Losses of seeds of temperate trees to soil fungi: effects of habitat and host ecology

Evidence suggests that impacts of fungal pathogens on tree recruitment tend to be greater in the forest understory than in openings, and that shade-tolerant trees are less vulnerable than shade-intolerant species. To investigate the role that harmful soil fungi may have in reducing regeneration of temperate trees, we applied fungicide to buried seeds of matched pairs of species differing in their relative shade tolerance and/or successional status (Acer negundo versus Acer saccharum, Prunus virginiana versus Prunus serotina, and Pinus strobus versus Tsuga canadensis), in three habitats that differed in their degree of openness (old field, forest gap, and forest understory). Our results indicated that soil fungi reduced germination of A. negundo, P. virginiana, P. serotina. and T. canadensis, and reduced viability of ungerminated seeds of P. strobus; no significant effects of fungi on seeds of A. saccharum were detected. However, we found seeds were not less likely to survive following burial in forest understory than in gaps. As well, results for only one species pair (A. negundo versus A. saccharum) were consistent with the prediction that shade-intolerant or successional species should be more susceptible to fungal attack than mature forest species. These results contrast with other studies of temperate and especially tropical forest trees.     

Growth, Allocation and Water Relations of Shade-grownQuercus rubraL. Saplings Exposed to a Late-season Canopy Gap

For understory saplings to exploit canopy gaps successfully,carbon gain must increase in the gap environment. We predictedthat total biomass of shade-grown red oak saplings would increaseafter exposure to a late-season canopy gap, and that increasedwater and nutrient demand within the canopy gap would drivechanges in the allocation of this carbon. Shade-grown red oaksaplings acclimated to gaps by increasing biomass during theseason of gap formation and increasing the potential for carbongain in the following summer. Within-season carbon gain didnot result from greater production of leaf area, so it mostlikely arose from higher photosynthetic rates of existing shade-developedfoliage, which may be linked to accumulation of leaf nitrogen.During the season of gap formation, shade-gap plants increasedallocation to storage of total non-structural carbohydrates(TNC), and to root growth. The increase in TNC storage suggeststhat shade-developed saplings exposed to gaps were also primedfor fast growth and carbon gain in the following summer. Theincrease in root growth suggests that higher nutrient and waterdemand drove allocation shifts to enhance the capacity for nutrientand water uptake in the gap. Plant hydraulic conductivity (K_a)of shade-grown plants was limited upon exposure to the gap,possibly because of embolism formation resulting from the abruptincrease in water demand. Greater water potential gradientscompensated for limitations to K_a, allowing saplings to maintainhigh transpiration rates, suggesting that actual water uptakeof shade-gap plants was unaffected by gap exposure. Acclimation; canopy gaps; carbon allocation; hydraulic conductivity; nitrogen allocation; non-structural carbohydrates;Quercus rubra L.; red oak; stomatal conductance; transpiration; water-relations     

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.     

Phylogeny and the patterns of leaf phenolics in gap-and forest-adapted Piper and Miconia understory shrubs

Summary The types and quantities of defense compounds found in plants occupying ecologically distinct habitats have received much theoretical and little empirical attention. Here we characterize the leaf phenolic chemistry of eight species in two genera of tropical rainforest shrubs; four species in each genus are typical of disturbed sites, and four are typical of mature forest understory. Two Miconia species growing in light gaps had significantly higher leaf tannin and total phenolic contents than congenors growing in the primary forest; this pattern was not found among the gap- and forest-adapted Piper species. Tannin patterns were not mirrored by leaf cinnamic acids. These results indicate that plant phylogeny must be considered when predicting plant defense investment.     

