events. Over half of the trees located in true gaps with intermediate crown classifications were Acer saccharum, A. rubrum, or Liriodendron tulipifera. Because the gaps were relatively small and close by lateral branch growth of perimeter trees, the most shade-tolerant A. saccharum has the greatest probability of becoming dominant in the canopy under the current disturbance regime. Half of the gap maker trees removed from the canopy were Quercus; however, Acer species are the most probable replacement trees. These data indicate that canopy gaps are important drivers of forest change prior to a complex stage of development. Even in relatively young forests, gaps provide the mechanisms for stands to develop a complex structure, and may be used to explain patterns of shifting species composition in secondary forests of eastern North America.     

Drought-induced mortality affects understory vegetation: release after death

In recent decades, severe droughts have become an important cause of canopy disturbance in forests, and have shown potential to cause rapid and pronounced vegetation shifts. Under dead canopy, undamaged understory could influence the nature of resource limitation for seedling growth and survival, limiting forest regeneration. We assessed the release response of understory vegetation after a severe drought event in temperate forests of northern Patagonia. Growth trends of dominant tree saplings, and changes in vegetation biodiversity and cover were compared between drought-dead and unaffected canopy. Nothofagus dombeyi undergo growth release after the climatic event in affected forests, and the response was evidenced immediately after the disturbance. For Austrocedrus chilensis, the growth release response was less evident, due mainly to a difference in age structure. In the understory the release response was barely discernable for some components. There was a tendency towards higher cover of the shrub layer in the understory of drought-affected forests, and an important presence of the exotic shrub Rosa rubiginosa. However, the clearest biotic response following drought mortality was the release in growth of understory dominant tree component. Those results strongly suggest that the environment under drought-dead canopy, and the die-off in woody sapling cohorts in a self-thinning process, could favor crown expansion and growth release of understory species that could help predict future forest trajectories in the context of the influence of climatic extreme events.     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

Comparison of the physiology, morphology, and leaf demography of tropical saplings with different crown shapes

Branch architecture, leaf photosynthetic traits, and leaf demography were investigated in saplings of two woody species, Homolanthus caloneurus and Macaranga rostulata, co-occurring in the understory of a tropical mountain forest. M. rostulata saplings have cylindrical crowns, whereas H. caloneurus saplings have flat crowns. Saplings of the two species were found not to differ in area-based photosynthetic traits and in average light conditions in the understory of the studied site, but they do differ in internode length, leaf emergence rate, leaf lifespan, and total leaf area. Displayed leaf area of H. caloneurus saplings, which have the more rapid leaf emergence, was smaller than that of M. rostulata saplings, which have a longer leaf lifespan and larger total leaf area, although M. rostulata saplings showed a higher degree of leaf overlap. Short leaf lifespan and consequent small total leaf area would be linked to leaf overlap avoidance in the densely packed flat H. caloneurus crown. In contrast, M. rostulata saplings maintained a large total leaf area by producing leaves with a long leaf lifespan. In these understory saplings with a different crown architecture, we observed two contrasting adaptation strategies to shade which are achieved by adjusting a suite of morphological and leaf demographic characters. Each understory species has a suite of morphological traits and leaf demography specific to its architecture, thus attaining leaf overlap avoidance or large total leaf area.     

Allometry, root/shoot ratio and root architecture in understory saplings of deciduous dicotyledonous trees in central Japan