Gap size and succession in cutover southern Appalachian forests: an 18 year study of vegetation dynamics

We used clearcut logging in establishing four replicated sizes of canopy openings (0.016, 0.08, 0.4, and 2.0 ha) in a southern Appalachian hardwood forest in 1981 to examine the long-term effects of disturbance size on plant community structure, biomass accumulation, aboveground net primary productivity (NPP), and mode of recovery. The reestablishment of NPP and biomass following logging was 6–7-fold greater in large than small openings by 17 years. Total biomass in the 2.0 ha openings (127.3 Mg ha⁻¹) recovered 59.5% as NPP (19.7 Mg ha⁻¹ yr⁻¹) reached 225% of precut forest levels. Biomass accumulation was 2.6–3.6-fold greater in interior than edge locations of all but the 0.016 ha gaps. The absence of significant patch size or edge vs. interior differences in tree densities suggests that growth rates of individual trees were enhanced in more insolated microenvironments. Sprouting (86–95% of tree NPP) was much more important than advance regeneration (4–10%) or seedling germination (<2%) during early recovery in all opening sizes. Canopy dominant Quercus and Carya trees exhibited limited sprouting following disturbance. Instead, shade-intolerant Robinia pseudoacacia and Liriodendron tulipifera were major sprouters that used N-fixation (Robinia) and rapid growth (Liriodendron) in attaining 7.4 and 5.9 fold greater biomass accumulation, respectively in 2.0 ha than 0.016 ha opening sizes. Seedling germination and understory production were extensive in all openings following logging, but declined rapidly as the young tree canopy began closing by 4–6 years. The relative importance of shade-intolerant tree biomass approximately doubled over 17 years as shade-tolerant tree seedlings, herbs, and shrubs gradually regained importance under the emerging canopy. Sprouting caused the persistence of a tree species composition in all openings that remained relatively similar to the precut forest. Large disturbances on mountain slopes of the southern Appalachians generally promote sprouting and rapid recovery, whereas small disturbances in low-elevation cove forests lead to a gradual recovery through seedling germination and/or advance regeneration. Continued logging in the southern Appalachians will increase the relative size and frequency of large disturbances, further the importance of sprouting of shade-intolerant species, and lead to more even-aged forest stands throughout the region.     

Sapling growth and survivorship as affected by light and flooding in a river floodplain forest of southeast Texas

We investigated the effects of light and flooding on growth and survivorship of saplings in a river floodplain forest of southeast Texas. Growth responses to light were consistent with the expectation that shade-intolerant species grow faster than shade-tolerant species in high light, and vice versa. Mortality risk was not associated with shade tolerance level unless high mortality risks associated with a period of high flooding were removed. These results support the hypothesis that shade-tolerant species in floodplains may be limited by flooding as previous studies suggested. Also, compared to their performance at a nearby mesic site, common species showed little intraspecific difference in shade tolerance, especially for shade-intolerant species. Finally, the positive correlation between low-light growth and survivorship suggests that carbon allocation to continued growth may be favored as a sapling strategy in floodplains.     

Growth,biomass distribution and CO2 exchange of northern hardwood seedlings in high and low light: relationships with successional status and shade tolerance

The physiology, morphology and growth of first-year Betula papyrifera Marsh., Betula alleghaniensis Britton, Ostrya virginiana (Mill.) K. Koch, Acer saccharum Marsh., and Quercus rubra L. seedlings, which differ widely in reported successional affinity and shade tolerance, were compared in a controlled high-resource environment. Relative to late-successional, shade-tolerant Acer and Ostrya species, early-successional, shade-intolerant Betula species had high relative growth rates (RGR) and high rates of photosynthesis, nitrogen uptake and respiration when grown in high light. Fire-adapted Quercus rubra had intermediate photosynthetic rates, but had the lowest RGR and leaf area ratio and the highest root weight ratio of any species. Interspecific variation in RGR in high light was positively correlated with allocation to leaves and rates of photosynthesis and respiration, and negatively related to seed mass and leaf mass per unit area. Despite higher respiration rates, early-successional Betula papyrifera lost a lower percentage of daily photosynthetic CO₂ gain to respiration than other species in high light. A subset comprised of the three Betulaceae family members was also grown in low light. As in high light, low-light grown Betula species had higher growth rates than tolerant Ostrya virainiana. The rapid growth habit of sarly-successional species in low light was associated with a higher proportion of biomass distributed to leaves, lower leaf mass per unit area, a lower proportion of biomass in roots, and a greater height per unit stem mass. Variation in these traits is discussed in terms of reported species ecologies in a resource availability context.     

Tree regeneration responses in a lowland Nothofagus-dominated forest after bamboo dieback in South-Central Chile

The bamboo Chusquea quila (Poaceae:Bambuseae) is the most abundant understory species ofNothofagus-dominated forests at low elevations inthe Chilean Lake District. Species of this genus strongly inhibit theestablishment and growth of tree species, especially those of the genusNothofagus. At intervals of many years, this bambooflowerssynchronously and dies, creating large-scale disturbances. The mainobjective of this study was to determine the influence of bamboo dieback ontreeregeneration, especially of shade-intolerant species. The forest studiedis a remnant stand of old-growth forest dominated by emergent individualsof N. obliqua and Eucryphiacordifoliawhich project about 10 m above a main canopy formed byAextoxicon punctatum, Laureliaphilippiana, and Podocarpus saligna. Treeseedlings that established prior to bamboo dieback were recorded in six gapsdensely covered by bamboo. After the dieback event in the early 1990's,recruitment, height growth and survivorship were monitored in four gaps duringtwo growing seasons. After the synchronous mass flowering and death ofChusqueaquila, both reorganization of advance regeneration and new seedlingrecruitment were observed in gaps. The advance regeneration consisted mainly ofthe shade-tolerant species A. punctatum,L. philippiana, and Amomyrtus luma.These species together with E. cordifolia accounted forover 90% of the total individuals in gaps. During the following twoseasons, c. 40% of the advance regeneration either died or was damaged.Root suckers of E. cordifolia and L.philippiana were taller and grew faster than the seedlings of otherspecies. Nothofagus obliqua was unable to establishadvanceregeneration in gaps formerly dominated by bamboo. New recruitment resulted inthe synchronized establishment of treeseedlings, especially less shade-tolerant species such as N.obliqua, Aristotelia chilensis,Rhaphithamnus spinosus, and E.cordifolia. These species recruited into the gaps almost entirelyduring the first season right after the bamboo dieback. However, by the end ofthe second season, these species had a lower density due to a combination oflower recruitment rate, shortened recruitment period, and greater mortalityratecompared to the shade-tolerant species. New bamboo seedlings grew fasterand were more abundant than tree seedlings, except for root suckers (bothnew establishment and advance regeneration) of E.cordifolia which is more likely to successfully recruit into thecanopy. Seedlings of N. obliqua compete poorly with bambooseedlings; its successful recruitment may require bamboo flowering coincidentwith a mast seed year for the tree species.     

Multiple canopy opening effects on recruited saplings in a typhoon-disturbed tropical rainforest,Taiwan

To investigate the influence of multiple canopy openings on the composition and diversity of recruited saplings in a forest frequently disturbed by typhoons. We conducted tree-by-tree censuses (diameter at breast height ≥ 1 cm) and mapped gaps (canopy height < 5 m) in 1993, 2000, 2008, and 2013 in a tropical mountain zonal foothill evergreen broad-leaved forest in Nanjenshan Nature Reserve, Taiwan. We analyzed the composition and diversity of recruited saplings within a 2.1 ha plot (840 sampling quadrats (5 m × 5 m)) with variable numbers of canopy openings recorded during the study period. Composition of recruited saplings was dissimilar between quadrats that stayed opened and those that stayed closed throughout the study period (pairwise similarity estimates C₀₂ = 0.52, 95% CI = 0.38–0.66). The quadrats under closed canopy had high diversity when weighting rare species (species richness), whereas quadrats with one or two gap opening records during the past 20 years had high diversity when weighting the abundance of species. Although canopy openings provided establishment conditions for saplings of some shade-intolerant species, due to the nature of small gap size, such habitats do not favor the dominance of shade-intolerant species. Even in a frequently disturbed forest, species composition and richness of recruited saplings were mainly contributed by shade-tolerant species. Although multiple canopy openings facilitated the establishment of shade-intolerant species, species diversity in the study forests is possibly mainly mediated by coexistence mechanisms of those shade-tolerant species rather than light-gap-related species strategies.     