Plant allometry that is related to plant architecture and biomass allocation strongly influences a plants ability to grow in shaded forest understory. Some allometric traits can change with plant size. The present study compared crown and trunk allometries, root/shoot biomass allometry, and root architecture among understory saplings (0.25--5m height, except for two trees > 5 < 7 m) of seven deciduous dicotyledonous species in central Japan. Associations of the crown and trunk allometries with several plant morphological attributes were analyzed. Branch morphology (plagiotropyvs orthotropy) and life size were correlated with sapling crown and trunk allometries. Both large leaves and orthotropic branches were associated with a narrow small crown and slender trunk. The root/shoot ratio decreased rapidly with increasing plant height for saplings shorter than about 1.5 m. Less shade-tolerant species tended to have smaller root/shoot ratios for saplings taller than 1.5 m. With an increase in plant height, the branch/trunk biomass ratio decreased for saplings with plagiotropic branches but increased for saplings with orthotropic branches. Four subcanopy species (Acer distylum, Carpinus cordata, Fraxinus lanuginosa and Acanthopanax sciadophylloides) had superficial root systems; a common understory species (Sapium japonica) had a deep tap root system; and a canopy species (Magnolia obovata) and a subcanopy species (Acer tenuifolium) had heart root systems of intermediate depth. The root depth was not related to shade tolerance. Among species of the same height, the difference in fine root length can be 30-fold.     

Comparative studies on acorn and seedling dynamics of fourQuercus species in an evergreen broad-leaved forest

The dynamics and regeneration probabilities ofQuercus acuta, Q. gilva, Q. salicina, andQ. sessilifolia were studied in a 0.48 ha plot in an old-growth evergreen broad-leaved forest, Kagoshima prefecture, southwestern Japan. The canopy was well closed and had only one small gap. Seventeen tree species occurred in the canopy. The fourQuercus species ranked second to fifth in basal area. Excluding the trees originated from sprouts,Q. acuta and Q. gilva lacked trees at the subcanopy layer, andQ. salicina andQ. sessilifolia had some trees at the same layer. For eachQuercus species, the amount of fallen acorns fluctuated from year to year. The number of current year seedlings was positively correlated with the number of fallen acorns in the previous year. A disproportion between the spatial distribution of fallen acorns and that of established seedlings has been shown. The establishment of seedlings was frequently observed where no acorns had fallen in the previous year. Saplings of four species survived for several years and formed sapling banks under the closed canopy. During the three years of the study, the mortality ofQ. salicina andQ. sessilifolia saplings was lower than that ofQ. acuta andQ. gilva. However, the differences in the average annual height growth of saplings between four species were not significant, and the most of them did not grow much. These results suggest that the saplings of the fourQuercus, especiallyQ. gilva andQ. acuta, have no chance to recruit to the canopy or subcanopy layer within a closed stand, and that some changes in the environment caused by gap formation are required for their recruitment.     

Methods for estimating understory light conditions using crown projection maps and topographic data

Four types of indices of canopy characteristics were compared to estimate understory light conditions using crown projection maps and topographic data. Crown area (CA) was formulated from the crowns in a focal sub-block (20 m×20 m). Canopy shade index (CSI) was formulated from the crowns in a focal sub-block and the surrounding sub-blocks with topographic effect. Site shade index (SSI) incorporated the shade of the slopes into CSI and represented the total physical closure of the site. Site light index (SLI) incorporated the azimuth of the site into SSI and evaluated the light condition of the understory. We tested the methods using the data on steep old-growth Cryptomeria japonica forests on Yakushima Island and the three-dimensional canopy structures of the forests were digitized on a geographic information system. The results were validated by the basic indices obtained by hemispherical photographs: canopy openness (CO) and gap light index (GLI). The three-dimensional structure of the surrounding canopies and topography had significant effects on the understory light conditions. CA had no significant correlation with CO, whereas CSI and SSI had significant negative correlations with CO. Although SLI had the highest negative correlation with GLI (R²=0.69), CSI had practical utility in the estimation of GLI (R²= 0.65). Our method is applicable to past crown projection maps that have been recorded at many forest research sites, thus making it possible to reconstruct past understory light conditions and to compare them with present conditions.     