小兴安岭不同耐荫性树种枝叶性状变异及权衡

随着植株生长,不同耐荫性树种枝叶性状对资源的响应策略存在差异。探究不同耐荫性树种在不同径级间枝叶性状变异及其相关关系,对理解植物功能性状种内和种间变异以及植物对资源的响应策略具有重要意义。以黑龙江凉水国家级自然保护区阔叶红松（Pinus koraiensis）林中不同径级（小树、中等树和大树）的喜光树种（白桦Betula platyphylla、枫桦Betula costata）和耐荫树种（春榆Ulmus japonica、紫椴Tilia amurensis、色木槭Acer mono）为研究对象,测定其出叶强度、枝横截面积、枝干重、单叶面积、总叶面积、总叶干重共6个枝叶性状。利用单因素方差（LSD）分析检验不同耐荫性树种在不同径级间其枝、叶性状是否存在显著差异;以标准化主轴估计（SMA）对不同耐荫性树种枝叶性状间相关关系进行分析。结果表明：除枝干重外,耐荫树种枝叶性状均大于喜光树种;不同耐荫性树种大树枝横截面积均最大,中等树叶面积均最小;喜光树种不同径级间枝干重、总叶干重和出叶强度无显著差异,而耐荫树种均存在显著差异。随径级增大,不同耐荫性树种枝横截面积与叶面积均呈正异速生长关系,且除耐荫树种枝横截面积与单叶面积外,均存在共同斜率;不同耐荫性树种出叶强度与单叶面积存在共同斜率为-0.51和0.47,小树和中等树枝横截面积与总叶面积和总叶干重的异速生长指数均与1存在显著差异,而大树与1无显著差异。结果表明：不同耐荫性树种在不同径级间对资源的获取策略存在差异,不同耐荫性树种小树阶段均表现出资源获取策略,大树阶段则表现出资源保守型策略。与耐荫树种相比,喜光树种表现为快速生长策略,而随着径级增大不同耐荫性树种这种差异逐渐消失,本研究结果为森林演替过程中种内和种间的相互作用提供了一个新的视角。     

When sex is not enough: ecological correlates of resprouting capacity in congeneric tropical forest shrubs

In moist tropical forests resprouting may be an important component of life history, contributing to asexual reproduction through the clonal spread of individuals derived from shoot fragments. However, in contrast to other ecosystems where resprouting is common, the ecological correlates of resprouting capacity in tropical forests remain largely unexplored. In this study we characterized shade tolerance, resprouting capacity and sexual reproductive success of eight co-occurring Piper species from lowland forests of Panama. In field experiments we found that shade-tolerant Piper species had a higher capacity to regenerate from excised or pinned stem fragments than light-demanding species in both gap and understory light conditions. In contrast, shade-tolerant species had lower recruitment probabilities from seeds, as a consequence of lower initial seed viability, and lower seedling emergence rates. All Piper species needed gap conditions for successful seedling establishment. Of 8,000 seeds sown in the understory only 0.2% emerged. In gaps, seed germination of light-demanding species was between 10 and 50%, whereas for shade-tolerant species it was 0.5–9.8%. We propose that the capacity to reproduce asexually from resprouts could be adaptive for shade-tolerant species that are constantly exposed to damage from falling litter in the understory. Resprouting may allow Piper populations to persist and spread despite the high rate of pre-dispersal seed predation and low seed emergence rates. Across Piper species, we detected a trade-off between resprouting capacity and the annual viable seed production per plant but not with annual seed mass produced per plant. This suggests that species differences in sexual reproductive success may not necessarily result from differential resource allocation. Instead we suggest that low sexual reproductive success in the understory may in part reflect reduced genetic diversity in populations undergoing clonal growth, resulting in self-fertilization and in-breeding depression.     

Ontogenetic Changes in Carbohydrate Storage and Sprouting Ability in Pioneer Tree Species in Peninsular Malaysia

Sprouting ability is highly variable among different tree species. In many cases, there are trade‐offs in carbon allocations between growth and storage in seedlings. However, this trade‐off is likely to change with growth stages from seedling to mature plant because carbon investments in reproductive activities and/or risk of disturbance also change by species and growth stage. To examine how sprouting ability and carbohydrate storage change with growth stage, we compared two tropical secondary‐forest trees, Macaranga bancana and M. gigantea, which have different ecological traits. Maximum tree size and growth rate are higher in M. gigantea. We monitored sprout growth and stored resources, including total non‐structural carbohydrate (TNC) and nitrogen in the root, among different tree sizes for 12 months following stem‐cutting treatment. Sprouting ability (total sprout mass) and TNC concentrations were significantly higher in small individuals than in larger specimens in both species. TNC concentration decreased in all size classes after stem cutting. Macaranga bancana had greater sprout survivorship than M. gigantea, which had higher sprouting ability in larger tree‐size classes. Thus, sprouting ability likely depends on root TNC concentration and tree‐size class in both Macaranga species. Higher TNC concentration and sprout survival rates in M. bancana may be related to greater carbon allocation in survival than in growth. This hypothesis is consistent with the ecological traits of M. bancana, such as its growth rate, which was lower than that of M. gigantea.     

Carbohydrate storage,survival, and growth of two evergreen <Emphasis Type="Italic">Nothofagus</Emphasis> species in two contrasting light environments

A number of traits have been attributed important roles in tolerance of shade by plants. Some explanations emphasize traits enhancing net carbon gain; others emphasize energy conservation traits such as storage of non-structural carbohydrates (NSC). To date, cross-species studies have provided mixed support for the role of NSC storage in low-light survival. We examined NSC status, survival, biomass, and growth of large seedlings of two evergreen species of differing shade tolerance (Nothofagus nitida and N. dombeyi) grown in deep shade and 50% light for two growing seasons. We expected to find higher NSC concentration in the more shade-tolerant N. nitida and since allocation to storage involves sacrificing growth, higher growth rate in the shade-intolerant N. dombeyi. NSC concentration of both species was >twofold higher in 50% light than in deep shade, and in roots and stems did not differ significantly between species in either environment. NSC contents per plant were also similar between dead and living plants in deep shade. N. dombeyi outgrew N. nitida in 50% light, while this pattern was reversed in deep shade. Survival in deep shade was not correlated with NSC concentration. Leaf mass fraction was similar between species in 50% light, but lower in N. dombeyi in deep shade. Results provide little evidence of a link between carbohydrate storage and low-light survival in Nothofagus species, and support the view that understorey survival is primarily a function of net carbon gain. Patterns of variation in NSC concentration of the temperate species we studied are likely dominated by more important influences than adaptation to shade, such as limitation of growth or adaptation to cold stress.     