Introduced deer reduce native plant cover and facilitate invasion of non-native tree species: evidence for invasional meltdown

Invasive species are a major threat to native communities and ecosystems worldwide. One factor frequently invoked to explain the invasiveness of exotic species is their release in the new habitat from control by natural enemies (enemy-release hypothesis). More recently, interactions between exotic species have been proposed as a potential mechanism to facilitate invasions (invasional meltdown hypothesis). We studied the effects of introduced deer on native plant communities and exotic plant species on an island in Patagonia, Argentina using five 400 m² exclosures paired with control areas in an Austrocedrus chilensis native forest stand. We hypothesized that introduced deer modify native understory composition and abundance and facilitate invasion of introduced tree species that have been widely planted in the region. After 4 years of deer exclusion, native Austrocedrus and exotic Pseudotsuga menziesii tree sapling abundances are not different inside and outside exclosures. However, deer browsing has strongly inhibited growth of native tree saplings (relative height growth is 77% lower with deer present), while exotic tree sapling growth is less affected (relative height growth is 3.3% lower). Deer significantly change abundance and composition of native understory plants. Cover of native plants in exclosures increased while cover in controls remained constant. Understory composition in exclosures after only 4 years differs greatly from that in controls, mainly owing to the abundance of highly-browsed native species. This study shows that introduced deer can aid the invasion of non-native tree species through negatively affecting native plant species.     

Architecture of Iberian canopy tree species in relation to wood density,shade tolerance and climate

Tree architecture has important consequences for tree performance as it determines resource capture, mechanical stability and dominance over competitors. We analyzed architectural relationships between stem and crown dimensions for 13 dominant Iberian canopy tree species belonging to the Pinaceae (six Pinus species) and Fagaceae (six Quercus species and Fagus sylvatica) and related these architectural traits to wood density, shade tolerance and climatic factors. Fagaceae had, compared with Pinaceae, denser wood, saplings with wider crowns and adults with larger maximal crown size but smaller maximal height. In combination, these traits enhance light acquisition and persistence in shaded environments; thus, contributing to their shade tolerance. Pinaceae species, in contrast, had low-density wood, allocate more resources to the formation of the central trunk rather than to branches and attained taller maximal heights, allowing them to grow rapidly in height and compete for light following disturbances; thus, contributing to their high light requirements. Wood density had a strong relationship with tree architecture, with dense-wooded species having smaller maximum height and wider crowns, probably because of cheaper expansion costs for producing biomechanically stable branches. Species from arid environments had shorter stems and shallower crowns for a given stem diameter, probably to reduce hydraulic path length and assure water transport. Wood density is an important correlate of variation in tree architecture between species and the two dominant families, with potentially large implications for their resource foraging strategies and successional dynamics.     

The limited role of canopy gaps in the successional dynamics of a mature mixed Quercus forest remnant

Question: Are canopy gap dynamics responsible for driving the structural and compositional changes that have occurred over a 26‐year period in a mature Quercus forest remnant? Location: Dobbs Natural Area, an unlogged 3.6‐ha forest preserve in west‐central Indiana, USA. Methods: We analyzed mapped permanent plot data for a site that illustrates a trend common in Quercus‐dominated forests in eastern North America, where recruitment of new stems is dominated by mesophytic, shade‐tolerant species such as Acer saccharum, rather than Quercus. We developed a GIS database from stand census measurements taken in 1974 and 2000, employing it to conduct tree‐by‐tree comparisons that allow direct determination of ingrowth, mortality and survivorship, and to relate the spatial patterns of subcanopy dynamics to canopy gap occurrence. Results: The re‐census shows modest changes in canopy composition, but much greater turnover in the subcanopy. Nearly half of all individuals originally present died; much of this mortality resulted from a major decline in subcanopy Ulmus americana. While overall density remained fairly constant, the subcanopy experienced substantial ingrowth of shade‐tolerant Acer saccharum, Fagus grandifolia, and Tilia americana. Canopy gaps, although forming at rates in the upper range of regional averages, did not significantly benefit subcanopy populations of Quercus spp. or most other taxa with limited shade tolerance. Conclusions: Canopy gaps play a minor role in driving the recent demographic trends of this stand. The spatial and temporal scales of light availability in gaps do not support regeneration of most shade‐intolerant species. Compositional change parallels a historical shift in light regimes.