The role of carbohydrate reserves in the growth, resilience, and persistence of cabbage palm seedlings (Sabal palmetto)

Sabal palmetto (Walt.) Lodd. ex Schultes (cabbage palm) is an arborescent palm common in many plant communities throughout Florida, U.S.A., and the Caribbean. Although its seedlings grow very slowly in forest understories, they survive damage and defoliation well, and the species may increase in dominance following disturbances such as fire, logging, and hurricanes. We investigated the potential importance of total nonstructural carbohydrate (TNC) pools in the ability of cabbage palm seedlings to recover from the loss of aboveground tissue such as that caused by fire, grazing, or shallow burial by storm debris. TNC concentrations in belowground organs of seedlings from a forest understory were high, and TNC pools were sufficient to theoretically replace >50% of a seedling's canopy. The largest fraction of the belowground TNC pool was in stem tissue, where TNC in unclipped plants accounted for 26–54% of stem dry mass. Experimental reduction of TNC pools by repeated defoliation slowed seedling regrowth, and seedlings with inherently smaller pools (smaller seedlings) suffered higher mortality after repeated defoliation than did larger seedlings. Although regrowth and recovery after the loss of aboveground tissue was related to the size of the TNC pool in belowground organs, even the smallest seedlings with the smallest pools had sufficient stores to withstand at least two defoliations at frequent (7-week) intervals. Large belowground TNC pools in S. palmetto seedlings appear to enable them to survive all but the most frequent defoliations (e.g., frequent grazing or mowing). Allocation of resources to these stores, however, may contribute to the slow growth rates of S. palmetto seedlings in natural communities. Received: 13 April 1998 / Accepted: 28 August 1998     

兴安落叶松天然林碳储量及其碳库分配特征

兴安落叶松天然林作为大兴安岭林区的主要植被类型,在森林生态系统碳循环中具有重要的作用。在大兴安岭林区选择不同林龄的兴安落叶松天然林,调查其乔木、灌草、枯落物和土壤,并结合已建立的单木异速生长方程分别计算其碳储量,以期为明确该地区碳库动态及其碳库分配特征提供参考。结果表明,兴安落叶松天然林总碳储量随林龄的增加逐渐增大,由幼龄林到过熟林分别为140.46、186.63、208.64、308.62和341.03 Mg C/hm2,整体表现为碳汇,这主要与乔木碳储量随林龄的增加逐渐增大有关;乔木碳库的变化范围为45.44—212.67 Mg C/hm2,且其占总碳储量的比例也随林龄的增加逐渐增大,由幼龄林的32.60%到过熟林的62.36%;灌草碳储量占总碳储量的比例较小,仅为0.48%—0.93%;枯落物碳库在过熟林中较多,为26.11Mg C/hm2,这与过熟林较少的人为干扰有关;土壤碳储量以幼龄林最小,成熟林最高,分别为78.06和131.93 Mg C/hm2,但这与我国其他地区天然林相比均较低,这与该地区较浅的土壤发生层有关;土壤碳储量随林龄的变化并不明显,但其占总碳储量的比例却随林龄的增加逐渐减小,由幼龄林的56.01%减小到过熟林的29.35%。     

Photosynthetic sunfleck utilization potential of understory saplings growing under elevated CO<Subscript>2</Subscript> in FACE

Few studies have evaluated elevated CO₂ responses of trees in variable light despite its prevalence in forest understories and its potential importance for sapling survival. We studied two shade-tolerant species (Acer rubrum, Cornus florida) and two shade-intolerant species (Liquidambar styraciflua, Liriodendron tulipifera) growing in the understory of a Pinus taeda plantation under ambient and ambient+200 ppm CO₂ in a free air carbon enrichment (FACE) experiment. Photosynthetic and stomatal responses to artificial changes in light intensity were measured on saplings to determine rates of induction gain under saturating light and induction loss under shade. We expected that growth in elevated CO₂ would alter photosynthetic responses to variable light in these understory saplings. The results showed that elevated CO₂ caused the expected enhancement in steady-state photosynthesis in both high and low light, but did not affect overall stomatal conductance or rates of induction gain in the four species. Induction loss after relatively short shade periods (<6 min) was slower in trees grown in elevated CO₂ than in trees grown in ambient CO₂ despite similar decreases in stomatal conductance. As a result leaves grown in elevated CO₂ that maintained induction well in shade had higher carbon gain during subsequent light flecks than was expected from steady-state light response measurements. Thus, when frequent sunflecks maintain stomatal conductance and photosynthetic induction during the day, enhancements of long-term carbon gain by elevated CO₂ could be underestimated by steady-state photosynthetic measures. With respect to species differences, both a tolerant, A. rubrum, and an intolerant species, L. tulipifera, showed rapid induction gain, but A. rubrum also lost induction rapidly (c. 12 min) in shade. These results, as well as those from independent studies in the literature, show that induction dynamics are not closely related to species shade tolerance. Therefore, it cannot be concluded that shade-tolerant species necessarily induce faster in the variable light conditions common in understories. Although our study is the first to examine dynamic photosynthetic responses to variable light in contrasting species in elevated CO₂, studies on ecologically diverse species will be required to establish whether shade-tolerant and -intolerant species show different photosynthetic responses in elevated CO₂ during sunflecks. We conclude that elevated CO₂ affects dynamic gas exchange most strongly via photosynthetic enhancement during induction as well as in the steady state. Received: 1 April 1999 / Accepted: 16 August 1999     

土壤碳库积累与分配对热带森林恢复的响应

为探明土壤有机碳沉积对热带森林恢复的响应过程与机理,选取西双版纳处于不同恢复阶段的热带森林类型(前期的白背桐群落、中期的崖豆藤群落、后期的高檐蒲桃群落)为研究对象,探讨土壤有机碳库各组分积累与分配(微生物量碳储量/总有机碳储量、易氧化有机碳储量/总有机碳储量、惰性有机碳储量/总有机碳储量)的时空变化规律,分析乔木与林下物种的丰富度和多样性、土壤温湿度、容重、pH及氮库(全氮、水解氮、铵氮、硝氮)对土壤有机碳库组分积累与分配的影响。结果表明:(1)热带森林恢复显著促进土壤碳库各组分的蓄积(P<0.05),相较于恢复前期,恢复中后期土壤总有机碳、微生物量碳、易氧化有机碳、惰性有机碳储量增幅达9.25%-50.84%;恢复促进了土壤微生物量碳和易氧化有机碳的分配(8.98%-25.36%)(P<0.05),但对惰性有机碳分配无显著影响;(2)不同恢复阶段热带森林土壤碳组分积累与分配的时空变化存在一定的差异。其中上述4种碳组分积累最大值均出现在6月、垂直变化均沿土层递减;土壤易氧化有机碳和微生物量碳分配最大值出现在6月、惰性有机碳分配则在12月最大,易氧化有机碳和微生物量碳分配沿土层递减、惰性有机碳分配无显著垂直变化;(3)土壤微生物量碳、易氧化有机碳、惰性有机碳的储量在土壤碳库储量的分配占比分别维持在2.40%-5.00%、18.22%-39.34%、18.50%-26.55%,土壤有机碳组分对总有机碳储量变化的解释率表现为:微生物量碳(83.71%)>惰性有机碳(82.17%)>易氧化有机碳(78.54%);(4)相较于恢复初期,恢复后期乔木与林下物种丰富度和Shannon多样性提升了42.78%-490.82%,氮库(全氮、水解氮、铵氮、硝氮)含量仅提升了12.73%-25.51%;(5)冗余分析表明,林下物种丰富度、温湿度、水解氮是影响土壤有机碳组分积累的主要驱动因子,而乔木香农多样性、湿度、容重则是影响土壤有机碳库组分分配的主控因子。因此,西双版纳热带森林恢复进程显著促进了土壤有机碳库组分积累与分配,影响程度取决于样地林下物种丰富度、乔木香农多样性、土壤温湿度、容重与水解氮的状况。     

Canopy cover and tree regeneration in old-growth cove forests of the Appalachian Mountains

Relationships between canopy cover and tree regeneration were determined for various species in cove forests of the Great Smoky Mountains. Old-growth stands were sampled with six plots covering a total area of 4.8 ha. Each plot was subdivided into contiguous 10×10 m quadrats. Canopy cover overlying each of the 480 quadrats was characterized with three different indices based on visual estimates of cover. Influences of: (1) overlying cover, (2) proximate openings, and (3) total area of proximate openings on quadrat regeneration densities were determined. Most species reproducing by seed and some species reproducing by vegetative means had higher densities in quadrats with openings, but only the intolerants were highly dependent on gaps. Tsuga canadensis, a very shade-tolerant species, was one of the few species with abundant regeneration beneath dense canopy cover. In general, understory areas near gaps had somewhat higher regeneration densities than other areas with overlying cover. Several shade-tolerant species showed a positive regeneration density response to canopy openings and an ability to regenerate in gaps 0.01–0.03 ha in area. These openings were too small for intolerant species. Many species exhibited a positive response to total size of the proximate opening(s). A sharp increase in regeneration density with area of the opening(s) was evident at approximately 0.04 ha for the shade-intolerant species.     

Fire-induced shifts in overstory tree species composition and associated understory plant composition in Glacier National Park,Montana

In Rocky Mountain forests, fire can act as a mechanism of change in plant community composition if postfire conditions favor establishment of species other than those that dominated prefire tree communities. We sampled pre and postfire overstory and postfire understory species following recent (1988–2006) stand-replacing fires in Glacier National Park (GNP), Montana. We identified changes in relative density of tree species and groups of species (xerophytes vs. mesophytes and reseeders vs. resprouters) in early succession. Postfire tree seedling densities were adequate to maintain prefire forest structure, but relative densities among species were variously changed. Changes were directly related to individual species’ response to severe fires. Most notably, relative density of the mesophytic resprouter quaking aspen (Populus tremuloides) and the xerophytic reseeder lodgepole pine (Pinus contorta) increased substantially following fire, with a concomitant decline in proportional abundance of other tree species that, in some cases, dominated stands before fire. Trends identified in our study suggest that forest community shifts toward those dominated by lodgepole pine and quaking aspen are occurring in GNP. Cover of understory species was not affected by tree species composition or density. These forest communities will likely change throughout succession with the addition of shade-intolerant species in early seral stages and shade-tolerant species later in succession. However, with increased fire frequency, the lodgepole pine-dominated postfire communities observed in our study may become more common throughout time.     

Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees

Among 13 tropical tree species on Barro Colorado Island, species with high seedling mortality rates during the first year in shade had higher reltive growth rates (RGR) from germination to 2 months in both sun (23% full sun) and shade [2%, with and without lowered red: far red (R:FR) ratio] than shade tolerant species. Species with higher RGR in sun also had higher RGR in shade. These interspecific trends could be explained by differences in morphological traits and allocation paterns among species. Within each light regime, seedlings of shade-intolerant species had lower root: shoot ratios, higher leaf mass per unit area, and higher leaf area ratios (LAR) than shade tolerant species. In contrast, leaf gas exchange characteristics, or acclimation potential in these traits, had no relationship with seedling mortality rates in shade. In both shade tolerant and intolerant species, light saturated photosynthesis rates, dark respiration, and light compensation points were higher for sungrown seedlings than for shade-grown seedlings. Differences in R:FR ratio in shade did not affect gas exchange, allocation patterns, or growth rates of any species. Survival of young tree seedlings in shade did not depend on higher net photosynthesis or biomass accumulation rates in shade. Rather, species with higher RGR died faster in shade than species with lower RGR. This trend could be explained if survival depends on morphological characteristics likely to enhance defense against herbivores and pathogens, such as dense and tough leaves, a well-established root system, and high wood density. High construction costs for these traits, and low LAR as a consequence of these traits, should result in lower rates of whole-plant carbon gain and RGR for shade tolerant species than shade-intolerant species in shade as well as in sun.